José D. Gilgado

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.

Historical perspective, new contributions and an enlightening dispersal mechanism for the endogean genus Typhlocharis Dieck 1869 (Coleoptera: Carabidae: Trechinae)

A historical review of the study of genus Typhlocharis and a synthesis of current knowledge are carried out, providing a historical context for the description of a new species in the province of Alicante (Spain). Relevant data on the biotic and abiotic environment where the species lives are also provided. With regard to its taxonomic position, it is affiliated with the Typhlocharis monastica group, although it is necessary to redefine that group. The most plausible hypothesis for the unusual presence of unguiform gonocoxites in the genus Typhlocharis and the implications for the systematics of the group are discussed. Finally, a dispersal mechanism for Typhlocharis that would explain its wide distribution in the Iberian Peninsula and North Africa is proposed.

Hypogenic versus epigenic subterranean ecosystem: lessons from eastern Iberian Peninsula

*alberto.sendra@uv.es epikarst, a very superficial habitat under the surface of the karst in the infiltration zone (Bakalowizc, 2005), which is an ecotone between surface water and groundwater (Pipan, 2005); and the hypotelminorheic habitat (Meštrov, 1962; Culver & Pipan, 2009b), a persistent shallow subsurface of wet spots fed by subsurface water in a slight depression (Fig. 1). The aquatic compartment of the subterranean ecosystem extends through the deepest aquifers such as the Edwards Aquifer in Texas (USA) at 600 to 1000 meters deep, to the shallow interstitial habitats composed of water filled spaces between unconsolidated sediments (Culver & Pipan, 2009a). Among these subterranean compartments, the subterranean-adapted fauna can inhabit from the shallow habitats in the subsurface, through to the deepest known galleries at more than -2000 m and from the giant subterranean chambers to the smallest fissures (Sendra & Reboleira, 2012). In the early 70’s the discovery of the hypogenic karst, firstly studied in Europe and North America INTRODUCTION

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.

Dima assoi Pérez Arcas 1872 (Coleoptera: Elateridae): from montane to hypogean life. An example of exaptations to the subterranean environment?

Summary The mesovoid shallow substratum (MSS) of two areas in the Moncayo Massif (Zaragoza, Spain) was sampled for a year with subterranean pitfall traps installed at one meter depth in stony debris. Throughout this period, a large contingent of imagoes of Dima assoi Pérez Arcas 1872 (Elateridae) was found; this is a species whose genus is disjunctively distributed at high altitudes, in the Himalayas and Europe. It frequently appeared in the four subterranean traps located in one of the two areas (a bare scree), as well as in some surface pitfall traps, but less frequently in the other area (a beech forest). This fact contrasts with the knowledge about this species, given its nocturnal nature and its sublapidicolous behavior, but seems to agree with its morphological features. Nevertheless, apart from anecdotal information, almost all other aspects of the biology of this species remain unknown. In this work we provide both biotic and abiotic data about the subterranean habitat where this species has been found. The implications of these findings are discussed in relation to its biology and to the significance of its possible exaptations to the subterranean environment as a result of adaptation to montane habitats.

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.

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

Abstract 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. RESUMEN. En la actualidad, una serie de estudios sistematizados en el Medio Subterráneo Superficial (MSS), se están llevando a cabo en diversos enclaves de la península Ibérica, lo que ha propiciado el descubrimiento de la enigmática hormiga Aphaenogaster cardenai Espadaler, 1981 en un macizo montañoso muy alejado de su área de distribución conocida. Esta especie ha sido considerada, hasta el momento, como una especie hipogea. Se discute su rol ecológico y su posible área de distribución real de acuerdo con este nuevo hallazgo, así como la morfología de las castas conocidas, corología, capacidad de vuelo de las formas sexuadas y su período de actividad. Esto ha permitido revisar y discutir el estado actual del conocimiento sobre las posibles adaptaciones y exaptaciones de los Formicidae a los ambientes subterráneos (sensu lato), y concretamente al MSS. De acuerdo con todo lo anterior, las hormigas podrían adaptarse a los ambientes hipogeos más profundos mediante la modificación de su estructura social, pero sin dichos cambios, el MSS sería su última frontera en su proceso de colonización de los ambientes hipogeos.

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.

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.

Biological notes and description of egg and first instar larva of Carabus (Oreocarabus) ghilianii La Ferté-Sénectère 1847 (Coleoptera: Carabidae)

Abstract In this work, the authors describe the egg and first instar larva, hitherto unknown, of Carabus (Oreocarabus) ghilianii La Ferté-Sénectère 1847, a threatened and protected species endemic to the Iberian Peninsula. With respect to the larval morphology, a comprehensive study of the chaetotaxy of the three tagmata is presented, accompanied by a detailed iconography. In addition, data on the biology of imagoes are provided, taken in its natural habitat and in captivity, highlighting the novel fact that this species produces winter larvae. Thus, reproduction begins in late spring. Both the eggs and the larvae were obtained after captive rearing of nine specimens collected in the Sierra de Guadarrama (Madrid, Spain).