Patterns of intra- and interspecific association in leaf-mining insects on three oak host species

Abstract. 1. We determined mortality and distributional patterns of leaf miners on three oak host species (Quercus falcata, Q.nigra and Q.hemisphaerica) in northern Florida, U.S.A.
2. Patterns of intra- and interspecific occurrence within leaves, and mortality of five most abundant leaf miner species were analysed as a test of competition.
3. Miners co-occurred on leaves more often that expected by chance (P<0.05) in six of ten possible species combinations and log-linear model analysis showed no negative higher-order interactions.
4. All five miner species had highly clumped distributions between leaves (P<0.01).
5. Leaf miner survival was less than expected for four of five species when co-occurring on leaves with conspecifics than when mining with heterospecifics or alone (P<0.05).
6. We conclude that interspecific competition is unapparent within this leaf miner guild and that intraspecific competition occurs in four of the five major leaf miner species. We discuss leaf miner selection of common leaves, perhaps based on chemical/physical leaf characters, as a cause of intra- and interspecific aggregation.     

Human land use, agriculture, pesticides and losses of imperiled species

Aim  Anthropogenic habitat loss is usually cited as the most important cause of recent species' extinctions. We ask whether species losses are in fact more closely related to habitat loss than to any other aspect of human activity such as use of agricultural pesticides, or human population density (which reflects urbanization).
Location  Canada.
Methods  We statistically compared areas in Canada where imperiled species currently occur, versus areas where they have been lost. Using multiple regressions, we relate the numbers of species that had suffered range reductions in an ecoregion to variables that represent present habitat loss, pesticide use and human population density.
Results  We find high losses of imperiled species in regions with high proportions of agricultural land cover. However, losses of imperiled species are significantly more strongly related to the proportion of the region treated with agricultural pesticides. The relationship between species losses and area treated with pesticides remains significant after controlling for area in agriculture.
Main conclusions  Our results are consistent with the hypothesis that agricultural pesticide use, or something strongly collinear with it (perhaps intensive agriculture more generally), has contributed significantly to the decline of imperiled species in Canada. Habitat conversion per se may be a less important cause of species declines than how that converted habitat is used.     

Ecological correlates of geographical range occupancy in North American birds

Aim  The degree to which a species is predictably encountered within its range varies tremendously across species. Understanding why some species occur less frequently within their range than others has important consequences for conservation and for analyses of ecological patterns based on range maps. We examined whether patterns in geographical range occupancy can be explained by species-level traits.
Location  North America.
Methods  We used survey data from 1993 to 2002 from the North American Breeding Bird Survey along with digital range maps produced by NatureServe to calculate range occupancy for 298 species of terrestrial birds. We tested whether species traits explained variation in range occupancy values using linear regression techniques.
Results  We found three species traits that together explained more than half of the variation in range occupancy. Population density and niche breadth were positively correlated with occupancy, while niche position was negatively correlated with occupancy.
Main conclusions  Our results suggest that high range occupancy will occur in species that are common at sites on which they occur, that tolerate a relatively wide range of ecological conditions and that tend to have ranges centred on areas with common environmental conditions. Furthermore, it appears that niche-based characteristics may explain patterns of distribution and abundance from local habitats up to the scale of geographical ranges.     

Specificity and host predictability: a comparative analysis among monogenean parasites of fish

1. This article compares generalist (parasite species found on two or more host species) and specialist (found on only one host species) monogenean parasite species of fish. The reduction of the host range – that is an increase in host specificity – may correspond with a better adaptation of the parasite to a more predictable host environment. A more predictable environment may allow the parasite species to develop specific adaptations.
2. We assume that the more predictable host environment can be evaluated by host body size, since numerous life-traits, such as longevity, are positively correlated with size.
3. We found that specialist parasites parasitize larger hosts species than generalist parasites. We also found a good relationship between host body size and parasite body size for specialist parasite species.
4. An adaptation to the mechanical problems encountered in the host's gill chamber may lead to an increase in parasite body size. The infection of a larger part of the host population in order to decrease the chances of local extinction due to fluctuations of host abundance may be another adaptive mechanism.
5. We found a negative correlation between parasite body size and prevalence for generalist parasite species. This relationship disappeared when using the comparative method controlling for phylogeny, which proved that it was a phylogenetic effect.     

Metapopulations of moths on islands: a test of two contrasting models

1. We describe a generalized mainland-island metapopulation model which includes migration among the island populations. We test model predictions with quantitative data on more than 200 species of moths in two contrasting networks of small islands. The data include a direct measure of migration rate, based on trapping of moths on rocky skerries with no local populations of the vast majority of species.
2. We predicted that moths which are strong fliers but uncommon on the islands have a higher incidence on scattered islands than on islands in a group, because the latter 'compete' for immigrants from the mainland. In contrast, we predicted that weakly flying species with potentially large local populations on the islands occur more frequently on islands in a group due to enhanced colonization rate.
3. Both predicted patterns were observed. Island occupancy increased significantly with the number of individuals caught on the rocky skerries, which is our measure of migration rate from the mainland, supporting the basic assumption that the species occur on the islands in a balance between colonizations and extinctions.
4. These results demonstrate that the moth metapopulations on islands represent a mixture of Levins's and mainland-island metapopulations, and that the mixture is different for different species in the same landscape.     

MATING FREQUENCY AND FECUNDITY IN INSECTS

1. The paper summarizes the published evidence on the relation between mating frequency and fecundity in insects. There is experimental evidence of varying quality for 63 species and non-experimental evidence for about 60.
2. Repeated mating may be universally necessary for full fecundity and fertility in female insects (in species in which the females normally mate more than once).
3. The evidence is remarkably poor. We need more properly designed experiments (and not just observations of natural variation), with sufficient sample sizes and statistics, and measurements of the fecundities and fertilities of singly and multiply mated females, when the multiple matings are separated by many days or weeks. Most of the existing experiments of this sort are defective in some way.
4. In species with greater total fecundity and longevity, multiple mating may be more likely to enhance fertility than in species with small fecundity and short life span.
5. Females in naturally monandrous species do not show increased fecundity or fertility with repeated mating, whereas females of polyandrous species do.
6. There is no obvious connexion between paternal investment, in so far as we know about it, and the increase of fecundity by repeated mating.
7. There is a small tendency for females to breed more quickly and be shorter lived if they mate repeatedly.     

The impact of shrub encroachment on savanna bird diversity from local to regional scale

Aim  Evidence is accumulating of a general increase in woody cover of many savanna regions of the world. Little is known about the consequences of this widespread and fundamental ecosystem structural shift on biodiversity.
Location  South Africa.
Methods  We assessed the potential response of bird species to shrub encroachment in a South African savanna by censusing bird species in five habitats along a gradient of increasing shrub cover, from grassland/open woodland to shrubland dominated by various shrub species. We also explored historical bird species population trends across southern Africa during the second half of the 20th century to determine if any quantifiable shifts had occurred that support an ongoing impact of shrub encroachment at the regional scale.
Results  At the local scale, species richness peaked at intermediate levels of shrub cover. Bird species composition showed high turnover along the gradient, suggesting that widespread shrub encroachment is likely to lead to the loss of certain species with a concomitant decline in bird species richness at the landscape scale. Finally, savanna bird species responded to changes in vegetation structure rather than vegetation species composition: bird assemblages were very similar in shrublands dominated by Acacia mellifera and those dominated by Tarchonanthus camphoratus .
Main conclusions  Shrub encroachment might have a bigger impact on bird diversity in grassland than in open woodland, regardless of the shrub species. Species recorded in our study area were associated with historical population changes at the scale of southern Africa suggesting that shrub encroachment could be one of the main drivers of bird population dynamics in southern African savannas. If current trends continue, the persistence of several southern African bird species associated with open savanna might be jeopardized regionally.     

Geographical patterns of species turnover in aquatic plant communities

1. A classic theory in biogeography predicts that high latitude communities are unstable. This may be because of decreased species richness or decreased environmental predictability and productivity towards the poles.
2. We studied latitudinal patterns in long-term community persistence of aquatic vascular plants in 112 Finnish lakes, situated within a 1000-km range from the northernmost to the southernmost lake.
3. Contrary to theoretical predictions, we found that the turnover rate of plant species in 45 years was inversely related to latitude. That is, plant communities in northern lakes were more persistent than communities in southern lakes. When we used multiple regression to find the best predictors of species turnover rate (TR), latitude was the only variable that was highly significantly related to species turnover rate. Area, species number, water transparency, pH and change in transparency did not notably explain the gradient observed.
4. The latitudinal trend was mainly because of lower species immigration rates at higher latitudes, whereas extinction rate did not so strongly decrease with increasing latitude. Immigrations and extinctions in the lakes were not in balance: the species numbers between the 1930s and 1980s increased more strongly in the southern than northern lakes.
5. We suggest that the inverse relationship between latitude and plant species TR in Finland is most probably caused by human influence on lakes, especially eutrophication and immigration of new species in southern latitudes. In addition, although species richness per lake did not decrease towards the north, the total species pool probably does, which means that in the north there are fewer species that can actually immigrate.     

Assessing conservation value of stream communities: a comparison of approaches based on centres of density and species richness

1. We used a recently developed Index of Centres of Density (ICD) to assess conservation value of thirty-nine sites in the upper Clinch River drainage of Virginia and Tennessee, U.S.A. The ICD incorporates information on the population density of each species at a site relative to sites throughout the region.
2. Value assessments based on the ICD were compared to those based on species richness. Species richness at sites was not related to ICD scores, but collective species lists from high-ranking sites were similar for both approaches. All sites with rare species had either a high ICD score or high species richness.
3. Four community types (defined by physiography and stream size) were better represented in sites with high-ranking ICDs than in sites with high-ranking species richness. Sites with high ICD scores were distributed uniformly throughout the region.
4. The ICD is a more powerful tool than species richness for assessing conservation value because the ICD identifies areas with regionally rare species, especially viable populations, or distinctive communities, all of which are key components of a region's biodiversity.     

Variation in bark thickness in a tropical forest community of Western Ghats in India

1. We examined the relation between bark thickness and girth in a large sample of trees from evergreen and semi-evergreen rain forest.
2. There is a significant tendency for bark thickness to increase with tree girth. Removing this tendency, we find a significant trend for species from more disturbed habitats to possess thicker bark.
3. Species from more disturbed habitats also have a greater propensity for secreting gums and resins.
4. Nine of the 29 species occurring in more than one habitat type and with a sample size of at least 11 individuals show a tendency for possession of a thicker bark by individuals in the more disturbed habitats.
5. We conclude that bark thickness and occurrence of gums and resins are physiognomic–structural attributes of value in characterizing tree communities created by different levels of disturbance.     

Salamander occupancy in headwater stream networks

1. Stream ecosystems exhibit a highly consistent dendritic geometry in which linear habitat units intersect to create a hierarchical network of connected branches.
2. Ecological and life history traits of species living in streams, such as the potential for overland movement, may interact with this architecture to shape patterns of occupancy and response to disturbance. Specifically, large-scale habitat alteration that fragments stream networks and reduces connectivity may reduce the probability a stream is occupied by sensitive species, such as stream salamanders.
3. We collected habitat occupancy data on four species of stream salamanders in first-order (i.e. headwater) streams in undeveloped and urbanised regions of the eastern U.S.A. We then used an information–theoretic approach to test alternative models of salamander occupancy based on a priori predictions of the effects of network configuration, region and salamander life history.
4. Across all four species, we found that streams connected to other first-order streams had higher occupancy than those flowing directly into larger streams and rivers. For three of the four species, occupancy was lower in the urbanised region than in the undeveloped region.
5. These results demonstrate that the spatial configuration of stream networks within protected areas affects the occurrences of stream salamander species. We strongly encourage preservation of network connections between first-order streams in conservation planning and management decisions that may affect stream species.     

Disentangling the effects of area, energy and habitat heterogeneity on boreal forest bird species richness in protected areas

Aim  One of the few general laws in ecology is that species richness is a positive function of area. However, it has been proposed that area would merely be a proxy for energy. Additionally, habitat heterogeneity has been found to be an important factor determining species richness. Yet the relative importance of those relationships is little known, and it is still unclear how they are brought about. We aimed to dissect which factors drive the species richness of boreal forest birds, and to identify the most probable mechanisms.
Location  Forested protected areas in Finland.
Methods  Using bird line census data collected in 104 protected areas, we ran simultaneous autoregressive models to explain the species richness of forest birds. We explored the value of forest area, tree volume, tree growth, mean degree days and habitat heterogeneity as explanatory variables and used the species richness within different species groups, based on the predictions of hypothesized mechanisms, as a response variable.
Results  Energy, rather than area or habitat heterogeneity, seems to be the main driver of species richness in boreal forest birds. More specifically, productive energy was a better predictor of total species richness than solar energy. Among the tested hypothetical mechanisms, the sampling hypothesis received strong support. After accounting for sampling, solar energy had an effect on species richness.
Main conclusions  As productive energy, such as tree volume, is associated with species richness, high-energy areas should be prioritized in forest conservation planning. Reductions in productive energy may first lead to the disappearance of the rarest species due to the random sampling process. Climate change may result in increased species richness due to increasing amount of productive and solar energy in forests. However, the range shifts of bird species may not be fast enough to keep up with the temperature increases.     

Latitudinal patterns of range size and species richness of New World woody plants

Aim  Relationships between range size and species richness are contentious, yet they are key to testing the various hypotheses that attempt to explain latitudinal diversity gradients. Our goal is to utilize the largest data set yet compiled for New World woody plant biogeography to describe and assess these relationships between species richness and range size.
Location  North and South America.
Methods  We estimated the latitudinal extent of 12,980 species of woody plants (trees, shrubs, lianas). From these estimates we quantified latitudinal patterns of species richness and range size. We compared our observations with expectations derived from two null models.
Results   Peak richness and the smallest- and largest-ranged species are generally found close to the equator. In contrast to prominent diversity hypotheses: (1) mean latitudinal extent of tropical species is greater than expected; (2) latitudinal extent appears to be decoupled from species richness across New World latitudes, with abrupt transitions across subtropical latitudes; and (3) mean latitudinal extents show equatorial and north temperate peaks and subtropical minima. Our results suggest that patterns of range size and richness appear to be influenced by three broadly overlapping biotic domains (biotic provinces) for New World woody plants.
Main conclusions  Hypotheses that assume a direct relationship between range size and species richness may explain richness patterns within these domains, but cannot explain gradients in richness across the New World.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Soil nitrogen and carbon heterogeneity in woodlands and grasslands: contrasts between temperate and tropical regions

Aim  Soil resource heterogeneity is linked to several ecological processes including invasion of woody species into grasslands. Studies from the temperate zone have demonstrated greater soil heterogeneity beneath woody vegetation than beneath grasslands. Woody species have a more widespread and coarser root system than herbaceous species, and may have a competitive advantage in relatively heterogeneous soils. We tested the global generality of greater soil heterogeneity beneath woody vegetation.
Location  Global.
Methods  We used data from published literature for soil nitrogen and carbon heterogeneity from paired woodland and grassland sites around the world.
Results  Woodland and grassland soil heterogeneities from paired observations were strongly correlated. There was, however, significant geographical variability in the relationship. Soils were more heterogeneous in woodlands than grasslands in temperate areas, but the opposite was true for tropical habitats. Grassland soils were more heterogeneous at lower than higher latitudes. Woodland soil heterogeneity did not vary with latitude.
Main conclusions  The previously described high soil heterogeneity in woody vegetation compared to grasslands holds only for temperate regions. Consequently, the relationship between soil resource heterogeneity and vegetation type is dependent on the study region. Macroecological studies should test the generality of relationships between soil and vegetation at the global scale.     

When does diversity fit null model predictions? Scale and range size mediate the mid‐domain effect

Aim  Recently, a flurry of studies have focused on the extent to which geographical patterns of diversity fit mid-domain effect (MDE) null models. While some studies find strong support for MDE null models, others find little. We test two hypotheses that might explain this variation among studies: small-ranged groups of species are less likely than large-ranged species to show mid-domain peaks in species richness, and mid-domain null model predictions are less robust for smaller spatial extents than for larger spatial extents.
Location  We analyse data sets from elevational, riverine, continental and other domains from around the world.
Methods  We use a combination of Spearman rank correlations and binomial tests to examine whether differences within and among studies and domains in the predictive power of MDE null models vary with spatial scale and range size.
Results  Small-ranged groups of species are less likely to fit mid-domain predictions than large-ranged groups of species. At large spatial extents, diversity patterns of taxonomic groups with large mean range sizes fit MDE null model predictions better than did diversity patterns of groups with small mean range sizes. MDE predictions were more explanatory at larger spatial extents than at smaller extents. Diversity patterns at smaller spatial extents fit MDE predictions poorly across all range sizes. Thus, MDE predictions should be expected to explain patterns of species richness when ranges and the scale of analysis are both large.
Main conclusions  Taken together, the support for these hypotheses offers a more sophisticated model of when MDE predictions should be expected to explain patterns of species richness, namely when ranges and the scale of analysis are both large. Thus the circumstances in which the MDE is important are finite and apparently predictable.     

Herbivory is related to taxonomic isolation, but not to invasiveness of tropical alien plants

Aim  The enemy release hypothesis is often invoked to explain why some alien plant species become invasive. Here, we investigated relationships between invasiveness, taxonomic isolation and leaf herbivory for tropical alien plant species introduced to a botanical garden in East Africa.
Location  Amani Botanical Garden, East Usambara mountains, northeast Tanzania.
Methods  We measured the proportion of leaves damaged, and the percentage leaf area damaged on individuals of 28 alien plant species. We extracted data on the presence/absence of native congeners and the number of native confamilial species from an inventory of the East Usambara flora. We also obtained data on planting effort for 26 species, from historical records. Linear and generalized linear models were used to analyse the relationships between invasiveness, herbivory and taxonomic isolation.
Results  Mean proportion of leaves damaged per species was significantly explained by taxonomic isolation; proportion of leaves damaged increased with the number of native confamilial species and was greater, on average, for species with native congeners than those without native congeners. The mean percentage of leaf area damaged per species could not be explained by any variables considered in this study. There was no relationship between the degree of herbivory or taxonomic isolation and alien plant species invasiveness, but more-invasive species did have a significantly greater planting effort than less-invasive species.
Conclusions  The role herbivores play in controlling alien plant invasions has been investigated relatively little in the tropics. In this study, although the amount of herbivory suffered by alien plants was related to taxonomic isolation, we found no evidence for leaf-feeding invertebrates having a significant role in invasion, suggesting that other factors may be responsible for differences in species success.     

Bird diversity and environmental gradients in Britain: a test of the species–energy hypothesis

1.  We tested the species diversity–energy hypothesis using the British bird fauna. This predicts that temperature patterns should match diversity patterns. We also tested the hypothesis that the mechanism operates directly through effects of temperature on thermoregulatory loads; this further predicts that seasonal changes in temperature cause matching changes in patterns of diversity, and that species' body mass is influential.
2.  We defined four assemblages using migration status (residents or visitors) and season (summer or winter distribution). Records of species' presence/absence in a total of 2362, 10 × 10-km, quadrats covering most of Britain were used, together with a wide selection of habitat, topographic and seasonal climatic data.
3.  We fitted a logistic regression model to each species' distribution using the environmental data. We then combined these individual species models mathematically to form a diversity model. Analysis of this composite model revealed that summer temperature was the factor most strongly associated with diversity.
4.  Although the species–energy hypothesis was supported, the direct mechanism, predicting an important role for body mass and matching seasonal patterns of change between diversity and temperature, was not supported.
5.  However, summer temperature is the best overall explanation for bird diversity patterns in Britain. It is a better predictor of winter diversity than winter temperature. Winter diversity is predicted more precisely from environmental factors than summer diversity.
6.  Climate change is likely to influence the diversity of different areas to different extents; for resident species, low diversity areas may respond more strongly as climate change progresses. For winter visitors, higher diversity areas may respond more strongly, while summer visitors are approximately neutral.     

Timing is everything? Phenological synchrony and population variability in leaf‐chewing herbivores of Quercus

Abstract.  1. Specialization on ephemeral resources (e.g. new leaves) should produce large annual variation in herbivore population size when the timing of availability of those resources is unpredictable. Despite considerable evidence for impacts of synchrony with budburst on survival of larval Lepidoptera, previous studies of adult Geometridae and Noctuidae found no correlations between insect phenology and population variability.
2. We surveyed larval Lepidoptera feeding on Quercus alba and Q. velutina in Missouri from 1993 to 2003 and examined population variability, measured as the coefficient of variation of population density (CV), in a subset of abundant species. We compared CV values among species whose larvae feed only in spring, early summer, mid-summer, late summer, or all season. We predicted that univoltine species whose larvae eclose and complete development in spring during leaf expansion would have higher variability than species feeding later in the season, having multiple generations, or having longer development times.
3. As predicted and consistent with hypotheses, spring-feeding species had CV values 32% higher than species feeding in summer months. Coefficients of variation were also 34% higher in leaf-rolling and mining guilds compared with free-feeders, suggesting that mobile species may compensate for asynchrony with budburst by dispersing to higher quality plants or plant parts. Multivoltine species, however, did not differ from univoltine species in population variability.
4. Our results suggest that asynchrony with plant phenology and factors that might exacerbate it, such as climate change, will have the largest impacts on the dynamics of spring-feeding Lepidoptera, particularly species with limited mobility.     

More species, fewer specialists: 100 years of changes in community composition in an island biogeographical study

Aim  We measured the changes in an island avifauna over more than 100 years (1898–2006), using community indices accounting for difference in expected species sensitivity to land-use and climate changes.
Location  Ouessant Island, France, Great Britain.
Methods  We assessed the temporal trend of the relative proportion of generalist species breeding on Ouessant island and whether high-temperature tolerant species have replaced less tolerant species over this time period. We further tested the relationship between the observed change in the avifauna composition, and long-term population species' trends measured independently in potential source regions of colonist species (France and Great Britain).
Results  During the whole study period, Ouessant island has experienced a strong increase in species richness (+41%), but a severe decline in specialist species. In contrast, we found no change in species composition in terms of their temperature-tolerance. The observed trend was highly correlated with species trends measured in the Great Britain.
Main conclusions  Our results revealed an ongoing biotic homogenization process towards more generalist species, coupled with a strong local increase in species richness. The observed trend was most likely driven by a strong habitat change in the island occurring during the period considered, favouring the colonization of generalist species. Our results show that an increase in species richness can be misinterpreted as a sign of conservation improvement and that assessing change in community composition using species-specific ecological traits provides more accurate insights for conservation planning purposes.