Can ant‐eating Zodarion spiders (Araneae: Zodariidae) develop on a diet optimal for euryphagous arthropod predators?

Abstract Little attention is paid to the behavioural and physiological adaptations of ant‐eating predators. It is expected that there should be a strong selection for traits related to prey handling, leading to the evolution of morphological, behavioural and nutritional adaptations. Such adaptations may then entail trade‐offs in handling and utilization of alternative prey. To investigate behavioural as well as nutritional adaptations and the occurrence of the corresponding trade‐offs in two ant‐eating spiders of the genus Zodarion [Zodarion atlanticum Pekár & Cardoso and Zodarion germanicum (C. L. Koch)], spiders are reared on two diets: ants (i.e. their preferred prey) and fruit flies (i.e. an alternative prey that is nutritionally optimal for euryphagous spiders). Food consumption is observed and several fitness‐related life‐history parameters are measured. Although spiders readily accept ants, more than one‐third of 35 spiders refuse to consume fruit flies and starve. Furthermore, severe hunger does not induce these individuals to accept fruit flies. Starving spiders die before moulting to the second stadium. Spiders that eat fruit flies increase only little and slowly in weight, and all of these die during the first two stadia. By contrast, spiders on an ant diet increase dramatically in weight, and develop up to the fourth stadium. These data indicate that fruit flies are not suitable for Zodarion, supporting the hypothesis that there are behavioural and nutritional trade‐offs. Taking into account the results of previous studies, it is suggested that nutritional trade‐offs are generally important for stenophagous spiders.     

Differences in venom composition between orb-weaving and wandering Hawaiian Tetragnatha (Araneae)

Spider venoms are complex mixtures of toxins that are used primarily for immobilizing prey. There is evidence of chemical variation in spider venoms among close relatives, yet few studies have analysed their evolution within an ecological and phylogenetic framework. On the Hawaiian archipelago, Tetragnatha, a cosmopolitan orb-weaving genus, has undergone a radiation in which a monophyletic lineage has abandoned web-building and become obligately wandering foragers. This study compares venom composition and details of feeding behaviour between orb-weaving and wandering Hawaiian Tetragnatha. Protein gel electrophoresis patterns indicated that relative to orb-weavers, wandering species had a reduced concentration of low molecular weight (<14kDa) components. Both orb-weaving and wandering Tetragnatha captured flying prey (adult lepidopterans, dipterans), but wandering spiders also captured non-flying prey (insect larvae, spiders). There were no distinct differences between orb-weavers and wanderers in prey capture and immobilization sequences, or in the paralytic effects of bites on prey. However, prey bitten by wanderers took longer to be permanently immobilized than prey bitten by orb-weavers. Contrary to predictions, there was no indication that web-loss in this group was associated with an increase in venom potency.     

Trophic specialisation in a predatory group: the case of prey‐specialised spiders (Araneae)

Predators appear to be less frequently specialised (i.e. adapted to restricted diet) on their prey than herbivores, parasites or parasitoids. Here, we critically evaluate contemporary evolutionary hypotheses that might be used to explain the evolution of specialised foraging in predators. We propose a unifying concept within which we define four types of trophic categories using ecological (diet breadth) and evolutionary (degree of adaptations) contexts. We use data on spiders (Araneae), the most diversified order of terrestrial predators, to assess applicability of frameworks and evolutionary concepts related to trophic specialisation. The majority of spider species are euryphagous but a few have a restricted prey range, i.e. they are stenophagous. We provide a detailed overview of specialisation on different prey types, namely spiders, crustaceans, moths, dipterans, ants, and termites. We also review the available evidence for trophic adaptations, classified into four categories: behavioural, morphological, venomic and metabolic. Finally, we discuss the ecological and evolutionary implications of trophic specialisation and propose avenues for future research.     

Isotopic niche differs between seal and fish‐eating killer whales (Orcinus orca) in northern Norway

Ecological diversity has been reported for killer whales (Orcinus orca) throughout the North Atlantic but patterns of prey specialization have remained poorly understood. We quantify interindividual dietary variations in killer whales (n = 38) sampled throughout the year in 2017–2018 in northern Norway using stable isotopic nitrogen (δ¹⁵N: ¹⁵N/¹⁴N) and carbon (δ¹³C: ¹³C/¹²C) ratios. A Gaussian mixture model assigned sampled individuals to three differentiated clusters, characterized by disparate nonoverlapping isotopic niches, that were consistent with predatory field observations: seal‐eaters, herring‐eaters, and lumpfish‐eaters. Seal‐eaters showed higher δ¹⁵N values (mean ± SD: 12.6 ± 0.3‰, range = 12.3–13.2‰, n = 10) compared to herring‐eaters (mean ± SD: 11.7 ± 0.2‰, range = 11.4–11.9‰, n = 19) and lumpfish‐eaters (mean ± SD: 11.6 ± 0.2‰, range = 11.3–11.9, n = 9). Elevated δ¹⁵N values for seal‐eaters, regardless of sampling season, confirmed feeding at high trophic levels throughout the year. However, a wide isotopic niche and low measured δ¹⁵N values in the seal‐eaters, compared to that of whales that would eat solely seals (δ_N‐measured = 12.6 vs. δ_N‐expected = 15.5), indicated a diverse diet that includes both fish and mammal prey. A narrow niche for killer whales sampled at herring and lumpfish seasonal grounds supported seasonal prey specialization reflective of local peaks in prey abundance for the two fish‐eating groups. Our results, thus, show differences in prey specialization within this killer whale population in Norway and that the episodic observations of killer whales feeding on prey other than fish are a consistent behavior, as reflected in different isotopic niches between seal and fish‐eating individuals.     

Trophic niche overlap between round sardinella (Sardinella aurita) and sympatric pelagic fish species in the Western Mediterranean

The northward expansion of round sardinella (Sardinella aurita) in the Mediterranean Sea, together with declines and fluctuations in biomass and landings of European sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus) observed in recent decades, may suggest potential inter‐specific competition in the pelagic domain. The coexistence of sympatric zooplanktivorous fish species might therefore be exposed in part to trophic niche overlap and competition for food. Combining visual diet characterization under the microscope with DNA metabarcoding from stomach contents of fish collected in spring results show that predation on relatively large krill is equally important for sardinella than for the other two niche overlapping species. Furthermore, an important overlap is found in their isotopic niche, especially with anchovy, using nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotopes in muscle tissue. In fact, the three fish species are able to feed effectively in the whole prey size spectrum available during the sampled season, from the smallest diatoms and copepods to the larger prey (i.e., decapods and euphausiids), including fish larvae. Moreover, effective predation upon other large prey like siphonophores, which is observed only when multi‐proxy analyses in stomach contents are applied, might also be relevant in the diet of sardinella. The overlapping diet composition in spring, together with the effective use of food resource by sardinella, can be of special interest in potential future scenarios with warmer water temperature leading to lower zooplankton and/or higher jellyfish availability, where sardinella may take advantage over other species due to its feeding plasticity.     

Two new Mallinella species from southern China (Araneae,Zodariidae)

Two new species of the spider genus Mallinella Strand, 1906 are reported from China: Mallinella sphaerica sp. n. (male, female) from Tianmu Mountain, Zhejiang Province and Mallinella pluma sp. n. (male) from Daming Mountain, Guangxi Zhuang Autonomous Region.     

The rove beetle Drusilla sparsa (Coleoptera: Staphylinidae) is a myrmecophilous species associated with a myrmicine ant,Crematogaster osakensis (Hymenoptera: Formicidae)

The rove beetle genus Drusilla includes some myrmecophilous species. The Japanese species Drusilla sparsa (Sharp, 1874) has been regarded as a non‐myrmecophilous beetle. In Kagawa Prefecture, Shikoku Island, western Japan, however, we often observed that D. sparsa adults were walking in the vicinity of foraging workers of the myrmicine ant Crematogaster osakensis Forel, 1990. The body color of the beetle is similar to C. osakensis as in other myrmecophilous beetles found near the trails of the host ants. To examine whether D. sparsa is myrmecophilous, we investigated the distribution of D. sparsa and C. osakensis in the field, as well as their behavior including prey preference of the beetle in the laboratory. Drusilla sparsa beetles were collected only in sites where C. osakensis ants occurred. When the beetles encountered the ant workers, they bent the abdominal tip toward the ants. The ants licked the abdominal tip, and then the beetles usually walked away. Such behavioral reaction of the ants was not observed when the beetles encountered workers of the formicine ant Nylanderia flavipes (Smith, 1874) that continuously attacked the beetles. Drusilla sparsa preferred to feed on dead workers of C. osakensis even when other ants were available as food, indicating that D. sparsa is a myrmecophilous species associated with C. osakensis. Crematogaster osakensis was frequently found in the stomach in the ant predator, the Japanese treefrog Hyla japonica Günther, 1859. Thus, the significance of body color similarity between the host ants and beetles is not a case of Batesian mimicry.     

Evolution of stenophagy in spiders (Araneae): evidence based on the comparative analysis of spider diets

Stenophagy (narrow diet breadth) represents an extreme of trophic specialization in carnivores, but little is known about the forces driving its evolution. We used spiders, the most diversified group of terrestrial predators, to investigate whether stenophagy (1) promoted diversification; (2) was phylogenetically conserved and evolutionarily derived state; and (3) was determined either by geographical distribution and foraging guild. We used published data on the prey of almost 600 species. Six categories of stenophagy were found: myrmecophagy, araneophagy, lepidopterophagy, termitophagy, dipterophagy, and crustaceophagy. We found that the species diversity of euryphagous genera and families was similar to stenophagous genera and families. At the family level, stenophagy evolved repeatedly and independently. Within families, the basal condition was oligophagy or euryphagy. Most types of stenophagy were clearly derived: myrmecophagy in Zodariidae; lepidopterophagy in Araneidae; dipterophagy in Theridiidae. In contrast, araneophagy was confined to basal and intermediate lineages, suggesting its ancestral condition. The diet breadth of species from the tropics and subtropics was less diverse than species from the temperate zone. Diet breadth was lower in cursorial spiders compared to web-building species. Thus, the evolution of stenophagy in spiders appears to be complex and governed by phylogeny as well as by ecological determinants.     

Habitat preferences of the urban wall spider Oecobius navus (Araneae, Oecobiidae)

Abstract  To assess the relative impact of a range of habitat variables on spider abundance, field and laboratory experiments were conducted on populations of the urban wall spider Oecobius navus in suburban Perth, Western Australia. Habitat characteristics investigated were: substrate type, wind speed, rainfall, sunlight exposure, relative humidity, air temperature, substrate temperature, artificial lighting and prey type/abundance. In the field, O. navus was found to be associated with high humidity, low air temperature and shelter from sunlight and rainfall. Oecobius navus was more abundant at sites with greater prey abundance. The most common prey item was the red meat ant Iridomyrmex chasei . Juvenile spiders were more abundant than adult spiders; however, patterns between spider abundance and habitat variables were similar for both adults and juveniles. Laboratory experiments showed that O. navus preferred to build webs on wooden substrates, and pitted limestone walls. These findings indicate that O. navus may be vulnerable to desiccation and/or thermal stress, and thus survives better on sheltered walls.     

Niche partitioning and niche filtering jointly mediate the coexistence of three closely related spider species (Araneae,Philodromidae)

1. Several mechanisms can mediate the coexistence of species, such as the neutral dynamic, niche filtering and niche partitioning. The present study investigated which of these mechanisms mediate the coexistence of closely related spider species at the scale of one locality. 2. The coexistence of three spider species of the genus Philodromus (Philodromidae) (Philodromus albidus, Philodromus aureolus and Philodromus cespitum) was studied. The study area comprised three habitat types, including a deciduous forest, a scrub and a plum tree stand. The spatial niche properties of the Philodromus species were explored by comparing their micro/macrohabitat preferences. The natural diet of these species was analysed. Laboratory experiments involving prey acceptance were conducted to investigate the trophic niche properties. The species' phenologies were studied to compare their temporal niches. 3. The philodromids had differentiated their trophic and habitat niches. The coexistence mechanisms were therefore assessed by studying the relationships between niche overlaps and the pairwise product of relative abundances. A negative relationship was observed between functional niche overlaps and the product of relative abundances, whereas a positive relationship was observed between spatial niche overlaps and the product of relative abundances. 4. The present results suggest that different mechanisms influence different niche dimensions. Niche partitioning influenced functional niches, whereas niche filtering influenced spatial niches. The results also suggest that the filtering process in one dimension was facilitated by niche differentiation in another dimension.     

Neuropharmacological profile of FrPbAII, purified from the venom of the social spider Parawixia bistriata (Araneae, Araneidae), in Wistar rats

The aims of the present study were to investigate the anticonvulsant activity and behavioral toxicity of FrPbAII using freely moving Wistar rats. Moreover, the effectiveness of this compound against chemical convulsants was compared to that of the inhibitor of the GABAergic uptake, nipecotic acid. Our results show that FrPbAII was effective against seizures induced by the i.c.v. injection of pilocarpine (ED(50) = 0.05 microg/animal), picrotoxin (ED(50) = 0.02 microg/animal), kainic acid (ED(50) = 0.2 microg/animal) and the systemic administration of PTZ (ED(50) = 0.03 microg/animal). The anticonvulsant effect of FrPbAII differed from that of nipecotic acid in potency, as the doses needed to block the seizures were more than 10 folds lower. Toxicity assays revealed that in the rotarod, the toxic dose of the FrPbAII is 1.33 microg/animal, and the therapeutic indexes were calculated for each convulsant. Furthermore, the spontaneous locomotor activity of treated animals was not altered when compared to control animals but differed from the animals treated with nipecotic acid. Still, FrPbAII did not induce changes in any of the behavioral parameters analyzed. Finally, when tested for cognitive impairments in the Morris water maze, the i.c.v. injection of FrPbAII did not alter escape latencies of treated animals. These findings indicate that the novel GABA uptake inhibitor is a potent anticonvulsant with mild side-effects when administered to Wistar rats.     

Natural history of the non-solitary sheetweaving spider Anelosimus jucundus (Araneae: Theridiidae)

Field observations were made on one Panamanian population of Anelosimus jucundus. Almost all webs were found in a relatively exposed area, on small shrubs and composites. This is in contrast to A. eximius , a colonial congeneric which in our study site built its large webs along moist, fern-covered banks. Web structure varied with different plant substrates; for example, only some of the webs incorporated a clearly defined lower sheet. Webs were built by penultimate instar females, and perhaps by adult females and males. In some cases, several adult females inhabited a web, but they remained under separate leaf canopies; during census observations co-operative efforts were not noted. Prey consisted mainly of winged insects. Juvenile sex ratio was about equal. The degree of sociality demonstrated by A. jucundus is discussed.     

Sensory structures involved in prey detection on the labial palp of the ant‐hunting beetle Siagona europaea Dejean 1826 (Coleoptera,Carabidae)

Giglio, A., Ferrero E.A., Perrotta, E., Talarico, F.F. and Zetto Brandmayr, T. 2010. Sensory structures involved in prey detection on the labial palp of the ant‐hunting beetle Siagona europaea Dejean 1826 (Coleoptera, Carabidae). —Acta Zoologica (Stockholm) 91 : 328–334 The ultrastructure and distribution of sensilla on the labial palps of a myrmecophagous carabid beetle, Siagona europaea, were investigated using scanning and transmission electron microscopy techniques. Five types of sensilla were identified: three types of sensilla basiconica on the apical sensory area and two types, one sensillum trichodeum and one coeloconicum, on the external palp surface. On morphological grounds, the s. basiconica type 1 were considered as olfactory, the type 2 as gustatory, the type 3 and the s. trichodeum as mechanoreceptive, and the s. coeloconicum as a thermo/hygroreceptor. Their function is discussed in relation to prey detection and habitat adaptations.     

Deciphering the diet of a wandering spider (Phoneutria boliviensis; Araneae: Ctenidae) by DNA metabarcoding of gut contents

Arachnids are the most abundant land predators. Despite the importance of their functional roles as predators and the necessity to understand their diet for conservation, the trophic ecology of many arachnid species has not been sufficiently studied. In the case of the wandering spider, Phoneutria boliviensis F. O. Pickard‐Cambridge, 1897, only field and laboratory observational studies on their diet exist. By using a DNA metabarcoding approach, we compared the prey found in the gut content of males and females from three distant Colombian populations of P. boliviensis. By DNA metabarcoding of the cytochrome c oxidase subunit I (COI), we detected and identified 234 prey items (individual captured by the spider) belonging to 96 operational taxonomic units (OTUs), as prey for this wandering predator. Our results broaden the known diet of P. boliviensis with at least 75 prey taxa not previously registered in fieldwork or laboratory experimental trials. These results suggest that P. boliviensis feeds predominantly on invertebrates (Diptera, Lepidoptera, Coleoptera, and Orthoptera) and opportunistically on small squamates. Intersex and interpopulation differences were also observed. Assuming that prey preference does not vary between populations, these differences are likely associated with a higher local prey availability. Finally, we suggest that DNA metabarcoding can be used for evaluating subtle differences in the diet of distinct populations of P. boliviensis, particularly when predation records in the field cannot be established or quantified using direct observation.     

Phylogenetic analysis and revision of the linyphiid spider genus Solenysa (Araneae: Linyphiidae: Erigoninae)

In this paper we carry out a taxonomic revision and phylogenetic analysis of the linyphiid spider genus Solenysa Simon, 1894. A total of 12 species is treated here, including five new species collected from China and Japan: Solenysa akihisai Tu sp. nov., Solenysa lanyuensis Tu sp. nov., Solenysa retractilis Tu sp. nov., Solenysa tianmushana Tu sp. nov. , and Solenysa yangmingshana Tu sp. nov. Solenysa circularis Gao, Zhu & Sha, 1993 is a junior synonym of Solenysa protrudens Gao, Zhu & Sha, 1993. We have assembled two different character matrices to reconstruct the phylogenetic relationships of Solenysa. In the first matrix (Matrix 1), five representative species of Solenysa were added to the morphological dataset of Miller & Hormiga to test the monophyly of the genus and its placement within Linyphiidae. The genitalic structures and somatic morphology of Solenysa were studied by means of scanning electron microscopy for the first time. To infer the species‐level phylogenetic relationships of Solenysa we produced a second matrix (Matrix 2) that includes all 12 Solenysa species and six outgroup species chosen from the results of the analysis of the first matrix. The two most parsimonious trees from the analysis of Matrix 1 support the monophyly of Solenysa and its placement within the ‘Distal Erigonines’ clade. The single most parsimonious tree resulting from the analysis of the second matrix suggests that the Solenysa clade includes four monophyletic groups, each group represented by a distinct genitalic pattern. The morphology of Solenysa, both somatic and genitalic, is highly autapomorphic. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 484–530.     

Phylogenetic analysis of the tropical wolf spider subfamily Cteninae (Arachnida,Araneae, Ctenidae)

A cladistic analysis based on parsimony was performed to test the monophyly of Cteninae. The data matrix comprises 72 terminal taxa scored for 88 characters from external morphology, female internal genitalia, and two behavioural characters. Cteninae, as currently delimited, is polyphyletic. The monoplyly of Cteninae requires the transfer of two Ctenus species to Enoploctenus (Acantheinae) and two Ctenus species to Asthenoctenus (Viridasiinae). Ctenus, the genus type, comprises approximately half of the described species of the family, and is polyphyletic as currently delimited. From the 32 Ctenus species added to this analysis, only 13 are recognized as Ctenus s.s.: (C. rectipes (C. pauloterrai (C. manauara, C. nigritus ((C. dubius, C. crulsi) ((C. amphora, C. minor) (C. medius, C. paubrasil (C. fernandae (C. ornatus, C. vehemens)))))))). The results suggest that at least eight new genera should be described to accommodate species misplaced in Ctenus. The result also refutes the monophyly of Amauropelma Raven, Stumkat & Gray, 2001, Celaetycheus Simon, 1897, Leptoctenus L. Koch, 1878, and Thoriosa Simon, 1910. The following taxonomic changes are proposed: Ctenus inazensis Strand, 1909 and Ctenus miserabilis Strand, 1916 are transferred to Enoploctenus; Ctenus tarsalis F.O. Pickard‐Cambridge, 1902 and Ctenus bulimus Strand, 1909 are transferred to Asthenoctenus Simon, 1897; Celaetycheus modestus Bryant, 1942 is transferred to Ohvida Polotow & Brescovit, 2009. © 2014 The Linnean Society of London     

Constraints affecting partial prey consumption by a crab spider,Diaea sp. indet. (Araneae: Thomisidae)

Summary The influence of feeding constraints on the feeding behaviour of Diaea sp. indet., was investigated. Diaea is a crab spider which ambushes its prey and practises extraintestinal digestion. A laboratory study was carried out using fruit flies, Drosophila immigrans, as prey. Diaea feeds from two sites on the prey — initially the head, followed by the posterior abdomen, with most of the prey's contents being extracted from the head. If additional prey are available, Diaea will, instead of switching to the posterior abdomen, catch a new prey item. The efficiency with which Diaea can extract food is influenced by changes which occur in the prey as a consequence of it being killed and fed on. Evaporative fluid loss from prey is an important constraint on food uptake because in influences the viscosity of the prey's contents. Regardless of whether a new prey item arrives, Diaea discards the prey item on which it is feeding before all of the available food has been extracted from it. The fluid content of the prey is not only part of the food the spider extracts, it is also a resource enabling efficient transfer of food from prey to predator. The value of the prey's fluid content as a resource decreases as a function of feeding time and as a consequence of the spider feeding on the prey.     

Prey preferences ofPortia fimbriata,an araneophagic,web-building jumping spider (Araneae: Salticidae) from Queensland

Portia fimbriata from Queensland, a previously studied jumping spider (Salticidae), routinely includes web-building spiders and cursorial salticids in its diet, both of these types of prey being dangerous and unusual prey for a salticid. The present paper is the first detailed study ofP. fimbriata's prey preferences. Three basic types of tests of prey preference were used, providing evidence that (1)P. fimbriata males and females prefer spiders (both web-building spiders in webs and salticids away from webs) to insects; (2)P. fimbriata males and females prefer salticids to web-building spiders; (3)P. fimbriata males and females prefer larger spiders to smaller spiders; (4) there are intersexual differences in the preferences ofP. fimbriata for prey size, females preferring larger prey and males preferring smaller prey; and (5)P. fimbriata's prey preferences are not affected by a prior period without food of 2 weeks. When preferences were tested for by using both living, active prey and dead, motionless lures, the same preferences were expressed, indicating thatP. fimbriata can distinguish among different types of prey independent of the different movement patterns of different prey.