Disjunct habitats as islands: genetic variability in the Caucasian rock lizard Lacerta portschinskii

Genetic diversity at 37 allozyme loci was surveyed in Lacerta portschinskii from contiguous populations and from a disjunct population. Indices of genetic diversity (heterozygosity, number of alleles per locus, and percentage of loci polymorphic) were greater in contiguous populations than in the smaller disjunct population. In this regard, disjunct populations appear to be similar to island populations. Indices of genetic diversity in Caucasian Lacerta are less than those reported from vagile lizard taxa and more similar to those of sit-and-wait predators. This revised version was published online in July 2006 with corrections to the Cover Date.     

Male mating success, reproductive success and multiple paternity in a natural population of sand lizards: behavioural and molecular genetics data

Sand lizard Lacerta agilis females characteristically mate with several males which, in staged mating experiments, results in multiple paternity of the offspring. In order to investigate multiple paternity in a natural population and interpret male reproductive behaviours in terms of sired young, we sampled the blood of females, potential fathers and hatchlings, and determined paternity using multilocus DNA fingerprinting as well as the variation at a single locus detected by the probe (TC) _n . The paternity analyses were preceded by a laboratory experiment in which we established that the parental alleles identified by the single locus probe were inherited in a Mendelian way. Our molecular data demonstrated that 12 out of 13 males (92%) that sired offspring were correctly identified from the 56 sexually mature males in the population. Also smaller males were accepted as sexual partners by the females, but sired fewer young in competition with larger males and were less able to maintain prolonged post-copulatory mate guarding. This may result in that some sexually successful males are only observed inside a female's home range, but never in pair-association with the female.     

Evolution in populations of Swedish sand lizards: genetic differentiation and loss of variability revealed by multilocus DNA fingerprinting

The Swedish sand lizard (Lacerta agilis) is a relict species from the period of warmth following the last glacial episode and has a fragmented distribution in central Sweden and a more continuous distribution in the southern part of the country. We used this model system of colonization–extinction for a study of genetic variability within and among Swedish populations from different parts of the distribution range using multilocus DNA fingerprinting. The results from the Swedish populations are then contrasted with those from a large Hungarian population in the centre of the species geographical distribution range, which is likely to closely resemble the ancestral founding population of Sweden. Swedish populations have a low level of genetic variability compared with the Hungarian reference population, which showed a genetic variability within the range described for outbred populations. Within the Swedish populations, the average bandsharing was 0.61, the mean heterozygosity 0.45 and the estimated number of alleles 2.7. The figures for the Hungarian population were a bandsharing of 0.19, a heterozygosity of 0.89 and an estimated number of alleles of 9.8. A population bottleneck, common to all Swedish sand lizards, is indicated by less than 20% of the alleles in the Hungarian population being retained in the Swedish populations, and higher bandsharing similarity between different Swedish populations (0.33) as opposed to the Hungarian population (0.19). The limited variability found in Swedish sand lizards is strongly subdivided between populations, with an average F_ST of 0.32, indicating a very limited gene flow between the isolated populations, as well as between populations in the region where the sand lizard has a more or less continuous distribution (F_ST = 0.41).     

A biogeographical analysis of the European Atlantic lowland heathlands

Questions: What is the climatic envelope of European Atlantic heathlands and the relationship between their floristic geographical variability and climatic parameters? Are the biogeographic patterns extracted from genuine heath plants comparable to those extracted from the accompanying flora? To what extent does the species composition extracted from phytosociological data support the current theory of refuge areas of heath vegetation in southern Atlantic Europe during the Pleistocene ice ages? Location: Atlantic Europe and NW Morocco. Methods: The geographical territory in which Atlantic heathlands occur was divided into 23 sectors following geographic and chorological criteria. A presence–absence table with 333 taxa was then constructed with the available phytosociological data. The taxa were classified into different groups according to their phytosociological affinity. Several types of numerical analysis were performed with this matrix and the climatic data obtained from meteorological sources. Results: Heathlands require a humid and oceanic climate and are limited by cold temperatures in the north and by summer droughts in the south. The highest floristic richness of this vegetation type is found in NW Iberia. Ordinations indicate a strong correlation between floristic composition of biogeographic sector and summer drought (Ios) and thermicity (It). Conclusions: The main climatic factors determining lowland heathland floristic distribution are thermicity and summer drought. The current optimal conditions for heath flora in NW Iberia suggest that there were southern refuges during the Pleistocene ice ages from which northward expansion has taken place.     

Vegetation and soil studies of the enclosed heathlands of the Lizard Peninsula,Cornwall

Summary The heathland vegetation of the Lizard Peninsula, Cornwall, which had been formerly enclosed for agricultural purposes and allowed to revert to heathland, was compared with unenclosed areas. The enclosed vegetation tended to be more complex and intermediate between two of the main heath types found on the Lizard, Short and Tall Heath (sensu Coombe & Frost 1956a). The concentrations of exchangeable calcium, sodium and magnesium in the soils of the enclosed heaths were also intermediate between those of the two unenclosed vegetation types, whilst exchangeable potassium and total phosphorus concentrations were higher, perhaps a relic of past management. The enclosed heaths are therefore distinctive entities in their own right, although they are related to the unenclosed vegetation types. The relevance to healthland conservation on the Lizard Peninsula is discussed.Species nomenclature follows Clapham, Tutin & Warburg (1962) for higher plants and Watson (1968) for bryophytes.We would like to thank Drs. D.E. Coombe and L.C. Frost for considerable assistance throughout this work. Professor P. Bannister and Mr. M.O. Hill assisted with numerical analysis, and Dr. D.F. Chamberlain confirmed the identification of bryophytes. Professor A.D. Bradshaw kindly allowed one of us (R.H.M.) the facilities of his department to complete this work. N.E.R.C. are thanked for financial support.     

Experiments with artificial nests on predation in reed habitats

We performed nest predation experiments with artificial nests in reedbeds investigating whether nest predation pressure is different at the water-reed edge and the grassland-reed edge compared with the reed interior. Furthermore, we tested the effects of vegetation structure (reed density, height and thickness) and the effect of other nest site characteristics (distance from edge, water depth) on the success of artificial nests. The experiments were completed 3 times during the breeding season in 2001 at Lake Neusiedl, Austria. Each artificial nest resembled Great Reed Warbler (Acrocephalus arundinaceus) nests and contained one plasticine and one Quail (Coturnix coturnix) egg and the predators were identified by marks left on the eggs. The potential predators were birds, probably the Marsh Harrier (Circus aeruginosus), gulls (Larus spp.) and reed warblers (Acrocephalus spp.). Nest survival data were analysed using the Mayfield method, and we performed a discriminant analysis for the data of vegetation and nest site characteristics. The nest predation was higher at the edges than in the reed interior, and was most pronounced in April, before the new reed sprouted. The reason for this finding was probably that after May the new reed contributed to greater concealment of the nests through the higher reed density and height.     

Phylogeography of western Palaearctic reptiles – Spatial and temporal speciation patterns

A phylogeographic analysis of eight species complexes of European reptiles was performed using different molecular methods. While mitochondrial genes (mainly cytochrome b sequences) enabled conclusions about phylogeography and differentiation, additional application of bisexually inherited markers provided information about speciation stages. As species with similar distribution patterns in southern and Central Europe were selected, matching phylogeographic patterns are useful for drawing general conclusions:

(1) The species complexes are in different stages of speciation. In some cases, cryptic species were detected.

(2) Highest genetic diversity occurs in southern Europe, the Near East and the Caucasus, regions corresponding with glacial refuges in the Iberian, Apennine and Balkan Peninsulas as well as in Turkey and the Caucasus. Often, several microrefugia must have existed in close neighbourhood. Additional microrefugia were located in southern France and in the Carpathian Basin.

(3) North Africa and the Middle East did not serve as glacial refuges for Central or northern European lineages and are typically inhabited by independent clades.

(4) Evidence for multiple range retractions and expansions, which were postulated for the times of Pleistocene climatic oscillations, could be found in the Balkans, but in Central Europe their traces have been wiped out by the last glacial. Only the Holocene invasion has left imprints in the genomes from this area.

(5) Central and northern Europe were recolonized from Balkan and Pontic refugia in the Holocene.

(6) Groups from the Iberian and Apennine Peninsulas rarely conquered other regions. This limitation can be attributed to the barrier function of the Pyrenees and the Alps.

Palaeoecological investigations towards the reconstruction of the history of forest clearances and coastal heathlands in south-western Norway

Palynological data collected over a period of 60 years have been compiled and re-interpreted in order to reveal the patterns of deforestation and health establishment in the south-western Norwegian coastal heathland. This heathland area has been divided into four sub-regions based on topography, bedrock and drift cover. The palynological investigations are from sites with pollen source areas of different sizes. The palynological signals are interpreted in terms of models that suggest an abrupt, gradual or stepwise deforestation which can be explained by terms of different pollen source areas. The deforestation seems to have been metachronous, leading to a regional mosaic pattern of different vegetation types. The deforestation process spanned more than 3600 calendar years (4000-400 B.C.), with three pronounced clearance periods at 4000-3600 B.C. (Mesolithic/Early Neolithic transition), 2500-2200 B.C. (Middle Neolithic II/Early Late Neolithic transition), and 1900-1400 B.C. (Late Neolithic to Bronze Age period II). The expansion of heathland has also been metachronous and took place over a period of ca. 4000 years between 4000-200 B.C., but was mainly completed by the end of the Bronze Age. Regional differences in the chronology of deforestation and heathland establishment are discussed. Deforestation with subsequent heathland expansion can best be explained in terms of the interaction between land-use history, topography and edaphic conditions under climatic conditions that favoured heathland development.     

Development of the plerocercoid I of Ophiotaenia europaea in reptiles

The existence of a two-host life-cycle in ophiotaeniid proteocephalideans was tested experimentally using Ophiotaenia europaea as a model. Three species of reptiles, Natrix natrix, Natrix tessellata and Lacerta viridis, were fed with experimentally infected copepods containing a large number of infective plerocercoids I. A few plerocercoids, most of which were dead, corresponding morphologically to the plerocercoid II developmental stage of O. europaea, were found encysted in the intestinal wall of N. natrix (8 days p.i.), N. tessellata (5 and 150 days p.i.) and L. viridis (40 days p.i.), while no plerocercoids or adult worms were recovered from their intestines. The results indicate that the infective plerocercoid I of O. europaea cannot undergo further development when ingested directly by the final host (a reptile), and that environmental temperature stimuli cannot initiate a reverse plerocercoid migration to the gut followed by strobilization.     

Short-term study of effects of fertilisation and cutting treatments on the vegetation dynamics of mountain heathlands in Spain

The influence of management and nutrient availability on the vegetation dynamics of heathlands characterised by Calluna vulgaris and Erica tetralix were studied in three mountain sites in Northern Spain. A total of 90 plots (1 m² each) received different combinations of cutting and twice the estimated background atmospheric deposition of nitrogen (56 kg ha⁻¹ yr⁻¹). One of the two dominant ericaceous species was selectively cut by hand at ground level and their regeneration compared in the presence or absence of the other. The results after 2 years showed significant effects of the fertiliser on the vegetation cover, mainly by favouring perennial herbaceous species. There were less noteworthy effects on the number of flowers and on the annual growth of the ericaceous species. It is concluded that, in the short term, increased nutrients alone, at twice the estimated current atmospheric deposition for the area, will not alter significantly the composition of the mountain heathlands. However, once the stands reach the mature phase, the capacity of the community to regenerate after a severe disturbance diminishes. A drastic impact, such as cutting may not result in re-growth of the same shrub species but in replacement by herbaceous species, which will also benefit from the increased nutrients.     

Vegetation study as a generator for population biological and physiological research on salt marshes

The paper deals with some views on the phytocoenose in relation to the functioning of vegetation and its plant-species populations in space and time. From these viewpoints the study of vegetation is seen as a field of tension between the organismic and reductionistic approaches. Both have their value, provided any dogmatism is avoided and either can be applied to the other.In the field of vegetation structure characteristic features of life-form spectra and species distribution, inversion phenomena in zonation, and community architecture in relation to production and decomposition are discussed. In this connection some remarks are made on habitat and niche differentiation with respect to the phytocoenose concept.Vegetation dynamics are discussed in relation to the introduction of Spartina anglica, the frequency of flooding by the tides, different environmental disturbances caused by heavy winter frost, rainfall and hot and dry periods, as well as to human interferences for agricultural and civil-technical purposes.It is suggested that salt-marsh plants may have found refuge areas in inland habitats as well as on more southerly coastal sites during glaciations.Nomenclature follows Tutin et al. (1964–1980).The author wishes to thank his collaborators Messrs B. P. Koutstaal and W. de Munck for much fieldwork, and Mrs M. J. van Leerdam-de Dreu and Messrs A. A. Bolsius and J. A. van den Ende for the preparation of text and figures.Communication No. 302 of the Delta Institute for Hydrobiological Research.     

Lack of congruence between morphometric evolution and genetic differentiation suggests a recent dispersal and local habitat adaptation of the Madeiran lizard Lacerta dugesii

Genetic differentiation among nine populations of the endemic lizard Lacerta dugesii Milne-Edwards 1829 (Lacertidae) from four groups of islands constituting the Archipelago of Madeira, was investigated by protein electrophoresis at 23 enzyme loci. Among twenty polymorphic loci, the total genetic diversity was due primarily to intra-population variation. The allele and genotypic frequencies among populations showed some heterogeneity, allowing the species to present a structuring pattern compatible with their geographical clustering. Some evidence suggests that selection acting on some loci in different ecological conditions may be responsible for the clustering of the populations studied. There was no apparent isolation effect expected under an "island" model of population divergence, and no correlation was found between genetic and geographic distances among populations. Morphological variation of the proposed three L. dugesii subspecies is not congruent with the allozyme analysis. This most probably suggests a rapid colonization of the islands followed by a strong effect of selection operating over the morphological characters used to define the subspecies.     

An ecological quality status assessment procedure for soft-sediment benthic habitats: Weighing alternative approaches

An assessment procedure for determining the ecological quality status of soft-sediment benthic habitats requires the following aspects: (1) habitat assignation of the samples (habitat approach), (2) reference or target conditions for the benthic parameters (reference approach), and (3) the selection of indicator tools to assess the relative quality status (indicator approach). For all these aspects, different approaches exist, and the indicator selection and reference approaches are largely documented. The aspect of the habitat approach is sometimes neglected, but is essential in determining the reference conditions per habitat type. Benthic habitats differ in structure and function, and as such will show wide variations in statistics or measures between habitats. A major problem, mainly in coastal soft-bottom systems, is to track the deviation lines within data of the different benthic habitat types. This study shows that both a classical community analysis and the use of habitat suitability maps seem to be appropriate tools, but further fine-tuning is necessary. For the second assessment step, an objective assessment of reference conditions is a challenge in areas lacking pristine or minimally disturbed sites, and areas of which historic data are not available. This can be remedied by using a dataset of the area with a good spatial and temporal coverage of the benthic data, thereby avoiding data originating from highly impacted areas. For the third aspect in the assessment procedure, we recommend the use of different indicators with different properties (parameters, algorithms) since it indicates their weaknesses and strengths in the local region.In general, this study showed us that it is valuable to test different existing approaches for EcoQ assessment in a local area, revealing strengths, weaknesses and shortcomings within the assessment regarding data, habitat identification, monitoring strategy, reference settings and indicator use. All these aspects need to be taken into account within an ecological quality status assessment of an area in order to improve its confidence.     

Power analysis to determine sample size for monitoring vegetation change in salt marsh habitats

Numerous initiatives are underway throughout New England and elsewhere to quantify salt marsh vegetation change, mostly in response to habitat restoration, sea level rise, and nutrient enrichment. To detect temporal changes in vegetation at a marsh or to compare vegetation among different marshes with a degree of statistical certainty an adequate sample size is required. Based on sampling 1 m² vegetation plots from 11 New England salt marsh data sets, we conducted a power analysis to determine the minimum number of samples that were necessary to detect change between vegetation communities. Statistical power was determined for sample sizes of 5, 10, 15, and 20 vegetation plots at an alpha level of 0.05. Detection of subtle differences between vegetation data sets (e.g., comparing vegetation in the same marsh over two consecutive years) can be accomplished using a sample size of 20 plots with a reasonable probability of detecting a difference when one truly exists. With a lower sample size, and thus lower power, there is an increased probability of not detecting a difference when one exists (e.g., Type II error). However, if investigators expect to detect major changes in vegetation (e.g., such as those between an un-impacted and a highly impacted marsh) then a sample size of 5, 10, or 15 plots may be appropriate while still maintaining adequate power. Due to the relative ease of collecting vegetation data, we suggest a minimum sample size of 20 randomly located 1 m² plots when developing monitoring designs to detect vegetation community change of salt marshes. The sample size of 20 plots per New England salt marsh is appropriate regardless of marsh size or permanency (permanent or non-permanent) of the plots.     

An Experimental Investigation of Bioconvection in Three Species of Microorganisms*

SYNOPSIS. A series of experiments was carried out on bioconvection using cultures of Polytomella agilis, Tetrahymena pyriformis and Chlamydomonas moewusii. During convection these microorganisms disperse into more and less dense regions causing visible patterns. These patterns, called the cell plan forms, were found to change with increasing Rayleigh numbers above Rc, the lowest concentration gradient at which bioconvection occurs. Hexagonal convection cells were observed near the critical Rayleigh number, Rc. As the Rayleigh number was increased the cell plan form changed from hexagons to 2-dimensional rolls and then to square convection cells. The size of the observed convection cells was less than expected for such a physical phenomenon. This appears to result from an increasing tendency of the microorganisms to circulate near the upper surface at higher Rayleigh numbers. The square convection cells appear to be unique to bioconvection. The inhibition of bioconvection is directly linked to an increased mortality rate. Observations on the nature of negative geotaxis were made which tend to support the statocyst hypothesis.     

Interspecific aggregation of ectoparasites on bats: importance of hosts as habitats supersedes interspecific interactions

Patterns of aggregation of species or individuals may result from combinations of interspecific interactions such as competition, facilitation, or apparent facilitation, as well as from equivalent responses to environmental factors. Host–parasite systems are ideal for the investigation of mechanisms that structure assemblages. Interspecific aggregation is documented for multiple groups that are ectoparasitic on mammals and host‐mediated apparent facilitation has been suggested to explain these aggregation patterns. To investigate the generality of this pattern and to determine likely structuring mechanisms, I analyzed species co‐occurrence, correlations of abundances, and nestedness for ectoparasite assemblages from each of 11 species of Neotropical bat. Ectoparasite assemblages on four of 11 host species exhibited significant positive co‐occurrence for the entire assemblage or for at least one pair of species in the assemblage; ectoparasites on two host species exhibited positive co‐occurrence that approached significance. There was no evidence of negative co‐occurrence. Nine species‐pairs exhibited positive abundance correlations, including seven of the eight species‐pairs that exhibited positive co‐occurrence. No species‐pair exhibited a negative correlation of abundances (i.e. density compensation). Ectoparasite assemblages from five of 11 host species exhibited nestedness, including all three assemblages that exhibited assemblage‐wide positive co‐occurrence. Multiple mechanisms associated with host characteristics may contribute to host aggregation in ectoparasite assemblages, including host body size, vagility, home range size, burrow or roost size and complexity, immunocompetence and social structure. In general, data in this study and elsewhere are not consistent with interspecific interactions among ectoparasites, including apparent facilitation, being primary structuring mechanisms of ectoparasite assemblages on mammalian hosts. Rather, host behavior and ecology are likely to affect the frequency of host–ectoparasite encounters and of conspecific host interactions that facilitate transfer of ectoparasites, thereby, molding patterns of ectoparasite co‐occurrence, abundance and species composition on mammalian hosts. Combinations of characteristics that are primarily responsible for molding ectoparasite assemblage composition likely are host‐taxon specific.     

The importance of novel and agricultural habitats for the avifauna of an oceanic island

Conservation management can no longer rely on protecting pristine habitats, but must consider the wider landscape. This is especially true on oceanic islands where endemic species are believed to be particularly susceptible to the extinction risks that accompany land conversion. Despite this, there is a paucity of studies examining how endemic communities on oceanic islands may be distributed across such human-modified habitats. Taking Príncipe Island in West Africa as a case study, we investigate how avian communities vary across the habitats (primary forest, secondary forest, agricultural areas) of this globally important centre of endemism. Here, recent policy reforms aimed at poverty alleviation and increased food production are rapidly altering the current land-use mosaic. Across all habitats, 27 bird species were encountered. Survey points in secondary forest and agricultural areas were, on average, more diverse and held higher overall abundances of birds than those within primary forest. This was true for both the entire avian assemblage and the endemic species alone. Nevertheless, two IUCN-listed species were restricted to primary forest, and many other endemics occurred at higher densities within this habitat. We demonstrate that agricultural areas and novel habitats, such as secondary forest, can hold high abundances of endemic species and thus have the potential to act as a resource for biodiversity conservation. A double-stranded approach to conservation is therefore required that both protects the integrity of the primary forest and controls the rapid changes in agricultural land-use to ensure that it continues to support a large component of the endemic avifauna.