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Underground. Variable Degrees and Variety of Reasons for Cave Penetration in Terrestrial Gastropods Naslednja Postaja: Podzemlje

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Underground. Variable Degrees and Variety of Reasons for Cave Penetration in Terrestrial Gastropods Naslednja Postaja: Podzemlje COBISS: 1.01 NEXT Stop: Underground. Variable degrees AND varietY of reasons for cave penetration in terrestrial gastropods Naslednja postaja: podzemlje. Različne stopnje in različni razlogi prodiranja kopenskih polžev V jame Alexander M. Weigand1,2 Abstract UDC 594.3:551.44 Izvleček UDK 594.3:551.44 Alexander M. Weigand: Next Stop: Underground. Variable Alexander M. Weigand: Naslednja postaja: podzemlje. Razli- degrees and variety of reasons for cave penetration in terres- čne stopnje in različni razlogi prodiranja kopenskih polžev v trial gastropods jame Cave-dwelling animals can be classified based on their occur- Podzemeljske živali lahko opredelimo glede na njihovo pojav- rence in and relationship to the subterranean environment. ljanje v podzemeljskem okolju in odnos do tega okolja. Podatki Subsurface distribution data and studies addressing the initial o razširjenosti živali v podzemlju in študije, ki obravnavajo causes for animals to enter underground habitats are sparse. By vzroke za kolonizacijo podzemlja so redki. Stopnja prodiranja retrieving occurrence data from two voluntary biospeleological kopenskih polžev v jame in morebitni evolucijski vzroki so bili collections in Central Germany, the degree of cave penetration proučevani na podlagi dveh biospeleoloških zbirk v osre dnji in terrestrial gastropods was investigated, thus to infer poten- Nemčiji. Skupno je bilo določenih 66 vrst polžev, ki zaidejo tial evolutionary drivers. In total, 66 identified gastropod spe- v podzemlje, od tega 23 vrst iz temnih predelov podzemlja. cies entered the subterranean environment with 23 of the spe- Čeprav polži kažejo različne stopnje prodiranja v jame, podze- cies also recorded from the dark zone. Gastropods possessed meljska oblika polžev ni bila ugotovljena. Najpogostejši vzorec variable degrees of cave penetration and no obligate cave forms razširjenosti je bil upadanje njihovega pojavljanja od vhoda were observed. A decline of occurrence records from the en- proti notranjosti, čeprav je bilo po nekaj vrst vedno ugotov- trance to the dark zone was the most prominent pattern. Nev- ljenih v vseh treh conah: na vhodu, na prehodu med svetlo in ertheless, several species were collected from all three light temno cono ter v temni coni. Možnih je več razlogov, zaradi zones (i.e. entrance, transition and dark). A variety of potential katerih se pojavijo v podzemlju: izkoriščanje alternativnih reasons can be inferred to explain their subsurface appearance: virov hrane, afotično gibanje na površju, paritvena mesta, exploitation of alternative food sources, aphotic above-ground začasno, sezo nsko ali dolgotrajno zatočišče ter priložnost. movement, mating sites, temporal, seasonal and long-term ref- Rezultati kažejo na pogosto migracijo vrst med posameznimi ugia, and chance. Moreover, the results imply a frequent migra- conami in površjem. Posledično bi bilo potrebno kopenske tion of species between the interconnected light zones and the polže upoštevati pri raziskavah izvora in vnosa alohtonih virov surface. Consequently, terrestrial gastropods should be consid- v jame. ered when investigating the origin and importation of alloch- Ključne besede: Gastropoda (polži), podzemeljsko okolje, thonous resources in caves. ekologija, podatki o razširjenosti, kolonizacija jam, speciacija. Keywords: Gastropoda, subterranean environment, ecology, distribution data, cave colonization, speciation. 1 Goethe-University, Institute for Ecology, Evolution and Diversity, Department of Phylogeny & Systematics, Max-von-Laue Str. 13, 60438 Frankfurt am Main 2 Croatian Biospeleological Society, Demetrova 1, 10000 Zagreb, Croatia, e-mail: [email protected] Received/Prejeto: 08.03.2013 ACTA CARSOLOGICA 43/1, 175–183, POSTOJNA 2014 ALEXander M. Weigand Introduction Since the first nominal description of a cave-dwelling an- tions, unsuitable environmental conditions on the sur- imal, the blind cave salamander Proteus anguinus (Lau- face (Climate Relict Hypothesis) and the exploitation of renti 1768), subterranean organisms have attracted peo- alternative resources (Adaptive Shift Hypothesis) have ple’s attention. Possessing a life in permanent darkness, been discussed as evolutionary drivers (for a review this curiosity immediately can be imagined. But even see Juan et al. 2010). Subsequently, speciation of under- more so, the perception is frequently underscored by a ground lineages may have been triggered by subsurface strongly modified, sometimes bizarre morphology. Cave- dispersal or vicariance events leading to the formation dwelling adaptive features include the (often complete) of allopatric populations. More recently, a combination reduction of eyes and body pigmentation (= reductive of both non-exclusive processes has been considered troglomorphies) resulting in ‘blind albinos’ or the elonga- (Culver et al. 2007; Weigand et al. 2013). Besides studies tion of body appendages and an improved extra-optical addressing speciation processes within the underground sensory system (= constructive troglomorphies; Protas realm or to a surface population, causes for the initial & Jeffery 2012). However, not all cave species exhibit a phase of cave colonizations have been studied less in- troglomorphic appearance (Bichuette & Trajano 2003). tensively (Camp & Jensen 2007) although frequently re- Subterranean organisms can be ecologically classified de- constructed from phylogenetic patterns (Leys et al. 2003; pending on their occurrence in and relationship to the Howarth & Hoch 2005; Cooper et al. 2007). subterranean habitat (Sket 2008). Obligate cave forms (= In this survey, the affinity of gastropod species to eutroglobionts) are restricted and highly adapted to a life the subterranean environment is investigated with the in darkness, whereas in some occasions, epigean animals aim to infer potential evolutionary drivers for their oc- may accidentally occur in the subterranean environment currence in underground habitats. Since gastropods are (= eutrogloxenes). Both extremes of this ecological con- chemically oriented organisms, they do not have to over- tinuum are connected by all levels of intermediate forms. come the disadvantage imposed by visual orientation in Based on the formation of stable or temporal subterra- caves (Culver & Pipan 2009). As a consequence, a certain nean populations, they are commonly referred to as eu- number of surface litter species are expected to occur in troglophiles or subtroglophiles, respectively. caves, thus representing multiple case studies. To explain the initial phases of cave colonization and speciation between surface and subsurface popula- Variable degrees of subterranean penetration Observation records of cave-dwelling terrestrial gastro- single species were collected in multiple zones during a pods were retrieved from two voluntary collections: from visit (i.e. N = 1 but more than one light zone with an oc- the Biospeleological Register of the Hesse Federation for currence record). Taxonomic identification is based on Cave and Karst Research (HES; Zaenker 2008; Reiss et al. conchological and/or anatomical characteristics and was 2009a) and for the region of Rhineland-Palatinate + Saar- primarily performed by S. Zaenker, D. Weber, K. Bogon land (RP + SAR; Weber 2012) (Fig. 1). The geology of the and H. Kappes. Taxonomic nomenclature is according to study area is very heterogeneous but primarily consists of Fauna Europaea (2012). For species with N ≥ 15 and oc- limestone, argillaceous shale, slate and red sandstone. In currence records from all three light zones, a Chi2-test was addition to natural objects (i.e. caves, deep fissures), spe- conducted in SPSS 12.0 (IBM) to test for equal frequency cies were recorded from artificial cavities (i.e. bunkers, distributions. In total, 66 gastropod species were found tunnels, mines) totaling 3352 investigated terrestrial ob- within the subterranean environment (Tab. 1, Fig. 2), jects (i.e. without springs, groundwater samples). During i.e. at least penetrating the entrance zone. An amount of each visit (N), terrestrial gastropods were recorded and 58% (38/66) of the identified species was shared by both classified based on their occurrence in the entrance (E), geographical regions (i.e. HES vs. RP + SAR). Notable transition (T) or dark zone (D) of the subterranean en- exceptions refer to Arion ater, Arion silvaticus, Ena mon- vironment. Thereby, species observations are qualitative tana and Pomatias elegans only recorded from HES and records for a given zone rather than representing count Arion rufus and Phenacolimax major, which were only data of specimens. In some occasions, specimens of a present in RP + SAR. Daudebardia rufa was almost en- 176 ACTA CARSOLOGICA 43/1 – 2014 NEXT Stop: Underground. Variable degrees and varietY of reasons for cave penetration in ... Fig. 1: Overview of the study area. A: The map indicates the geographical location of the two voluntary biospeleological collections for Hesse (HES) and Rhineland-Palatinate + Saarland (RP + SAR) in Germany (colored in red). B: Detailed geographical overview of the study area. tirely restricted to HES (73 observations) with a single constituted 13% of the total records (706/5589), Boett- specimen found in SAR + RP. The dark zone was pen- gerilla pallens and Oxychilus spp. accounted for 56% of etrated by 35% (23/66) of all species. Although they only all dark zone observations. Fig. 2: Terrestrial gastropod species penetrating the subterranean environment. A: Discus rotundatus; B: Monachoides incarnatus; C: Pomatias
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