Specializations and polyphagy of Plebejus argus (Lepidoptera: Lycaenidae) in North Wales

Abstract. 1. The polyphagous butterfly, Plebejus argus L., was found to have specialized requirements: eggs are laid along vegetation/bare ground margins, larvae specialize on tender meristematic and reproductive growth, larvae and pupae are tended by Lasius ants, and warm microclimates are favoured.
2. These requirements are met in early successional habitats within several bio topes. These habitats are locally distributed, as is P. argus.
3. Populations on limestone, heathland and mossland biotopes did not differ greatly in ecologies. Given a choice of foodplants in captivity, larvae from heathland and limestone did show slight differences in preference, selecting some of the hostplants they normally encounter in the wild. Regardless of biotope, all these larvae were plant taxa generalists (on three plant families), but specialists on plant parts.
4. The decline of P. argus in Britain is attributed to loss of biotopes and to declining traditional management of them. Fewer habitats are available, and the distances between them are greater than formerly. As some of these habitats are short lived, and P. argus rarely flies far, the butterfly's decline is a compound effect of reduced areas of habitat and inability to colonize them.     

Methods for the activation of the resting eggs of Daphnia

SUMMARY. 1. The conditions required to initiate development of resting eggs of thirty-six clones of Daphnia representing seven species were investigated.
2. The temperature of both dark incubation and subsequent light treatment are shown to affect hatch success. By varying these parameters the majority of resting eggs from each test clone were stimulated to develop. Arctic clones required a low hatching temperature (7°C), whereas clones from warmer climates hatched best at 14–21°C.
3. Variation in hatching cues existed between conspecific individuals from different collection sites. These differences suggest that research determining macro- and microgeographic patterns in hatching phenology would be fruitful.     

Three ways of assessing metapopulation structure in the butterfly Plebejus argus

1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus . First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area.
2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m.
3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale.
4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations.
5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur.
6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.     

Do leaf margins of the temperate forest flora of southern South America reflect a warmer past?

Aim   Our aims were: (1) to characterize the linear relationship between the proportion of woody dicotyledonous species with entire-margined leaves ( E ) and mean annual temperature (MAT) from a southern temperate flora that still harbours many lineages that originated under warmer climates; (2) to compare this relationship with those developed from floras of different regions of the world; and (3) to contrast temperature predictions based on leaf margins of the native southern flora versus the naturalized alien flora, mostly of boreal origin.
Location   The temperate forest of southern South America (TFSA).
Methods   At each 1° latitudinal band, we estimated E based on species latitudinal ranges and MAT from both an isotherm map and a global temperature grid. We also calculated E from five local floras located between 40 and 43° S, and from the naturalized alien flora of Nahuel Huapi National Park in southern Argentina.
Results   We found a close relationship between E and MAT for the TFSA. Equations developed from floras of the Northern Hemisphere overestimated extant temperatures of this biome by 6–10 °C at both geographical and local spatial scales. On the other hand, MAT predictions from leaf margins of the alien flora were similar to the actual MAT. A published regression between E and MAT from tropical South America was remarkably similar to the one we estimated from the TFSA. This tropical equation predicted accurately the temperatures observed for this temperate biome based on leaf margins of the native flora.
Main conclusions   Despite massive plant extinction due to environmental cooling and biogeographical isolation during the Tertiary, leaf-margin analysis reveals that the flora of the TFSA still reflects its original development under the warmer conditions of western Gondwana and its past connections with low-latitude forest floras of tropical South America.     

Elevational parallels of latitudinal variation in the proportion of lianas in woody floras

Aim  To determine if elevational variation in the proportion of lianas in woody floras parallels the variation observed on latitudinal gradients. This is to be expected if the poleward decrease in the importance of lianas is related to the vulnerability of their wide vessels to freeze embolism.
Location  Coastal ranges of south-central Chile (latitude 37°–40° S) and western South Island of New Zealand (41°–43° S).
Methods  The presence of all woody species was recorded in plots of 2500 m² (Chile) or 100–400 m² (New Zealand) on four elevational gradients in temperate rain forest. Each species was classified as a tree, shrub or liana. Original data were obtained from 22 plots at two sites in Chile. In New Zealand, two surveys comprising a total of 296 plots were extracted from the National Vegetation Survey data base.
Results  Liana species richness declined more or less monotonically on all four gradients, whereas richness of trees and shrubs showed more varied elevational patterns. The proportion of woody species contributed by the liana life-form was negatively correlated with elevation on all four gradients, falling from 15 to 35% of the woody flora at c . 200 m a.s.l. to nil well below the tree line. The elevational and latitudinal limits of liana species were marginally significantly correlated in Chile, but not in New Zealand.
Main conclusions  The elevational parallel of the well-documented decline in liana representation with increasing latitude is consistent with the hypothesis that cold intolerance is a strong control on the global distribution of the liana life-form.     

Does prescribed burning result in biotic homogenization of coastal heathlands?

Biotic homogenization due to replacement of native biodiversity by widespread generalist species has been demonstrated in a number of ecosystems and taxonomic groups worldwide, causing growing conservation concern. Human disturbance is a key driver of biotic homogenization, suggesting potential conservation challenges in seminatural ecosystems, where anthropogenic disturbances such as grazing and burning are necessary for maintaining ecological dynamics and functioning. We test whether prescribed burning results in biotic homogenization in the coastal heathlands of north‐western Europe, a seminatural landscape where extensive grazing and burning has constituted the traditional land‐use practice over the past 6000 years. We compare the beta‐diversity before and after fire at three ecological scales: within local vegetation patches, between wet and dry heathland patches within landscapes, and along a 470 km bioclimatic gradient. Within local patches, we found no evidence of homogenization after fire; species richness increased, and the species that entered the burnt Calluna stands were not widespread specialists but native grasses and herbs characteristic of the heathland system. At the landscapes scale, we saw a weak homogenization as wet and dry heathland patches become more compositionally similar after fire. This was because of a decrease in habitat‐specific species unique to either wet or dry habitats and postfire colonization by a set of heathland specialists that established in both habitat types. Along the bioclimatic gradient, species that increased after fire generally had more specific environmental requirements and narrower geographical distributions than the prefire flora, resulting in a biotic ‘heterogenisation’ after fire. Our study demonstrates that human disturbance does not necessarily cause biotic homogenization, but that continuation of traditional land‐use practices can instead be crucial for the maintenance of the diversity and ecological function of a seminatural ecosystem. The species that established after prescribed burning were heathland specialists with relatively narrow geographical ranges.     

Habitat associations of species show consistent but weak responses to climate

Different vegetation types can generate variation in microclimates at local scales, potentially buffering species from adverse climates. To determine if species could respond to such microclimates under climatic warming, we evaluated whether ectothermic species (butterflies) can exploit favourable microclimates and alter their use of different habitats in response to year-to-year variation in climate. In both relatively cold (Britain) and warm (Catalonia) regions of their geographical ranges, most species shifted into cooler, closed habitats (e.g. woodland) in hot years, and into warmer, open habitats (e.g. grassland) in cooler years. Additionally, three-quarters of species occurred in closed habitats more frequently in the warm region than in the cool region. Thus, species shift their local distributions and alter their habitat associations to exploit favourable microclimates, although the magnitude of the shift (approx. 1.3% of individuals from open to shade, per degree Celsius) is unlikely to buffer species from impacts of regional climate warming.     

Forest cover reduces thermally suitable habitats and affects responses to a warmer climate predicted in a high-elevation lizard

Warmer climates have affected animal distribution ranges, but how they may interact with vegetation patterns to affect habitat use, an important consideration for future wildlife management, has received little attention. Here, we use a biophysical model to investigate the potential thermal impact of vegetation pattern on the habitat quality of a high-elevation grassland lizard, Takydromus hsuehshanensis, and to predict the thermal suitability of vegetation for this species in a future warmer climate (assuming 3 °C air temperature increase). We assess the thermal quality of vegetation types in our study area (Taroko National Park in areas >1,800 m) using three ecologically relevant estimates of reptiles: body temperature (T _b), maximum active time, and maximum digestive time. The results show that increasing forest canopy gradually cools the microclimates, hence decreasing these estimates. In the current landscape, sunny mountain-top grasslands are predicted to serve as high quality thermal habitat, whereas the dense forests that are dominant as a result of forest protection are too cold to provide suitable habitat. In simulated warmer climates, the thermal quality of dense forests increases slightly but remains inferior to that of grasslands. We note that the impact of warmer climates on this reptile will be greatly affected by future vegetation patterns, and we suggest that the current trend of upslope forest movement found in many other mountain systems could cause disadvantages to some heliothermic lizard species.     

Temperature effects in relation to the patterns of distribution and abundance of three species in the Drosophila affinis subgroup

Abstract. 1. The effect of temperature on several aspects of the biology of Drosophila affinis, D.algonquin and D.athabasca were investigated in order to gain insight as to why these closely related species differ in their spatial and temporal patterns of occurrence.
2. No differences exist between the three species in the effect of temperature on egg hatchability.
3. Productivities of the three species were measured in a temperature gradient device. These measurements indicated that D.athabasca has an advantage in productivity over D.affinis at cooler temperatures (below 25°C). At warmer temperatures (above 25°C) the greater productivity of D.affinis makes it the better species. Drosophila algonquin , like D.athabasca , was characterized by its productivity curve as a species more adapted to cooler climates.
4. The competitive abilities of the three species were measured at 25°C, at 18°C, and in an environment which fluctuated between 18 and 25°C on a daily cycle. Competitive ability was estimated on the basis of the performance of the species in two- and three-species populations.
5. In general, D.affinis exhibited its highest competitive ability relative to the other two species at the highest experimental temperature. D.athabasca had its highest competitive ability at the lowest temperature. The competitive ability of Ddgonquin was intermediate; less than that of D.affinis but greater than D.athabasca .
6. The dependence of competitive ability and productivity on temperature is thought to be partly responsible for the differences between the species with respect to their geographic distributions and their spatial and temporal patterns in natural populations.     

Determinants of species richness in patches of grassland and heathland in Himmerland (Denmark)

Island biogeographical theory predict species richness to increase with habitat area and to decrease with isolation from colonisation sources. This theory has been applied to habitat fragments, and the predictions tested and found valid in empirical studies on fragments of deciduous woodland in northern Europe. However, previous results on fragments of grassland have been ambiguous. In the present study the species richness of vascular plants was investigated in 63 patches of grassland and heathland scattered in an agricultural and forested landscape using multiple linear regression. The relationship between species richness and patch area, isolation and local habitat conditions including heterogeneity were examined. Area was an important determinant of species richness, both in the full data set and in a subset of small habitat patches. In contrast, spatial isolation and habitat heterogeneity were not important factors determining species richness. Differences in soil acidity were accounting for a large proportion of the variance in species richness. This result is probably due to differences in the size of the regional species pools of grasslands with different levels of soil pH.     

Overwintering survivorship of pupae of the mimosa web worm, Homadaula anisocentra (Lepidoptera: Plutellidae), in an urban landscape

ABSTRACT. 1. Overwintering survivorship of pupae of the mimosa webworm, Homadaula anisocentra Meyrick (Lepidoptera: Plutellidae), was examined in several urban habitats in central Iowa during the winters of 1981–82, 1982–83 and 1983–84.
2. Survivorship and supercooling point temperatures were determined throughout the winters. Corroborative laboratory studies were conducted during the winter of 1982–83.
3. Minimum ambient temperatures that equalled or were below the supercooling point of the insect, at any time, were lethal.
4. Prolonged cold exposure below 0°C and above the supercooling point resulted in high mortality levels. To quantify this relationship, a concept of minimum-temperature exposure was developed by tabulating the number of degrees that the daily minimum temperature was below 0°C for a given sampling period.
5. Some mimosa webworm pupae were found to overwinter in highly protected sites (2.5–5.0°C warmer than the ambient air temperatures) in the urban environment, resulting in less minimum-temperature exposure and reducing the probability of reaching the lethal supercooling point temperature.     

Synchrony in small mammal populations of montane forest patches in southern India

1. Small mammal populations were studied in montane evergreen forests in the Nilgiris, southern India, using live-trapping from January 1994 to September 1996. Two sites were selected, one with a single large forest patch and adjoining plantations, and the other with several small forest patches separated by grassland.
2. Nine species were recorded, of which eight were trapped in the forest patches, two in the grasslands and five in plantations. Rattus rattus was the most abundant species (2–36 individuals ha⁻¹) followed by Suncus montanus (0–11 individuals ha⁻¹). Densities of seven other species recorded were very low.
3. Synchrony in various population variables – density, biomass, mean weight, density of adults and adult females, and proportion of adults, adult females and sub-adults – was examined for Rattus rattus populations in the forest patches and plantations. Density and biomass were studied in seven other species recorded at these sites. Spearman's rank order correlation was used as a measure of synchrony between the population variables.
4. Within-site synchrony was higher than between-site synchrony in population characters. Synchrony was also higher between plots within the unfragmented site than they were between plots in the fragmented site. Relatively high synchrony in proportions of adults, adult females and juveniles in the forest patches implied that breeding is probably influenced by climate and food availability, which are seasonal in this habitat.
5. Given the small patch sizes (≈1–10 ha) and low population sizes, asynchrony is likely to be an outcome of demographic and environmental stochasticity, and low dispersal rates may impede establishment of synchrony.     

Latitudinal compensation in oyster ciliary activity

1. Theories of latitudinal compensation predict that individuals living in colder temperature regimes should physiologically compensate for the slowing of standard physiological rates, owing to the relatively low temperature of their local environments, by increasing their metabolic rate in colder water temperatures relative to individuals living in warmer water temperature regimes.
2. This hypothesis was tested with oyster strains originally from geographically separated populations that were raised in a common environment for seven generations. The physiological parameter measured was ciliary activity across a temperature gradient.
3. Support for the latitudinal compensation hypothesis was found: the strain originally from the colder temperature regime had more active cilia at lower experimental temperatures than individuals originally from the warmer temperature regime. Ciliary activity of the more northern Long Island Sound oysters was significantly greater than activity in the more southern Delaware Bay oysters at temperatures of –1, 2 and 6 °C.
4. These results suggest that there is genetically based physiological differentiation between these populations of oysters consistent with the latitudinal compensation for local temperature regime.     

Over-wintering stage polymorphism of a bruchine beetle: geographical variation in optimal diapause strategy

Abstract.  1. Most temperate insects undergo diapause at a specific developmental stage to overcome severe winters. The bruchine beetle Bruchidius dorsalis in a warmer region in Japan, however, has diverse over-wintering stages – non-diapausing larvae, diapausing larvae, and diapausing adults, whereas in a cooler region, it over-winters only as the final instar larva or adult in diapause.
2. The geographical pattern of over-wintering stages in 12 populations was investigated over a wide range along the mainland of Japan. It revealed that a distinct geographical pattern of over-wintering stages exists in relation to temperature conditions. In regions with warmer climates, the proportion of non-diapausing larvae increased and B. dorsalis had a more complex over-wintering stage structure.
3. Life cycles were also compared between two areas of Japan by field experiments. In the cooler area, the first generation over-wintered in the diapausing larval or adult stage. Conversely, in the warmer area, diapause was induced later and some of the first-generation adults produced second-generation offspring before over-wintering.
4. Based on the geographical cline of climates and the differences in cold hardiness among stages, we can demonstrate that the over-wintering stage variation among and within populations results from an adaptive timing of diapause induction in each region, because the late larval or adult diapauses protect pupae or eggs – which unlike other stages are not cold hardy – from being produced late in the season.     

Plant species richness and composition of heathland relics in north‐western Belgium: evidence for a rescue‐effect?

Aim To study the effects of the degree of fragmentation of heathland patches on their species richness and species composition, and to infer the ecological mechanisms behind the observed patterns. Location The heathland patches of the north‐western part of Flanders, Belgium. During the last 200 years, the heathland area in this study area has been reduced from c. 10,000 to c. 40 ha, yielding c. 150 small and highly isolated relic fragments. Methods Different isolation measures were calculated for each of the 153 inventoried heathland patches. The influence of isolation, area and habitat diversity on species richness was investigated using correlation techniques. Community composition of the patches was tested for nestedness, and the mechanisms potentially underlying this pattern were determined. Results and main conclusions Both the analyses at the species richness and community composition level yielded evidence for a positive species–area relation. This relation was not caused by higher habitat heterogeneity in larger patches. Patch isolation, measured in different ways, however seemed much more important in explaining species richness and community composition than fragment area. Our results indicate that area effects are overcompensated by the rescue‐effect: if a patch is close to other patches, species can disperse between them and prevent the species from going extinct. Species having a short living seed bank were also more sensitive to isolation than species with a long living seed bank, indicating that the latter most probably depend on the seed bank to survive periods when environmental conditions are harsh. Analogously to the spatial rescue‐effect, the existence of a persistent seed bank may lead to a sort of temporal rescue‐effect, where the extinction of a plant species is prevented through survival in the seed bank of a patch.     

Geographical variation in Japan in egg development of the mayfly, Ephoron shigae (Ephemeroptera: Polymitarcyidae)

1. The effect of temperature on embryonic development was compared in four populations, two bisexual and two unisexual, of Ephoron shigae , including one each near the northern and southern periphery of the species range in Japan.
2. Eggs from every population were chilled at 4, 8 or 12 °C for diapause development after 50 days at 20 °C for pre-diapause development (experiment I). Some eggs hatched during chilling at 8 °C or 12 °C, whereas no eggs hatched at 4 °C. The rate of hatching in a given condition of chilling was higher for the eggs from warmer winter environments.
3. Chilling at 4 or 8 °C effectively facilitated diapause development. Chilling at 12 °C was, in general, not so effective, but relatively effective for the eggs from warmer winter environments.
4. Eggs were incubated at 8, 12, 15 or 20 °C after chilling at 4 °C to examine the effect of temperature on post-diapause development (experiment II). The eggs incubated at higher temperature after chilling hatched quicker and more synchronously and had higher hatching success.
5. The relationship between temperature and the days required for hatching after chilling was well described by the power function. There was no significant difference in the slope of the regression lines (i.e. temperature dependency) among local populations. However, a longer time was required for hatching at a given temperature for the population from the colder winter environment.
6. There was no detectable difference in the observed intraspecific variations between unisexual and bisexual populations.     

Genetic erosion in a stenotopic heathland ground beetle (Coleoptera: Carabidae): a matter of habitat size?

Two centuries ago large areas of north-west Europe were covered by coherent heathlands which hosted numerous specialized species. Changes in land use made heathlands fragmented and rare, consequently, they are in the focus of nature conservation efforts today. But how large should remaining heathland patches be in order to secure the survival of populations of specialized species? We investigated the genetic diversity at five allozyme loci of Poecilus lepidus, a flightless and stenotopic heathland ground beetle. 29 populations from differently sized heathland patches in north-west Germany were analyzed. Results show a weak but significant genetic differentiation and no evidence for isolation by distance or other patterns of spatial autocorrelation. Linear regression analysis revealed significant relationships between patch size, allelic richness, number of alleles and expected heterozygosity. These findings are explained by severe habitat fragmentation together with strong fluctuations in population size which have been reported for this species in the past. To conserve the vast majority of the species’ genetic diversity for a period of 100 years we suggest to maintain heathland patches of at least 50 ha in size.     

Seasonal development of annual shoots in some terrestrial orchids from Russia and Japan

Annual shoots of 46 terrestrial orchid species commonly found in wide ranges of temperate climates in Russia and Japan change their patterns of growth recurrence from the dormancy state, through formation and growth, to the next dormancy state during the course of yearly response to seasonal cycles of environmental conditions. Each of the species has its own strategy in seasonal development of aerial shoots, rhizomes, tubers and roots, and shows seasonal differentiation of shoot morphogenesis at the early stage of new shoot apex formation in accordance with its growth habit, habitat and range size of geographical distribution. Perennial orchids with sympodial growth patterns and primitive life forms are characterized by long duration of shoot and inflorescence development inside the bud. Among the species studied, the orchids that have annually regenerating root-stem tubers have the shortest duration of root and shoot morphogenesis. The species that have predominant patterns of monopodial growth show variability in duration of lateral shoot growth due to the energy budget of the mother plant. The species which have latitudinally long ranges of distribution from northern colder regions to southern warmer regions tend to take longer for shoot development inside the bud, and aerial shoots have a shorter life-span in the northern regions than those in the south.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

Climate change,extinction, and Sky Island biogeography in a montane lizard

Around the world, many species are confined to “Sky Islands,” with different populations in isolated patches of montane habitat. How does this pattern arise? One scenario is that montane species were widespread in lowlands when climates were cooler, and were isolated by local extinction caused by warming conditions. This scenario implies that many montane species may be highly susceptible to anthropogenic warming. Here, we test this scenario in a montane lizard (Sceloporus jarrovii) from the Madrean Sky Islands of southeastern Arizona. We combined data from field surveys, climate, population genomics, and physiology. Overall, our results support the hypothesis that this species' current distribution is explained by local extinction caused by past climate change. However, our results for this species differ from simple expectations in several ways: (a) their absence at lower elevations is related to warm winter temperatures, not hot summer temperatures; (b) they appear to exclude a low‐elevation congener from higher elevations, not the converse; (c) they are apparently absent from many climatically suitable but low mountain ranges, seemingly “pushed off the top” by climates even warmer than those today; (d) despite the potential for dispersal among ranges during recent glacial periods (~18,000 years ago), populations in different ranges diverged ~4.5–0.5 million years ago and remained largely distinct; and (e) body temperatures are inversely related to climatic temperatures among sites. These results may have implications for many other Sky Island systems. More broadly, we suggest that Sky Island species may be relevant for predicting responses to future warming.