Taxonomic,phylogenetic and ecological diversity of Niphargus (Amphipoda: Crustacea) in the Hölloch cave system (Switzerland)

Groundwater belongs to the spatially most extensive, but least explored freshwater systems. On a global scale, the species richness of several subterranean invertebrate taxa parallels species richness found in surface waters, while on a local scale species richness hardly exceeds 20 species. This results in a high contribution of groundwater ecosystems to regional β- and γ-diversity, and to a smaller degree to α-diversity, and deserves focused attention. In general, more species are to be found in large cave systems. The second largest cave system in Europe is Hölloch in Switzerland. In this paper we revised the taxonomic, phylogenetic and ecological diversity of the amphipod community in the Hölloch cave system. While previous records listed five geographically widespread species of the genus Niphargus for this cave system, we could not confirm the presence of any of those species, but rather found three highly distinct species new to science. In this paper we describe Niphargus styx sp. nov., Niphargus murimali sp. nov., and Niphargus muotae sp. nov., and suggest that previous records from that cave were probably misidentifications. Although amphipod species richness in this cave system seems to be lower than previously thought in terms of absolute numbers, the cave retained its regional and international importance in terms of nature conservation for multiple reasons. First, all newly described species are probably endemic to this cave system. Second, they are phylogenetically distantly related and exhibit moderate to high phylogenetic diversity. Third, the species, as inferred from their functional morphology, are also ecologically highly divergent. Based on geographic distribution of their nearest relatives, we hypothesize that the cave system was most likely independently colonized from North, West and South and that the pre-adapted ancestors occupied different ecological niches within the system.

http://zoobank.org/urn:lsid:zoobank.org:pub:A19309E5-C06B-4844-A4D8-7571F05F25C9     

ECOMORPHOLOGICAL CONVERGENCE OF CAVE COMMUNITIES

Extreme selective environments are commonly believed to funnel evolution toward a few predictable outcomes. Caves are well‐known extreme environments with characteristically adapted faunas that are similar in appearance, physiology, and behavior all over the world, even if not closely related. Morphological diversity between closely related cave species has been explained by difference in time since colonization and different ecological influence from the surface. Here, we tested a more classical hypothesis: morphological diversity is niche‐based, and different morphologies reflect properties of microhabitats within caves. We analyzed seven communities with altogether 30 species of the subterranean amphipod (crustacean) genus Niphargus using multivariate morphometrics, multinomial logit models cross‐validation, and phylogenetic reconstruction. Species clustered into four distinct ecomorph classes—small pore, cave stream, cave lake, and lake giants—associated with specific cave microhabitats and of multiple independent phylogenetic origins. Traits commonly regarded as adaptations to caves, such as antenna length, were shown to be related to microhabitat parameters, such as flow velocity. These results demonstrate that under the selection pressure of extreme environment, the ecomorphological structure of communities can converge. Thus, morphological diversity does not result from adaptive response to temporal and ecological gradients, but from fine‐level niche partitioning.     

The subterranean genus Niphargus (Crustacea, Amphipoda) in the Middle East: A faunistic overview with descriptions of two new species

The Middle East is at the southeastern borderline of the range of the subterranean amphipod Niphargus. The review of new and published data identified two new species and set the guidelines for the future research in the area. The genus in this part of the world seems to be insufficiently studied. The taxonomic status of Niphargus valachicus, population identified as N. spoeckeri and some populations, identified as N. nadarini, need to be reviewed. According to present data, we expect the highest diversity in Western Turkey. The eventual new records of the genus in the Middle East can be expected from those areas where even the longest periods of drought in the recent geological history did not affect the water supply.     

The effect of light on oxygen consumption in two amphipod crustaceans-the hypogean Niphargus stygius and the epigean Gammarus fossarum

The effects of light on the metabolic rates of the hypogean amphipod Niphargus stygius and the epigean amphipod Gammarus fossarum were compared by measuring oxygen consumption and respiratory electron transport system (ETS) activity. They were exposed to light intensities of 720 and 4700 lx at 10°C. Oxygen consumption increased significantly in N. stygius exposed to both low and high intensities of light, but no significant increase was observed in G. fossarum at either intensity. The increase of oxygen consumption in N. stygius was significantly greater at the higher light intensity. This indicates a stress response in which exploitation of half the metabolic potential for energy production in N. stygius during exposure to high light intensity constitutes an adverse effect on its metabolism, since this species usually uses less than 25% of its total metabolic potential for standard metabolic demands.     

Comparative study of electron transport system activity and oxygen consumption of amphipods from caves and surface habitats

1. Respiratory electron transport system (ETS) activity and oxygen consumption were measured at 10 °C for the hypogean amphipods Niphargus stygius (from two epikarst locations) and N. krameri (from a sinking river in a cave) and the epigean amphipod Gammarus fossarum (inhabiting a spring and a river). 2. ETS activity and oxygen consumption were compared between caves and the two surface locations, and between hypogean and epigean amphipods. ETS activities were found not to differ between animals from different locations, or between epigean and hypogean amphipods. As expected, the oxygen consumption of N. krameri and N. stygius was lower than that of G. fossarum. 3. The high ratio between ETS activity and oxygen consumption (ETS/R ratio) showed that N. krameri and N. stygius possess high metabolic potential that can be exploited for energy recovery and rapid restoration of body reserves immediately following the appearance of favourable conditions (food and/or oxygen). In contrast, G. fossarum exhibited a low ETS/R ratio, indicating exploitation of a great proportion of the metabolic potential for standard metabolism. 4. Feeding in N. stygius increased oxygen consumption but not ETS activity, and thus fed N. stygius had a much lower ETS/R ratio than starved ones. 5. A relatively high metabolic potential (i.e. high ETS/R ratio) may be an adaptation to hypogean life in amphipods, in that it improves survival under poor food and/or oxygen conditions.     

Taxonomy and biogeography of <Emphasis Type="Italic">Niphargus steueri</Emphasis> (Crustacea: Amphipoda)

Niphargus steueri (Niphargidae) comprises a complex of four subspecies (N. s. steueri, N. s. kolombatovici, N. s. subtypicus, and N. s. liburnicus), the morphology and distribution of which have been poorly known until now. New diagnostic characters of the species and its four subspecies are presented and illustrated. The species is distributed along the major part of the Dinaric Karst, between Slovenia in the northwest and Bosnia and Herzegovina in the southeast. The distribution of the four subspecies approximately resembles the distribution of the evolutionary lineages of the subterranean amphibian Proteus anguinus and the Dinaric lineage of the cave shrimp Troglocaris agg. anophthalmus. Niphargus s. steueri is restricted to the Istran Peninsula; N. s. subtypicus is distributed in southeast Slovenia and northwest Croatia; N. s. liburnicus is known from two disjunctive localities, one on the island of Krk (Croatia) and the other in Gorizia (Italy); and N. s. kolombatovici is restricted to Dalmacija and Herzegovina. The somewhat variable putative synapomorphies of N. steueri probably suggest that the group is old and that the present distribution pattern is a result of historical events, possibly events in the Miocene Dinaride Lake system. Two populations of N. s. kolombatovici and one population of N. s. subtypicus deviate from the general distributional pattern and may belong to cryptic taxa that cannot be distinguished on the basis of morphology. Both hypotheses corroborate with the estimated times of divergence and with the number of independent lineages in the similarly distributed but unrelated stygobionts Proteus and Troglocaris.     

Can heterochrony help explain the high morphological diversity within the genus Niphargus (Crustacea: Amphipoda)?

Postembryonic development of nine species of Niphargus (Crustacea: Amphipoda) was studied, with spine development and shifts in allometric growth being combined in developmental sequences that were compared across species. The developmental sequences show high diversity with respect to the position of individual events in the sequence, as well as a high frequency of events being inapplicable in some species. Within Niphargus, the highest degree of independence between events occurs mainly in early mid-aged instars, where the shifts in position are both the largest in magnitude and also the most frequent in occurrence. Constructive troglomorphic features of subterranean species were inferred to develop more because of accelerated growth rather than a delayed offset of growth. Shifts in both relative timing and growth rate appeared to have played a role in the evolution of sexually dimorphic elongated appendages. Growth patterns differed greatly between species for individual sexually dimorphic and troglomorphic traits, hinting at their possible independent origin. The independence between developmental events that was generally apparent might indicate the existence of an important genetic basis for the extreme intrageneric morphological variation arising from numerous highly variable body parts being combined in a ‘mosaic’ manner. Moreover, we suggest that the overall diversity of the genus might actually be underestimated in light of possible convergent features accompanied by ‘cryptic’ speciation. We conclude that both sequential and growth heterochrony appear to have played a key role in the evolution of Niphargus, the most diverse genus of freshwater amphipods. A preliminary list of heterochronic characters is provided.     

A gradient of interstitial Niphargus rhenorhodanensis populations in two karst/floodplain transition zones of the French Jura

The Bou-Rouch technique was used to sample two small alluvial plains in two different karst areas. One was mainly supplied by karst water (Verna), the other by both karst and surface water (Pissoir).Physical and chemical characteristics varied with time (seasons) and location of sampling sites at both stations.The distribution of Niphargus rhenorhodanensis populations was influenced by the geographical characteristics of the outlets. Only a small number of individuals was found near the outlet and none in the floodplain when the outlet was located at a base-level (Verna). Individuals were more numerous and their presence constant at all sites and dates when the outlet was located at the overflow level of the massif (Pissoir).