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.

An annotated checklist and a family key to the pseudoscorpion fauna (Arachnida: Pseudoscorpiones) of Sri Lanka.

Sri Lanka is part of the Western Ghats & Sri Lanka biodiversity hotspot. Thus, the conservation of Sri Lanka's unique biodiversity is crucial. The current study is part of an ongoing survey of pseudoscorpion fauna of Sri Lanka. We carried out an island-wide survey of pseudoscorpions using a range of collection methods to sample a diverse set of habitats around the country. This produced 32 species, four of which might be new to science, belonging to 25 genera. The family Cheiridiidae was discovered on the island for the first time. One new combination, Indogarypus ceylonicus (Beier, 1973) comb. nov., is proposed. Out of the 47 species now recorded, 20 (43 %) are potentially endemic to Sri Lanka. We provide a checklist of all known species, document their distribution and give a key to the families.

On the crab spider genus Angaeus Thorell, 1881 and its junior synonym Paraborboropactus Tang and Li, 2009 (Araneae: Thomisidae).

Morphological comparison of the genera Angaeus Thorell, 1881 and Paraborboropactus Tang and Li, 2009 (Araneae: Thomisidae) shows that the latter should be relegated to a junior synonym of the former. Further, I propose the following new synonyms: Angaeus leucomenus (Thorell, 1895), Stephanopis weyersi Simon, 1899 and Paraborboropactus leguminaceus Tang and Li, 2009 = Angaeus rhombifer Thorell, 1890 syn. nov., Paraborboropactus oblatus Tang and Li, 2010 = Angaeus lenticulosus Simon, 1903 syn. nov. The following species are transferred from Paraborboropactus: Angaeus canalis (Tang & Li, 2010) comb. nov., Angaeus liangweii (Tang & Li, 2010) comb. nov., Angaeus rhombus (Tang & Li, 2009) comb. nov. and Angaeus zhengi (Tang & Li, 2009) comb. nov. The holotype of Angaeus pudicus (type species of the genus) is illustrated, and the male and female of Angaeus rhombifer are redescribed and illustrated based partly on fresh material from Singapore. A new species, Angaeus christae sp. nov. is described based on both sexes, and a syntype of Angaeus comatulus Simon, 1909 is illustrated.

Local stochastics and ecoclimatic situation shape phytophagous chafer assemblage composition.

Very little is known about factors determining the assemblage structure of megadiverse polyphagous-herbivore scarab chafers in the tropics (Coleoptera: Scarabaeidae). Here, we examined the composition of Sri Lankan chafer assemblages and investigated whether it is influenced more by the general ecoclimatic situation, macrohabitat, or indetermined stochastic biotic and abiotic factors of each locality. We also explored the influence of the latter on separate lineages and general body size. Based on dedicated field surveys conducted during the dry and wet seasons, we examined 4847 chafer individuals of 105 species sampled using multiple UV-light traps in 11 localities covering different forest types and altitudinal zones. Assemblages were assessed for compositional similarity, species diversity, and abundance within four major eco-spatial partitions: forest types, elevational zones, localities, and macrohabitats. Our results revealed that assemblages were shaped mainly by locality stochastics (i.e., multi-factor ensemble of all biotic and abiotic environmental conditions at local scale), and to a minor extent by ecoclimatic conditions. Macrohabitat had little effect on the assemblage composition. This was true for the entire chafer assemblage as well as for all single lineages or different body size classes. However, in medium and large species the contrasts between localities were less pronounced, which was not the case for individual lineages of the assemblage. Contrasts of assemblage similarity between localities were much more evident than those for forest types and elevation zones. Significant correlation between species composition and geographic distance was found only for the assemblage of small-bodied specimens. Seasonal change (dry-wet) in species composition was minor and only measurable in a few localities. The strong turnover between examined localities corroborates with the high degree of endemism in many phytophagous chafers, particularly in Sericini. Connected with their hypothetic poor habitat specificity and polyphagy, this might also explain why so many chafer crop pests in the Asian tropics are endemics.

Origin and diversification of free-living stick spiders of Sri Lanka including the description of four new species of Rhomphaea L. Koch, 1872 and two new species of Neospintharus Exline, 1950.

Sri Lanka is a biologically diverse South Asian island, and together with the Western Ghats (Southern India) is one of the 36 world's most biologically diverse areas. Here, we investigated the origin and diversification of Rhomphaea and Neospintharus of Sri Lanka using sequences of three genes: mitochondrial cytochrome c oxidase I (COI) and 16S rRNA (16S); and nuclear 28S rRNA (28S). Our phylogeny included 32 taxa (30 ingroup and 2 outgroup). We used Bayesian Inference and maximum likelihood methods to reconstruct the placement of species, divergence time estimations and their foraging behavior with an emphasis on species from Sri Lanka. Our phylogenetic hypothesis support the monophyly of Argyrodinae as well as the monophyly of Rhomphaea, where Rhomphaea is a sister group of Neospintharus. Further, our analysis also suggests that Sri Lanka was colonized by Argyrodinae several times. Additionally, the following new species are described: Rhomphaea shanthi sp. nov., Rhomphaea jacko sp. nov., Rhomphaea martini sp. nov., Rhomphaea marani sp. nov., Neospintharus kandelensis sp. nov. and Neospintharus ohiyiaensis sp. nov.

Multiple species delimitation approaches with COI barcodes poorly fit each other and morphospecies - An integrative taxonomy case of Sri Lankan Sericini chafers (Coleoptera: Scarabaeidae).

DNA taxonomy including barcoding and metabarcoding is widely used to explore the diversity in biodiversity hotspots. In most of these hotspot areas, chafers are represented by a multitude of species, which are well defined by the complex shape of male genitalia. Here, we explore how well COI barcode data reflect morphological species entities and thus their usability for accelerated species inventorization. We conducted dedicated field surveys in Sri Lanka to collect the species-rich and highly endemic Sericini chafers (Coleoptera: Scarabaeidae). Congruence among results of a series of protocols for de novo species delimitation and with morphology-based species identifications was investigated. Different delimitation methods, such as the Poisson tree processes (PTP) model, Statistical Parsimony Analysis (TCS), Automatic Barcode Gap Discovery (ABGD), Assemble Species by Automatic Partitioning (ASAP), and Barcode Index Number (BIN) assignments, resulted in different numbers of molecular operational taxonomic units (MOTUs). All methods showed both over-splitting and lumping of morphologically identified species. Only 18 of the observed 45 morphospecies perfectly matched MOTUs from all methods. The congruence of delimitation between MOTUs and morphospecies expressed by the match ratio was low, ranging from 0.57 to 0.67. TCS and multirate PTP (mPTP) showed the highest match ratio, while (BIN) assignment resulted in the lowest match ratio and most splitting events. mPTP lumped more species than any other method. Principal coordinate analysis (PCoA) on a match ratio-based distance matrix revealed incongruent outcomes of multiple DNA delimitation methods, although applied to the same data. Our results confirm that COI barcode data alone are unlikely to correctly delimit all species, in particular, when using only a single delimitation approach. We encourage the integration of various approaches and data, particularly morphology, to validate species boundaries.

Distributional and taxonomic notes on the crab spider genus Talaus Simon, 1886 with description of a new species (Araneae: Thomisidae).

Taxonomic notes on the crab spider genus Talaus Simon, 1886 are provided. One new species, T. beccarii sp. nov. is described. Further four species are redescribed: T. nanus Thorell, 1890, T. triangulifer Simon, 1886, T. oblitus O. Pickard-Cambridge, 1899 and T. opportunus (O. Pickard-Cambridge, 1873). Talaus now contains 13 species.

First record of Synagelides Strand, 1906 (Araneae: Salticidae) from Sri Lanka: description of four endemic species from tropical wet forest of the island.

Synagelides Strand, 1906 is recorded for the first time in Sri Lanka. Four new species are described and illustrated: S. hortonensis sp. nov., S. lakmalii sp. nov., S. rosalindae sp. nov. and S. orlandoi sp. nov. A key to the four new species is given.

A phylogenetic and taxonomic review of baviine jumping spiders (Araneae, Salticidae, Baviini).

The systematics and taxonomy of the tropical Asian jumping spiders of the tribe Baviini is reviewed, with a molecular phylogenetic study (UCE sequence capture, traditional Sanger sequencing) guiding a reclassification of the group's genera. The well-studied members of the group are placed into six genera: Bavia Simon, 1877, Indopadilla Caleb & Sankaran, 2019, Padillothorax Simon, 1901, Piranthus Thorell, 1895, Stagetillus Simon, 1885, and one new genus, Maripanthus Maddison, gen. nov. The identity of Padillothorax is clarified, and Bavirecta Kanesharatnam & Benjamin, 2018 synonymized with it. Hyctiota Strand, 1911 is synonymized with Stagetillus. The molecular phylogeny divides the baviines into three clades, the Piranthus clade with a long embolus (Piranthus, Maripanthus), the genus Padillothorax with a flat body and short embolus, and the Bavia clade with a higher body and (usually) short embolus (remaining genera). In general, morphological synapomorphies support or extend the molecularly delimited groups. Eighteen new species are described: Bavia nessagyna, Indopadilla bamilin, I. kodagura, I. nesinor, I. redunca, I. redynis, I. sabivia, I. vimedaba, Maripanthus draconis (type species of Maripanthus), M. jubatus, M. reinholdae, Padillothorax badut, P. mulu, Piranthus api, P. bakau, P. kohi, P. mandai, and Stagetillus irri, all sp. nov., with taxonomic authority W. Maddison. The distinctions between baviines and the astioid Nungia Zabka, 1985 are reviewed, leading to four species being moved into Nungia from Bavia and other genera. Fifteen new combinations are established: Bavia maurerae (Freudenschuss & Seiter, 2016), Indopadilla annamita (Simon, 1903), I. kahariana (Prószynski & Deeleman-Reinhold, 2013), I. sonsorol (Berry, Beatty & Prószynski, 1997), I. suhartoi (Prószynski & Deeleman-Reinhold, 2013), Maripanthus menghaiensis (Cao & Li, 2016), M. smedleyi (Reimoser, 1929), Nungia hatamensis (Thorell, 1881), N. modesta (Keyserling, 1883), N. papakula (Strand, 1911), N. xiaolonghaensis (Cao & Li, 2016), Padillothorax casteti (Simon, 1900), P. exilis (Cao & Li, 2016), P. flavopunctus (Kanesharatnam & Benjamin, 2018), Stagetillus banda (Strand, 1911), all comb. nov. One combination is restored, Bavia capistrata (C. L. Koch, 1846). Five of these new or restored combinations correct previous errors of placing species in genera that have superficially similar palps but extremely different body forms, in fact belonging in distantly related tribes, emphasizing that the general shape of male palps should be used with caution in determining relationships. A little-studied genus, Padillothorus Prószynski, 2018, is tentatively assigned to the Baviini. Ligdus Thorell, 1895 is assigned to the Ballini.

The crab spider genus Tarrocanus Simon, 1895 with notes on the generabr />Alcimochthes Simon, 1895 and Domatha Simon, 1895 (Araneae:Thomisidae).

The crab spider genus Tarrocanus Simon, 1895 currently includes two species: T. capra Simon, 1895 and T. viridis Dyal, 1935. Recent field work revealed the presence of a new species, providing an opportunity to review the genus. The new species is described as T. jaffnaensis sp. nov. Furthermore, the male of T. capra is described for the first time, and taxonomic notes on Alcimochthes Simon, 1885 and Domatha Simon, 1895 are given, both presumably being close relatives of Tarrocanus. Tarrocanus viridis Dyal, 1935 is considered as nomen dubium.

Giant assassin in the cave: a new species of the genus Myiophanes from Sri Lanka (Hemiptera: Heteroptera: Reduviidae: Emesinae).

A new thread-legged assassin bug species, Myiophanes (Myiophanes) wygodzinskyi sp. nov. (Hemiptera: Heteroptera: Reduviidae: Emesinae: Emesini), collected from Ravana Cave in Uva Province of Sri Lanka, is described. It is the largest species of the genus described so far.

Repeated Diversification of Ecomorphs in Hawaiian Stick Spiders.

Insular adaptive radiations in which repeated bouts of diversification lead to phenotypically similar sets of taxa serve to highlight predictability in the evolutionary process [1]. However, examples of such replicated events are rare. Cross-clade comparisons of adaptive radiations are much needed to determine whether similar ecological opportunities can lead to the same outcomes. Here, we report a heretofore uncovered adaptive radiation of Hawaiian stick spiders (Theridiidae, Ariamnes) in which different species exhibit a set of discrete ecomorphs associated with different microhabitats. The three primary ecomorphs (gold, dark, and matte white) generally co-occur in native forest habitats. Phylogenetic reconstruction mapped onto the well-known chronosequence of the Hawaiian Islands shows both that this lineage colonized the islands only once and relatively recently (2-3 mya, when Kauai and Oahu were the only high islands in the archipelago) and that the distinct ecomorphs evolved independently multiple times following colonization of new islands. This parallel evolution of ecomorphs matches that of "spiny-leg" long-jawed spiders (Tetragnathidae, Tetragnatha), also in Hawaii [2]. Both lineages are free living, and both have related lineages in the Hawaiian Islands that show quite different patterns of diversification with no evidence of deterministic evolution. We argue that repeated evolution of ecomorphs results from a rugged adaptive landscape, with the few peaks associated with camouflage for these free-living taxa against the markedly low diversity of predators on isolated islands. These features, coupled with a limited genetic toolbox and reduced dispersal between islands, appear to be common to situations of repeated evolution of ecomorphs.

A new species of Angaeus from Malaysia with possible affinity to related genera within Stephanopinae (Araneae: Thomisidae).

The crab spider genus Angaeus Thorell, 1881 currently contains 10 described species (Benjamin 2013; WSC 2017). All species of the genus are restricted to tropical Asia. The aim of this correspondence is to illustrate and describe a new species of the genus characterized by a number of features previously found in the genera Angaeus, Borboropactus Simon, 1884, Epidius Thorell, 1877 and Geraesta Simon, 1889. The most unusual feature is the elongated tibia of the male palp that was previously thought to be diagnostic of Epidius (Figs 1, 2, 8; character 1 in Benjamin 2011; Benjamin 2017). However, the new species lacks tibial macrosetae (Figs 1, 2, 8) and lacks a flexibly attached MA, both also being characteristics of Epidius (characters 2 and 18 in Benjamin 2011). Furthermore, this new species differs considerably in general appearance from all known species of Epidius.

Description of three new species of the tropical Asian jumping spider genus Onomastus Simon, 1900 from high altitude cloud forests of Sri Lanka (Araneae: Salticidae).

Spiders of the tropical Asian jumping spider genus Onomastus Simon, 1900 are small to medium-sized, delicate, translucent, commonly found inhabitants of Asian evergreen forest foliage. In this paper, three new species of the genus, O. jamestaylori sp. nov. (♂♀), O. corbetensis sp. nov. (♂♀) and O. maskeliya sp. nov. (♂♀) are described from Sri Lanka. The three new species are added to the matrix of a previous study to assess their phylogenetic position. The resulting cladistic analysis, based on 35 morphological characters from 18 taxa (13 Onomastus species and 5 outgroups) supports the monophyly of the genus. Additionally, a monophyletic, well-supported South Asian clade (India, Sri Lanka), which is restricted to the Sri Lanka-Western Ghats biodiversity hotspot, is recovered in most analysis. The three newly described species might be endangered due to their small population size and restricted distribution in high altitude cloud forest.

The Goblin spider genus Xestaspis in Sri Lanka (Araneae: Oonopidae).

Three new species of the genus Xestaspis Simon, 1884: X. nuwaraeliya sp. nov., X. padaviya sp. nov. and X. pophami sp. nov. are described based on both sexes. Xestaspis kandy Eichenberger, 2012 is illustrated based on new material. A key and a distribution map for Sri Lankan Xestaspis is provided. Phylogenetic placement of Sri Lankan Xestaspis spp. obtained by the analysis of 43 morphological characters is presented and discussed. In all our analysis Gamasomorpha species appear paraphyletic suggesting that Xestaspis is most probably a junior synonym of Gamasomorpha. Thus, forming monophyletic group of oriental hard bodied oonopids.

A review of Sri Lankan Brignolia including the description of four new species (Araneae: Oonopidae).

Sri Lankan species of the genus Brignolia Dumitrescu & Georgescu, 1983 are reviewed and four new species are described: B. carlmulleri sp. nov., B. meemure sp. nov., B. ondaatjei sp. nov. and B. shyami sp. nov. All new species are described based on both sexes. New data on B. ambigua (Simon, 1893), B. parumpunctata (Simon, 1893), B. ratnapura Platnick et al., 2011 and B. trichinalis (Benoit, 1979) are given. A total of 10 species are now known from Sri Lanka. B. ambigua, B. carlmulleri sp. nov., B. meemure sp. nov., B. ondaatjei sp. nov., B. ratnapura, B. shyami sp. nov. and B. sinharaja are endemic to the island. A key and a distribution map for Sri Lankan Brignolia are provided.

Two new species of Pharta Thorell, 1891 with the description of Ibana senagang gen. et sp. nov. (Araneae: Thomisidae).

Two new species of Pharta, P. sudmannorum sp. nov. (♂♀, Borneo) and P. koponeni sp. nov. (♂, Thailand) are described. Furthermore, Ibana senagang gen. nov. & sp. nov. from Malaysia is described based on its exceptional palp, which has a reduced, movable conductor and thick-long spines on the distal, ventral surface of the tibia, reminiscent of Epidius Thorell, 1877.

Phylogenetic affinities of Phobetinus to other pirate spider genera (Araneae: Mimetidae) as indicated by spinning field morphology.

Spinnerets from Phobetinus sagittifer and an undescribed Phobetinus species were examined by scanning electron microscopy to gain a better understanding of this genus' relationships to other genera in the family Mimetidae. Consistent with placement of Phobetinus in Mimetinae, females possessed two synapomorphies of this subfamily; enlarged cylindrical silk gland spigots with domed shafts and a single cylindrical spigot per posterior lateral spinneret (PLS). Spinning field features overall suggest Phobetinus is most closely related to Mimetus, followed by Australomimetus, then Ero. A possible synapomorphy of a clade including Mimetus and Phobetinus is a pair of modified piriform silk gland spigots on each anterior lateral spinneret of adult males located adjacent to the secondary major ampullate silk gland tartipore. These spigots were present in P. sagittifer; however, similarly positioned spigots in the undescribed species were not obviously modified (i.e., wider or with larger openings relative to the other piriform spigots). Close affinity to Mimetus was also indicated by tartipore-accommodated PLS aciniform silk glands in both Phobetinus species. These have been consistently observed in Mimetus, but not in Australomimetus or Ero. Somatic and genitalic drawings of P. sagittifer are provided to aid identification and similarities are noted between male pedipalps of Mimetus and Phobetinus.