Subterranean ants: the case of Aphaenogaster cardenai (Hymenoptera: Formicidae).

Recently, a series of systematized studies of the Milieu Souterrain Superficiel (MSS) are being carried in several enclaves of the Iberian Peninsula, which have entailed the finding of the enigmatic ant Aphaenogaster cardenai Espadaler, 1981, hitherto considered as hypogean, in a mountain range far away from its known distribution area. Its ecological role and its possible area of distribution are discussed due to this finding, as well as its known morphology, distribution, habitat use, flight ability of the sexual forms, and moment of activity. This enabled reviewing and discussing the actual knowledge on the possible adaptations and exaptations of the Formicidae to the subterranean environments in wide sense and concretely to the MSS. According to all above, ants might adapt to the deepest hypogean environments by means of changes in their social structure, but without those changes, the MSS would be their last frontier in their process of colonization of hypogean environments.

A new hypogean Trechus Clairville (Coleoptera, Carabidae, Trechini) discovered in a non-calcareous Superficial Subterranean Habitat of the Iberian System (Central Spain).

A new hypogean species of Trechus Clairville, Trechus arrecheai sp. nov., is described from the Iberian Peninsula. It was captured by subterranean pitfall traps in a non-calcareous Superficial Subterranean Habitat from the Moncayo Massif (Zaragoza, Spain). Data on the accompanying fauna are provided and the biogeographical implications of this discovery are discussed. A synthesis of the data about the known distribution of the Trechus angusticollis species group is provided.

A new mute species of the genus Nemobius Serville (Orthoptera, Gryllidae, Nemobiinae) discovered in colluvial, stony debris in the Iberian Peninsula: A biological, phenological and biometric study.

Sampling of a Mesovoid Shallow Substratum (MSS) of a scree in the Guadarrama mountains (Madrid, Spain) revealed a population of crickets of the genus Nemobius Serville. A detailed morphological study revealed that the cricket was a new species, Nemobius interstitialis sp. nov., which is principally characterized by the absence of a tympanum in the outer margin of the foreleg tibiae and a peculiar design of venation of the forewing of the male. Sampling of this environment over 1 year using surface and MSS pitfall traps, set at a depth of one meter, allowed study of population dynamics. A population maximum is attained in August. Abiotic (temperature and humidity) and biotic (accompanying fauna) data are given to contextualize the habitat of this new species.

Postfire biodiversity database for eastern Iberia.

In the summer of 2012, two fires affected Mediterranean ecosystems in the eastern Iberian Peninsula. The size of these fires was at the extreme of the historical variability (megafires). Animals are traditionally assumed to recolonize from source populations outside of the burned area (exogenous regeneration) while plants recover from endogenous regeneration (resprouting and seeding). However, there is increasing evidence of in situ fire survival in animals. To evaluate the effect of large-scale fires on biodiversity and the mechanism of recovery, in 2013, we set up 12 plots per fire, covering burned vegetation at different distances from the fire perimeter and unburned vegetation. In each plot, we followed the postfire recovery of arthropods, reptiles (including some of their parasites), and plants for 2 to 5 years. Here we present the resulting database (POSTDIV) of taxon abundance. POSTDIV totals 19,906 records for 457 arthropod taxa (113,681 individuals), 12 reptile taxa (503 individuals), 4 reptile parasites (234 individuals), and 518 plant taxa (cover-abundance). We provide examples in the R language to query the database.

Six groups of ground-dwelling arthropods show different diversity responses along elevational gradients in the Swiss Alps.

Elevational gradients along mountain slopes offer opportunities to study key factors shaping species diversity patterns. Several environmental factors change over short distances along the elevational gradient in predictable ways. However, different taxa respond to these factors differently resulting in various proposed models for biodiversity patterns along elevational transects. Using a multi-taxa approach, we investigated the effects of elevation, area, habitat and soil characteristics on species richness, individual abundance and species composition of six groups of ground-dwelling arthropods along four transect lines in the Swiss National Park and its surroundings (Eastern Alps). Spiders, millipedes, centipedes, ants, ground beetles and rove beetles were sampled using standardized methods (pitfall traps, cardboard traps, visual search) in 65 sites spanning an elevational range from 1800 to 2750 m a.s.l.. A total of 14,782 individuals comprising 248 species were collected (86 spider, 74 rove beetle, 34 ground beetle, 21 millipede, 19 centipede and 14 ant species). Linear mixed model-analysis revealed that rarefied species richness in five out of the six arthropod groups was affected by elevation (the quadratic term of elevation provided the best fit in most cases). We found three different patterns (linear decrease in centipedes, low elevation plateau followed by a decrease in ants and rove beetles, and midpoint peak in spiders and millipedes). These patterns were only partially mirrored when considering individual abundance. Elevation influenced species composition in all groups examined. Overall, elevation was the most important factor explaining the diversity patterns, while most local habitat and soil characteristics have little influence on these patterns. Our study supports the importance of using multi-taxa approaches when examining effects of elevational gradients. Considering only a single group may result in misleading findings for overall biodiversity.

Functional diversity and habitat preferences of native grassland plants and ground-dwelling invertebrates in private gardens along an urbanization gradient.

Urbanization is occurring around the globe, changing environmental conditions and influencing biodiversity and ecosystem functions. Urban domestic gardens represent a small-grained mosaic of diverse habitats for numerous species. The challenging conditions in urban gardens support species possessing certain traits, and exclude other species. Functional diversity is therefore often altered in urban gardens. By using a multi-taxa approach focused on native grassland plants and ground-dwelling invertebrates with overall low mobility (snails, slugs, spiders, millipedes, woodlice, ants, rove beetles), we examined the effects of urbanization (distance to city center, percentage of sealed area) and garden characteristics on functional dispersion, functional evenness, habitat preferences and body size. We conducted a field survey in 35 domestic gardens along a rural-urban gradient in Basel, Switzerland. The various groups showed different responses to urbanization. Functional dispersion of native grassland plants decreased with increasing distance to the city center, while functional dispersion of ants decreased with increasing percentage of sealed area. Functional evenness of ants increased with increasing distance to the city center and that of rove beetles decreased with increasing percentage of sealed area. Contrary to our expectation, in rove beetles, the proportion of generalists decreased with increasing percentage of sealed area in the surroundings, and the proportion of species preferring dry conditions increased with increasing distance to the city center. Body size of species increased with distance to city center for slugs, spiders, millipedes, ants, and rove beetles. Local garden characteristics had few effects on functional diversity and habitat preferences of the groups examined. Our study supports the importance of using multi-taxa approaches when examining effects of environmental change on biodiversity. Considering only a single group may result in misleading findings for overall biodiversity. The ground-dwelling invertebrates investigated may be affected in different ways from the more often-studied flying pollinators or birds.

Description of the female morphology of the relict troglobiont millipede Cantabrodesmus lorioli Mauriès 1971 (Diplopoda, Polydesmida, Chelodesmidae), and new records in the deepest pit of Spain.

The troglobiont millipede, Cantabrodesmus lorioli Mauriès, 1971, inhabits a few caves in a small region of Northern Spain. It is considered a relict species because of its geographical isolation compared to others in its family, and its to some extent uncertain systematic position. Cantabrodesmus lorioli has been placed in three different families, and now is considered a member of the Chelodesmidae, a family with only one other known native European representative. There are few records of this species, and almost nothing is known about its biology. The only available information on this species is the description of the male and a list of caves where it has been captured. In the present work, we provide the first description of the female morphology, illustrated by scanning electron microscopy, and a series of new records from caves in a region where it had never been captured before, including the deepest known pit in Spain, Torca del Porrón. The female morphology does not show many differences from that of the male, but several characters are discussed. All of the known information on the morphology of the species is used to discuss the implications for its biology and systematic position.

Ground-dwelling invertebrate diversity in domestic gardens along a rural-urban gradient: Landscape characteristics are more important than garden characteristics.

Urbanisation is increasing worldwide and is regarded a major driver of environmental change altering local species assemblages. Private domestic gardens contribute a significant share of total green area in cities, but their biodiversity has received relatively little attention. Previous studies mainly considered plants, flying invertebrates such as bees and butterflies, and birds. By using a multi-taxa approach focused on less mobile, ground-dwelling invertebrates, we examined the influence of local garden characteristics and landscape characteristics on species richness and abundance of gastropods, spiders, millipedes, woodlice, ants, ground beetles and rove beetles. We assume that most of the species of these groups are able to complete their entire life cycle within a single garden. We conducted field surveys in thirty-five domestic gardens along a rural-urban gradient in Basel, Switzerland. Considered together, the gardens examined harboured an impressive species richness, with a mean share of species of the corresponding groups known for Switzerland of 13.9%, ranging from 4.7% in ground beetles to 23.3% in woodlice. The overall high biodiversity is a result of complementary contributions of gardens harbouring distinct species assemblages. Indeed, at the garden level, species richness of different taxonomical groups were typically not inter-correlated. The exception was ant species richness, which was correlated with those of gastropods and spiders. Generalised linear models revealed that distance to the city centre is an important driver of species richness, abundance and composition of several groups, resulting in an altered species composition in gardens in the centre of the city. Local garden characteristics were important drivers of gastropod and ant species richness, and the abundance of spiders, millipedes and rove beetles. Our study shows that domestic gardens make a valuable contribution to regional biodiversity. Thus, domestic urban gardens constitute an important part of green infrastructure, which should be considered by urban planners.

Out of the dark void: Ommatoiulus longicornis n. sp., a new julid from Spain (Diplopoda, Julida) with notes on some troglobiomorphic traits in millipedes.

A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habitat in eastern Spain, and the elongation of its antennae is discussed in relation to the habitat where it dwells. We take the occasion to discuss the elongation of appendages as a troglobiomorphic character in millipedes and shed the light on the various, sometimes controversial, hypotheses that could have led to these modifications in millipedes and other subterranean arthropods.

Distinctive Collembola communities in the Mesovoid Shallow Substratum: First data for the Sierra de Guadarrama National Park (Central Spain) and a description of two new species of Orchesella (Entomobryidae).

Two new species of the genus Orchesella Templeton, 1836 have been identified following intensive sampling in the Colluvial Milieu Souterrain Superficiel (Mesovoid Shallow Substratum, or MSS) of the Sierra de Guadarrama using Subterranean Sampling Devices (SSD). The data were obtained from the first extraction of the traps between May and October of 2015. During a study of the Collembola taxon, 32 different genera (61 species) were identified. The highest representative genus presence in almost all traps was Orchesella, with two new species. One of the two species described had been misidentified until this study was carried out, indicating that their preferential habitat had not been sampled; the second species had never been identified. The community of the Orchesella species in the Colluvial MSS was investigated, leading to the conclusion that this environment has its own assemblage of characteristic species. The opportunity to study specimens that belong to five species of the genus Orchesella, including three previously recollected, has allowed for obtaining reliable information regarding their macrochaetotaxy. A part of this chaetotaxy is proposed as a useful diagnostic tool for the species of the genus. In conclusion, it can be affirmed that this study has demonstrated that the Colluvial Mesovoid Shallow Substratum (Colluvial MSS) has its own fauna, and it supports the hypothesis that it constitutes a new biotope, at least for Collembola.

A new genus and species of Haplobainosomatidae (Diplopoda: Chordeumatida) from the MSS of the Sierra de Guadarrama National Park, central Spain.

The chordeumatidan fauna of the Iberian Peninsula is far from being well known, but recent efforts are improving that knowledge. Samplings carried out in the Milieu Souterrain Superficiel (also known as the Mesovoid Shallow Substratum) on several screes of the Sierra de Guadarrama National Park (central Spain) have led to the discovery of a new species of chordeumatidan millipede. This new species belongs to the family Haplobainosomatidae, which is mainly known from the northern Iberian Peninsula, and thus this is the first record of the family in central Spain. However, the extremely simple gonopods of this species are so different from all other known species of the family that it must be placed in a new genus. In this work, a complete description of the species, named as Guadarramasoma ramosae gen. & sp. nov., with a detailed iconography based on scanning electron microscopy images is provided together with a distribution map and a brief discussion of the implications of this new finding.

Intra- and inter-population polymorphism in Coletinia maggii (Grassi, 1887) (Zygentoma: Nicoletiidae), an inhabitant of soil, mesovoid shallow substratum (MSS) and caves--A challenge for the strict classification of subterranean fauna?

New locations of Coletinia maggii (Grassi, 1887) have been discovered in the center of the Iberian Peninsula in different types of subterranean environments, such as a stony layer in the subsoil of an alluvial plain, an alluvial Mesovoid Shallow Substratum or Milieu Souterrain Superficiel (MSS) and a gypsum cave. This is the first record of both an alluvial MSS in the center of the Iberian Peninsula and of a subterranean species living in it. The high number of specimens captured allowed the first detailed study of the morphological intra- and inter-population variations of this species. The implications of its presence in these different environments, its wide distribution area across Europe, and the relevance of the morphological variation in the characters for the taxonomy of this species are discussed. Based on the results, Coletinia hernandoi Molero, Bach & Gaju, 2013 is proposed as a new synonym of C. maggii.

The hypogean Iberian genus Typhlopsychrosoma Mauriès, 1982 (Diplopoda, Chordeumatida, Vandeleumatidae): distribution map, key to species, first record in a Mesovoid Shallow Substratum (MSS) and detailed iconography of T. baeticaense (Mauriès, 2013).

The troglobiont millipede Typhlopsychrosoma baeticaense (Mauriès, 2013) is recorded in the Mesovoid Shallow Substratum (MSS) of two screes in the Aitana and Bernia mountains (Eastern Iberian Peninsula), far away from its known distribution area. A detailed Scanning Electron Microscopy iconography provides additional information on gonopod morphology, as well as other details such as the inconspicuous evaginations of the cuticle at the place of the eyes in the anophthalmous specimens. We present an updated distribution map and a key to species of the genus, with illustrations of the gonopods of all species described so far. The implications of the appearance of this hypogean species in an MSS are discussed.

Sierra Nevada (Granada, Spain): a high-altitude biogeographical crossroads for millipedes (Diplopoda), with first data on its MSS fauna and description of a new species of the genus Ceratosphys Ribaut, 1920 (Chordeumatida: Opisthocheiridae).

Millipedes (Diplopoda), with a few notable exceptions, are poor dispersers, showing a very high degree of endemicity, not the least in mountains. The first samplings of the Mesovoid Shallow Substratum (MSS) of the higher altitudes of the Sierra Nevada Mountains (Baetic System, Southern Spain) have led to the discovery of a high number of millipedes, each of the species present showing a different degree of establishment in this subterranean environment. An update of the knowledge on the millipedes of this region, the first data of the millipede communities in the MSS and the description of Ceratosphys cryodeserti Gilgado, Mauriès & Enghoff n. sp. are here provided, as well as the first data on the humidity and temperature fluctuations in the MSS of this high mountain. The new species is similar to other Baetico-Riffan species, while the only previously known congener from the region, C. soutadei Mauriès, 1969, has more similarities to certain Pyrenean species. Biogeographical relationships of all the captured species are also discussed.

The "Alluvial Mesovoid Shallow Substratum", a new subterranean habitat.

In this paper we describe a new type of subterranean habitat associated with dry watercourses in the Eastern Iberian Peninsula, the "Alluvial Mesovoid Shallow Substratum" (alluvial MSS). Historical observations and data from field sampling specially designed to study MSS fauna in the streambeds of temporary watercourses support the description of this new habitat. To conduct the sampling, 16 subterranean sampling devices were placed in a region of Eastern Spain. The traps were operated for 12 months and temperature and relative humidity data were recorded to characterise the habitat. A large number of species was captured, many of which belonged to the arthropod group, with marked hygrophilous, geophilic, lucifugous and mesothermal habits. In addition, there was also a substantial number of species showing markedly ripicolous traits. The results confirm that the network of spaces which forms in alluvial deposits of temporary watercourses merits the category of habitat, and here we propose the name of "alluvial MSS". The "alluvial MSS" may be covered or not by a layer of soil, is extremely damp, provides a buffer against above ground temperatures and is aphotic. In addition, compared to other types of MSS, it is a very unstable habitat. It is possible that the "alluvial MSS" may be found in other areas of the world with strongly seasonal climatic regimes, and could play an important role as a biogeographic corridor and as a refuge from climatic changes.