Matías A. Izquierdo

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.

Taxonomic Review of the Goblin Spiders of the Genus Dysderoides Fage and Their Himalayan Relatives of the Genera Trilacuna Tong and Li and Himalayana, New Genus (Araneae: Oonopidae)

ABSTRACT The study of many museum specimens of goblin spiders from the Himalayan range and neighboring countries allows for the description of new taxa of the family Oonopidae. The genus Dysderoides Fage is taxonomically reviewed. It comprises small, blind, loricate troglobitic spiders: the type species (D. typhlos Fage, from India) and at least five new species from northern India (D. synrang Grismado and Deeleman) and Thailand (D. muang Grismado and Deeleman, D. kaew Grismado and Deeleman, D. kanoi Grismado and Deeleman, and D. lawa Grismado and Deeleman). The genus Trilacuna, previously known from China, Thailand, Malaysia, and Sumatra, is newly diagnosed by the loss of the furrow connecting the posterior spiracles in males, and is represented in the Himalayan region by seven species: T. aenobarba (Brignoli), from Bhutan (here transferred from Epectris Simon), and six new: four from northern India (T. meghalaya Grismado and Piacentini, T. besucheti Grismado and Piacentini, T. mahanadi Grismado and Piacentini, and T. loebli Grismado and Piacentini), one from India and Nepal (T. bangla Grismado and Ramírez), and one from Pakistan (T. hazara Grismado and Ramírez). The new genus Himalayana Grismado comprises species very similar to those of Trilacuna, but differs in the characters of the postepigastric scuta and by having an additional acute dorso-prolateral projection on the male palpi. Six new species are assigned to Himalayana: H. kathmandu Grismado (type species), H. castanopsis Grismado, H. parbat Grismado, and H. martensi Grismado (all from Nepal); and H. siliwalae Grismado and H. andreae Grismado (from India). The study of the internal female genitalia of T. meghalaya and T. bangla revealed a complex copulatory system, and an entelegyne condition, apparently uniform for the entire genus and probably for Dysderoides and Himalayana as well. The males of the three genera have a complex set of paraembolic laminae with brushes of filiform structures, among which discharges a gland through a thin, tortuous cuticular tube. The genitalic and somatic morphology of the three genera suggest that they conform a monophyletic group, here named “Dysderoides complex,” and that their closer relatives can be found among Prethopalpus Baehr et al., and other genera related to Silhouettella Benoit. The loss of the membranous diagonal area on the base of the anterior lateral spinnerets is proposed as a synapomorphy of an advanced group of loricatae oonopids usually referred as gamasomorphines. Furthermore Triaeris glenniei Fage, described from a single female from a cave in Uttarakhand, is redescribed and transferred to Camptoscaphiella Caporiacco.

Description of the female of Orsolobus pucara Forster & Platnick 1985, with comments on the functional morphology of the female genitalia in Dysderoidea (Araneae: Dysderoidea: Orsolobidae)

Abstract The female of Orsolobus pucara Forster & Platnick 1985 is described and its genitalia examined using the scanning electron microscope (SEM). A small phylogenetic matrix with female genital and sexual behavior characters was made with the aim to study the evolution of these characters in the superfamily Dysderoidea. This is the first time that the female genitalia of a species of the family Orsolobidae have been studied in detail with SEM. The anterior portion of the female genitalia is a sclerotized structure with gland ducts and sites for muscle attachments. The posterior portion has a membranous receptaculum and a sclerotized plate that serves as attachment for muscles. We discuss the probable function of genital characters in a phylogenetic context. The anterior sclerotized elements of the female genitalia of some Dysderidae, Orsolobidae and Oonopidae species and the anterior receptaculum in the Segestriidae seem to be homologous structures because of the presence of gland ducts and sperm. However, both of these characteristics are lost in some species of these families, the anterior portion of the female genitalia being transformed into a highly modified structure serving mainly as attachment for muscles implicated in sexual behavior mechanisms.

The Amazonian Goblin Spiders of the New Genus Gradunguloonops (Araneae: Oonopidae)

A new genus of soft-bodied oonopids, Gradunguloonops, is established for a group of goblin spiders found in the Amazonian rainforests of northern South America. Members of this genus differ from other oonopids in that the proclaw of tarsi I and II is notably larger than the corresponding retroclaw, a putative synapomorphy of the group. Gradunguloonops comprises twelve species, all new and described in this contribution: G. mutum (type species) from Brazil and Peru, G. bonaldoi, G. amazonicus, G. urucu, G. pacanari, G. juruti from Brazil, G. erwini from Peru, G. orellana and G. nadineae from Ecuador, G. benavidesae and G. florezi from Colombia, and G. raptor from Venezuela. Two preliminary intrageneric groups are proposed on the basis of their female genital morphology: the bonaldoi group, to which are assigned the species with the anterior section comprising only a single anterior sclerite, and the mutum group, with a more complex, tripartite anterior section.

Male Genital Mutilation in the High-Mountain Goblin Spider, Unicorn catleyi

Abstract Male genital mutilation is a common mechanism by which males reduce sperm competition by plugging female insemination ducts with different parts of its own genital system. This behavior is frequent in many spider families but is uncommon in Haplogynae. The reproductive biology of Dysderoidea is not well studied and the data is fragmentary; male genital mutilation has been reported only for one species of Oonopidae. This study provides evidence of male genital mutilation in Unicorn catleyi Platnick and Brescovit (Araneae: Oonopidae). Pieces of the embolus were found in the female posterior receptaculum. This behavior is a strategy used by the males in order to guarantee their paternity and not for escape from female attacks as has been reported for other species of Araneae, since cannibalism is unlikely in this species. The presence of embolus in the posterior receptaculum suggests this is the first place where sperm is received. The similarity of the female genitalia of U. catleyi to those of Orsolobidae, along with sclerotization of the seminal duct in the male copulatory bulb that is also present in Orchestina, Xiombarg, and Orsolobidae, provide strong evidence of the basal position of this genus in the family Oonopidae.

Taxonomic revision of the jumping goblin spiders of the genus Orchestina Simon, 1882, in the Americas (Araneae, Oonopidae).

ABSTRACT The genus Orchestina Simon is distributed worldwide and is characterized by having an enlarged fourth femur with which these species are capable of jumping. The genus is also characterized by having a well-sclerotized sperm duct, a near H-shaped arrangement of the eye group, a 4-4-3-3 pattern of raised receptors on the tarsal organs of the legs I–IV, respectively, and by lacking spines on all the legs. All these characters, together with molecular evidence, support the monophyly of the genus, as well as its placement as sister group of Oonopinae. Most American species of Orchestina inhabit the forest foliage and canopy, but in unforested areas they also occur in shrubs and grasses. In this work, we revise the American representatives of Orchestina in a comprehensive context for the first time. In the past, six species were known from the Americas: five from the United States and only one from South America, described from Venezuela. After the study of the principal collections of the world and several field trips to several South American countries, we describe 85 new species and redescribe all previously known species. Matching sexes was occasionally problematic; while females are very homogeneous in somatic traits, males may have modifications on different parts of the body, making the matching very difficult. Therefore, in this review 56 of the species are described from only one sex, whereas 20 unmatched species are informally described as morphospecies, pending the discovery of conspecific sexes. Two species, O. pavesiiformis Saaristo and O. dentifera Simon, originally known from Israel and Sri Lanka, respectively, are here reported as introduced in several countries in the Americas and other continents. O. justini Saaristo described from the Seychelles is here considered a synonym of O. dentifera. One species, O. truncata Wunderlich, previously known as a subfossil spider from Colombian Copal is here tentatively redescribed based on recent material from Costa Rica, Colombia, and Ecuador; the female is also described for the first time. The species list by country is as follows (numbers refer to records, independently of the locality of the type material): United States (9 species: O. utahana Chamberlin and Ivie, O. moaba Chamberlin and Ivie, O. obscura Chamberlin and Ivie, O. saltitans Banks, O. nadleri Chickering, the introduced O. pavesiiformis Saaristo, and three new species, O. quasimodo, O. kamehameha, and O. auburndalensis); Mexico (3 species: O. utahana Chamberlin and Ivie, and two new species, O. nahuatl and O. chaparrita); Guatemala (1 new species: O. guatemala); Costa Rica (3 new species: O. laselva, O. griswoldi, and O. chiriqui; and the previously known O. truncata Wunderlich); Panama (5 new species: O. chiriqui, O. labarquei, O. pan, O. campana, and O. galapagos); Jamaica (2 species, the introduced O. dentifera Simon and O. galapagos); Haiti and Dominican Republic (only the introduced O. dentifera Simon); Colombia (6 new species: O. filandia, O. zingara, O. arboleda, O. cali, O. platnicki, O. pakitza; and O. truncata Wunderlich, plus the morphospecies OMI020 and OMI038); Venezuela (7 species: O. saltabunda; and 6 new, O. venezuela, O. aragua, O. bolivar, O. maracay, O. ranchogrande, and O. neblina); Trinidad and Tobago (1 new species: O. kairi); Guyana (1 morphospecies: OMI026); Ecuador (18 new species: O. galapagos, O. fernandina, O. erwini, O. ecuatoriensis, O. sotoi, O. magna, O. shuar, O. golem, O. waorani, O. tzantza, O. predator, O. goblin, O. yanayacu, O. otonga, O. santodomingo, O. quijos, O. mayo, O. laselva, and O. truncata Wunderlich, plus the morphospecies OMI020, OMI021, OMI022, OMI023, OMI024, OMI025, OMI026, OMI027, OMI029, OMI036, and OMI038); Peru (9 new species: O. cajamarca, O. comaina, O. atocongo, O. mancocapac, O. silvae, O. madrededios, O. pakitza, O. losamigos, O. golem, and the morphospecies OMI020, OMI023, OMI026, and OMI030); Brazil (25 new species: O. cristinae, O. coari, O. moura, O. valquiria, O. aproeste, O. caxiuana, O. para, O. taruma, O. retiro, O. divisor, O. juruti, O. platnicki, O. iemanja, O. bonaldoi, O. rapaz, O. itapety, O. catarina, O. leon, O. saudade, O. sarava, O. goblin, O. sotoi, O. golem, O. waorani, O. ucumar, the introduced O. pavesiiformis Saaristo, and O. dentifera Simon, plus the morphospecies OMI021, OMI023, OMI024, OMI025, OMI026, OMI027, OMI029, OMI032, OMI033, OMI034, OMI035, OMI036, OMI037, OMI039, and OMI040); Bolivia (3 new species: O. moyuchi, O. grismadoi, and O. ucumar); Chile (12 new species: O. pandeazucar, O. caleta, O. totoralillo, O. pizarroi, O. molles, O. granizo, O. quenies, O. curico, O. cachai, O. nahuelbuta, O. jaiba, and O. osorno); and Argentina (6 new species: O. ucumar, O. andianavarroi, O. luispi, O. cristinae, O. platnicki, O. jaiba, and the introduced O. pavesiiformis Saaristo). Although the internal relationships of the genus are still not clear, we suggest some groups of species based on morphological characters and geographic distributions. Species from the western United States share the presence of a net-shaped pattern of coloration and, in some species, modifications of carapace or first legs of males. Females of this group are characterized by the presence of a thin, generally long and twisted anterior receptaculum and external pockets on the epigastric region. This group inhabits in a geographic region that remained isolated during the late Cretaceous and is still isolated by geographic and climatic conditions. Species groups in South American members were more difficult to establish, given their wide diversity. However, some species are unique in having internal pockets in female genitalia and apophyses on the male copulatory bulb. The Chilean species are probably a monophyletic group; they are separated from the rest by the combined presence of external pockets and ridges on the epigastric region of female, whereas males are very similar in the morphology of copulatory bulb, which is very simple, with long embolus and sometimes with additional spineshaped apophysis. This group may have experienced events of isolation during and after the elevation of the Andes range. As in other members of the family the genus is interesting for the presence of secondary sexual characters in males whereas females are practically homogenous in somatic characters. However, female genital morphology is variable, which may suggest a coevolution with somatic characters in males and probably mechanical interactions. Although this work reveals the incredible diversity of this genus, many important geographic regions remain undersampled and records for some countries are entirely lacking.

Description of a new species of the genus Neotrops from Colombia (Araneae: Oonopidae)

A new species of the genus Neotrops (Araneae: Oonopidae) from Colombia, N. ramirezi, is described on the basis of male and female specimens collected in Yotoco Natural Reserve (Valle de Cauca). It is the third species of the genus reported for the country. Una nueva especie del género Neotrops (Araneae: Oonopidae) de Colombia, N. ramirezi se describe en base a especímenes machos y hembras colectados en la Reserva Natural Yotoco (Valle de Cauca). Es la tercera especie del género reportada para el país.

A new orb-weaving spider from the Argentinean flooding pampas grasses: Aculepeira morenoae new species (Araneae, Araneidae)

A new species of the orb-weaving spider genus Aculepeira Chamberling & Ivie 1942, A. morenoae new species, is described and illustrated based on male and female specimens from the Argentinean natural flooding pampas grasses.