On the significance of dispersal power for populations of carabid-beetles (Coleoptera,Carabidae)

Summary By various observations on carabid populations the author attempts to give an impression of the quantitative occurrence of dispersal and of the relation between dispersal and the chance of founding populations (dispersal power). Pitfall-catches in the recently reclamed Zuiderzee-polder E-Flevoland demonstrate that within seven years individuals of a number of monomorphic macropterous and dimorphic species had founded populations there. From the very high frequency of full-winged individuals within the latter populations it follows that full-winged carabid individuals generally must have a much greater power of dispersal than flightless ones. Therefore, winged individuals of dimorphic species were about equally able to reach E-Flevoland as were those of monomorphic macropterous ones, whereas individuals of monomorphic brachypterous species obviously are seriously hampered. The early appearance of individuals of riparian species on the shores of an artificial lake in the dune area Meijendel suggests that particularly populations living in unstable environments extensively invest in dispersal. It appears, however, that an important investment in dispersal apparently is not restricted to species from unstable environments; at least some sparse populations living in more stable environments also sacrifice relatively great numbers of individuals for dispersal (Pterostichus strenuus). The hypothesis is proposed, that populations facing a high risk of extinction generally will have a sufficient chance of founding populations (high turnover) when investing extensively in dispersal. Not only macropterous but — at least in some populations — also brachypterous individuals participate in migration, although in the populations studied the dispersal power of flightless individuals is found to be very small (Carabus problematicus). Under certain conditions the dispersal of full-winged individuals from wing-dimorphic populations may ultimately lead to a decrease or even a loss of dispersal power by a decrease of the frequency of macropterous individuals. It is assumed, however, that under certain natural conditions also brachypterous individuals may contribute to the spreading of risk within and between populations. The dispersal power of monomorphic macropterous, dimorphic and monomorphic brachypterous populations in a cultivated countryside like Drenthe is discussed. The connection between the dispersal power of different kinds of carabid populations and the resulting chance of survival under different conditions is discussed. Some suggestions for nature preservation management are given.Communication No. 144     

Conservation value of roadside verges for stenotopic heathland Carabidae: corridors or refugia?

It has been shown that, in The Netherlands, sandy roadside verges may aid the survival of stenotopic Carabidae associated with open habitats such as heathland and sandy grassland. It is suggested that roadsides serve as corridors linking heathland fragments, thus maintaining a metapopulation structure, which helps avoid regional extinction. However, a counter-argument is that verges encourage dispersal into unsuitable habitats, and function as sinks in which individuals die without reproducing. The stenotopic species recorded from roadside verges but not in adjacent open habitats all had high powers of dispersal. The least successful dispersers were found only in open habitats and in verges closely connected to them. From this, the conclusion that verges function as corridors can be drawn. We reinterpret these data to suggest that stenotopic species were breeding on the roadside verges, and that they acted as refugia rather than corridors. We also present evidence from a roadside verge in the Breckland region of East Anglia, England. We show that many stenotopic species breed on the verge, which thus functions as a refugium. For Carabidae dependent on early successional habitats, survival may be more likely on disturbed roadside verges than on nature reserves in the absence of regular management disturbance.     

The relationships between the structure of paddy levees and the plant species diversity in cultural landscapes on the west side of Lake Biwa,Shiga, Japan

Paddy levees form networks of narrow linear habitats and play various roles in cultural landscapes. Traditional landscapes on the west side of Lake Biwa consist of paddy field terraces and both stone and soil levees that have been maintained by paddy field management using local resources. Paddy levees in this study site are principally classified into five different types. Our study points out how differences in paddy levee structure as well as in management practices influence the plant species. Seventeen paddy levee transects were split into four habitat types based on their species components by TWINSPAN. Spatial characteristics and physical structures of paddy levees depended on natural conditions and human activities. The species–area curves of each levee type showed a clear distinction: the soil, stone and abandoned curves were steep, while the concrete and consolidated ones were gentle. The vegetation on consolidated levees was utterly different from the vegetation on traditional levee types from the aspect of species richness and species components. Soil type levees contained various woody plant species and included more diverse and indigenous plant species than abandoned type levees.     

Beetles (Coleoptera) on brownfield sites in England: An important conservation resource?

A total of 78 brownfield (post-industrial and urban) sites were surveyed for beetles between 1991 and 2001 throughout England using pitfall traps. The distribution of ground, rove and phytophagous beetle assemblages was investigated using ordination and classification analyses. Site drainage and vegetation cover had a profound effect on the distribution of ground and rove beetle assemblages but site location was also important for phytophagous beetle assemblages. A total of 182 records of 46 nationally rare and scarce species (16 ground, 10 rove and 20 phytophagous species) were generated. A number of these species are more usually associated with other, more natural habitats such as riverine sediments, sandy heaths and chalk grassland. Brownfield sites provide habitat conditions similar to more natural habitats and they may help maintain populations of some rare and scarce species. The results indicate that brownfield sites are important habitats for beetles and there is evidence that the situation is similar for other invertebrate groups. There should be no further assumptions that post-industrial and urban sites have no conservation interest.     

Density compensation in island avifaunas

Summary This paper analyzes factors which determine the extent of density compensation on islands; i.e., is the summed population density of all species on an island equal to the summed mainland density? A graphical analysis allows quantitative comparisons of density compensation studies. Two hypotheses which are generally applicable predict the extent of density compensation on islands: (1) Niche theory predicts that summed population densities should be low if island species number is low. (2) The habitat appropriateness hypothesis predicts that summed population densities should be low if island populations occupy unfamiliar habitat. Both hypotheses successfully explain variability in the extent of density compensation on islands. Relative to the mainland, summed population densities on islands are high when islands support a large number of species and those species occupy familiar habitats. Summed population densities on islands are low when islands support few species and those species occupy novel habitats.     

Reproductive patterns in annual legume species on an aridity gradient

Reproductive patterns are analysed in annual legumes of west Asia, and their relationships to increasing aridity determined by multivariate analysis. Dormancy, seed size, dispersal and fecundity are shown to be partially substitutable in terms of their effect on survival and population growth.The range of patterns show greatest diversity under mesic conditions in coastal, mediterranean areas with high winter rainfall, low incidence of frost and long growing season. Increasing aridity leads to increasing reproductive homogeneity, in which a subset of patterns-those composed of high levels of seed dormancy, high seed to pod ratios, restricted dispersal capabilities and early flowering-become predominant. These findings corroborate earlier theoretical and empirical evidence concerning desert annuals.The majority of widespread species are shown to possess arid-type patterns. This exerts considerable influence on both the type and degree of ecotypic differentiation within species. The only feature that responds consistently to climatic change is flowering time, particularly among species characterized by high seed dormancy. In the few widespread species that do not display arid-type characteristics, ecotypic differentiation appears more frequently in a larger suite of traits.The unequal environmental demands made on species showing different levels of dormancy (e.g. variation in length of growing season from year to year and place to place) leads to strong asymmetries governing the relationships between reproduction and ecological amplitude. In this respect arid-type strategists with high seed dormancy appear to have greater chances of expansion than others. This has implications when choosing pasture legumes to improve mediterranean grasslands.     

Can the acoustic adaptation hypothesis predict the structure of Australian birdsong?

The acoustic adaptation hypothesis (AAH) predicts that long-distance signals will be structured so as to maximize their transmission fidelity. Previous studies testing the hypothesis on birdsong have provided equivocal support. The best support comes from large-scale comparative studies and those studies where habitat is characterized as open versus densely vegetated. In the first case, sufficient statistical power is present to detect even small effects on song structure, whereas in the later case the effect size of the habitat may be sufficiently large. Most studies have focused on Holarctic or Neotropical species, which may ultimately share a common evolutionary history. In this study, Australian birds were chosen for a phylogenetically independent test of key predictions of the AAH. Specifically, birds in open habitats were predicted to sing songs with higher frequencies, greater bandwidth, have a greater probability of having overtones, and be emitted at a quicker rate than birds inhabiting densely forested habitats. Acoustic measurements were made on commercially available recordings of 121 species of Australian birds from 41 different families. Analyses controlled for variation explained by body mass (using ANCOVA), and phylogeny (using genus pairs analyses). We found only modest support for the AAH. Our finding that birds in open habitats produced higher frequency vocalizations and greater bandwidth vocalizations is also consistent with hypotheses about signal structure facilitating auditory distance assessment. Forest birds may therefore rely on cues other than frequency-dependent attenuation for ranging.     

Day-night differences in the number of pineal “synaptic” ribbons in two diurnal rodents,the chipmunk (Tamias striatus) and the ground squirrel (Spermophilus richardsonii)

Summary Daytime numbers of pineal synaptic ribbons higher than reported in the pineal gland of any other mammalian species were observed in two diurnal rodents, the eastern chipmunk and Richardson's ground squirrel. The number of synaptic ribbons was lower during the daytime and higher at night in both of these species.     

Structure,composition and species diversity in an altitude-substrate matrix of rain forest tree communities on Mount Kinabalu,Borneo

We studied forest structure, composition and tree species diversity of eight plots in an environmental matrix of four altitudes (700, 1700, 2700 and 3100 m) and two types of geological substrates (ultrabasic and non-ultrabasic rocks) on Mount Kinabalu, Borneo. On both substrate series, forest stature, mean leaf area and tree species diversity (both 4.8 cm and 10 cm diameter at breast height [dbh]) decreased with altitude. The two forests on the different substrate series were similar at 700 m in structure, generic and familial composition and tree species diversity, but became dissimilar with increasing altitude. The decline in stature with altitude was steeper on the ultrabasic substrates than on the non-ultrabasic substrates, and tree species diversity was generally lower on ultrabasic substrates than on non-ultrabasic substrates at 1700 m. The forests on non-ultrabasic substrates at higher altitudes and those on ultrabasic substrates at the lower altitudes were similar in dbh versus tree height allometry, mean leaf area, and generic and familial composition at 1700 m. These contrasting patterns in forest structure and composition between the two substrate series suggested that altitudinal change was compressed on the ultrabasic substrates compared to the non-ultrabasic substrates. Tree species diversity was correlated with maximum tree height and estimated aboveground biomass, but was not with basal area, among the eight study sites. We suggest that forests with higher tree species diversity are characterized by greater biomass allocation to height growth relative to trunk diameter growth under more productive environment than forests with lower tree species diversity.     

Local distribution of the lycaenid butterfly,Jalmenus evagoras,in response to host ants and plants

Summary The caterpillars of Jalmenus evagoras are tended by ants as they feed upon Acacia trees. In the area of Brisbane, Australia, J. evagoras require ants of the Iridomyrmex anceps species group; predation and parasitism are so intense that larvae and pupae deprived of attendant ants cannot survive (Pierce 1983). We investigated the efficiency with which J. evagoras locate and exploit the host ant resource by sampling 737 quadrats in 30 sampling grids and six study sites containing appropriate host plants; ants were collected at baits located in the center of each quadrat. J. evagoras was found in all habitats where I. anceps cooccurred with host Acacia. Nine of the ten sampling grids which had three or more I. anceps/Acacia host quadrats also had colonies of J. evagoras present (or immediately adjacent), including sites as far as 35 km apart. Of 19 sampling grids on which host quadrats were rare (i.e., less than three quadrats), none had J. evagoras (P<0.001). Within sample grids, I. anceps was distributed indepedently from Acacia trees, suggesting that they are not dependent for their survival on either Acacia or on J. evagoras. Within montane pasture habitats, I. anceps and at least one other ground-dwelling Iridomyrmex species were distributed in mutually exclusive ant mosaic territories which were stable during a one month period. I. anceps did not colonize or tend pupae of J. evagoras experimentally placed in adjacent territories of a different, nontending species of Iridomyrmex, demonstrating the integrity of territory boundaries. Sampling of ants in Acacia trees revealed that, in the absence of J. evagoras, Iridomyrmex workers are not common above ground level, and that their numbers decline in larger trees (P=0.02). In I. anceps territories, eight of nine J. evagoras pupae placed in trees over 3.0 m tall were not found after 24 h whereas all ten controls placed in low trees were found and tended (P=0.00012). This may explain why J. evagoras tends to oviposit in trees less than 2.0 m tall. An alternative hypothesis, that smaller trees have higher content of total nitrogen, and are threfore more nutritious, was not supported. We conclude that the local distribution and host tree selection by J. evagoras is dependent upon the distribution, patchiness, and foraging behavior of the host ant, I. anceps, and its spatial overlap with a number of species of host Acacia.     

Segregation analysis in hereditary retinoblastoma

Summary Segregation analysis was performed on 211 nuclear families belonging to 166 pedigrees of hereditary retinoblastoma found in a number of series which have been gathered from the literature. Bilaterally affected carriers appear homogeneous. The segregation ratio in their offspring is 0.49, and the proportion of bilateral cases among affected offspring is 0.87. Both unilaterally affected and unaffected carriers appear heterogeneous. The very low segregation ratio (0.08) in the offspring of unilateral carriers who are not detected through an affected child, suggests the possiblity of two types of carriers, high and low transmitters. The proportions of low transmitters was estimated as 0.14 among all familial unilateral carriers and as 0.45 among all detected unaffected carriers. Unilateral and unaffected high transmitters give a significantly lower segregation ratio than bilaterally affected carriers.On the one hand, the existence of these two different types of carriers provides arguments in support of the hypothesis of delayed mutation. On the other hand, the differences in penetrance among high transmitters, according to their phenotype, supports the hypothesis of host resistance. Under the two-mutation hypothesis, the possibility that the mutation rate is variable among individuals and partly genetically determined, is suggested.     

Contribution of tree species to the biodiversity of a 1ha Old World rainforest in Brunei, Borneo

Tree biodiversity data is presented for a 1-ha permanent study plot on a low slope position of the Kuala Belalong valley in Temburong District, Brunei, Borneo. This is the first study in the region to document the tree flora of a low slope. All trees 5cm at 1.3m were mapped, measurements of dbh and height taken, and identified to species and morpho-species level. The total and relative values of each family for density, diversity, dominance and Importance Value Indices were calculated. A total of 1062 stems were recorded on the plot (1019 live, 43 dead) with a total of 278 species representing 110 genera in 49 families. The most ecologically significant family as determined by the Family Importance Value (FIV) was Euphorbiaceae with a value of 40 and'the'most significant species, using the Importance Value (IV), was morpho-speciesFordia splendissima with an IV of 29. The total basal area of all living trees was 31.43m². Forest structure was marked by a few tall (>45m) emergents, for example,Koompassia excelsa, Crypteronia griffithii and two species of the Dipterocarpaceae family, however, lower understorey and sub-canopy species dominated the plot with the average height of all individuals over the hectare being 11m.     

Adaptive energy metabolism in four chromosomal species of subterranean mole rats

Summary Gross energy intake and apparent dry matter digestibility of animals fed carrots ad lib in the laboratory, were measured in the four chromosomal species of the Spalax ehrenbergi superspecies in Israel. Gross energy intake of 132.8 and 155.9 kJ/day was measured for the 2n=52 and 2n=58 chromosomal mesic species, while in the 2n=54 and 2n=60 chromosomal xeric species it measured only 80.3 and 75.0 kJ/day, respectively. Dry matter digestibility ranged between 92.3 and 95.6% in thefour chromosomal species. The differences in gross energy intake between the mesic and xeric species, appeared to reflect adaptive energy metabolism variation associated with geographic variation in climate, habitat productivity, and food availability.     

Cache site selection by chipmunks (Tamias spp.) and its influence on the effectiveness of seed dispersal in Jeffrey pine (Pinus jeffreyi)

The effectiveness of Jeffrey pine (Pinus jeffreyi) seed dispersal performed by seed-caching yellow pine chipmunks (Tamias amoenus) and lodgepole chipmunks (Tamias speciosus) was compared to that of wind dispersal in the Sierra Nevada of western Nevada. Wind-dispersed seeds typically fall under or near the parent tree. Chipmunks removed 90 and 97% of 1064 radioactive seeds from each of two simulated wind-dispersed seed shadows in less than 24 h. Wind-dispersed seeds were deployed within 12 m of the two source trees, but chipmunk caches were found from 2–69 m from the trees. Chipmunks carried nearly all seeds away from source trees, greatly reducing the density of seeds under and near source trees. Caches contained from 1–35 seeds and most were buried 7–21 mm deep. Chipmunks cached in open bitterbrush shrubland with mineral soils much more than expected and cached in closed-canopy Jeffrey pine and lodgepole pine forests with thick needle litter much less than expected. Many Jeffrey pine seedlings and saplings grow in the bitterbrush habitat and few grow in the pine forests. Ten and 20% of the original caches survived until April, the time of seed germination, at the two sites. The movement of wind-dispersed seeds is random relative to environmental variables important in seedling survival, and the wind in coniferous forests cannot quickly bury seeds. The quality of seed dispersal rendered by chipmunks was superior to that provided by the wind because the chipmunks quickly harvested seeds on the ground, moved them away from source trees, and buried them in the ground in habitats and microhabitats where they were more likely to establish new seedlings. The increased quality of seed dispersal provided by animals relative to the wind may help explain why over twenty species of pines have evolved seeds and cones that are adapted for dispersal by seed-caching animals.     

Dispersal patterns of endemic alpine butterflies with contrasting population structures:<Emphasis Type="Italic"> Erebia epiphron</Emphasis> and<Emphasis Type="Italic"> E. sudetica</Emphasis>

We studied population sizes and mobility of Erebia epiphron and Erebia sudetica, two high mountain butterflies forming endemic subspecies in the Hrubý Jeseník Mountains, Czech Republic. E. epiphron formed two continuous populations containing 100,000 and 4,500 individuals on alpine grasslands. The butterflies moved freely within their habitats, but movements between the two populations were highly unlikely. E. sudetica formed a system of colonies at timberline sites on valley headwalls and in forest clearings. Two such colonies studied in detail contained 4,500 and 450 adults and were interconnected by limited dispersal. The negative exponential function and the sigmoid function (this assumes flat decrease of movements over short distances) were superior to the inverse power function in fitting mobility data for both species. For E. sudetica, the functions describing movements within a habitat differed significantly from total movements, suggesting different behaviours of dispersing individuals. The habitats of E. epiphron are uniform and highly isolated, favouring free within-habitat mobility but prohibiting leaving their boundaries. The habitats of E. sudetica are diverse and disturbance-dependent; leaving such habitats is less risky, and a source-sink model may explain the persistence of the species in the mountains.     

A Cytogenetic Study of the Leaf Beetle Genus Cyrtonus (Coleoptera, Chrysomelidae)

The chromosomes of ten species of Cyrtonus and the genome sizes of six are surveyed. Among the total of 15 chromosomally studied species, 11 have 2n=28 chromosomes and a 13+Xy_p male meioformula, three have 2n=40 and 19+Xy_p and one 2n=46 and 22+Xy_p. All but one species with 28 chromosomes show only metacentric or submetacentric chromosomes, whereas the species with 40 and 46 chromosomes display some telocentrics or subtelocentrics, that are probably derived from the former by centric fissions. However, since the number of major chromosome arms is strikingly higher in these latter species (NF=70 and 78) than in the 28-chromosome species (mostly NF=56), other chromosomal rearrangements such as pericentric inversions or heterochromatin accretions could also be involved. The genome sizes display a narrow range, from 1C=0.6–1.22pg, and they are not significantly correlated with the chromosome numbers. Some possible factors implied in the rough chromosomal evolution of Cyrtonus are discussed in relation to a few other genera of the subfamily Chrysomelinae.     

Meiotic studies ofElymus nutans andE. jacquemontii (Poaceae,Triticeae) and their hybrids withPseudoroegneria spicata and seventeenElymus species

Hybridizations ofElymus nutans andE. jacquemontii were carried out with one species ofPseudoroegneria (S genome), and 20Elymus species, each containing either of the SH, SY, SYH, or SYW genomes. Chromosome configurations were analysed at metaphase I of the two target taxa and their interspecific hybrids. It is concluded that (i)E. nutans is an allohexaploid containing the SYH genomes, andE. jacquemontii is an allotetraploid having the SY genomes; (ii) the genomic affinity is associated with the geographic distance between the species studied; (iii) minor genomic structural rearrangements have occurred within the hexaploid taxon ofE. nutans.     

The occurrence of former heathland vegetation in the coastal areas of the south-east Baltic sea,in particular Lithuania: a review

Previously published pollen diagrams from the coastal area of the south-east Baltic Sea including parts of the Polish and Lithuanian peninsulas, Mierzeja Wilana (Frische Nehrung) and Kuri Nerija (Courland Spit) respectively, document the former presence of heathland vegetation and an associated burning practice. This type of human influence can be dated back to ca. 1200 B.P. at the latest, and was maintained until the 18th century. It is suggested that this former Calluna-dominated vegetation belonged to the known west to north-west European heathland vegetation. Several dwarf shrub taxa, such as Genista spp. and Erica tetralix, normally connected to heathland vegetation, may have had a former wider distribution.     

Estimation of the retention times and distances of seed dispersed by two monkey species, Alouatta seniculus and Lagothrix lagotricha, in a Colombian forest

Isolated-bout method to estimate the retention times and dispersal distances was applied to the seed dispersal by red howler monkeys (Alouatta seniculus) and Humboldts woolly monkeys (Lagothrix lagotricha) in a lowland tropical forest at La Macarena, on the border of the Macarena and Tinigua National Parks, the Department of Meta, Colombia. Continuous observations were made on the feeding and ranging behavior of well-habituated troops of howler monkeys and woolly monkeys as well as continuous collection of monkeys feces. We selected out the isolated-bout as a feeding bout on the specific species that was only once recorded within 48 h before the seeds of that species appeared in the feces of monkeys. In that case, the seeds were strongly suggested to come from that isolated bout. Then retention times, route seed dispersal distances and direct seed dispersal distances were estimated. Howler monkeys, which are regarded as generalist herbivores, showed longer retention times and dispersal distances along monkeys route than did woolly monkeys, which are specialist frugivores. However, the direct distances that seeds were carried from the mother tree were not significantly greater for howler monkeys than for woolly monkeys. This shows that both retention time and movement patterns by the monkeys, especially the total ranging area, influence the direct distance that seeds are carried from the mother tree.     

Nitrate,nitrate reduction and organic nitrogen in plants from different ecological and taxonomic groups of Central Europe

Summary 48 plant species of the families Asteraceae, Chenopodiaceae, Ericaceae, Fabaceae, Lamiaceae, Polygonaceae and Urticaceae were investigated in 14 natural habitats of Central Europe having different nitrate supplies, with respect to their nitrate content, nitrate reductase activity (NRA) and organic nitrogen content. Plants that were flowering were selected where possible for analysis. The plants were subdivided into flowers, laminae, petioles+shoot axes and below-ground organs. Each organ was analyzed separately. Differences among species were found for the three variables investigated. Apart from the Fabaceae, which had particularly high concentrations of organic N, these differences reflect mainly the ecological behaviour, i.e. high nitrate and organic N contents and NRA values per g dry weight were found in species on sites rich in nitrate, and vice versa. Nitrate content, NRA and organic N content were correlated with nitrogen figures of Central European vascular plants defined by Ellenberg (1979). By use of regression equations this correlation was tested with species from other systematic groups. Some species were attributed with calculated N figures for the first time.