Research Article |
Corresponding author: Wayne P. Maddison ( wmaddisn@mail.ubc.ca ) Academic editor: Jeremy Miller
© 2020 Wayne P. Maddison, David R. Maddison, Shahan Derkarabetian, Marshal Hedin.
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, Maddison DR, Derkarabetian S, Hedin M (2020) Sitticine jumping spiders: phylogeny, classification, and chromosomes (Araneae, Salticidae, Sitticini). ZooKeys 925: 1-54. https://doi.org/10.3897/zookeys.925.39691
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The systematics of sitticine jumping spiders is reviewed, with a focus on the Palearctic and Nearctic regions, in order to revise their generic classification, clarify the species of one region (Canada), and study their chromosomes. A genome-wide molecular phylogeny of 23 sitticine species, using more than 700 loci from the arachnid Ultra-Conserved Element (UCE) probeset, confirms the Neotropical origins of sitticines, whose basal divergence separates the new subtribe Aillutticina (a group of five Neotropical genera) from the subtribe Sitticina (five genera of Eurasia and the Americas). The phylogeny shows that most Eurasian sitticines form a relatively recent and rapid radiation, which we unite into the genus Attulus Simon, 1868, consisting of the subgenera Sitticus Simon, 1901 (seven described species), Attulus (41 described species), and Sittilong Prószyński, 2017 (one species). Five species of Attulus occur natively in North America, presumably through dispersals back from the Eurasian radiation, but an additional three species were more recently introduced from Eurasia. Attus palustris Peckham & Peckham, 1883 is considered to be a full synonym of Euophrys floricola C. L. Koch, 1837 (not a distinct subspecies). Attus sylvestris Emerton, 1891 is removed from synonymy and recognized as a senior synonym of Sitticus magnus Chamberlin & Ivie, 1944. Thus, the five native Attulus in North America are Attulus floricola, A. sylvestris, A. cutleri, A. striatus, and A. finschi. The other sitticines of Canada and the U.S.A. are placed in separate genera, all of which arose from a Neotropical radiation including Jollas Simon, 1901 and Tomis F.O.Pickard-Cambridge, 1901: (1) Attinella Banks, 1905 (A. dorsata, A. concolor, A. juniperi), (2) Tomis (T. welchi), and (3) Sittisax Prószyński, 2017 (S. ranieri). All Neotropical and Caribbean “Sitticus” are transferred to either Jollas (12 species total) or Tomis (14 species). Attinella (three species) and Tomis are both removed from synonymy with Sitticus; the synonymy of Sitticus cabellensis Prószyński, 1971 with Pseudattulus kratochvili Caporiacco, 1947 is restored; Pseudattulus Caporiacco, 1947 is synonymized with Tomis. Six generic names are newly synonymized with Attulus and one with Attinella. Two Neotropical species are described as new, Jollas cupreus sp. nov. and Tomis manabita sp. nov. Forty-six new combinations are established and three are restored. Three species synonymies are restored, one is new, and two are rejected. Across this diversity of species is a striking diversification of chromosome complements, with X-autosome fusions occurring at least four times to produce neo-Y sex chromosome systems (X1X2Y and X1X2X3Y), some of which (Sittisax ranieri and S. saxicola) are sufficiently derived as to no longer preserve the simple traces of ancestral X material. The correlated distribution of neo-Y and a base autosome number of 28 suggests that neo-Y origins occurred preferentially in lineages with the presence of an extra pair of autosomes.
Amycoida, karyotype, molecular phylogeny, Salticinae, sex chromosomes
The jumping spider species long placed in the genus Sitticus Simon, 1901 are well known in both Eurasia and the Americas as prominent members of habitats as diverse as boreal forests, marshes, deserts and human habitations (e.g., Locket and Milledge 1951; Prószyński 1968, 1971, 1973, 1980;
Subtribe Aillutticina (1–4) and the Jollas-Tomis clade of the subtribe Sitticina (5–14) 1–4 Aillutticus nitens, Uruguay (-34.877, -56.023): 1–3 male 4 female 5, 6 Tomis palpalis male and female, Ecuador (-0.1996, -77.7023) 7, 8 Jollas species: 7 J. cupreus male, Ecuador (-0.675, -76.397) 8 Jollas sp. female, Ecuador (-0.7223, -77.6408) 9 J. leucoproctus, Uruguay (-34.94, -54.95) 10 J. flabellatus, Uruguay (-34.426, -55.195) 11–14 Attinella dorsata male (11–13) and female (14), Canada (48.870, -123.379). Also included in the Jollas-Tomis clade is Sittisax (Figs
This work’s three goals are to resolve sitticine phylogeny, to review the taxonomy of sitticines of one region (Canada), and to describe the remarkably diverse chromosomes of sitticines. Our immediate (and urgent) purpose in studying the group’s phylogeny is to settle its turbulent generic classification, which has seen, for instance, some well-known species change names three times in two years, for example, from Sitticus floricola (C. L. Koch, 1837) to Sittiflor floricola (by Prószyński 2017a) to Calositticus floricola (by
Until the last few years, most sitticines were placed in the single widespread and species-rich genus Sitticus Simon, 1901 (e.g.,
Neither Prószyński’s “pragmatic” classification nor Breitling’s COI-based classification have promoted taxonomic stability in sitticines. Prószyński’s intentionally non-phylogenetic approach is particularly problematical. The great majority of systematists no longer use such “pragmatic” non-evolutionary classifications, as they are not anchored to a broadly predictive external reality: they are subject to the whims of biologists’ interests and the character systems they focus on. A taxon delimited for this sense of pragmatism carries with it no promise of meaning or utility, other than the promise it will bear the diagnostic characters chosen. Different choices of diagnostic characters would lead to different classifications, with no basis for selecting among different authors’ approaches except the weight of authority – in the end, not as pragmatic as a stable phylogenetic classification, which, by the implications of genetic descent, will predict trait distributions across the genome. Breitling’s approach might have dampened the instability, as it is phylogenetic and uses explicit data and analysis, but his choice of the single gene COI, without supporting morphological information, has yielded a classification in which we can have little confidence. Prószyński’s and Breitling’s reclassifications might have been steps forward had they been done in a group of salticids with almost no previous attention, but the sitticines are reasonably well studied and often mentioned in the literature. These sudden, comprehensive, conflicting, and largely baseless rearrangements of Sitticus have yielded taxonomic instability in a well-known group.
Taxonomic instability yields confusion in ecological and other biodiversity literature about the identity of species studied, and damages the reputation of the taxonomic enterprise. We are now sufficiently capable of resolving phylogeny that we do not need to rely on the “pragmatic” choices of one authority or on a single misbehaving gene. Our goal is to provide stronger evidence, explicitly analyzed, for phylogenetic relationships in order to stabilize the classification of sitticines.
Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Most of the coquille drawings were done in 1977 or 1978 using a reticle grid in a stereomicroscope. Colour drawings were done in 1974 through 1977 with a stereomicroscope and reticle grid. Pen and pencil drawings were made recently using a drawing tube on a Nikon ME600L compound microscope. Because some images were made decades ago, we are unable to supply scale bars on many. Terms used are standard for Araneae. All measurements are given in millimeters. 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”); RTA, retrolateral tibial apophysis of the male palp.
Specimens were examined from the collections of the American Museum of Natural History (
Authors of nomenclatural acts in this paper vary by rank. For acts affecting the synonymy of genera (viz., reinstatement of Attinella and Tomis; synonymies of Sitticus, Pseudattulus and Sittiab), the authors are those of the paper itself. For all other acts, the author is W. Maddison. These include the establishment of the Aillutticina, new subtribe, acts that affect the synonymy and placement of species (new synonyms, restored synonyms, new combinations), and new species.
If not otherwise indicated, the authors of species names are given in the Classification section.
Taxa were sampled to cover a diversity of sitticine species groups from Eurasia, North America, and South America (Table
Specimens from which UCE sequence data gathered. “UCE loci” indicates number of loci from Phyluce. “Reads Pass QC” indicates number of reads retained after quality control and adapter removal via Illumiprocessor.
Species | Specimen | sex | Locality | Reads Pass QC | Contigs | UCE loci |
---|---|---|---|---|---|---|
Aillutticus nitens | d475 | f | Uruguay: Canelones: -34.867, -56.009 | 946351 | 207743 | 434 |
Attinella dorsata | d490 | m | U.S.A.: California: 37.2834, -120.8515 | 1617332 | 360661 | 480 |
Attulus ammophilus | d482 | m | Canada: British Columbia: 49.7963, -119.5338 | 1471891 | 351670 | 588 |
A. burjaticus | RU18-7302 | f | Russia: Tuva: 50.205, 95.135 | 529905 | 151897 | 627 |
A. distinguendus | RU18-6432 | f | Russia: Tuva: 50.746, 93.142 | 406186 | 90846 | 626 |
A. fasciger | d487 | m | Canada: Ontario: 43.35074, -79.75928 | 1370738 | 299273 | 564 |
A. finschi | d480 | m | Canada: British Columbia: 49.0261, -114.0611 | 1489551 | 303924 | 579 |
A. floricola | d488 | m | Canada: Saskatchewan: 52.4898, -107.3843 | 1466702 | 303612 | 606 |
A. inexpectus | RU18-6799 | m | Russia: Tuva: 50.669, 92.9844 | 261947 | 60612 | 653 |
A. longipes | ARV4504 | m | Italy: Stilfs | 16385503 | 42677 | 515 |
A. mirandus | RU18-7308 | f | Russia: Tuva: 50.205, 95.135 | 468358 | 110900 | 649 |
A. pubescens | d483 | m | Canada: British Columbia: 49.2, -123.2 | 1316697 | 279173 | 503 |
A. rupicola | d491 | m | Poland: Cisna near Lesko | 187507 | 58418 | 312 |
A. rupicola | d492 | m | Poland: Bukowksa Kopa | 418777 | 137114 | 397 |
A. saltator | d512 | m | Germany: Saxony: 51.607, 12.711 | 416618 | 113416 | 591 |
A. sylvestris | d489 | m | U.S.A.: California: 36.3646, -121.5544 | 1289981 | 278727 | 506 |
A. terebratus | RU18-5346 | m | Russia: Novosibirsk Oblast: 53.73, 77.866 | 306744 | 72547 | 668 |
A. zimmermanni | d493 | m | Poland: Grabarka 52.417, 23.005 | 338718 | 113167 | 408 |
A. zimmermanni | RU18-5156 | m | Russia: Novosibirsk Oblast: 53.721, 77.726 | 435654 | 93640 | 627 |
Breda bicruciata | d471 | f | Uruguay: Lavalleja: -34.426, -55.195 | 646088 | 248616 | 549 |
Colonus hesperus | d472 | m | U.S.A.: Arizona: 34.5847, -112.5707 | 1015130 | 250378 | 448 |
Jollas cellulanus | d479 | f | Argentina: Neuquén: -37.0679, -69.7566 | 981935 | 268639 | 497 |
J. cupreus | d473 | m | Ecuador: Orellana: -0.526, -77.418 | 1419103 | 289905 | 469 |
J. cupreus | d474 | m | Ecuador: Orellana: -0.526, -77.418 | 3513351 | 723782 | 607 |
J. leucoproctus | d478 | f | Uruguay: Maldonado: -34.94, -54.95 | 121131 | 61298 | 109 |
Sittisax ranieri | d481 | m | U.S.A.: Oregon: 44.0322, -121.6722 | 1529835 | 322636 | 536 |
Tomis manabita | d476 | m | Ecuador: Manabí: -1.5497, -80.8104 | 2524270 | 710859 | 651 |
T. palpalis | d477 | m | Ecuador: Napo: -0.1996, -77.7023 | 1211674 | 256367 | 582 |
For most samples, DNA was extracted from multiple legs using the Qiagen DNeasy Blood and Tissue Kit (Qiagen, Valencia, CA) following manufacturer’s protocol. Specimens d491 and d492 of Attulus rupicola and d493 of A. zimmermanni were extracted using standard phenol-chloroform methods. UCE library preparation followed methods previously used in arachnids (e.g.,
In the resulting set of loci, most taxa have over 100,000 base pairs of sequence data, but some are less thoroughly sequenced. The less thoroughly sequenced taxa are: J. leucoproctus d478 (13,943 bp), Attulus rupicola d491 (46,660 bp), Attulus rupicola d492 (65,500 bp), and A. zimmermanni d493 (68,285 bp). The last species is represented by an alternative well-sequenced specimen, the others by well-sequenced close relatives. Although we did analyses with the entire set of taxa (“All Taxa”), we were concerned that the weakly sequenced taxa would disrupt resolution. Therefore, we rely primarily on analyses (and bootstrap values) that exclude these and use only the remaining well-sequenced taxa (“Core Taxa”). The Core Taxa dataset also excludes the less thoroughly sequenced of the two specimens of Jollas cupreus (d473, 92,549 bp).
This pipeline therefore resulted in two collections of genes, one of 968 loci for all the taxa (“All Taxa”), the other of 957 loci for the core set of well-sequenced taxa (“Core Taxa”). A filter of occupancy was then applied, eliminating all loci which had sequences for fewer than seven of the 20 well-sequenced taxa of the ingroup (Jollas, Attinella, Tomis, Sittisax, Attulus), resulting in 810 loci in the All Taxa dataset and 803 in the Core Taxa dataset. Preliminary analyses of these loci revealed some whose gene trees strongly suggested two paralogs or chimeras were included: a single very long branch isolating a few taxa (which for all other considerations and subsequent analyses showed no indication of being so distinctive or related to one another), whose sequences differed from the others extensively and consistently. Out of caution we chose to discard a locus if its preliminary gene tree (RAxML 8.2.8,
Maximum likelihood phylogenetic analyses were run using IQ-TREE version 1.6.7.1 (
A separate small phylogenetic analysis was done to explore the distinction in Attulus floricola between hemispheres, using data of other specimens in Genbank and BOLD (boldsystems.org), to blend with our data. Insofar as only COI barcode data are available online, and this gene struggles to reconstruct salticid phylogeny (Hedin and Maddison 2001; Maddison et al. 2008,
Sequence reads are deposited in the Sequence Read Archive (BioProject submission ID PRJNA605426, http://www.ncbi.nlm.nih.gov/bioproject/605426). Alignments and trees are deposited in the Dryad data repository (https://doi.org/10.5061/dryad.cjsxksn2q).
Chromosomes were studied in 17 taxa of Sitticina. The specific identity of the specimen labelled “A. rupicola/floricola” is ambiguous because the voucher specimen has not been located, and the first author is not confident he was able to distinguish the two species in the 1980s. Although its specific identity is not known, it can be confidently placed within the floricola group, and so can play a role in phylogenetic interpretation.
Meiotic chromosomes were observed in testes of adult and subadult males using Feulgen staining, following the methods of
Evidence for scoring chromosome complement of each species is described in Chromosome observations. Most chromosome scoring was done from meiotic nuclei in first metaphase or diakinesis showing chromosomes that are well separated, or, if overlapping, easily interpretable. Although well-spread mitotic nuclei would have added useful data, we judge meiotic chromosomes to be sufficient as they show distinctive features, e.g. when they are oriented by the centromere pulling toward the pole on the metaphase plate. Metacentrics show an obvious bend at the centromere where the second arm hangs loose like a dog’s ear (Fig.
In describing chromosome complements, we use “a” and “m” to indicate one-armed (acrocentric/telocentric) and two-armed (metacentric/submetacentric) chromosomes respectively. Thus, “26a+XaXa0” would mean “26 acrocentric autosomes plus two X’s, both of which are acrocentric”. In all cases, the multiple Xs of a male are interpreted as not being homologous, and therefore it would be more proper to refer to the systems as X1X20, X1X2Y, or X1X2X3Y rather than as XX0, XXY, or XXXY. However, the “1”, “2”, “3” will be left implicit, omitted for ease of reading, to avoid overly complex labels like Xa1Xa2Xa3Ym.
The maximum likelihood tree from the UCE data is shown in Figure
The phylogeny of Sitticina shows two major groups, the Jollas-Tomis clade and Attulus. The Jollas-Tomis clade is distributed entirely in the Americas except for the two species of Sittisax; Attulus is entirely Eurasian except for 8 species in North America. The only previously published comprehensive phylogeny of sitticines, of Prószyński (1983), is substantially similar in placing Sittisax and Attinella outside of the major clade of the floricola, distinguendus and penicillatus species groups. The most notable differences between his arrangement and ours are the placements of Attulus pubescens and A. dzieduszycki. Prószyński’s more recent (2017a) classification into genera, however, is discordant in many respects with our results, as can be seen in the many combinations that we establish or reinstate below in order to achieve monophyly of genera and subgenera. This discord may have arisen partly because Prószyński was not attempting to create a taxonomy that reflected phylogenetic relationships, but rather the distribution of a few diagnostic characters (Prószyński 2017b).
Our UCE phylogeny differs in several respects from
Although Attulus includes some Nearctic members, it is considerably more species-rich in Eurasia, and is most parsimoniously interpreted as having radiated there. The few Nearctic members of this clade are likely recent returns from the Palearctic, insofar as they are Holarctic (Attulus floricola, A. cutleri, A. finschi), close relatives of Eurasian species (A. sylvestris within the A. floricola group, A. striatus close to A. rivalis), or recent introductions (A. ammophilus, A. fasciger, A. pubescens: see Prószyński 1976, 1983 and
The deep branches of the Eurasian Radiation are short, suggesting the group diversified rapidly. Nonetheless, the monophyly of subgenus Sitticus is well supported by a bootstrap percentage of 100 in our primary Core Taxa analysis (Fig.
The relationships among Attulus species are concordant with morphological expectations with one exception: the placement of A. burjaticus with A. zimmermanni, suggesting that the longer embolus of A. zimmermanni and the floricola group are convergent. Otherwise, the floricola group holds together, as do the morphologically similar pairs of A. ammophilus/distinguendus and mirandus/saltator. The placement of A. pubescens nested within the terebratus group indicates that the very short embolus of the former is a derivation from the very long embolus of the latter.
Jollas and Tomis together form a Neotropical radiation and share (typically) an RTA that appears displaced basally, so as to appear to arise closer to the patella, as well as anteriorly placed epigynal openings.
The phylogenetic results lead us to revise the generic division of sitticines. Unless we are to put all Sitticina into a single genus, perhaps palatable for the shallow-diverging Eurasian fauna, but not for the deep Neotropical lineages, then Tomis must be restored for many of the Neotropical species. Given that, Sittisax must be separated from Attulus/Sitticus, rejecting Breitling’s synonymy of this taxon with Sitticus. These choices are relatively easy. The more difficult choices concern the Eurasian Radiation.
Here we give a taxonomic review of the tribe, focussing especially on the species in Canada, and the two new species used in the molecular phylogeny (Jollas cupreus and Tomis manabita). In order to faciliate the use of figures for identification and comparison of species in North America, the sequence of taxa in figures will be different from that in the text, with a series of standardized plates placing images of all of the Canadian species in a block (Figs
Amycoid salticids with fourth legs much longer than third and retromarginal cheliceral tooth lacking. Ancestrally they were ground-dwellers in the Neotropics, later diversifying in Eurasia to include species that live on tree trunks (e.g., A. finschi) and up in vegetation (e.g., Attulus floricola).
Eleven genera are here recognized in the Sitticini, including one (Semiopyla Simon, 1901) whose placement is unclear, and thus remains incertae sedis within the tribe. Two genera are in Eurasia (Attulus and Sittisax), while a disjunct set of eight genera are in South America (the five aillutticines, plus Tomis, Jollas, and Semiopyla). This geographical partitioning matches a phylogenetic division approximately, but not precisely, for the Holarctic Sittisax is phylogenetically a member of the Neotropical radiation. North America has four genera, one arising from the Eurasian radiation (Attulus), and three from the Neotropical radiation (Attinella, Sittisax, and Tomis).
Despite the synonymy of Sitticus with Attulus, the names Sitticini and Sitticina can persist (ICZN Article 40.1).
Aillutticus Galiano, 1987
This group of five Neotropical genera was first recognized by
Aillutticus Galiano, 1987
Amatorculus Ruiz & Brescovit, 2005
Capeta Ruiz & Brescovit, 2005
Gavarilla Ruiz & Brescovit, 2006
Nosferattus Ruiz & Brescovit, 2005
There are no known morphological synapomorphies of this subtribe, but the molecular data show clearly that the five genera listed here form a clade. There are two major subgroups according to the UCE phylogeny: the genus Attulus, a primarily Eurasian radiation, and the Jollas-Tomis clade (Attinella, Jollas, Sittisax, Tomis), a primarily Neotropical radiation. We divide the taxonomy below into those two major groups, and under each discuss the genera, describe the Canadian species and two new Ecuadorian species used in the molecular work.
Attulus Simon, 1868 (type species Attus helveolus Simon, 1871)
Sitticus Simon, 1901 (type species Araneus terebratus Clerck, 1757)
Sitticulus F. Dahl, 1926 (type species Attus saltator O. Pickard-Cambridge, 1868), syn. nov.
Calositticus Lohmander, 1944 (type species Attus caricis Westring, 1861), syn. nov.
Hypositticus Lohmander, 1944 (type species Aranea pubescens Fabricius, 1775), syn. nov.
Sittipub Prószyński, 2016 (type species Aranea pubescens Fabricius, 1775), syn. nov.
Sittiflor Prószyński, 2017 (type species Euophrys floricola C.L. Koch, 1837), syn. nov.
Sittilong Prószyński, 2017 (type species Attus longipes Canestrini, 1873), syn. nov.
We unite the primary Eurasian radiation under the single genus Attulus because of the recency of the radiation, the very short phylogenetic branches separating the subgroups, and the clade’s morphological homogeneity. The total phylogenetic depth of Attulus is far less than that of its sister group (Fig.
Our choice to consider all but two Eurasian species as belonging to Attulus is informed partly by their phylogenetic context among Neotropical salticids. From a Palearctic perspective, the Eurasian radiation of sitticines may seem to represent a lineage of salticids so distinctive and species-rich that they deserve splitting into many genera, especially since the sister group of sitticines among the Old World salticids is the huge clade Salticoida (
The appropriate name for this unified genus is Attulus, as it is far older than Sitticus, and has been used continuously, though for only a few species. Two proposals have been made to ignore priority and instead use Sitticus, the generic name used for most of the species until Prószyński’s (2016, 2017) splitting. Prószyński himself had proposed to the ICZN in 2008 suppression of Attulus in favour of Sitticus, but in 2018 apparently withdrew that proposal (
However, there is value in offering a weaker recognition of three subgroups of Attulus, as subgenera, given that there are names available: Attulus, Sitticus, and Sittilong. Our results support reciprocal monophyly of the subgenera Attulus and Sitticus, and a placement of Sittilong outside of both. Monophyly of subgenus Attulus has variable bootstrap support (72% to 95%, Fig.
The three subgenera have subtle but mostly consistent morphological differences. Attulus s. str. tends to have smaller and more compact bodies, with roundish carapaces (Figs
Attulus includes 49 species in three subgenera:
Subgenus Attulus Simon, 1868, with 41 species:
Attulus (Attulus) albolineatus (Kulczyński, 1895), comb. nov., transferred from Sitticus
Attulus (Attulus) ammophilus (Thorell, 1875)
Attulus (Attulus) ansobicus (Andreeva, 1976)
Attulus (Attulus) atricapillus (Simon, 1882), comb. nov., transferred from Calositticus
Attulus (Attulus) avocator (O. Pickard-Cambridge, 1885)
Attulus (Attulus) barsakelmes (Logunov & Rakov, 1998), comb. nov., transferred from Sitticus
Attulus (Attulus) burjaticus (Danilov & Logunov, 1994)
Attulus (Attulus) caricis (Westring, 1861), comb. nov., transferred from Calositticus
Attulus (Attulus) clavator (Schenkel, 1936)
Attulus (Attulus) cutleri (Prószyński, 1980), comb. nov., transferred from Calositticus
Attulus (Attulus) damini (Chyzer, 1891)
Attulus (Attulus) distinguendus (Simon, 1868) (= type species Attus helveolus Simon, 1871)
Attulus (Attulus) dubatolovi (Logunov & Rakov, 1998)
Attulus (Attulus) dudkoi (Logunov, 1998), comb. nov., transferred from Calositticus
Attulus (Attulus) dzieduszyckii (L. Koch, 1870), comb. nov., transferred from Sittisax
Attulus (Attulus) eskovi (Logunov & Wesolowska, 1995), comb. nov., transferred from Sitticus
Attulus (Attulus) floricola (C. L. Koch, 1837), comb. nov., transferred from Calositticus
Attulus (Attulus) goricus (Ovtsharenko, 1978)
Attulus (Attulus) hirokii Ono & Ogata, 2018
Attulus (Attulus) inexpectus (Logunov & Kronestedt, 1997), comb. nov., transferred from Calositticus
Attulus (Attulus) inopinabilis (Logunov, 1992)
Attulus (Attulus) karakumensis (Logunov, 1992)
Attulus (Attulus) kazakhstanicus (Logunov, 1992)
Attulus (Attulus) mirandus (Logunov, 1993)
Attulus (Attulus) monstrabilis (Logunov, 1992), comb. nov., transferred from Calositticus
Attulus (Attulus) nenilini (Logunov & Wesolowska, 1993)
Attulus (Attulus) nitidus Hu, 2001, comb. nov., transferred from Sitticus
Attulus (Attulus) niveosignatus (Simon, 1880)
Attulus (Attulus) penicillatus (Simon, 1875)
Attulus (Attulus) penicilloides (Wesolowska, 1981)
Attulus (Attulus) pulchellus (Logunov, 1992), comb. nov., transferred from Calositticus
Attulus (Attulus) rivalis (Simon, 1937), comb. nov., and removed from synonymy with A. striatus (Emerton).
Attulus (Attulus) rupicola (C. L. Koch, 1837), comb. nov., transferred from Calositticus
Attulus (Attulus) saltator (O. Pickard-Cambridge, 1868)
Attulus (Attulus) sinensis (Schenkel, 1963)
Attulus (Attulus) striatus (Emerton, 1911), comb. nov., transferred from Calositticus
Attulus (Attulus) sylvestris (Emerton, 1891), comb. nov., transferred from Sitticus, removed from synonymy with A. palustris
Attulus (Attulus) talgarensis (Logunov & Wesolowska, 1993)
Attulus (Attulus) vilis (Kulczyński, 1895)
Attulus (Attulus) zaisanicus (Logunov, 1998)
Attulus (Attulus) zimmermanni (Simon, 1877), comb. nov., transferred from Calositticus
Subgenus Sitticus Simon, 1901, with seven species:
Attulus (Sitticus) fasciger (Simon, 1880), comb. nov., transferred from Sitticus
Attulus (Sitticus) finschi (L. Koch, 1879), comb. nov., transferred from Sitticus
Attulus (Sitticus) godlewskii (Kulczyński, 1895), comb. nov., transferred from Sitticus
Attulus (Sitticus) pubescens (Fabricius, 1775), comb. nov., transferred from Sitticus
Attulus (Sitticus) relictarius (Logunov, 1998), comb. nov., transferred from Sitticus
Attulus (Sitticus) tannuolana (Logunov, 1991), comb. nov., transferred from Sitticus
Attulus (Sitticus) terebratus (Clerck, 1757) (type species of Sitticus), comb. nov., transferred from Sitticus
Subgenus Sittilong Prószyński, 2017, with one species:
Attulus (Sittilong) longipes (Canestrini, 1873) (type species of Sittilong), comb. nov., transferred from Sittilong
Figures
Attulus Simon, 1868 (type species Attus helveolus Simon, 1871).
Sitticulus F. Dahl, 1926 (type species Attus saltator O. Pickard-Cambridge, 1868).
Calositticus Lohmander, 1944 (type species Attus caricis Westring, 1861).
Sittiflor Prószyński, 2017 (type species Euophrys floricola C.L. Koch, 1837).
Body generally more compact than in subgenus Sitticus, with a wider carapace. The spermatheca is a simple tube, folded near the middle. From the point at which the copulatory ducts enter the spermatheca, the spermatheca extends medially to the fertilization duct, but also laterally and then posteriorly (floricola group) or medially (most others) to a separate posterior lobe. Most Attulus (Attulus) have the embolus short, arising near the basal prolateral corner of the bulb, and the tegulum with basal edge more or less straight (not rounded). Several species have a rounder bulb and longer embolus, representing two or three lineages: the floricola group (A. caricis, A. floricola, A. inexpectus, A. rupicola, A. sylvestris), the striatus group (A. striatus, A. rivalis, A. cutleri, A. dudkoi) and the zimmermanni group (A. zimmermanni, A. atricapillus). These also have the folded spermathecae rotated slightly compared to the other Attulus, with the posterior lobe pointing posteriorly, rather than medially. The placement of A. niveosignatus in Attulus (Attulus) is somewhat doubtful, as the position of the tibial apophysis and the anterior medial epigynal openings both resemble those of Sittisax and Attulus subgenus Sittilong. We are reluctant to move it, however, until it is better studied.
Five species of Attulus (Attulus) are known from North America, all of which occur in Canada, as follows.
Attulus subgenus Attulus 15–17 male and female A. distinguendus, Tuva (50.746, 93.142) 18–20 male and female A. mirandus, Tuva (50.205, 95.135) 21–23 A. burjaticus: 21 male, Tuva (50.68, 92.99) 22 male, Tuva (50.205, 95.135) 23 female, Tuva (50.68, 92.99) 24–26 A. zimmermanni: 24, 25 male Novosibirsk Oblast (53.721, 77.726) 26 female Novosibirsk Oblast (53.730, 77.865) 27–30 A. ammophilus: 27 male Tuva (50.6690, 92.9844) 28 male Ontario, Oakville 29 female Ontario, Hamilton 30 male British Columbia (49.08, -119.52). For additional images of A. ammophilus, see Figs
Attulus subgenus Attulus, continued (floricola group) 31, 32 Attulus sylvestris: 31 male, Ontario, Ottawa 32 male, Maryland, Dorchester Co 33–35 A. floricola: 33 male, Ontario, Port Cunnington 34 male, Ontario (46.9300, -79.7268) 35 male, Ontario, Gravenhurst 36–38 A. inexpectus: 36, 37 male, Tuva (50.6690, 92.9844) 38 female, Tuva (51.316, 94.495). For additional images of the floricola group, see Figs
Attus ammophilus Thorell, 1875
Attulus ammophilus is part of the species-rich distinguendus group that is otherwise unrepresented in North America. We have collected it from rocks on the ground in Ontario, British Columbia, and Utah, on litter among marsh plants along the edge of a lake in Siberia, and occasionally from buildings. It was introduced into North America during the 20th century (Prószyński 1976, 1983).
(all in UBC-SEM): Canada: Ontario: Hamilton (69 males, 35 females), Oakville (4 males, 3 females), Toronto (1 male), Windsor (1 male, 2 females); British Columbia: 49.7963, -119.5338 (1 male, 2 females), 49.95, -119.401 (3 males, 2 females); U.S.A.: Utah: 40.7482, -112.1856 (5 males, 7 females), 40.7672, -112.1575 (2 males).
Euophrys floricola C. L. Koch, 1837.
Attus palustris
Peckham & Peckham, 1883 (specimens in
Attus morosus
Banks, 1895 (synonymized by Prószyński 1980; confirmed here by examination of holotype female in
A widespread Holarctic species often found in retreats in dry flower heads in wetter areas such as marshes, A. floricola is distinctive for the sharp white lines around the eyes of males, forming an apparent mask (Fig.
We treat the North American populations as full floricola, not a distinct subspecies. While Nearctic populations were long recognized as a separate species palustris, Prószyński (1980) suggested they are conspecific with the Eurasian populations. He maintained them as a distinct subspecies, but he expressed doubt as to whether even that distinction was warranted. We concur with his skepticism. If any consistent differences exist between the continents, they are no more visible than any differences that might exist between the Eurasian and North American populations of other species for which we don’t recognize subspecies such as Sittisax ranieri, Attulus cutleri, Dendryphantes nigromaculatus (Keyserling, 1885), Pellenes ignifrons (Grube, 1861), and Pellenes lapponicus (Sundevall, 1833).
The results of our COI analysis of Palearctic and Nearctic floricola group (Fig.
Within North America, the characterization of A. floricola has been muddied by confusion with a second species, A. sylvestris. Attulus sylvestris, long synonymized with palustris, is a distinctively different species. Attulus floricola is larger-bodied, has a much more contrasting colour pattern, and longer legs. Attulus floricola has a different angle of the spermaphore loop (subtle but consistent; Fig.
A more serious confusion apparently occurred with the labelling of type specimens of Attus palustris. The description by
At stake is not the name used for the common white-striped species (which would be floricola regardless), but the name for the uncommon dusty brown species, which would be palustris were we to accept these specimens as its types. However, as argued above, they are not the types. We therefore treat palustris as a synonym of floricola, and sylvestris as the name for the dusty brown species. To settle the mislabelling properly, a male specimen of the white-striped species from Wisconsin (the type locality) should be designated as the neotype or lectotype of palustris. We have not yet done so as we await reexamination of the full Peckham collection in case specimens can be located that might be identifiable as from the true type series.
Canada: British Columbia: Richmond (2 females), 49.66, -114.73 (1 female), 49.45, -115.08 (3 males, 6 females); Alberta: 52.46, -113.94 (1 male); Ontario: Richmond (2 males, 1 female), Gravenhurst (3 males), Port Cunnington (1 female); Dwight (2 males, 5 females), Batchawana Bay (1 female), Woodstock (3 females), 46.9300, -79.7268 (2 males, 1 female), 42.53, -80.12 (1 female), 43.2626, -80.5636 (1 male), 49.0852, -81.3237 (1 female); Quebec: Touraine (1 male); Nova ScotiA: 44.4318, -64.6075 (1 male); U.S.A.: Washington: 46.43, -123.86 (2 males); Colorado: Jackson Lake State Rec. Area (1 male); Nebraska: 41.88, -103.09 (1 female).
Attus sylvestris
Emerton, 1891 (Holotype male in
Sitticus magnus Chamberlin & Ivie, 1944, syn. nov.
Sitticus rupicola – Prószyński, 1980, figs 58, 59 (misidentification), specimen from Texas.
A widespread but little-known Nearctic species, A. sylvestris can be found on partially shaded ground where the males stand out for their tiny bouncing bright white spots (the white tuft of setae on the palp’s tibia). We have found them on rocks and leaf litter along a forest edge in Ontario, on the ground at the edge of a creek in a forest in California, and on forest leaf litter in Maryland. See discussion under A. floricola about why we judge A. sylvestris to be the proper name of this species, at issue because of confusion over the type specimens of Attus palustris.
Both males and females have shorter legs and less contrasting markings than in A. floricola, but the distinction of markings is most notable in the male, which lacks the high-contrast white stripes on dark brown seen in A. floricola. The white setae on the male’s palp are concentrated on just the tibia and end of the femur. The bulb of the palp is rotated slightly more than in A. floricola, and thus the spermophore’s path shows an upturn (i.e., the loop is angled to point distally instead of basally as in floricola), and the female’s copulatory ducts arrive further to the posterior before looping back anteriorly to enter the spermathecae. In these regards the genitalia resemble those of the Eurasian A. rupicola, A. caricis, and A. inexpectus (Logunov and Kronsestedt 1997). Attulus sylvestris is most similar to A. caricis in appearance (low-contrast brown markings), in having a small loop of the copulatory duct, and small body size, but differs in brighter markings on the palp, a more anteriorly-placed junction where the ducts enter the spermatheca, a larger epigynal RTA coupling pocket, and a more distinctly swollen bulb of the spermatheca. (They are also distinct on the COI tree, Fig.
(all in UBC-SEM except as indicated): Canada: Ontario: Ottawa, Britannia Bay, 45.374, -75.796 (26 males, 3 females), Long Point, 42.53, -80.12 (2 females); U.S.A.: Maryland: Dorchester Co. (1 male 1 female,
Sitticus striatus Emerton, 1911
Attulus striatus is a small-bodied Northern species with distinctively striped males, from sphagnum bogs. Although we were unable to obtain molecular data for it or the similar A. rivalis and A. cutleri, these three species can be placed into subgenus Attulus with some confidence, based on their boxy carapaces (resembling the other Attulus (Attulus) rather than Attulus (Sitticus)), and the genitalic similarities with subgenus Attulus, including the two small posterior openings of the epigyne on either side of a narrow triangular RTA coupling pocket. Prószyński (1980) considered them close to the floricola group in particular.
We reinstate S. rivalis Simon, 1937 as a distinct species (contra Prószyński 2017a), accepting
(all UBC-SEM): Canada: Alberta: S. Islay (3 female), Beaverhill Lake (1 female); Ontario: 48.3260, -76.8365 (1 female); 3 km S. Richmond (6 males, 2 females); New Brunswick: Chipman (1 male, 1 female); U.S.A.: New Hampshire: Ponemah Bog (1 female).
Sitticus cutleri Prószyński, 1980
Sitticus gertschi Prószyński, 1980
A Sibero-American boreal species that is little collected, resembling closely A. striatus but differing in having less striped legs, a less rotated bulb of the male palp, more medially placed epigynal openings. Collected on “leaf litter under small Salix just above stream” (D. Maddison, June 1981, Inuvik).
Canada: Northwest Territories: Wrigley (1 female,
Figures
Sitticus Simon, 1901 (type species Araneus terebratus Clerck 1757)
Hypositticus Lohmander, 1944 (type species Aranea pubescens Fabricius, 1775)
Sittipub Prószyński, 2016 (type species Aranea pubescens Fabricius, 1775)
The species placed here, despite having palpi with very different embolus lengths, share a similarly narrow and high body with relatively long legs (Figs
Attulus subgenus Sitticus 39, 40 A. fasciger, male, Ontario (43.3508, -79.7593) 41, 42 A. finschi: 41 male, Saskatchewan (55.31, -105.11) 42 male body, Ontario, 4 miles S of Wawa 44, 45 A. terebratus: 44 male, Novosibirsk Oblast (53.730, 77.865) 45 female, Novosibirsk 46, 47 A. relictarius male, Stavropol Krai, (43.88, 42.70). For additional images of Attulus (Sitticus), see Figs
Maximum likelihood phylogeny from 757 concatenated UCE loci (average 113231 base pairs/taxon) analyzed primarily for the 23 Core Taxa in black (IQ-TREE, partitioned by locus). Topology is identical in unpartitioned analyses, with nearly identical branch lengths. Bootstrap percentage values from 1000 replicates shown for each clade. Where two numbers are shown, the first is the bootstrap percentage for the partitioned analysis, the second for the unpartitioned analysis. Where one number is shown, both analyses yielded the same percentage. An analysis of the All Taxa dataset, including the weakly-sequenced taxa in grey, yielded the same topology.
Sitticines of Canada: Attulus subgenus Attulus (for A. ammophilus, see Figs
Attus finschii L. Koch, 1879
Euophrys cruciatus Emerton, 1891
The natty contrasting black-and-white markings distinguish Attulus finschi from the closely related A. fasciger. Attulus finschi is the only Sitticus that has likely been in the Americas for thousands of years; it also lives in Siberia. It is found in boreal habitats on tree trunks.
(all UBC-SEM): Canada: Saskatchewan: 55.31, -105.11 (1 male, 1 female), 55.27, -105.19 (1 female); Ontario: Wawa (1 male), Nipigon (1 female), 48.9143, -80.9446 (2 females); New Brunswick: Doaktown (1 male).
Attus fasciger Simon, 1880
This species, introduced to North America apparently in the middle of the 20th century (
(all in UBC-SEM): Canada: Ontario: Burlington (3 males, 6 females), 43.35074, -79.75928 (25 males, 14 females); U.S.A.: Missouri: Dogtown (3 males, 4 females); Massachusetts: Cambridge (1 female).
Aranea pubescens Fabricius, 1775
Although closely related to A. fasciger and A. terebratus, which have among the longest emboli and copulatory ducts in sitticines, Attulus pubescens has among the shortest known in sitticines. The very large RTA is distinctive. Introduced to North America in the 20th century (
(All in UBC-SEM): Canada: British Columbia: Vancouver (1 male 1 female); U.S.A.: Massachusetts: Cambridge (3 males, 3 females), Boston (2 males), Milton (2 males), Arlington (1 female).
Sitticines of Canada: Attulus, continued 69–73 Attulus (Attulus) ammophilus: 69palp (Ontario, Oakville) 70, 71 ventral view of epigyne, dorsal view of cleared vulva (Ontario, Hamilton) 72 male (British Columbia, 49.08, -119.52) 73 female (British Columbia, 49.08, -119.52) 74–78 A. (Sitticus) fasciger (Ontario, 43.3508, -79.7593): 74 palp 75, 76 ventral view of epigyne, dorsal view of cleared vulva 77 male 78 female 79–83 A. (S.) finschi: 79 palp (Ontario, Wawa) 80, 81 ventral view of epigyne, dorsal view of cleared vulva (Saskatchewan, 55.31, -105.11) 82 male (Saskatchewan, 55.31, -105.11) 83 female (Saskatchewan, 55.27, -105.19) 84–88 A. (S.) pubescens: 84 palp (Massachusetts, Milton) 85, 86 ventral view of epigyne, dorsal view of cleared vulva (Massachusetts, Arlington) 87 male (Massachusetts, Cambridge) 88 female (Massachusetts, Cambridge). For other images of Attulus (Sitticus), see Figs
Sittilong Prószyński, 2017 (type species Attus longipes Canestrini, 1873)
The single species Attulus (Sittilong) longipes of the European Alps is peculiar for its flat body and long first legs in the male, as well as its genitalia. Like Sittisax and other members of the Jollas-Tomis clade, the RTA is offset basally, and the epigynal openings are anterior and medial. The little-studied Attulus niveosignatus has somewhat similar genitalia and may also belong in Sittilong.
We have chosen not to subdivide the Neotropical Sitticina more finely than into two genera, Tomis and Jollas, primarily because the fauna is poorly enough known that it is as yet unclear what coarseness of division would be most useful. We might have synonymized their respective Nearctic offshoots (Sittisax into Tomis, and Attinella into Jollas), but by retaining them as distinct, we facilitate the eventual splitting of both Tomis and Jollas as their species become better known. We choose splitting in the Jollas-Tomis clade, in contrast to lumping with Attulus, because the phylogenetic divergences are so much deeper in the former compared to the latter.
The Jollas-Tomis clade includes four genera with 31 species:
Attinella Banks, 1905, with three species:
Attinella concolor (Banks, 1895), comb. nov., transferred from Sitticus
Attinella dorsata (Banks, 1895), combination restored, transferred from Sitticus (type species)
Attinella juniperi (Gertsch & Riechert, 1976), comb. nov., transferred from Sittiab
Jollas Simon, 1901, with 12 species:
Jollas amazonicus Galiano, 1991
Jollas cellulanus (Galiano, 1989), comb. nov., transferred from Sitticus
Jollas cupreus W. Maddison, sp. nov.
Jollas flabellatus (Galiano, 1989), comb. nov., transferred from Sitticus
Jollas geniculatus Simon, 1901 (type species)
Jollas hawkeswoodi Makhan, 2007
Jollas leucoproctus (Mello-Leitão, 1944), comb. nov., transferred from Sitticus
Jollas manantiales Galiano, 1991
Jollas paranacito Galiano, 1991
Jollas pompatus (Peckham & Peckham, 1894)
Jollas puntalara Galiano, 1991
Jollas richardwellsi Makhan, 2009
Sittisax Prószyński, 2017, with two species:
Sittisax ranieri (Peckham & Peckham, 1909)
Sittisax saxicola (C. L. Koch, 1846) (type species)
Tomis F.O. Pickard-Cambridge, 1901, with 14 species
Tomis canus Galiano, 1977, combination restored, transferred from Sitticus
Tomis kratochvili (Caporiacco, 1947), comb. nov., transferred from Pseudattulus
Tomis manabita W. Maddison, sp. nov.
Tomis mazorcanus (Chamberlin, 1920), comb. nov., transferred from Sitticus
Tomis mona (Bryant, 1947), comb. nov., transferred from Sidusa
Tomis palpalis F. O. Pickard-Cambridge, 1901, combination restored, transferred from Sitticus (type species)
Tomis pavidus (Bryant, 1942), comb. nov., transferred from Sidusa
Tomis phaleratus (Galiano & Baert, 1990), comb. nov., transferred from Sitticus
Tomis pintanus (Edwards & Baert, 2018, comb. nov., transferred from Sitticus
Tomis tenebricus (Galiano & Baert, 1990), comb. nov., transferred from Sitticus
Tomis trisetosus (Edwards & Baert, 2018), comb. nov., transferred from Sitticus
Tomis uber (Galiano & Baert, 1990), comb. nov., transferred from Sitticus
Tomis vanvolsemorum (Baert, 2011), comb. nov., transferred from Sitticus
Tomis welchi (Gertsch & Mulaik, 1936), comb. nov., transferred from Sitticus
Attinella Banks, 1905 (type species Attus dorsatus Banks, 1895)
Sittiab Prószyński, 2017 (type species Sitticus absolutus Gertsch & Mulaik, 1936), syn. nov.
Small species from southern North America, related to the Jollas of South America. Except for the thin longitudinal stripes of A. dorsata, their bodies are more or less unmarked. Like many other members of the Jollas-Tomis clade, the RTA is long and thin, paralleling the axis of the palp, the tibia is robust, and the embolus is fairly short. The first leg’s trochanter is unusually long in at least some males (note angles in Fig.
Attus concolor
Banks, 1895 (holotype examined; see
Sittacus cursor Barrows, 1919, synonymy restored
Sitticus floridanus Gertsch & Mulaik, 1936
A small unmarked leaf litter species, known best from the southeastern United States, but recently reported from Canada in the BOLD barcode database (Ratnasingam and Hebert 2007, 2013), from the extreme southern point in Ontario (Point Pelee National Park, specimens PPELE142-11, PPELE183-11, CNPPE2332-12, PPELE666-11, PPELE644-11).
Prószyński (2017a) rejected
U.S.A.: Florida: Gainesville (1 male, 1 female, UBC-SEM).
Sitticines of Canada: the Jollas-Tomis clade, represented by the genera Attinella and Sittisax 89–93 Attinella concolor: 89 palp (Florida, Gainesville) 90, 91 ventral view of epigyne, dorsal view of cleared vulva (Florida, Gainesville) 92 male (Texas, 30.10, -97.25) 93 female (Texas, 30.10, -97.25) 94–98 Attinella dorsata: 94 palp (California, San Diego County) 95, 96 ventral view of epigyne, dorsal view of cleared vulva (British Columbia, Nanaimo) 97 male (California, Siskiyou County) 98 female (British Columbia, 48.870, -123.379) 99–103 Sittisax ranieri: 99 palp (Northwest Territories, Tuktoyaktuk) 100, 101 ventral view of epigyne, dorsal view of cleared vulva (Nunavut, Baffin Island) 102 male (Saskatchewan, 55.27, -105.19) 103 female (Ontario, Old Woman Bay).
Relationships among Attulus floricola mitochondrial COI sequences in the context of the floricola group. Specimens in bold had their relationships constrained by the UCE phylogeny of Fig.
Attus dorsatus
Banks, 1895 (holotype female in
Sitticus absolutus Gertsch & Mulaik, 1936, synonymy restored
Sitticus callidus Gertsch & Mulaik, 1936, synonymy restored
While females of this small Southwestern desert-dwelling species are indistinctly unmarked, males tend to be reddish with a narrow central longitudinal stripe (Figs
Canada: British Columbia: Summerland (1 male,
Figures
Jollas Simon, 1901 (type species Jollas geniculatus Simon, 1901)
Oningis Simon, 1901 (type species Neon pompatus Peckham & Peckham, 1893)
A Neotropical group, consisting of two species groups, the small glabrous or shiny geniculatus group (
Jollas cupreus, sp. nov. (except 112, J. puntalara) 108, 109 Left palp of holotype 108 ventral view 109 retrolateral view 110 ventral view of epigyne of paratype 111 dorsal view of same, cleared 112 palp of holotype of J. puntalara Galiano 113–115 holotype male 116 male from Yasuní, Ecuador (-0.675, -76.397) 117 holotype male in alcohol 118, 119 paratype female.
Male holotype and 2 male, 3 female paratypes from Ecuador: Orellana: Río Bigal Reserve, main camp area. 0.5251, -77.4177. 950 m elev. 1–5 November 2010. W & D Maddison, M Vega, M Reyes. WPM#10-041c. The holotype (specimen ECU2010-2060) pertains to the Museum of Zoology, Pontificia Universidad Católica, Quito, Ecuador (
Refers to the copper colour of males.
A species common in eastern Ecuador on disturbed open grassy ground. It was used in the molecular phylogenetic study of
Differs from the very similar Jollas puntalara Galiano, 1991 in the thinner and straighter RTA and the angle at which the embolus arises. The RTA is more or less straight until a curl at the tip, but it narrows dramatically for its terminal three quarters (Fig.
Male (holotype). Carapace length 1.37; abdomen length 1.16. Carapace orange with a black ocular area, mostly glabrous, with only a few scattered setae. Clypeus orange-brown. Chelicerae vertical, orange. Palp orange-brown except for dark brown cymbium, with dark setae except brilliant white patch of setae dorsally on patella. Legs long, especially the first and fourth. Legs honey coloured to orange-brown except for a strong black line on prolateral-ventral face of first patella, tibia and metatarsus. Embolus arises at ca. 5 o’clock and curls half-way around bulb. Tibia somewhat bulbous, broad, with bases of setae on retrolateral side forming row of tubercles. RTA begins broad but then narrows abruptly at ca. one quarter its length, from which point it proceeds straight until just before the tip, where it curls. Abdomen orange-brown, with black scalloped patch covering dorsum, covered with metallic scales. A patch of bright white setae sits above the anal tubercle.
Female (paratype). Carapace length 1.36; abdomen length 1.89. Much darker than the male in body and appendages (Figs
22 males and 6 females from: Ecuador: Napo: Tarapoa. 23 June – 1 July 1988 W. Maddison WPM#88-002 (1 male); Ecuador: Napo: bridge over Rio Cuyabeno on road to Tipishca. 25–30 June 1988 W. Maddison WPM#88-004 (1 male 1 female); Ecuador: Napo: bridge over Rio Cuyabeno on road to Tipishca. 29–30 July 1988 W. Maddison WPM#88-018 (4 males 2 females); Ecuador: Napo: Reserva Faunistica de Cuyabeno, Laguna Grande, Sendero La Hormiga. 2–5 August 1988 W. Maddison WPM#88-023 (2 males); Ecuador: Napo: Reserva Faunistica de Cuyabeno, Laguna Grande, PUCE field station. 1–7 August 1988 W. Maddison WPM#88-025 (1 male); Ecuador: Napo: bridge over Rio Cuyabeno on road from Lago Agrio to Tipishca. 8–9 August 1988 W. Maddison WPM#88-027 (1 male); Ecuador: Sucumbios: Reserva Faunistica Cuyabeno, Laguna Grande, PUCE field station. 0.002, -76.172. 21–29 July 1989 W. Maddison WPM#89-032 (1 male); Ecuador: Sucumbios: bridge over Rio Cuyabeno on road between Tarapoa and Tipishca, 0.025, -77.308. 29 July 1989 W. Maddison WPM#89-036 (1 male); Ecuador: Sucumbios: Reserva Faunistica Cuyabeno, Nuevo Mundo cabins along Rio Cuyabeno at jcn with Lago Agrio-Tipishca HWY 19–29 April 1994 W. Maddison WPM#94-021 (3 males); Ecuador: Sucumbios: Reserva Faunistica Cuyabeno, Nuevo Mundo cabins, jcn Rio Cuyabeno & Lago Agrio-Tipishca HWY tree trunks 19–29 April 1994 W. Maddison WPM#94-023 (1 male); Ecuador: Morona Santiago: km 3 from Limón towards Gualaceo. 2.9663, -78.4209; 1250 m el. 12 July 2004 Maddison, Agnarsson, Iturralde, Salazar. WPM#04-030 (1 male 2 females); Ecuador: Morona Santiago: km 4 from Limón towards Gualaceo. 2.9808, -78.4414; 1380 m el. 12 July 2004 Maddison, Agnarsson, Iturralde, Salazar. WPM#04-031 (2 males); Ecuador: Orellana: Yasuní Res.Stn.area, Station area 0.675, -76.397 210–280 m elev. 26 July – 13 Aug 2011 Maddison/Piascik/Vega WPM#11-015 (2 males); Ecuador: Orellana: Yasuní Res.Stn.area, Station area 0.674, -76.397 210–280 m elev. Clearings, forest edge 8–9.Aug.2011 Maddison/ Piascik/Vega. WPM#11-104 (1 male); Ecuador: Orellana: Río Bigal Reserve, boundary along road. 0.541, -77.424. 970 m elev. 5 November 2010. M Vega, D & W Maddison, M Reyes. WPM#10-048 (1 female). (Note: the province Sucumbios was established after 1988; the 1988 localities listed as Napo Province would now all be in Sucumbios.).
Sittisax Prószyński, 2017 (type species Euophrys saxicola C.L. Koch, 1846)
Attus lineolatus Grube, 1861 (junior homonym)
Sittacus ranieri Peckham & Peckham, 1909
The Holarctic Sittisax ranieri is a widespread boreal species, which in North America follows the high elevations of the Western Cordillera to the south, living on rocks and litter. It is dark in colour, large-bodied, and with distinctive genitalia.
Canada: Northwest Territories: Tuktoyaktuk (1 male); Nunavut: Baffin Island (1 female); British Columbia: Downton Creek (2 males 2 females), 49.026, -114.061 (1 male), 59.8, -127.5 (1 male), Pink Mountain (1 male); Yukon: km 72 Dempster Highway (2 males, 2 females); km 75.6 Dempster Highway (1 female); Saskatchewan: 55.27, -105.19 (2 males), Ontario: Old Woman Bay (1 female); New Brunswick: 65.336, -69.4 (6 males, 3 females); U.S.A.: Washington: Spray Park (1 males, 2 females); Oregon: 45.261, -117.178 (1 female); Colorado: 39.803, -105.782 (1 male).
Figures
Tomis F.O. Pickard-Cambridge, 1901 (type species Tomis palpalis F.O. Pickard-Cambridge, 1901)
Pseudattulus Caporiacco, 1947 (type species Pseudattulus kratochvili Caporiacco, 1947), syn. nov.
A Neotropical group whose male spermophore (with some possible exceptions) has a “shortcut loop”. That is, the large loop of the spermophore that normally occupies much of the visible face of the tegulum, and which points basally in many sitticines (e.g., Fig.
The phylogeny strongly places T. palpalis, T. manabita, and Sittisax ranieri together. Although the phylogeny gives us the freedom to synonymize Sittisax into Tomis, this deep clade will eventually deserve at least two genera, and so we tentatively retain the boundary between the Neotropical Tomis and the Holarctic Sittisax, based on the apparent difference in spermophore loops. The other species are included in Tomis because of their apparent relationship with T. palpalis and T. manabita. The palpalis group (T. palpalis, T. canus, T. mazorcanus, T. phaleratus, T. vanvolsemorum, and T. uber) is delimited by a flattened cymbium (Galiano, 1991a) and well-separated epigynal openings. The remaining species all are known from coastal areas of the Caribbean or South America, and at least some live on beaches. They might not form a monophyletic group, as some show a long thin RTA, others not. T. pavidus and T. mona appear especially close to T. manabita by similarities in the palps. The others can be tentatively included in Tomis because they share with T. palpalis and T. manabita the shortcut spermophore loop.
The placement of Sitticus welchi Gertsch & Mulaik, 1936 in Tomis is tentative. The holotype female (
We synonymize Pseudattulus (see
Male holotype, 10 male and 8 female paratypes from Ecuador: Manabí: Puerto Rico, Cabañas Alandaluz 5 May 1994 W. Maddison WPM#94-031. The holotype (specimen UBC-SEM AR00217) pertains to the Museum of Zoology, Pontificia Universidad Católica, Quito, Ecuador (
Based on the type locality; the form is the adjective in Spanish (masculine or feminine).
A species on the beaches of coastal Ecuador, resembling Attulus in habitus. It was used in the molecular phylogenetic study of
Palp closely resembles that of Tomis pavidus, from which it differs in the smaller tibia and longer RTA.
Male (holotype). Carapace length 1.58; abdomen length 1.51. Carapace (Figs
Female (paratype # UBC-SEM AR00218). Carapace length 1.76; abdomen length 2.22. Carapace (Figs
15 males and 7 females from Ecuador: Manabí: Machalilla National Park, Salaite, between HWY and coast 6 May 1994 W. Maddison WPM#94-032 (4 males, 2 females); Ecuador: Manabí: Machalilla National Park, Salaite, 1 km inland along trail from HWY. 6 May 1994 W. Maddison WPM#94-033 (3 males); Ecuador: Manabí: Machalilla National Park, trail between Agua Blanca & San Sebastien 50–400 m dry forest 7 May 1994 W. Maddison WPM#94-034 (1 male); Ecuador: Manabí: Crucita. 30 August 1988 W. Maddison WPM#88-040 (2 males 4 females); Ecuador: Del Oro: Jambelí 13 August 1989 W. Maddison WPM#89-040 (3 males); Ecuador: Manabí: Puerto Lopez. 1.5497, -80.8104; 5 m el. 1–5 August 2004 W. Maddison. WPM#04-067 (2 males 1 female).
The following species are not sitticines, indicated by the presence of retromarginal cheliceral teeth (lacking in the Sitticini, a synapomorphy) or characteristic euophryine genitalia.
The following three are members of the Euophryini. They are left within sitticine genera because it is unclear to which genus they should be transferred.
Jollas armatus (Bryant, 1943)
Jollas crassus (Bryant, 1943)
Jollas minutus (Petrunkevitch, 1930)
The following two species described in Sitticus are also euophryines (see Prószyński 2017a). They are tentatively placed in a likely genus, Chinophrys:
Chinophrys taiwanensis (Peng & Li, 2002), comb. nov.
Chinophrys wuae Peng, (Tso & Li, 2002), comb. nov.
The following species can be moved out of Sitticus to their appropriate genera. The type specimens of both, in the
Heliophanus designatus (Peckham & Peckham, 1903), comb. nov. – bears the stridulatory apparatus characteristic of chrysillines (
Mexigonus peninsulanus (Banks, 1898), comb. nov. – appears as a typical Mexigonus with euophryine genitalia.
Table
Chromosome complements observed for males of 17–18 species of sitticines. The autosomal counts represent diploid complement, and thus 26a means 13 pairs of acrocentric autosomes. In the chromosome counts, a = acrocentric (one-armed), m = metacentric (two-armed). Exx. is the number of specimens; nuc. is the number of nuclei showing the full chromosome complement; +nuc sex is the number of additional nuclei showing the sex chromosomes (though not clearly the autosomes). Uncertainties about scoring, in particular about Attinella dorsata, Attulus burjaticus and the specimen labelled “Attulus rupicola/floricola” are explained under Chromosome observations.
Species | Autosomes | ♂ Sex chrom. | Y present | Locality | exx | nuc | +nuc sex |
Jollas-Tomis clade | |||||||
Attinella concolor | 14m | Xm0 | no | U.S.A.: Gainesville, 29.63, -82.37 | 1 | 6 | 2 |
A. dorsata | 26a? | XaXa0? | ? | U.S.A.: Dillon Cr., 41.57, -123.54 | 3 | 11 | |
Jollas cupreus | 26a | XaXa0 | no | Ecuador: Tarapoa, -0.12, -76.34 | 1 | 2 | 1 |
Sittisax ranieri | 24a | XmXaYm | yes | Canada: Leguil Creek, 59.8, -127.5 | 2 | 10 | 1 |
Canada: Inuvik, 68.31, -133.49 | 1 | 1 | |||||
Canada: Wawa, 47.79, -84.90 | 2 | 8 | |||||
U.S.A.: Mt. Monadnock, 42.87, -72.11 | 1 | 3 | |||||
Sittisax saxicola | 24a | XaXaXaYm or XmYaYaYa | yes | Switzerland: Flims, 46.9, 9.2 | 3 | 14 | 10 |
Tomis manabita | 26a | XaXa0 | no | Ecuador: Crucita, -0.9, -80.5 | 1 | 3 | 2 |
Attulus | |||||||
Attulus (Attulus) ammophilus | 26a | XaXa0 | no | Canada: Toronto, 43.65, -79.32 | 3 | 9 | 1 |
Russia: Uvs Nuur, 50.6690, 92.9844 | 2 | 11 | 6 | ||||
A. (A.) burjaticus | ? (28a?) | XaXa0 | no | Russia: Uvs Nuur, 50.677, 92.99 | 1 | 1 | 7 |
A. (A.) caricis | 26a | XaXa0 | no | 38.6, 34.8 ( |
– | ||
A. (A.) cutleri | 26a | XaXaYa | yes | Canada: Inuvik, 68.35, -133.70 | 1 | 3 | 4 |
A. (A.) floricola | 28a | XaXa0 | no | Canada: Barrie, 44.43, -79.65 | 1 | 7 | 7 |
U.S.A.: Naselle, 46.43, -123.86 | 1 | 2 | |||||
A. (A.) rupicola/floricola | 24a? | XaXaXaYm? | yes | Switzerland: Flims, 46.9, 9.2 | 1 | 3 | 5 |
A. (A.) inexpectus | 26a | XaXa0 | no | Russia: Uvs Nuur, 50.6690, 92.9844 | 2 | 13 | 5 |
A. (A.) striatus | 24a | XaXaXaYm | yes | U.S.A.: Ponemah, 42.82, -71.58 | 1 | 5 | 6 |
Attulus (Sitticus) fasciger | 26a | XaXa0 | Canada: Burlington, 43.351, -79.759 | 3 | 16 | ||
A. (S.) finschi | 28a | XaXa0 | no | Canada: Nipigon, 49.01, -88.16 | 1 | 4 | |
Canada: Sault Ste. Marie, 46.94, -84.55 | 1 | 1 | |||||
Canada: Chinook L., 49.67, -114.60 | 1 | 8 | 3 | ||||
A. (S.) pubescens | 26a | XaXmYa | yes | U.S.A.: Cambridge, 42.38, -71.12 | 4 | 10 | 9 |
A. (S.) terebratus | 26a | XaXa0 | no | Russia: Karasuk, 53.730, 77.866 | 1 | 9 | 14 |
26a | XaXa0 |
|
– |
Attinella concolor: 14m+Xm0 (Figs
Attinella dorsata: 26a+XaXa0 (uncertain). Scored as 26a+XaXa0 in notes from the 1980s, the slides are too faded and degraded for precise re-count, but re-examination shows they have at least 13 acrocentric bivalents, and what looks like XX0. Although we might have abandoned the score entirely, we include it here to show that it is at least similar to the typical salticid complement, and not at all what is seen in the close relative Attinella concolor.
Jollas cupreus: 26a+XaXa0. One first division nucleus appears clearly as 13 autosomal bivalents plus two acrocentric Xs, while two more show the typical Xs side by side.
Sittisax ranieri: 24a+XmXaYm (Figs
Sittisax saxicola: 24a+XaXaXaYm or 24a+XmYaYaYa (good quality, though ambiguous in interpretation; Figs
Chromosomes of first meiotic division in males of the Jollas-Tomis clade 129, 130 Attinella concolor, with only seven pairs of autosomes, but each two-armed, 14m+Xm0, Florida (29.63N, 82.37W) 131 Tomis manabita, showing the two Xs off to one pole, and 13 acrocentric bivalents on the metaphase plate, Ecuador (0.9S, 80.5W) 132–136 Sittisax ranieri, whose distinctive XmXaYm appears as a rabbit head with a droopy ear. White triangles show points where two bivalents are apparently linked together 134–136 details of XXY of S. ranieri 137–139 Sittisax saxicola, with sex chromsomes, interpreted tentatively as XaXaXaYm, appearing as a rabbit head with three ears, Switzlerland (46.9N, 9.2E).
All four sex chromosomes of S. saxicola come together in a quintuple junction. This and the quadruple junction of S. ranieri are unusual, possibly formed because mutual translocations or repeated sequences generate a knit pattern of pairing.
Tomis manabita: 26a+XaXa0 (Fig.
Attulus (Attulus) ammophilus: 26a+XaXa0 (Figs
Attulus (Attulus) burjaticus: ?+XaXa0 (autosome count uncertain; Fig.
Chromosomes of first meiotic division of Attulus subgenus Attulus 140, 141 Attulus ammophilus, Tuva (50.6690N, 92.9844E): 140 four nuclei, three showing the two X chromosomes toward one pole 141 two nuclei showing two Xs and thirteen pairs of acrocentric autosomes 142 Attulus burjaticus, showing the two X chromosomes toward one pole, Tuva (50.677N, 92.99E). The three large spots to the lower right are spermatids.
Attulus (Attulus) cutleri: 26a+XaXaYa (Figs
Attulus (Attulus) floricola: 28a+XaXa0, with one autosome much smaller (Figs
Attulus (Attulus) rupicola/floricola (Switzerland): 24a+XaXaXaYm (uncertain in details, though the presence of at least one Y is secure; Figs
Attulus (Attulus) inexpectus: 26a+XaXa0 (Figs
Chromosomes of meiosis of Attulus subgenus Attulus, continued 143–146 Attulus floricola, with an extra small bivalent (s) to make 28a+XaXa0, Ontario (44.43, -79.65): 143, 144 first metaphase 145 second division, showing one nucleus with 14 acrocentrics, the other with 14 acrocentrics and the two condensed Xs 147–149 Attulus inexpectus, showing 13 acrocentric bivalents and the sex chromosomes (26a+XaXa0), Tuva (50.6690, 92.9844) 150, 151 Attulus sp. (ambiguously identified, either A. rupicola or floricola), tentatively intepreted as having 24a+XaXaXaYm, Switzerland (46.9, 9.2): 151 same, sex chromosomes from another nucleus 152 Attulus cutleri, with 26a+XaXaYa, Canada (68.35, -133.70) 153 same, sex chromosomes from another nucleus
Attulus (Attulus) striatus: 24a+XaXaXaYm. The slides are too faded to score now even under phase contrast, and so for this we rely entirely on notes from 1985. Those notes give good evidence to consider the interpretation secure. The slides were then clear enough to score chiasma localization in the acrocentric autosomes (in 14 nuclei with at least ten autosomes scorable, the numbers of proximal vs. interstitial vs. terminal chiasmata were 76:12:50 respectively). Five of these nuclei showed a clear count of 14 acrocentric autosomes. The sex chomosomes were clear in several nuclei, consisting of a “V” shaped trivalent with a metacentric at the point of the “V”, to each arm of which was paired an acrocentric. One of those acrocentrics was decondensed (heteropycnotic) in its centromeric half, and lying alongside it achiasmately was a decondensed acrocentric, thus in total making a figure of four. The achiasmate pairing and heteropycnosis suggest those acrocentrics have ancestral X material, as in the XXXY Habronattus (
Attulus (Sitticus) fasciger: 26a+XaXa0 (Fig.
Attulus (Sitticus) finschi: 28a+XaXa0, with one autosome much smaller. This score relies primarily on old notes, which indicate 28 acrocentric autosomes, one much smaller than the others, and two acrocentric Xs. From the Chinook Lake specimen we have been able to re-score eight nuclei in first division with 15 figures, all appearing as acrocentrics, and one much smaller than the others. The quality of those nuclei is now too poor to distinguish the Xs. However, three other metaphase nuclei in which the autosomes are not countable show clearly the two acrocentric Xs heteropycnotic and lying side by side and toward one pole.
Attulus (Sitticus) pubescens: 26a+XaXmYa (Figs
Attulus (Sitticus) terebratus: 26a+XaXa0 (Fig.
Chromosomes of meiosis of Attulus subgenus Sitticus 154 Attulus fasciger, three nuclei, one showing the two Xs together and toward a pole, Canada (43.351N, 79.759W) 155–163 Attulus pubescens, with XaXmYa sex chromosomes, Massachusetts (42.38N, 71.12W) 157–161 XmYa sex chromosomes from other nuclei; the second X is often not paired with them 162, 163 Second division nuclei, all having 14 acrocentrics, and some having in addition a metacentric (m) 164 Attulus terebratus, two nuclei (26a+XaXa0), Novosibirsk Oblast (53.730N, 77.866E).
While salticids are fairly conservative in basic chromosome complement, with most species showing 26 acrocentric autosomes and X1X20 sex chromosomes (
Chromosome evolution in sitticines. Ancestral nodes show the most parsimonious reconstruction of the evolution of Y via X-autosome fusions (black) from the X1X20 sex chromosome system (white). Phylogeny from Figure
The ancestral autosome number in sitticines is unclear. Among the species with XX0, some have 26 autosomes, others have 28. Assessing a comparable autosome number with neo-Y species requires interpretation, as the neo-Y system itself binds one or more autosomal pairs with the X chromosomes, as indicated in part by distinctive condensation patterns. If (as in Habronattus,
An unanticipated but consistent correlation between base autosome number and the presence of neo-Y is seen in Fig.
If these small chromosomes are supernumerary (B) chromosomes, it is possible that there is considerably more variation within species than our small sample sizes can detect. Undetected intraspecific variation in autosomes or sex chromosomes would not negate our basic evolutionary conclusions. Were we to find species variable with respect to the presence of a neo-Y chromosome, for example, it would point to even more transitions between XX0 and XXY/XXXY.
Our uncertainty about chromosome complement in some species does not strongly affect our conclusions about homoplasy or correlations, though it could affect a detailed reconstruction of the evolution of autosome number, or of particular fusions involved in a neo-Y system. For instance, if we delete autosome number for Attinella dorsata and Attulus burjaticus (the two species with uncertain counts) from Fig.
Chromosome evolution of sitticines will not be well understood, however, until a larger sample of species and specimens is obtained, given the high diversity seen in our small sample. Our data hint to the possibility of rapid evolution provoked by special mechanisms.
We are grateful to several colleagues who made special efforts to provide us access to specimens: to Galina Azarkina for greatly facilitating collecting in Siberia, to Gergin Blagoev for preparing and taking photographs of Attulus sylvestris specimens used for barcoding, and to Cristian Grismado for taking photographs of the holotype of Jollas puntalara Galiano. We thank Petra Kranebitter of the Naturmuseum Südtirol/ Museo di Scienze Naturali dell’Alto Adige for lending us the specimen of Attulus (Sittilong) longipes, Simone Ballini for collecting it, and Tobias Bauer for leading us to it. Laura Leibensperger and Jennifer Zaspel both helped generously in our attempts to trace what are the true types of Attus palustris Peckham and Peckham. Laura Leibensperger and Gonzalo Giribet of the