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1. Range size is one of the most important criteria for determining species conservation status. It is an outcome of multiple factors, including a species' ecological tolerance and propensity to disperse, availability and temporal stability of suitable habitat patches, and between-patch connectivity. Understanding the determinants of range size could help identify the main threats, which may differ among species. 2. Subterranean species typically have smaller ranges than their surface counterparts. However, the determinants of their ranges are poorly known. The aim of this study was to test three potential determinants of species range size (habitat type, body size, and species age) using the species of the amphipod genus Niphargus. The study was performed within the Dinarides (south-east Europe), a global subterranean hotspot. 3. The only significant determinant of range size was the habitat in which the species lived. The species with the smallest ranges inhabit fissure systems in the upper layers of the carbonate massifs with predominantly vertical water flow. The species with the largest ranges live in the deepest zone – the permanently flooded phreatic with lateral water flow. These results imply that differences in connectivity among habitats determine the range sizes of aquatic subterranean species. 4. The vertical gradient in habitat connectivity indicates the vulnerability of the species to climate change. Desertification and longer periods of drought will probably have a greater effect on fauna in fissures than in the phreatic zone. In addition, lowering the groundwater table might exacerbate the fragmentation of intermediate habitats, possibly enhancing local extinction. 5. The habitat ecology of subterranean species should be considered in International Union for Conservation of Nature assessments. Given that the most narrowly distributed aquatic subterranean species live in habitats subjected to climatic drying, the protection of subterranean fauna depends on the management of surface habitats such as forests.

Speciation and dispersal are among key processes that drive biodiversity. Adaptive radiation (AR), a monophyletic species proliferation accompanied by ecological diversification, may be generating much of Earth’s biodiversity, but it has never been tested in subterranean environment. We reconstructed deep evolutionary history of amphipods. Next, we comprehensively analysed the speciation and ecomorphological diversification of the subterranean genus Niphargus, on continental scale, within a subterranean hotspot, and within one community. Amphipods evolved in several diversification pulses that correspond to ecological opportunities. Niphargus followed the course of AR that unfolded in the time of emerging mountain ranges. Several clades independently radiated, adaptively and nonadaptively. The clades adapted to distinct sets of adaptive optima with limited convergence. The speciation and dispersal contributed differently within the Dinarides. The diversity of the south-eastern Dinarides emerged mostly through local diversification of several clades. They acted as a donor area. The north-western hotspot arose through a combination of dispersal and local diversification. Although the general patterns support the AR hypothesis, explicit tests for ecological speciation are lacking. A community in Melissotrypa Cave is a potential case of ecological speciation of Niphargus. AR is thus a universal phenomenon, present also in subterranean environment, but more research is needed to understand the contribution of different processes to biodiversity, especially on microevolutionary level. Speciacija in disperzija sta ključna procesa, ki poganjata biotsko pestrost. Adaptivna radiacija (AR), hitro vznikanje vrst iz skupnega prednika, pospremljeno z ekološko diverzifikacijo, bi lahko bila ključen proces nastanka biotske pestrosti na Zemlji, a še ni bila testirana v podzemlju. Najprej smo rekonstruirali globoko evolucijsko zgodovino postranic. Nato smo celovito analizirali vzorce speciacije in ekomorfološke diverzifikacije slepih postranic iz rodu Niphargus, na kontinentalnem nivoju, znotraj vroče točke podzemne biotske pestrosti in znotraj ene združbe. Postranice so se razvile v več diverzifikacijskih pulzih, ki sovpadajo z ekološkimi priložnostmi. Slepe postranice so diverzificirale v procesu AR, na območju dvigajočih se kraških gorstev. Več kladov je neodvisno radiiralo, adaptivno in neadaptivno. Čeprav deloma konvergirajo, so se prilagodili na različne adaptivne optimume. Prispevek speciacije in disperzije se vzdolž Dinaridov spreminja. Biotska pestrost jugovzhodnih Dinaridov je večinoma posledica lokalne diverzifikacije več kladov in deluje kot prispevno območje vrst. Severozahodna vroča točka pa je posledica kompleksne kombinacije disperzije in lokalne diverzifikacije. Čeprav splošni vzorci pritrjujejo hipotezi AR, pa ekološka speciacija še ni bila primerno testirana. Združba slepih postranic v jami Melissotrypa bi lahko predstavljala primer ekološke speciacije. AR je tako univerzalni fenomen, prisoten tudi v podzemlju. Za razumevanje prispevka različnih procesov, še posebej na mikroevolucijskem nivoju, pa bodo potrebne dodatne raziskave.

Sex allocation theory predicts that the proportion of daughters to sons will evolve in response to ecological conditions that determine the costs and benefits of producing each sex. All else being equal, the adult sex ratio (ASR) should also vary with ecological conditions. Many studies of subterranean species reported female-biased ASR, but no systematic study has yet been conducted. We test the hypothesis that the ASR becomes more female-biased with increased isolation from the surface. We compiled a dataset of ASRs of 35 species in the subterranean amphipod Niphargus, each living in one of three distinct habitats (surface-subterranean boundary, cave streams, phreatic lakes) representing an environmental gradient of increased isolation underground. The ASR was female-biased in 27 of 35 species; the bias was statistically significant in 12 species. We found significant difference in the ASR among habitats after correction for phylogeny. It is most weakly female-biased at the surface-subterranean boundary and most strongly female-biased in phreatic lakes. Additional modelling suggests that the ASR has evolved towards a single value for both surface-subterranean boundary and cave stream dwelling species, and another value for 9 of 11 phreatic lake dwellers. We suggest that a history of inbreeding in subterranean populations might lower inbreeding depression such that kin selection favours mating with siblings. This could select for a female-biased offspring sex ratio due to local mate competition among brothers. The observed patterns in sex ratios in subterranean species make them a group worthy of more attention from those interested in sex allocation theory. peerReviewed

Plusiocampa (Plusiocampa) grandii Silvestri, 1933 Plusiocampa grandii Silvestri, 1933b: 1, figs i–ii. Diagnosis (Silvestri 1933b; Condé 1954a) Troglomorphic species. Antennae with 33–38 antennomeres. Pronotum with 1+1 ma, 2+2 la, 2+2 lp macrosetae; mesonotum with 1+1 ma, 2+2 la, 2+2 lp, 1+1 mp macrosetae; metanotum with 1+1 ma, 1+1 la, 2+2 lp, 1+1 mp macrosetae; long, barbed notal macrosetae; abundant clothing setae. Three ventral tibial macrosetae; unequal claws (1.25) with large lateral crests, posterior claw with a backward overhang. Urotergites I–II with 1+1 post macrosetae; urotergite III with 1+1 la, 1+1 post macrosetae; urotergite IV with 1+1 la, 5+5 post macrosetae; urotergites V–VII with 2+2 la, 5+5 post macrosetae; urotergite VIII with 6+6 post macrosetae; urosternite I with 7+7, urosternites II–VII with 6+6, urosternite VIII with 2+2 macrosetae. Male unknown; female with subcylindrical appendages bearing glandular a 1 setae., Published as part of Sendra, Alberto, Antić, Dragan, Barranco, Pablo, Borko, Špela, Christian, Erhard, Delić, Teo, Fadrique, Floren, Faille, Arnaud, Galli, Loris, Gasparo, Fulvio, Georgiev, Dilian, Giachino, Pier Mauro, Lukić, Marko, Marcia, Paolo, Miculinić, Kazimir, Nicolosi, Giuseppe, Palero, Ferran, Paragamian, Kaloust, Pérez, Toni, Polak, Slavko, Prieto, Carlos E., Turbanov, Ilya, Vailati, Dante & Reboleira, Ana Sofia P. S., 2020, Flourishing in subterranean ecosystems: Euro-Mediterranean Plusiocampinae and tachycampoids (Diplura, Campodeidae), pp. 1-138 in European Journal of Taxonomy 591 on page 51, DOI: 10.5852/ejt.2020.591, http://zenodo.org/record/3659823, {"references":["Silvestri F. 1933 a. Nuovi contributi alla conoscenza della fauna delle isole italiane dell'Egeo. Bolletino del Laboratorio di Zoologia generale e agraria in Portici 27: 61 - 111.","Silvestri F. 1933 b. Descrizione di una nuova specie cavernicola di Campodeidae (Thysanura Entothropha) del Trentino. Bollettino del Laboratorio di Entomologia agraria de Bologna 6: 1 - 4.","Conde B. 1954 a. Ordnung: Entotropha (Diplura). In: Franz H. (ed.) Der Nordost im Spiegel ihrer Landtierwelt: 644 - 649. Universitatsverlag Wagner, Innsbruck, Austria."]}

Plusiocampa (Plusiocampa) gadorensis Sendra, 2001 Plusiocampa gadorensis Sendra, 2001: 73, figs 1–4, tables i–ii. Diagnosis (Sendra 2001) Troglomorphic species. Antennae with 46–52 antennomeres; 15–18 complex olfactory chemoreceptors in cupuliform organ; small protruding frontal process with a few slightly tubercular setae. Pronotum with 1+1 ma, 2+2–3+3 la, 2+2 lp macrosetae; mesonotum with 1+1 ma, 2+2 la, 2+2 (2+3) lp macrosetae; metanotum with 1+1 ma, 2+2 la, 2+2 (2+3) lp macrosetae; long, barbed notal macrosetae; short notal clothing setae. Two or three ventral tibial macrosetae; slightly unequal claws (1.15) with large lateral crests, posterior claw with no backward overhang. Urotergite II with 0+0–1+1 (0+1) post macrosetae; urotergite III with 0+0–1+1 la, 1+1–2+2 (0+1) post macrosetae; urotergite IV with 1+1 (1+2) la, 3+3– 4+4 (4+5) post macrosetae; urotergite V with 1+1 (1+2) la, 4+4–5+5 (4+5) post macrosetae; urotergite VI with 1+1 (1+2) la, 5+5 (4+5) post macrosetae; urotergite VII with 1+1 la, 5+5 (4+5) post macrosetae; urotergite VIII with 6+6 (6+5) post macrosetae; abdominal segment IX with 8+8 post macrosetae; urosternite I with 7+7–8+8 (8+9), urosternites II–VII with 5+5, urosternite VIII with 2+2 macrosetae. Male with an area of glandular g 1 setae, subcylindrical appendages with glandular a 1 setae; female and male appendages identical. Habitat and distribution Subterranean species endemic to the isolated karst of Sierra de Gádor, in the Peni-Baetic System, southeastern Iberian Peninsula. It is known from seven natural caves and one mine., Published as part of Sendra, Alberto, Antić, Dragan, Barranco, Pablo, Borko, Špela, Christian, Erhard, Delić, Teo, Fadrique, Floren, Faille, Arnaud, Galli, Loris, Gasparo, Fulvio, Georgiev, Dilian, Giachino, Pier Mauro, Lukić, Marko, Marcia, Paolo, Miculinić, Kazimir, Nicolosi, Giuseppe, Palero, Ferran, Paragamian, Kaloust, Pérez, Toni, Polak, Slavko, Prieto, Carlos E., Turbanov, Ilya, Vailati, Dante & Reboleira, Ana Sofia P. S., 2020, Flourishing in subterranean ecosystems: Euro-Mediterranean Plusiocampinae and tachycampoids (Diplura, Campodeidae), pp. 1-138 in European Journal of Taxonomy 591 on pages 50-51, DOI: 10.5852/ejt.2020.591, http://zenodo.org/record/3659823, {"references":["Sendra A. 2001. Dipluros campodeidos (Diplura: Campodeidae) de las grutas almerienses (Almeria, Espana). Zoologica Baetica 12: 71 - 82."]}