Holocene mammal extinctions in the Carpathian Basin: a review

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Characteristics of mature male parr in the northernmost Atlantic salmon populations

The incidence, relative abundance and biological characteristics of mature male parr of Atlantic salmon Salmo salar were studied in two subarctic (69–70 N°) river systems in the northernmost part of Scandinavia. The proportion of mature males was low (0–25%) among ≥1 year‐old parr, and clearly lower than reported earlier in more southern rivers. The ages of mature males ranged from 1+ to 6 + year, the most abundant age group was 3 + year. Both age structure and size of the mature male parr varied between rivers and years. Mature male parr in the main stem were younger and also smaller than the parr of the same age in the tributaries.     

Natural and anthropogenic influences on the population structure of white‐tailed eagles in the Carpathian Basin and central Europe

European populations of the white‐tailed eagle Haliaeetus albicilla suffered a drastic decline during the 20th century. In many countries, only a few dozen breeding pairs survived or the species disappeared completely. By today, the populations have recovered, naturally or through restocking (e.g. in Scotland or the Czech Republic). In the Carpathian Basin, which is now a stronghold in southern Europe for the species in the southern part of the distribution range with more than 500 breeding pairs, only about 50 pairs survived the bottleneck. This region provides important wintering places for individuals arriving from different regions of Eurasia. In the present study, we investigated 249 DNA samples from several European countries, using 11 microsatellites and mitochondrial control region sequences (499 bp), to answer two main questions: 1) did the Carpathian Basin population recover through local population expansion or is there a significant gene flow from more distant populations as well? 2) Does the Czech population show signs in its genetic structure of the restocking with birds of unknown origin? Our microsatellite data yielded three genetically separate lineages within Europe: northern, central and southern, the latter being present exclusively in the Carpathian Basin. Sequencing of mitochondrial DNA revealed that there is one haplotype (B12) which is not only exclusive to the Carpathian Basin but it is frequent in this population. Our results suggest that in accordance with the presumably philopatric behaviour of the species, recovery of the Carpathian Basin population was mainly local, but some of the wintering birds coming from the northern and central populations contributed to its genetic composition as well. We detected considerably higher proportions of northern birds within the Czech Republic compared to the neighbouring areas, making it likely that parents of the reintroduced birds came from northern populations.     

Phorophyte preferences of epiphytic bryophytes in a stream valley in the Carpathian Basin

Abstract

Epiphytic bryophytes were studied on two tree species with different bark characteristics (Alnus glutinosa and Carpinus betulus) in two vertical ranges (0–60 and 60–200 cm height above ground on the trunk) in a stream valley in the Carpathian Basin (N.W. Hungary). There is a significant difference in the species composition of the trees. Dicranum montanum and Platygyrium repens occurred significantly more often on Alnus glutinosa, whereas Frullania dilatata and Radula complanata were more often on Carpinus betulus in both vertical ranges. Orthotrichum species were typical only in the vertical range 60–200 cm on Carpinus betulus, whereas Plagiothecium nemorale and Metzgeria furcata were typical of Alnus glutinosa and Carpinus betulus, respectively, in the vertical range 0–60 cm. Although most of the bryophyte species have significant phorophyte preferences, Hypnum cupressiforme (in both vertical ranges) and Isothecium alopecuroides (in vertical range 0–60 cm) are indifferent in this respect. Our findings showed clearly that phorophyte preference and vertical position on the trunk are not independent variables. We investigated the independence of the frequencies and abundances of bryophyte species in the two vertical ranges separately on the two tree species. The species form four different groups: (1) species with greater abundance in the higher vertical range; (2) species with smaller abundance in the higher vertical range; (3) indifferent species with nearly uniform abundances in both vertical ranges; (4) species present only in one vertical range. It was found that even a single bryophyte species can show different vertical preferences depending on tree species. No significant relationships were found between the abundances of bryophytes and distance from the stream, slope of the bark surface and trunk circumference. Species-specific traits appear to explain individual phorophyte preferences; however, in several cases chemically mediated effects of the bark appeared to be important.     

Genetic variation and population structure in the endangered greater horseshoe bat Rhinolophus ferrumequinum

Following a dramatic decline last century, the British population of the endangered greater horseshoe bat Rhinolophus ferrumequinum is highly fragmented. To examine the consequences of fragmentation and limited dispersal on patterns of genetic structure and variation, we used microsatellite markers to screen bats from around 50% of the known maternity colonies in Britain, and two areas from continental Europe. Analyses revealed that Welsh and English colonies were genetically isolated. This, and lower variability in Britain than north France, may result from either genetic drift, or the species' colonization history. Gene flow among most neighbouring colonies was not generally restricted, with one exception. These findings have important implications for the ongoing conservation management of this species.     

Invasion Gateways and Corridors in the Carpathian Basin: Biological Invasions in Hungary

Biological invasions in Hungary are causing severe problems as a result of recent introductions and rapid land use changes. Poorly managed agricultural and rural, disturbed areas, and aquatic ecosystems are the most prone to plant invasions. Dry grasslands and semi-natural forests are less prone to invasions. A few plant species have led to human health (allergenic) problems. Some insect species have caused economic problems to crop production. A number of monitoring networks and control measures are in place for selected plants and insects. This revised version was published online in July 2006 with corrections to the Cover Date.     

Teleconnections and local weather orchestrate the reproduction of tit species in the Carpathian Basin

Variation in climatic conditions is an important driving force of ecological processes. Populations are under selection to respond to climatic changes with respect to phenology of the annual cycle (e.g. breeding, migration) and life‐history. As teleconnections can reflect climate on a global scale, the responses of terrestrial animals are often investigated in relation to the El Niño‐Southern Oscillation and North Atlantic Oscillation. However, investigation of other teleconnections and local climate is often neglected. In this study, we examined over a 33‐year period the relationships between four teleconnections (El Niño‐Southern Oscillation, North Atlantic Oscillation, Arctic Oscillation, East Atlantic Pattern), local weather parameters (temperature and precipitation) and reproduction in great tits Parus major and blue tits Cyanistes caeruleus in the Carpathian Basin, Hungary. Furthermore, we explored how annual variations in the timing of food availability were correlated with breeding performance. In both species, annual laying date was negatively associated with the Arctic Oscillation. The date of peak abundance of caterpillars was negatively associated with local temperatures in December–January, while laying date was negatively related to January–March temperature. We found that date of peak abundance of caterpillars and laying date of great tits advanced, while in blue tits clutch size decreased over the decades but laying date did not advance. The results suggest that weather conditions during the months that preceded the breeding season, as well as temporally more distant winter conditions, were connected to breeding date. Our results highlight that phenological synchronization to food availability was different between the two tit species, namely it was disrupted in blue tits only. Additionally, the results suggest that in order to find the climatic drivers of the phenological changes of organisms, we should analyze a broader range of global meteorological parameters.     

Mitochondrial sequence variation in ancient horses from the Carpathian Basin and possible modern relatives

Movements of human populations leave their traces in the genetic makeup of the areas affected; the same applies to the horses that move with their owners This study is concerned with the mitochondrial control region genotypes of 31 archaeological horse remains, excavated from pre-conquest Avar and post-conquest Hungarian burial sites in the Carpathian Basin dating from the sixth to the tenth century. To investigate relationships to other ancient and recent breeds, modern Hucul and Akhal Teke samples were also collected, and mtDNA control region (CR) sequences from 76 breeds representing 921 individual specimens were combined with our sequence data. Phylogenetic relationships among horse mtDNA CR haplotypes were estimated using both genetic distance and the non-dichotomous network method. Both methods indicated a separation between horses of the Avars and the Hungarians. Our results show that the ethnic changes induced by the Hungarian Conquest were accompanied by a corresponding change in the stables of the Carpathian Basin.     

Feeding ecology of migrating waders (Charadrii) at sodic-alkaline pans in the Carpathian Basin

Capsule Waders fed on the most abundant invertebrates: visual foragers preferred nektonic bugs, while tactile foragers had more diverse food preferences.     

Biogeography of the smooth snake (Coronella austriaca): origin and conservation of the northernmost population

Understanding historical range expansions and population demography can be crucial for the conservation and management of endangered species. In doing so, valuable information can be obtained regarding, for example, the identification of isolated populations, associations to particular habitats and distribution range shifts. As poikilotherms, snakes are vulnerable to environmental changes that can greatly shape their distribution ranges. Here we used mitochondrial data to elucidate the origin of the smooth snake population in Åland island, which is the northernmost location where the species is found. In Åland, we used mitochondrial and microsatellite data to fine‐map its spatial genetic structure, infer its demographic dynamics and determine its effective population size. We found three independent lineages, which expanded north from Iberian, the Balkans and Caucasus regions. The central lineage originating in the Balkans was the only one that reached Scandinavia. The Åland population belongs to this lineage and potentially colonized the island from the west via Sweden. This population appeared to be critically small and fragmented into two genetically isolated subpopulations. We discuss our results in light of previous findings regarding colonization routes in Europe and Scandinavia. Moreover, we discuss the origin and current genetic status of the Åland population relative to other co‐occurring snakes and suggest conservation measures based on our findings. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 426–435.     

Contrasting the ancestry patterns of three distinct population groups from the northernmost region of South America

Colombia, located in the north of the South American subcontinent is a country of great interest for population genetic studies given its high ethnic and cultural diversity represented by the admixed population, 102 indigenous peoples and African descent populations. In this study, an analysis of the genetic structure and ancestry was performed based on 46 ancestry informative INDEL markers (AIM-INDELs) and considering the genealogical and demographic variables of 451 unrelated individuals belonging to nine Native American, two African American, and four multiple ancestry populations. Measures of genetic diversity, ancestry components, and genetic substructure were analyzed to build a population model typical of the northernmost part of the South American continent. The model suggests three types of populations: Native American, African American, and multiple ancestry. The results support hypotheses posed by other authors about issues like the peopling of South America and the existence of two types of Native American ancestry. This last finding could be crucial for future research on the peopling of Colombia and South America in that a single origin of all indigenous communities should not be assumed. It then would be necessary to consider other events that could explain their genetic variability and complexity throughout the continent.     

Population structure of honey bees in the Carpathian Basin (Hungary) confirms introgression from surrounding subspecies

Carniolan honey bees (Apis mellifera carnica) are considered as an indigenous subspecies in Hungary adapted to most of the ecological and climatic conditions in this area. However, during the last decades Hungarian beekeepers have recognized morphological signs of the Italian honey bee (Apis mellifera ligustica). As the natural distribution of the honey bee subspecies can be affected by the importation of honey bee queens or by natural gene flow, we aimed at determining the genetic structure and characteristics of the local honey bee population using molecular markers. All together, 48 Hungarian and 84 foreign (Italian, Polish, Spanish, Liberian) pupae and/or workers were used for mitochondrial DNA analysis. Additionally, 53 sequences corresponding to 10 subspecies and the Buckfast hybrid were downloaded from GenBank. For the nuclear analysis, 236 Hungarian and 106 foreign honey bees were genotyped using nine microsatellites. Heterozygosity values, population‐specific alleles, FST values, principal coordinate analysis, assignment tests, structure analysis, and dendrograms were calculated. Haplotype and nucleotide diversity values showed moderate values. We found that one haplotype (H9) was dominant in Hungary. The presence of the black honey bee (Apis mellifera mellifera) was negligible, but a few individuals resembling other subspecies were identified. We proved that the Hungarian honey bee population is nearly homogeneous but also demonstrated introgression from the foreign subspecies. Both mitochondrial DNA and microsatellite analyses corroborated the observations of the beekeepers. Molecular analyses suggested that Carniolan honey bee in Hungary is slightly affected by Italian and black honey bee introgression. Genetic differences were detected between Polish and Hungarian Carniolan honey bee populations, suggesting the existence of at least two different gene pools within A. m. carnica.     

The northernmost sauropod record in the Northern Hemisphere

Isolated sauropod teeth from the Early Cretaceous Teete locality in Yakutia (Eastern Siberia, Russia) are the only evidence that sauropods lived in high latitudes (palaeolatitude estimate of N 62°) in the Northern Hemisphere. The spatulate broad tooth crowns of adult individuals lack marginal denticles while these are present in a juvenile tooth. The teeth have overlapping facets and likely belong to a basal macronarian. The juvenile tooth indicates that sauropods reproduced in high latitudes and possibly stayed there around the year. The Teete vertebrate assemblage comprises both endothermic, or presumably endothermic tetrapods (theropod dinosaurs, tritylodontids and mammals), and ectothermic tetrapods (salamanders, turtles, choristoderes and lizards), but no crocodyliforms. This suggests a temperate climate, with an annual mean temperature well above freezing level but below 14°C.     

A population on the edge: genetic diversity and population structure of the world's northernmost harbour seals (Phoca vitulina)

It is crucial to examine the genetic diversity and structure of small, isolated populations, especially those at the edge of their distribution range, because they are vulnerable to stochastic processes if genetic diversity is low and the isolation level is high, and because such populations provide insight into the consequences of population declines in a broader conservation context. The harbour seal (Phoca vitulina) population in Svalbard is the world's northernmost P. vitulina population. Nothing is known about the genetic diversity, distinctiveness, or origin of this small, marginalized mammalian population. Thus, the present study investigated its genetic status in the context of nearby P. vitulina in Iceland, south‐east Greenland, and northern Norway: this species is depleted/threatened in all of these regions. A total of 174 samples distributed between the four locations were analysed using 15 polymorphic microsatellites and variation in the displacement loop region (D‐loop). Each of the four locations was a genetically distinct population. The Svalbard population was highly genetically distinct, had reduced genetic diversity, received limited gene flow, had a rather low effective population size, and showed an indication of having experienced a bottleneck resulting from a recent population decline. The significant heterozygote excess observed in the Svalbard sample might be attributed to the low effective population size, which could initiate future population inbreeding effects. This phenomenon has not been reported earlier from other P. vitulina populations, but if the Svalbard population is experiencing inbreeding, this could reduce its resilience to climate change, disease outbreaks, or other perturbations. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 420–439.     

Integrating population genetics to define conservation units from the core to the edge of Rhinolophus ferrumequinum western range

The greater horseshoe bat (Rhinolophus ferrumequinum) is among the most widespread bat species in Europe but it has experienced severe declines, especially in Northern Europe. This species is listed Near Threatened in the European IUCN Red List of Threatened Animals, and it is considered to be highly sensitive to human activities and particularly to habitat fragmentation. Therefore, understanding the population boundaries and demographic history of populations of this species is of primary importance to assess relevant conservation strategies. In this study, we used 17 microsatellite markers to assess the genetic diversity, the genetic structure, and the demographic history of R. ferrumequinum colonies in the western part of its distribution. We identified one large population showing high levels of genetic diversity and large population size. Lower estimates were found in England and northern France. Analyses of clustering and isolation by distance suggested that the Channel and the Mediterranean seas could impede R. ferrumequinum gene flow. These results provide important information to improve the delineation of R. ferrumequinum management units. We suggest that a large management unit corresponding to the population ranging from Spanish Basque Country to northern France must be considered. Particular attention should be given to mating territories as they seem to play a key role in maintaining high levels of genetic mixing between colonies. Smaller management units corresponding to English and northern France colonies must also be implemented. These insular or peripheral colonies could be at higher risk of extinction in the near future.     

Discrimination of insect wingbeat-frequencies by the batRhinolophus ferrumequinum

Summary Five Greater Horseshoe bats,Rhinolophus ferrumequinum, were trained in a two-alternative forced-choice procedure to discriminate between artificial echoes of insects fluttering at different wingbeat rates. The stimuli were electronically produced phantom targets simulating fluttering insects with various wingbeat frequencies (Figs. 3, 4). Difference thresholds for wingbeat rates of 50 Hz and 100 Hz were determined. For an S+ of 50 Hz the difference threshold values lay between 2.8 and 4.6 Hz for individual bats; with an S+ of 100 Hz they increased to between 9.8 and 12.0 Hz (Figs. 5, 6, Table 1).Three bats, previously trained to discriminate between a S+ of 50 Hz and a S– with a lower wingbeat rate, were tested with higher frequency stimuli. When they had to decide between their old S+ of 50 Hz and either a 60 or 70 Hz echo two bats continued to select the 50 Hz stimulus while the third bat now preferred the faster fluttering insects (Table 2).During the discrimination task the echolocation behavior of the bats was monitored. When the phantom targets were presented all bats increased their duty-cycle of sound emission from about 40% to sometimes near 70%. They did so by either emitting longer echolocation calls or by increasing the sound repetition rate (Figs. 7, 8).The results show that Greater Horseshoe bats can determine the wingbeat rate of flying insects with an accuracy between 6 and 12%. Possible cues for flutter rate determination by cf-fm bats from natural and artificial insect echoes are discussed.Abbreviations DC duty-cycle - PD pulse duration - PI pulse interval - cf constantfrequency - fm frequency modulation     

马铁菊头蝠对同种咀嚼声的反应研究

咀嚼是哺乳动物食物吸收的重要组成部分,对动物的生存和繁殖极其重要。动物咀嚼时会发出低频低强度的咀嚼声,研究表明人类的咀嚼声可以增强自身或他人的食欲和愉悦度。蝙蝠作为哺乳动物中的第二大类群,其咀嚼声的特征和功能仍不清楚。本研究以吉林省集安市治安村的马铁菊头蝠Rhinolophus ferrumequinum为研究对象,通过回放蝙蝠咀嚼声和空白对照实验,同步录制蝙蝠的进食行为和回声定位声波,试图阐明蝙蝠咀嚼声对其进食行为和回声定位声波的影响。结果表明:进食次数在2种回放条件下的差异无统计学意义,但捕食尝试在2种回放条件下的差异有统计学意义。因此,咀嚼声能够显著地提高蝙蝠的进食欲望。此外,在蝙蝠咀嚼声的刺激下,其回声定位声波的频率增加,持续时间延长,脉冲速率减慢。这可能是因为蝙蝠在咀嚼声的影响下改变发声动机,从而改变了其回声定位声波的频谱时间结构。本研究第一次报道了蝙蝠咀嚼声对其进食行为的影响,为进一步阐明蝙蝠咀嚼声的功能和进化具有重要意义。     

Glacial-relict symptoms in the Western Carpathian flora

Glacial relicts have been regionally more common in glacial than in recent times. A rigorous assessment of which species are indeed glacial relicts is extremely difficult because direct evidence is untraceable or equivocal for many species. We aimed to identify species of the Western Carpathian flora (vascular plants, bryophytes and terrestrial lichens) that display apparent biogeographical and ecological symptoms, suggesting a wider regional or supra-regional distribution during glacial times, or at least before the middle-Holocene climate optimum. We worked with the premise that exemplary relict species should tolerate continental and/or arctic climates, should have large distribution ranges with disjunctions, being regionally rare and ecologically conservative nowadays, should be associated with habitats that occurred during glacial times (tundra, steppe, peatland, open coniferous forest) and should display a restriction of ecological niches in the study region. The assessed species were primarily those with boreo-continental or artcic-alpine distribution. We demonstrated a conspicuous gradient of glacial-relict symptoms, with Carex vaginata, Betula nana, Trichophorum pumilum, Nephroma arcticum, Saxifraga hirculus and Cladonia stellaris topping the ranking. Based on the arbitrary ranking, 289 taxa can be considered high-probability relicts. For only a minority of them, there are any phylogeographical and/or palaeoecological data available from the study area. Biogeographical and ecological symptoms of 144 taxa suggest that they retreated rapidly after the Last Glacial Maximum whereas other species probably retreated later. The first principal component of biogeographical symptoms sorted species from circumpolar arctic-alpine species of acidic peatlands and wet tundra to strongly continental species of steppe, steppe-tundra and mineral-rich fens. This differentiation may mirror the altitudinal zonation of glacial vegetation in the Western Carpathians.     

Recovery process of genetic diversity through seed and pollen immigration at the northernmost leading-edge population of Fagus crenata

The range expansion of a plant species begins with colonization of ecological empty patches from posterior source populations. This process involves stochastic loss of genetic diversity. However, the founder population could restore genetic diversity by gene flow from posterior populations via seeds and pollen and its recovery affects evolutionary potential for species expansion. To clarify the recovery process of genetic diversity during species range expansion, gene flow via seeds and pollen was investigated at the expansion front of Fagus crenata. Based on eight nuclear microsatellite genotypes of a total of 150 individuals and 225 seeds at the northernmost leading-edge population, genetic diversity, fine-scale spatial genetic structure (FSGS), and genetic differentiation from other five northern populations were investigated. Moreover, both seed and pollen immigration and their effects on genetic diversity at different successional stages were analyzed. The leading-edge population showed lower genetic diversity and substantial genetic differentiation, reflecting its strong genetic drift. Non-significant FSGS and a negative inbreeding coefficient for mature trees may indicate that the earliest generation consisted of founders from foreign seed sources. The significant proportion of seed and pollen immigration increased the number of different alleles for later successional stages. The effective number of pollen parents from foreign sources (20.8) was markedly higher than that from the local source (2.1). These results indicated that pollen immigration incorporated new and rare alleles and increased the genetic diversity of the population. However, the proportion of foreign gene flow decreased during succession, probably due to the increased reproductive success of local individuals as they reached maturity and grew in size.