Miquel A. Arnedo

Tangled in a sparse spider web: single origin of orb weavers and their spinning work unravelled by denser taxonomic sampling

In order to study the tempo and the mode of spider orb web evolution and diversification, we conducted a phylogenetic analysis using six genetic markers along with a comprehensive taxon sample. The present analyses are the first to recover the monophyly of orb-weaving spiders based solely on DNA sequence data and an extensive taxon sample. We present the first dated orb weaver phylogeny. Our results suggest that orb weavers appeared by the Middle Triassic and underwent a rapid diversification during the end of the Triassic and Early Jurassic. By the second half of the Jurassic, most of the extant orb-weaving families and web designs were already present. The processes that may have given origin to this diversification of lineages and web architectures are discussed. A combination of biotic factors, such as key innovations in web design and silk composition, as well as abiotic environmental changes, may have played important roles in the diversification of orb weavers. Our analyses also show that increased taxon sampling density in both ingroups and outgroups greatly improves phylogenetic accuracy even when extensive data are missing. This effect is particularly important when addition of character data improves gene overlap.

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.

Mitochondrial markers reveal deep population subdivision in the European protected spider Macrothele calpeiana (Walckenaer, 1805) (Araneae, Hexathelidae)

The funnel-web spider genus Macrothele is the only representative of the mygalomorph family Hexathelidae not found in Australia or New Zealand. Its 26 species occur in Central Africa and the Oriental region. Two Macrothele species are found in Europe: M. cretica Kulczynski, 1903 from Crete, and M. calpeiana (Walckenaer, 1805) type species of the genus and the largest European spider, whose distribution extends across the south-eastern Iberian Peninsula, and in two localities of North Africa. Macrothele calpeiana is the only spider protected under European legislation. The fragmentation and destruction of the cork oak forest, with which M. calpeiana was thought to be closely associated, prompted the inclusion of this species in the Bern Convention. Some authors, however, have challenged this view and consider M. calpeiana to be neither a cork oak forest bioindicator nor an endangered species. By contrast, other observations suggest that the distribution of the species is extremely fragmented and that most local populations should be considered as threatened. In this paper, we examine aspects of the conservation status of M. calpeiana in the light of molecular phylogenetic analyses based on mitochondrial markers of sample specimens from major populations. Our data confirm the fragmented distribution of M. calpeiana and reveal high levels of genetic differentiation across its populations. Local population growth cannot be ruled out, though the lineage as a whole has apparently not undergone population growth. Lineage age estimates suggest that M. calpeiana colonized the Iberian Peninsula during the Messinian salinity crisis and that the current population fragmentation originates from the Pliocene and Pleistocene. We argue that the fragmentation and deep genetic divergence across populations, along with evolutionary singularity and endemicity in one of Europe’s main biodiversity hotspots, support the preservation of its legally protected status.

Higher‐level phylogenetics of linyphiid spiders (Araneae, Linyphiidae) based on morphological and molecular evidence

This study infers the higher‐level cladistic relationships of linyphiid spiders from five genes (mitochondrial CO1, 16S; nuclear 28S, 18S, histone H3) and morphological data. In total, the character matrix includes 47 taxa: 35 linyphiids representing the currently used subfamilies of Linyphiidae (Stemonyphantinae, Mynogleninae, Erigoninae, and Linyphiinae (Micronetini plus Linyphiini)) and 12 outgroup species representing nine araneoid families (Pimoidae, Theridiidae, Nesticidae, Synotaxidae, Cyatholipidae, Mysmenidae, Theridiosomatidae, Tetragnathidae, and Araneidae). The morphological characters include those used in recent studies of linyphiid phylogenetics, covering both genitalic and somatic morphology. Different sequence alignments and analytical methods produce different cladistic hypotheses. Lack of congruence among different analyses is, in part, due to the shifting placement of Labulla, Pityohyphantes, Notholepthyphantes, and Pocobletus. Almost all combined analyses agree on the monophyly of linyphioids, Pimoidae, Linyphiidae, Erigoninae, Mynogleninae, as well as Stemonyphantes as a basal lineage within Linyphiidae. Our results suggest independent origins of the desmitracheate tracheal system in micronetines and erigonines, and that erigonines were primitively haplotracheate. Cephalothoracic glandular specializations of erigonines and mynoglenines apparently evolved independently. Subocular sulci of mynoglenines and lateral sulci (e.g. Bathyphantes) evolved independently but glandular pores in the prosoma proliferated once. The contribution of different character partitions and their sensitivity to changes in traditional analytical parameters is explored and quantified.

The dark side of an island radiation: systematics and evolution of troglobitic spiders of the genus Dysdera Latreille (Araneae : Dysderidae) in the Canary Islands

The spider genus Dysdera Latreille is an excellent model for the study of the evolution of cave life: ten species are known to exist exclusively in the subterranean environment of the Canary Islands, where the genus has undergone local diversification. In the present paper, two new troglobitic species (Dysdera madai, sp. nov. and D. sibyllina, sp. nov.) and the previously unknown sex of five additional species are described and illustrated: the males of D. gollumi Ribera & Arnedo, 1994, D. hernandezi Arnedo & Ribera, 1999 and D. labradaensis Wunderlich, 1991; and the females of D. andamanae Arnedo & Ribera, 1997 and D. gibbifera Wunderlich, 1991. The first direct evidence of troglobitic members of Dysdera in micro- and mesocaverns are reported. The evolution of cave life as hypothesised following a combined morphological and molecular phylogeny is investigated. Troglobitic Canarian Dysdera species have colonised the under- ground on eight independent occasions. The Dysderidae groundplan represents a preadaptation to cave life and has facil- itated the colonisation of caves. Canarian members of Dysdera have a predominantly parapatric mode of speciation, although postspeciation changes in distribution may have obscured allopatric processes. Eye regression and, to a lesser extent, larger body size and appendage elongation characterise troglobitic species. The different levels of troglobiomor- phism are interpreted as local adaptations to heterogeneous subterranean conditions. The high levels of sympatry among troglobites are explained by trophic segregation and changes in prey capture strategy were involved in the single identi- fied case of subterranean speciation in the group.

From morphology and karyology to molecules. New methods for taxonomical identification of asexual populations of freshwater planarians. A tribute to Professor Mario Benazzi

Abstract Asexual reproduction by fissiparity is a frequent mode of reproduction in freshwater planarians (Turbellaria, Tricladida, Paludico‐la). Asexual populations pose a main taxonomical problem: to assign them to particular sexual species. We review here two case studies of asexual populations of freshwater triclads belonging to the genera Schmidtea and Dugesia. We show that karyotypes and morphology of the reproductive system of rare ex‐fissiparous specimens led to identifing as Schmidtea mediterranea the few asexual populations of this genus. Instead, morphology and karyology were unable to deal with the frequent asexual populations of Dugesia. We used the internal transcribed spacer region (ITS‐1) of ribosomal DNA to study 20 populations, sexual and asexual, of this genus. The pattern of sequence variation enabled the unambiguous assignment of all asexual populations to particular sexual species. The origin of asexual populations is briefly discussed.

Phylogeny of the parasitic wasp subfamily Charipinae (Hymenoptera, Cynipoidea, Figitidae)

The Charipinae are a major group of hyperparasitoids of Hemiptera. Here, we present the first cladistic analysis of this subfamilys internal relationships, based on 96 morphological characters of adults. The data matrix was analysed using uniformly weighted parsimony. The effects of using alternative weighting schemes were explored by performing additional searches employing implied weights criteria. One of the caveats of implied weights analysis is that it lacks an objective criterion for selecting the value of the concavity function. In the present study, differential weighting was used to explore the sensitivity of our results to the alternative assumptions made in the analysis and to select one of the most parsimonious trees under equal weights, which we regard as being the hypothesis that minimizes the amount of ad hoc assumptions. The validity of the two existing tribes and the monophyly of all the genera of Charipinae were tested, in particular the cosmopolitan and highly species‐rich Alloxysta and Phaenoglyphis, which appear repeatedly in ecological and biochemical studies of host–parasitoid associations. The evolution of several major characters and the relationships between genera are discussed. On the basis of the phylogenetic results, we discuss a number of taxonomic issues. A new classification of the subfamily is proposed in which no tribes are maintained, Carvercharips is synonymyzed with Alloxysta, and the creation of a new genus from Nepal is justified. Our analysis points to the need for a world revision of the basal genus Phaenoglyphis, which is shown as paraphyletic.

Colonization and diversification of the spider genus Pholcus Walckenaer, 1805 (Araneae, Pholcidae) in the Macaronesian archipelagos: evidence for long-term occupancy yet rapid recent speciation.

Macaronesian archipelagos stand apart from other oceanic islands reputed as laboratories for the study of evolution by their proximity to the mainland, lack of subsidence, and steep ecological gradients. The genus Pholcus Walckenaer, 1805, commonly known as daddy-long-leg spiders, is one of the most speciose arthropod groups in the region, with 25 endemic species. In the present study, we use information from four mitochondrial genes, along with morphological data, to examine the phylogenetic relationships and diversification patterns of the genus in the region. Phylogenetic analyses support monophyly of Macaronesian Pholcus including the Moroccan species Pholcus vachoni and hence a single colonization event in the archipelagos. Madeira colonizers most likely originated from the Canaries, and a back-colonization of the nearby mainland receives further support. Estimated lineage divergence times suggest a long-time presence of Pholcus in the region, but also reveal that most present-day species are the result of recent, and probably rapid, speciation events. Diagnostic characters among Macaronesian Pholcus are confined to structures involved in copulation. Coupled with the extremely high diversification rate, the highest recorded for spiders, these copulatory characters suggest that sexual selection has played a key role in the local diversification of Pholcus in Macaronesia.

SYSTEMATICS OF THE GENUS DYSDERA (ARANEAE, DYSDERIDAE) IN THE EASTERN CANARY ISLANDS

Abstract The circum-Mediterranean spider genus Dysdera has undergone an outsanding species radiation in the volcanic archipelago of the Canary Islands. The present study deals with the endemic species that inhabit the older and ecologically distinct islands of Fuerteventura, Lanzarote and their nearby islets. A new species, Dysdera sanborondon, is described. The male of D. spinidorsum Wunderlich 1991, is described for the first time. Five species are redescribed: D. alegranzaensis Wunderlich 1991; D. lancerotensis Simon 1907; D. longa Wunderlich 1991; D. nesiotes Simon 1907, and D. spinidorsum Wunderlich 1991. The species D. liostethus Simon 1907 is proposed to be a senior synonym of D. clavisetae Wunderlich 1991 and its presence in the eastern islands is considered to be doubtful. A neotype is designated for D. nesiotes. The distribution of D. alegranzaensis is extended to Lanzarote and the other northern islets. Dysdera nesiotes is reported for the first time in the eastern Canaries. Morphological affinities and distribution patterns are discussed. The remarkably lower number of endemic species harbored by the eastern islands, when compared with other Canarian islands similar in size but younger in age, is proposed to be the result of a major extinction event in the eastern Canaries due to climatic change.

Living on the edge: demographic and phylogeographical patterns in the woodlouse-hunter spider Dysdera lancerotensis Simon, 1907 on the eastern volcanic ridge of the Canary Islands

The Eastern Canary Islands are the emerged tips of a continuous volcanic ridge running parallel to the northeastern African coast, originated by episodic volcanic eruptions that can be traced back to the Miocene and that, following a major period of quiescence and erosion, continued from the Pliocene to the present day. The islands have been periodically connected by eustatic sea‐level changes resulting from Pleistocene glacial cycles. The ground‐dwelling spider Dysdera lancerotensis Simon, 1907 occurs along the entire ridge, except on recent barren lavas and sand dunes, and is therefore an ideal model for studying the effect of episodic geological processes on terrestrial organisms. Nested clade and population genetic analyses using 39 haplotypes from 605 base pairs of mitochondrial DNA cytochrome c oxidase I sequence data, along with phylogenetic analyses including two additional mitochondrial genes, uncover complex phylogeographical and demographic patterns. Our results indicate that D. lancerotensis colonized the ridge from north to south, in contrast to what had been expected given the SSW‐NNE trend of volcanism and to what had been reported for other terrestrial arthropods. The occurrence of several episodes of extinction, recolonization and expansion are hypothesized for this species, and areas that act as refugia during volcanic cycles are identified. Relaxed molecular clock methods reveal divergence times between main haplotype lineages that suggest an older origin of the northern islets than anticipated based on geological evidence. This study supports the key role of volcanism in shaping the distribution of terrestrial organisms on oceanic islands and generates phylogeographical predictions that warrant further research into other terrestrial endemisms of this fascinating region.