Composition and distribution of ground-dwelling beetles among oak fragments and surrounding pine plantations in a temperate forest of North China

In this study, we compared ground-dwelling beetle assemblages （Coleoptera） from a range of different oak fragments and surrounding conifer plantations to evaluate effects of forest size and surrounding matrix habitat in a temperate forest of north China. During 2000, beetles were sampled via pitfall traps within two large oak fragments （ca. 2.0-4.0 ha）, two small oak fragments （ca. 0.2-0.4 ha） and two surrounding matrices dom- inated by pine plantations （〉4 ha） in two sites of different aspects. Overall, no significantly negative effects from forest patch size and the surrounding matrix habitat were detected in total species number and abundance of ground-dwelling beetles. However, compared with small oak patches or pine plantations, more species were associated with an affinity for at least one large oak patch of the two aspects. Multivariate regression trees showed that the habitat type better determined the beetle assemblage structure than patch size and aspect, indicating a strong impact of the surrounding matrix. Linear mixed models indicated that species richness and abundance of all ground-dwelling beetles or beetle families showed different responses to the selected environmental variables. Our results suggest that more disturbed sites are significantly poorer in oak forest specialists, which are usually more abundant in large oak fragments and decrease in abundance or disappear in small fragments and surrounding matrix habitats. Thus, it is necessary to preserve a minimum size of forest patch to create conditions characteristic for forest interior, rather than the more difficult task of increasing habitat connectivity.     

Patch size of forest openings and arthropod populations

Summary Five sizes of canopy openings (0.016 ha to 10 ha) were established in the Southern Appalachian Mountains in early 1982 to examine the initial patterns of plant and arthropod establishment across a size range of forest disturbances. Vegetation standing crop after the first growing season was considerably higher in large than small openings in apparent response to greater resource release (e.g., sunlight) in larger openings. Woody stump and root sprouts were the dominant mode of revegetation in each patch size. Forest dominants such as Quercus rubra, Q. prinus and Carya spp. were less important as sprouters in openings than several minor forest components (e.g., Robinia pseudo-acacia, Acer rubrum, Halesia carolina and Cornus florida). Arthropod abundance and community composition varied across the size range of forest openings. Arthropods from the surrounding forest readily utilized the smallest canopy openings (0.016 ha). All feeding guilds were well represented in these small openings and herbivore biomass and load (mg of herbivores/g of foliage) were much higher than in larger patches. In contrast, arthropod abundance and species richness were significantly lower in mid-size than smaller patches. The relatively sparse cover and high sunlight in mid-size openings may have promoted surface heat buildups or soil surface/litter moisture deficits which restricted arthropod entry from the surrounding forest. Arthropod abundance and species richness were higher in large than mid-size patches. The greater vegetation cover in larger openings may have minimized the deleterious effects on arthropod populations. However, the absence of population increases among these arthropod species maintained herbivore loads at very low levels in large patches. Our results suggest that arthropod abundance and diversity in sprout-dominated forest openings are highly dependent on the extent of environmental differences between patch and surrounding forest.     

Avian responses to forest fragmentation during the breeding and non-breeding seasons

Forest fragmentation represents a threat to several bird species worldwide. Several factors can change across seasons (e.g. bird perception of the landscape, weather conditions, biotic interactions), which can modify the response of bird populations to forest fragmentation. However, most studies have been conducted only during the breeding season. Here we assessed the relationship between forest fragmentation (patch area and patch isolation) with population abundances of resident species during both the breeding and the non-breeding seasons. Bird population abundances (all species in the community, subsets of forest and habitat generalist species and for individual species) were estimated across a gradient of area-isolation in a semi-arid forest in Cordoba, Argentina. Population abundance of the overall avian community and of the subset of forest species declined with patch area reduction independently of the season. By contrast, the subset of habitat generalist species was not affected by patch area reduction or by the increase in patch isolation, either during the breeding or during the non-breeding season. When the analyses were carried out for individual species, we found four forest species and one habitat generalist species whose responses (the relationship between population abundance and patch area or with isolation) were different between breeding and non-breeding seasons. The negative effects of forest fragmentation were found mainly during the breeding season. Our results suggest that reduction of patch area may lead to a reduction of more than 65% of the population abundance of forest bird species, during both the breeding and the non-breeding season. Therefore, there is an urgent need to conserve large forest patches within the region as irreplaceable elements for the conservation of populations of several species.     

Population density and area: the role of between- and within-patch processes

The relationship between population density and the size of host plant patches was investigated for the red milkweed beetle Tetraopestetraophthalmus inhabiting unmanipulated patches of Asclepias syriaca. The resource concentration hypothesis proposes that density-area patterns, specifically that of increasing herbivore density with patch size, are primarily a function of movement between host plant patches. This research investigated the degree to which movement accounted for density-area patterns. Poisson regression analysis of beetle abundance versus milkweed patch size revealed that beetle density tended to increase with patch size. The pattern of density and patch size resulted from local reproduction and residence time. The density of emerging beetles tended to increase with patch size while emigration rates were unrelated to patch size. Immigration rates were constant with patch size for male beetles, and decreased with patch size for female beetles. Net flux of beetles (immigration – emigration) did not vary with patch size for male beetles and decreased with patch size for female beetles. Comparisons are made between this system and previously studied systems where movement plays a significant role in forming density area patterns. Additionally, several hypotheses are presented which may account for greater in situ recruitment and residence time in large patches. Received: 23 February 1996 / Accepted: 8 January 1997     

Importance of patch scale vs landscape scale on selected forest birds

The management and protection of natural areas have primarily occurred in isolation from surrounding land management. The structure of surrounding land cover, however, may be important to the abundance and reproductive success of birds within a habitat patch. We investigated the relative importance of forest patch area, within patch habitat and surrounding landscape forest cover on the abundance of three Neotropical migrant bird species thought to be area-sensitive (ovenbird [ Seiurus aurocapillus ], wood thrush [ Hylocichla mustelina ] and red-eyed vireo [ Vireo olivaceus ]), and on pairing success of the ovenbird. We selected 31 isolated forest patches of differing sizes, and three 80-ha plots in continuous forest each centered within non-overlapping 200-ha landscapes, such that patch area and landscape forest cover were uncorrelated among landscapes. Each study plot was surveyed to estimate abundances of territorial males and ovenbird pairing success. Landscape forest cover ( p <0.05) explained the most variation in ovenbird abundance, while percent deciduous forest cover within patches ( p <0.05) and patch size ( p <0.05) explained the most variation in red-eyed vireo and wood thrush abundance, respectively. Patch size was a significant ( p <0.05) predictor of abundance for all three study species; however, density for all species decreased significantly ( p <0.05) with patch size. Ovenbird pairing success was higher in continuous forest plots than in forest patches ( p =0.018). This study's findings suggest that the relative importance of within patch characteristics, patch size and landscape forest cover varies for different bird species, and that conservation efforts would benefit from the inclusion of all three factors.     

Combined effects of patch size and plant nutritional quality on local densities of insect herbivores

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.     

Tree leaf litter composition drives temporal variation in aquatic beetle colonization and assemblage structure in lentic systems

Tree leaf litter inputs to freshwater systems are a major resource and primary drivers of ecosystem processes and structure. Spatial variation in tree species distributions and forest composition control litter inputs across landscapes, but inputs to individual lentic habitat patches are determined by adjacent plant communities. In small, ephemeral, fishless ponds, resource quality and abundance can be the most important factor affecting habitat selection preferences of colonizing animals. We used a landscape of experimental mesocosms to assess how natural populations of aquatic beetles respond over time to variation in tree leaf litter composition (pine or hardwood). Patches with faster-decomposing hardwood leaf litter were initially colonized at higher rates than slower-decomposing pine pools by most species of Hydrophilidae, but this pattern reversed later in the experiment with higher colonization of pine pools by hydrophilids. Colonization did not differ between pine and hardwood for dytiscids and the small hydrophilid Paracymus, but there were distinct beetle assemblages between pine and hardwood patches both early and late in the experiment. Our data support the importance of patch quality and habitat selection as determinants of species abundances, richness, and community structure in freshwater aquatic systems, not only when new habitat patches are formed and initial conditions set, but as patches change due to interactions of processes such as decomposition with time.     

Importance of habitat patch size for occupancy and density of aspen-associated saproxylic beetles

In Fennoscandian boreal forests, aspen (Populus tremula) is one of the most important tree species for biodiversity. In this study we explore how occupancy and density of beetles associated with dead aspen are related to habitat patch size and connectedness in a 45,000 ha boreal managed forest landscape in central Sweden. Patch size was estimated as amount of breeding substrate and connectedness as crown cover of living aspen in the surrounding landscape. The beetles were sampled by sieving of bark or by inspection of species-characteristic galleries in 56 patches with dead aspen. Six of nine aspen-associated species (Xylotrechus rusticus, Ptilinus fuscus, Mycetophagus fulvicollis, Cyphaea curtula, Homalota plana and Endomychus coccineus) showed a positive significant relationship between habitat patch size and occupancy. For all these species, except C. curtula, there was also a significant positive relationship between patch size and density. Connectedness was not retained as a significant variable in the analyses. Species not defined as aspen-associated constituted a significantly larger proportion of the total density of individuals of saproxylic beetles in smaller habitat patches than in larger patches. Richness of aspen-associated species was positively related to habitat patch size. Efforts in the managed forest should be directed towards preserving and creating larger patches of living and dead aspen trees and increasing the amount of aspen at the landscape level.     

Host plant growth form and diversity: Effects on abundance and feeding preference of a specialist herbivore,Acalymma vittata (Coleoptera: Chrysomelidae)

Summary Abundances of the specialist herbivore, Acalymma vittata (Fab.) (Coleoptera: Chrysomelidae), were assessed in small experimental plots with three levels of plant diversity (cucumber monoculture, cucumber/corn, and cucumber/tomato) and two levels of host plant growth form (horizontal on the ground and vertical, staked up or growing up other plant species). Host plant growth form more strongly affected beetle abundances than did plant diversity; greater numbers were found on vertically growing than on horizontally growing cucumber plants. The combination of cucumber monoculture and vertical growth form supported significantly greater herbivore abundances than did any other type of plot, emphasizing a strong interaction between diversity and growth form. Beetles were not more common in monocultures with horizontal growth forms than in mixed species plots, and beetles did not respond differently to plots with corn and plots with tomatoes.Feeding experiments demonstrated that the plant diversity under which a host plant is grown strongly influenced herbivore feeding preference. Beetles given a choice of cucumber leaves grown in monoculture and in plots with tomatoes exhibited individual differences in their food selection behavior, however, a significantly greater number of beetles preferred monoculture leaves. Those individuals preferring monoculture leaves and those individuals preferring leaves from plots with tomatoes did not differ in either absolute or relative amounts of feeding damage per leaf.Neither plant size nor the date on which plots were colonized by beetles explained the differences in herbivore abundance. It is suggested that differences in movement patterns and plant quality contributed to the greater numbers of beetles on plants growing vertically in monocultures.     

Resource partitioning by three species of hemipteran herbivores on the basis of host plant density

Summary Three hemipteran herbivores partition a common resource plant, Senecio smallii, on the basis of bud density and plant patch size. The monophagous herbivore, Neacoryphus bicrucis, is most abundant in larger patches where bud density is greatest. The oligophagous herbivore, Harmostes reflexulus, is most abundant in small patches of high bud density. The polyphagous species, Lygus pratensis, is most abundant in small patches where abundance is independent of bud density. The aggressive nature of the monophagous herbivore appears to mediate the responses of the other species.     

The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions

Diet selection by mammalian herbivores is often influenced by plant community composition, and numerous studies have focused on the relationships between herbivore foraging decisions and food/plant species abundance. However, few have examined the role of neighbour palatability in affecting foraging of a target plant by large mammalian herbivores. We used a large-scale field dataset on diet selection by red deer Cervus elaphus in Fiordland National Park, New Zealand to: (1) estimate the palatability of native forest plant species to introduced deer from observed patterns of browse damage; and (2) examine whether intraspecific variation in browsing of plants can be related to variation in the local abundance of alternative forage species. Overall, 21 of the 53 forest species in our dataset were never browsed by deer. At a community level, plants were more likely to be browsed if they were in a patch of vegetation of high forage quality, containing high abundances of highly palatable species and/or low abundances of less-palatable species. Our findings suggest that deer make foraging decisions at both a coarse-grain level, selecting vegetation patches within a landscape based on the overall patch quality, and at a fine-grain level by choosing among individual plants of different species.     

A hierarchical multi‐scale analysis of the spatial relationship between parasitism and host density in urban habitats

Studies on spatial density dependence in parasitism have paid scarce attention to how changes in host density at different hierarchical scales could influence parasitism in an herbivore at a particular scale. Here, we evaluated if rates of parasitism per leaf (by the whole parasitic complex and by dominant species) of the specialist leaf miner Liriomyza commelinae (Diptera: Agromyzidae) respond to variations in host density at the leaf, plant patch and site levels in an urban setting. We used multi‐level Bayesian models that incorporate the spatial hierarchy occurring in this system, as well as habitat factors previously found to have an effect on the L. commelinae parasitoid community in an urban context (patch size, patch isolation and urbanization level). According to the fitted model, overall parasitism rates decreased with increasing number of mines per leaf, being independent of host‐density variations at patch and site level. Patch structure was found to have a strong effect on parasitism rates per leaf. The analysis of parasitism by parasitoid species separately showed consistent results with the response at community level. These results suggest that parasitism of the parasitoid community here studied would be sensitive to hierarchical cues related to the host at the leaf level and to the host habitat at the patch level.     

Spatial and temporal dynamics of habitat selection across canopy gradients generates patterns of species richness and composition in aquatic beetles

Abstract 1. Colonisation is a critical ecological process influencing both population and community level dynamics by connecting spatially discrete habitat patches. How communities respond to both natural and anthropogenic disturbances, furthermore, requires a basic understanding of how any environmental change modifies colonisation rates. For example, disturbance‐induced shifts in the quantity of forest cover surrounding aquatic habitats have been associated with the distribution and abundance of numerous aquatic taxa. However, the mechanisms generating these broad and repeatable field patterns are unclear. 2. Such patterns of diversity could result from differential spatial mortality post colonisation, or from colonisation alone if species select sites non‐randomly along canopy coverage gradients. We examined the colonisation/oviposition dynamics of aquatic beetles in experimental ponds placed under both open and closed forest canopies. 3. Canopy coverage imposed a substantial behavioural filter on the colonisation and reproduction of aquatic beetles representing multiple trophic levels, and resulted in significantly higher abundance, richness, and oviposition activity in open canopy ponds. These patterns strengthened overtime; although early in the experiment, the most abundant beetle had similar abundance in open and closed ponds. However, its abundance subsequently declined and then most other species heavily colonised open canopy ponds. 4. The primary response of many aquatic species to disturbances that generate canopy coverage gradients surrounding aquatic ecosystems is behavioural. The magnitude of the colonisation responses reported here rivals, if not exceeds, those produced by predators, suggesting that aquatic landscapes are behaviourally assessed and partitioned across multiple environmental gradients. The community level structure produced solely by selective colonisation, is predicted to strongly modify how patch area and isolation affect colonisation rates and the degree to which communities are linked by the flux of individuals and species.     

Landscape context outweighs local habitat quality in its effects on herbivore dispersal and distribution

Past studies with spatially structured herbivore populations have emphasized the primacy of intrinsic factors (e.g., patch quality), patch geometry (e.g., patch size and isolation), and more recently landscape context (e.g., matrix composition) in affecting local population abundance and dispersal rate. However, few studies have examined the relative importance of each factor, or how they might interact to affect herbivore abundance or dispersal. Here, we performed a factorial field experiment to examine the independent and interactive effects of patch quality (plant biomass, leaf protein, leaf phenolics) and matrix composition [mudflat or non-host grass (Bromus inermis)] on planthopper (Prokelisia crocea) emigration from host-plant patches (prairie cordgrass, Spartina pectinata). In addition, a field survey was conducted to examine the relative importance of patch quality, geography, and matrix composition on planthopper occupancy and density. In the experiment, we found that rates of emigration from low and intermediate quality patches were, on average, 21% percent higher for patches embedded in brome than mudflat. In contrast, the emigration rate was unaffected by matrix composition in nutrient-rich patches. Within matrix types, plant quality had little effect on emigration. In the survey, planthopper density and the patch occupancy rate of planthoppers increased nonadditively with increasing patch size and the percentage of the surrounding matrix composed of mudflat. This study suggests that landscape-level factors, such as the matrix, may be more important than factors intrinsic to the patches.     

垫状植物囊种草对群落物种多样性的影响

在甘肃盐池湾国家级自然保护区内海拔4137 m处,选择典型的囊种草垫状植被设置研究样地,研究了垫状植物囊种草对群落物种组成和群落物种多样性的影响,并且定量的研究了囊种草对群落物种丰富度的影响能力和维持潜力。研究结果表明:囊种草为群落中增加了新的植物种类,并且提高了部分生境一般种的多度;囊种草的出现提高了群落物种密度和物种丰富度,进而提高了群落物种多样性;囊种草斑块的增加将会引起景观水平物种丰富度的增加,表明囊种草具有为群落中引入新的植物种类进而提高群落物种丰富度的能力;在景观水平,囊种草所创造的生境多样性则成为一种保障,可以维持景观中物种丰富度从而降低物种损失的风险,表明囊种草具有较高的群落物种丰富度维持潜力。     

Trait-specific response of ground beetles (Coleoptera: Carabidae) to forest fragmentation in the temperate region in Korea

The objective of this study was to examine the trait-specific response of ground beetles in terms of abundance, species richness, and composition for habitat fragmentation in temperate forests. In addition, we examined how different ecological groups and species of ground beetles responded to environmental variables. During middle May to early November in 2013, ground beetles were sampled using pitfall traps in 27 sites (18 patches and 9 continuous forests) in central Korea. A total of 51 species were identified from 17,845 ground beetles in the 27 study sites. Continuous forests had generally higher abundance and species richness of ground beetles than forest patches. Canonical correspondence analysis for ground beetles was significant, and the proportion of variance explained by environmental variables, such as patch size, elevation, organic matter (C), and soil moisture and pH, was 43.61%. In addition, species composition of ground beetles in continuous forests was grouped distinctively away from those in medium and small-sized patches. In conclusion, both small (1.1–9.6 ha) and medium patches (12.8–51.2 ha) were failed to preserve similar ground beetle biodiversity compared to continuous forests. However, our study revealed that medium-sized forest patches clearly had higher conservation value for forest specialist ground beetles than small-sized forest patches irrespective of forest type.     

Landscape mosaic of Araucaria forest and forest monocultures influencing understorey spider assemblages in southern Brazil

Abstract This study investigates how abundance, diversity and composition of understorey spiders were influenced by four different forest habitats in a southern Brazilian Araucaria forest. The study area encompasses a landscape mosaic comprised of Araucaria forest, Araucaria plantation, Pinus plantation, and Eucalyptus plantation. Understorey spiders were collected by beating the vegetation inside three patches of each forest habitat. To assess possible predictors of spider assemblage structure, several patch features were analysed: potential prey abundances, estimation of vegetation cover, diversity index of vegetation types, patch ages, patch areas, and geographical distance between patches. To assess the influence of high‐level taxa approaches on spider assemblage patterns, analyses were carried out individually for family, genera and species levels. Additionally, Mantel tests were carried out in underlying similarity matrices between each taxon. Significant differences in spider abundances among forest habitats were found. Pinus plantations showed the highest abundance of spiders and Eucalyptus plantations showed the lowest abundance. Spider abundance was significantly influenced by patch ages, geographical distance and vegetation cover. Expected numbers of families, genera and species did not vary among forest habitats. Spider composition of two Eucalyptus patches differed from the other forest patches, probably due to their low vegetation cover and isolation. Genera composition was the best correlate of species composition, showing that a higher‐level surrogate can be an alternative to the species approach. The understorey spider diversity in this managed area could be maintained when suitable habitat structures are provided, thus ensuring the connectivity between different habitat types. Further studies should focus on individual species responses to the conversion of native forest to monocultures.     

Distribution and colonisation ability of three parasitoids and their herbivorous host in a fragmented landscape

Habitat fragmentation can disrupt communities of interacting species even if only some of the species are directly affected by fragmentation. For instance, if parasitoids disperse less well than their herbivorous hosts, habitat fragmentation may lead to higher herbivory in isolated plant patches due to the absence of the third trophic level. Community-level studies suggest that parasitoids tend to have limited dispersal abilities, on the order of tens of metres, much smaller than that of their hosts, while species-oriented studies document dispersal by parasitoids on the scale of kilometres. In this study the distribution patterns of three parasitoid species with different life histories and their moth host, Hadena bicruris, a specialist herbivore of Silene latifolia, were compared in a large-scale network of natural fragmented plant patches along the rivers Rhine and Waal in the Netherlands. We examined how patch size and isolation affect the presence of each species. Additionally, experimental plots were used to study the colonisation abilities of the species at different distances from source populations.In the natural plant patches the presence of the herbivore and two of the parasitoids, the gregarious specialist Microplitis tristis and the gregarious generalist Bracon variator were not affected by patch isolation at the scale of the study, while the solitary specialist Eurylabus tristis was. In contrast to the herbivore, the presence of all parasitoid species declined with plant patch size. The colonisation experiment confirmed that the herbivore and M. tristis are good dispersers, able to travel at least 2 km within a season. B. variator showed intermediate colonisation ability and E. tristis showed very limited colonisation ability at this spatial scale. Characteristics of parasitoid species that may contribute to differences in their dispersal abilities are discussed.     

Distribution and growth of benthic macroinvertebrates among different patch types of the littoral zones of two arctic lakes

1. To evaluate the effect of habitat patch heterogeneity on abundance and growth of macroinvertebrates in arctic lakes, macroinvertebrate abundance, individual biomass, and potential food resources were studied in three patch types in two arctic lakes on the Alaskan North Slope near the Toolik Lake Field Station. An experiment was conducted to determine which sediment patch type supported higher growth rates for Chironomus sp., a commonly occurring macroinvertebrate. 2. Potential organic matter (OM) resources were significantly higher in both rock and macrophyte patches than in open‐mud patches. Total macroinvertebrate densities in both lakes were highest in rock patches, intermediate in macrophytes and lowest in open‐mud. The open‐mud patches also had lower species richness compared with other patch types. Additionally, individual biomass for one clam species and two chironomid species was significantly greater in rock patches than in open‐mud. 3. In a laboratory experiment, Chironomus showed two to three times greater mass increase in sediments from macrophyte and rock patches than from open‐mud patches. Rock and macrophyte experimental sediments had at least 1.5 × the percentage OM as open‐mud sediments. 4. Chlorophyll a appeared to be the best predictor for invertebrate abundances across all patch types measured, whereas OM content appeared to be the variable most closely associated with Chironomus growth. 5. Our results combined with previous studies show that the relationships between macroinvertebrate community structure, individual growth, and habitat heterogeneity are complex, reflecting the interaction of multiple resources, and biotic interactions, such as the presence or absence of a selective vertebrate predator (lake trout, Salvelinus namaycush).