Phylogeny and biogeography of harmochirine jumping spiders (Araneae: Salticidae).

We use ultraconserved elements (UCE) and Sanger data to study the phylogeny, age, and biogeographical history of harmochirine jumping spiders, a group that includes the species-rich genus Habronattus, whose remarkable courtship has made it the focus of studies of behaviour, sexual selection, and diversification. We recovered 1947 UCE loci from 43 harmochirine taxa and 4 outgroups, yielding a core dataset of 193 UCEs with at least 50 % occupancy. Concatenated likelihood and ASTRAL analyses confirmed the separation of harmochirines into two major clades, here designated the infratribes Harmochirita and Pellenita. Most are African or Eurasian with the notable exception of a clade of pellenites containing Habronattus and Pellenattus of the Americas and Havaika and Hivanua of the Pacific Islands. Biogeographical analysis using the DEC model favours a dispersal of the clade's ancestor from Eurasia to the Americas, from which Havaika's ancestor dispersed to Hawaii and Hivanua's ancestor to the Marquesas Islands. Divergence time analysis on 32 loci with 85 % occupancy, calibrated by fossils and island age, dates the dispersal to the Americas at approximately 4 to 6 million years ago. The explosive radiation of Habronattus perhaps began only about 4 mya. The phylogeny clarifies both the evolution of sexual traits (e.g., the terminal apophyses was enlarged in Pellenes and not subsequently lost) and the taxonomy. Habronattus is confirmed as monophyletic. Pellenattus is raised to the status of genus, and 13 species moved into it as new combinations. Bianor stepposus Logunov, 1991 is transferred to Sibianor, and Pellenes bulawayoensis Wesolowska, 1999 is transferred to Neaetha. A molecular clock rate estimate for spider UCEs is presented and its utility to inform prior distributions is discussed.

Phylogeny with introgression in Habronattus jumping spiders (Araneae: Salticidae).

BACKGROUND: Habronattus is a diverse clade of jumping spiders with complex courtship displays and repeated evolution of Y chromosomes. A well-resolved species phylogeny would provide an important framework to study these traits, but has not yet been achieved, in part because the few genes available in past studies gave conflicting signals. Such discordant gene trees could be the result of incomplete lineage sorting (ILS) in recently diverged parts of the phylogeny, but there are indications that introgression could be a source of conflict. RESULTS: To infer Habronattus phylogeny and investigate the cause of gene tree discordance, we assembled transcriptomes for 34 Habronattus species and 2 outgroups. The concatenated 2.41 Mb of nuclear data (1877 loci) resolved phylogeny by Maximum Likelihood (ML) with high bootstrap support (95-100%) at most nodes, with some uncertainty surrounding the relationships of H. icenoglei, H. cambridgei, H. oregonensis, and Pellenes canadensis. Species tree analyses by ASTRAL and SVDQuartets gave almost completely congruent results. Several nodes in the ML phylogeny from 12.33 kb of mitochondrial data are incongruent with the nuclear phylogeny and indicate possible mitochondrial introgression: the internal relationships of the americanus and the coecatus groups, the relationship between the altanus, decorus, banksi, and americanus group, and between H. clypeatus and the coecatus group. To determine the relative contributions of ILS and introgression, we analyzed gene tree discordance for nuclear loci longer than 1 kb using Bayesian Concordance Analysis (BCA) for the americanus group (679 loci) and the VCCR clade (viridipes/clypeatus/coecatus/roberti groups) (517 loci) and found signals of introgression in both. Finally, we tested specifically for introgression in the concatenated nuclear matrix with Patterson's D statistics and DFOIL. We found nuclear introgression resulting in substantial admixture between americanus group species, between H. roberti and the clypeatus group, and between the clypeatus and coecatus groups. CONCLUSIONS: Our results indicate that the phylogenetic history of Habronattus is predominantly a diverging tree, but that hybridization may have been common between phylogenetically distant species, especially in subgroups with complex courtship displays.

Co-occurrence of related asexual, but not sexual, lineages suggests that reproductive interference limits coexistence.

We used randomizations to analyse patterns of co-occurrence of sexual and apomictic (asexual) members of the North American Crepis agamic complex (Asteraceae). We expect strong asymmetry in reproductive interactions in Crepis: apomicts produce clonal seeds with no need for pollination and are not subject to reproductive interference from co-occurring relatives. However, because they still produce some viable pollen, apomicts can reduce reproductive success of nearby sexual relatives, potentially leading to eventual local exclusion of sexuals. Consistent with this, randomizations reveal that sexuals are over-represented in isolated sites, while apomicts freely co-occur. Incorporation of taxonomic and phylogenetic evidence indicates that this pattern is not driven by local origins of asexuals. Our evidence that patterns of local co-occurrence are structured by reproductive interference suggests an underappreciated role for these interactions in community assembly, and highlights the need for explicit tests of the relative contributions of ecological and reproductive interactions in generating patterns of limiting similarity.

The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling.

We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher-level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb-weavers, compatible with their non-monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, with Amphinecta, Mamoea, Maniho, Paramamoea and Rangitata (transferred from Amphinectidae); Ischaleinae, with Bakala and Manjala (transferred from Amaurobiidae) and Ischalea (transferred from Stiphidiidae); Metaltellinae, with Austmusia, Buyina, Calacadia, Cunnawarra, Jalkaraburra, Keera, Magua, Metaltella, Penaoola and Quemusia; Porteriinae (new rank), with Baiami, Cambridgea, Corasoides and Nanocambridgea (transferred from Stiphidiidae); and Desinae, with Desis, and provisionally Poaka (transferred from Amaurobiidae) and Barahna (transferred from Stiphidiidae). Argyroneta is transferred from Cybaeidae to Dictynidae. Cicurina is transferred from Dictynidae to Hahniidae. The genera Neoramia (from Agelenidae) and Aorangia, Marplesia and Neolana (from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, with Gasparia, Gohia, Hulua, Neomyro, Myro, Ommatauxesis and Otagoa (transferred from Desidae); and Toxopinae, with Midgee and Jamara, formerly Midgeeinae, new syn. (transferred from Amaurobiidae) and Hapona, Laestrygones, Lamina, Toxops and Toxopsoides (transferred from Desidae). We obtain a monophyletic Oval Calamistrum clade and Dionycha; Sparassidae, however, are not dionychans, but probably the sister group of those two clades. The composition of the Oval Calamistrum clade is confirmed (including Zoropsidae, Udubidae, Ctenidae, Oxyopidae, Senoculidae, Pisauridae, Trechaleidae, Lycosidae, Psechridae and Thomisidae), affirming previous findings on the uncertain relationships of the "ctenids" Ancylometes and Cupiennius, although a core group of Ctenidae are well supported. Our data were ambiguous as to the monophyly of Oxyopidae. In Dionycha, we found a first split of core Prodidomidae, excluding the Australian Molycriinae, which fall distantly from core prodidomids, among gnaphosoids. The rest of the dionychans form two main groups, Dionycha part A and part B. The former includes much of the Oblique Median Tapetum clade (Trochanteriidae, Gnaphosidae, Gallieniellidae, Phrurolithidae, Trachelidae, Gnaphosidae, Ammoxenidae, Lamponidae and the Molycriinae), and also Anyphaenidae and Clubionidae. Orthobula is transferred from Phrurolithidae to Trachelidae. Our data did not allow for complete resolution for the gnaphosoid families. Dionycha part B includes the families Salticidae, Eutichuridae, Miturgidae, Philodromidae, Viridasiidae, Selenopidae, Corinnidae and Xenoctenidae (new fam., including Xenoctenus, Paravulsor and Odo, transferred from Miturgidae, as well as Incasoctenus from Ctenidae). We confirm the inclusion of Zora (formerly Zoridae) within Miturgidae.

Stark sexual display divergence among jumping spider populations in the face of gene flow.

Gene flow can inhibit evolutionary divergence by eroding genetic differences between populations. A current aim in speciation research is to identify conditions in which selection overcomes this process. We focused on a state of limited differentiation, asking whether selection enables divergence with gene flow in a set of Habronattus americanus jumping spider populations that exhibit three distinct male sexual display morphs. We found that each population is at high frequency or fixed for a single morph. These strong phenotypic differences contrast with low divergence at 210 AFLP markers, suggesting selection has driven or maintains morph divergence. Coinciding patterns of isolation by distance and 'isolation by phenotype' (i.e. increased genetic divergence among phenotypically contrasting populations) across the study area support several alternative demographic hypotheses for display divergence, each of which entails gene flow. Display-associated structure appears broadly distributed across the genome and the markers producing this pattern do not stand out from background levels of differentiation. Overall, the results suggest selection can promote stark sexual display divergence in the face of gene flow among closely related populations.

Hivanua, a new genus of harmochirine jumping spiders from the Marquesas Islands (Araneae, Salticidae, Harmochirina).

The genus Hivanuagen. nov. is established for the harmochirine jumping spiders of the Marquesas Islands, formerly placed in Habronattus F.O. Pickard-Cambridge, 1901 and Havaika Prószynski, 2002. The type species, Hivanuatekaosp. nov. is described, and five species described by Berland are re-illustrated and moved into the genus: Hivanuaflavipes (Berland, 1933), comb. nov., Hivanuanigrescens (Berland, 1933), comb. nov., Hivanuanigrolineata (Berland, 1933), comb. nov., Hivanuarufescens (Berland, 1934), comb. nov., and Hivanuatriangulifera (Berland, 1933), comb. nov. The female epigyne is much like that of Habronattus, Bianor Peckham & Peckham, 1896, and other harmochirines, with a centrally placed coupling pocket and two atria with crescent-shaped edges. The terminal apophysis of the male palp, which is variable throughout the pellenine subgroup of the Harmochirina, is absent in H.rufescens but present in H.tekaosp. nov., in which it is elbowed much as in Habronattus. These Pacific Island harmochirines, like the Havaika of Hawaii, appear to be largely foliage dwellers, unlike most of their continental relatives.

Ghatippuspaschima, a new species and genus of plexippine jumping spider from the Western Ghats of India (Salticidae, Plexippini, Plexippina).

We propose a new genus of plexippine jumping spiders from the Western Ghats of India based on the new species Ghatippuspaschimagen. et sp. nov. While it bears a superficial resemblance to Pancorius in body form and Hyllus in membrane-bearing embolus, our UCE phylogenomic data-the first to resolve broad relationships within the Plexippina-as well as morphological features justify its status as a new genus. In addition to the molecular data and morphological descriptions, we provide photographs of living specimens of Ghatippuspaschimagen. et sp. nov. and information on their natural history.

On the enigmatic jumping spider genus Ogdenia Peckham, 1908 (Araneae, Salticidae, Chrysillini).

The monotypic genus Ogdenia Peckham, 1908, is redefined based on the redescription of the holotype of O.mutilla (Peckham & Peckham, 1907), along with the newly discovered male specimens and intraspecific variation from China, Malaysia, Singapore, and Vietnam. Description, illustrations, and photographs are provided.

Molecular phylogeny, divergence times and biogeography of spiders of the subfamily Euophryinae (Araneae: Salticidae).

We investigate phylogenetic relationships of the jumping spider subfamily Euophryinae, diverse in species and genera in both the Old World and New World. DNA sequence data of four gene regions (nuclear: 28S, Actin 5C; mitochondrial: 16S-ND1, COI) were collected from 263 jumping spider species. The molecular phylogeny obtained by Bayesian, likelihood and parsimony methods strongly supports the monophyly of a Euophryinae re-delimited to include 85 genera. Diolenius and its relatives are shown to be euophryines. Euophryines from different continental regions generally form separate clades on the phylogeny, with few cases of mixture. Known fossils of jumping spiders were used to calibrate a divergence time analysis, which suggests most divergences of euophryines were after the Eocene. Given the divergence times, several intercontinental dispersal events are required to explain the distribution of euophryines. Early transitions of continental distribution between the Old and New World may have been facilitated by the Antarctic land bridge, which euophryines may have been uniquely able to exploit because of their apparent cold tolerance. Two hot-spots of diversity of euophryines are discovered: New Guinea and the Caribbean Islands. Implications of the molecular phylogeny on the taxonomy of euophryines, and on the evolution of unusual genitalic forms and myrmecophagy, are also briefly discussed.

NeXML: rich, extensible, and verifiable representation of comparative data and metadata.

In scientific research, integration and synthesis require a common understanding of where data come from, how much they can be trusted, and what they may be used for. To make such an understanding computer-accessible requires standards for exchanging richly annotated data. The challenges of conveying reusable data are particularly acute in regard to evolutionary comparative analysis, which comprises an ever-expanding list of data types, methods, research aims, and subdisciplines. To facilitate interoperability in evolutionary comparative analysis, we present NeXML, an XML standard (inspired by the current standard, NEXUS) that supports exchange of richly annotated comparative data. NeXML defines syntax for operational taxonomic units, character-state matrices, and phylogenetic trees and networks. Documents can be validated unambiguously. Importantly, any data element can be annotated, to an arbitrary degree of richness, using a system that is both flexible and rigorous. We describe how the use of NeXML by the TreeBASE and Phenoscape projects satisfies user needs that cannot be satisfied with other available file formats. By relying on XML Schema Definition, the design of NeXML facilitates the development and deployment of software for processing, transforming, and querying documents. The adoption of NeXML for practical use is facilitated by the availability of (1) an online manual with code samples and a reference to all defined elements and attributes, (2) programming toolkits in most of the languages used commonly in evolutionary informatics, and (3) input-output support in several widely used software applications. An active, open, community-based development process enables future revision and expansion of NeXML.

Two new genera of Euophryini from southern China and Malaysia (Araneae, Salticidae, Salticinae).

Two new genera of euophryine jumping spiders in the Laufeia group are established for species from Southeast Asia: Amoenema Yu & Zhang, gen. nov. and Lokina Yu, Maddison & Zhang, gen. nov., with descriptions of 12 new species: Amoenema erhai Yu & Zhang, sp. nov. (♂♀, type species), A. liuae Yu & Zhang, sp. nov. (♂♀), A. zizhongi Yu & Zhang, sp. nov. (♂♀), Lokina blombergi Yu, Maddison & Zhang, sp. nov. (♂♀), L. chiyou Yu & Zhang, sp. nov. (♂♀, type species), L. fuxi Yu & Zhang, sp. nov. (♂♀), L. kubah Yu, Maddison & Zhang, sp. nov. (♂), L. nyuewa Yu & Zhang, sp. nov. (♂♀), L. pixi Yu, Maddison & Zhang, sp. nov. (♂), L. tamasi Yu & Zhang, sp. nov. (♂♀), L. wuyi Yu & Zhang, sp. nov. (♂), L. zhishengi Yu & Zhang, sp. nov. (♂). In addition, three new combinations are proposed: Amoenema robusta (Lei & Peng, 2012) comb. nov. (transferred from Euophrys C. L. Koch, 1834), Lokina eximia (Zhang & Maddison, 2012) comb. nov. (transferred from Laufeia Simon, 1889) and L. tengchongensis (Lei & Peng, 2012) comb. nov. (transferred from Chalcovietnamicus Marusik, 1991).

Taxonomic revision of Orcevia Thorell, 1890, with description of fifteen new species (Araneae, Salticidae, Euophryini).

The Asian euophryine genus Orcevia Thorell, 1890 is revised, and fifteen new species from southern China and Southeast Asia are described: Orcevia bokoblin Yu, Maddison & Zhang, sp. nov. (), O. calcicola Yu, Maddison & Zhang, sp. nov. (), O. deelemanae Yu & Zhang, sp. nov. (), O. feitongae Yu & Zhang, sp. nov. (), O. gongae Yu & Zhang, sp. nov. (), O. jinping Yu & Zhang, sp. nov. (), O. meinei Yu, Maddison & Zhang, sp. nov. (), O. mercuryi Yu, Maddison & Zhang, sp. nov. (), O. nietzschei Yu, Maddison & Zhang, sp. nov. (), O. pakse Yu & Zhang, sp. nov. (), O. shuyuani Yu & Zhang, sp. nov. (), O. timburtoni Yu, Maddison & Zhang, sp. nov. (), O. wuliang Yu & Zhang, sp. nov. (), O. yahaha Yu, Maddison & Zhang, sp. nov. (), O. zabkai Yu, Maddison & Zhang, sp. nov. () and O. zu Yu & Zhang, sp. nov. (). Three new combinations and one new synonym are proposed: Orcevia sokoli (Prszyski & Deeleman-Reinhold, 2013) comb. nov. (transferred from Echeclus Thorell, 1890), O. sica (Wu & Yang, 2008) comb. nov. (=Laufeia longapophysis Lei & Peng, 2012 syn. nov.; transferred from Laufeia Simon, 1889), and Magyarus terrestris (Logunov, 2021) comb. nov. (transferred from Orcevia). The previously unknown females of Orcevia sica (Wu & Yang, 2008) comb. nov. and O. proszynskii (Song, Gu & Chen, 1988) are described. O. perakensis (Simon, 1901) is herein listed as a species inquirenda.

Review of Chalcovietnamicus Marusik, 1991, with description of four new species (Araneae, Salticidae, Euophryini).

The euophryine genus Chalcovietnamicus Marusik, 1991, a member of the Laufeia group of genera, is reviewed, and four new species from southern China and Southeast Asia are described with both sexes: Chalcovietnamicus logunovi Yu, Maddison & Zhang, sp. nov., C. marusiki Yu, Maddison & Zhang, sp. nov., C. terbakar Yu, Maddison & Zhang, sp. nov. and C. weihangi Yu & Zhang, sp. nov.. The monotypic genus Junxattus Prszyski & Deeleman-Reinhold, 2012 syn. nov. is considered as a junior synonym of Chalcovietnamicus Marusik, 1991, and Parvattus Zhang & Maddison, 2012 is revalidated and removed from synonymy with Chalcovietnamicus Marusik, 1991. The previously unknown female of C. vietnamensis (abka, 1985) is described based on specimens from Vietnam and Singapore. All species of Chalcovietnamicus are assigned to two species groups (daiqini-group and vietnamensis-group), except C. naga Logunov, 2020 that is herein considered as an incertae sedis.

The biogeography and age of salticid spider radiations (Araneae: Salticidae).

Globally distributed, jumping spiders (Salticidae) are species-rich and morphologically diverse. Recent molecular phylogenetic work has revealed that major clades are largely isolated to particular continental regions, suggesting their radiations postdated Mesozoic continental break up, but corroboration from a multi-gene time-calibrated phylogeny has been lacking, and an important tropical forest region, Central and West Africa, has been largely unsampled. Newly sampled species, many from Gabon, were included among taxa sequenced for the genes 28s, Actin 5C, 16sND1, and CO1. Likelihood and Bayesian analyses show that most of the Gabonese species from forest habitats fall into a single large clade, which we name the Thiratoscirtinae (new subfamily), within the broader Aelurilloida. The aelurilloids, together with the plexippoids, euophryines, heliophanines and smaller groups (e.g. Leptorchestae, Hasarieae, Philaeus group, Salticus), form a large clade that we name the Saltafresia. Most saltafresian diversification appears to have occured in Afro-Eurasia, with the exception of the euophryines (largely Neotropical, Australasian and Southeast Asian) and two radiations in the New World (Habronattus, freyines). Using Bayesian relaxed molecular-clock methods, calibrated by amber fossil data and a geological constraint, we estimate that most recent common ancestor of the family occurred 47-57 million years ago, when the continents would have already separated substantially. The Salticoida is dated to 41-50 million years, and its four major subclades Amycoida, Astioida, Marpissoida, and Saltafresia are each dated to 29-44 million years. By these inferred dates, salticids were radiating while the earth was warmer than today, with expanded megathermal forests and, most likely, diverse insect herbivores. Our phylogeny indicates mixing of radiating faunas from isolated regions has been limited, yet some long-range dispersal events, such as the arrival of the genus Habronattus to the New World, have occurred. Four African species formerly in Viciria are moved to Telamonia, establishing the new combinations Telamonia besanconi (Berland and Millot), Telamonia fuscimana (Simon), Telamonia longiuscula (Thorell), Telamonia thoracica (Thorell). The Marpissoida is expanded to include the Ballinae.

Revision of the genus Charippus Thorell, 1895, with descriptions of eight new species (Araneae, Salticidae, Euophryini).

The euophryine genus Charippus Thorell, 1895 is revised and eight new species are described: Charippus asper Yu, Maddison Zhang, sp. nov. ( ), C. bukittimah Yu, Maddison Zhang, sp. nov. ( ), C. callainus Yu, Maddison Zhang, sp. nov. ( ), C. denjii Yu, Maddison Zhang, sp. nov. ( ), C. heishiding Yu, Maddison Zhang, sp. nov. ( ), C. kubah Yu, Maddison Zhang, sp. nov. ( ), C. minotaurus Yu, Maddison Zhang, sp. nov. ( ) and C. wanlessi Yu, Maddison Zhang, sp. nov. ( ). A new combination for Charippus yunnanensis (Cao Li, 2016) comb. nov. (ex. Cytaea Keyserling, 1882) is proposed, and its female is described for the first time. Diagnostic illustrations and photographs are provided for all known species.

Three new and notes on two other jumping spider species of the genus Stenaelurillus Simon, 1886 (Salticidae: Aelurillina) from the Deccan Plateau, India.

We describe three new species of Stenaelurillus Simon, 1886 from the Deccan Plateau of India, and report on populations of S. sarojinae Caleb Mathai, 2014 and S. marusiki Logunov, 2001. One of the new species, S. shwetamukhi Marathe, Sanap, Maddison, sp. nov., has black-and-white markings, characteristic of several other Indian Stenaelurillus species. The other two new species, S. tamravarni Marathe Maddison, sp. nov., and S. vyaghri Sanap, Joglekar, Caleb, sp. nov., are colourful and with fringed male abdomens, like several other Indian species including S. sarojinae. The population of S. sarojinae from Mysuru, Karnataka, shows colours distinct from the population at the type locality. The female of S. marusiki is described for the first time.

Kelawakaju gen. nov., a new Asian lineage of marpissine jumping spiders (Araneae, Salticidae, Marpissina).

The genus Kelawakaju Maddison & Ruiz, gen. nov., is described for a lineage of bark-dwelling Asian marpissine jumping spiders that represent a dispersal to Eurasia separate from that of the Marpissa-Mendoza lineage, according to the phylogeny recovered from analysis of four gene regions. All species of Kelawakaju are new to science except Kelawakajufrenata (Simon, 1901), comb. nov., which is transferred from Ocrisiona Simon, 1901. Kelawakajufrenata is known from Hong Kong, Guangdong, Guangxi, and likely Taiwan. The five new species are Kelawakajumulu Maddison & Ruiz, sp. nov. (type species of Kelawakaju, from Sarawak, Malaysia, ♂♀), K.intexta Maddison & Ruiz, sp. nov. (from Sarawak, ♂), K.leucomelas Maddison & Ng, sp. nov. (Singapore and Johor Bahru, ♂♀), K.sahyadri Vishnudas, Maddison, & Sudhikumar, sp. nov. (India, ♂♀), and K.singapura Maddison & Ng, sp. nov. (Singapore, ♂♀).

Estimating trait-dependent speciation and extinction rates from incompletely resolved phylogenies.

Species traits may influence rates of speciation and extinction, affecting both the patterns of diversification among lineages and the distribution of traits among species. Existing likelihood approaches for detecting differential diversification require complete phylogenies; that is, every extant species must be present in a well-resolved phylogeny. We developed 2 likelihood methods that can be used to infer the effect of a trait on speciation and extinction without complete phylogenetic information, generalizing the recent binary-state speciation and extinction method. Our approaches can be used where a phylogeny can be reasonably assumed to be a random sample of extant species or where all extant species are included but some are assigned only to terminal unresolved clades. We explored the effects of decreasing phylogenetic resolution on the ability of our approach to detect differential diversification within a Bayesian framework using simulated phylogenies. Differential diversification caused by an asymmetry in speciation rates was nearly as well detected with only 50% of extant species phylogenetically resolved as with complete phylogenetic knowledge. We demonstrate our unresolved clade method with an analysis of sexual dimorphism and diversification in shorebirds (Charadriiformes). Our methods allow for the direct estimation of the effect of a trait on speciation and extinction rates using incompletely resolved phylogenies.

Sitticine jumping spiders: phylogeny, classification, and chromosomes (Araneae, Salticidae, Sitticini).

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 subtribeAillutticina (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 Prószynski, 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 Prószynski, 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 Prószynski, 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.