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A synthesis on cave-dwelling spiders in Europe
Journal of Zoological Systematics and Evolutionary Research
Abstract
We provide the first overview on spiders living in subterranean habitats in Europe, including the first European subterranean spider checklist. In Europe, there are 486 spider species known to dwell in caves and other subterranean habitats, distributed across 22 families. Despite a few species being able to colonize caves across the whole continent, approximately 90% of the species show a restricted distribution, occurring exclusively in one or two countries. From a biogeographic perspective, southern Europe emerges as the main hot spot of subterranean spider diversity, showing the highest richness of endemic species. Compared to other temperate regions of the world, some families appear to be well represented and other poorly represented (or lacking) in European subterranean habitats. Overall, it appears that the taxonomical knowledge on subterranean spiders in Europe is sufficient, but not evenly distributed. As this checklist represents a useful baseline for advances in this field, we point out specific areas of interest for future research.
Supplementary resources (2)
Data
December 2017
... In total, there are 14 publications that have provided 11 spider species for the Crimean caves. Compared to other karst regions of the Alpine-Mediterranean foldbelt, the Crimean caves are characterised by lower spider diversity (Turbanov et al. 2016b;Mammola et al. 2018). Yet, the overall spider diversity in Crimea is equal/comparable to that of neighbouring regions of similar size (Kovblyuk and Kastrygina 2015;Nentwig et al. 2024). ...
... This species is likely to be a facultative synanthrope that can inhabit underground biotopes as a subtroglophile. Charitonov, 1947 Figs 1B, 3-5 Tegenaria taurica Charitonov, 1947: Charitonov 1947aCharitonov 1947b: 1;Birstein 1963: 128;Tyshchenko 1971: 23, 156, 161, 163;Mikhailov 1997: 145;Mikhailov 1998: 22;Esyunin and Farzalieva 2001: 261-263, figs 1-5;Kovblyuk 2002: 105;Amelichev et al. 2004: 136, 140;Evtushenko 2004: 66, 68;Kovblyuk 2004a: 214;Kovblyuk 2004b: 43, 45, 47-48, figs 2, 3(1) ;Kovblyuk 2004c: 254, 256;Zagorodniuk and Vargovitsh 2004: 207;Kovblyuk 2010: 224;Mikhailov 2013: 141;Bolzern et al. 2013: 776, 803, 818, 846;Kovblyuk 2014: 34, 44, 51;Kovblyuk and Kastrygina 2015: 6;Turbanov et al. 2016bTurbanov et al. : 1283Prokopov and Turbanov 2017: 101;Mammola et al. 2018: : Charitonov 1932: 21;Charitonov 1939: 197. Tegenaria sp.: Evtushenko 2004Zagorodniuk and Vargovitsh 2004: 208;Turbanov et al. 2019a: 218. ...
... Ecology. A troglophile (Mammola et al. 2018). There is a single record of T. taurica from an anthropogenic biotope (Kovblyuk 2004b), which in fact refers to T. parietina (1 ♂ (TNU 1630/1), Yalta, indoors, 17.X.2001 ...
Based on a critical review of the literature and study of the authors’ own collections, a survey of cave spiders of the Crimean Mountains has been conducted, resulting in 20 reliable species records in eight families. Nine species have been discovered in the Crimean caves for the first time, of which two are described as new to science. A classification of spiders by ecological groups depending on their cave lifestyle is provided; a troglomorphic spider is found and described from Crimea for the first time. The most likely scenarios of spider colonization into underground habitats of Crimea are discussed. Most species arrived during multiple Pleistocene-Holocene regressions of the Black Sea basin, when zoogeographic corridors on the exposed shelf connected Crimea with the Caucasus and the Balkans. However, four synanthropic species entered Crimean caves in historical times. High relative humidity and temperature are considered key factors that enable caves to serve as refugia for Pleistocene araneofauna. A zoogeographical analysis of cave spiders is carried out. The majority of the spider species considered, totalling 16 species, are widespread, with ranges including cosmopolitan, Holarctic, trans-Palaearctic, West and Central Palaearctic, East European, and East Mediterranean. Three species are endemic to Crimea: Tegenaria taurica, Bisetifer tactussp. nov., and Troglohyphantes exspectatussp. nov.Bisetifer gruzin is a Crimean-Caucasian subendemic species.
... Species well distributed at European level, especially in warm localities, it is often found in buildings. Troglophile species (Mammola et al. 2018) observed for the first time in the cave "Addaura III", from Roewer (1960a), data taken from Brignoli (1972) who also reports the species for the cave "Grotta dei Bovidi" (located near the cave mentioned above) and still Dresco (1963) also for the "Grotta dei Bovidi". ...
... Species linked to caves, occasionally found under large stones. Troglophile species (Mammola et al., 2018). Species reported for "Grotta Addaura III" (Denis, 1959;Roewer, 1960;Strinati, 1962;Brignoli, 1972) and for "Grotta Caprara" (Brignoli, 1972). ...
... Troglophile species (Mammola et al., 2018) reported for "Grotta Caprara" or "Antro nero" from Dresco (1963) and Brignoli (1972). DISTRIBUTION. ...
In this work I report on the presence of some Araneae species new or interesting for Sicily and a preliminary checklist of the Araneofauna of the mountain massif "Monte Pellegrino" located in the territory of Palermo (Sicily, Italy) including: Altella lucida (Simon, 1874) (Dictnydae), Agyneta saxatilis (Blackwall, 1844) (Linyphiidae), Anatolidion gentile (Simon, 1881) (Theridiidae) and Heliophanus auratus C.L. Koch, 1835 (Salticidae). Additional biological, literature, chorotype and distribution are indicated.
... Taxonomic and geographical coverage. We initially updated the checklist of European subterranean spiders provided in ref. 29 to obtain a list of species for which to collect relevant traits. Specifically, we: I). ...
... Detailed explanation of traits, including their hypothesized functional meaning, is given in Table 2. The dataset consists of 64 traits (some examples of trait distributions are given in Fig. 2) for 520 species belonging to 20 families associated with caves (Table 3)-34 species more than in the previous checklist 29 . The family comprising most species is Linyphiidae (224 species, almost half of them belonging to a single genus Troglohyphantes), followed by Dysderidae (62 species), Leptonetidae (60), Nesticidae (56), and Agelenidae (43). ...
... The remaining families are all represented by up to 30 species and encompass spiders with diverse levels of subterranean specialisation. We refer the reader to ref. 29 . for an in-depth taxonomic and biogeographical account. ...
Species traits are an essential currency in ecology, evolution, biogeography, and conservation biology. However, trait databases are unavailable for most organisms, especially those living in difficult-to- access habitats such as caves and other subterranean ecosystems. We compiled an expert-curated
trait database for subterranean spiders in Europe using both literature data (including grey literature published in many different languages) and direct morphological measurements whenever specimens were available to us. We started by updating the checklist of European subterranean spiders, now including 512 species across 20 families, of which at least 192 have been found uniquely in subterranean habitats. For each of these species, we compiled 64 traits. The trait database encompasses morphological measures, including several traits related to subterranean adaptation, and ecological traits referring to habitat preference, dispersal, and feeding strategies. By making these data freely available, we open up opportunities for exploring different research questions, from the quantification of functional dimensions of subterranean adaptation to the study of spatial patterns in functional diversity across European caves.
... It is very common to find troglophile arachnids in the entrance of Levantine caves, as in caves of other regions of the world. Arachnids (and among them spiders) are often numerically dominant in caves and are considered dominant predators in many cave foodwebs [31][32][33][34]. Of the 11 extant terrestrial arachnid orders, five orders (i.e., Araneae, Opiliones, Palpigradi, Pseudoscorpiones, and Scorpiones) and the polyphyletic sub-class Acari were reported to have troglobite species around the world [35][36][37][38], while troglobite species of three orders (i.e., Amblypygi [39], Ricinulei, and Schizomida) were reported only from subtropical and tropical regions [35]. ...
... We assigned each of the 62 spider species in our survey to one of three categories: troglobite, troglophile, and accidental, according to their distribution and the known use of caves as a habitat in Israel and Europe [33]. We also categorized them based on troglomorphic phenotype. ...
... Based on our results, 20% of the families known from Israel and Palestine include troglobite and/or troglophile species, compared to 34% in Europe (22 out of 64 families reported from Europe [33,49]). From our current data, only about 5% of the spider species known from Israel and Palestine are cave-dwellers (40 species) with approximately 2% endemics. ...
Caves share unique conditions that have led to convergent adaptations of cave-dwelling animals. In addition, local factors act as filters on regional species-pools to shape the assemblage composition of local caves. Surveys of 35 Levantine caves, distributed along a climate gradient from the mesic in the north of Israel to hyper-arid areas in the south of Israel, were conducted to test the effect of cave characteristics, location, climate, bat presence, and guano level on the spider assemblage. We found 62 spider species and assigned four species as troglobites, 28 as troglophiles, and 30 as accidentals. Precipitation, elevation, latitude, minimum temperature, and guano levels significantly affected the composition of cave-dwelling spider assemblages. Caves situated in the Mediterranean region had higher species richness and abundance, as well as more troglobite and troglophile arachnids. These discoveries contribute to the knowledge of the local arachnofauna and are important for the conservation of cave ecosystems. By comparing spider assemblages of Levantine caves to European caves, we identified gaps in the taxonomic research, focusing our efforts on spider families that may have additional cryptic or yet to be described cave-dwelling spider species. Our faunistic surveys are crucial stages for understanding the evolutionary and ecological mechanisms of arachnid speciation in Levantine caves.
... To study community assembly rules, we leveraged the unprecedented amount of data available for subterranean spiders in Europe (36), namely community composition data for selected caves across the continent (37), and standardized traits for all species (38). A previous analysis of the taxonomic component of this dataset demonstrated a quick turnover in the taxonomic diversity of subterranean spiders across Europe, mediated primarily by geographic distance among caves, and secondarily by the climatic conditions and availability of karst. ...
... Note that we focused solely on "subterranean spiders" (36,38), excluding "accidental" surface species (55) occasionally found underground. ...
Understanding how species assemble into communities is a central tenet in ecology. One of the most elusive questions is the relative contribution of environmental filtering versus limiting similarity. Important advances in this area have been achieved by looking at communities through a functional lens (i.e., the traits they express), so as to derive principles valid across species pools. Yet, even using traits in lieu of taxonomy, the issue remains controversial because i) environmental filtering and limiting similarity often act simultaneously in shaping communities; and ii) their effect is scale-dependent. We exploited the experimental arena offered by caves, island-like natural laboratories characterized by largely constant environmental gradients and a limited diversity of species and interactions. Leveraging uniquely available data on distribution and traits for European cave spiders, we tested explicit hypotheses about variations in community assembly rules across ecological gradients and scales. We demonstrate that environmental filtering and limiting similarity shape cave communities acting on trait evolution in opposing directions. These effects are strongly scale dependent, varying along multiple environmental gradients. Conversely, the effect of geography on trait composition is weak, indicating that trait turnover in space happens primarily by substitution of species pursuing similar functions due to strong environmental filters. Our findings reconcile contrasted views about the relative importance of the two main mechanisms shaping patterns of biodiversity, and provide a conceptual foundation to account for scaling effects in the study of community assembly.
... Such observations have to our knowledge never previously been reported in the literature. Given the relatively high abundance of Meta menardi in temperate caves and the relatively large number of studies on their distribution and ecology (see Mammola et al. 2018; this is perhaps initially surprising. However, this can likely be explained by the scarcity of walking prey inside caves, the low likelihood of walking prey getting near and interacting with the webs of M. menardi and the notorious challenges posed by conducting detailed behavioural observations in caves (Mammola et al. 2021). ...
The European cave spider ( Meta menardi ) builds orb webs in the entrance and twilight zone of caves. However, the scarcity of flying prey inside caves means that only about half of its diet consists of flying insects, with walking prey making up the remaining half. The capture of non‐flying prey is not generally known from aboveground orb web building spiders, which exclusively capture flying prey in their webs. It is currently a mystery how the cave spider manages to capture prey walking on the cave wall. A number of hypotheses have been suggested, including that the spider leaves its web completely ( the off‐web hunting hypothesis ), that walking prey slips on the smooth cave ceiling and falls into the horizontally inclined webs ( the prey slip hypothesis ) or that the unique geometry of the cave spider's orb web, which lacks frame threads, means that the radii that attach directly to the cave wall are used as tripwires to alert the spider of passing prey ( the radius tripwire hypothesis ). Here we report on three in situ observations of M. menardi responding to radius stimulation from artificial and walking prey by running towards the cave wall. Although the opportunistic observations of responses to real prey both involved potentially dangerous prey (a centipede and another large M. menardi ) and were unsuccessful prey capture events, the clear response to walking prey nonetheless constitutes the first strong observational evidence in support of the radius tripwire hypothesis.
... It is a known phenomenon that in some Harpactea species adapted to subterranean life, the eyes reduce in size or disappear altogether (Mammola et al. 2018). For example, Harpactea karaschkhan Kunt, Özkütük, Elverici, Marusik & Karakaş, 2016, known from caves in Türkiye, is completely eyeless, while Harpactea kalavachiana Gücel, Charalambidou, Göçmen & Kunt, 2019, from caves in neighboring Cyprus, bears smaller eyes and the anterior eyes are more separated than usual (Gücel et al. 2019;Kunt et al. 2016). ...
Two new Harpactea species, H. cosari sp. nov. (male female) and H. subterranea sp. nov. (male female), are described from
the Manisa Province of Türkiye. The two new species belong to the rubicunda species group based on the characteristics
of the male and female copulatory organs. H. subterranea sp. nov. was sampled from the mesovoid shallow substratum
using special traps. An apparent reduction of the eyes was observed as an adaptation to living underground.
In addition to the new species, a new population of Harpactea cruriformis Bosmans, 2011, previously described from
the Greek island of Chios, was discovered in İzmir province and reported as new to the spider fauna of Türkiye. Also, the
unknown female of H. cruriformis is described for the first time. The female copulatory organ of the species resembles
that of Harpactea terveli Lazarov, 2009, as it is the male. Bulb and vulva images of Harpactea pugio Varol & Akpınar,
2016 and H. terveli, along with bulb images of H. kencei Kunt, Elverici, Özkütük & Yağmur, 2011 and the holotype of H.
cruriformis are included for comparison.
... Dysderids are mostly found in forested areas, but they are not uncommon in open habitats. They are also among the most frequent and diverse groups of spiders in Mediterranean caves (Deltshev, 1999;Culver and Sket, 2000;Ribera, 2004;Mammola et al., 2018). Some species have evolved morphological adaptations to the underground habitat (i.e. ...
The family Dysderidae is a highly diverse group of nocturnal ground‐dwelling and active‐hunter spiders. Dysderids are mostly restricted to the Western Palearctic, and particularly rich and abundant around the Mediterranean region. Interestingly, the distribution of species richness among its 24 genera and three subfamilies is highly biased—80% of its 644 documented species belong to just two genera, Dysdera (326) and Harpactea (211). Dysderidae provides an excellent study case for evolutionary and ecological research. It includes cases of trophic specialization, which are uncommon among spiders, and exhibit other remarkable biological (e.g. holocentric chromosomes), behavioural (e.g. cryptic female choice), evolutionary (e.g. adaptive radiation) and ecological features (e.g. recurrent colonization of the subterranean environment). The lack of a quantitative hypothesis on its phylogenetic structure has hampered its potential as a testing ground for evolutionary, biogeographical and ecological hypotheses. Here, we present the results of a target, multi‐locus phylogenetic analysis, using mitochondrial (cox1, 16s and 12s) and nuclear genes (h3, 28s and 18s), of the most exhaustive taxonomic sample within Dysderidae (104 spp.) to date and across related families (Synspermiata) (83 spp.). We estimate divergence times using a combination of fossil and biogeographic node calibrations and use this timeline to identify shifts in diversification rates. Our results support the monophyly of the Dysderidae subfamilies Rhodinae and Dysderinae but reject Harpacteinae as currently defined. Moreover, the clades recovered within Harpacteinae do not support its current taxonomy. The origin of the family most likely post‐dated the break‐up of Pangea, and cave colonization may be older than previously considered. After correcting for the taxonomic artefacts, we identified a significant shift in diversification rates at the base of the genus Dysdera . Although the unique coexistence of specialist and generalist diets within the lineage could be suggested as the potential driver for the rate acceleration, further quantitative analyses would be necessary to test this hypothesis.
... Certaines vivent sur le pierrier, telles que Drassodes lesserti, Pardosa nigra, Philodromus laricium, Heliophanus aeneus (Hahn, 1832), Attulus rupicola. D'autres, trouvées en profondeur, sont considérées comme troglophiles par (Mammola et al. 2018) : Pholcus opilionoides (Schrank, 1781) et Rugathodes bellicosus. Il est fort probable que d'autres taxons puissent compléter cette liste. ...
Un premier inventaire des araignées de la réserve naturelle nationale de Chastreix-Sancy incluant l’espace naturel sensible (ENS) de la montagne du Mont a été réalisé en 2021 et 2022. Huit stations d’échantillonnage ont été définies : haut marais évolué, tourbière haute, hêtraie-sapinière, pelouse vivace sèche, prairie subalpine, lande montagnarde ainsi qu’une zone humide forestière et une zone rase après restauration sur l’ENS de la montagne du Mont. De plus, des relevés sur divers habitats ont été réalisés (aulnaie-frênaie, saulaie, hêtraie, tourbière subalpine, pierrier, ripisylve, etc.) et ont permis de largement compléter cet inventaire. Au total, 302 espèces d’araignées (dont trois au niveau générique) ont été identifiées. Au regard de leur rareté, de leur degré de spécialisation et/ou du niveau de menace qui pèse sur elles, 23 espèces peuvent être considérées comme remarquables, dont 14 inscrites sur la liste des araignées menacées (huit en danger d’extinction [EN], dont deux spécifiquement dans le Massif central et six quasi-menacées [NT]). Quarante espèces sont citées pour la première fois d’Auvergne, quatre d’Auvergne-Rhône-Alpes. Les espèces observées sont majoritairement associées aux milieux ouverts. Les cortèges associés aux tourbières et milieux humides sont parmi les plus remarquables. Les pelouses vivaces sèches non pâturées contribuent à la liste de la réserve par un cortège riche et à affinité xérophile. Des espèces montagnardes, pour certaines d’origine pyrénéenne, ont été mises en évidence. Certaines, présentes uniquement sur les sommets, sont menacées à moyen terme par le changement climatique. Enfin, l’absence de taxons spécialistes et relictuels parmi les cortèges forestiers semble refléter la forte pression anthropique subie par ces milieux au xixe siècle.
... In this country, Argiope and Cyclosa are currently represented by a single species each, and Araneus by five. A total of 248 spider species are currently listed for Kosovo, but considering that Kosovo's spider fauna is poorly known, this number is expected to increase dramatically (Vrenozi & Jäger, 2013;Mammola et al., 2018, Geci & Naumova, 2021a, 2021bGrapci-Kotori et al., 2022;Berisha & Geci, 2023;Geci et al., 2023). This study's aim is to present new findings for the Araneidae spider fauna of Kosovo and to update the local species list for this family. ...
In this paper, we provide additional data on the orb-weaver spider fauna (Araneae: Araneidae) of Kosovo. Three species are reported for the first time from this country: Araneus grossus (C.L. Koch, 1844), Argiope lobata (Pallas, 1772) and Cyclosa oculata (Walckenaer, 1802). In consideration of the findings and the updated checklist of Araneidae in this study, the documented number of spider species in Kosovo has now increased to 251. Given that the spider fauna of Kosovo is still not fully explored, our study emphasizes the necessity of ongoing exploration to better comprehend the diversity of spider species within the region.
... Members of this genus are small (less than 1 cm in body length) and brownish, and their eyes are small or absent. Some Cybaeodes species are adapted to subterranean environments, being found in the MSS (Mesovoid Shallow Substratum) and caves (Mammola et al. 2018), like C. dosaguas Ribera & De Mas, 2015, C. magnus Ribera & De Mas, 2015 indalo Ribera & De Mas, 2015, and C. liocraninus (Simon, 1913). The species C. mallorcensis was originally described from specimens collected in three localities in Majorca (Llac Cúber, Valldemosa, and Sóller) (Wunderlich 2008), and was subsequently recorded in the Cova Novella de na Llebrona (Ribera & De Mas 2015) and in four localities in Minorca (Barrientos & Febrer 2017). ...
The spider genus Cybaeodes Simon, 1878 is found in the Western
Mediterranean region. It belongs to the family Liocranidae,
but its exact phylogenetic position has been debated due to its
unique morphological features. Here we present the first records
of the species Cybaeodes mallorcensis Wunderlich, 2008 outside
of the Balearic Islands, specifically in the Iberian Peninsula,
and speculate on the possible causes of this disjunct distribution.
Additionally, we publish the first genetic data for the genus
Cybaeodes and use them to interrogate about the phylogenetic
position of this remarkable genus within the spider tree of life.
The moderate genetic differentiation found among some of the
individuals sampled in the Iberian Peninsula suggests that these
may be native populations, and not the result of introductions
from the Balearic Islands. However, sequencing specimens from
the islands would help shed some light on their origin. Finally,
the phylogenetic tree containing the new genetic data of
Cybaeodes renders Liocranidae paraphyletic, the genus
Cybaeodes being more closely related to the family
Cithaeronidae and two liocranid genera, albeit with low supports.
Our results highlight the need for a more comprehensive
phylogeny to determine the placement of this obscure genus.
... Despite traditional studies assuming that animals specialized to live underground are strictly bound to the subterranean environment (Barr Jr & Holsinger, 1985;Mammola et al., 2018;Poulson & White, 1969), there is growing evidence that the ecological separation between cave-dwelling and surface animals is not always clearcut, with some surface species actively exploiting underground environments for specific tasks/stages (Lunghi et al., 2014). As an example, it is well documented that populations of typical surface species, like the Pyrenean newt Calotriton asper, regularly exploit groundwaters (Guillaume, 2022;Uiblein et al., 1995). ...
Despite traditional studies assuming that animals specialized
to live underground are strictly bound to the subterranean
environment there is
growing evidence that the ecological separation between
cave-dwelling and surface animals is not always clearcut,
with some surface species actively exploiting underground
environments for specific tasks/stages
... Spiders are considerably more abundant, diverse, and functionally important among all terrestrial invertebrates (Mammola et al 2017). Furthermore, they are prevalent in various habitats, as well as being crucial to conserving biodiversity (Michalko et al 2019, Milano et al 2021. ...
... According to the latest overview of European subterranean spiders (Mammola et al. 2018a), all European species of Typhlonesticus are obligate troglobionts, being exclusively found in subterranean habitats. This ecological preference parallels a suite of morphological and physiological adaptations to the subterranean environment (i.e., troglomorphism; see Christiansen 2012) pertaining, among others, size, leg elongation, loss of pigmentation and eye regression. ...
The spider genus Typhlonesticus Kulczyński, 1914 (Araneae: Nesticidae) includes seven species, most of which exhibiting strict preference for caves, abandoned mines and other subterranean habitats. In Italy the genus is represented by two species: T. morisii (Brignoli, 1975), an extremely narrow endemic species from SW-Alps with a very high level of subterranean adaptation and T. idriacus (Roewer, 1931), showing a much wider distribution in NE-Italy and poor adaptations to subterranean life. Our recent biospele-ological surveys in the Alps lead to the discovery of new populations of highly troglomorphic Typhlonesticus that proved to belong to two new species based on morphological and molecular data. Considering the rarity of these new species, we provide general information on their ecology and distribution, including a comparative analysis of troglomorphic traits in Typhlonesticus in relation to biogeographic factors. Information on the conservation status, useful for assessing their extinction risk based on International Union for Conservation of Nature (IUCN) guidelines, is also provided.
... They have yet to be studied taxonomically. Interpreting a spider as a troglobiont or troglophile can be difficult, even in temperate lineages [50]. In MDL-HC, the diversity and euedaphomorphic morphology of the many small species from soil or caves makes the task particularly complicated, even with parallel sampling. ...
The southern part of the Mekong Delta Limestones of Vietnam (MDL-HC or Hon Chong karst) comprises numerous small limestone hills. It is a hotspot of biodiversity for soil and cave invertebrates. Here, we synthesize the results of biological surveys carried out in Hang Mo So, the richest MDL-HC cave for troglobionts, and in surrounding karsts. Methodologies for the ecological characterization of species are discussed, with emphasis on parallel sampling (external soil plus cave). Hang Mo So has 27 troglobionts, including many still undescribed. An additional 40 cave-obligate species are known from other caves of MDL-HC. Among them, several are expected to be found in Hang Mo So. Most troglobionts of MDL-HC are endemic. Several relictual taxa without close relatives in Southeast Asia occur in Hang Mo So and in MDL-HC, reflecting an ancient origin of the fauna. The reasons for this richness are uncertain, but the cause of its current destruction—quarrying—is all too evident. Most of the original 4 km² of the MDL-HC karst has been destroyed or soon will be, ultimately leaving only 1.6 km² unquarried. Endemic species linked to karst habitats are, therefore, under clear threat of extinction. The Hon Chong karst (MDL-HC) was listed among the ten most endangered karsts on the planet 25 years ago. Today it would probably top the list.
... In addition, E. pallida is often recorded in caves. The subterranean habitat host numerous endemic and specialized arthropod species, and represents a unique and delicate ecosystem that can easily be disrupted by the introduction of alien invasive species (Mammola et al. 2018, Nicolosi et al. 2023. Previous studies investigated and documented the negative effect of alien spider species establishment in Europe (Nentwig 2015), and particularly assessed the danger of congeneric competition in cave habitats (Mammola 2017). ...
Updated non-native distributional data of the invasive spider Eidmannella pallida (Emerton, 1875), Nesticidae, are herein summarized together with the morphological characters of the species. We report all the known localities from the literature. Furthermore, we present and discuss recent findings for the Italian peninsula. In particular, the species has been consistently found in vineyards and annual crop fields in North Italy and in olive groves in South Italy. We include a redescription of the species, and detailed illustrations of the diagnostic characters of both males and females, based on stereomicroscope and SEM images. The spreading potential of the species and its possible impacts on local communities are discussed. Our data suggest that E. pallida has high invasiveness potential and several new findings in non-native areas were recorded in the last years. We found that E. pallida can establish permanent populations in crop-dominated landscapes and has a tendency to exploit vulnerable ecosystems such as the subterranean environment. We thus recommend new and more comprehensive studies on this species to investigate in deeper detail its phylogeny, ecology, and micro-habitat preferences. A high effort should be placed to assess the potential negative effects of E. pallida on the endemic fauna in invaded areas, especially in caves.
... L'espèce se trouve dans des grottes, des mines, mais aussi en forêt sous des pierres (Le Peru, 2011). Dans la présente étude, cette Theridiidae aux moeurs troglophiles (Mammola et al. 2018) a été capturée par C. Besuchet (conservateur au MHNG jusqu'en 1992) qui collectait principalement les coléoptères par tamisage du sol au Winkler, méthode qui lui a permis de trouver de nombreuses espèces intéressantes dans le canton et même dans le monde (Hlaváč 2011). Bien que la méthode de collecte de cette araignée ne soit pas explicitement mentionnée, il est très probable qu'il s'agisse d'un tamisage. ...
Five first and one second record of spiders for Switzerland (Araneae). − Araniella proxima (Kulczyński, 1885), Ballus rufipes (Simon, 1868), Porrhoclubiona leucaspis (Simon, 1932), Robertus mazaurici (Simon, 1901) and Steatoda nobilis (Thorell, 1875) are mentioned for the first time in Switzerland, and Olios argelasius (Walckenaer, 1806) for the second time. These are recent captures in the canton of Geneva made between 2019 and 2022, except for one which comes from a 1973 capture and was discovered in the Natural History Museum of Geneva. The distribution, ecology, circumstances of capture and a discussion of each species are presented.
... For subterranean spiders, we included the entirety of the Alps range from France in the east extending westward through Switzerland, Italy, Liechtenstein, Austria, Germany and Slovenia, by integrating available data in Global Biodiversity Information Facility (GBIF) , Spiders of Europe (Nentwig et al., 2022), Araneae.it (Pantini & Isaia, 2019) and other literature sources (reviewed in Mammola et al., 2018;Mammola, Pavlek, et al., 2022). ...
1. Biodiversity conservation is a central imperative of the 21st century. Subterranean ecosystems deliver critical nature's contributions to people and harbour a broad diversity of poorly understood specialised organisms. However, the subterranean biome is still largely overlooked in global biodiversity targets.
2. We assessed how well subterranean biodiversity is represented in protected areas (Natura 2000 and Emerald networks) in two global hotspots of subterranean biodiversity (the Pyrenees and the Alps). For this, we used two comprehensive databases of terrestrial subterranean taxa, that is, leiodids (beetles) from the Pyrenees and spiders from the Alps, and identified priority areas in each region using both species richness and geographic rarity patterns.
3. Our results show the incapacity of surface-protected area networks to represent subterranean biodiversity, as more than 70% and 90% of the identified priority areas (and 40% and 22% of the species) are not effectively covered by protected areas in the Pyrenees and the Alps, respectively.
4. These findings call for developing an urgent plan for subterranean biodiversity conservation within the European Biodiversity Strategy for 2030.
... At the collection localities, the cave has an almost constant temperature of ~8°C (Cossu 2022, personal communication). In lack of noticeable troglomorphic traits and in accordance with Mammola et al. (2018), we regard the species as troglophile -not a troglobiont as reported by Lana et al. (2016). Interestingly, despite the eyes being normally developed in all specimens (Fig. 5), the abdomen of the specimens collected in the deepest sector of the cave is paler compared to the others. ...
Knowledge on Sardinian cave spiders started with Angela Gozo’s 1908 description of the female of Lepthyphantes sardous. Because Gozo’s original specimens appear to be lost, we visited Gozo’s type location (Su Marmuri cave, Ogliastra, Sardinia) in an effort to find the as yet unknown male and to collect topotypic specimens on which a new diagnosis and a possible revision of the taxonomic position of the species could be based. Both efforts were successful and the results are given below. Our finding of the unknown male now allows the placement of this species within the genus Tenuiphantes Saaristo & Tanasevitch, 1996. Specimens of Tenuiphantes sardous (Gozo, 1908) n. comb. have been collected on the cave ground, among rocks in areas mostly covered with guano. In lack of proper troglomor-phic traits we regard the species as troglophile. As far as the distribution is concerned, the species is currently known exclusively for the type locality. However, the troglophile habit and the general lack of arachnological prospections in Sardinian caves suggests a wider distribution on the island.
... Although often unnoticed by speleologists and cavers alike, the sheet-web spiders in the genus Troglohyphantes Joseph (Araneae: Linyphiidae) are some of the most frequent inhabitants of Southern European caves (Deeleman-Reinhold 1978, Mammola et al. 2018c, 2022. This genus is presently known to include 131 species and 5 subspecies (World Spider Catalog 2022), primarily distributed through the European mountain ranges, from the Cantabrian Mountains in the Iberian Peninsula on the west, to the Caucasus on the east. ...
Spiders of the genus Troglohyphantes (Araneae: Linyphiidae) exemplify one of the largest subterranean adaptive radiation across European mountain ranges, counting over 130 species and representing about one fifth of total species richness of cave spiders in Europe. Despite the emerging potential of Troglohyphantes as a biogeographical model, no attempt has been made to reconstruct the geological events underlying the current distribution patterns of the genus. By coupling traditional taxonomy with target gene sequence data and comparative functional trait analyses, we describe a new species of Troglohyphantes, the first reported from the island of Corsica (France). The species is characterized by a high level of subterranean adaptation and distinct morphological affinities with geographically distant congenerics. By means of time-stamped phylogenies, we tested contrasting hypotheses about the origin of the new species. The most parsimonious explanation suggests that the species diverged from an ancestral group of species originating in the Adriatic plate in the lower Miocene (~19 Ma), colonizing Corsica from the east. In the absence of relevant fossil records, the well-known geochronology of Corsica and the Western Mediterranean basin can be used in future studies to reconstruct the biogeography of the whole genus and for inferring the timeline of its diversification.
... Of these, 1,945 spider species belonging to 493 genera in 61 families have been reported in India (Caleb and Sankaran, 2023;WSC, 2023). They have unique patterns of abundance, diversity, biomass, and functional roles in different ecosystems (Foelix, 2011;Mammola et al., 2018). ...
A detailed investigation of the spider diversity in Salem district, Tamil Nadu was carried out across different habitats for a period of five years. A total of 184 spider species belonging to 97 genera in 29 families were recorded, which represented nearly 10% of Indian and 65% of Tamil Nadu spider diversity. Among them 25 spider species are endemic to India. From the 29 families, the three most abundant families based on number of specimens sampled were Lycosidae (21%), Araneidae (18%), and Eresidae (17%), constituting 56% of the spider species. Simpson diversity indices ranged between 0.88 to 0.30 for all the studied habitats. The species richness was highest in bamboo fields (2.78), and the lowest was observed in grasslands (0.76). The highest abundance of spider species was observed in the sugarcane fields (0.83), followed by the grasslands (0.57) and paddy fields (0.53). Further, these spiders were categorised into 9 types based on their foraging guilds. Among them, the highest species richness was observed in foliage runners. A maximum of nine spider guilds were observed in Shrub ecosystem. Natural ecosystems such as shrublands, treescapes, and grasslands had higher spider diversity than altered agricultural and domestic ecosystems. This is the first report on spider diversity in Salem District, Tamil Nadu revealing the varying spider diversity along with their guild types across different habitats
... Within arachnids, several groups are known to be associated with caves, and troglobitic arachnid lineages account for key components of cave ecosystems worldwide (Mammola et al., 2018a(Mammola et al., , 2018b(Mammola et al., , 2018c. One such group is the order Amblypygi (commonly, "whip spiders" or "tailless whip scorpions"), which are notable for their unusual appearance and their characteristic antenniform first pair of walking legs (the "whips"). ...
Caves constitute ideal study systems for investigating adaptation and speciation, as the abiotic conditions shared by aphotic habitats exert a set of environmental filters on their communities. Arachnids constitute an important component of many cave ecosystems worldwide. We investigated the population genomics of two whip spider species: Sarax ioanniticus, a widely distributed parthenogenetic species found across the eastern Mediterranean; and S. israelensis, a recently described troglomorphic species that is endemic to caves in Israel. Here, we show that S. israelensis is completely genetically distinct from S. ioanniticus and most likely also constitutes a parthenogen. Counterintuitively, despite the lack of genetic variability within S. ioanniticus and S. israelensis, we discovered considerable variation in the degree of median eye reduction, particularly in the latter species. Natural history data from captive-bred specimens of S. israelensis validated the interpretation of parthenogenesis. Our results are most consistent with a scenario of a sexual ancestral species that underwent speciation, followed by independent transitions to apomictic parthenogenesis in each of the two daughter species. Moreover, the lack of genetic variability suggests that variation in eye morphology in S. israelensis is driven exclusively by epigenetic mechanisms.
... The specimen was found wandering in the evening in the small garden of a house. Troglophile species (Mammola et al., 2018). REMARKS. ...
New spiders (Arachnida Araneae) from Sicily (Italy) are reported, some new for Italy, in this paper. Particularly, twelve genera and sixteen new species are examined: Agelena orientalis
... As such, caves and the cave-obligate taxa inhabiting them tend to have high potential for conservation attention. Similarly, the geographic regions that contain large numbers of caves, typically karst habitats (eroded regions of limestone that hold sinkholes, caves, associated mesocaverns, etc.), also deserve conservation attention for their highly localized endemic diversity (e.g., Culver et al. 2003;Mammola et al. 2018a;Rabelo et al. 2018) and because these regions are often threatened by human development (e.g., Parise and Gunn 2007;Fleury 2007;Souza-Silva et al. 2015). The Balcones Escarpment region of central Texas includes karst features that are home to many unique, endemic, and often endangered cave-limited animals. ...
Genomic-scale data for non-model taxa are providing new insights into landscape genomic structuring and species limits, leading to more informed conservation decisions, particularly in taxa with extremely restricted microhabitat preferences and small geographic distributions. This study applied sequence capture of ultraconserved elements (UCEs) to gather genomic-scale data for two federally endangered Texella harvester species distributed in Edwards Formation cave and karst habitats of central Texas, near Austin. We gathered UCE data for 51 T. reyesi specimens from 46 different caves, seven T. reddelli specimens from five caves, and from relevant outgroup species. For these UCE data we applied a combination of phylogenomic, multispecies coalescent phylogenetic, and single-nucleotide polymorphism machine-learning analyses. We found that samples of T. reddelli and T. reyesi together form a single clade in phylogenetic analyses, but that T. reddelli samples are not recovered as monophyletic. Instead, T. reddelli samples from three northern caves are embedded within a larger T. reyesi genetic clade. Significantly, the genetic structuring of all samples closely follows geologic barriers defined for the region and formalized as karst fauna regions (KFRs). One exception is the Jollyville Plateau KFR, which includes two divergent, non-sister genetic lineages. Levels of troglomorphy, here assessed by a simple scoring of corneal and retinal development, also closely follows clade (and geographic) boundaries, implying that divergent genetic lineages might also have distinct ecologies. Overall, our study has important taxonomic implications, is the first to explore (and validate) regional KFR boundaries using intraspecific genetic data, and provides essential data for future management decisions involving these federally endangered species.
... Spiders in the family Leptonetidae are included in the group of haplogyne spiders because they have relatively simple reproductive organs. They are small (1.0-3.0 mm), light coloured (pale) and with 6 eyes; in some troglobiont species, the eyes are reduced or may be completely absent (Mammola et al. 2018). The eyes are distributed as four in the anterior and two in the posterior. ...
Leptonetela turcica sp. n. is described and illustrated based on material collected in the Kahramanmaraş Province of Turkey. Differences between the new species and the closely related species are discussed.
... Indeed, the collected specimen was found inside a house. It may occupy the entrance of caves (Zamani et al. 2014, Mammola et al. 2018). Its bite does not cause pain but could cause injuries, since it is equipped with necrotic venom. ...
A first checklist of the spiders (Araneae) of the Ionian islands of Cephalonia and Ithaka (Greece) is provided. Argiope bruennichi (Scopoli, 1772), Metellina merianae (Scopoli, 1763), Frontinellina frutetorum (C.L. Koch, 1835), Leptorchestes berolinensis (C.L. Koch, 1846), and Menemerus semilimbatus (Hahn, 1827) are recorded for the first time for Cephalonia island, and Philaeus chrysops (Poda, 1761) for Ithaka island.
... The arachnofauna of caves in the southern Levant, however, is poorly characterised compared to its epigean habitats, and to that of European caves [17,18]. We have previously demonstrated that southern Levantine caves harbour diverse assemblages of troglophile and troglobite arachnids, with 62 observed spider species in 35 caves in Israel and Palestine, including 32 troglobite and troglophile spiders [19]. ...
Caves possess a continuum of ecological zones that differ in their microhabitat conditions, resulting in a gradient of nutrients, climate, and illumination. These conditions engender relatively rapid speciation and diverse assemblages of highly specialised spider fauna. It is unclear, however, how zonation of these caves affects spider assemblage composition and structure. Surveys of 35 Levantine caves were conducted to compare the assemblages of spiders between their different ecological zones. The diverse spider assemblages of these caves differed between the entrance, twilight, and dark zones, with troglophiles and accidental species occupying the cave entrance, endemic troglobites occupying the dark zones, and hybrid assemblages existing in the twilight zones. The progression of assemblage composition and divergence throughout cave zones is suggestive of processes of ecological specialisation, speciation, and adaptation of cave-endemic troglobites in the deepest zones of caves, while cave entrance assemblages are composed of relatively common species that can also be found in epigean habitats. Moreover, the cave entrance zone assemblages in our study were similar in the different caves, while the cave dark zone assemblages were relatively distinct between caves. Cave entrance assemblages are a subset of the regional species pool filtered by the cave conditions, while dark zone assemblages are likely a result of adaptations leading to local speciation events.
... Species names and species validity have been checked from [15] and public databases [16,17]. Species ecological status has been inferred from the taxonomic literature, from [18], and from [19] for spiders. ...
Located in Northern Pyrenees, in the Arbas massif, France, the system of the Coume Ouarnède, also known as Réseau Félix Trombe—Henne Morte, is the longest and the most complex cave system of France. The system, developed in massive Mesozoic limestone, has two distinct resurgences. Despite relatively limited sampling, its subterranean fauna is rich, composed of a number of local endemics, terrestrial as well as aquatic, including two remarkable relictual species, Arbasus caecus (Simon, 1911) and Tritomurus falcifer Cassagnau, 1958. With 38 stygobiotic and troglobiotic species recorded so far, the Coume Ouarnède system is the second richest subterranean hotspot in France and the first one in Pyrenees. This species richness is, however, expected to increase because several taxonomic groups, like Ostracoda, as well as important subterranean habitats, like MSS (“Milieu Souterrain Superficiel”), have not been considered so far in inventories. Similar levels of subterranean biodiversity are expected to occur in less-sampled karsts of central and western Pyrenees.
... Our discovery of C. dobrogicus from Tavrida Cave is the first reliable finding of this spider species in Russia. The species is considered as troglophilic (Mammola et al., 2018: Table S1). In addition to caves, it inhabits sandy steppes, sea coasts and agrocenoses (Polchaninova, Prokopenko, 2013). ...
Preliminary results of studying the biota of the recently discovered Tavrida Cave (Foothill Crimea) are presented. By now, six species of arthropods (Arthropoda) have been identified, i.e. one species of spiders (Aranei), one species of diplura (Diplura), two species of springtails (Collembola) and two species of flies (Diptera). The obtained results indicate the relative youth of the recent fauna of Tavrida Cave, despite Eopleistocene age cave, and have a significant faunal value, since a number of discovered species are specified for the first time for the Crimea and Russia. For the troglobiont diplura Campodea (Dicampa) taurica, discovered in Tavrida Cave, a probable scenario of its penetration into the karst cavities of the Crimea in connection with global climate changes in the Late Pleistocene time is presented. However, all other found species are most likely to have entered Tavrida Cave at the present time, after its opening as a result of road construction work.
... Tegenaria faniapollinis Brignoli, 1978(Deltshev 2008a Tegenaria sp. Leptonetidae • ■ Leptonetela thracia Gasparo, 2005(Gasparo 2005b, Wang & Li 2011, Naumova et al. 2016, Mammola et al. 2017, Wang et al. 2017 Linyphiidae Centromerus milleri Deltshev, 1974, Gasparo 2005b, Naumova et al. 2016 ...
In this report, we present the results of the action "Inventory of habitat 8310 in the project sites" (action A1.2) which aims to collect and analyze data to describe and evaluate the conservation of habitat 8310 (Caves not open in the public) in 8 sites of the project [caves Tzani and Psimaki (Crete), cave Limon (Achaia), cave Za (Naxos island), Aliartos sinkhole (Voiotia), Spilaiovarathro cave (Pella), Zesta Nera cave (Serres) and Maroneia cave (Komotini)] the relevant invertebrate fauna and the typical species. In addition, a brief reference is made to some interesting species in the lower Panagopoula’s tunnel although as a technical work is irrelevant with habitat 8310.
... Troglophile species (Mammola et al. 2018), (Van Helsdingen, 2020). ...
Motivation
Groundwater ecosystems sustain a unique and globally important biodiversity but remain understudied due to sampling and exploration challenges, as well as a shortage of taxonomic experts. Groundwater ostracods, like other groundwater taxa, exhibit a high degree of endemism, rarity and subterranean specialisation, positioning them as potentially vulnerable organisms. To better understand biodiversity patterns and the conservation needs of this highly diverse group, we assembled a team of experts to gather the most comprehensive information available about groundwater ostracods in Europe. We present a dataset comprising 2065 occurrence records of 110 species, 11 undescribed species and 5 subspecies of groundwater ostracods. This open dataset may support future research on the distribution, evolutionary pathways and conservation needs of European groundwater ostracods, as well as inspire targeted sampling efforts in regions with currently limited data available.
Main Types of Variables Contained
Occurrence records of groundwater ostracods, with details about taxonomy, source of records, occurrence locality, habitat type and species dependence on groundwater (obligate [stygobiont] versus facultative groundwater‐dwellers [stygophile]).
Spatial Location and Grain
Geographical Europe, spanning 32 countries. Occurrence records were assigned decimal degrees coordinates (EPSG:4326). Most occurrence records are at 100 m resolution.
Time Period
1915–2024.
Major Taxa and Level of Measurement
Crustacea: Ostracoda. Most records have species or subspecies‐level identification, while some are identified to genus or family levels.
Software Format
Comma‐separated values file (.csv) and Excel file (.xlsx), with UTF‐8 encoding and metadata provided following the Darwin Core standard.
The strawberry spider Araneus alsine (Walckenaer, 1802) is a Palearctic species that has been reported from most European countries, including several in the Balkans. In this contribution, for the first time, we report the presence of this species in Kosovo. A single specimen was observed on 24 August, 2023 in the Blinajë hunting reserve, in the municipality of Lipjan. The specimen was found on grass near a forest.
Aim
Quantifying the relative contribution of environmental filtering versus limiting similarity in shaping communities is challenging because these processes often act simultaneously and their effect is scale‐dependent. Focusing on caves, island‐like natural laboratories with limited environmental variability and species diversity, we tested: (i) the relative contribution of environmental filtering and limiting similarity in determining community assembly in caves; (ii) how the relative contribution of these driving forces changes along environmental gradients.
Location
Europe.
Time period
Present.
Major taxa studied
Subterranean spiders.
Methods
We used data on distribution and traits for European cave spiders (n = 475 communities). We estimated the trait space of each community using probabilistic hypervolumes, and obtained estimations of functional richness independent of the species richness of each community via null modelling. We model functional diversity change along environmental gradients using generalized dissimilarity modelling.
Results
Sixty‐three percent of subterranean spider communities exhibited a prevalence of trait underdispersion. However, most communities displayed trait dispersion that did not depart significantly from random, suggesting that environmental filtering and limiting similarity were both exerting equally weak or strong, yet opposing influences. Overdispersed communities were primarily concentrated in southern latitudes, particularly in the Dinaric karst, where there is greater subterranean habitat availability. Pairwise comparisons of functional richness across caves revealed these effects to be strongly scale‐dependent, largely varying across gradients of cave development, elevation, precipitation, entrance size and annual temperature range. Conversely, geographical distance weakly affected trait composition, suggesting convergence in traits among communities that are far apart.
Main conclusions
Even systems with stringent environmental conditions maintain the potential for trait differentiation, especially in areas of greater habitat availability. Yet, the relative influence of environmental filtering and limiting similarity change with scale, along clear environmental gradients. The interplay of these processes may explain the assembly of species‐poor subterranean communities displaying high functional specialization.
Invasive non-native species are recognized as a serious threat to the native biodiversity of the areas they colonize. The subterranean spider Howaia mogera (Yaginuma 1972) (syn. Nesticella mogera) is considered a highly invasive non-native species rapidly expanding its geographical range from Asia across the European continent and to remote oceanic islands. Due to its preference for moist, dark, and climatically stable habitats, including caves, this spider represents a potential threat to the endemic fauna living in the forest litter and subterranean environments. Nevertheless, the origin, biogeography, patterns of colonization, and niche preference of H. mogera remain unexplored. In this study, we reconstructed the phylogeography of the species combining a multi-locus phylogenetic analysis and a niche modeling approach. Our results confirm the center of origin of H. mogera in southern China, from where the species naturally spread and diversified in eastern Asia during the Pleistocene Epoch. Its expansion into non-native areas occurred only recently, possibly as a consequence of human-mediated passive transportation and in conjunction with a shift in the habitat preference of the species. Non-native populations have shifted from an original preference for a subterranean lifestyle to more generalist conditions. This change has allowed them to exploit a wider breadth of habitats and has facilitated their expansion in Europe and remote oceanic islands. Yet, the retention of the original subterranean habitat preferences in these non-native populations poses a conservation threat to specialized and fragile subterranean ecosystems, which H. mogera can efficiently exploit in invaded areas. Our study underscores the importance of comprehending phylogeography and niche dynamics in invasive non-native species to predict and manage their future expansions. We further highlight the urgency of monitoring H. mogera's spread worldwide, particularly in Southern Europe, to protect fragile invaded ecosystems.
En Canarias el medio subterráneo terrestre habitable ocupa una elevada proporción del terreno e incluye hábitats diversos como los tubos de lava y otros tipos de cavidades volcánicas, la red profunda de grietas, el medio subterráneo superficial o los campos de piroclastos. Su extensión y frecuente interconexión permite la presencia de una gran diversidad de especies adaptadas a este medio (troglobiontes). Todas las arañas troglobiontes encontradas son endemismos canarios y cada isla, a excepción de Lanzarote, tiene sus propias especies. Tras los coleópteros, las arañas son el grupo de artrópodos con más troglobiontes en el archipiélago: 39 especies pertenecientes a 9 familias distintas. En este trabajo se aporta un listado de las arañas troglobiontes del archipiélago canario, se comentan datos relevantes de las distintas familias representadas, y se compara dicha fauna con la de otros archipiélagos oceánicos. Finalmente, se abordan las principales amenazas que afectan a estas especies y a la conservación del medio subterráneo en Canarias.
The subterranean heritage includes both natural and built sites with a strong cultural and historical fingerprint, some of each being enriched with painted surfaces. These semi-confined environments shelter specific and fragile biodiversity. This paper is focused on the case of a Roman painting (2nd-3rd century AD) located in an underground archaeological site in Marino Laziale, near Rome, which was opened to the public for the first time in 2021. The painted Mithraic scene is in a good state of conservation. The methodological approach included on site and laboratory investigations aimed to screen the main biological components associated to this hypogeum monument. The observed biodiversity included heterotrophic and chemolithotrophic microorganisms, and a mesofauna composed of eutroglophile and subtroglophile species, characteristic for many subterranean environments. The ecological mechanisms and the conservation state of the work of art were analyzed for planning the best fruition practices. The aesthetic change, the possible mechanical damages induced by various organisms, and the presence of significant amounts of organic matter, represent the main risks for painting conservation. These aspects, beside the new possible risks associated with the presence of visitors, are under a constant and ongoing conservation surveillance program.
We revise the Portuguese spiders of the genus Harpactea Bristowe, 1939 (Araneae: Dysderidae). The following seven new species are described: H. adicensis sp. nov., H. crespoi sp. nov., H. dolanskyi sp. nov., H. henriquesi sp. nov., H. korenkoi sp. nov., H. krejcii sp. nov. and H. pekari sp. nov. In three species, H. algarvensis Ferrández, 1990, H. minoccii Ferrández, 1982 and H. tavirensis Wunderlich, 2020 the females are described for the first time. The Portuguese Harpactea spiders belong to two species groups, namely the hombergi group and the corticalis group (sensu Deeleman-Reinhold 1993). The majority of the Portuguese representatives of the corticalis group probably constitute a monophyletic group endemic for the Iberian peninsula, the minoccii subgroup Ferrández, 1990. The synapomorphy of this clade is the unique position of partners during copulation: the male grasps one of the femaleʼs chelicerae between his chelicerae. The malesʼ chelicerae are morphologically adapted for this behaviour. Due to this unusual position during copulation the female genitalia are more distant from the male than in other Harpactea species. Therefore, the pedipalps are more elongated . Furthermore, the males of this subgroup possess larger basal part of male copulatory organ (the tegulum), where the ejaculate is stored before copulation. We observed that the males of this subgroup invest significantly more time into single copulation than the other representatives of the corticalis group with smaller tegulum. Therefore, enlargement of the tegulum might reflect different sperm competition strategy, in which males invest more ejaculate in each copulation. In Portugal, Harpactea spiders are frequently found under woody plants that produce slowly decomposing leaf litter, usually Quercus spp., or introduced Eucalyptus sp. These spiders require slightly humid substratum.
Simple Summary
The subterranean fauna is an important component of global biodiversity. However, research on the subterranean fauna of the Yunnan–Guizhou Plateau in Southwest China, one of three important karst landforms in the world, is limited. In this study, we performed a population genetic analysis and reconstructed the phylogenetic tree of six populations of Trogloneta yunnanensis in South China Karst. The results showed that there was high genetic divergence among six populations, and the divergence of these six populations can be traced back to the late Pleistocene. Our results suggested that isolation was a pivotal factor affecting the biodiversity of cave faunas, and the biodiversity of cave-dwelling faunas needs to be studied as soon as possible.
Abstract
Subterranean karst caves can contain unexpected biodiversity, but few studies related to spider population genetics have been conducted in the karst area of Southern China. In this study, we investigated the population genetic structure of Trogloneta yunnanensis (Song & Zhu, 1994) based on 73 spider samples from six underground populations in South China Karst. Population genetic structure analysis showed a clear divergence (FST > 0.9 and Nm < 0.05) among populations according to mitochondrial genes. The phylogenetic gene tree constructed by BI and ML methods recovered six geographic clades. Divergence time estimation indicated that the divergence of these six populations can be traced back to the late Pleistocene. We supposed that the geographic isolation led to the extreme population structure. According to this study and previous studies about troglobites living in this region, the subterranean habitats of the Yunnan-Guizhou Plateau may contain many organisms with similar genetic structures. The subterranean biodiversity in the karst area of Southern China needs to be re-evaluated and protected.
Alien species are a significant threat to natural ecosystems and human economies. Despite global efforts to address this challenge, the documented number of alien species is rapidly increasing worldwide. However, the magnitude of the impact of alien species may vary significantly across habitats. For example, some habitats are naturally less prone to biological invasions due to stringent abiotic and biotic characteristics, selecting for a limited number of introduced species possessing traits closely related to the native organisms. Subterranean ecosystems are quintessential examples of habitats with strong environmental filters (e.g. lack of light and scarcity of food), driving convergent adaptations in species that have successfully adapted to life in darkness. Despite these stringent environmental constraints, the number of records of alien species in subterranean ecosystems has increased in recent decades, but the relevant literature remains largely fragmented and mostly anecdotal. Therefore, even though caves are generally considered very fragile ecosystems, their susceptibility to impacts by alien species remains untested other than for some very specific cases. We provide the first systematic literature survey to synthesise available knowledge on alien species in subterranean ecosystems globally. This review is supported by a database summarising the available literature, aiming to identify gaps in the distribution and spread of alien invertebrate species in subterranean habitats, and laying the foundations for future management practices and interventions. First, we quantitatively assessed the current knowledge of alien species in subterranean ecosystems to shed light on broader questions about taxonomic biases, geographical patterns, modes of dispersal, pathways for introductions and potential impacts. Secondly, we collected species-specific traits for each recorded alien species and tested whether subterranean habitats act as ecological filters for their establishment, favouring organisms with pre-adaptive traits suitable for subterranean life. We found information on the presence of 246 subterranean alien species belonging to 18 different classes. The dominant alien species were invertebrates, especially insects and arachnids. Most species were reported in terrestrial subterranean habitats from all continents except Antarctica. Palaearctic and Nearctic biogeographic regions represented the main source of alien species. The main routes of introductions into the recipient country are linked to commercial activities (84.3% of cases for which there was information available). Negative impacts have been documented for a small number of case studies (22.7%), mostly related to increased competition with native species. For a limited number of case studies (6.1%), management strategies were reported but the effectiveness of these interventions has rarely been quantified. Accordingly, information on costs is very limited. Approximately half of the species in our database can be considered established in subterranean habitats. According to our results, the presence of suitable traits grants access to the stringent environmental filter posed by subterranean environments, facilitating establishment in the new habitat. We recommend that future studies deepen the understanding of invasiveness into subterranean habitats, raising public and scientific community awareness of preserving these fragile ecosystems.
The conservation of biodiversity is a central imperative of the 21st century. Subterranean ecosystems deliver critical nature's contributions to people and harbour a broad diversity of poorly-understood specialized organisms that are of interest from both a conservation and evolutionary perspective. However, the subterranean biome is still systematically overlooked in global biodiversity targets and conservation agendas. The main objective of this study was to assess how far subterranean biodiversity is represented in protected areas (Natura 2000 and Emerald networks) in two global hotspots of subterranean biodiversity (the Pyrenees and the Alps). For this, we used the most complete databases of terrestrial subterranean biodiversity known to us, i.e., leiodids (beetles) from the Pyrenees and spiders from the Alps, and identi ed priority areas in each region using both species richness and geographic rarity patterns. Our results show the incapacity of surface protected area networks to represent subterranean fauna, as more than 70 and 90% of the identi ed priority areas (and the 40 and 22% of the species) are not effectively covered by protected areas in the Pyrenees and the Alps, respectively. These ndings call for urgent policies and would be key to developing a coherent plan for subterranean biodiversity conservation within the European Biodiversity Strategy for 2030.
Lesvos is the third biggest island of Greece, part of the East Aegean Islands and located near the coastline of Anatolia peninsula. In contrast to other islands, such as Crete, Corfu, Samos and Rhodes, the island’s arthropod fauna was understudied and with many literature gaps and for this reason, extensive research was required. Samplings took place from April 2019 to January 2022 by using pitfall traps, collecting specimens by hand and collecting and isolating galls. At first, samplings with pitfall traps took place between 2nd March 2021 and 5th May 2021 from 28 sampling stations in the area of Pirgoi Thermis. In addition, arthropods were collected by hand throughout the island of Lesvos between April 2019 and January 2022. At the same time, from March 2021 to January 2022, gall samplings from various host plants took place and they were isolated in order to collect emerged adults, parasitoids and inquilines. Consequently, all collected arthropods were photographed with a camera and, then, the photos were uploaded on the online platform iNaturalist in order to fully record the place and the date of each observation. In total, 339 different arthropod species were recorded from 29 orders, from which 12 species are probably new for science and 20 species are new for Greece. Moreover, 33 alien and invasive arthropod species were collected as well as 16 endemic species. Finally, whether Lesvos is an area of endemism is under discussionas well as the value of studying the local fauna to the benefit of agriculturists, doctors and citizens.
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.
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 km ² and AOO < 2,000 km ² , meeting the thresholds for the inclusion in the threatened categories. Five species have a more widespread distribution (EOO > 20,000 km ² ), 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.
We investigated 11 pillbox bunkers in Dibër Municipality and one military tunnel near Tirana (Albania). We found 101 spider specimens belonging to 15 species, of which two species are reported for the first time from Albania: Leviellus thorelli (Ausserer, 1871) (Araneidae) and Meta bourneti Simon, 1922 (Tetragnathidae). This paper contributes new information on the diversity, distribution and natural history of spider fauna in the bunkers of Albania.
en This paper provides an overview on troglobitic springtails found in European caves, including a checklist at species level. The paper also reviews what is currently known about Collembola, which occur in caves of the most important mountain ranges in Europe. Only troglobitic species were included since many troglophiles were of uncertain ecological status. A total of 338 troglobitic species of Collembola is recorded from European caves, distributed across 12 families. Spain and France appear to host the highest richness of species, including endemics. From a biogeographic perspective, troglobitic species are unevenly distributed in Europe, especially in the most important mountain ranges, like the Alps, the Carpathians, the Pyrenees, the Caucasus, and other European mountains. Troglobitic springtails are far more abundant in temperate zones than in the tropics. Despite this, several genera of Collembola appear to be well represented, while some are poorly represented (or lacking) in European caves. Many advances in knowledge of subterranean springtails have been made, particularly in the description of new species. However, there are still major gaps in the knowledge of the biology, environmental requirements, and impacts on subterranean fauna. This paper highlights the need for further research and provides baseline data for such efforts.
Resumen
es En este trabajo se proporciona una visión general de los colémbolos troglobios encontrados en las cuevas europeas, incluyendo un listado a nivel de especie. También se revisa lo que hasta ahora se conoce sobre los colémbolos que se encuentran en las cuevas de las cadenas montañosas más importantes de Europa. Se incluyen solamente las especies troglobias ya que el estado ecológico de muchos troglófilos es dudoso. Se registran 338 especies troglobias de Collembola en cuevas europeas, distribuidas en 12 familias. España y Francia parecen albergar la mayor riqueza de especies, incluidas las endémicas. Desde una perspectiva biogeográfica, las especies troglobias se distribuyen de forma desigual en Europa, especialmente en las cadenas montañosas más importantes, como los Alpes, los Cárpatos, los Pirineos, el Cáucaso y otras montañas europeas. Los colémbolos troglobios son mucho más abundantes en las zonas templadas que en los trópicos. A pesar de esto, varios géneros de Collembola parecen estar bien representados, mientras que otros están poco representados (o no existen) en las cuevas europeas. Últimamente se han realizado muchos avances en el conocimiento de los colémbolos subterráneos, particularmente en la descripción de nuevas especies. Sin embargo, aún existen importantes lagunas sobre su biología, los requisitos ambientales y los impactos sobre la fauna subterránea. Este trabajo destaca la necesidad de realizar más investigaciones y aporta datos de referencia en este sentido.
We present the first record of Attulus saltator (O. Pickard-Cambridge, 1868) for Spain. The single male was collected in Blanes,
Catalonia. Additionally, we offer nine new records for the Spanish region of Galicia (province of Lugo) and seven for Catalonia (province
of Girona), one of which is Cepheia longiseta (Simon, 1881), which represents the first record of the family Synaphridae in the region of
Catalonia. Furthermore, a redescription of the male of Amaurobius occidentalis Simon, 1893 is provided.
The aims of the study were to investigate the Arthropoda fauna of Zindan Cave (Aksu, Isparta Province, Turkey) and to consider some ecological characteristics of the collected species such as feeding habits, cave-dwelling categories, and zone distributions in caves. Twenty-seven species of Arthropoda were determined between May 2015 and December 2016, from Zindan Cave. The species found belong to the following orders: nine to Collembola, six to Coleoptera, five to Araneae, two to Diplopoda, and one to Orthoptera, Trichoptera, Diptera, Trombidiformes and Isopoda. Nine species are endemic of Turkey. Traegaardhia distosolenidia Zacharda, 2010 (Acari: Rhagidiidae) and Folsomia asiatica Martynova, 1971 (Collembola: Isotomidae) are newly recorded for the fauna of Turkey. The presence of Heteromurus sexoculatus Brown, 1926 (Collembola: Entomobryidae) in Turkey is confirmed. Twenty-four of the 27 species were from the dark zone and half of the collected species (6 troglobites and 8 troglophiles) are ecologically adapted to cave ecosystems. Species can be divided into three groups according to trophic preferences: 15 scavengers, 10 predators, and two omnivores. Food habits, cave-dwelling categories, and zone distributions of collected species are discussed.
The Republic of Kosovo is the territory with the least known spider fauna within the
Balkans. The present list of spiders is based on all published records available to the authors and
also includes original unpublished data. The checklist comprises only 159 species belonging to 29
families and 108 genera. This low number is due to different reasons but mostly because the spiders
herein has never been studied in their entirety and because we excluded all the records from the
“Balkans”, “Yugoslavia”, “Serbia”, and the border mountain “Kopaonik”, for which cannot
certainly be argued that relate to Kosovo, so we worked only with reports, containing reliable data
on the spider fauna from there. The aim of this study is to presents annotated preliminary checklist
of the spiders of Kosovo with additional faunistic data.
Despite their ecological importance and diversity, spiders (Arachnida: Araneae) are underrepresented in conservation policies in comparison to other groups. We review all extant conservation tools focusing on spiders in Europe, highlighting general patterns, limitations, gaps, and future directions. We assembled a comprehensive online database reporting all available information concerning the legal protection and conservation status of 4,154 spider species. Existing international legislation has limited coverage, with only one species listed in the Bern Convention and EU Habitats Directive. At the national and subnational levels, 178 species are formally mentioned in the legislation of 19 European countries. Moreover, the International Union for Conservation of Nature (IUCN) includes assessments for 301 species worldwide, 164 of these threatened and eight native to Europe. In addition, spiders are mentioned in Regional Red Lists and Red Books in 28 out of 42 European countries considered in this review. Northern and Central European countries have the highest percentage of species assessed at the regional level in Red Lists and Red Books. The Mediterranean basin has the highest spider diversities in Europe but conservation efforts are lacking, both in terms of assessments and national or subna-tional legislation. Among European species, Dolomedes plantarius, Argyroneta aquatica and Eresus kollari are the most frequently mentioned in European conservation measures, possibly due to their ecological traits and their strict association with declining habitats. Considering the current threats to spiders in Europe, the protection of large areas of suitable habitat should be considered as the most effective approach to spider conservation.
Dysderid spiders are common in ground habitats, especially leaf litter, and are often found in caves (Chatzaki & Arnedo 2006). The genus Harpactea Bristowe currently includes 5 species in Serbia, i.e. Harpactea hombergi (Scopoli), Harpactea lepida (C.L. Koch), Harpactea pr. incerta Brignoli, Harpactea rubicunda (C.L. Koch), Harpactea saeva (Herman) (Deltshev et al. 2003). They are known from the territory of Europe, widespread mainly in central, southern and eastern parts (Platnick 2011). Harpactea complicata sp. nov. (male/female) is here described and illustrated. The new species was collected in South Western Serbia, Javor Mountain, also in caves. To clear the situation with Harpactea pr. incerta needs additional investigations.
Background
The North Atlantic archipelagos of Madeira and Selvagens present a unique biological diversity including, presently, 56 endemic spider species. Several recent projects provide valuable information on their distribution across most islands and habitats. To date, the only endemic spider assessed according to the IUCN Red List criteria is Hognaingens. The objective of this paper is to assess all remaining endemic species and advise on possible future conservation actions critical for the survival of endangered species.
New information
Seven species were found to have a continuing decline in either range or population size. Their decline can be mostly attributed to habitat destruction or degradation, invasive plant species that reduce quality of habitat, forest fires at high mountain regions and possible competition for resources from invasive congeners. The tetragnathid M.barreti is considered as possibly extinct due to the suspected impact of a competing species. Although most endemic spiders from the Madeira and Selvagens archipelagos have relatively low extinction risk due to the good condition and protection of the laurisilva forests where many live, there are a number of species requiring urgent attention and protection measures. These include all cave and mountain-restricted species as well as those threatened by competing congeners or invasive plants. Extending current protected areas, restoring original habitats of threatened species and the control of invasive taxa should remain a priority for species survival.
Over the last two decades there has been an exponential increase in the use of correlative species distribution models (SDMs) to address a variety of topics in ecology, biogeography, evolution, and conservation biology. Conversely, the use of these statistical methods to study the potential distribution of subterranean organisms has lagged behind, relative to their above-ground (epigean) counterparts. The reason for this is possibly related to a number of peculiarities of subterranean systems, which pose important limits, but also opportunities, for these correlative models. The aim of this forum is to explore the caveats that need to be made when generalizing these statistical techniques to caves and other subterranean habitats. We focus on the typical bias in spatial datasets of cave-dwelling species, and provide advice for selecting the model calibration area. In parallel, we discuss the potential use of different large scale surface variables to represent the subterranean condition. A more widespread adoption of these statistical techniques in subterranean biology is highly attractive and has great potential in broadening our knowledge on a variety of ecological topics, especially in the fields of climate change and biodiversity conservation. Their use would especially benefit the study of the biogeographic patterns of subterranean fauna and the impact of past and future climate change on subterranean ecosystems.
Mygalomorph spiders are rarely found in caves and most of the records appear as accidental or restricted to small populations. The present study took place in iron formations in Carajás region, southeastern Pará state, eastern Brazilian Amazon rainforest. Each cave was sampled twice: dry season (from May to October) and wet season (from November to April). Of the 242 caves sampled, we found mygalomorphs in 98 (40%). The survey yielded 254 specimens, 223 (87.8%) juveniles. Of the 14 species recorded, nine were represented by adults: Dolichothele tucuruiense (Guadanucci, 2007), Hapalopus aymara Perdomo, Panzera & Pérez-Miles, 2009, Acanthoscurria geniculata (CL Koch, 1841), Theraphosa blondi (Latreille, 1804), Nhandu coloratovillosus (Schmidt, 1998), Fufius minusculus Ortega, Nagahama, Motta & Bertani, 2013, and three new are species described here: Guyruita metallophila n. sp., Hapalopus serrapelada n. sp., and Idiops carajas n. sp. Five other “morphospecies” were represented by juveniles only: Ummidia sp. (Ctenizidae), Paratropis sp. (Paratropididae), Bolostromus sp. (Cyrtaucheniidae), Diplura sp. (Dipluridae), and Idiophtalma sp. (Barychelidae). The high number of juveniles suggests two alternatives: some species could be using the hypogean environment as reproductive shelter; the hypogean environment is used as a refuge by immatures. Two species appear to be troglophiles: G. metallophila and H. aymara.
https://zoobank.org/urn:lsid:zoobank.org:pub:C92F29CB-4B9C-402D-8AA6-F8251DE49529.
The threshold zones between the epigean and hypogean environments are generally characterized by less harsh ecological conditions than deep subterranean habitats, and usually support a greater abundance of organisms. Transitional habitats such as these should be more easily colonised by alien species, especially by those possessing exaptations suitable for subterranean life. In spite of this, few studies have been conducted to unravel the ecological dynamics between native and alien species in the habitats situated at the epigean/hypogean interface. A unique test case is offered by cave-dwelling Meta orb-weaver spiders in Great Britain (Araneae: Tetragnathidae). One species, M. menardi, is a widespread native, whilst M. bourneti is believed to be a recently introduced (1940s) species, that has since become established in the south-eastern part of the country. Species distribution models (SDM) were used to predict current and future habitat suitability for the two species, generating hypotheses regarding their distribution in different global warming scenarios. Model projections indicate that the two species respond to similar environmental variables. Seasonal temperature variations at the surface and elevation are the main factors explaining the distribution of both species, whereas annual precipitation, daily temperature range and limestone distribution contributed little to the model performance. It is predicted that due to climate change, there will be poleward shifts in the ranges of both species. However, the native species M. menardi will primarily be able to exploit suitable areas which will appear northward to their current distribution, and M. bourneti will colonise empty niches left available by its congeneric. The analytical framework employed in this paper may be easily adapted to other subterranean systems and species, stimulating future studies focusing on the distribution of native and alien species in extreme environments.
Caves and cave-dwelling biota have fascinated scientists for centuries. Nevertheless, there is a considerable lack of information on subterranean realms and the ecosystems they host. Nematoda, for example, is a group of invertebrates that plays an important role in the functioning of epigean ecosystems, but whether or not the same is true for subterranean ecosystems remains unknown. For this reason it was decided to conduct an in-depth review of all reports related to cave-dwelling nematodes in order to provide a sound basis for future studies. A literature survey of 41 scientific works from over the last 138 years revealed 295 unique taxa reported from 78 different cave systems. The historical trends in cave nematology, peculiar findings from important studies and an ecological classification system are discussed. Lastly, the trophic distribution of the reported taxa is presented, whilst nematodes from other (non-cave) subterranean environments are also considered.
World experts of different disciplines, from molecular biology to macro-ecology, recognize the value of cave ecosystems as ideal ecological and evolutionary laboratories. Among other subterranean taxa, spiders stand out as intriguing model organisms for their ecological role of top predators, their unique adaptations to the hypogean medium and their sensitivity to anthropogenic disturbance. As the description of the first eyeless spider (Stalita taenaria), an array of papers on subterranean spider biology, ecology and evolution has been published, but a comprehensive review on these topics is still lacking. We provide a general overview of the spider families recorded in hypogean habitats worldwide, we review the different adaptations of hypogean spiders to subterranean life, and we summarize the information gathered so far about their origin, population structure, ecology and conservation status. Finally, we point out the limits of the knowledge we currently have regarding hypogean spiders, aiming to stimulate future research.
Subterranean organisms always attracted the attention of humans using caves with various purposes, due to the strange appearance of several among them and life in an environment considered extreme. According to a classification based on the evolutionary and ecological relationships of these organisms with subterranean habitats, first proposed by Schiner in 1854 and emended by Racovitza in 1907, three categories have been recognized: Troglobites, troglophles and trogloxenes. The Schiner-Racovitza system has been discussed, criticized, emended, the categories have been redefined, subdivided, original meanings have changed, but it is used until now. Herein we analyze in a conceptual framework the main ecological classifications of subterranean organisms, from Schiner to Trajano, in 2012, so far the last author to introduce a relevant conceptual change on the categories definitions, incorporating the source-sink population model. Conceptual inconsistencies are pointed, especially with regards to the generally ill-defined trogloxene category, and the correspondence between categories according to the original sense and in alternative classifications is discussed. Practical criteria for distinction between these categories and difficulties for their application are presented. The importance of rightly classifying subterranean populations according to the Schiner-Racovitza system for conservation of these fragile and mostly threatened habitats is discussed.
Animal life in caves has fascinated researchers and the public alike because of the unusual and sometimes bizarre morphological adaptations observed in numerous troglobitic species. Despite their worldwide diversity, the adaptations of cave millipedes (Diplopoda) to a troglobitic lifestyle have rarely been examined. In this study, morphological characters were analyzed in species belonging to four different orders (Glomerida, Polydesmida, Chordeumatida, and Spirostreptida) and six different families (Glomeridae, Paradoxosomatidae, Polydesmidae, Haplodesmidae, Megalotylidae, and Cambalopsidae) that represent the taxonomic diversity of class Diplopoda. We focused on the recently discovered millipede fauna of caves in southern China. Thirty different characters were used to compare cave troglobites and epigean species within the same genera. A character matrix was created to analyze convergent evolution of cave adaptations. Males and females were analyzed independently to examine sex differences in cave adaptations. While 10 characters only occurred in a few phylogenetic groups, 20 characters were scored for in all families. Of these, four characters were discovered to have evolved convergently in all troglobitic millipedes. The characters that represented potential morphological cave adaptations in troglobitic species were: (1) a longer body; (2) a lighter body color; (3) elongation of the femora; and (4) elongation of the tarsi of walking legs. Surprisingly, female, but not male, antennae were more elongated in troglobites than in epigean species. Our study clearly shows that morphological adaptations have evolved convergently in different, unrelated millipede orders and families, most likely as a direct adaptation to cave life.
Subterranean ecosystems present ideal opportunities to study mechanisms underlying responses to changes in climate because species within them are often adapted to a largely constant temperature. We have characterized the thermal conditions of caves in the Western Alps, and related these hypogean climate data to the occurrence of Troglohyphantes spiders (Araneae, Linyphiidae). Our data indicated that present distributions reflect Pleistocene glaciation events and also pointed to specific responses as a consequence of changes in temperature. Constant temperatures recorded inside caves provide an approximation of the mean annual temperature outside, thus we extended the results to a regional scale. We used ecological niche modeling to predict habitat suitability both in the Pleistocene and under future global warming scenarios. These analyses pointed toward a future decline in habitat suitability for subterranean spiders and the potential extinction of the most restricted endemic species. When compared with other species that live in confined habitats such as islands and mountains, we expect cave species to be as much, if not more, vulnerable to climate change.
This paper describes and illustrates a new genus and a new species belonging to the family Nesticidae based on morphology and supported by molecular data. The new genus, Kryptonesticus gen. nov., groups eight species spread from Bulgaria and Turkey to Croatia, including Montenegro, Bosnia and Herzegovina and Crete. As a result, seven new combinations are proposed: K. eremita (Simon, 1879) comb. nov., K. arenstorffi (Kulczyński, 1914) comb. nov., K. fagei (Kratochvíl, 1933) comb. nov., K. beroni (Deltshev, 1977) comb. nov., K. beshkovi (Deltshev, 1979) comb. nov., K. henderickxi (Bosselaers, 1998) comb. nov. and K. dimensis (López-Pancorbo, Kunt & Ribera, 2013) comb. nov., all ex Nesticus. Kryptonesticus deelemanae gen. et sp. nov. is described on the basis of both sexes and its phylogenetic relationships with closely related species are discussed based on morphological and molecular data (the cox1, rrn and H3 genes). In addition, the species of this new genus (except for K. eremita) are clear candidates for protection: they have highly restricted ranges and some of them show a high degree of adaptation to the subterranean environment.
Pimoa delphinica Mammola, Hormiga & Isaia, 2016 is a troglophile araneoid spider endemic of the high Varaita valley (Western Alps, Province of Cuneo, NW Italy). In spite of relatively intense field research and examination of museum collections, the species is restricted to ten localities, thus showing a reduced extent of occurrence (EOO; 26 km) and area of occupancy (AOO; 12 km). Although the habitat and the subpopulations of P. delphinica are not currently threatened, the species is potentially exposed due to its extremely narrow geographic distribution range, subdivision in subpopulations and low dispersal capacity.
Pimoidae is a small family of araneoid spiders, hitherto represented in Europe by two species with disjunct distribution in the Alps and in the Cantabrian Mountains of northern Spain. Here we report the description of two additional European species of Pimoa, discovered within the range of the only former alpine species, P. rupicola: P. graphitica sp. nov. and P. delphinica sp. nov. The new species are distinguished from the latter by genitalic characters as well as by molecular characters. On the basis of the re-examination of old and recent abundant material collected in caves and other subterranean habitats, we revise the distribution patterns of the genus Pimoa in the Alps and outline the species distribution ranges. Molecular data suggest the existence of gene flow between populations of the two new species when in sympatry. The different species probably originated in the alpine region as a result of range contractions following dramatic climatic changes in the Alps since the mid Miocene. We interpreted the present-day overlapping distribution in light of a possible postglacial expansion. Finally, we provide insights on the natural history and life cycles of the new species and discuss their phylogenetic relationships within Pimoidae.
Background and purpose: The approaches to the impact of climate
change on carabid beetles are highly differentiated. We review here most
studies that tried to compare past situations with present time conditions
that may concern ground beetle species, populations and phenology, area
and upslope shifts. Both epigean and hypogean domains have taken in con-
sideration and, whenever possible, all time scales addressed by the authors.
Materials and methods: The methods adopted vary in relation to the
time scale. Geo-period studies cover the largest time lapse, and compare
Plio-Pleistocene fossil species assemblages with present ones. Further ap-
proaches concern long-term observations based on museum materials and
the response of carabids to climate or man-conditioned habitat changes.
Primarily devoted to the understanding of climate change impacts on com-
munities are the space-for-time and the time-per-time approach, that imply
the comparison of year samples collected in the same site after some decades.
Other long-term studies are devoted to the phenology changes. The impact
on cave dwelling carabids has studied by recording the new taxa discoveries
during the last two centuries.
Results: Geo-period studies reveal strong area expansions restrictions or
local extinctions of presently living ground beetles. The old idea of a stability
of taxonomic status of such beetles seems to give way to a more dynamic vision
of speciation events during the Quaternary, as suggested by recent population
genetics studies on Trechines and some Carabus species. Long term studies
based on museum materials reveal population and area declines of large and
brachypterous species of open lands, but the climate responses are difficult to
disentangle from anthropogenous habitat and landscape modifications. The
space-for-time approach focuses especially glacier forelands but seems promis-
ing also in forecasting species declines starting from climate gradients in forests.
The time-for-time (time lapse) monitoring of ground beetle assemblages has
adopted so far in Mediterranean mountains and Dolomites. It reveals that
in the last three decades a strong uphill shift has observed especially in open
land assemblages. In the Dolomites local extinctions and decline of species’
diversity has recorded around or above the tree line. The long-term monitor-
ing of carabids in defined habitats provides extremely rich databases that can
be useful in explain the relationships between climate change, population
decline or increase and species phenology. Hypogean obligate troglobitic cara-
bids revealed to be extremely sensitive to global warming, as testified by the
continuous appearance of new highly evolved taxa that concentrate in the
periods of fastest temperature rise of the last century.
Conclusions: Epigean and hypogean carabid beetles are excellent and
„multitask” indicators of climate change. Moreover, their response time to
climate changes seems to be shorter than for plants.
Forty-three new species of Nesticidae are described from China, Indonesia, Philippines, Singapore,
Thailand, Vietnam and Madagascar, and two new junior synonyms are suggested. A new genus, Spele-
oticus gen. n., is described with Nesticus navicellatus Liu & Li, 2013 as the type species, and four species
are transferfed from Nesticus, i.e., Speleoticus globosus (Liu & Li, 2013), comb. n., S. libo (Chen & Zhu,
2005), comb. n., S. navicellatus (Liu & Li, 2015), comb. n. and S. uenoi (Yaginuma, 1972), comb. n.
The new species described in this paper belong to four genera and are: Hamus cornutus sp. n. (♂♀),
H. kangdingensis sp. n. (♂), H. luzon sp. n. (♀), H. mangunensis sp. n. (♂), Nescina kohi sp. n. (♂♀),
Nesticella baiseensis sp. n. (♂♀), N. baobab sp. n. (♂), N. caeca sp. n. (♂♀), N. chongqing sp. n. (♀), N.
dazhuangensis sp. n. (♂♀), N. fuliangensis sp. n. (♂♀), N. gazuida sp. n. (♀), N. gongshanensis sp. n. (♀),
N. griswoldi sp. n. (♂♀), N. hongheensis sp. n. (♂♀), N. huomachongensis sp. n. (♂♀), N. jingpo sp. n.
(♀), N. kaohsiungensis sp. n. (♂♀), N. lisu sp. n. (♂♀), N. liuzhaiensis sp. n. (♀), N. nandanensis sp. n.
(♂♀), N. phami sp. n. (♂♀), N. potala sp. n. (♀), N. qiaoqiensis sp. n. (♀), N. qiongensis sp. n. (♂♀),
N. robusta sp. n. (♂♀), N. rongtangensis sp. n. (♂), N. sanchaheensis sp. n. (♂♀), N. sulawesi sp. n. (♀),
N. sumatrana sp. n. (♂), N. tibetana sp. n. (♂♀), N. vanlang sp. n. (♀), N. wanzaiensis sp. n. (♂♀),
N. xiongmao sp. n. (♂♀), N. xixia sp. n. (♂♀), N. yanbeiensis sp. n. (♂♀), N. yao sp. n. (♀), N. zhi-
yuani sp. n. (♂♀), Pseudonesticus dafangensis sp. n. (♂♀), P. miao sp. n. (♂♀), P. spinosus sp. n. (♂♀), P.
wumengensis sp. n. (♀), P. ziyunensis sp. n. (♂♀). Nesticella inthanoni (Lehtinen & Saaristo, 1980), syn. n. is synonymised with N. mollicula (Thorell, 1898); N. taiwan Tso & Yoshida, 2000, syn. n. is synonymised
with N. odonta (Chen, 1984). The female of Nesticella connectens Wunderlich, 1995, so far unknown, is
described and recorded from Thailand. Nesticidae are reported from Madagascar for the first time. Nesti-
cella nepalensis (Hubert, 1973) is recorded for the first time from China. Types of Nesticella odonta (Chen,
1984), N. songi Chen & Zhu, 2004 and N. yui Wunderlich & Song, 1995 are re-examined and photo-
graphed. The entire genus Nesticella is reviewed, and four species groups are recognised. DNA barcodes of
the new species are obtained to confirm their correct identifications.
Endemism, which is typically high on islands and in caves, has rarely been studied in the cave entrance ecotone. We investigated the endemism of the spider genus Uthina at cave entrances. Totally 212 spiders were sampled from 46 localities, from Seychelles across Southeast Asia to Fiji. They mostly occur at cave entrances but occasionally appear at various epigean environments. Phylogenetic analysis of DNA sequence data from COI and 28S genes suggested that Uthina was grouped into 13 well-supported clades. We used three methods, the Bayesian Poisson Tree Processes (bPTP) model, the Bayesian Phylogenetics and Phylogeography (BPP) method, and the general mixed Yule coalescent (GMYC) model, to investigate species boundaries. Both bPTP and BPP identified the 13 clades as 13 separate species, while GMYC identified 19 species. Furthermore, our results revealed high endemism at cave entrances. Of the 13 provisional species, twelve (one known and eleven new) are endemic to one or a cluster of caves, and all of them occurred only at cave entrances except for one population of one species. The only widely distributed species, U. luzonica, mostly occurred in epigean environments while three populations were found at cave entrances. Additionally, eleven new species of the genus are described.
The ground-dweller spider genus Dysdera shows very high species richness on the oceanic archipelago of the Canary Islands, providing one of the most outstanding examples of island radiation among spiders, only paralleled by Tetragnatha spiders on the Hawaiian archipelago. A georeferenced database of the 48 Dysdera species occurring in the Canary Islands was assembled to facilitate ongoing and future research on this remarkable lineage. All species are endemic to the archipelago except for the cosmopolitan Dysderacrocata. The dataset consists of 794 distributional records documented from 1971 to 2015, each locality being represented only once per species. Distribution maps are provided for each species, along with basic diversity and distribution information. The database and geographical maps included in this article stand for the most updated, accurate and complete information on the distribution of the spider genus Dysdera in the Canary Islands.
The term Milieu Souterrain Superficiel (MSS) has been used since the early 1980s in subterranean biology to categorize an array of different hypogean habitats. In general terms, a MSS habitat represents the underground network of empty air-filled voids and cracks developing within multiple layers of rock fragments. Its origins can be diverse and is generally covered by topsoil. The MSS habitat is often connected both with the deep hypogean domain---caves and deep rock cracks---and the superficial soil horizon. A MSS is usually characterized by peculiar microclimatic conditions, and it can harbor specialized hypogean, endogean, and surface-dwelling species. In light of the many interpretations given by different authors, we reviewed 235 papers regarding the MSS in order to provide a state-of-the-art description of these habitats and facilitate their study. We have briefly described the different types of MSS mentioned in the scientific literature (alluvial, bedrock, colluvial, volcanic, and other types) and synthesized the advances in the study of the physical and ecological factors affecting this habitat---i.e., microclimate, energy flows, animal communities, and trophic interactions. We finally described and reviewed the available sampling methods used to investigate MSS fauna.
Turinyphia cavernicola Wunderlich, 2008 is one of the two endemic cavernicolous spiders known from the Azorean archipelago (Portugal). In spite of relatively intensive searches, this species is known from only three caves, but with a single sustainable subpopulation restricted to the Algar do Carvão show cave. With an observed extent of occurrence (EOO) of 2 km² and an area of occupancy (AOO) around 0,75 km², it presents a restricted range, severe fragmentation, and continuing decline in EOO, AOO, habitat quality, number of locations and subpopulations caused by the ongoing threat from pasture intensification and tourist activities in the cave systems.
One of the most challenging fauna to study in situ is the obligate cave fauna because of the difficulty of sampling. Cave-limited species display patchy and restricted distributions, but it is often unclear whether the observed distribution is a sampling artifact or a true restriction in range. Further, the drivers of the distribution could be local environmental conditions, such as cave humidity, or they could be associated with surface features that are surrogates for cave conditions. If surface features can be used to predict the distribution of important cave taxa, then conservation management is more easily obtained. We examined the hypothesis that the presence of major faunal groups of cave obligate species could be predicted based on features of the earth surface. Georeferenced records of cave obligate amphipods, crayfish, fish, isopods, beetles, millipedes, pseudoscorpions, spiders, and springtails within the area of Appalachian Landscape Conservation Cooperative in the eastern United States (Illinois to Virginia and New York to Alabama) were assigned to 20 x 20 km grid cells. Habitat suitability for these faunal groups was modeled using logistic regression with twenty predictor variables within each grid cell, such as percent karst, soil features, temperature, precipitation, and elevation. Models successfully predicted the presence of a group greater than 65% of the time (mean = 88%) for the presence of single grid cell endemics, and for all faunal groups except pseudoscorpions. The most common predictor variables were latitude, percent karst, and the standard deviation of the Topographic Position Index (TPI), a measure of landscape rugosity within each grid cell. The overall success of these models points to a number of important connections between the surface and cave environments, and some of these, especially soil features and topographic variability, suggest new research directions. These models should prove to be useful tools in predicting the presence of species in understudied areas.
For management strategies in the context of global warming, accurate predictions of species response are mandatory. However, to date most predictions are based on niche (bioclimatic) models that usually overlook biotic interactions, behavioral adjustments or adaptive evolution, and assume that species can disperse freely without constraints. The deep subterranean environment minimises these uncertainties, as it is simple, homogeneous and with constant environmental conditions. It is thus an ideal model system to study the effect of global change in species with poor dispersal capabilities. We assess the potential fate of a lineage of troglobitic beetles under global change predictions using different approaches to estimate their thermal niche: bioclimatic models, rates of thermal niche change estimated from a molecular phylogeny, and data from physiological studies. Using bioclimatic models, at most 60% of the species were predicted to have suitable conditions in 2080. Considering the rates of thermal niche change did not improve this prediction. However, physiological data suggest that subterranean species have a broad thermal tolerance, allowing them to stand temperatures never experienced through their evolutionary history. These results stress the need of experimental approaches to assess the capability of poor dispersal species to cope with temperatures outside those they currently experience.
The spider family Mysmenidae is reported from Tibet for the first time. Two new species, Chanea voluta sp. n. (male and female) and Mysmena lulanga sp. n. (male and female) are found in eastern Tibet in high altitude. Morphological descriptions, diagnoses and comparative photos are provided for the two new species.
Three new species of the spider genus Leptonetela collected from caves in Greece are described: L. arvanitidisi sp. n. (male & female), L. paragamiani sp. n. (male & female) and L. penevi sp. n. (male & female). Detailed illustrations of the new species are provided. DNA barcodes were obtained for future use.
Spiders occurring in soils and fissured rocks were investigated using pipe traps. Four microphthalmic species, namely Hahnia microphthalma, Porrhomma egeria, P. microps and P. cambridgei were collected. Hahnia microphthalma is reported from the Czech Republic for the first time. The importance of collecting material by complex pipe traps (consisting of a perforated pipe and a set of removable cups) to record the depth distribution of spiders in subterranean habitats is stressed. The importance of the soil and fissure network formed by sandy marlite bedrock and of alluvial soils for the life of subterranean spiders is documented.
The unique morphology of cave animals has interested biologists at least since the time of Lamarck. After a number of non-adaptive explanations for the morphology of cave animals, especially with respect to eye and pigment loss, a neo-Darwinian explanation, emphasizing constructive morphological changes rather than losses, was put forward by Barr, Christiansen, and Poulson in the 1960’s. Emphasizing convergent evolution (troglomorphy), this paradigm has recently been challenged, with evidence of divergent selection and the widespread occurrence of morphologically similar species in other subterranean habitats, including ones with abundant food and relatively strong environmental fluctuations. An emerging paradigm emphasizes the central role of darkness in convergent evolution, and the role of habitat size and interspecific competition in divergent evolution of subterranean species.
Sprememba paradigme o evoluciji jamskega življenja
Edinstvena morfologija jamskih živali, predvsem v povezavi z izgubo oči in pigmenta, je zanimala biologe že od časa Lamarcka. Kasneje so bile predstavljene tudi druge, t.im. neodarvinistične razlage, s poudarkom na konstruktivnih morfoloških spremembah namesto izgubah, povzete s strani Barra, Christiansena in Poulsona v 60. letih 20. stoletja. Z novejšimi raziskavami o divergentni selekciji in razširjeni prisotnosti morfološko podobnih vrst v drugih podzemeljskih habitatih, vključno s tistimi, kjer je veliko hrane in z relativno velikimi okoljskimi nihanji, se je zamajala paradigma, temelječa na konvergentni evoluciji (troglomorfija). Novo nastajajoča paradigma poudarja osrednjo vlogo teme v konvergentni evoluciji, ter vlogo velikosti habitata in medvrstne kompeticije v divergentni evoluciji podzemeljskih vrst.
David C. Culver & Tanja Pipan: Climate, abiotic factors, and the evolution of subterranean life Climate, and more generally the physical conditions in caves and other subterranean habitats have a profound influence on the biota. At longer time scale (centuries), climate change can force and/or isolate species in subterranean habitats. Not only Pleistocene climate changes, but earlier ones as well, such as the Messinian salinity crisis were important in this regard. While many speleobiologists assume that caves are nearly constant environmentally and with scarce organic carbon, this is not the case, especially in non-cave subterranean habitats. Many shallow subterranean habitats, such as epikarst, seepage springs, and talus harbor highly modified organisms, ones without eyes and pigment and with elongated appendages. Yet these habitats are highly variable with respect to temperature and other environmental factors, and often have high levels of organic carbon. Overall, the role of these shallow subterranean habitats in the evolution and biogeography of subterranean species may be crucial. On smaller spatial scales, environmental differences, such as differences in chemistry of epikarst water, may be important in allowing large numbers of species to coexist.
The spider family Symphytognathidae has never been recorded from Europe, being mostly present in tropical regions. Different collecting trips to a cave system in Portugal revealed several specimens of a new species of Symphytognathidae here described, Anapistula ataecina sp. n. This is one of the smallest spiders described to date. The species is almost exclusively known from the Frade Cave System in Portugal which is partly endangered by limestone quarries. No males were found to date despite many collecting trips to the caves during more than three years. Parthenogenesis could therefore be responsible for the species reproduction. Its web, with a sheet-like structure, seems atypical for the family and for the genus. Details on the eggsacs and spinneret morphology are also given.
Caves and other subterranean habitats with their often strange (even bizarre) inhabitants have long been objects of fascination, curiosity, and debate. The question of how such organisms have evolved, and the relative roles of natural selection and genetic drift, has engaged subterranean biologists for decades. Indeed, these studies continue to inform the general theory of adaptation and evolution. Subterranean ecosystems generally exhibit little or no primary productivity and, as extreme ecosystems, provide general insights into ecosystem function. The Biology of Caves and other Subterranean Habitats offers a concise but comprehensive introduction to cave ecology and evolution. Whilst there is an emphasis on biological processes occurring in these unique environments, conservation and management aspects are also considered. The monograph includes a global range of examples from more than 25 countries, and case studies from both caves and non-cave subterranean habitats; it also provides a clear explanation of specialized terms used by speleologists. This accessible text will appeal to researchers new to the field and to the many professional ecologists and conservation practitioners requiring a concise but authoritative overview. Its engaging style will also make it suitable for undergraduate and graduate students taking courses in cave and subterranean biology. Its more than 650 references, 150 of which are new since the first edition, provide many entry points to the research literature.
AimTo describe the distribution of two congeneric species of cave-dwelling spiders under current climatic conditions and future warming scenarios, keeping into account their bioclimatic requirements, dispersal capacity and competition dynamics. LocationEurope and North Africa. Methods
We gathered detailed occurrence data for Meta menardi and M. bourneti (Araneae: Tetragnathidae) and modelled their distribution across their bioclimatic range. We evaluated the breadth of the bioclimatic niche of the two species and assessed the potential overlap between their multidimensional Hutchinsonian hypervolumes. We compared these results with their current distribution maps obtained by species distribution modelling (SDMs) keeping into account dispersal and competition dynamics, and we projected future trends of distribution according to different climatic scenarios. ResultsThe overall size of the niche hypervolumes of the two species was very similar and highly overlapping. However, when accounting for competition in the model, the predicted ranges of the two species showed only minor areas of overlap. On the base of SDM analysis, we predicted a significant poleward shift in the distribution of both species, with the appearance of a new contact area especially in Central Europe, southern UK, France and northern Spain. Main conclusionsInterspecific competition keeps the current ranges of the two European Meta species apart, although their fundamental bioclimatic niches are very similar. Rapid poleward shifts of the two species are likely due to the high dispersal capacity of the two spiders. The case of Meta spiders exemplifies rapid shift in range of distribution associated with global warming, which are unlikely to occur in the case of most cave-dwelling arthropods.
The theraphosine genus Hemirrhagus Simon 1903 is revised based on the examination of the type specimens and additional material collected in Mexico. Eight species were redescribed and illustrated. The males of Hemirrhagus ocellatus, Hemirrhagus papalotl, and Hemirrhagus stygius, formerly unknown, are described for the first time. Five new species were recognized and are newly described and illustrated. Hence, Hemirrhagus comprises 21 valid species, all endemic to Mexico. All species are keyed and mapped. New taxonomic features are included in the descriptions and different types of stridulatory organs are described for the first time for the genus. It is reported for the first time that Hemirrhagus is the only known Theraphosinae that lays fixed egg-sacs. Hemirrhagus embolulatus sp. nov. is described as the only known Hemirrhagus that possesses embolus keels present in other Theraphosinae genera. Information on species habitat and reproduction are included. © 2014 The Linnean Society of London
Stony debris represent a typical island biotope in the C European landscape. This abrupt contrast between the temperature and moisture conditions on the surface and those in the inner zone is typical. Thermophilous and psychrophilous species occur simultaneously in bare insolated debris. Both the rare, relict species and the common, opportune species live side by side in debris. Amongst the most noteworthy arachnological finds in Czechoslovakia in recent years are the glacial relicts Bathyphantes eumenis and Wubanoides longicornis. -from Author
With 128 described species and five subspecies, the spider genus Troglohyphantes (Araneae, Linyphiidae) is a remarkable example of species diversification in the subterranean environment. In this paper, we conducted a systematic revision of the Troglohyphantes species of the Italian Alps, with a special focus on the Lucifuga complex, including the description of two new species (T. lucifer sp. nov. and T. apenninicus n. sp.). In addition, we provided new diagnostic drawings of the holotype of T. henroti (Henroti complex) and established three new synonymies within the genus. The molecular analysis of the animal DNA barcode confirms the validity of this method of identification of the Alpine Troglohyphantes and provides additional support for the morphology based species complexes. Finally, we revised the known distribution range of additional Troglohyphantes species, as well as other poorly known alpine cave-dwelling spiders.
Troglohyphantes lucifer http://www.zoobank.org/urn:lsid:zoobank.org:act:AB4B644A-D18A-4EB7-99E7-B5117AED5389
Troglohyphantes apenninicus http://www.zoobank.org/urn:lsid:zoobank.org:act:3C86E78E-7145-4F0C-803E-2A909201579C
Four new species are described in a previously monotypic genus Pinelema Wang et Li, 2012. P. curcici sp. nov. (male and female), P. huobaensis sp. nov. (male and female), P. yaosaensis sp. nov. (male and female) from Yunnan, and P. xiushuiensis sp. nov. (male and female) from Guangxi in southern China.
The subterranean environment has a set of unique characteristics, including low thermic variation, high relative humidity, areas with total absence of light and high dependence on nutrient input from the epigean environment. Such characteristics promote distinct ecological conditions that support the existence of unique communities. In this work, we studied seven caves in the Presidente Olegário municipality, Minas Gerais state, Southeast Brazil, to determine their richness of predatory species, to understand how they are spatially distributed in the cave and whether their distribution is influenced by competition and/or predation. We carried out five surveys of the caves, with each cave divided into sampling plots. We collected fauna within the plots using a manual search method. The collected animals were fixed in 70% ethanol for later identification. We performed a canonical correspondence analysis to verify the spatial distribution and substrate preference of each species, and selected five species for agonistic interaction testing in the laboratory. We found a great richness of predators in the caves, with 79 species distributed among 22 families of spiders, five families of pseudoscorpions, three families of chilopods, two families of opilionids and one family each of scorpions and heteroptera. Spiders were the most species diverse and abundant of all arthropods we found in the caves. We recorded evidence of competition among some pairs of species but, in general, the spatial distribution of the predatory community in the interior of the caves seems to be unrelated to interspecies competition. The laboratory pairings support our field observations that most species merely share space, rather than exhibiting aggressive or predatory behaviour.
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Eight new species of Telema collected from caves in Guangxi (China) are described and illustrated: T. adunca sp. nov., T. biyunensis sp. nov., T. cordata sp. nov., T. cucurbitina sp. nov., T. mikrosphaira sp. nov., T. renalis sp. nov., T. yashanensis sp. nov. and T. zonaria sp. nov. All species have a clearly pigmented body, six eyes, and relatively short legs. They differ from congeners and each other in the detailed structure of the spermatheca and male palp.
Diagnostic characters of Stenochilus and Colopea are reviewed and corrected. Nine species of Colopea are regarded valid, including 6 new species C. malayana (Malayan peninsula), C. unifoveata (Borneo) , C. silvestris (Papua New Guinea), C. xerophila (Papua New Guinea), C. romantica (Bali & Sumatra), and C. virgata (Central Thailand & Vietnam). The range of Stenochilidae extends from NW India to Fiji. All species of Stenochilus prefer xerophilic conditions in open habitats, and a few species of Colopea are ecologically similar, but most Colopea spp. are regularly found in the leaf litter of tropical forests.-from Author
The cave dwelling spider Porrhomma rosenhaueri (L. Koch) is unique to the British fauna as it is considered to be the only species of troglobiontspider present. This spider has a very limited distribution in the UK and is known only from two cave sites, both of which are in South Wales: Ogof y Ci near Merthyr Tydfil and Lesser Garth Cave near Cardiff. Monitoring populations of cavemicolous species is very difficult to achieve and the aim of this survey was to set up a Common Standards Monitoring (CSM) protocol thatwould allow meaningful data to be compiled as to the size and viability of the spider population in the Lesser Garth Cave complex. A defined transect of the main passage in the cave was surveyed, carefully searching the accessible parts of the passage for both live spiders and webs in good condition but with no obvious spider present. The first visit was made in September 2009 and 17 live spiders were recorded, removing one to confirm identification. During a follow up survey in December 2009 only 6 live spiders were recorded. Possible reasons for this are discussed in this paper. The overall impression is that, despite living next to an active quarry, the spider Porrhomma rosenhaueri is still well-established in the Lesser Garth Cave. The cave itself has a rich diversity of cave associated species and is thus an important site for speleobiology in the UK.
The material included 20 dipteran, 11 beetle, 10 springtail, 6 spider, 4 stonefly, 3 cryptostigmatid and 2 prostigmatid mite, 1 mayfly and 1 caddisfly species. Several species of insects which as larvae prefer cold, fast-running rivers occurred in this cave, many of them in great numbers. -from Authors
Two new species of the spider family Leptonetidae from caves of Balkan Peninsula are diagnosed, described, and illustrated, i.e., Cataleptoneta lingulata sp. nov. from Northern Dalmatia, Croatia, and Cataleptoneta semipinnata sp. nov. from Island Kythira, Greece.