M. Nyffeler’s research while affiliated with University of Basel and other places

In this paper the findings of a global literature and social media survey of spider mycoses are presented. Our survey revealed that spider mycoses occur in the geographic belt between latitude 78.00 N and 52.00 S, and that more than 40 out of the known 132 spider families (>30%) are attacked by fungal pathogens. Jumping spiders (Salticidae), cellar spiders (Pholcidae), and sheet web spiders (Linyphiidae) are the families most frequently reported to be attacked by fungal pathogens (combined >40% of all reported cases). Ninety-two percent of the infections of spiders can be attributed to pathogens in the order Hypocreales / phylum Ascomycota (i.e., almost exclusively families Cordycipitaceae and Ophiocordycipitaceae). Within the Hypocreales, the anamorph genus Gibellula is an historically species-rich and widespread genus of specific spider-pathogenic fungi. For ≈70 species of spider-pathogenic fungi their hosts could be identified at least to family level. The data presented here reaffirms the findings of previous studies that spider-pathogenic fungi are most common and widespread in tropical and subtropical forested areas, with free-living cursorial hunters (especially Salticidae) being most frequently infected. Free-living cursorial hunters (especially Salticidae) and subterranean cellar spiders (Pholcidae) are the most frequently fungus-infected spiders in North America, whereas web-weavers (especially Linyphiidae and Pholcidae) are the most common spider hosts in Europe. Our survey implies that spider-pathogenic fungi are an important mortality factor of spiders which has hitherto been underestimated.

In this review, we report on harvestmen and spiders feeding on fungi, fruits, and seeds. Fungivory in harvestmen is widespread, with most reports referring to tropical species in the family Sclerosomatidae, which consume mainly small forest mushrooms (families Marasmiaceae and Mycenaceae). In contrast, consumption of fungal material by spiders apparently occurs only if airborne spores trapped in the viscid threads of orb-webs (e.g., Araneidae and Tetragnathidae) are ingested along with old webs prior to the construction of new webs. Consumption of fruit pulp by harvestmen is also widespread, with several records of Leiobunum spp. (Sclerosomatidae) feeding on Rubus spp. berries and other lipid-poor fruits in the Holarctic region. In Neotropical forests, harvestmen in the families Cosmetidae and Gonyleptidae feed on lipid-poor pulp of fallen fruits. Among spiders, we document several cases of synanthropic species opportunistically feeding on fruit waste (e.g., pieces of banana, papaya, watermelon, or orange pulp) inside houses or disposed in yards. Only one case of a spider feeding on a wild fruit in the field was found in our search. Finally, we report several cases of harvestmen and spiders feeding on elaiosomes or arils (i.e., lipid-rich seed appendages). In conclusion, harvestmen consume mushrooms, fruit pulp, seeds, and seed appendages more frequently than spiders probably because they are “solid food feeders”, which means they can ingest solid tissues by biting off small pieces. In turn, spiders are “fluid feeders” and feed on vegetable matter most frequently in the form of fluids (e.g., nectar, stigmatic exudate, plant sap, and honey dew), rather than fungal or plant tissues.

According to a recent global literature survey, a total of 39 out of the 129 known spider families (30%) contain species capable of capturing vertebrate prey. The finding that the percentage of spider families engaged in vertebrate predation is so high is novel. Two groups of vertebrate-eating spiders are distinguished: habitual vertebrate-eaters vs. occasional vertebrate-eaters. The habitual vertebrate-eaters comprise ten spider families (Araneidae, Atracidae, Ctenidae, Lycosidae, Nephilidae, Pisauridae, Theraphosidae, Theridiidae, Trechaleidae, and Sparassidae) to which can be attributed 91% of all reported vertebrate predation incidents. The habitual vertebrate-eaters have evolved prey-capture adaptations such as (1) sufficient physical strength coupled with large body size, (2) the use of potent venoms, and (3) the use of highly efficient prey-catching webs. By contrast, unexpected feeding on vertebrates by the occasional vertebrate-eaters (i.e., Actinopodidae, Agelenidae, Amaurobiidae, Anyphaenidae, Barychelidae, Clubionidae, Corinnidae, Ctenizidae, Cyrtaucheniidae, Deinopidae, Desidae, Dipluridae, Eresidae, Filistatidae, Gnaphosidae, Haplonoproctidae, Linyphiidae, Liocranidae, Miturgidae, Oxyopidae, Pholcidae, Porrhothelidae, Salticidae, Selenopidae, Sicariidae, Sparassidae, Tetragnathidae, and Thomisidae) might be considered as chance events that took place when a tiny vertebrate crossed the path of an opportunistic spider. For a few families (e.g., Idiopidae) their status as habitual or occasional vertebrate predators is still unclear. In conclusion, our survey unveiled a large number of spider taxa previously not anticipated to feed on vertebrate prey. These findings improve our general understanding of spider feeding ecology and provide a first assessment of the significance of vertebrate prey as a food source for spiders.

In this paper, we present an update on our knowledge on egg predation (oophagy) by spiders. Based on a survey of 233 reports, ghost spiders (Anyphaenidae), lynx spiders (Oxyopidae), jumping spiders (Salticidae), and yellow sac spiders (Cheiracanthiidae) were the most prominent groups of spiders engaged in oophagy. Around 75% of the reports referred to the consumption of lepidopteran and spider eggs worldwide. Another 10% referred to the consumption of eggs/embryos of anurans especially predation upon embryos of glass frogs (Centrolenidae) by spiders from the families Anyphaenidae and Trechaleidae in the Neotropics. The remaining 17% included rare instances of feeding on eggs of coleopterans, dermapterans, dipterans, heteropterans, homopterans, hymenopterans, acarids, neuropterans, opilionids, and squamates. Our study demonstrates that oophagy in spiders is much more widespread than previously thought, both geographically and taxonomically. The finding that spiders feed on eggs/embryos from so many different invertebrate and vertebrate taxa is novel.

In this paper, vertebrate predation by jumping spiders (Salticidae) was revisited, taking into account incidents of this kind recently published in the scientific literature or on the internet. Our study revealed that vertebrate predation by salticids is more widespread than previously thought, geographically and taxonomically. Roughly ninety percent of all reported cases refer to predation on anurans (Hylidae and Ranixalidae) and lizards (Dactyloidae and Gekkonidae) by salticids from the subfamily Salticinae (Hyllus spp., Phidippus spp., and an unidentified species presumably related to Hasarius Simon, 1871). In the remaining cases, salticids from the subfamily Salticinae (Paraphidippus cf. aurantius (Lucas, 1833) and Phidippus audax (Hentz, 1845)) were observed attacking bird hatchlings (families Paridae and Trochilidae), weighing 46 times more than the spiders. In two instances, the spiders were observed biting the hatchlings, but only in one single case, a salticid was seen feeding on a hatchling.

In this paper, 319 incidents of snake predation by spiders are reported based on a comprehensive global literature and social media survey. Snake-catching spiders have been documented from all continents except Antarctica. Snake predation by spiders has been most frequently documented in USA (51% of all incidents) and Australia (29%). The captured snakes are predominantly small-sized with an average body length of 25.9 ± 1.3 cm (median = 27 cm; range: 5.8–100 cm). Altogether >90 snake species from seven families have been documented to be captured by >40 spider species from 11 families. About 60% of the reported incidents were attributable to theridiids (≈0.6–1.1 cm body length), a spider family that uses strong tangle webs for prey capture. Especially the Australian redback spider (Latrodectus hasselti Thorell, 1870), the African button spider (Latrodectus indistinctus O. Pickard-Cambridge, 1904), an Israeli widow spider (Latrodectus revivensis Shulov, 1948), and four species of North American widow spiders (Latrodectus geometricus C.L. Koch, 1841, Latrodectus hesperus Chamberlin & Ivie, 1935, Latrodectus mactans (Fabricius, 1775), and Latrodectus variolus Walckenaer, 1837) – equipped with a very potent vertebrate-specific toxin (α-latrotoxin) – have proven to be expert snake catchers. The use of vertebrates as a supplementary food source by spiders represents an opportunity to enlarge their food base, resulting in enhanced survival capability. Interestingly, the snakes captured by spiders also encompasses some species from the families Elapidae and Viperidae known to be highly toxic to humans and other vertebrates. Not only do spiders sometimes capture and kill snakes, quite often the tables are turned – that is, a larger number of arthropod-eating snake species (in particular nonvenomous species in the family Colubridae) include spiders in their diets.

In this paper, 374 incidents of frog predation by spiders are reported based on a comprehensive global literature and social media survey. Frog-catching spiders have been documented from all continents except for Antarctica (>80% of the incidents occurring in the warmer areas between latitude 30° N and 30° S). Frog predation by spiders has been most frequently documented in the Neotropics, with particular concentration in the Central American and Amazon rain forests and the Brazilian Atlantic forest. The captured frogs are predominantly small-sized with an average body length of 2.76 ± 0.13 cm (usually ≈0.2–3.8 g body mass). All stages of the frogs' life cycle (eggs/embryos, hatchlings, tadpoles, emerging metamorphs, immature post-metamorphs, adults) are vulnerable to spider predation. The majority (85%) of the 374 reported incidents of frog predation were attributable to web-less hunting spiders (in particular from the superfamilies Ctenoidea and Lycosoidea) which kill frogs by injection of powerful neurotoxins. The frog-catching spiders are predominantly nocturnal with an average body length of 2.24 ± 0.12 cm (usually ≈0.1–2.7 g body mass). Altogether >200 frog species from 32 families (including several species of bitter tasting dart-poison frogs) have been documented to be hunted by >100 spider species from 22 families. Our finding that such a high diversity of spider taxa is utilizing such a high variety of frog taxa as prey is novel. The utilization of frogs as supplementary food increases the spiders' food supply (i.e., large diet breadth), and this is presumed to enhance their chance of survival. Studies from Australia and South America indicate that frogs might be a substantial component in the diet of some mygalomorph spiders (i.e., families Atracidae, Idiopidae, and Theraphosidae). Many more quantitative investigations on the natural diets of tropical spiders are needed before reliable conclusions on the importance of frogs as spider food can be drawn.

Aerial web-spinning spiders (including large orb-weavers), as a group, depend almost entirely on flying insects as a food source. The recent widespread loss of flying insects across large parts of western Europe, in terms of both diversity and biomass, can therefore be anticipated to have a drastic negative impact on the survival and abundance of this type of spider. To test the putative importance of such a hitherto neglected trophic cascade, a survey of population densities of the European garden spider Araneus diadematus—a large orb-weaving species—was conducted in the late summer of 2019 at twenty sites in the Swiss midland. The data from this survey were compared with published population densities for this species from the previous century. The study verified the above-mentioned hypothesis that this spider’s present-day overall mean population density has declined alarmingly to densities much lower than can be expected from normal population fluctuations (0.7% of the historical values). Review of other available records suggested that this pattern is widespread and not restricted to this region. In conclusion, the decline of this once so abundant spider in the Swiss midland is evidently revealing a bottom-up trophic cascade in response to the widespread loss of flying insect prey in recent decades.

Aerial web-spinning spiders (including large orb-weavers) depend, as a group of insectivores, completely on flying insects as a food source. The recent widespread loss of flying insects across large parts of western Europe, both in terms of diversity and biomass, can therefore be anticipated to have a drastic negative impact on survival and abundance of this type of spiders. To test the putative importance of such a to date neglected trophic cascade, a survey of population densities of the European garden spider Araneus diadematus (a large orb-weaving spider) was conducted in late summer 2019 on twenty sites of the Swiss midland. The data from this survey were compared with published population densities for this species from the previous century. The study verifies above-mentioned hypothesis that this spider's present-day overall mean population density has declined alarmly to densities much lower than can be expected from normal population fluctuations (0.7% of the historical densities). Review of other available records suggests this pattern is widespread and not restricted to this region. In conclusion, the here documented abundance decline of this once so abundant spider in the Swiss midland is evidently revealing a bottom-up trophic cascade in response to the widespread loss of flying insect prey in recent decades.