Relative importance of quantity, quality and isolation of patches for butterfly diversity in fragmented urban forests

The majority of forests in urban areas are small and isolated. Improving habitat quality of small forests instead of increasing habitat size and connectivity could be an effective means of conserving the biodiversity of such highly fragmented landscapes. In this study, we investigated the relative importance of habitat quantity, quality and isolation on butterfly assemblages in urban fragmented forests in Tokyo, Japan. We used four habitat geographic parameters: (1) fragment size, (2) shape index, (3) isolation (distance to the mainland), and (4) connectivity; and three habitat quality parameters: (1) herbaceous nectar plant abundance, (2) herbaceous nectar plant diversity, and (3) larval host plant diversity. We surveyed butterfly assemblages along transects in 20 forest fragments that ranged in size from 1 to 122 ha. We used generalized linear models to relate the number of species in a fragment to four habitat geographic parameters and three habitat quality parameters. The averaged models based on AIC_c showed that fragment size had a strong positive effect on butterfly species richness. There was also a positive effect of herbaceous nectar plant abundance on species diversity. These findings suggest that improving the habitat quality of small and isolated forests in highly fragmented landscapes may be capable of maintaining levels of butterfly diversity comparable to those of large fragments.     

Relative importance of habitat and landscape scales on butterfly communities of urbanizing areas

Agricultural decline and urbanization entail rapid alterations of the patterns of organization of rural landscapes in Europe. The spread of the urban footprint to the adjacent countryside contributes to the development of new anthropogenic ecosystems in formerly rural hinterlands. In this study, butterflies are considered as biological indicators of these rapid environmental changes. Our purpose is to better understand changes in biodiversity related to the evolution of available habitats in a mutating landscape. In this study, we investigate butterfly communities of four land-use types (fallow lands, gardens, vineyards, woodlands) within different landscape contexts. Our results reveal that variations in structure and functional composition of these communities are related to different levels of human disturbance at both landscape scale and habitat scale.     

Patch occupancy in the endangered butterfly Lycaena helle in a fragmented landscape: effects of habitat quality, patch size and isolation

While there is agreement that both habitat quality and habitat network characteristics (such as patch size and isolation) contribute to the occupancy of patches by any given species, the relative importance of these factors is under debate. This issue is of fundamental ecological importance, and moreover of special concern for conservation biologists aiming at preserving endangered species. Against this background we investigated patch occupancy in the violet copper Lycaena helle, one of the rarest butterfly species in Central Europe, in the Westerwald area (Rhineland-Palatinate, Western Germany). Occupied (n = 102) differed from vacant (n = 128) patches in altitude, size, connectivity, availability of wind shelter, in the abundance of the larval host-plant, in the abundance of a grass species indicating favorable habitat conditions and in the abundance of nitrophilous plants. Overall, patch occupancy was primarily determined by patch size, connectivity and the abundance of the larval host plant, while all other parameters of habitat quality were of subordinate importance. Therefore, our findings suggest that even for extremely sedentary species such as L. helle habitat networks are decisive and—next to the preservation of habitat quality—need to be an integral part of any conservation management for this species.     

When habitat management can be a bad thing: effects of habitat quality,isolation and climate on a declining grassland butterfly

The conservation of most temperate grassland habitats and their characteristic fauna and flora requires regular low-intensive forms of land-use to counteract natural succession. Although many species tolerate moderate disturbance regimes, some are known to be susceptible to grazing or mowing, thereby causing a management dilemma. One of these species is the Woodland Ringlet butterfly, Erebia medusa. In this study, we analysed which environmental factors determine the occurrence of E. medusa in the Diemel Valley (Central Germany). Furthermore, we conducted microclimatic measurements during the winter months to investigate the role of the litter layer as a microclimatic buffer. Patch occupancy in the Diemel Valley was well explained by the amount of litter present in a patch and connectivity to other inhabited patches. The role of local climatic conditions could not be clarified, due to inter-correlations with connectivity. During the winter, the air temperature inside the litter layer was significantly less variable than above it. We conclude that the current distribution of E. medusa in the Diemel Valley is caused by the combined effect of habitat quality and connectivity, and perhaps also by climatic factors. The importance of the litter layer reflects the dependence of E. medusa on low-intensive or absent land-use. In addition, the litter layer possibly constitutes an essential habitat element, as it buffers temperature fluctuations and thus probably reduces the energy consumption of overwintering larvae. Given the species’ preference for abandoned grasslands, the conservation of E. medusa requires a low-intensity habitat management, for example, by rotational grazing or mowing of small parts of the sites. On the landscape level, the preservation of well-connected habitat networks is important.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.     

Relative importance of habitat area, isolation and quality for the occurrence of middle spotted woodpecker Dendrocopos medius (L.) in Sweden

In a stepwise regression analysis, oak habitat area and habitat isolation were the most important parameters explaining numbers, density, regularity, and probability of presence in two successive years of middle spotted woodpeckers. No estimate of habitat quality contributed significantly to the equations. A 50% regularity of occurrence during the study period was found in oak areas of 75 ha. It is suggested that, for a small population, habitat fragmentation that gives rise to partially isolated subpopulations will make the population more vulnerable. The large area required by Swedish middle spotted woodpeckers compared with Central European ones, is believed to be a result either of an 'extinction disequilibrium' or of poorer habitats in the periphery of the distribution area of the species.     

Relative importance of isolation, area and habitat heterogeneity for vascular plant species richness of temporary wetlands in east-German farmland

The purpose of this investigation was to explore the determinants of vascular plant species richness for temporary, isolated wetland habitats which are influenced by hydrologic gradients and characterized by variation in habitat area. The dependent variables total plant species richness and the number of obligate wetland species were analyzed consecutively. In regression analyses habitat area explained between 11 and 15% of the variation in the dependent variables. Habitat area was correlated with the heterogeneity of the hydroperiods between the upper and lower parts of the hydrologic gradients. In multivariate regression analyses, habitat heterogeneity accounted for 70-77% of the variation in the dependent variables, and habitat area did not have a significant impact. The results are most consistent with the habitat heterogeneity hypothesis. I therefore concluded that area is a surrogate variable for habitat heterogeneity which directly enhances vascular plant species diversity. There was no significant impact of isolation on species richness. The data suggest that the expanses of agricultural fields are not an effective barrier to the dispersal of the studied plant species. Only 10 of,52 wetland species were negatively influenced by isolation. This group of species did not differ from the other wetland species with respect to dispersal strategies and longevity of seed banks. However, the longevity of the seed banks was generally high, and there was a dominance of species whose propagules are transported with the soil clinging to the feet of birds. The results are discussed in the context of accurate dispersal strategies and remnant populations, which may counteract the effects of isolation.     

Effects of seasonality,isolation and patch quality for habitat selection processes by mute swans Cygnus olor in a fishpond landscape

Foragers in patchy environments do not only select sites for single patch characteristics, but also have to consider the local environment of such patches. We studied habitat selection by mute swans Cygnus olor in a wide and heterogeneous fishpond region (the Dombes, eastern France). In this study, we considered fishpond isolation, resource quality within fishponds and breeding status of mute swans during both summer and winter. Mute swans did not select aquatic habitat randomly within the landscape. During summer, the population spread preferentially on medium to large fishponds, in subregions with numerous or closely related waterbodies, without generating a clumped distribution of birds. In addition to a positive effect of local fishpond number (2 km radius), breeding birds also responded positively to fishpond size. Non‐breeders selected fishponds mainly according to their size. Intraspecific territoriality did not appear to limit the presence of non‐breeders (i.e. moulting flocks), since both breeders and non‐breeders could coexist on the larger fishponds. During winter, mute swans used medium to large reflooded fishponds after summer drainage. The surrounding aquatic environment of fishponds played a minor role in determining flocking, compared to actual patch quality. Flocking occurred on large fishponds that had reflooded after having dried the summer before, whatever the agricultural cultivation practiced in the summer following drainage. The results suggest that geographical aspects should be taken into account when considering the potential impact of this expanding species within such ecosystems, and also in more general management policies dealing with aquatic habitats for waterbird populations.     

栖息地质量对两种网蛱蝶集合种群结构和分布的影响

在河北省赤城县研究了栖息地质量对大网蛱蝶Melitaea phoebe和金堇蛱蝶Euphydryas aurinia两种网蛱蝶集合种群结构和分布的影响。这两种网蛱蝶在约10 km_²的区域内共存,成虫期的蜜源植物几乎相同,大网蛱蝶的发生峰期比金堇蛱蝶晚约一个月,两者只有不到一周左右的时间重叠。大网蛱蝶和金堇蛱蝶幼虫的寄主植物分别是: 祁州漏芦（菊科）和华北蓝盆花（川续断科）。蜜源植物的丰度与两种网蛱蝶的局域种群大小呈正相关;祁州漏芦的密度对大网蛱蝶的局域种群大小影响很大,金堇蛱蝶的局域种群大小则与其寄主植物华北蓝盆花的高度正相关;斑块内平均植被高度与两种网蛱蝶的局域种群大小均呈正相关,植物多样性、植物均匀性和植被盖度均与金堇蛱蝶的局域种群大小负相关,与大网蛱蝶的关系不大。同时分析了其他因子如斑块的坡向、坡度等的影响。主要结论是：1）幼虫寄主植物的不同和成蝶飞行峰期的分离允许两种网蛱蝶在这样一个小的斑块区域内共存;2）蜜源是重要的限制因子,并且受气候随机性的影响很大,蜜源的波动可以很好地解释网蛱蝶集合种群在年度间的动态变化;3）大网蛱蝶和金堇蛱蝶的飞行、食物搜寻能力的不同以及各自寄主植物的生物学特性、空间分布的不同决定了它们具有不同的集合种群结构： 金堇蛱蝶是经典的集合种群,大网蛱蝶是源-汇集合种群;4）斑块质量和昆虫行为共同决定了两种网蛱蝶的集合种群结构和分布。     

Relative importance of landscape and species characteristics on extinction debt,immigration credit and relaxation time after habitat turnover

Habitat turnover concomitantly causes destruction and creation of habitat patches. Following such a perturbation, metapopulations harbor either an extinction debt or an immigration credit, that is the future decrease or increase in population numbers due to this disturbance. Extinction debt and immigration credit are rarely considered simultaneously and disentangled from the relaxation time (time to new equilibrium). In this contribution, we test the relative importance of two potential drivers of time-delayed metapopulation dynamics: the spatial configuration of the habitat turnover and species dispersal ability. We provide a simulation-based investigation projecting metapopulation dynamics following habitat turnover in virtual landscapes. We consider two virtual species (a short-distance and a long-distance disperser) and five scenarios of habitat turnover depending on net habitat loss or gain and habitat aggregation. Our analyses reveal that (a) the main determinant of the magnitude of the extinction debt or immigration credit is the net change in total habitat area, followed by species dispersal distance and finally by the post-turnover habitat aggregation; (b) relaxation time weakly depends on the magnitude of the immigration credit or of the extinction debt; (c) the main determinant of relaxation time is dispersal distance followed by the net change in total habitat area and finally by the post-turnover habitat aggregation. These results shed light on the relative importance of dispersal ability and habitat turnover spatial structure on the components of time-delayed metapopulation dynamics.     

Relative importance of colonist quantity,quality, and arrival frequency to the extinction of two zooplankton species

Introduced plants can positively affect population viability by augmenting the diet of native herbivores, but can negatively affect populations if they are subpar or toxic resources. In organisms with complex life histories, such as insects specializing on host plants, the impacts of a novel host may differ across life stages, with divergent effects on population persistence. Most research on effects of novel hosts has focused on adult oviposition preference and larval performance, but adult preference may not optimize offspring performance, nor be indicative of host quality from a demographic perspective. We compared population growth rates of the Baltimore checkerspot butterfly, Euphydryas phaeton, on an introduced host, Plantago lanceolata (English plantain), and the native host Chelone glabra (white turtlehead). Contrary to the previous findings suggesting that P. lanceolata could be a population sink, we found higher population growth rates (λ) on the introduced than the native host, even though some component parameters of λ were higher on the native host. Our findings illustrate the importance of moving beyond preference–performance studies to integrate vital rates across all life stages for evaluating herbivore–host plant relationships. Single measures of preference or performance are not sufficient proxies for overall host quality nor do they provide insights into longer term consequences of novel host plant use. In our system, in particular, P. lanceolata may buffer checkerspot populations when the native host is limiting, but high growth rates could lead to crashes over longer time scales.     

Relative importance of density-dependent regulation and environmental stochasticity for butterfly population dynamics

The relative contribution of density-dependent regulation and environmental stochasticity to the temporal dynamics of animal populations is one of the central issues of ecology. In insects, the primary role of the latter factor, typically represented by weather patterns, is widely accepted. We have evaluated the impact of density dependence as well as density-independent factors, including weather and mowing regime, on annual fluctuations of butterfly populations. As model species, we used Maculinea alcon and M. teleius living in sympatry and, consequently, we also analysed the effect of their potential competition. Density dependence alone explained 62 and 42% of the variation in the year-to-year trends of M. alcon and M. teleius, respectively. The cumulative Akaike weight of models with density dependence, which can be interpreted as the probability that this factor should be contained in the most appropriate population dynamics model, exceeded 0.97 for both species. In contrast, the impacts of inter-specific competition, mowing regime and weather were much weaker, with their cumulative weights being in the range of 0.08–0.21; in addition, each of these factors explained only 2–5% of additional variation in Maculinea population trends. Our results provide strong evidence for density-dependent regulation in Maculinea, while the influence of environmental stochasticity is rather minor. In the light of several recent studies on other butterflies that detected significant density-dependent effects, it would appear that density-dependent regulation may be more widespread in this group than previously thought, while the role of environmental stochasticity has probably been overestimated. We suggest that this misconception is the result of deficiencies in the design of most butterfly population studies in the past, including (1) a strong focus on adults and a neglect of the larval stage in which density-dependent effects are most likely to occur; (2) an almost exclusive reliance on transect count results that may confound the impact of environmental stochasticity on butterfly numbers with its impact on adult longevity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population structure of a monophagous moth in a patchy landscape

1. The population structure of a monophagous noctuid moth, Abrostola asclepiadis , living on a patchily distributed perennial herb, Vincetoxicum hirundinaria is described. The study took place over 5 years at a landscape scale (about 12 km²).
2. Patch occupancy rates and population densities were studied in relation to patch size, degree of patch isolation, level of sun exposure and distance from the coast. In addition, flight tests in the laboratory were performed to estimate the potential dispersal capacity of the moth.
3. Occupancy rates were high and the likelihood of extinction depended on patch size. Small patches were less likely to be occupied than were large patches (> 10 m²). Sun-exposed patches were occupied for a lower proportion of years than were shaded patches. No distance effects could be discerned at the spatial scale of study, presumably because the insect is a strong flier.
4. Population densities in occupied patches decreased with increasing patch size. Furthermore, insect densities tended to increase with distance from the coast. Density changes in patches were synchronized.
5. The studied insect population can be described as a 'patchy population' sensu Harrison (1991) with spatially correlated population dynamics. These dynamics are superimposed on a landscape gradient.     

Additive partitioning of butterfly diversity in a fragmented landscape: importance of scale and implications for conservation

Aim Most of the Atlantic Forest in Brazil occurs in fragments of various sizes. Previous studies indicate that forest fragmentation affects fruit‐feeding butterflies. Conservation strategies that seek to preserve organisms that are distributed in high‐fragmented biomes need to understand the spatial distribution of these organisms across the landscape. In view of the importance of understanding the fauna of these forest remnants, the objective of the present work is to investigate the extent to which the diversity of this group varies across spatial scales ranging from within‐forest patches to between landscapes. Location South America, south‐eastern Brazil, São Paulo State. Methods We used bait traps to sample fruit feeding butterflies at 50 points in 10 fragments in two different landscapes during a period of 12 months. Total species richness and Shannon index were partitioned additively in diversity at trap level, and beta diversity was calculated among traps, among forest patches, and between landscapes. We used permutation tests to compare these values to the expected ones under the null hypothesis that beta diversity is only a random sampling effect. Results There was significant beta diversity at the smallest scale examined; however, the significance at higher scales depends on the diversity measurement used. Beta diversity with Shannon index was smaller than expected by chance among fragments, whereas species richness was not. Among landscapes, only beta diversity in richness was higher than expected by chance. Main conclusions The results observed occur because there is great variability in species composition among forest patches in the same landscape, changing this diversity even though the communities are formed from the same pool of species. At the largest scale evaluated (between landscapes), these pattern changes and differences in beta diversity in richness were detectable. This difference is probably caused by the presence of rare species. Thus, a conservation strategy that seeks to preserve as many species as possible per unit of area in high‐fragmented biomes should give priority to protecting fragments in different landscapes, rather than more fragments in the same landscape.     

Effects of habitat fragmentation on the genetic structure of the monophagous butterfly Polyommatus coridon along its northern range margin

Population genetic patterns of species at their range margin have important implications for species conservation. We performed allozyme electrophoresis of 19 loci to investigate patterns of the genetic structure of 17 populations (538 individuals) of the butterfly Polyommatus coridon, a monophagous habitat specialist with a patchy distribution. The butterfly and its larval food plant Hippocrepis comosa reach their northern distribution margin in the study region (southern Lower Saxony, Germany). Butterfly population size increased with host plant population size. The genetic differentiation between populations was low but significant (FST = 0.013). No isolation-by-distance was found. Hierarchical F-statistics revealed significant differentiation between a western and an eastern subregion, separated by a river valley. The combination of genetic and ecological data sets revealed that the expected heterozygosity (mean: 18.5%) decreased with increasing distance to the nearest P. coridon population. The population size of P. coridon and the size of larval food plant population had no effect on the genetic diversity. The genetic diversity of edge populations of P. coridon was reduced compared to populations from the centre of its distribution. This might be explained by (i). an increasing habitat fragmentation towards the edge of the distribution range and/or (ii). a general reduction of genetic variability towards the northern edge of its distribution.     

Analysis of genetic diversity in a peatland specialist butterfly suggests an important role for habitat quality and small habitat patches

Understanding effects of habitat and landscape features on genetic variation is a prerequisite for the development of habitat and landscape management strategies aimed at conserving genetic diversity. While there has been considerable research on the effects of landscape structure on the genetics of populations, a recent review identified key biases in this body of work. The majority of landscape genetic studies investigate the intervening matrix’s influence on differentiation and gene flow among populations. Although characteristics of local habitat patches may be important determinants of genetic diversity, fewer studies have examined these relationships. Here we use node- and neighbourhood-based approaches to analyze correlates of genetic diversity in the bog copper (Lycaena epixanthe), a specialist butterfly endemic to temperate Nearctic peatlands that is threatened in parts of its range. Based on 190 repeatable and polymorphic amplified fragment length polymorphism loci, we found that genetic diversity was higher in habitat patches that were smaller and surrounded by more open water. Our results indicate that valuing small peatlands and preserving the surrounding water table may be important for conservation of genetic diversity in this highly specialized species. Our study highlights the importance of variables affecting habitat quality for conservation genetics.     

The influences of landscape structure on butterfly distribution and movement: a review

We review the literature on the influence of landscape structure on butterfly distribution and movement. We start by examining the definition of landscape commonly used in spatial ecology. Landscape-level processes are reviewed before focusing on the impact of the geometry and spatial arrangement of habitat patches on butterflies e.g. the nature of the matrix, patch size and shape, minimum area requirements, immigration and emigration, and temporal habitat dynamics. The role of landscape elements is reviewed in terms of corridors (and stepping-stones), barriers, nodes, environmental buffers, and prominent landmark features.     

The distribution and decline of a widespread butterfly Lycaena phlaeas in a pastoral landscape

1. Ecological specialists are often regarded as most likely to be threatened by anthropogenic habitat changes but few relevant data are available on changes in the status of widespread species. 2. Grid square distribution maps have been used widely to measure rates of decline and target conservation resources but it is known that coarse grain mapping is not appropriate to identify declines in widespread species that initially contain numerous local populations per grid cell. Changes in the status of widespread species need to be quantified. 3. Present‐day habitat associations, determined from over 2000 transect counts, combined with data on historical and present‐day habitat distributions, reveal that the area of occupancy and population‐level rate of decline of the Small Copper butterfly Lycaena phlaeas is likely to have been of the order of 92 and 89% respectively, in 35 km² of North Wales. Similar data on the species' major host plants Rumex acetosa and R. acetosella indicate possible declines in area occupied of 48 and 91%. If a 1‐km² grid was applied to the landscape, and if L. phlaeas, R. acetosa, and R. acetosella had occupied all 1‐km² cells in the study area in 1901 (non‐limestone cells for R. acetosella only), their declines would only have been recorded as 15, 9, and 35% respectively. 4. Many declining ecological specialists are threatened with extinction because of their initial rarity. At a population level, however, they may or may not be declining faster than less specialised species. The results presented here illustrate that some widespread species may have declined as much as many of Britain's rarities.