Are island and mainland biotas different? Richness and level of generalism in parasitoids of a microlepidopteran in Macaronesia

Island communities are exposed to several evolutionary and ecological processes that lead to changes in their diversity and structure compared to mainland biotas. These phenomena have been observed for various taxa but not for parasitoids, a key group in terms of community diversity and functioning. Here we use the parasitoid communities associated with the moth Acroclita subsequana (Lepidoptera: Tortricidae) in the Macaronesian region, to test whether species richness differs between islands and mainland, and whether island parasitoid faunas are biased towards generalist species. Host larvae were collected on several islands and adjacent mainland, carefully searched for ectoparasitoid larvae and dissected to recover any endoparasitoids. Parasitoids were classified as idiobionts, which usually have a wide host range (i.e. generalists), or koinobionts that are considered specialists. Mainland species richness was lower than expected by chance, with most of the species being koinobionts. On the other hand, island communities showed a greater proportion of idiobiont species. Overall parasitism rates were similar between islands and mainland, but islands had higher rates of parasitism by idiobionts than expected by chance, and mainland areas showed the highest koinobiont parasitism rates. These results suggest that island parasitoid communities are dominated by generalists, in comparison to mainland communities. Several hypotheses may explain this pattern: (1) generalist parasitoids might have better dispersal abilities; (2) they may be less constrained by ‘sequential dependencies’; and (3) island parasitoids probably have fewer competitors and/or predators, thus favouring the establishment of generalists. New studies including multiple hosts, other habitats, and/or more islands are necessary to identify which of these processes shape island parasitoid communities.     

Host diversity and latitude drive trematode diversity patterns in the European freshwater fauna

Aim We investigated the relationship between host and parasite diversity as well as latitudinal gradients in parasite diversity on a continental scale in European freshwater trematodes. Location European freshwaters. Methods We extracted distributional data for 564 freshwater trematodes across 25 biogeographical regions in Europe from the Limnofauna Europaea and used multiple regression analyses to test for correlations between the diversity of definitive (vertebrates) or first intermediate (gastropods) hosts and that of trematodes, and for latitudinal gradients in trematode diversity. In particular, we investigated patterns in beta diversity among latitudinal bands and between trematode species that parasitize host groups with low (autogenic) and high (allogenic) dispersal capacity. We also tested for a latitudinal gradient in the proportional representation of these two trematode groups within regional faunas. Results Latitude or first intermediate host richness had no effect on trematode richness, but definitive host richness was a strong predictor of trematode richness, among both allogenic and autogenic parasites. We found that beta diversity of trematode faunas within latitudinal bands decreased to the north, with similar values for allogenic and autogenic trematodes. Finally, we observed an increasing proportion of autogenic species toward the north of Europe. Main conclusions The richness of definitive hosts appears to be the driver of trematode diversity at a continental scale. The latitudinal gradient in beta diversity reflects patterns observed in free‐living species and probably results from recolonization in the aftermath of the ice ages. The similar beta‐diversity patterns of allogenic and autogenic trematodes and the increasing proportion of autogenic trematodes with increasing latitude are surprising. We suggest that the geographical scale of our analysis or confounding factors such as differences in habitat utilization and specialization may partly explain these patterns.     

Latitudinal gradients in the phenetic diversity of New World bat communities

Although the examination of latitudinal gradients of species richness is common, little attention has been devoted to other components of biodiversity such as phenetic diversity. Because the phenotype reflects aspects of an organism's environment, ecological relationships and evolutionary history, measures of phenetic diversity likely provide complimentary information to that of species richness, and may provide unique insights for understanding the mechanistic basis to patterns of biodiversity. Herein, we evaluate latitudinal gradients in the phenetic diversity of 32 New World bat communities. Seven morphological characters were used to estimate phenotypic variation among bat species within local communities. Principal components analysis decomposed this variation into axes of size and shape. Three measures of phenetic diversity were calculated separately for size and for shape axes. The range of species scores on a particular axis described the amount of phenetic variation encompassed by species in a community. The standard deviation of minimum spanning‐tree segment lengths described uniformity of species. Average nearest‐neighbor distances described local packing. We separately regressed these six measures on local species richness and latitude separately. Variation in species richness accounted for a significant amount of variation in each measure of phenetic diversity. Latitude also accounted for significant variation in phenetic diversity except for the standard deviation of minimum‐spanning tree segment lengths and the average nearest‐neighbor distance on the shape axis. More importantly, gradients in phenetic diversity were significantly different than would be expected as a consequence of latitudinal gradients in species richness. Nonetheless, when variation among communities regarding the richness and composition of their regional faunas was taken into consideration, differences between empirical and simulated gradients were nonsignificant. Thus, factors that determine the composition of regional faunas have a great impact on the phenetic diversity of communities and ultimately the latitudinal gradient in biodiversity.     

Patterns of functional diversity across an extensive environmental gradient: vertebrate consumers, hidden treatments and latitudinal trends

Over the last two decades, although much has been learned regarding the multifaceted nature of biodiversity, relatively little is known regarding spatial variation in constituents other than species richness. This is particularly true along extensive environmental gradients such as latitude. Herein, we describe latitudinal gradients in the functional diversity of New World bat communities. Bat species from each of 32 communities were assigned to one of seven functional groups. Latitudinal gradients existed for the richness, diversity and scaled‐dominance of functional groups. No significant patterns were observed for evenness of functional groups. Measures of functional diversity were different in magnitude and increased towards the equator at a faster rate than expected given the underlying spatial variation in species richness. Thus, latitudinal gradient in species richness alone do not cause the latitudinal gradient in functional diversity. When variation in species composition of the regional fauna of each community was incorporated into analyses, many differences between observed and simulated patterns of functional diversity were not significant. This suggests that those processes that determine the composition of regional faunas strongly influence the latitudinal gradient in functional diversity at the local level. Nonetheless, functional diversity was lower than expected across observed sites. Community‐wide responses to variation in the quantity and quality of resources at the local level probably contribute to differences in functional diversity at local and regional scales and enhance beta diversity.     

Species pool structure determines the level of generalism of island parasitoid faunas

Aim To examine whether island parasitoid faunas are biased towards generalists when compared with the mainland and their species pool, and to evaluate the effects of climate, island characteristics and regional factors on the relative proportions of idiobionts (i.e. generalists) and koinobionts (i.e. specialists) of two parasitic wasp families, Braconidae and Ichneumonidae. Location Seventy‐three archipelagos distributed world‐wide. Methods We used data on the distribution and biology obtained from a digital catalogue and several literature sources. We related level of generalism, measured as the ratio between the number of idiobiont and koinobiont species, to climatic, physiographic and regional factors using generalized linear models. We compared models by means of Akaike weighting, and evaluated the spatial structure of their residuals. We used partial regressions to determine whether the final models account for all latitudinal structure in the level of generalism. Results Islands host comparatively more idiobionts than continental areas. Although there is a latitudinal gradient in the level of generalism of island faunas correlating with both environmental factors and island characteristics, the most important determinant of island community structure is their source pool. This effect is stronger for ichneumonids, where generalism is higher in the Indomalayan region, arguably due to the higher diversity of endophytic hosts in its large rain forests. Main conclusions The level of generalism of island parasitoid faunas is largely constrained by regional factors, namely by the structure of the species pool, which emphasizes the importance of including regional processes in our understanding of the functioning of ecological communities. The fact that generalist species are more predominant in islands with a large cover of rain forests pinpoints the importance of the indirect effects of ecological requirements on community structure, highlighting the complex nature of geographical gradients of diversity.     

Among‐species overlap in rodent body size distributions predicts species richness along a temperature gradient

Temperature is widely regarded as a major driver of species richness, but the mechanisms are debated. Niche theory suggests temperature may affect richness by filtering traits and species in colder habitats while promoting specialization in warmer ones. However, tests of this theory are rare because niche dimensions are challenging to quantify along broad thermal gradients. Here, we use individual‐level trait data from a long‐term monitoring network spanning a large geographic extent to test niche‐based theory of community assembly in small mammals. We examined variation in body size among 23 communities of North American rodents sampled across the National Ecological Observatory Network (NEON), ranging from northern hardwood forests to subtropical deserts. We quantified body size similarity among species using a metric of overlap that accounts for individual variation, and fit a structural equation model to disentangle the relationships between temperature, productivity, body size overlap, and species richness. We document a latitudinal gradient of declining similarity in body size among species towards the tropics and overall increase in the dimensions of community‐wide trait space in warmer habitats. Neither environmental temperature nor net primary productivity directly affect rodent species richness. Instead, temperature determines the community‐wide niche space that species can occupy, which in turn alters richness. We suggest a latitudinal gradient of trait space expansion towards the tropics may be widespread and underlie gradients in species diversity.     

Untangling latitudinal richness gradients at higher taxonomic levels: familial perspectives on the diversity of New World bat communities

Aims (i) To describe at the level of local communities latitudinal gradients in the species richness of different families of New World bats and to explore the generality of such gradients. (ii) To characterize the relative effects of changes in the richness of each family to the richness of entire communities. (iii) To determine differences in the rate and direction of latitudinal gradients in species richness within families. (iv) To evaluate how differences among families regarding latitudinal gradients in species richness influence the latitudinal gradient in species richness of entire communities. Location Continental New World ranging from the northern continental United States (Iowa, 42° N) to eastern Paraguay (Canindeyú, 24° S). Methods Data on the species composition of communities came from 32 intensively sampled sites. Analyses focused on species richness of five of nine New World bat families. Multivariate analysis of variance and discriminant function analysis determined and described differences among temperate, subtropical, and tropical climatic zones regarding the species richness of bat families. Simple linear regression described latitudinal gradients in species richness of families. Path analysis was used to describe: (i) the direct effect of latitude on species richness of communities, (ii) the indirect effects of latitude on the species richness of communities through its effect on the species richness of each family, (iii) the relative effects of latitude on the species richness of bat families, and (iv) the relative contribution of each family to variation in the species richness of communities. Results Highly significant differences among climatic zones existed primarily because of a difference between the temperate zone and the tropical and subtropical zones combined. This difference was associated with the high number of vespertilionids in the temperate zone and the high number of phyllostomids in the tropical and subtropical zones. Latitudinal gradients in species richness were contingent on phylogeny. Although only three of the five families exhibited significant gradients, all families except for the Vespertilionidae exhibited indistinguishable increases in species richness with decreases in latitude. The Emballonuridae, Phyllostomidae and Vespertilionidae exhibited significant latitudinal gradients whereby the former two families exhibited the classical increase in species richness with decreasing latitude and the latter family exhibited the opposite pattern. Variation in species richness of all families contributed significantly to variation in the species richness of entire communities. Nonetheless, the Phyllostomidae made a significantly stronger contribution to changes in species richness of communities than did all other families. Much of the latitudinal gradient in species richness of communities could be accounted for by the effects of latitude on the species richness of constituent families. Main conclusions Ecological and evolutionary differences among higher taxonomic units, particularly those differences involving life‐history traits, predispose taxa to exhibit different patterns of diversity along environmental gradients. This may be particularly true along extensive gradients such as latitude. Nonetheless, species rich taxa, by virtue of their greater absolute rates of change, can dominate and therefore define the pattern of diversity at a higher taxonomic level and eclipse differences among less represented taxa in their response to environmental gradients. This is true not only with respect to how bats drive the latitudinal gradient in species richness for all mammals, but also for how the Phyllostomidae drives the latitudinal gradient for all bats in the New World. Better understanding of the mechanistic basis of latitudinal gradients of diversity may come from comparing and contrasting patterns across lower taxonomic levels of a higher taxon and by identifying key ecological and evolutionary traits that are associated with such differences.     

Latitudinal gradient in the taxonomic composition of parasite communities

Although latitudinal gradients in diversity have been well studied, latitudinal variation in the taxonomic composition of communities has received less attention. Here, we use a large dataset including 950 surveys of helminth endoparasite communities in 650 species of vertebrate hosts to test for latitudinal changes in the relative contributions of trematodes, cestodes, nematodes and acanthocephalans to parasite assemblages. Although the species richness of helminth communities showed no consistent latitudinal variation, their taxonomic composition varied as a function of both host type and latitude. First, trematodes and acanthocephalans accounted for a higher proportion of species in helminth communities of fish, whereas nematodes achieved a higher proportion of the species in communities of bird and especially mammal hosts. Second, the proportion of trematodes in helminth communities of birds and mammals increased toward higher latitudes. Finally, the proportion of nematodes per community increased toward lower latitudes regardless of the type of host. We present tentative explanations for these patterns, and argue that new insights in parasite community ecology can be gained by searching for latitudinal gradients not only in parasite species richness, but also in the taxonomic composition of parasite assemblages.     

A macroecological perspective of diversity patterns in the freshwater realm

1. The aim of this paper is to review literature on species diversity patterns of freshwater organisms and underlying mechanisms at large spatial scales. 2. Some freshwater taxa (e.g. dragonflies, fish and frogs) follow the classical latitudinal decline in regional species richness (RSR), supporting the patterns found for major terrestrial and marine organism groups. However, the mechanisms causing this cline in most freshwater taxa are inadequately understood, although research on fish suggests that energy and history are major factors underlying the patterns in total species and endemic species richness. Recent research also suggests that not all freshwater taxa comply with the decline of species richness with latitude (e.g. stoneflies, caddisflies and salamanders), but many taxa show more complex geographical patterns in across‐regions analyses. These complexities are even more profound when studies of global, continental and regional extents are compared. For example, clear latitudinal gradients may be present in regional studies but absent in global studies (e.g. macrophytes). 3. Latitudinal gradients are often especially weak in the across‐ecosystems analyses, which may be attributed to local factors overriding the effects of large‐scale factors on local communities. Nevertheless, local species richness (LSR) is typically linearly related to RSR (suggesting regional effects on local diversity), although saturating relationships have also been found in some occasions (suggesting strong local effects on diversity). Nestedness has often been found to be significant in freshwater studies, yet this pattern is highly variable and generally weak, suggesting also a strong beta diversity component in freshwater systems. 4. Both geographical location and local environmental factors contribute to variation in alpha diversity, nestedness and beta diversity in the freshwater realm, although the relative importance of these two groups of explanatory variables may be contingent on the spatial extent of the study. The mechanisms associated with spatial and environmental control of community structure have also been inferred in a number of studies, and most support has been found for species sorting (possibly because many freshwater studies have species sorting as their starting point), although also dispersal limitation and mass effects may be contributing to the patterns found. 5. The lack of latitudinal gradients in some freshwater taxa begs for further explanations. Such explanations may not be gained for most freshwater taxa in the near future, however, because we lack species‐level information, floristic and faunistic knowledge, and standardised surveys along extensive latitudinal gradients. A challenge for macroecology is thus to use the best possible species‐level information on well‐understood groups (e.g. fish) or use surrogates for species‐level patterns (e.g. families) and then develop hypotheses for further testing in the freshwater realm. An additional research challenge concerns understanding patterns and mechanisms associated with the relationships between alpha, beta and gamma components of species diversity. 6. Understanding the mechanistic basis of species diversity patterns should preferably be based on a combination of large‐scale macroecological and landscape‐scale metacommunity research. Such a research approach will help in elucidating patterns of species diversity across regional and local scales in the freshwater realm.     

Pattern and process in the distribution of North American freshwater fish

Published species lists were analysed to determine the contributions of dispersal, habitat preference, river channel size, body size, and glacial history to large‐scale patterns in freshwater fish species richness in North America, north of central Mexico. Total species richness declines to the north and west but the pattern for endemics differs from that of widespread species. Mississippi Basin regions are more species rich than more isolated, coastal, regions. Richness declines more rapidly with increasing latitude in riverine specialist than in habitat generalist species. Levels of endemism are greatest in species found in small‐ to medium‐sized river channels. The strong Rapoport effect, more marked in migratory than resident species, is correlated with habitat preference, channel size, and glacial history. Body size increases with latitude, largely as a result of a trend from small resident to large migrant species. In unglaciated regions, ancestral species survived in large habitats because these are longer‐lived, more extensive, less isolated and more stable than headwaters, permitting larger populations and lower extinction levels. Reduced levels of gene flow in small, peripheral, channels isolated by larger downstream habitats have resulted in the production of many, small range, small‐bodied species. The latitudinal richness gradient is a consequence of speciation and extinction events in unglaciated faunas and an increasing domination of faunas by generalist, large bodied, large channel, recolonizing species in more northern regions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 46–61.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Explaining the latitudinal gradient anomaly in ichneumonid species richness: evidence from butterflies

1. Despite the profusion of Lepidoptera and other potential hosts in the tropics, the Ichneumonidae are not more species-rich in the lowland tropics than in temperate communities. Alternate but partially complementary hypotheses emphasizing resource fragmentation, predation, or chemical defence have been proposed to explain this paradox.
2. Because different diversity gradients occur in the various subgroups of the Ichneumonidae, as well as among their hosts, general explanations are unlikely to account for this 'anomalous diversity.' Hypotheses of ecological causation are best tested at the taxonomic level where the pattern of interest applies.
3. Phylogenetic trends in the distribution, abundance, and palatability of butterflies (Papilionoidea), a group that increases dramatically in species richness towards the tropics, are presented as a case study for evaluating hypotheses of parasitoid species richness. Evidence is presented that supports the 'nasty host hypothesis' as an explanation for the lack of diversity of associated parasitoids.
4. An examination of hypotheses and the approaches used to test them suggests that the nasty-host hypothesis is best supported by the limited evidence available, while evidence in favour of other hypotheses is either lacking or ambiguous. In particular, use of the koinobiont/idiobiont dichotomy and parasitoid assemblage-size comparisons may result in contradictory conclusions.     

Latitudinal gradients of parasite richness: a review and new insights from helminths of cricetid rodents

The latitudinal diversity gradient (LDG), or the trend of higher species richness at lower latitudes, has been well documented in multiple groups of free‐living organisms. Investigations of the LDG in parasitic organisms are comparatively scarce. Here, I investigated latitudinal patterns of parasite diversity by reviewing published studies and by conducting a novel investigation of the LDG of helminths (parasitic nematodes, trematodes and cestodes) of cricetid rodents (Rodentia: Cricetidae). Using host–parasite records from 175 parasite communities and 60 host species, I tested for the presence and direction of a latitudinal pattern of helminth richness. Additionally, I examined four abiotic factors (mean annual temperature, annual precipitation, annual temperature range and annual precipitation range) and two biotic variables (host body mass and host diet) as potential correlates of parasite richness. The analyses were performed with and without phylogenetic comparative methods, as necessary. In this system, helminths followed the traditional LDG, with increasing species richness with decreasing latitude. Nematode richness appeared to drive this pattern, as cestodes and trematodes exhibited a reverse LDG and no latitudinal pattern, respectively. Overall helminth richness and nematode richness were higher in areas with higher mean annual temperatures, annual precipitation and annual precipitation ranges and lower annual temperature ranges, characteristics that often typify lower latitudes. Cestode richness was higher in areas of lower mean annual temperatures, annual precipitation and annual precipitation ranges and higher annual temperature ranges, while trematode richness showed no relationship with climate variables when phylogenetic comparative methods were used. Host diet was significantly correlated with cestode and trematode species richness, while host body mass was significantly correlated with nematode species richness. Results of this study support a complex association between parasite richness and latitude, and indicate that researchers should carefully consider other factors when trying to understand diversity gradients in parasitic organisms.     

Large-scale species-richness gradients in the Atlantic Ocean

The increase in species richness from the poles to the Equator has been observed in numerous terrestrial and aquatic taxa. A number of different hypotheses have been put forward as explanations for this trend, e.g. area and energy availability. However, whether these hypotheses apply to large spatial scales in marine environments remains unclear. The present study shows a clear latitudinal gradient from high to low latitude (from 80 degrees N to 70 degrees S) in marine species richness for 6643 species (fishes and invertebrates) in 10 different taxa dwelling in benthic and pelagic habitats on both sides of the Atlantic. The patterns in benthic taxa are strongly influenced by coastal hydrographic processes, with marked peaks and troughs, and consequently the gradients are not symmetric along both Atlantic sides. Pelagic taxa show a plateau-shaped distribution and the influence from coastal events on gradients could not be demonstrated. The relationships between species richness and different environmental factors indicate that area size does not explain the latitudinal pattern in benthic species richness on a large spatial scale. Sea-surface temperature (positive relationship) is the best predictor of this pattern for benthic species, and nitrate concentration (negative relationship) is the best predictor for pelagic species. The results call into question the existence of a single primary cause that would explain the pattern in marine species richness on a large spatial scale.     

Antagonistic interaction networks are structured independently of latitude and host guild

An increase in species richness with decreasing latitude is a prominent pattern in nature. However, it remains unclear whether there are corresponding latitudinal gradients in the properties of ecological interaction networks. We investigated the structure of 216 quantitative antagonistic networks comprising insect hosts and their parasitoids, drawn from 28 studies from the High Arctic to the tropics. Key metrics of network structure were strongly affected by the size of the interaction matrix (i.e. the total number of interactions documented between individuals) and by the taxonomic diversity of the host taxa involved. After controlling for these sampling effects, quantitative networks showed no consistent structural patterns across latitude and host guilds, suggesting that there may be basic rules for how sets of antagonists interact with resource species. Furthermore, the strong association between network size and structure implies that many apparent spatial and temporal variations in network structure may prove to be artefacts.     

Body size determines the strength of the latitudinal diversity gradient

In most groups of organisms, the species richness decreases from the tropics to the poles. The mechanisms causing this latitudinal diversity gradient are still controversial. We present data from a comprehensive weighted meta-analysis on the strength of the latitudinal gradient in relation to body size. We sampled literature data on the correlation between species richness and latitude for a variety of organisms, ranging from trees to protozoa. In addition, own data on the presence of large-scale diversity patterns for diatoms were included, both for local and regional species richness. The strength of the latitudinal gradient was positively correlated to the size of the organisms. Strongest decreases of species richness to the poles was found for large organisms like trees and vertebrates, whereas meiofauna, protozoa and diatoms showed weak or no correlations between species richness and latitude. These results imply that latitudinal gradients are shaped by non-equilibrium (regional) processes and are persistent under conditions of dispersal limitation.     

Diversity patterns of leaf-associated aquatic hyphomycetes along a broad latitudinal gradient

Information about the global distribution of aquatic hyphomycetes is scarce, despite the primary importance of these fungi in stream ecosystem functioning. In particular, the relationship between their diversity and latitude remains unclear, due to a lack of coordinated surveys across broad latitudinal ranges. This study is a first report on latitudinal patterns of aquatic hyphomycete diversity associated with native leaf-litter species in five streams located along a gradient extending from the subarctic to the tropics. Exposure of leaf litter in mesh bags of three different mesh sizes facilitated assessing the effects of including or excluding different size-classes of litter-consuming invertebrates. Aquatic hyphomycete evenness was notably constant across all sites, whereas species richness and diversity, expressed as the Hill number, reached a maximum at mid-latitudes (Mediterranean and temperate streams). These latitudinal patterns were consistent across litter species, despite a notable influence of litter identity on fungal communities at the local scale. As a result, the bell-shaped distribution of species richness and Hill diversity deviated markedly from the latitudinal patterns of most other groups of organisms. Differences in the body-size distribution of invertebrate communities colonizing the leaves had no effect on aquatic hyphomycete species richness, Hill diversity or evenness, but invertebrates could still influence fungal communities by depleting litter, an effect that was not captured by the design of our experiment.     

Multiregional comparison of the ecological and phylogenetic structure of butterfly species richness gradients

Aim To examine butterfly species richness gradients in seven regions/countries and to quantify geographic mean root distance (MRD) patterns. My primary goal is to determine the extent to which an explanation for butterfly richness patterns based on tropical niche conservatism and the evolution of cold tolerance, proposed for the fauna of Canada and the USA, applies to other parts of the world. Location USA/Canada, Mexico, Europe/NW Africa, Transbaikal Siberia, Chile, South Africa and Australia. Methods Digitized range maps for butterfly species in each region were used to map richness patterns in summer (for all areas) and winter (for USA/Canada, Europe/NW Africa and Australia). A phylogeny resolved to subfamily was used to map the geographic MRD patterns. Regression trees and general linear models examined climatic and vegetation correlates of species richness and MRD within and among regions. Results Various combinations of climate and vegetation were strong predictors of species richness gradients within regions, but unresolved ‘regional’ factors contributed to the multiregional pattern. Regionally based differences in phylogenetic structure also exist, but MRD is negatively correlated with temperature both within and across areas. MRD patterns consistent with tropical niche conservatism occur in most areas. With a possible partial exception of Mexico, faunas in cold climates and in mountains are more derived than faunas in lowlands and tropical/subtropical climates. In USA/Canada, Europe and Australia, winter faunas are more derived than summer faunas. Main conclusions The phylogenetic pattern previously found in the USA and Canada is widespread in both the Northern and Southern Hemispheres, and niche conservatism and the evolution of cold tolerance is the likely explanation for the development of the global butterfly species richness gradient over evolutionary time. Contemporary climate also influences species richness patterns but is unlikely to be a complete explanation globally. The importance of climate is also manifested in the seasonal loss of more basal butterfly elements outside the tropics in winter.     

Species diversity and structure of phytophagous beetle assemblages along a latitudinal gradient: predicting the potential impacts of climate change

Abstract.  1. Assemblages of phytophagous beetles on Acacia were examined along a 1150 km latitudinal gradient in eastern Australia to investigate the potential effects of climate change on insect communities. The latitudinal gradient was used as a surrogate for differences in temperature. Several possible confounding variables were held constant by selecting comparable sites and emphasising a single host-plant species.
2. Total species richness increased towards the tropics, but there were no significant differences among latitudes for average species density, species richness, Fisher's α , or average Chao-1 index.
3. Beetles sampled along the gradient were classified into four climate change response groups, depending on their latitudinal range and apparent host specificity: cosmopolitan species, generalist feeders , climate generalists , and specialists . These four groups might respond differently to shifting climate zones. Cosmopolitan species (22% of community, found at more than one latitude and on more than one host plant) may be resilient to climate change. Generalist feeders (16%, found only at one latitude but found on more than one Acacia species) may well feed on several species but will have to move with their climatic envelope. Climate generalists (6%, found only on Acacia falcata but found at more than one latitude) may be constrained by the host species' ability to either cope with the changing climate or move with it. Finally, specialists (55%, found only on A. falcata and at only one latitude) may be forced to move poleward concurrently with their host species, or go extinct.
4. The analyses indicate that community structure may be fairly resilient to temperature change. The displacement or local extinction of species, especially the species that are found at only one latitude and on only one host plant, however, may lead to significant changes in community composition.     

Can the tropical conservatism hypothesis explain temperate species richness patterns? An inverse latitudinal biodiversity gradient in the New World snake tribe Lampropeltini

Aim  A latitudinal gradient in species richness, defined as a decrease in biodiversity away from the equator, is one of the oldest known patterns in ecology and evolutionary biology. However, there are also many known cases of increasing poleward diversity, forming inverse latitudinal biodiversity gradients. As only three processes (speciation, extinction and dispersal) can directly affect species richness in areas, similar factors may be responsible for both classical (high tropical diversity) and inverse (high temperate diversity) gradients. Thus, a modified explanation for differential species richness which accounts for both patterns would be preferable to one which only explains high tropical biodiversity.
Location  The New World.
Methods  We test several proposed ecological, temporal, evolutionary and spatial explanations for latitudinal diversity gradients in the New World snake tribe Lampropeltini, which exhibits its highest biodiversity in temperate regions.
Results  We find that an extratropical peak in species richness is not explained by latitudinal variation in diversification rate, the mid-domain effect, or Rapoport's rule. Rather, earlier colonization and longer duration in the temperate zones allowing more time for speciation to increase biodiversity, phylogenetic niche conservatism limiting tropical dispersal and the expansion of the temperate zones in the Tertiary better explain inverse diversity gradients in this group.
Main conclusions  Our conclusions are the inverse of the predictions made by the tropical conservatism hypothesis to explain higher biodiversity near the equator. Therefore, we suggest that the processes invoked are not intrinsic to the tropics but are dependent on historical biogeography to determine the distribution of species richness, which we refer to as the 'biogeographical conservatism hypothesis'.