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Diplurans are among the earliest hexapods in the geological record. These primitively wingless relatives of insects are infrequently encountered despite being pervasive in soil habitats. Two groups have disparate adaptations for hunting, one mechanical and the other chemical, in Japygoidea and Projapygoidea, respectively. Here, we report three genera (two new) and four species of fossil Projapygidae preserved in Mesozoic and Cenozoic ambers: Electroprojapyx alchemicus gen. et sp. nov. in mid-Cretaceous Myanmar amber, and Symphylurinopsis punctatus gen. et sp. nov. and two species (unnamed) of Symphylurinus in Miocene Dominican amber. The exceptionally preserved specimens possess cerci that are morphologically specialized for expelling a glandular substance on prey. The new Cretaceous E. alchemicus is a stem group to all living species of the family and provides conclusive Mesozoic evidence for a rare type of predation involving offensive (vs. defensive) secretions. The specimen was fossilized in a preying-and-spraying position with its presumptive meal, a Symphypleona springtail (Collembola), reflecting the behavioural predatory repertoire of modern relatives. Apart from one Cenozoic species, these rare specimens represent the only fossil record of the family Projapygidae. Our findings demonstrate that these basal hexapods were predators of small arthropods of ancient forest soils and enrich our understanding of palaeoecological associations and behavioural strategies paramount for the survival of species.

A new genus of Plusiocampinae (Diplura, Campodeidae), Anatoliacampa diclensis Sendra, Tusun & Satar gen. et sp. nov., is described from specimens found in the Southeastern Anatolian Peninsula in Dicle cave (Bozaba village, Dicle district, Diyarbakır province, Turkey). Based on its morphological characteristics and molecular analysis, the new taxon undoubtedly belongs to the well-established subfamily Plusiocampinae and occupies, like most species of the subfamily, cave habitats. Anatoliacampa diclensis has two important features: its unique pretarsus and the abundant distribution of urosternal macrosetae. The finding of this new taxon is of high palaeogeographic interest, as it provides new evidence for establishing Anatolia as a land bridge for the colonization of the Euro-Mediterranean lands by cave-adapted Plusiocampinae and their few soil-dwelling relatives.

1. Diplura is the sister group to insects and one of the three basal hexapod groups with unique entognathan mouthparts. The order is divided into 10 families, which include 1008 species in 141 genera, with a high proportion of monotypic genera. They are ubiquitous in soils and subsurface terrestrial habitats, as well as have an important role in overall biogeochemical cycles. 2. We present the first comprehensive review of the global biodiversity and ecology of Diplura. We highlight four aspects of this basal hexapod group: diversity in morpho logical body plans and sizes; ecology in terrestrial environments from soil to caves; food preference and trophic levels, and their biogeographical and paleobiogeographical significance. 3. Diplura depends on high humidity and moderate temperatures. They are presum ably very sensitive to anthropogenic pressures and climate change, and therefore are a suitable model for ecophysiological studies and evident priority targets for conservation. 4. We conclude that the future efforts should focus on establishing a molecular phy logeny to clarify the relationships between and within families, as well as to reveal global biogeographical patterns. This will require an increase in sampling effort in several regions of the globe, especially in tropical regions.

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Soil organisms drive major ecosystem functions by mineralising carbon and releasing nutrients during decomposition processes, which supports plant growth, aboveground biodiversity and, ultimately, human nutrition. Soil ecologists often operate with functional groups to infer the effects of individual taxa on ecosystem functions and services. Simultaneous assessment of the functional roles of multiple taxa is possible using food-web reconstructions, but our knowledge of the feeding habits of many taxa is insufficient and often based on limited evidence. Over the last two decades, molecular, biochemical and isotopic tools have improved our understanding of the feeding habits of various soil organisms, yet this knowledge is still to be synthesised into a common functional framework. Here, we provide a comprehensive review of the feeding habits of consumers in soil, including protists, micro-, meso- and macrofauna (invertebrates), and soil-associated vertebrates. We have integrated existing functional group classifications with findings gained with novel methods and compiled an overarching classification across taxa focusing on key universal traits such as food resource preferences, body masses, microhabitat specialisation, protection and hunting mechanisms. Our summary highlights various strands of evidence that many functional groups commonly used in soil ecology and food-web models are feeding on multiple types of food resources. In many cases, omnivory is observed down to the species level of taxonomic resolution, challenging realism of traditional soil food-web models based on distinct resource-based energy channels. Novel methods, such as stable isotope, fatty acid and DNA gut content analyses, have revealed previously hidden facets of trophic relationships of soil consumers, such as food assimilation, multichannel feeding across trophic levels, hidden trophic niche differentiation and the importance of alternative food/prey, as well as energy transfers across ecosystem compartments. Wider adoption of such tools and the development of open interoperable platforms that assemble morphological, ecological and trophic data as traits of soil taxa will enable the refinement and expansion of the multifunctional classification of consumers in soil. The compiled multifunctional classification of soil-associated consumers will serve as a reference for ecologists working with biodiversity changes and biodiversity–ecosystem functioning relationships, making soil food-web research more accessible and reproducible.

Samples collected in Central Asia, Kyrgyzstan, have revealed a hitherto unknown diversity of Campodeidae (Diplura) in soil and cave habitats, including a new genus and species, Kyrgyzstancampa sanare Sendra & Ferreira gen. et sp. nov., Turkmenocampa edaphica Sendra & Sánchez-García sp. nov. and a previously recognized soil-dwelling species, Campodea (Dicampa) catalana Denis, 1930. Kyrgyzstancampa sanare Sendra & Ferreira gen. et sp. nov. was collected in the deep zone of an interesting geological and cultural cave, Ak-Turpak Cave, located near the western margin of Kadamjay District, Batken Province. This genus belongs to the subfamily Campodeinae, sharing the morphology of the pretarsus with Eutrichocampa and other related genera, but differing from them in the shape of the claws and the laminar lateral processes, in addition to its unique cupuliform organ and the macrosetal pattern on the thorax and abdomen. Turkmenocampa edaphica Sendra & Sánchez-García sp. nov. was found in humid edaphic habitats, under stones or near roots, and is morphologically and geographically very similar to the cave-dwelling species Turkmenicampa mirabilis Sendra & Stoev, 2017, which occurs in an isolated cave in the nearby country of Turkmenistan.

Five new species are described, Plusiocampa (Didymocampa) cvijici Sendra & Antić, sp. nov., Plusiocampa (Plusiocampa) atom Sendra & Antić, sp. nov., Plusiocampa (Stygiocampa) barethae Sendra & Rađa, sp. nov., Plusiocampa (Stygiocampa) dulcici Sendra & Rađa, sp. nov. and Plusiocampa (Venetocampa) pirnati Sendra & Borko, sp. nov. This brings the number of cave-adapted species of campodeids known from the Dinarides, Eastern Alps, Balkan System and Rodope Massif to 28 (one species in the genus Campodea and 27 in Plusiocampa). Among Plusiocampa, four out of five subgenera are present (Pentachaetocampa is not present in the region studied): Didymocampa (one species), Plusiocampa s. str. (15 species), Stygiocampa (eight species) and Venetocampa (three species), whereby Stygiocampa and Venetocampa are endemic to the studied region. These results reveal the importance of the Dinarides karst region as a centre of diversification for campodeids and for cave animals in general. A monophyletic subgroup, consisting of Stygiocampa, Venetocampa and Plusiocampa s. str. and characterized by the absence of medial posterior thoracic macrosetae, presumably colonized the Dinaric plate during the middle of the Cenozoic and occurs in that area since then.

A new dipluran species, Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov., from the deep zone in three caves in the Imereti region, Georgia, is described. This new troglobitic Plusiocampa is an addition to four others known Diplura from around the Black Sea region, two Dydimocampa and two Plusiocampa s. str. The present study also provides the first CO1 sequences for the Plusiocampinae taxa and the first molecular data for cave-dwelling Plusiocampa species. Although bootstrap values were low, the maximum-likelihood phylogenetic tree grouped Plusiocampa (P.) imereti Sendra & Barjadze sp. nov. with two Plusiocampa s. str. species from Eastern Europe. Morphologically, P. (P.) imereti Sendra & Barjadze sp. nov. is closely related to two cave-dwelling species: Plusiocampa (Plusiocampa) glabra Condé, 1984 and Plusiocampa (P.) chiosensis Sendra & Gasparo, 2020. The new species can be distinguished by the presence of lateral anterior macrosetae on metanotum, more uneven claws, and the presence of 2+2 lateral anterior macrosetae on middle urotergites. The five species currently known for the Black Sea region inhabit caves located at low altitude but with no influence from former glacial or permafrost processes.

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Este artículo contiene 138 páginas, 213 figuras, 14 tablas., Diplura is a group of entognathous hexapods, often considered a sister group to insects. They play an important role in recycling organic matter in soil and subterranean terrestrial ecosystems. The Campodeidae is the most diverse family, divided into four subfamilies. The subfamily Plusiocampinae has a subterranean life-style with many species distributed in the Euro-Mediterranean area. The incertae sedis tachycampoids (“lignée Tachycampoïde”) is a group within the family Campodeidae that share with the Plusiocampinae a strong preference for subterranean habitats and several morphological characters, such as slender body shape, elongated appendages, considerable increment in the number of antennomeres and cercal articles, and complexity of sensorial structures. The present monograph provides a taxonomic revision of the subfamily Plusiocampinae and the genera belonging to the tachycampoid lineage from Europe and the Mediterranean region. It comprises detailed morphological descriptions and illustrations together with data on the habitats and distributions of 87 species, 10 subspecies and 11 affinis forms. Seven new species are described among those, namely: Plusiocampa (Plusiocampa) apollo Sendra, Giachino & Vailati sp. nov., P. (P.) chiosensis Sendra & Gasparo sp. nov., P. (P.) dublanskii Sendra & Turbanov sp. nov., P. (P.) hoffmanni Sendra & Paragamian sp. nov., P. (P.) rhea Sendra sp. nov., P. (P.) ternovensis Sendra & Borko sp. nov. and P. (Venetocampa) ferrani Sendra & Delić sp. nov., DS’s field trips were funded by the Serbian Ministry of Education, Science and Technology (Grant 173038); KP was partially funded by the HISR project “Conservation of the Cave Fauna of Greece” funded by the MAVA Foundation and WWF Greece; PMG and DV carried out sampling in Greece using permits from the Ministry of Environment no. 124085/1362/2015, 135366/373/2016 and 166238/248/2018; IT was funded within the framework of the state assignment of FASO Russia (themes no. АААА-А18-118012690106-7 and АААА-А18-118012690105), supported in part by RFBR (project no. 17-54-40017Абх_а); sampling efforts by AF and colleagues resulted from various projects on the evolution of cave fauna diversity, some of them funded by the German Research Foundation, DFG (DFG FA 1042/1-1 and DFG BA 2152/14-1); PMG and DV were partly supported by the program “Research Missions in the Mediterranean Basin” sponsored by the World Biodiversity Association onlus XLI contribution; ASPSR was supported by a research grant (15471) from Villum Fonden.

Over the last three decades, intensive sampling of the MSS in the Iberian Peninsula has revealed new records of Leistus (Pogonophorus) puncticeps Fairmaire & Laboulbène, 1854 in previously unknown areas. These new records extend the species geographic range towards southern Mediterranean localities, showing a discontinu­ous distribution pattern across the eastern third of the Iberian Peninsula. Also, some slight variability in the shape of the apical lamina of the edeago is revealed. Abrupt landscape, especially in stony slopes, arises as a favorable habitat for the species. In general, the new localities for L. (P.) puncticeps have low and torrential precipitation, an average rainfall of around 600-700 mm, and remarkable thermal amplitude. The species is collected for the first time in the Mesovoid Sallow Substratum (MSS) using subterranean sampling devices (SSD). The pres­ence of this species in this hypogean habitat across several iberian areas (Sierra del Moncayo, mountains of the north of Alicante province and Sierra de María) indicates its (sub)troglophile condition. The new records, in addition to previously available data, suggest the same evolutionary history for L. (P.) puncticeps as for Leistus (Pogonophorus) montanus Stephens, 1827 and Leistus (Pogonophorus) parvicollis Chaudoir, 1869, being the three species glacial relicts as a result of vicariant speciation. The environmental changes that occurred during the postglacial time period could explain the discontinuous distribution that L. (P.) puncticeps shows at present and its tendency to occupy the MSS. Numerosos muestreos realizados por el territorio peninsular, a lo largo de estas tres últimas décadas, han deparado el hallazgo de Leistus (Pogonophorus) puncticeps Fairmaire & Laboulbène, 1854 en áreas geográfi­cas en las que se desconocía su presencia hasta la fecha. Las nuevas citas amplían la distribución ibérica de la especie hacia enclaves mediterráneos meridionales, perfilando un patrón de distribución discontinuo por el tercio oriental. Al mismo tiempo, estos hallazgos revelan cierta variabilidad, muy leve, en la forma de la lámina apical del edeago. Por otro lado, se confirman los parajes quebrados y abruptos como enclaves potencialmente favorables para la supervivencia de esta especie, especialmente si disponen de laderas pedregosas. En general, las nuevas localidades en donde se ha hallado L. (P.) puncticeps, cuentan con precipitación escasa y torrencial, una pluvio­sidad media que ronda los 600-700 mm, y una notable amplitud térmica. Como novedad, y utilizando Estaciones de Muestreo Subterráneo (EMS), se ha colectado en el Medio Subterráneo Superficial (MSS). El hallazgo de esta especie en este singular medio hipogeo y en diversos enclaves ibéricos (Sierra del Moncayo, formaciones montañosas del norte de la provincia de Alicante y Sierra de María), pone de manifiesto su tendencia troglófila, y más probablemente de tipo subtroglófila. Los nuevos hallazgos, sumados a los ya conocidos, sugieren que L. (P.) puncticeps ha seguido la misma historia evolutiva que Leistus (Pogonophorus) montanus Stephens, 1827 y Leistus (Pogonophorus) parvicollis Chaudoir, 1869, constituyendo una triada de relictos glaciales, resultado de un proceso de especiación por vicarianza. Los cambios acaecidos en el período postglacial explicarían la distribución discontinua que muestra en la actualidad L. (P.) puncticeps, y su tendencia a explorar, en estos enclaves, el MSS.

Diplurans (Hexapoda) are considered the ‘ideal cavernicolous animal’ having one of the highest ratios of cave-adapted vs. non-cave-adapted species. They are successful colonizers of subterranean habitats, thriving in all cryptic, dark, terrestrial environments. Diplurans play an important role in the decomposition of organic matter below the ground and are sensitive to anthropogenic pressures. We present the first comprehensive review about cave Diplura diversity, ecology, evolution, distribution and biogeography. We provide a roadmap for research questions regarding the ecology, aimed at stimulating the pursuit of new studies on this fascinating group. Filling these current knowledge gaps will contribute to conservation efforts for cave ecosystems.

Two new dipluran species of the family Campodeidae have been unearthed in the Canary Islands. Remycampa herbanica sp. nov. was found in a highly threatened lava tube on Fuerteventura island. It is related to the soil-dwelling northwest African Remycampa launeyi that also inhabits four of the Canary Islands. The two known Remycampa species are characterized by a torsion of the labial palps. They differ chiefly in the distribution of macrosetae and in the features of cave adaptation of R. herbanica, i.e. elongation of body and appendages, and a higher number of olfactory chemoreceptors with a coniform shape unique within campodeids. Spaniocampa relicta sp. nov. was collected in the mesovoid shallow substratum (MSS) and has been assigned to a formerly monotypic genus that includes the soil-dwelling Spaniocampa prima from the Republic of Guinea. The two species differ in the number of abdominal macrosetae. Females of S. relicta sp. nov. have small setae arranged in groups along the posterior border of the first urosternite. These structures of unknown function have never been described in other campodeid species. Sequencing the COI barcode region of R. herbanica has been produced but it proved insufficient to identify closest relatives. The two new hexapods from subterranean habitats raise the Canarian campodeid fauna to six species. Five of them are living in soil and/or MSS, whereas the cave-adapted R. herbanica is known only from a single, particularly endangered lava tube distant from other caves.

Four species of Campodeidae have been recorded from Sarnena Gora Mts. region from which Campodea (Dicampa) caucasica Rusek, 1965 and Campodea (Campodea) magna Ionescu, 1955 are new records to the fauna of Bulgaria.

The mesovoid shallow substratum (MSS) can act as a climatic refuge for invertebrates, as a biogeographic corridor to deeper substrates or as a permanent habitat for some species. This study characterizes the seasonal invertebrate diversity and abundance of MSS ecosystems in central Portugal focusing on Diplopoda, Diplura, Orthoptera and Coleoptera during one year. Sampling was performed with standard MSS pitfalls in scree slopes (colluvial MSS) of karst areas and environmental parameters (temperature, pH, conductivity, water content, organic carbon, nitrate, phosphate and ammonium) were quantified. Our results show that winter was the season with the highest arthropod abundance and that the MSS acts as a permanent habitat for chordeumatidan millipedes and as a climatic refuge for orthopterans and most beetles. All Diplura collected belong to a single species known previously from surface habitats in the Iberian Peninsula, which does not seem to use the Portuguese MSS as a refuge. MSS habitats in central Portugal, classified as western Mediterranean and thermophile deposits protected by the Natura 2000 network based on plant communities and geology, revealed an abundant and diverse invertebrate community that urges characterization and protection.

The aim of this study was to unravel the relative role played by speleogenesis (i.e., the process in which a cave is formed), landscape‐scale variables, and geophysical factors in the determination of species richness in caves. Biological inventories from 21 caves located in the southeastern Iberian Peninsula along with partial least square (PLS) regression analysis were used to assess the relative importance of the different explanatory variables. The caves were grouped according to the similarity in their species composition; the effect that spatial distance could have on similarity was also studied using correlation between matrices. The energy and speleogenesis of caves accounted for 44.3% of the variation in species richness. The trophic level of each cave was the most significant factor in PLS regression analysis, and epigenic caves (i.e., those formed by the action of percolating water) had significantly more species than hypogenic ones (i.e., those formed by the action of upward flows in confined aquifers). Dissimilarity among the caves was very high (multiple‐site βsim = 0.92). Two main groups of caves were revealed through the cluster analysis, one formed by the western caves and the other by the eastern ones. The significant—but low—correlation found between faunistic dissimilarity and geographical distance (r = .16) disappeared once the caves were split into the two groups. The extreme beta‐diversity suggests a very low connection among the caves and/or a very low dispersal capacity of the species. In the region under study, two main factors are intimately related to the richness of terrestrial subterranean species in caves: the amount of organic material (trophic level) and the formation process (genesis). This is the first time that the history of a cave genesis has been quantitatively considered to assess its importance in explaining richness patterns in comparison with other factors more widely recognized.

A description of a new troglobitic Lepidocampa species from La Reunion Island is given. Certain taxonomical attributes of this new species suggest that the subgenera traditionally established within Lepidocampa genus should be eliminated. Next, the main troglobiomorphic features in campodeid diplurans (large body size, long appendages, high number of antennomeres and cercal articles, presence of lateral crests and laminar processes on telotarsus, high number of complex antennal sensilla, and absence of a seasonal reproductive cycle) are reviewed in order to clarify their adaptive value to the subterranean milieu. Finally, the growth of cerci in relation to body size in campodeids is explored using data compiled from the literature and standardised major axis regression analysis. A large body and long appendages may be especially relevant in campodeids when dwelling in the subterranean fissures. Yet, the allometric pattern of cerci length in relation to body size among subfamilies points to a phylogenetic signal, probably related to the habitat preferences of each group, and which suggests the importance of the physical space in which the different families preferentially dwell.

The species Paraphaenops breuilianus (Jeannel, 1916) is an Iberian iconic cave beetle with “aphaenopsian” facies, monospecific until now. An extensive field work over the last years allowed the revision of this peculiar genus using the combinations of morphological, molecular and ecological approaches and led to the description of new taxa Paraphaenops fadriquei Ortuno and Faille sp. nov., Paraphaenops breuilianus espanoli Ortuno and Faille ssp. nov. and the full larval diagnosis of the third larval instar of the type species. The sequencing of two mitochondrial (cox1, rrnl + tRNA-Leu + nad1) and one nuclear (LSU) gene fragments evidenced a strong divergence for all markers considered between P. breuilianus and the new species P. fadriquei sp. nov. The strong genetic differences between the two taxa contrast singularly with the extreme morphological homogeneity of the genus. The ecological data recorded for Paraphaenops Jeannel, 1916 species show that they are stenotherms and stenohygrobionts, perfectly adapted to the subterranean environment, living at temperatures between 4.4 and 10 °C with high relative humidity. The new localities and taxa enlarge the former distribution area from 72 to 5091 square kilometres sprinkled between twelve caves: six in the Mola de Cati (P. breuilianus breuilianus); three in the aforementioned karst region towards the south, crossing the geographical barrier “Barranc de Regatxol” (P. breuilianus espanoli ssp. nov.); and three, 60 km towards the west, in a different karst region: the “Sierra de la Dehesas” (P. fadriquei sp. nov.) This geographical range overlaps a geostructural area quoted as “junction area” between the Iberian System and the Catalan coastal Mountain range, an area up to 1000 meters above sea level.

A description is given of a biogeographically interesting new species of Campodeidae (Diplura), Plusiocampa (Plusiocampa) tinoamorei sp. nov., a troglobiotic species found in the Sicilian Villasmundo and Scrivilleri caves. A second subterranean species already known from three caves in northwest Sicily near Palermo, Campodea (Campodea) majorica sicula Condé, 1957, also was studied. Both species were characterized with SEM. Each species belongs to a different monophyletic group: Plusiocampa s. str., with thoracic medial posterior macrosetae, and the group related to Campodea (Campodea) grassi Silvestri, 1912. Both groups are widely distributed on almost all of the large western Mediterranean islands. Nevertheless, although these two monophyletic groups have a different origin both could be dispersed via land connections between the mainland and the islands during the Messinian Age. This new discovery shows the great value of Sicilian caves that hold species with unique features and of high biogeographic significance.

Extensive fieldwork has been carried out in two protected caves (Villasmundo and Monello) in eastern Sicily in 2018–2019. The caves, located respectively within the Strict Nature Reserves “Complesso Speleologico Villasmundo-S. Alfio” and “Grotta Monello”, are two of the most important karst systems of the Hyblean area. They are both remarkably developed and possess speleothem specimens of great scientific value such as stalactites, stalagmites, and columns. The study has further increased our knowledge about the existing cave fauna, and focused on the optimization of good conservation practices of hypogean environments. Systematic investigations have confirmed the presence of a rich arthropod community, and, above all, have highlighted species of great importance, including ones new to science. In particular, two pselaphids (Coleoptera: Staphylinidae) of the genus Tychobythinus Ganglbauer, 1896 and a dipluran (Entognatha) of the genus Plusiocampa Silvestri, 1912 have been identified and described.

A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zygentoma) in the collection of the Natural History Museum of Denmark, University of Copenhagen is presented. The NHMD collection harbours type material of 175 species, and it is composed of 132 holotypes, 4 neotypes, 3 syntypes, 32 lectotypes, and 39 paratypes. For Diplura one neotype is newly designated. The order Protura is represented by 5 families, Collembola by 7 families, Diplura by 6 families, Archaeognatha by 1 family and Zygentoma by 2 families.

Although Iberian subsurface terrestrial habitats have been sampled for a half century, they remain poorly known. During the last five years much more sampling of these subsurface habitats has been made, mainly in scree slopes (also called colluvial Mesovoid Shallow Substratum habitats, MSS) but also in alluvial debris of temporal watercourses (alluvial MSS). In our study, diplurans, a basal hexapod group, were extracted from two hundred traps installed in 69 locations in the mountain ranges of six different regions of the Iberian Peninsula, from north to south: Cantabrian, Pyrenees, Iberic System, Central System, Prebaetic and Penibaetic Mountains. A total of 1251 specimens in fifteen dipluran species: thirteen described Campodeidae, one described Japygidae and one new Campodea species inhabiting the alluvial MSS habitats of the watercourses of Prebaetic Mountains. A few populations of these dipluran species show troglobiomorphic features as a consequence of the medium-sized voids of the MSS habitats, such as Campodea grassii Silvestri, 1912, collected in a scree slope connected with a deep subterranean system in Penyas Roset, Prebaetic Mountains. Most species found in MSS habitats are endogean or epiedaphic species living in the area, but this is not the case in Sierra de Guadarrama, where three species (Campodea propinqua Silvestri, 1932, Campodea neusae Sendra & Moreno, 2006 and Campodea zuluetai Silvestri, 1932) unknown in the soil of these mountains have appeared in these subsurface terrestrial habitats., This work has been supported by the research projects CGL2010-19924 and 1143/2014. V.M.O. was supported by the I3 program of the Spanish Ministry of Education and Science, and AJ.-V. was supported by the MINECO Ramón y Cajal Program (RYC-2013-14441).

Over the last 40 years, hypogenic karst/caves have become well known and hypogene speleogenesis has been used to explain the formation of some of the largest subterranean maze caves. These hypogenic systems involve confined aquifers with upwards flow, responsible for their karstification. Such spaces begun and could be remain isolated from the surface and the contiguous subterranean habitats, including the shallow ones. We studied the invertebrate fauna and the geology/speleogenesis of two recently opened caves from Eastern Iberian Peninsula, which clearly showed features of hypogenic origin. Their scarce faunistic composition, formed by non cave-adapted fauna, is similar to other hypogenic caves from Europe and North America, suggesting that there are restrictions for faunal colonization of the hypogenic subterranean ecosystems. We propose differentiating the hypogenic from the epigenic subterranean ecosystem based on the fact that the first one lacks fauna during its genesis. This new scenario may have consequences in the regional biodiversity patterns of troglobiont species.

Six new taxa of Campodeidae (Diplura) are described in the genera Litocampa, Juxtlacampa, Oncinocampa, and Tachycampa. We also redescribe the interesting species Juxtlacampa juxtlahucensis Wygodzinsky, 1944 from Juxtlahuaca cave in Guerrero, Mexico. All of these taxa are cave-dwelling species with more or less noticeable troglobiomorphic features They inhabit the subterranean ecosystem in six limestone massifs and one lava tube cave in the central states of Mexico. Four of these species are included in the "tachycampoide" group and one species in the "podocampoide" group (sensu BarethConde). Nine species already known in Central and South America of the "tachycampoide" group, in such poorly-sampled regions compared with the eight species in the well-sampled Mediterranean region (Ibero-Sardinia and north Africa), suggest an American origin for this group.

Resume Les Anillini hypoges se trouvent cantonnes dans les massifs karstiques de lest de la peninsule Iberique, avec cinq especes du genre Speleotyphlus, une du genre monospecifique Iberanillus (I. vinyasi) et un troglobie exceptionnel du genre Aphaenotyphlus, lui aussi monospecifique. La singularite de ce dernier genre (Aphaenotyphlus alegrei Espanol & Comas 1985) nous a amene a nous interesser a l’etude detaillee de son anatomie, en incluant la description du mâle, jusqu’a present inconnue, et a approfondir l’ecologie et la biogeographie de ce taxon. A. alegrei appartient a la sous-tribu Scotodipnina et semble etre proche de Microtyphlus et de Speleotyphlus en raison de certains caracteres morphologiques partages, parmi lesquels la separation du labium par rapport au prebasilaire. Cependant, A. alegrei constitue une forme plus specialisee au milieu souterrain comme il semble se deduire a partir de son habitus, avec un certain “modelage aphaenopsien“. Nous proposons une cle de genres des Anillini iberiqu...

© 2015, Springer International Publishing Switzerland. In this study, the invertebrate fauna of a colluvial Mesovoid Shallow Substratum (MSS) is Eastern Spain was characterized, and the importance of the MSS as refuge for endemic and rare species was assessed. Ten pitfall traps were buried up to 1 m deep inside multiperforated PVC tubes across a stone debris. Several environmental variables were measured in each sampling point. The completeness of the inventory was assessed, and different diversity patterns—variation in abundance, species richness and species composition—were analyzed. 4150 individuals and 164 species were captured, most of them arthropods. The number of individuals and species varied considerably among traps. Half of the species were represented by one or two individuals, and neither the species accumulation curves nor the curves for the non-parametric estimators showed any sign of stabilizing. Individuals of abundant species aggregated in one or a few traps. No consistent diversity patterns were found, except that distance among traps partially explained the similarity in species composition when considering only the most abundant species. This is a very rich but uneven assemblage that shows a high degree of species turnover between traps. One-sixth of the species can be classified as either troglobionts or troglophiles, but there is a significant presence of edaphic species. Obtaining complete fauna inventories in the MSS has proven to be a challenging task. The study of the MSS reveals important faunistic information, providing new taxa to science as well as interesting records of poorly known species., This research Project (CGL2010-19924) was funded by the Spanish Ministry of Science and Innovation. A. J.-V. was supported by the CSIC JAE-Doc Program which is partially financed by the European Social Fund.

La position taxonomique d’Ildobates neboti Espanol, 1966, un paleoendemique hypoge de la Cordillere Iberique, a ete confuse depuis sa decouverte. Dans la description originale, il a ete assigne, av...

Two new species and one new subspecies of campodeids (Diplura) are described from caves of the northeastern Iberian Peninsula: Campodea (Campodea) cossetana sp. n., Campodea (Campo - dea) maestrazgoensis sp. n. and Plusiocampa bonneti condei ssp. n. Ten additional forms are studied, all of them belonging to the same biogeographic area. We finally discuss some hypotheses concerning the colonization's chronology of the northeastern Iberian Peninsula.

The discovery of several members of the genus Coletinia Wygodzinsky, 1980, from subterranean habitats (endogean and troglobiont), prompted the review of this genus in the Iberian Peninsula. Most of the samples came from caves of the Mediterranean basin of Spain, from Cádiz to the Tarragona province. As a result of this revision, nine new species have been established: C. herculea n. sp., an endogean from Cádiz; C. vergitana n. sp. from the Gádor calcareous mountains in Almería; C. calaforrai n. sp. from the gypsum karst in Almería; C. intermedia n. sp. from caves in Murcia and Alicante; C. diania n. sp., found in the north of the province of Alicante; C. longitibia n. sp. and C. tessella n. sp., both troglobites from Valencia; C. redetecta n. sp. from Castellón caves and finally C. hernandoi n. sp., an endogean from Tarragona. Moreover, Coletinia maggii (Grassi, 1887) is reported for the first time in the Iberian Peninsula, and new data are presented regarding C. mendesi, C. tinautiand C. capolongoi that widen their geographic distribution and enhance the information about their anatomic characteristics and biology. These results increase the number of known species of this genus to 14 in the region and to 21 in the world. The new species are described and compared with the most closely related previously known species of the genus. Characters with the most taxonomic relevance are discussed using optical and scanning microscope studies. A key for the identification of the Iberian Coletinia species and a distribution map including all of them are also provided.