Resource complementation and the response of an insect herbivore to habitat area and fragmentation

Few studies have disentangled the effects of the area and fragmentation of a focal habitat type on species that use multiple habitat types within a landscape. We experimentally investigated the effects of habitat area, habitat fragmentation, and matrix composition on the movement and distribution of Melanoplus femurrubrum. Adults of this grasshopper feed preferentially on grasses, but oviposit almost exclusively in soil dominated by forbs. We compared population densities among plots that were made to vary in the area and fragmentation of clover habitat and composition of the matrix (grass or bare ground) within which clover habitat was embedded. In addition, a mark-recapture survey was conducted to examine effects of habitat area, fragmentation, and matrix composition on loss of individuals from a plot’s clover habitat and movement between clover subplots within plots. Overall densities of adult M. femurrubrum (averaged over clover and matrix) were 2.2× higher in plots where the matrix was composed of grass as compared to bare ground, and 1.8× higher in plots with 64 compared to 16 m² of clover habitat. Overall densities of nymphs were also positively influenced by greater clover area, but were unaffected by matrix composition. Within focal clover habitat embedded in grass matrix, adult densities were 2.1× higher in small clover subplots than large clover subplots. We conclude that the grass matrix had a positive effect on adult densities, but not nymph densities, because grass and forb-dominated habitats likely provide complementary resources only for adults. The aggregation of adults on small clover subplots within grass matrix was mainly attributed to a greater rate of emigration loss per unit area. In addition, this study emphasizes that a species’ response to changes in the area of a focal habitat type can depend significantly on the availability of complementary resources in the surrounding landscape. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of habitat fragmentation and soil quality on reproduction in two heathland Genista species

Habitat fragmentation decreases plant population size and increases population isolation, as well as altering patterns of plant–animal interactions, all of which may reduce plant fitness. Here, we studied effects of habitat fragmentation (in terms of population size and isolation) and soil quality on the reproduction of two rare legume species, Genista anglica (13 populations) and Genista pilosa (14 populations), confined to remnants of acidic and nutrient‐poor Calluna heathlands. Single individuals of the Genista plants are impossible to distinguish; population size was therefore estimated according to the area occupied (referred to as population size hereafter). We collected soil samples in all heathland sites to determine content of soil water, C, N, P, Ca, K and Mg. In both species values of soil pH and C/N ratio, as well as content of soil P and base cations, reflected the highly acidic and nutrient‐poor environment of the heathlands. Population sizes were unrelated to soil quality. Although the two Genista species are similar in morphology and ecology, effects of explanatory variables on reproduction were largely inconsistent across species. In G. anglica, population size had a positive impact on all reproductive variables except germination rate, which, in contrast, was the only variable affected positively by population size in G. pilosa. In both species, mean total reproductive output, calculated as the product of total seed mass per shoot and total germination, increased with increasing water content and decreased with increasing P. In G. anglica, we found positive effects of the C/N ratio on all reproductive variables except mean single and total seed mass per shoot. In summary, in both species reproductive success per shoot decreased with increasing soil nutrient availability in the heathland sites. The infestation of two large populations of G. pilosa with the pre‐dispersal, seed‐predating weevil Apion compactum had no significant effect on reproduction of the populations.     

黄河三角洲景观组分判定与景观破碎化分析

以地理信息系统（ＡＲＣ／ＩＮＦＯ）为手段,利用遥感卫星图片（ＴＭ５）及其他相关图件,以地貌、土壤与植被作为景观分类指标,把黄河三角洲的景观分成８大类型,３０个景观类型．在此基础上,以斑块的周长面积比值、相对面积及与其他景观类型之间的空间相关关系作为识别基质的指标,判定黄河三角洲景观的基质是柽柳芦苇潮盐土斜平地景观;根据斑块的周长面积比值识别廊道,并进行定量化研究．采用斑块密度对黄河三角洲景观的破碎化进行分析,发现生态交错带内斑块密度大;老河道附近景观破碎化严重     

The effect of periodic habitat fluctuations on a nonlinear insect population model

 Laboratory data show that populations of flour beetles (Tribolium), when grown in a periodically fluctuating volume of flour, can exhibit significant increases in numbers above those attained when grown in a constant volume (of the same average). To analyze and explain this phenomenon a discrete stage-structured model of Tribolium dynamics with periodic environmental forcing is introduced and studied. This model is an appropriately modified version of an experimentally validated model for flour beetle populations growing in a constant volume of flour, in which cannibalism rates are assumed inversely proportional to flour volume. This modeling assumption has been confirmed by laboratory experiments. Theorems implying the existence and stability of periodic solutions of the periodically forced model are proved. The time averages of periodic solutions of the forced model are compared with the equilibrium levels of the unforced model (with the same average flour volume). Parameter constraints are determined for which the average population numbers in the periodic environment are greater than (or less than) the equilibrium population numbers in the associated constant environment. Sample parameter estimates taken from the literature show that these constraints are fulfilled. These theoretical results provide an explanation for the experimentally observed increase in flour beetle numbers as a result of periodically fluctuating flour volumes. More generally, these integrated theoretical and experimental results provide the first convincing example illustrating the possibility of increased population numbers in a periodically fluctuating environment. Received 23 April 1996; received in revised form 28 March 1997     

The effects of energy input,immigration and habitat size on food web structure: a microcosm experiment

It has been hypothesised that larger habitats should support more complex food webs. We consider three mechanisms which could lead to this pattern. These are increased immigration rates, increased total productivity and spatial effects on the persistence of unstable interactions. Experiments designed to discriminate between these mechanisms were carried out in laboratory aquatic microcosm communities of protista and bacteria, by independently manipulating habitat size, total productivity and immigration rate. Larger habitats supported more complex food webs, with more species, more links per species and longer maximum and mean food chains, even in the absence of differences in total energy input. Increased immigration rate resulted in more complex food webs, but habitats with higher energy input per unit area supported less complex food webs. We conclude that spatial effects on the persistence of unstable interactions, and variation in immigration rates, are plausible mechanisms by which habitat size could affect food web structure. Variation in total productivity with habitat area seems a less likely explanation for variation in food web structure.     

Contrasting effects of habitat fragmentation,population density,and prey availability on body condition of two orb‐weaving spiders

1. Habitat fragmentation is a major threat to biodiversity because it disrupts movement between habitat patches. In addition, arthropod fitness may be reduced in fragmented habitats, e.g. due to reduced prey availability. 2. We studied the relationship of spider body condition with habitat fragmentation, population density, and prey availability. We expected that prey availability and population density of spiders would be affected by landscape composition and patch isolation. Body condition should be enhanced by high prey availability, but negatively affected by population density due to competition. 3. We sampled spiders on 30 groups of cherry trees that varied independently in the level of isolation from other woody habitats and in the percentage of woody habitat within 500 m radius. As a measure of body condition, we used residuals of the relationship between individual body mass/opisthosoma width and prosoma width of the two most common orb‐weaving spider species, Nuctenea umbratica Clerck and Araniella opisthographa Kulczynski. 4. Body condition of A. opisthographa was positively correlated with the abundance of flies, which increased with the percentage of forest in the landscape. In contrast, body condition of N. umbratica was reduced at high population densities, presumably due to intraspecific competition. In addition, body condition and population density of A. opisthographa was lower at isolated sites. 5. Our study suggests that effects of landscape fragmentation on body condition vary strongly between spider species, depending on the relative role of food limitation and intraspecific competition.     

The influence of habitat heterogeneity on host-pathogen population dynamics

The influence of spatial heterogeneity on the population dynamics of a naturally occurring invertebrate host-pathogen system was experimentally investigated. At ten week intervals over a two year period, I quantified the spatial distribution of natural populations of the terrestrial isopod crustacean Porcellio scaber infected with the isopod iridescent virus (IIV). During the seasonally dry periods of summer and early fall in central California, isopod populations were highly aggregated and the degree of patchiness and distance between inhabited patches was greatest. Coincident with increased patchiness and patch spacing was an increase in isopod density within patches. During the wet seasons of winter and spring, isopod population patchiness, inter-patch spacing, and within-patch density was low. Seasonal changes in virus prevalence were negatively correlated with within-patch density, patchiness, and inter-patch spacing. The influence of the spatial distribution of isopods on virus prevalence was also tested in field experiments. The virus was introduced into arrays of artificial habitat patches colonized by isopods in which interpatch distance was varied. The prevalence of resulting infections was monitored at weekly intervals. In addition, dispersal rates between artificial patches and natural patches were quantified and compared. The results showed that isopods in treatments with the smallest inter-patch spacing had the highest virus prevalence, with generally lower prevalence among isopods in more widely spaced patches. The spacing of experimental patches significantly affected virus prevalence, although the experiments did not resolve a clear relationship between patch spacing and virus prevalence. Rates of dispersal between patches decreased with increased patch spacing, and these rates did not differ significantly from dispersal between natural patches. The results suggest that rates of dispersal between isopod subpopulations may be an important component of the infection dynamics in this system. I discuss the consequences of these findings for host-pathogen dynamics in fragmented habitats, and for other ecological interactions in spatially heterogeneous habitats.     

Habitat quality predicts the distribution of a lizard in fragmented woodlands better than habitat fragmentation

Organisms often face a higher risk of local extinction in fragmented than in continuous habitat. However, whether populations are affected by reduced size and connectivity of the habitat or by changes in habitat quality in fragmented landscapes remains poorly investigated. We studied the regional distribution and microhabitat selection of the lacertid lizard Psammodromus algirus in a fragmented landscape where the existence of deciduous and evergreen woodlands brought about variation in habitat quality. Lizards never occupied any fragment smaller than 0.5 ha. However, above that limit fragment size no longer predicted lizard occurrence, which was explained by woodland type instead, with lizards being more frequently found in deciduous than in evergreen woodlands. Lizards selected microhabitats that had structural features favouring thermoregulation, foraging and predator avoidance, and we identified better conditions for thermoregulation and food acquisition in deciduous than in evergreen woodlands. Our results support the idea that variation in habitat quality can sometimes override the effect of habitat fragmentation on animal populations. We consider the implications of our study for the conservation of Mediterranean lizards, discussing our results in a broader context framed by previous studies conducted in nearby areas.     

The effect of habitat fragmentation on finescale population structure of wood frogs (Rana sylvatica)

We examined the impact of recent anthropogenic habitat fragmentation on the genetic structure of wood frog (Rana sylvatica) breeding sites in Wellington County of Ontario, Canada. In addition to geographic distance (average pairwise distance ~22 km, greatest distance ~50.22 km), four landscape features hypothesized to contribute to genetic differentiation between breeding sites were considered: road density, a major highway (highway 401), canopy cover, and watershed discontinuity. Analysis of data from 396 samples across nine breeding sites using eight microsatellite DNA loci, revealed a small degree of significant genetic structure between breeding sites. The presence of highway 401 and road density were correlated with small but statistically significant structure observed between several groups of sites. One outlier breeding site outside of Wellington County located within the city of Toronto, had significantly lower allelic richness and much larger population differentiation with the Wellington sites. Our data suggest that recent fragmentation has had an effect on wood frog population structure and also demonstrate the importance of dispersal for this species in maintaining levels of genetic diversity.     

Estimation of effective population size and detection of a recent population decline coinciding with habitat fragmentation in a ground beetle

We assess the impact of habitat fragmentation on the effective size (N(e)) of local populations of the flightless ground beetle Carabus violaceus in a small (<25 ha) and a large (>80 ha) forest fragment separated by a highway. N(e) was estimated based on the temporal variation of allele frequencies at 13 microsatellite loci using two different methods. In the smaller fragment, N(e) estimates ranged between 59 and a few hundred, whereas values between 190 and positive infinity were estimated for the larger fragment. In both samples, we detected a signal of population decline, which was stronger in the small fragment. The estimated time of onset of this N(e) reduction was consistent with the hypothesis that recent road constructions have divided a continuous population into several isolated subpopulations. In the small fragment, N(e) of the local population may be so small that its long-term persistence is endangered.     

Effects of environment,habitat configuration and forest continuity on the distribution of forest plant species

In present day European landscapes many forest plant species are restricted to isolated remnants of a formerly more or less continuous forest cover. The two major objectives of this study were (1) to determine the relative importance of habitat quality (mainly in terms of soil parameters), habitat configuration (patch size and isolation) and habitat continuity for the distribution of herbaceous forest plant species in a highly fragmented landscape and (2) to examine if groups of species with different habitat requirements are affected differently. Deciduous forest patches in northwestern Germany were surveyed for the presence of a large set of forest species. For each patch, habitat quality, configuration and continuity were determined. Data were analysed by Redundancy Analysis with variation partitioning for effects on total species composition and multivariate logistic regression for effects on individual species, for two different data sets (base‐rich and base‐poor forest patches). Overall, we found strong effects of habitat quality (particularly of soil pH, water content and topographic heterogeneity in the base‐rich forest patches; and of calcium content and disturbance in the base‐poor patches), but only relatively weak effects of habitat configuration and habitat continuity. However, a number of species were positively affected by patch area and negatively affected by patch isolation. Furthermore, the relative importance of habitat configuration tended to be higher for species predominantly growing in closed forests compared to species occurring both in the forest and in the open landscape.     

The effect of habitat fragmentation on cyclic population dynamics: a reduction to ordinary differential equations

Habitat fragmentation is known to be a key factor affecting population dynamics. In a previous study by Strohm and Tyson (Bull Math Biol 71:1323?C1348, 2009), the effect of habitat fragmentation on cyclic population dynamics was studied using spatially explicit predator?Cprey models with four different sets of reaction terms. The difficulty with spatially explicit models is that often analytical tractability is lost and the mechanisms behind the behaviour of the models are difficult to analyse. In this study, we employ a simplification procedure based on a Fourier series first-term truncation of the spatially explicit models Strohm and Tyson (Bull Math Biol 71:1323?C1348, 2009) to obtain spatially implicit models. These simpler models capture the main features of the spatially explicit models and can be used to explain the dynamics observed by Strohm and Tyson. We find that the spatially implicit models and the spatially explicit models produce similar responses to habitat fragmentation for larger high-quality patch sizes. Additionally, we find that the critical patch size of the spatially implicit models provides an upper bound on the critical patch size of the spatially explicit models. Finally, we derive an approximation of the multi-patch habitat by a single-patch habitat with partial flux boundary conditions which allows for a lower bound on the critical patch size to be calculated.     

A demographic analysis of vole population responses to fragmentation and destruction of habitat

 Fragmentation and destruction of natural habitats is currently considered to be the major threat to wildlife populations. We here perform a comprehensive analysis of the demographic effects of habitat fragmentation and destruction on 14 populations of the root vole. The experiment was divided into two consecutive periods. During the first period, we contrasted populations with the same initial size and structure in continuous and fragmented habitat. During the second period, we fragmented the continuous habitat into the same configuration as the permanently fragmented habitat so that the effect of habitat destruction could be evaluated. We estimated survival and fecundity parameters and combined them into population projection matrices to evaluate their relative impact on population growth. In the first period of the experiment there was no difference in population growth rate between fragmented and continuous populations, although litter size was significantly higher in the continuous populations. In the second period, we found higher population growth rates in populations that had experienced habitat destruction. By applying the transition matrix model to empirical estimates of demographic parameters, we demonstrate that the difference in population growth rate in the second period of the experiment was the result of a nonsignificant difference in adult survival. Movements out of the habitat patches were significantly lower in populations that had experienced habitat destruction. We conclude that predator-caused mortality of animals moving out of the habitat patches was the main determinant of demographic variation in this system. Received: January 31, 2002 / Accepted: March 25, 2003     

Synergistic effects of an extreme weather event and habitat fragmentation on a specialised insect herbivore

Habitat fragmentation is considered to be one of the main causes of population decline and species extinction worldwide. Furthermore, habitat fragmentation can decrease the ability of populations to resist and to recover from environmental disturbances such as extreme weather events, which are expected to occur at an increasing rate as a result of climate change. In this study, we investigated how calcareous grassland fragmentation affected the impact of the climatically extreme summer of 2003 on egg deposition rates, population size variation and survival of the blue butterfly Cupido minimus, a specialist herbivore of Anthyllis vulneraria. Immediately after the 2003 summer heat wave, populations of the host plant declined in size; this was paralleled with decreases in population size of the herbivore and altered egg deposition rates. In 2006 at the end of the monitoring period, however, most A. vulneraria populations had recovered and only one population went extinct. In contrast, several butterfly populations had gone extinct between 2003 and 2006. Extinction probability was significantly related to initial population size, with small populations having a higher risk of extinction than large populations. These results support the prediction that species of higher trophic levels are more susceptible to extinction due to habitat fragmentation and severe disturbances.     

Population size,habitat fragmentation,and the nature of adaptive variation in a stream fish

Whether and how habitat fragmentation and population size jointly affect adaptive genetic variation and adaptive population differentiation are largely unexplored. Owing to pronounced genetic drift, small, fragmented populations are thought to exhibit reduced adaptive genetic variation relative to large populations. Yet fragmentation is known to increase variability within and among habitats as population size decreases. Such variability might instead favour the maintenance of adaptive polymorphisms and/or generate more variability in adaptive differentiation at smaller population size. We investigated these alternative hypotheses by analysing coding-gene, single-nucleotide polymorphisms associated with different biological functions in fragmented brook trout populations of variable sizes. Putative adaptive differentiation was greater between small and large populations or among small populations than among large populations. These trends were stronger for genetic population size measures than demographic ones and were present despite pronounced drift in small populations. Our results suggest that fragmentation affects natural selection and that the changes elicited in the adaptive genetic composition and differentiation of fragmented populations vary with population size. By generating more variable evolutionary responses, the alteration of selective pressures during habitat fragmentation may affect future population persistence independently of, and perhaps long before, the effects of demographic and genetic stochasticity are manifest.     

Effects of habitat fragmentation and road density on the distribution pattern of the moor frog Rana arvalis

1. The effects of habitat fragmentation on the distribution pattern of the moor frog Rana arvalis were investigated. Also, the possible isolation effects of the road network were taken into account.
2. Indications were found that habitat fragmentation partly explains the distribution pattern of the moor frog. The statistical models showed a positive effect of pond size (or marsh area) and a negative effect of road density on the probability of occupation of a moorland pond.
3. Because of the strong correlation between habitat quality variables and isolation variables, no unambiguous effects of isolation, described as the amount of suitable terrestrial habitat (moorland) in the surroundings of a moorland pond in a radius of 100–2000 m, could be demonstrated.
4. Spatial differences in road density can play a role in the selection of optimal locations for nature protection areas. The regression model used in this study predicts a reduced occupation probability in 55% of the study area. In the part of the study area adjacent to a motorway, occupation probability is lowered to less than 30%.
5. European studies of habitat fragmentation on amphibian species revealed a mean distance between occupied ponds of <1 km in all studies. This could be a general rule of thumb for persistent amphibian populations. Effects of pond size on the probability of occupation were more variable.
6. When discussing the effects of habitat fragmentation on amphibians and other ground dwelling species, the negative effects of roads are often underestimated.