A new pseudoscorpion genus (Garypinoidea: Garypinidae) from the Eocene supports extinction and range contraction in the European paleobiota.

During the Paleogene, the Holarctic experienced drastic climatic oscillations, including periods of extensive glaciation. These changes had a severe impact on both the flora and fauna causing widespread extinction and range shifts with some taxa retreating to refugia in the Mediterranean Basin. Here we provide evidence for this hypothesis using fossils from the pseudoscorpion family Garypinidae Daday, 1889 (Arachnida: Pseudoscorpiones). This family comprises 21 extant genera from all continents except Antarctica but is restricted to low mid-latitudes (<44°N) in the Northern Hemisphere. We provide the second record of garypinids from the European succinite ambers of the Eocene by describing the first extinct genus in Garypinidae, Baltamblyolpium gen. nov., which includes two species: Baltamblyolpium gizmotum sp. nov. from Baltic amber and Baltamblyolpium grabenhorsti sp. nov. from Bitterfeld amber. The new genus exhibits a morphology that closely resembles Neoamblyolpium Hoff, 1956 from western North America and the genus Amblyolpium Simon, 1898, which is widespread but includes taxa restricted to Mediterranean refugia in Europe. The discovery of a new fossil genus of Garypinidae from Europe confirms that the family was found at more northerly latitudes during the Eocene, however, extinction and range contraction resulted in their present-day relictual distribution in southern Europe like many other lineages that once thrived in the European "Baltic amber forest" of the Eocene.

Neglected no longer: Phylogenomic resolution of higher-level relationships in Solifugae.

Advanced sequencing technologies have expedited resolution of higher-level arthropod relationships. Yet, dark branches persist, principally among groups occurring in cryptic habitats. Among chelicerates, Solifugae ("camel spiders") is the last order lacking a higher-level phylogeny and have thus been historically characterized as "neglected [arachnid] cousins". Though renowned for aggression, remarkable running speed, and xeric adaptation, inferring solifuge relationships has been hindered by inaccessibility of diagnostic morphological characters, whereas molecular investigations have been limited to one of 12 recognized families. Our phylogenomic dataset via capture of ultraconserved elements sampling all extant families recovered a well-resolved phylogeny, with two distinct groups of New World taxa nested within a broader Paleotropical radiation. Divergence times using fossil calibrations inferred that Solifugae radiated by the Permian, and most families diverged prior to the Paleogene-Cretaceous extinction, likely driven by continental breakup. We establish Boreosolifugae new suborder uniting five Laurasian families, and Australosolifugae new suborder uniting seven Gondwanan families using morphological and biogeographic signal.

Molecular analyses of pseudoscorpions in a subterranean biodiversity hotspot reveal cryptic diversity and microendemism.

Nested within the Mediterranean biodiversity hotspot, the Dinaric Karst of the western Balkans is one of the world's most heterogeneous subterranean ecosystems and renowned for its highly diverse and mostly endemic fauna. The evolutionary processes leading to both endemism and diversity remain insufficiently understood, and large-scale analyses on taxa that are abundant in both subterranean and surface habitats remain infrequent. Here, we provide the first comprehensive molecular study on Croatian pseudoscorpions, a lineage of arachnids that is common and diverse in both habitats. Phylogenetic reconstructions using 499 COI sequences derived from 128 morphospecies collected across the Dinaric Karst show that: (i) occurrence in karstic microhabitats boosters speciation and endemism in the most diverse genera Chthonius C.L. Koch, 1843 (37 morphospecies) and Neobisium Chamberlin, 1930 (34 morphospecies), (ii) evidence for ongoing diversification is found in many species and species complexes through low optimal thresholds (OTs) and species delineation analyses, and (iii) landscape features, such as mountain ranges, correlate with patterns of genetic diversity in the diverse genus Neobisium. We present two synonymies: Protoneobisium Curcic, 1988 = Neobisium, syn. nov., and Archaeoroncus Curcic and Rada, 2012 = Roncus L. Koch, 1873, syn. nov. Overall, our study suggests that karstic microhabitats promote diversification in soil- and cave-dwelling arthropods at all taxonomic levels, but also provide important refugia for invertebrates in past and present periods of environmental change.

Rainforest conversion to rubber and oil palm reduces abundance, biomass and diversity of canopy spiders.

Rainforest canopies, home to one of the most complex and diverse terrestrial arthropod communities, are threatened by conversion of rainforest into agricultural production systems. However, little is known about how predatory arthropod communities respond to such conversion. To address this, we compared canopy spider (Araneae) communities from lowland rainforest with those from three agricultural systems in Jambi Province, Sumatra, Indonesia, i.e., jungle rubber (rubber agroforest) and monoculture plantations of rubber and oil palm. Using canopy fogging, we collected 10,676 spider specimens belonging to 36 families and 445 morphospecies. The four most abundant families (Salticidae N = 2,043, Oonopidae N = 1,878, Theridiidae N = 1,533 and Clubionidae N = 1,188) together comprised 62.2% of total individuals, while the four most speciose families, Salticidae (S = 87), Theridiidae (S = 83), Araneidae (S = 48) and Thomisidae (S = 39), contained 57.8% of all morphospecies identified. In lowland rainforest, average abundance, biomass and species richness of canopy spiders was at least twice as high as in rubber or oil palm plantations, with jungle rubber showing similar abundances as rainforest, and intermediate biomass and richness. Community composition of spiders was similar in rainforest and jungle rubber, but differed from rubber and oil palm, which also differed from each other. Canonical Correspondence Analysis showed that canopy openness, aboveground tree biomass and tree density together explained 18.2% of the variation in spider communities at family level. On a morphospecies level, vascular plant species richness and tree density significantly affected the community composition but explained only 6.8% of the variance. While abundance, biomass and diversity of spiders declined strongly with the conversion of rainforest into monoculture plantations of rubber and oil palm, we also found that a large proportion of the rainforest spider community can thrive in extensive agroforestry systems such as jungle rubber. Despite being very different from rainforest, the canopy spider communities in rubber and oil palm plantations may still play a vital role in the biological control of canopy herbivore species, thus contributing important ecosystem services. The components of tree and palm canopy structure identified as major determinants of canopy spider communities may aid in decision-making processes toward establishing cash-crop plantation management systems which foster herbivore control by spiders.

Evolutionary morphology of coxal musculature in Pseudoscorpiones (Arachnida).

Pseudoscorpions are an ancient and globally distributed lineage of arachnids with more than 4000 species. Despite being present in virtually all terrestrial habitats, their morphology and anatomy has rarely been studied to date, which hampers homology statements both within and between other arachnid orders. All pseudoscorpions share a morphological peculiarity, the fixation of the coxae of all the walking legs. The same morphological condition is seen in certain other arachnid taxa, such as Solifugae or Scorpiones - potential sistergroups of Pseudoscorpiones. To investigate the musculature apparatus of this unusual feature, we reconstructed the musculature in the coxae of walking legs in three species of pseudoscorpions that represent the three major clades within this order. Using micro-computed tomography (µCT), we show that pseudoscorpions have the highest number of coxal muscles amongst the arachnid orders (12 vs. fewer than 10 in others), and that the muscular composition of the first two legs differs from that in the hind legs, correlating with the difference in function, i.e. pulling in the front legs and pushing in the hind legs. Pseudoscorpions are also unique amongst the arachnids in lacking endoskeletal structures (coxal apodeme or costa coxalis) inside the coxae. We observed that within pseudoscorpions, there is a trend towards a reduction of the number of coxal muscles, with the most basal-branching taxon having the highest number and more derived taxa exhibiting lower counts. We hypothesize the muscular ground pattern for Pseudoscorpiones and discuss the evolution of this system by comparing it to the (scanty) data on other arachnids available in the literature.

Biogeographical and diversification analyses of Indian pseudoscorpions reveal the Western Ghats as museums of ancient biodiversity.

The tropical forests of the Western Ghats (WG) of India are considered 'refugia' harbouring highly diverse and endemic taxa but these refugia are under immense anthropogenic pressure. Most phylogenetic studies have explained diversity patterns across the WG using vertebrates, however, the processes impacting the highly endemic invertebrate fauna are still poorly understood. Here we investigate the evolutionary history of an ancient and widespread arachnid lineage (Pseudoscorpiones: Chthoniidae: Tyrannochthoniini) in the WG through a variety of biogeographical and diversification analyses, including sequence data from three markers for 45 terminals from the WG and 22 from Africa, East and Southeast Asia, Australasia and the Neotropics. Our results show that WG Tyrannochthoniini are rendered paraphyletic by African taxa, a result consistent with continental drift. WG Tyrannochthoniini are further split into two major clades, a southern WG clade and an African-central WG clade, due to vicariance along a major biogeographical barrier, the Palghat Gap. Central WG pseudoscorpions diverged from their African relatives at the Jurassic-Cretaceous boundary, coinciding with the separation of India from Africa. Despite ancient vicariance, six dispersal events occurred across the Palghat Gap, beginning in the Early Cretaceous as India was rafting towards Southeast Asia and ending by the early Miocene, implying that the Palghat Gap became impermeable to dispersal for these humid-adapted organisms during Miocene aridification. Although the ancestor of WG Tyrannochthoniini originated in lowland WG forests, three dispersal events into high montane Shola forests occurred most likely as a result of Late Cretaceous orogenesis and Neogene uplift, allowing lineages to occupy niches at higher elevations. An exponentially declining diversification rate, typical of older lineages, supports the 'museum' model of diversification for WG Tyrannochthoniini. Our study sheds light on the historical biogeography of relictual soil-arthropods in the WG and emphasises the role of WG forest refugia in preserving ancient invertebrate biodiversity.

From the lowlands to the highlands of Ecuador, a study of the genus Masteria (Araneae, Mygalomorphae, Dipluridae) with description of seven new species.

Dipluridae represent a small Mygalomorphae family of South American origin, the family includes two subfamilies Diplurinae and Masteriinae although the placement of the latter in Dipluridae is still under debate. The family has a predominantly South American distribution although the genus Masteria L. Koch, 1873 presents an interesting distribution with representatives found in Fiji, Micronesia, New Caledonia, New Guinea and Australia. This genus is diverse at the species level in tropical South America and the Caribbean but no species have been described from Ecuador to date. Ongoing field work as part of the BIO-GEEC Projecta consortium established by several Ecuadorian and German institutionshas resulted in the discovery of several new species from both lowlands and highland habitats in Ecuador. Herein we described seven new species of Masteria from Ecuador: M. jatunsacha n. sp. (male); M. machay n. sp. (female); M. chalupas n. sp. (male); M. papallacta n. sp. (male and female); M. pasochoa n. sp. (male and female); M. lasdamas n. sp. (male); and M. otongachi n. sp. (male). The type species of the genus, Masteria hirsuta L. Koch, 1873 from Fiji, is redescribed and re-illustrated, from the original type specimen.

Twenty years, eight legs, one concept: describing spider biodiversity in Zootaxa (Arachnida: Araneae).

Zootaxa published more than a thousand papers on Araneae from 2002 to the present, including descriptions of 3,833 new spider species and 177 new genera. Here we summarise the key contributions of Zootaxa to our current knowledge of global spider diversity. We provide a historical account of the researchers that have actively participated as editors, and recognize the more than 1,000 reviewers without whom none of this would have been possible. We conduct a simple analysis of the contributions by authors and geographic region, which allows us to uncover some of the underlying trends in current spider taxonomy. In addition, we examine some of the milestones in twenty years of spider systematic research in Zootaxa. Finally, we discuss future prospects of spider taxonomy and the role that Zootaxa and its younger sister journal Megataxa will play in it. We would like to dedicate this contribution to the memory of Norman I. Platnick, a crucial figure in the advancement of spider systematics.

An annotated type catalogue of the centipedes (Myriapoda: Chilopoda) held in the Zoological Museum Hamburg.

Centipedes (Myriapoda: Chilopoda) are carnivorous arthropods that live in leaf litter and soil habitats on all continents except Antarctica. Roughly 3,300 species have been described to date but the taxonomy is still progressing and requires a thorough documentation of type material and type deposits. In this paper we provide an annotated catalogue of the centipede type collection at the Zoological Museum (ZMH) in Hamburg. This collection comprises 490 type specimens belonging to 141 taxa originally described as species, subspecies and varieties. More than half of these taxa were described by the pioneering myriapodologists Carl Graf Attems (Vienna) and Karl Kraepelin (Hamburg). The collection includes material representing 93 valid species or subspecies and contains specimens predominantly from Australia (33%), Asia (22%), Africa (20%) and South America (20%). Taxonomically, the focus is on the centipede orders Scolopendromorpha (56 valid species) and Geophilomorpha (28 valid species) with fewer types for the Lithobiomorpha (8 species). A total of 48 taxa (34% of type species) have been synonymized since they were originally described and the type material for 12 additional species at the ZMH is lost. We provide a brief historical overview of this collection from its origin at the Museum Godeffroy and the Johanneum in Hamburg, including the most relevant taxonomists and collectors, followed by an annotated taxonomic catalogue with details on all species that are or were represented by type material.

Temperature fluctuations during embryonic development implicated in a naturally occurring instance of abnormal spinnerets in the spider Australomimetus maculosus (Araneae, Mimetidae).

We record developmental abnormalities of the spinnerets in a field-collected adult male specimen of Australomimetus maculosus. These include (1) a supernumerary right posterior lateral spinneret (PLS), (2) ectopic piriform silk gland spigots and tartipores on the left PLS that are normally restricted to anterior lateral spinnerets (ALSs), and (3) what appear to be ectopic ALS sensilla on the left posterior median spinneret (PMS). Published results of teratological experiments and climate data for the collection site indicate that fluctuating sub- and supra-optimal temperatures during embryogenesis may have been responsible for these anomalies. This specimen thus supports the view that spinneret abnormalities, among other aberrations, may be induced when embryos of entelegyne spiders are exposed to fluctuations between high and low temperatures, whether in the laboratory or, as here, in nature. To our knowledge, the ectopic structures seen on the left PLS and left PMS have not been observed previously. Their locations are consistent with a hypothesis by which only the lateral portion of the araneomorph ALS is serially homologous to the PLS, while the remainder of the ALS, along with the colulus/cribellum, is homologous to the PMS.

Hers and his: Silk glands used in egg sac construction by female spiders potentially repurposed by a 'modern' male spider.

Cylindrical silk gland (CY) spigots distinguish a large clade of modern spiders, the CY spigot clade, which includes all entelegyne spiders and their closest relatives. Following a widespread paradigm, CYs and their spigots are only known to occur in female spiders and they produce silk used in the construction of egg sacs. Here we report the occurrence of a CY spigot or CY nubbin on each posterior median spinneret (PMS) in males (5th stadium and later) of the spider Australomimetus maculosus. Late juvenile males had a CY spigot on each PMS, whereas adult males either had a CY spigot or, more often, a non-functional CY nubbin. This indicates that potential CY use by males is at least largely limited to late juvenile instars and is not involved with egg sac construction. Despite the presence of CY spigots in both sexes, sexual dimorphism with respect to CYs was still evident since males lacked the CY spigot on each posterior lateral spinneret present in late juvenile and adult females, and CY spigots of males never had the wide shaft and opening of adult females. This study adds to our knowledge of spinning apparatus variability in modern spiders and demonstrates an exception to the paradigm that, in the CY spigot clade, such spigots are restricted to female spiders.

Functional losses in ground spider communities due to habitat structure degradation under tropical land-use change.

Deforestation and land-use change in tropical regions result in habitat loss and extinction of species that are unable to adapt to the conditions in agricultural landscapes. If the associated loss of functional diversity is not compensated by species colonizing the converted habitats, extinctions might be followed by a reduction or loss of ecosystem functions including biological control. To date, little is known about how land-use change in the tropics alters the functional diversity of invertebrate predators and which key environmental factors may mitigate the decline in functional diversity and predation in litter and soil communities. We applied litter sieving and heat extraction to study ground spider communities and assessed structural characteristics of vegetation and parameters of litter in rainforest and agricultural land-use systems (jungle rubber, rubber, and oil palm monocultures) in a Southeast Asian hotspot of rainforest conversion: Sumatra, Indonesia. We found that (1) spider density, species richness, functional diversity, and community predation (energy flux to spiders) were reduced by 57-98% from rainforest to oil palm monoculture; (2) jungle rubber and rubber monoculture sustained relatively high diversity and predation in ground spiders, but small cryptic spider species strongly declined; (3) high species turnover compensated losses of some functional trait combinations, but did not compensate for the overall loss of functional diversity and predation per unit area; (4) spider diversity was related to habitat structure such as amount of litter, understory density, and understory height, while spider predation was better explained by plant diversity. Management practices that increase habitat-structural complexity and plant diversity such as mulching, reduced weeding, and intercropping monocultures with other plants may contribute to maintaining functional diversity of and predation services provided by ground invertebrate communities in plantations.

Description of four new species of armoured spiders (Araneae, Tetrablemmidae) from Sumatra, Indonesia.

Four new species of armoured spiders from Sumatra, Indonesia are described. Three species are described in the genus Ablemma Roewer, 1963 and one species in the genus Brignoliella Shear, 1978; Ablemmaandriana sp. n. (male), Ablemmacontrita sp. n. (male and female), Ablemmakelinci sp. n. (male) and Brignoliellapatmae sp. n. (male and female). The female of Ablemmasingalang Lehtinen, 1981 is described here for the first time. The first record of Brignoliella for Sumatra is also presented.

Description of three new species of Aposphragisma Thoma, 2014 (Araneae: Oonopidae) from Sumatra, Indonesia.

Three species from the family Oonopidae are newly described from leaf litter habitats in Sumatra, Indonesia based on male and female morphology. All three species belong to the genus Aposphragisma Thoma, 2014: Aposphragismaglobosum sp. n., Aposphragismajambi sp. n., and Aposphragismasumatra sp. n.

An annotated type catalogue of the camel spiders (Arachnida: Solifugae) held in the Zoological Museum Hamburg.

Solifuges are an enigmatic and poorly studied group of arachnids. Commonly referred to as camel spiders or sun spiders, these animals are voracious predators of small animals and found in arid biomes of the Old World and the Americas. In this paper, we provide a catalogue for the solifuges (Arachnida: Solifugae) that are held at the Center of Natural History in Hamburg. The collections in Hamburg are predominantly historical and were accumulated by Karl Kraepelin between 1889 and 1914 with the help of other famous arachnologists such as Ferdinant Karsch and Eugène Simon. The re-study of these collections indicates that there are 38 type species and 65 type specimens from 10 families. We provide a detailed account of this material, including collection data, taxonomic updates, measurements and high-resolution images for species that are either poorly or not at all illustrated. Most specimens (70%) were collected in Africa as part of colonial expeditions or field surveys but there are also types from Western Asia (11%), and North and South America (19%). We provide an overview of the history of this collection, including a summary of the field surveys during which the specimens were collected and the arachnologists who described the material. Overall, this is the third-largest collection of solifuges in Germany with a distinct biogeographical focus and one of the largest collections of camel spiders in Europe.

Four new Mouse Spider species (Araneae, Mygalomorphae, Actinopodidae, Missulena) from Western Australia.

Four new species of the Mouse Spider genus Missulena Walckenaer, 1805 (family Actinopodidae) are described from Western Australia based on morphological features of adult males. Missulena leniae sp. n.(from the Carnarvon and Yalgoo biogeographic regions), Missulena mainae sp. n. (Carnarvon), Missulena melissae sp. n. (Pilbara) and Missulena pinguipes sp. n. (Mallee) represent a broad spectrum of morphological diversity found in this genus and differ from other congeners by details of the male copulatory bulb, colour patterns, eye sizes, leg morphology and leg spination. Two of the species, M. pinguipes sp. n. and M. mainae sp. n., are characterised by swollen metatarsi of the fourth legs in males, a feature not previously recorded in the family. A key to males of all named Missulena species from Australia is presented and allows their identification based on external morphology.

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.

New species of mouse spiders (Araneae: Mygalomorphae: Actinopodidae: Missulena) from the Pilbara region, Western Australia.

Two new species of Mouse Spiders, genus Missulena, from the Pilbara region in Western Australia are described based on morphological features of males. Missulena faulderi sp. nov. and Missulena langlandsi sp. nov. are currently known from a small area in the southern Pilbara only. Mitochondrial cytochrome c oxidase subunit I (COI) sequence divergence failed in clearly delimiting species in Missulena, but provided a useful, independent line of evidence for taxonomic work in addition to morphology.