Rich and abundant spider communities result from enhanced web capture breadth and reduced overlap in urban greenspaces.

Urbanization is a key contributor to biodiversity loss, but evidence is mounting that cities can support rich arthropod communities, including rare and threatened species. Furthermore, greenspace is growing within hundreds of "shrinking cities" that have lost population resulting in a need to demolish an overabundance of infrastructure creating vacant land. Efforts are underway to transform vacant lots, often viewed as blighted areas, into habitats that promote biodiversity and generate ecosystem services, such as urban agroecosystems. To understand how reconfiguring these greenspaces might influence species conservation, elucidation of the factors that drive the distribution of an urban species pool is needed. In particular, the importance of species interactions in structuring urban communities is poorly understood. We tested hypotheses that (1) greater breadth of prey captured by web-building spiders and reduced overlap of prey capture among individuals facilitates the conservation of genera richness and abundance and (2) heterogeneity within a greenspace patch facilitates enhanced dietary niche breadth and greater resource partitioning. In 2013 and 2014, the abundance, breadth and degree of overlap in prey capture of sheet web spiders (Linyphiidae) was measured using web mimic traps at 160 microsites (0.25 m2 ) situated in four urban vacant lots and four urban farms in the city of Cleveland, Ohio, USA. Within a subset of 40 microsites, we used vacuum sampling and hand collection to measure the abundance and genera richness of Linyphiidae. Spider richness and abundance were significantly reduced within urban farms relative to vacant lots. The distribution of spiders and prey was explained by habitat structure, with microsites dominated by tall grasses and flowering plants, with a high bloom abundance and richness, supporting greater prey capture and a higher genera richness and abundance of spiders. In 2014, web capture overlap was significantly greater within microsites dominated by bare ground. These findings illustrate that urban greenspace conservation efforts that focus on reducing bare ground and incorporating a diversity of grasses and flowering plant species can promote linyphiid spiders, potentially by relaxing exploitative competition for shared prey.

Non-pest prey do not disrupt aphid predation by a web-building spider.

A generalist predator's ability to contribute to biological control is influenced by the decisions it makes during foraging. Predators often use flexible foraging tactics, which allows them to pursue specific types of prey at the cost of reducing the likelihood of capturing other types of prey. When a pest insect has low nutritional quality or palatability for a predator, the predator is likely to reject that prey in favour of pursuing alternative, non-pest prey. This is often thought to limit the effectiveness of generalist predators in consuming aphids, which are of low nutritional quality for many generalist predators. Here, we report behavioural assays that test the hypothesis that the generalist predator, Grammonota inornata (Araneae: Linyphiidae), preferentially forages for a non-pest prey with high nutritional quality (springtails), and rejects a pest prey with low nutritional quality (aphids). In no-choice assays, molecular gut-content analysis revealed that spiders continued to feed on the low-quality aphids at high rates, even when high-quality springtails were readily available. When provided a choice between aphids and springtails in two-way choice tests, spiders did not show the expected preference for springtails. Decision-making by spiders during foraging therefore appears to be sub-optimal, possibly because of attraction to the less frequently encountered of two preys as part of a dietary diversification strategy. These results indicate that behavioural preferences alone do not necessarily compromise the pest-suppression capacity of natural enemies: even nutritionally sub-optimal pest prey can potentially be subject to predation and suppression by natural enemies.

Intraguild predation in pioneer predator communities of alpine glacier forelands.

Pioneer communities establishing themselves in the barren terrain in front of glacier forelands consist principally of predator species such as carabid beetles and lycosid spiders. The fact that so many different predators can co-inhabit an area with no apparent primary production was initially explained by allochthonous material deposited in these forelands. However, whether these populations can be sustained on allochthonous material alone is questionable and recent studies point towards this assumption to be flawed. Intraguild predation (IGP) might play an important role in these pioneer predator assemblages, especially in the very early successional stages where other prey is scarce. Here, we investigated IGP between the main predator species and their consumption of Collembola, an important autochthonous alternative prey, within a glacier foreland in the Ötztal (Austrian Alps). Multiplex PCR and stable isotope analysis were used to characterize the trophic niches in an early and late pioneer stage over 2 years. Results showed that intraguild prey was consumed by all invertebrate predators, particularly the larger carabid species. Contrary to our initial hypothesis, the DNA detection frequency of IGP prey was not significantly higher in early than in late pioneer stage, which was corroborated by the stable isotope analysis. Collembola were the most frequently detected prey in all of the predators, and the overall prey DNA detection patterns were consistent between years. Our findings show that IGP appears as a constant in these pioneer predator communities and that it remains unaffected by successional changes.

Immigration of Lygus spp. (Hemiptera: Miridae) and predaceous natural enemies to trap-cropped organic strawberry.

Lygus spp. are polyphagous pests that overwinter in weedy vegetation. In the spring on the central coast of California, Lygus spp. emigrate from weeds into strawberry fields. Subsequent feeding on strawberry flowers causes fruit deformation that precludes sale on the fresh market. Use of alfalfa (Medicago sativa L.) (Fabales: Fabaceae) as a trap crop has been hypothesized to prevent Lygus spp. colonization in strawberries. We examined the movement of Lygus spp. and associated predators from weeds to strawberry fields with alfalfa trap crops using a protein mark-capture technique. Insects and spiders were collected from weeds, strawberry, and alfalfa 1 day, 2 days, and ~2 wk after an albumin protein mark was applied to weeds bordering strawberry fields. For marked Lygus spp. that emigrated from weeds, the majority (79%) of adults were recovered from alfalfa trap crops; however, all nymphs immigrated to strawberry. Most protein-marked predators immigrated to strawberry, rather than trap crops, resulting in a marked predator-to-Lygus spp. ratio of 5:1. Trap cropping effectively reduced the colonization of Lygus adults in strawberry. Converting weedy areas to native perennial plantings could further mitigate the risk of pest migration, while simultaneously conserving beneficial insects.

Dispersal syndrome and landscape fragmentation in the salt-marsh specialist spider Erigone longipalpis.

Dispersal and its evolution play a key role for population persistence in fragmented landscapes where habitat loss and fragmentation increase the cost of between-habitat movements. In such contexts, it is important to know how variation in dispersal and other traits is structured, and whether responses to landscape fragmentation are aligned with underlying dispersal-trait correlations, or dispersal syndromes. We, therefore, studied trait variation in Erigone longipalpis, a European spider species specialist of (often patchy) salt marshes. We collected spiders in two salt-marsh landscapes differing in habitat availability. We then reared lab-born spiders for two generations in controlled conditions, and measured dispersal and its association with various key traits. Erigone longipalpis population densities were lower in the more fragmented landscape. Despite this, we found no evidence of differences in dispersal, or any other trait we studied, between the two landscapes. While a dispersal syndrome was present at the among-individual level (dispersers were more fecund and faster growing, among others), there was no indication it was genetically driven: among-family differences in dispersal were not correlated with differences in other traits. Instead, we showed that the observed phenotypic covariations were mostly due to within-family correlations. We hypothesize that the dispersal syndrome is the result of asymmetric food access among siblings, leading to variation in development rates and carrying over to adult traits. Our results show we need to better understand the sources of dispersal variation and syndromes, especially when dispersal may evolve rapidly in response to environmental change.

Species conservation profiles of the endemic spiders Troglohyphantes (Araneae, Linyphiidae) from the Alps and the north-western Dinarides.

Background: The genus Troglohyphantes Joseph, 1882 (Araneae, Linyphiidae) includes 131 species, mainly distributed across the main European mountain ranges. The Alps and the north-western Dinarides account for 66 species, most of them showing narrow or even point-like distributions. The majority of Troglohyphantes spiders dwell in subterranean habitats including caves, mines, soil litter, rocky debris and other moist and shaded retreats. Despite being intensively studied from taxonomic, ecological and biogeographic standpoints, knowledge on the status of conservation and on the potential risk of extinction of these spiders is lagging. To date, only three species have been included in the global IUCN Red List, but their status has not been updated ever since their last assessment in 1996. The aim of this contribution is to assess the Alpine and north-western Dinaric species of the genus Troglohyphantes and to re-assess the species previously evaluated, according to the last version of the IUCN Red List Categories and Criteria. New information: Amongst the 66 species here considered, 62 had sufficient data to allow the quantification of their Extent Of Occurrence (EOO) and Area Of Occupancy (AOO). Most of the species have a narrow distribution range, with an estimated EOO < 20,000 km2 and AOO < 2,000 km2, meeting the thresholds for the inclusion in the threatened categories. Five species have a more widespread distribution (EOO > 20,000 km2), extending across multiple countries. The quality of the data on distribution of four species was not sufficient to provide a reliable estimation of the distribution range.A continuing decline in EOO, AOO and habitat quality was inferred for 30 species. The majority of them were subterranean specialised species, with a reduced thermal tolerance and a low dispersal ability. Accordingly, changes in subterranean microclimatic conditions due to climate change represent a major threat for these species. Land-use change and habitat alteration were identified as additional relevant threats for several species.A considerable proportion of the species here assessed was found in protected areas and in sites of the Natura 2000 network. In addition, 14 species are formally protected by national and sub-national legislation. At present, 25 species are listed in the regional Red Lists.Long-term monitoring programmes, management plans for both the species and their habitats, expansion of the extant protected areas and designation of new ones, should be considered as the most effective approaches to species conservation.

Unexpected sensitivity of the highly invasive spider Mermessus trilobatus to soil disturbance in grasslands.

The dwarf spider Mermessus trilobatus (Araneae: Linyphiidae), native to North America, has expanded its range over large parts of Europe within less than fifty years. It is notable for occurring in a wide range of mostly agricultural habitats, while most other invasive spiders in Europe are associated with human buildings. As in other invasive invertebrates and plants, the tremendous colonisation success of Mermessus trilobatus might be related to anthropogenic habitat disturbance. Here we aim to test if the invasion success of Mermessus trilobatus in Europe is associated with high tolerance towards soil disturbance. We sampled spiders from eight grasslands experimentally disturbed with superficial soil tillage and eight undisturbed grasslands without tillage. Opposite to our expectation, Mermessus trilobatus densities decrease sharply with soil disturbance. This is in contrast to several native species such as Oedothorax apicatus, which becomes more abundant in the fields after superficial soil tillage. Our study suggests that invasion success of Mermessus trilobatus is not connected to a ruderal strategy. The ecological and evolutionary processes behind colonisation success of Mermessus trilobatus need to be further investigated.

Prey naïveté rather than enemy release dominates the relation of an invasive spider toward a native predator.

Ecosystems may suffer from the impact of invasive species. Thus, understanding the mechanisms contributing to successful invasions is fundamental for limiting the effects of invasive species. Most intuitive, the enemy release hypothesis predicts that invasive species might be more successful in the exotic range than resident sympatric species owing to the absence of coevolution with native enemies. Here, we test the enemy release hypothesis for the invasion of Europe by the North American spider Mermessus trilobatus. We compare the susceptibility of invasive Mermessus trilobatus and a native species with similar life history to a shared predator with which both species commonly co-occur in Europe. Contrary to our expectations, invasive Mermessus trilobatus were consumed three times more frequently by native predators than their native counterparts. Our study shows that invasive Mermessus trilobatus is more sensitive to a dominant native predator than local sympatric species. This suggests that the relation between the invasive spider and its native predator is dominated by prey naïveté rather than enemy release. Further studies investigating evolutionary and ecological processes behind the invasion success of Mermessus trilobatus, including testing natural parasites and rapid reproduction, are needed to explain its invasion success in Europe.

Spatial Distribution of a Tree Trunk Specialist Spider: Relative Role of Landscape Versus Microhabitat Drivers.

By completely censusing a 1 ha forest dynamics plot it was possible to identify the variables (spider mass, size, sex and tree species, size, and bark roughness) that influenced the spatial distribution of adult Drapetisca alteranda Chamberlin 1909 (Araneae: Linyphiidae), a sheet web spider that specializes in lower tree trunks in North American forests. To account for spatial autocorrelation, a conditional autoregressive random effect was included in the zero-inflated Poisson generalized linear mixed model. Parameters estimated were produced by Bayesian inference with vague prior probability distributions and the best of 16 models were selected using Watanabe-Akaike Information Criterion. The best model showed that larger diameter trees located at higher plot elevations were more likely to have D. alteranda present. Smooth bark tree species such as paper birch and American basswood tended to have the most spiders while rough bark species had the least. The relationship between tree diameter and D. alteranda abundance also varied by tree species. Paper birch and quaking aspen tend to produce a greater slope compared to the other species, indicating that as these trees get larger, the abundance of D. alteranda increases at a higher rate than on other tree species. Spider sex and size were not associated with height on the trunk or tree species selection, nor were they associated with microhabitats such as bark furrow depth. Landscape-level factors largely predict D. alteranda abundance and distribution, suggesting that spatial autocorrelation should be considered when modeling the abundance of even small organisms, such as spiders.

Effects of reindeer grazing and recovery after cessation of grazing on the ground-dwelling spider assemblage in Finnish Lapland.

The effect of reindeer Rangifer tarandus L. grazing on the ground-dwelling spider assemblage in Northern Finland was studied. Changes in species richness, abundance and evenness of spider assemblages were analyzed in relation to changes in vegetation and environmental factors in long term grazed and ungrazed sites as well as sites that had recently switched from grazed to ungrazed and vice versa. Grazing was found to have a significant impact on height and biomass of lichens and other ground vegetation. However, it seemed not to have an impact on the total abundance of spiders. This is likely caused by opposing family and species level responses of spiders to the grazing regime. Lycosid numbers were highest in grazed and linyphiid numbers in ungrazed areas. Lycosidae species richness was highest in ungrazed areas whereas Linyphiidae richness showed no response to grazing. Four Linyphiidae, one Thomisidae and one Lycosidae species showed strong preference for specific treatments. Sites that had recovered from grazing for nine years and the sites that were grazed for the last nine years but were previously ungrazed resembled the long term grazed sites. The results emphasize the importance of reindeer as a modifier of boreal forest ecosystems but the impact of reindeer grazing on spiders seems to be family and species specific. The sites with reversed grazing treatment demonstrate that recovery from strong grazing pressure at these high latitudes is a slow process whereas reindeer can rapidly change the conditions in previously ungrazed sites similar to long term heavily grazed conditions.

A new species of the spider genus Solenysa from China (Araneae, Linyphiidae).

The genus Solenysa Simon, 1894 belongs to Linyphiidae Blackwall, 1859, which is a species-rich group, including 608 genera and 4,571 species (World Spider Catalog 2018). Solenysa currently includes 14 species from China, Japan and the Korean Peninsula (Simon 1894; Namkung 1986; Li Song 1992; Gao, Zhu Sha 1993; Tu, Ono Li 2007; Ono 2011; Tu Hormiga 2011; Wang, Ono Tu 2015). The linyphiid phylogeny based on molecular data shows that Solenysa species forms one of the seven main clades within Linyphiidae (Wang et al. 2015). According to the phylogenetic analysis based on morphological data, Tu and Hormiga (2011) divided the genus Solenysa into four species groups, each having a unique genital type comprised by series genital characters. As an old branch with a long evolutionary history, Solenysa spiders have accumulated a long list of synapomorphies (Tu Hormiga 2011), not only having a unique somatic appearance, but also specific genitalic characters that distinguish them from all other linyphiids.

First description of the male of the linyphiid spider Disembolus corneliae (Chamberlin Ivie, 1944) (Araneae, Linyphiidae).

The erigonine linyphiid genus Disembolus currently contains 24 described species (World Spider Catalog 2018). Disembolus was erected by Chamberlin and Ivie (1933) to accommodate their new species D. stridulans Chamberlin and Ivie from Utah. Since then, 23 species have been added to the genus. Chamberlin (1949) described D. zygethus Chamberlin, but the majority of species were added to the genus by Millidge (1981). Millidge (1981) described 16 species from museum specimens at the American Museum of Natural History and transferred five species from the genera Tapinocyba and Soudinus (D. alpha (Chamberlin 1949), D. kesimbus (Chamberlin 1949), D. phanus (Chamberlin 1949), D. sacerdotalis (Crosby Bishop 1933), and D. corneliae (Chamberlin Ivie 1944)). The last species added to the genus was D. bairdi Edwards described by Edwards in 1999. Disembolus corneliae Chamberlin and Ivie was described from the female only by Chamberlin and Ivie (1944) as Soudinus corneliae. The species was then transferred to Disembolus by Millidge (1981). Herein, I describe the male of the species for the first time.

A new species and new records of the spider genus Taranucnus (Araneae, Linyphiidae) from the Ukrainian Carpathians.

Taranucnus Simon, 1884 is a small Holarctic genus of the spider family Linyphiidae that contains only five species (WSC, 2018). In Europe and in Ukraine it is represented by three taxa: T. setosus (O. P.-Cambridge, 1863), T. bihari Fage, 1931 and T. carpaticus Gnelitsa, 2016 (WSC, 2018). The type species of the genus is T. setosus, widely distributed in the Palaearctic, T. bihari is known from a few localities from the Carpathians within Poland, Slovakia, and Romania (Nentwig et al. 2017). Recently, T. carpaticus was recorded from the two mountain massifs (Gorgany and Chornohora) of the Ukrainian Carpathians (Gnelitsa 2016). In this paper a fourth European species, Taranucnus beskidicus new species, is described.

A review of some Neriene spiders (Araneae, Linyphiidae) from China.

The genus Neriene Blackwall, 1833 from China is reviewed. Two new species are described: Neriene lushanensis n. sp., Neriene orthocera n. sp. Ketambea liupanensis (Tang Song, 1992) n. comb. and Ketambea nigripectoris (Oi, 1960) n. comb. are transferred from the genus Neriene. Nineteen known species are redescribed or diagnosed and discriminated from related species: Neriene aquilirostralis Chen Zhu, 1989, Neriene birmanica (Thorell, 1887), Neriene calozonata Chen Zhu, 1989, Neriene cavaleriei (Schenkel, 1963), Neriene clathrata (Sundevall, 1830), Neriene compta Zhu Sha, 1986, Neriene decormaculata Chen Zhu, 1988, Neriene emphana (Walcknear, 1841), Neriene hammeni (Van Helsdingen, 1963), Neriene japonica (Oi, 1960), Neriene limbatinella (Bosenberg Strand, 1906), Neriene longipedella (Bosenberg Strand, 1960), Neriene macella (Thorell, 1898), Neriene nitens Chen Zhu, 1991, Neriene oidedicata (Van Helsdingen, 1969), Neriene poculiforma Liu Chen, 2010, Neriene radiata (Walckenear, 1841), Neriene strandia (Blauvelt, 1936), and Neriene zhui Chen Li, 1995.

A DNA barcode-assisted annotated checklist of the spider (Arachnida, Araneae) communities associated to white oak woodlands in Spanish National Parks.

BACKGROUND: A large scale semi-quantitative biodiversity assessment was conducted in white oak woodlands in areas included in the Spanish Network of National Parks, as part of a project aimed at revealing biogeographic patterns and identify biodiversity drivers. The semi-quantitative COBRA sampling protocol was conducted in sixteen 1-ha plots across six national parks using a nested design. All adult specimens were identified to species level based on morphology. Uncertain delimitations and identifications due to either limited information of diagnostic characters or conflicting taxonomy were further investigated using DNA barcode information. NEW INFORMATION: We identified 376 species belonging to 190 genera in 39 families, from the 8,521 adults found amongst the 20,539 collected specimens. Faunistic results include the discovery of 7 new species to the Iberian Peninsula, 3 new species to Spain and 11 putative new species to science. As largely expected by environmental features, the southern parks showed a higher proportion of Iberian and Mediterranean species than the northern parks, where the Palearctic elements were largely dominant. The analysis of approximately 3,200 DNA barcodes generated in the present study, corroborated and provided finer resolution to the morphologically based delimitation and identification of specimens in some taxonomically challenging families. Specifically, molecular data confirmed putative new species with diagnosable morphology, identified overlooked lineages that may constitute new species, confirmed assignment of specimens of unknown sexes to species and identified cases of misidentifications and phenotypic polymorphisms.

Diplocephalus komposchi n. sp., a new species of erigonine spider (Araneae, Linyphiidae) from Austria.

The erigonine cladistic analyses of Hormiga (2000) and Miller & Hormiga (2004) demonstrated unambiguous support for a sister-taxon relationship between the genera Diplocephalus and Savignia. These genera, in addition to others, are commonly placed in the Savignia-group. Although the Savignia-group is not monophyletic as it was originally circumscribed by Millidge (1977), it contains a monophyletic core of genera that has been supported in various cladistic analyses, starting with Hormiga (2000). According to the most recent phylogenetic study (Frick et al. 2010), a clade within the Savignia-group included Diplocephalus along with Araeoncus, Dicymbium, Erigonella, Glyphesis and Savignia. Frick et al. (2010) included three Diplocephalus species - cristatus, latifrons and picinus - in their cladistic analyses. While D. latifrons and D. picinus were found to be the most basal species of the Savignia-group, D. cristatus was the most distal one.

Do seasonal changes in numbers of aerially dispersing spiders reflect population density on the ground or variation in ballooning motivation?

Groups of linyphiid spiders (Erigone spp.) (Araneae, Linyphiidae), collected at intervals from arable land, were tested in laboratory bioassays to determine the proportion of individuals that exhibited ballooning behaviour on each field sampling occasion. There was no significant variation in the proportions of spiders in each test group ballooning in the laboratory over a year. Investigations of ground density and aerial dispersal, in a grass field and a winter-wheat field, confirmed that peaks in numbers of spiders observed ballooning in the field generally coincided with population peaks in the summer and autumn. Significant correlations between ground populations and aerial catches were found for total spiders, immature spiders, and Bathyphantes gracilis (Blackwall) in both fields, adult spiders in the grass field, Erigone spp. in the grass, and Meioneta rurestris (C.L. Koch) in the wheat. The other groups analysed, Lepthyphantes tenuis (Blackwall) in both fields, Erigone spp. in wheat, and M. rurestris in grass, showed similar but non-significant trends. No significant difference was found between overall ground-to-air ratios for males compared to females, but adult spiders were more likely to balloon than immatures.

The effect of food supply on the colonisation of barley by aerially dispersing spiders (Araneae).

A field experiment was carried out to determine whether different levels of food availability affected the retention rate of ballooning spiders landing in trays of seedling barley plants, half of which were infested with aphids from laboratory cultures. The trays were placed within bases in the field, then collected sequentially and spider numbers assessed in each tray. Deposition trays, containing trapping fluid only, were used to measure ballooning activity throughout the experimental period. The experiment was repeated four times. Overall, ballooning spiders were more likely to be retained in trays where aphid prey were present, with a total of 340 spiders found in the infested trays and 251 in the aphid-free trays, over the four experiments. Most of the spiders found were of the family Linyphiidae. In the second and fourth experiments the increased retention of spiders in the aphid-infested trays was statistically significant. Immature linyphiids alone also showed significantly higher retention in the infested trays in those two experiments and in the fourth experiment were largely responsible for the higher numbers found in the infested trays. There was also a statistically significant trend for a higher retention rate of female spiders, compared to males, in the barley trays than would have been expected from the ratios of females to males caught in the deposition trapping trays alone.

Ambengana Millidge & Russell-Smith, 1992, a synonym of Neriene Blackwall, 1833 (Araneae, Linyphiidae).

The taxonomic status of the genus Ambengana Millidge & Russell-Smith, 1992, is revised on the basis of its original description, illustrations and re-examination of the type species. A new synonymy is proposed: Ambengana complexipalpis Millidge & Russell-Smith, 1992 (the type species of Ambengana) syn. n. with Neriene birmanica (Thorell, 1887). Therefore,the genus Ambengana Millidge & Russell-Smith, 1992 syn. n. is synonymized with Neriene Blackwall, 1833. A morphological re-description, diagnosis and comparative illustrations are provided for Neriene birmanica as well.