Spatial patterns of diatom diversity and community structure in ancient Lake Ohrid

For effective lakes’ management, high-frequent water quality data on a synoptic scale are essential. The aim of this study is to test the suitability of the latest generation of satellite sensors to provide information on lake water quality parameters for the five largest Italian subalpine lakes. In situ data of phytoplankton composition, chlorophyll-a (chl-a) concentration and water reflectance were used in synergy with satellite observations to map some algal blooms in 2016. Chl-a concentration maps were derived from satellite data by applying a bio-optical model to satellite data, previously corrected for atmospheric effects. Results were compared with in situ data, showing good agreement. The shape and magnitude of water reflectance from different satellite data were consistent. Output chl-a concentration maps, show the distribution within each lake during blooming events, suggesting a synoptic view is required for these events monitoring. Maps show the dynamic of bloom events with concentration increasing from 2 up to 7 mg m^?3 and dropping again to initial value in less than 20 days. Latest generation sensors were shown to be valuable tools for lakes monitoring, thanks to frequent, free of charge data availability over long time periods.     

Rapid evolution of an ancient lake species flock: Freshwater limpets (Gastropoda: Ancylidae) in the Balkan Lake Ohrid

Ancient lakes have long been recognized as evolutionary theatres and hot spots of endemism; the evolution of their morphologically often highly diverse species flocks has received much attention. However, as each ancient lake has its own geological and evolutionary history, modes of speciation may differ from system to system. Ancient lakes can act as evolutionary reservoirs that assure the survival of relict species, but at the same time extant species may evolve through intralacustrine speciation. Other aspects of interest are the actual rates of immigration, diversification or extinction as well as the temporal framework of morphological change. Many of these questions have been addressed in the African (e.g. Lake Tanganyika) and Asian (e.g. Lake Baikal) ancient lakes. For an European ancient lakes (e.g. Lakes Ohrid and Prespa), such studies are largely missing. In the present paper, extraordinarily shaped endemic freshwater limpets of the genus Ancylus from the Balkan Lake Ohrid are used in a phylogeographic and phylogenetic context to test whether they represent an ancient lake species flock, to study the mode of speciation, and to assess the timing of morphological change. Based on DNA data from two mitochondrial genes (COI, LSU rDNA), it has been found that the Lake Ohrid Ancylus species form an endemic monophyletic group. In addition, the lake's feeder springs are inhabited by another, undescribed Ancylus species. All other studied waterbodies within the watershed do not support their own Ancylus lineages but are inhabited by a widespread Mediterranean taxon. The split between the species endemic to the lake and its sister taxon is dated to 1.4±0.6 million years ago. The study presents the first genetic confirmation for the existence of a species flock in a European ancient lake. Contrary to the prevailing opinion it shows that, concerning Ancylus, Lake Ohrid represents a site of intralacustrine speciation rather than an evolutionary reservoir. Moreover, it provides the first evidence for rapid morphological change in an European ancient lake species flock. See also Electronic Supplement at: http://www.senckenberg.de/odes/06-12.htm.     

Ancient Lake Ohrid: biodiversity and evolution

Worldwide ancient lakes have been a major focal point of geological, biological, and ecological research, and key concepts in, for example, evolutionary biology are partly based on ancient lake studies. Ancient lakes can be found on most continents and climate zones with most actual or putative ancient lakes in Europe being restricted to the Balkan Region. The arguably most outstanding of them is the oligotrophic and karstic Lake Ohrid, a steep-sided graben of rift formation origin situated in the central Balkans. Here, an attempt is made to summarize current knowledge of the geological, limnological, and faunal history of Lake Ohrid. Additionally, existing data on biodiversity and endemism in Lake Ohrid are updated and evaluated, and patterns and processes of speciation are reviewed in the context of the Ohrid watershed, including its sister lake, Lake Prespa. Whereas the geological history of the Ohrid Graben is relatively well studied, there is little knowledge about the limnological and biotic history of the actual lake (e.g., the age of the extant lake or from where the lake first received its water, along with its first biota). Most workers agree on a time frame of origin for Lake Ohrid of 2–5 million years ago (Mya). However, until now, the exact limnological origin and the origin of faunal or floral elements of Lake Ohrid remain uncertain. Two largely contrasting opinions either favour the theory of de novo formation of Lake Ohrid in a dry polje with a spring or river hydrography or a palaeogeographical connection of Lake Ohrid to brackish waters on the Balkan Peninsula. Whereas neither theory can be rejected at this point, the data summarized in the current review support the de novo hypothesis. An assessment of the fauna and flora of Lake Ohrid confirms that the lake harbours an incredible endemic biodiversity. Despite the fact that some biotic groups are poorly studied or not studied at all, approximately 1,200 native species are known from the lake, including 586 animals, and at least 212 species are endemic, including 182 animals. The adjusted rate of endemicity is estimated at 36% for all taxa and 34% for Animalia. In terms of endemic biodiversity, Lake Ohrid is with these 212 known endemic species and a surface area of 358 km² probably the most diverse lake in the world, taking surface area into account. Preliminary phylogeographical analyses of endemic Lake Ohrid taxa indicate that the vast majority of respective sister taxa occurs in the Balkans and that therefore the most recent common ancestors of Ohrid- and non-Ohrid species likely resided in the region when Lake Ohrid came into existence. These data also indicate that there is relatively little faunal exchange and overlap between Lake Ohrid and its sister lake, Lake Prespa, despite the fact that the latter lake is a major water supplier for Lake Ohrid. Studies on selected species flocks and scatters, mostly in molluscs, point towards the assumption that only few lineages originally colonized Lake Ohrid from the Balkans and that the majority of endemic species seen today probably started to evolve within the lake during the early Pleistocene. Within the Ohrid watershed, endemism occurs at different spatial and taxonomic scales, ranging from species endemic to certain parts of Lake Ohrid to species endemic to the whole watershed and from subspecies to genus level and possibly beyond. Modes of speciation in the Ohrid watershed are largely affected by its degree of isolation. Observational evidence points towards both allopatric (peripatric) and parapatric speciation. Though sympatric speciation within a habitat is conceivable, so far there are no known examples. Today, the lake suffers from increasing anthropogenic pressure and a “creeping biodiversity crisis”. Some endemic species presumably have already gone extinct, and there are also indications of invasive species penetrating Lake Ohrid. The comparatively small size of Lake Ohrid and the extremely small range of many endemic species, together with increasing human pressure make its fauna particularly vulnerable. It is thus hoped that this review will encourage future research on the ecology and evolutionary biology of the lake’s taxa, the knowledge of which would ultimately help protecting this unique European biodiversity hot spot. Guest editors: T. Wilke, R. V?in?l? & F. Riedel Patterns and Processes of Speciation in Ancient Lakes: Proceedings of the Fourth Symposium on Speciation in Ancient Lakes, Berlin, Germany, September 4–8, 2006     

Concurrent evolution of ancient sister lakes and sister species: the freshwater gastropod genus <Emphasis Type="Italic">Radix</Emphasis> in lakes Ohrid and Prespa

Ancient sister lakes are considered to be ancient lakes lying in close geographic proximity, sharing a related origin and significant time of co-existence, usually having hydrological connection as well as a balanced degree of faunal overlap and distinctness. A paradigm for studying sister lake relationships are the ancient lakes Ohrid and Prespa in the Balkans, which are characterized by high degrees of endemicity. Three general patterns of endemic species can be distinguished for these lakes: (1) taxa that are endemic to either lake, with no close relatives in the respective sister lake, (2) closely related but distinct endemic taxa in both lakes (sister species) and (3) shared endemic taxa occurring in both lakes. In the present paper, two endemic freshwater pulmonate gastropod species, Radix relicta (Lake Ohrid) and R. pinteri (Lake Prespa), are used to study the evolution of presumed sister species based on biogeographical and comparative DNA data from world-wide Radix taxa. Phylogenetic, phylogeographical and parametric bootstrap analyses all suggest a sister group relationship of R. relicta and R. pinteri (pattern 2 of endemic diversity). Sister to these two taxa is the widespread R. ampla, which does not occur in the vicinity of lakes Ohrid and Prespa. The southern feeder spring complexes of Lake Ohrid are inhabited by another lineage (Radix sp. 1), which resembles Radix relicta in morphology/anatomy. For Lake Prespa, the widespread R. auricularia was reported in addition to the endemic R. pinteri. Comparative phylogenetic data favour a western Adriatic zoogeographical affinity of lakes Ohrid and Prespa over an Aegean-Anatolian faunal connection. The status of lakes Ohrid and Prespa as sister lakes is evaluated in the light of current knowledge on gastropod speciation and endemism in these hotspots of biodiversity.     

Amassing diversity in an ancient lake: evolution of a morphologically diverse parthenogenetic gastropod assemblage in Lake Malawi

Exceptional ecological niche diversity, clear waters and unique divergent selection pressures have often been invoked to explain high morphological and genetic diversity of taxa within ancient lakes. However, it is possible that in some ancient lake taxa high diversity has arisen because these historically stable environments have allowed accumulation of lineages over evolutionary timescales, a process impossible in neighbouring aquatic habitats undergoing desiccation and reflooding. Here we examined the evolution of a unique morphologically diverse assemblage of thiarid gastropods belonging to the Melanoides polymorpha'complex' in Lake Malawi. Using mitochondrial DNA sequences, we found this Lake Malawi complex was not monophyletic, instead sharing common ancestry with Melanoides anomala and Melanoides mweruensis from the Congo Basin. Fossil calibrations of molecular divergence placed the origins of this complex to within the last 4 million years. Nuclear amplified fragment length polymorphism markers revealed sympatric M. polymorpha morphs to be strongly genetically differentiated lineages, and males were absent from our samples indicating that reproduction is predominantly parthenogenetic. These results imply the presence of Lake Malawi as a standing water body over the last million years or more has facilitated accumulation of clonal morphological diversity, a process that has not taken place in more transient freshwater habitats. As such, the historical stability of aquatic environments may have been critical in determining present spatial distributions of biodiversity.     

Invaders or endemics? Molecular phylogenetics,biogeography and systematics of Dreissena in the Balkans

1. Zebra mussels and their relatives (Dreissena spp.) have been well studied in eastern, central and western Europe as well as in North America, because of their invasiveness and economic importance. Much less is known about the biology and biogeography of indigenous (endemic) taxa of Dreissena, in the Balkans. A better knowledge of these taxa could help us (i) understand the factors triggering invasiveness in some taxa and (ii) identify other potentially invasive species. 2. Using a phylogenetic approach (2108 base pairs from three gene fragments), Dreissena spp. from natural lakes in the Balkans were studied to test whether invasive Dreissena populations occur in such lakes on the Balkan Peninsula, whether Dreissena stankovici really is endemic to the ancient Lakes Ohrid and Prespa, and to infer the phylogenetic and biogeographical relationships of Balkan dreissenids. 3. No invasive species of Dreissena, such as Dreissena polymorpha, were recorded. The supposedly ‘endemic’D. stankovici is not restricted to the ancient Lakes Ohrid and Prespa, but is the most widespread and dominant species in the west‐central Balkans. Its southern sister taxon, Dreissena blanci, occurs sympatrically with D. stankovici in Lakes Prespa, Mikri Prespa and Pamvotis. Both species are classified into the subgenus Dreissena (Carinodreissena) of which the subgenus Dreissena (Dreissena) (which includes the invasive D. polymorpha) is the sister taxon. Dreissena blanci and D. stankovici are considered to represent distinct species. 4. On a global scale, the two Balkan species have small ranges. An early Pliocene time frame for the divergence of the subgenera Carinodreissena and Dreissena is discussed, as well as potential colonization routes of the most recent common ancestor of Carinodreissena spp. 5. The ambiguous taxonomy of dreissenids in the Balkans is addressed. As nominal D. blanci presbensis from Lake Prespa has nomenclatural priority over D. stankovici, the correct name for the latter taxon should be Dreissena presbensis.     

Palaearctic gastropod gains a foothold in the dominion of endemics: range expansion and morphological change of Lymnaea (Radix) auricularia in Lake Baikal

In and around the endemic-dominated Lake Baikal, palaearctic species are generally restricted to shallow, sheltered bays and in- and out-flowing river floodplains. However, we observed populations of the palaearctic snail Lymnaea (Radix) auricularia on the steep, rocky littoral of Lake Baikal proper. We compared the morphology of 542 shells sampled from this new habitat with potential source populations from conventional habitats. A size-free Discriminant Analysis indicated a strong morphological differentiation of the newly established populations from their likely sources. The new populations had a more compact shell shape with a wide aperture, which may be advantageous in wave-exposed habitats where a firm attachment to the substrate is needed. Shells from the conventional habitats were more elongated, with a narrow aperture, which may be advantageous in habitats that have a dry period where retreating into the mud is required and water loss should be limited. These results may suggest that selection is acting on shell shape in Lake Baikal. The apparent recent arrival of this pandemic gastropod in a habitat previously dominated by endemics constitutes a potential ecological threat and an alert to possible ecological change.     

Invaders eating invaders: exploitation of novel alien prey by the alien shimofuri goby in the San Francisco Estuary,California

The shimofuri goby (Tridentiger bifasciatus), which is native to Asian estuaries, was recently introduced to the San Francisco Estuary, California, USA. We conducted gut content analyses to examine the gobys feeding ecology in this highly invaded estuary. Shimofuri gobies were generalist predators on benthic invertebrates, consuming seasonally abundant prey, especially amphipods (Corophium spp.). In addition, shimofuri goby utilized two novel prey items not exploited by other resident fishes – hydroids (Cordylophora caspia) and barnacle (Balanus improvisus) cirri, both of which are alien. The shimofuri gobys feeding ecology appears well-suited to the fluctuating environment of the San Francisco Estuary and may partially explain observed increases in shimofuri goby abundance compared with declines in populations of some native species.     

Optimal conservation strategy in fluctuating environments with species interactions: resource-enhancement of the native species versus extermination of the alien species

Alien species are often a major threat to native species. We consider optimal conservation strategies for a population whose viability is affected both by an alien species (such as a competitor, a predator, or a pathogen) and by random fluctuations of the environment (e.g. precipitation, temperature). We assume that the survivorship of the native population can be improved by providing resources such as food and shelter, and also by an extermination effort that decreases the abundance of the alien species. These efforts decrease the extinction probability of the native population, but they are accompanied by economic costs. We search for the optimal strategy that minimizes the weighted sum of the extinction probability and the economic costs over a single year. We derive conditions under which investment should be made in both resource-enhancement and extermination, and examine how the optimal effort levels change with parameters. When the optimal strategy includes both types of efforts, the optimal extermination effort level turns out to be independent of the density and economic value of the native species, or the variance of the environmental fluctuation. Furthermore, the optimal resource-enhancement effort is then independent of the density of the alien species. However, the parameter dependencies greatly change if one of the efforts becomes zero. We also examine the situation in which the impact of the alien species is uncertain. The optimal extermination effort increases with the uncertainty of this impact except when the cost of extermination is very high.     

Patterns of genetic differentiation and population history of endemic isopods (Asellidae) from ancient Lake Ohrid: combining allozyme and mtDNA data

Ancient lakes as places of extensive speciation processes have been characterized by a high degree of endemicity and biodiversity. The most outstanding European ancient lake is the oligotrophic and karstic Balkan Lake Ohrid. The lake is inhabited by a number of endemic species, but their evolutionary history is largely unresolved. in the present study, the genetic structure, gene genealogy and demographic history of the representatives of the Ohridian endemic Proasellus species were studied using both biparentally (allozyme loci) and maternally (partial mitochondrial cytochrome oxidase subunit I gene) inherited markers. Both data sets gave similar results and supported discrepancies among genetic differentiation, the current morphology-based taxonomy and bathymetric segregation. Horizontal distribution of endemic Proasellus species (Lake Ohrid vs adjacent feeder springs) within the lake presumably promote parapatric speciation whereas the main role of vertical barriers into diversification processes was not fully supported. The analyses of demographic history suggested the decline of endemic isopod populations. The radiation of endemic Proasellus populations within the lake could have started from the sublittoral/profundal zone towards the littoral or in the opposite direction — from the littoral to the profundal. Our analyses did not exclude both possibilities.     

Larval apical sensory organ in a neritimorph gastropod,an ancient gastropod lineage with feeding larvae

The Neritimorpha is an ancient clade of gastropods that may have acquired larval planktotrophy independently of the evolution of this developmental mode in other gastropods (caenogastropods and heterobranchs). Neritimorphs are therefore centrally important to questions about larval evolution within the Gastropoda, but there is very little information about developmental morphology through metamorphosis for this group. We used immunolabeling (antibodies binding to acetylated α-tubulin and serotonin) and serial ultrathin sections for transmission electron microscopy to characterize the apical sensory organ in planktotrophic larvae of a marine neritimorph. The apical sensory organ of gastropod larvae is a highly conserved multicellular sensory structure that includes an apical ganglion and often an associated ciliary structure. Surprisingly, the apical ganglion of Nerita melanotragus (Smith, 1884) does not have typical ampullary neurons, a type of sensory neuron consisting of a cilia filled inpocketing that has been described in all other major gastropod groups. N. melanotragus has cilia-filled pockets embedded within the apical ganglion, but these so-called “sensory cups” are cassettes of multiple cells: one supporting cell and up to three multiciliated sensory cells. We suggest that an internalized pocket that is filled with cilia and open to the exterior via a narrow pore may be essential architectural features for whatever sensory cues are detected by ampullary neurons and sensory cups; however, morphogenesis of these features at the cellular level has undergone evolutionary change. We also note a correlation between the number of sensory elements consisting of cilia-filled pockets within the larval apical sensory organ of gastropods and morphological complexity of the velum or length of the trochal ciliary bands.     

A gastropod from the tropical Atlantic becomes an established alien in the Mediterranean

We report the occurrence of an established population of the gastropod Mitrella psilla (Duclos, 1846), native on west African coasts of Mauritania, Senegal and Angola, on the shore of La Goulette, Bay of Tunis. The molluscan species of tropical West African affinity hitherto ascertained in the Mediterranean are not aliens, but have a native range which encompasses at least part of the Alboran Sea since historical times. Mitrella psilla appears therefore as the first instance of a mollusc with a tropical Atlantic origin being added to the Mediterranean fauna. The species was found near an important commercial harbour and is presumed to have been introduced by shipping.     

Eradications of invasive alien species in Europe: a review

Eradication of alien species is a key conservation tool to mitigate the impacts caused by biologic invasions. The aim of the present paper is to review the eradications successfully completed in Europe and to discuss the main limits to a wider application of this management option in the region. On the basis of the available literature – including conference proceedings, national reports to the Bern Convention, etc. – a total of 37 eradication programmes have been recorded. Thirty-three eradications were carried out on islands and four on the mainland. The rat (Rattus spp.) has been the most common target (n = 25, 67%), followed by the rabbit (n = 4). In many cases, these eradications determined a significant recovery of native biodiversity. Differently to other regions of the world, no eradications of alien invertebrates and marine organisms have been recorded; regarding invasive alien plants, it appears that only some very localized removals have been completed so far in Europe. The limited number of eradications carried out in Europe so far is probably due to the limited awareness of the public and the decision makers, the inadequacy of the legal framework, and the scarcity of resources. Synthetic guidelines for improving the ability of European states to respond to aliens incursions are presented. This revised version was published online in July 2006 with corrections to the Cover Date.     

The neglected side of speciation in ancient lakes: phylogeography of an inconspicuous mollusc taxon in lakes Ohrid and Prespa

The morphologically remarkable endemic fauna within ancient lakes has received much attention in the literature. More inconspicuous taxa, however, often lack detailed molecular and morphometrical examination, although their proportion of the endemic fauna of an ancient lake must not be underestimated. Consequently, a better understanding of evolutionary patterns and processes within these lakes requires more knowledge about the often-neglected inconspicuous taxa. In the present study, we focus on the notoriously cryptic pea clam genus Pisidium (Bivalvia: Sphaeriidae). Though the genus is widely distributed, most endemic species are reported only from ancient lakes, including the European ancient sister lake system of Ohrid and Prespa on the Balkan Peninsula. Here we test for the first time hypotheses on the evolution of the endemic pea clams in this European biodiversity hotspot by molecular means. Combining a broad 16S phylogeny (comprising most European pea clam species), network analyses and morphometrical analyses, we found interesting biogeographical patterns and provide evidence for cryptic species in both lakes. Furthermore, we confirmed the proposed sister-species relationship of the endemics P. edlaueri in Lake Ohrid and P. maasseni in Lake Prespa, and we suggest scenarios of the endemic pea clam evolution within both lakes. The patterns of speciation found in the genus Pisidium are compared to patterns in morphologically distinct molluscan groups in lakes Ohrid und Prespa.     

Tramps, narrow endemics and morphologically cryptic species in the epiphyllous liverwort Diplasiolejeunea

Diplasiolejeunea is a pantropical, epiphytic genus of leafy liverworts that occurs from the lowlands to more than 4000m altitude. Phylogenetic analyses of a molecular dataset consisting of three markers (nuclear ribosomal ITS region, plastidic trnL-F region and rbcL gene) and 122 accessions (plus two outgroups, Colura and Cololejeunea) indicate that the evolutionary diversity of Diplasiolejeunea is underestimated by current morphology-based classification. Four morphologically semi-cryptic species have been recovered. The molecular phylogenies support a deep split into a Neotropical and a Paleotropical clade, the latter structured into Australasian, Asian and Afromadacascan lineages. Presented results confirm the ranges of two pantropical species (D. cavifolia, D. rudolphiana), provide evidence for dispersal from the Neotropics into the Paleotropics, indicate speciation along altitudinal gradients and demonstrate extensive morphological homoplasy. We propose a revised supraspecific classification of Diplasiolejeunea into a predominantly Paleotropical subgenus Physolejeunea and predominantly Neotropical subgenera Austrolejeuneopsis and Diplasiolejeunea, the former containing mainly epiphytic species, the latter mainly epiphylls. Several clades are supported by combinations of morphological character states, and could be assigned to sections at some later point. This is the first comprehensive phylogeny of a largely epiphyllous genus of liverworts.     

Invaders on the move: parasitism in the sexual galls of four alien gall wasps in Britain (Hymenoptera: Cynipidae)

Four alien, host-alternating cynipid gall wasps are established in the British Isles: Andricus kollari, A. lignicola, A. quercuscalicis and A. corruptrix. Their current distributions divide Britain into four zones; all four invaders are present in the south and south-east of Britain, whereas only A. kollari is present in the far north of Scotland. The rank order, according to how far north the distributions of the four invading cynipid species reach, is: A. corruptrix→ A. quercuscalicis→ A. lignicola→ A. kollari. The life cycles of all four cynipid species involve a sexual generation in spring on Quercus cerris, and an agamic generation in autumn on Q. robur. Here we studied the parasitoid attack by four pteromalid species on the sexual generations of the invaders. We collected and reared the galls of all four species at eight sites from the south to the north of the country (two sites per zone). The geographical locations of the sites reflect the invasion history and thus the residence time of the alien species in the four zones. At each site we measured the densities of all host galls on Q. cerris and of both obligatory host-tree species. We also took a series of measures, such as host-tree density and mean host-tree size, to further characterize the tree stands. These measures are referred to as local parameters. Host densities varied between sites and between years. In A. kollari, galling rates were highest in the middle of the country (zone 2) in 1994, whereas in 1995 they increased from the south to the north. In A. lignicola, galling rates in both years were lowest at the sites in zone 3 (closest to its distribution boundary). In A. quercuscalicis, galling rates were found to be lowest at the site most to the north-west in both years, again the one furthest away from the area where this species was first recorded. Mortality caused by parasitoid attack differed from less than 10% to as high as 70% and varied between host species, sites and years. In four out of six cases the historical/regional variables (north/south and east/west) correlated significantly with parasitoid attack rates that were characteristically lowest at sites close to the distribution boundaries. Of the local factors, we found parasitoid attack rates correlated negatively in one case with host density, whereas they correlated positively in four cases with the density of alternative hosts of the parasitoids. In one of the models the local density of Q. cerris trees correlated negatively with parasitoid attack on A. quercuscalicis. For all three host species the terms retained in the minimal adequate models obtained for 1994 and 1995 differed, which might indicate that these communities of native parasitoids and invading host have not yet settled in any definite structure.     

Hypoxia tolerance of three native and three alien species of bitterling inhabiting Lake Kasumigaura,Japan

Differences in hypoxia tolerance among three native and three alien bitterling species were examined by means of field surveys and aquarium experiments. I caught fish in minnow traps and measured environmental factors such as dissolved oxygen and current velocity once a month between June and November 2006 at 30 points around Lake Kasumigaura, Japan. In addition, aquarium experiments were carried out to compare aquatic surface respiration thresholds among the bitterling species. Much more individuals of an alien species, Rhodeus ocellatus ocellatus, was caught at the least oxygen range than expected in the field, and showed the highest hypoxia tolerance in the laboratory experiments. Another alien bitterling, Acheilognathus rhombeus, also showed higher hypoxia tolerance than the three native bitterling species. Lake Kasumigaura is well known for eutrophication and water masses with low dissolved oxygen have often been observed there. Differences in hypoxia tolerance among bitterling species might have affected bitterling assemblage structure in Lake Kasumigaura, which is currently characterized by the dominance of R. o. ocellatus.     

The role of barriers and gradients in differentiation processes of pyrgulinid microgastropods of Lake Ohrid

Ancient Lake Ohrid is characterized by vertical (bathymetrical) zones within the lake, presumably promoting allopatric speciation due to barriers or parapatric speciation along gradients. Examples within the lake include the belt of Chara algae as well as the shell zone, both presumably impeding migrations of benthic invertebrates. Three potential cases of vertical differentiation leading to distinct depth forms have been reported for the gastropod subfamily Pyrgulinae (Caenogastropoda: Hydrobiidae): Ginaia munda ssp., Macedopyrgula spp. and Ochridopyrgula macedonica ssp. Based on DNA data of the COI gene from a total of 145 specimens, this article aims at investigating the vertical differentiation within these depth forms and thus patterns of speciation in Lake Ohrid. An initial morphometric analysis showed a clear correlation of shell shape and collecting depth for Ginaia munda ssp. and Macedopyrgula spp. This morphological trend is largely reflected in the genetic structure of the respective taxa. The data presented here indicate the existence of strong gradients of abiotic and biotic factors in Lake Ohrid rather than distinct barriers. Therefore, parapatric speciation may be the predominant form of differentiation of benthic invertebrates in the lake. Incomplete lineage sorting, hybridization and phenotypic plasticity possibly caused by epigenetic mechanisms are discussed as possible reasons for the incongruence between geno- and phenotype observed in few specimens of Ginaia munda ssp. and Macedopyrgula spp. For the third taxon, Ochridopyrgula macedonica ssp., morphometric and genetic analyses revealed only weak support for the previously proposed depth forms. However, a horizontal differentiation of lake and spring populations was revealed instead, and parapatric and allopatric differentiations are discussed in this taxon.