Using habitat models to map diversity: pan-African species richness of ticks (Acari: Ixodida)

Aims To show how logistic regression models for individual species can be used to produce improved estimates of species richness at a continental scale; to present these data for African ticks (Acari: Ixodida); and to address the question of whether there is a latitudinal gradient in tick species richness. Location Africa. Methods A database of 34,060 collection records for African ticks is used to produce a pan‐African map of known tick species richness at 0.25 × 0.25‐degree resolution. The likely distributions of seventy‐three species are then estimated from environmental factors using logistic regression, and localities where there is a suitably high probability of occurrence for a given species are added to the original data for that species. These augmented data are combined to produce a map of the predicted pan‐African distribution of tick species richness. The relationship of species richness to latitude is considered along a transect placed across some of the more extensively collected areas. Results Maps of known and predicted pan‐African tick species richness are presented, and deficiencies in the available data are highlighted. Correlations using both known and predicted estimates of tick species richness suggest that ticks follow similar species richness patterns to those described for African mammals and birds, with a latitudinal gradient and highest species richness in east equatorial Africa. Tick species ranges are log‐normally distributed. Main conclusions Carefully constructed probability surfaces offer a more powerful approach to mapping species ranges than simple presence‐absence maps. Such models are a useful extension to current biogeographical methods and have a wide range of potential applications in ecology, epidemiology and conservation. Tick species richness at a continental scale follows similar trends to those reported for mammals and birds.     

Equipped to cope with climate change: traits associated with range filling across European taxa

In order to understand the ecological effects of climate change it is essential to forecast suitable areas for species in the future. However, species’ ability to reach potentially suitable areas is also critical for species survival. These ‘range‐shift’ abilities can be studied using life‐history traits related to four range‐shift stages: emigration, movement, establishment, and proliferation. Here, we use the extent to which species’ ranges fill the climatically suitable area available (‘range filling’) as a proxy for the ability of European mammals and birds to shift their ranges under climate change. We detect which traits associate most closely with range filling. Drawing comparisons with a recent analysis for plants, we ask whether the latitudinal position of species’ ranges supports the assertion that post‐glacial range‐shift limitations cause disequilibrium between ranges and climate. We also disentangle which of the three taxonomic groups has greatest range filling. For mammals, generalists and early‐reproducing species have the greatest range filling. For birds, generalist species with high annual fecundity, which live longer than expected based on body size, have the greatest range filling. Although we consider traits related to the four range‐shift stages, only traits related to establishment and proliferation ability significantly influence range filling of mammals and birds. Species with the greatest range filling are those whose range centroid falls in the latitudinal centre of Europe, suggesting that post‐glacial range expansion is a leading cause of disequilibrium with climate, although other explanations are also possible. Range filling of plants is lower than that of mammals or birds, suggesting that plants are more range‐limited by non‐climatic factors. Therefore, plants might be face greater non‐climatic restraints on range shifts than mammals or birds.     

Beta diversity and latitude in North American mammals: testing the hypothesis of covariation

Several hypotheses attempt to explain the latitudinal gradient of species diversity, but some basic aspects of the pattern remain insufficiently explored, including the effect of scales and the role of beta diversity. To explore such components of the latitudinal gradient, we tested the hypothesis of covariation, which states that the gradient of species diversity should show the same pattern regardless of the scale of analysis. The hypothesis implies that there should be no gradients of beta diversity, of regional range size within regions, and of the slope of the species-area curve. For the fauna of North American mammals, we found contrasting results for bats and non-volant species. We could reject the hypothesis of covariation for non-volant mammals, for which the number of species increases towards lower latitudes, but at different rates depending on the scale. Also, for this group, beta diversity is higher at lower latitudes, the regional range size within regions is smaller at lower latitudes, and z, the slope of the species-area relationship is higher at lower latitudes. Contrarily bats did not show significant deviations from the predictions of the hypothesis of covariation: at two different scales, species richness shows similar trends of increase at lower latitudes, and no gradient can be demonstrated for beta diversity, for regional range size, or for the slopes of the species-area curve. Our results show that the higher diversity of non-volant mammals in tropical areas of North America is a consequence of the increase in beta diversity and not of higher diversity at smaller scales. In contrast, the diversity of bats at both scales is higher at lower latitudes. These contrasting patterns suggest different causes for the latitudinal gradient of species diversity in the two groups that are ultimately determined by differences in the patterns of geographic distribution of the species.     

Fleshy fruit characteristics in a temperate deciduous forest of Japan: how unique are they?

This study investigated the fleshy fruit characteristics of 28 woody species in a Japanese temperate forest where large sedentary seed-dispersing mammals are present. We tested whether the findings in previous studies in temperate forests of Europe and North America are universal or not. Results have suggested that fruits of all species were eaten both by birds and mammals except for four species with larger fruits, which were eaten only by mammals. A gradient was found from a syndrome characterized by small, oily, and large-seeded fruits to a syndrome characterized by large, succulent, non-oily, and small-seeded fruits. The sizes and colors of the fruits were not conspicuously different from previous findings in Europe and North America. On the other hand, nitrogen and lipids in the fleshy part did not show seasonally increasing trends, or even seasonally decreasing trends in terms of dry weight. This result, suggesting the absence of community-level adaptation of fruit traits to migratory bird dispersers, contrasted with findings in Europe and North America. Large sedentary arboreal or tree-climbing mammals may have a greater effect on the evolution of fruit-disperser relations than opportunistic migratory birds.     

Global patterns of geographic range size in birds

Large-scale patterns of spatial variation in species geographic range size are central to many fundamental questions in macroecology and conservation biology. However, the global nature of these patterns has remained contentious, since previous studies have been geographically restricted and/or based on small taxonomic groups. Here, using a database on the breeding distributions of birds, we report the first (to our knowledge) global maps of variation in species range sizes for an entire taxonomic class. We show that range area does not follow a simple latitudinal pattern. Instead, the smallest range areas are attained on islands, in mountainous areas, and largely in the southern hemisphere. In contrast, bird species richness peaks around the equator, and towards higher latitudes. Despite these profoundly different latitudinal patterns, spatially explicit models reveal a weak tendency for areas with high species richness to house species with significantly smaller median range area. Taken together, these results show that for birds many spatial patterns in range size described in geographically restricted analyses do not reflect global rules. It remains to be discovered whether global patterns in geographic range size are best interpreted in terms of geographical variation in species assemblage packing, or in the rates of speciation, extinction, and dispersal that ultimately underlie biodiversity.     

NOTES ON SOME BIRDS FROM THE HIGHLANDS OF BASUTOLAND

Dean, W. R. J. &; MacDonald, I. A. W. 1981. A review of African birds feeding in association with mammals. Ostrich 52:135–155.

Feeding associations between African birds and mammals are defined. 96 species of birds show, or possibly show, some form of feeding associations with mammals. Five major categories of feeding association are identified, which differ in relative importance in the various habitat types of Africa. In the most widespread and common feeding association the bird obtains cryptic prey items flushed by the mammal, The opportunistic nature of most bird/mammal feeding associations is stressed. Hypothetical courses In the evolution of the more complex feeding associations are discussed. The possible significance of bird/mammal feeding associations in their biology and conservation is outlined. The lack of quantitative information on these feeding associations limits the conclusions that can be drawn, although their significance for the obligate ectoparasite gleaners, Buphagus species, appears to be clear.     

Of mice and wrens: the relation between abundance and geographic range size in British mammals and birds

We examine the relation between population size and geographic range size for British breeding birds and mammals. As for most other assemblages studied, a strong positive interspecific correlation is found in both taxa. The relation is also recovered once the phylogenetic relatedness of species has been controlled for using an evolutionary comparative method. The slope of the relation is steeper for birds than for mammals, but this is due in large part to two species of mammals that have much higher population sizes than expected from their small geographic ranges. These outlying mammal species are the only ones in Britain to be found only on small offshore islands, and so may be exhibiting density compensation effects. With them excluded, the slope of the abundance–range size relation for mammals is not significantly different to that for birds. However, the elevation of the relation is higher for mammals than for birds, indicating that mammals are approximately 30 times more abundant than birds of equivalent geographic range size. An earlier study of these assemblages showed that, for a given body mass, bats had abundances more similar to birds than to non-volant mammals, suggesting that the difference in abundance between mammals and birds might be due to constraints of flight. Our analyses show that the abundance–range size relation for bats is not different for that from other mammals, and that the anomalously low abundance of bats for their body mass may result because they have smaller than expected geographic extents for their size. Other reasons why birds and mammals might have different elevations for the relation between population size and geographic range size are discussed, together with possible reasons for why the slopes of these relations might be similar.     

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.     

Macroecological patterns in British breeding birds: covariation of species'geographical range sizes at differing spatial scales

The availability of high quality data on the distribution and abundance of British birds at the national scale means that this fauna is the basis for a growing body of macroecological study. Nevertheless. questions remain about how representative of wider patterns the distributions and abundances of birds in Britain may be. Here, we use data on the British. European and global breeding distributions of British birds to show that species that are widespread in Britain also tend to be widespread across larger regions. These results hold for both residents and migrants separately, and when controlling for the phylogenetic related ness of species. Species with wide latitudinal spans in Europe also tend to have large British ranges, with the largest British ranges exhibited by species inhabiting mid-latitudes in Europe. These results demonstrate that the distributions of birds within Britain are not simply idiosyacratic. but do reflect aspects of their broader distributions.     

Genetic diversity within vertebrate species is greater at lower latitudes

The latitudinal gradient of species diversity is one of the oldest recognized patterns in biology. While the cause of the pattern remains debated, the global signal of greater diversity toward the tropics is widely established. Whether the pattern holds for genetic diversity within species, however, has received much less attention. We examine latitudinal variation of intraspecific genetic diversity by contrasting nucleotide distance within low- and high-latitude animal groups. Using mitochondrial DNA markers across 72 vertebrate species that together span six continents, two oceans, and 129 degrees of latitude, we found significantly greater genetic diversity at low latitudes within mammalian species, and trends consistent with this pattern in reptiles, amphibians, fish, and birds. The signal held even after removing species whose current geographic ranges include areas recently covered by glaciers during the late Pleistocene and which presumably have experienced colonization bottlenecks in high latitudes. Higher genetic diversity within species was found at low latitudes also for genera that do not possess higher species richness toward the tropics. Moreover, examination of a subset of species with sufficient sampling across a broad geographic range revealed that genetic variation demonstrates a typical gradient, with mid-latitude populations intermediate in genetic diversity between high and low latitude ones. These results broaden the pattern of the global latitudinal diversity gradient, to now include variation within species. These results are also concordant with other studies indicating that low latitude populations and species are on different evolutionary trajectories than high latitude ones, and we speculate that higher rates of evolution toward the equator are driving the pattern for genetic diversity within species.     

The missing Madagascan mid-domain effect

Species richness varies enormously across geographical gradients, a well-known phenomenon for which there are many hypothesized explanations. One recent hypothesis uses null models to demonstrate that random re-distribution of species' ranges within a given domain leads to a 'mid-domain effect' (MDE): increasing species richness towards the centre of the area. Madagascar is especially well-suited for empirical evaluation of mid-domain models by virtue of its large endemic fauna and its clearly defined boundaries. Lees et al. [ Biol. J. Linn. Soc. 67 (1999) 529] observed patterns of species richness consistent with MDEs in the Madagascan rainforest (a slim, north–south belt). In this study, we test one-dimensional and two-dimensional mid-domain model predictions for the birds and mammals of the entire island of Madagascar. When only latitudinal extents of species' distribution are considered, patterns of richness in Madagascar show an MDE. However, this pattern disappears for both taxa after accounting for the tendency of latitudinal bands nearer the middle of the country to be larger. Two-dimensional mid-domain model predictions of species richness are qualitatively opposite to observed patterns. Instead, island-wide spatial gradients of species richness in Madagascar relate strongly to patterns of primary productivity and amount of remaining natural habitat. Earlier work that showed a mid-domain peak within the rainforest biome (effectively after controlling for climate and natural habitat) seems likely to have reflected methodological artefacts. The classic case in which MDEs should occur is, in fact, inconsistent with the mid-domain hypothesis.     

Latitudinal patterns of mammalian species richness in the New World: the effects of sampling method and faunal group

Abstract. Although the latitudinal gradient of species richness for mammals in North America is well documented, few investigators have quantified the relationship in South America. We examined the pattern in North and South America, at two spatial scales (2.5° and 5°) for each of two sampling methods (quadrats and latitudinal bands). A scale effect was evident for quadrats but not for bands. Significant linear relationships between species richness and latitude were found for three faunal groups: all mammals, nonvolant species, and bats. Effects of area confound the latitudinal relationship. By statistically removing such effects, we found that the latitudinal gradient is not an artifact of the species-area relationship, and that the latitudinal gradients for North and South America were statistically indistinguishable. Our data suggest that both faunal subgroups, nonvolant species and bats, contributed substantially to the overall mammalian pattern. Further, multiple regression analyses showed that only latitude is a necessary variable to explain bat richness; for nonvolant species, in addition to latitude, area and longitude may be important.     

Population structure and migratory directions of Scandinavian bluethroats Luscinia svecica– a molecular, morphological and stable isotope analysis

Many species of birds show evidence of secondary contact zones and subspeciation in their Scandinavian distribution range, presumably resulting from different post-glacial recolonization routes. We investigated whether this is the case also in the Scandinavian bluethroat Luscinia svecica , a species that has been suggested to consist of two separate populations: one SW-migrating and long-winged ( L. s. gaetkei ) breeding in southern Norway, and one shorter-winged ESE-migrating ( L. s. svecica ) in northern Scandinavia. We sampled males at eleven breeding sites from southern Norway to northernmost Sweden. There were no morphological differences or latitudinal trends within the sample, neither were there any genetic differences or latitudinal trends as measured by variation in AFLP and microsatellite markers. Stable isotope ratios of throat feathers moulted on the wintering grounds showed no, or possibly marginal differences between birds from southern Norway and northern Sweden. We also re-measured old museum skins that in previous studies were classified as L. s. gaetkei , and found marginally longer wings in birds from the southern part of the Scandinavian breeding range. The difference, however, was much smaller than proposed in earlier studies. We conclude that there is no evidence of a genetic population structure among Scandinavian bluethroats that would suggest the presence of a zone of secondary contact. Finally we discuss whether the presumed subspecies gaetkei ever existed.     

Body size and latitudinal gradients in regional diversity of New World birds

Are latitudinal gradients in regional diversity random or biased with respect to body size? Using data for the New World avifauna, I show that the slope of the increase in regional species richness from the Arctic to the equator is not independent of body size. The increase is steepest among small and medium‐sized species, and shallowest among the largest species. This is reflected in latitudinal variation in the shape of frequency distributions of body sizes in regional subsets of the New World avifauna. Because species are added disproportionately in small and medium size classes towards low latitudes, distributions become less widely spread along the body size axis than expected from the number of species. These patterns suggest an interaction between the effects of latitude and body size on species richness, implying that mechanisms which vary with both latitude and body size may be important determinants of high tropical diversity in New World birds.     

A meta‐analysis of global avian survival across species and latitude

Tropical birds are purported to be longer lived than their temperate counterparts, but it has not been shown whether avian survival rates covary with latitude worldwide. Here, we perform a global‐scale meta‐analysis of 949 estimates from 204 studies of avian survival and demonstrate that a latitudinal survival gradient exists in the northern hemisphere, is dampened or absent for southern hemisphere species, and that differences between passerines and nonpasserines largely drive these trends. We also show that while extrinsic factors related to climate were poor predictors of apparent survival compared to latitude alone, the relationship between apparent survival and latitude is strongly mediated by intrinsic traits – large‐bodied species and species with smaller clutch size had the highest apparent survival. Our findings reveal that differences among intrinsic traits and whether species were passerines or nonpasserines surpass latitude and its underlying climatic factors in explaining global patterns of apparent avian survival.     

Bird population trends are linearly affected by climate change along species thermal ranges

Beyond the effects of temperature increase on local population trends and on species distribution shifts, how populations of a given species are affected by climate change along a species range is still unclear. We tested whether and how species responses to climate change are related to the populations locations within the species thermal range. We compared the average 20 year growth rates of 62 terrestrial breeding birds in three European countries along the latitudinal gradient of the species ranges. After controlling for factors already reported to affect bird population trends (habitat specialization, migration distance and body mass), we found that populations breeding close to the species thermal maximum have lower growth rates than those in other parts of the thermal range, while those breeding close to the species thermal minimum have higher growth rates. These results were maintained even after having controlled for the effect of latitude per se. Therefore, the results cannot solely be explained by latitudinal clines linked to the geographical structure in local spring warming. Indeed, we found that populations are not just responding to changes in temperature at the hottest and coolest parts of the species range, but that they show a linear graded response across their European thermal range. We thus provide insights into how populations respond to climate changes. We suggest that projections of future species distributions, and also management options and conservation assessments, cannot be based on the assumption of a uniform response to climate change across a species range or at range edges only.     

Variability and size in mammals and birds

Body size, its variability, and their ecological correlates have long been important topics in evolutionary biology. Yet, the question of whether there is a general relationship between size and size-relative variability has not previously been addressed. Through an analysis of body-mass and length measurements from 65 074 individuals from 351 mammalian species, we show that size-relative variability increases significantly with mean species body size. Analysis of mean body mass and standard deviations for 237 species of birds revealed the same pattern. We present three plausible alternatives explanations and eliminate several others. Of these, the hypothesis that the increase in size-relative variability with mean body mass is related to the scaling of body mass components is most strongly supported. In effect, larger mammals and birds are more variable because their body mass is composed to greater relative degree of components with higher intrinsic variability (bone, fat, and muscle). In contrast, smaller mammals and birds have lower body mass variability because they are composed to a greater relative extent of components (viscera and nervous system) in which size variation is more highly constrained by energetic and functional factors.     

Break the pattern: breakpoints in beta diversity of vertebrates are general across clades and suggest common historical causes

The use of correlative analyses might be insufficient to understand the processes that control biodiversity, because the variables accounting for different hypotheses (e.g. current climate, past climate change, post‐glacial dispersal limitation) are mutually correlated. We suggest here that, in order to gain insight, it could be useful to search for latitudinal thresholds that could provide information about qualitative changes in the way biodiversity varies in space. Such tipping points could inform about higher‐level processes that are not reflected in correlative analyses. We test whether similar breakpoints in latitudinal beta‐diversity patterns exist for different vertebrate groups with diverse life histories and dispersal abilities. In birds, bats and non‐volant mammals we find breakpoints similar to those of amphibians. Differences in species composition are mainly due to species replacement from the equator to the breakpoint, but are dominated by nested species losses from the breakpoint to higher latitudes. Thus, marked thresholds discriminate two world regions where different processes appear to drive biodiversity.