Rapoport's Rule Revisited: Geographical Distributions of Human Languages

One of the most well studied ecological patterns is Rapoport''s rule, which posits that the geographical extent of species ranges increases at higher latitudes. However, studies to date have been limited in their geographic scope and results have been equivocal. In turn, much debate exists over potential links between Rapoport''s rule and latitudinal patterns in species richness. Humans collectively speak nearly 7000 different languages, which are spread unevenly across the globe, with loci in the tropics. Causes of this skewed distribution have received only limited study. We analyze the extent of Rapoport''s rule in human languages at a global scale and within each region of the globe separately. We test the relationship between Rapoport''s rule and the richness of languages spoken in different regions. We also explore the frequency distribution of language-range sizes. The language-range area distribution is strongly right-skewed, with 87% of languages having range areas less than 10,000 km², and only nine languages with range areas over 1,000,000 km². At a global scale, language-range extents and areas are positively correlated with latitude. At a global scale and in five of the six regions examined, language-range extent and language-range area are strongly correlated with language richness. Our results point to group boundary formation as a critical mediator of the relationship between Rapoport''s rule and diversity patterns. Where strong group boundaries limit range overlap, as is the case with human languages, and range sizes increase with latitude, latitudinal richness gradients may result.     

Environmental correlates of body size distributions of European springtails (Hexapoda: Collembola)

Aim Species–body size distributions (SBDs) are plots of species richness across body size classes. They have been linked to energetic constraints, speciation–extinction dynamics and to evolutionary trends. However, little is known about the spatial variation of size distributions. Here we study SBDs of European springtails (Collembola) at a continental scale and test whether minimum, average and maximum body size and the shapes of size distributions change across latitudinal and longitudinal gradients and whether SBDs of islands and mainlands differ. We also test whether the island rule and the positive body size–range size relationship of vertebrates also holds for Collembola. Location Europe. Methods We use a unique data set on the spatial distributions of 2102 species of European springtails across 52 countries and larger islands together with associated data on body size, area, climate variables, longitude and latitude. Differences in the central moments of SBDs are inferred from simultaneous spatial autoregression models. Results The SBD of the European Collembola and its largest suborder Entomobryomorpha is unimodal and symmetrical. Average, minimum and maximum body weight and the skewness of the mainland/island SBDs peaked at intermediate latitudes. We could not find simple latitudinal gradients in minimum and maximum body weight. Average and maximum body size increased with country/island area in accordance with the island rule in vertebrates, while minimum body size did not significantly differ between islands and mainlands. Finally, we found a weak but statistically significant positive correlation of range size and body size. Main conclusions We provide evidence for differences in body size distributions between islands and mainlands that are in part in line with the island rule in invertebrates. We also find evidence for an interspecific body size–range size relationship similar to that of vertebrates although the vertebrate pattern is much stronger than the springtail pattern. Our results on latitudinal gradients of maximum and average body size imply the need to account for species richness and area effects in the study of latitudinal gradients in body size. We recommend implementing sample size and area effects in the study of body size distributions on islands and mainlands.     

Patterns in species richness, size, and latitudinal range of East Atlantic fishes

Examination of latitudinal patterns in species richness, size, and distributional range of East Atlantic fish, based on a compilation of data encompassing the full latitudinal and depth distribution of 1746 East Atlantic fish species, showed that species richness declined towards higher latitudes at a rate of c 1 % of the number of species present, in five-degree bands, for each degree of latitude for both teleosts and elasmobranchs, regardless of habitat However, the latitudinal patterns in maximum fish size and latitudinal range differed between teleosts and elasmobranchs, and changed with habitat No clear evidence was obtained that the latitudinal range occupied increased with latitude, indicating that Rapoport's rule does not apply to E Atlantic fishes Rather, the latitudinal patterns in species richness, size, and distributional range of benthic Atlantic fish were depth-dependent, because species richness, average maximum size, and the average latitudinal range increased with depth and declined with latitude The importance of accounting for this depth-latitude covariation in the distribution of marine fish demonstrated here, together with recent evidence obtained for deep-sea benthic macrofauna, points to depth and latitude as the main factors in the distribution of marine animals     

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.     

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.     

Patterns of turtle species'geographic range size and a test of Rapoport's rule

There has been a recent resurgence of interest in documenting and explaining patterns of species' range sizes with the goal of determining if general patterns exist. Much recent debate has centered on whether Rapoport's rule, the tendency for range size to increase with increasing latitude, is a general rule or a local effect. I calculated the sizes of turtle species ranges from distribution maps and used published natural history data to examine how range size varies with latitude, and to determine if differences in range size exist among continents, and if correlations with body size, available land area, habitat and diet breadth exist. The distribution of turtle range sizes extended over six orders of magnitude and formed a lognormal distribution with many species having moderate or small ranges and few species having large ranges. Range size was positively correlated with available land area, habitat breadth, diet breadth, and body size. Multiple regression accounted for only 39% of the variance in range size indicating that other important factors remain unknown. At both global and continental scales, range size is largest near the equator and decreases with increasing latitude, the apposite of Rapoport's rule. However, range size did increase latitudinally above 25-30°N in both the Neararctic and Palearctic suggesting that the pattern would be more accurately considered a local effect than a general rule. Larger range sizes at low latitudes may occur because more land area in the tropics provides much suitable habitat for ectotherms and there are few large scale physical barriers to dispersal. Rapoport patterns result from the occurrence of a small number of wide-ranging cold tolerant species that have reinvaded northern latitudes after Pleistocene glaciation. Patterns of the longitudinal and latitudinal extents of species ranges and their positions illustrate the importance of climate, mountain ranges, deserts, and coastlines, as barriers potentially affecting range size.     

Geographic variation in body size and sexual size dimorphism of a seed-feeding beetle

Body size of many animals varies with latitude: body size is either larger at higher latitudes (Bergmann's rule) or smaller at higher latitudes (converse Bergmann's rule). However, the causes underlying these patterns are poorly understood. Also, studies rarely explore how sexual size dimorphism varies with latitude. Here we investigate geographic variation in body size and sexual size dimorphism of the seed-feeding beetle Stator limbatus, collected from 95 locations along a 38 degrees range in latitude. We examine 14 variables to test whether clines in environmental factors are adequate to explain geographic patterns of body size. We found that body size and sexual size dimorphism of S. limbatus varied considerably with latitude; beetles were smaller but more dimorphic at lower latitudes. Body size was not correlated with a gradient in mean temperature, contrary to the commonly accepted hypothesis that clines are produced by latitudinal gradients in temperature. Instead, we found that three factors were adequate to explain the cline in body size: clinal variation in host plant seed size, moisture (humidity), and seasonality (variance in humidity, precipitation, and temperature). We also found that the cline in sexual size dimorphism was partially explainable by a gradient in moisture, though moisture alone was not sufficient to explain the cline. Other ecological or environmental variables must necessarily contribute to differences in selection on male versus female body size. The main implications of our study are that the sexes differ in the magnitude of clinal variation in body size, creating latitudinal variation in sexual size dimorphism, and that clines in body size of seed beetles are likely influenced by variation in host seed size, water availability, and seasonality.     

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.     

Knitting patterns of biodiversity,range size and body size in aquatic beetle faunas: significant relationships but slightly divergent drivers

1. Ecogeographical rules refer to recurring patterns in nature, including the latitudinal diversity gradient (LDG), Rapoport's rule and Bergmann's rule, amongst others. In the present study, the existence of these rules was examined for diving beetles (Coleoptera: Dytiscidae), a family of aquatic predatory beetles. 2. Assemblage‐level data were analysed for diving beetles, focusing on species richness, local contribution to beta diversity (LCBD), mean range size and mean body size across the biogeographical provinces of Northern Europe. First, each of these variables was correlated with latitude, and then variation in each variable was modelled using actual environmental variables in boosted regression tree analysis. 3. Species richness was found to decrease with latitude, LCBD increased with latitude, mean range size did not show a significant relationship with latitude, and mean body size decreased with latitude. The latter finding was in contrast to Bergmann's rule. The actual environmental variables best predicting variation in these four response variables varied among the models, although they generally included temperature‐related and land use variables as the most influential ones. 4. The results obtained in the present study suggest that diving beetles conformed to the LDG, did not follow Rapoport's rule, and showed a reversed latitudinal gradient in the context of Bergmann's rule. In addition, species‐poor provinces harboured ecologically most unique faunas, suggesting that species richness and LCBD are complementary measures of biodiversity. 5. Even though general support was not found for most of the ecogeographical rules examined, the findings of the present study are interesting because they suggest that aquatic ectothermic invertebrates may show patterns different from those originally described for terrestrial endothermic vertebrates.     

Diversity and distribution of small mammals in the South American Dry Andes

The Andean mountain range has played an important role in the evolution of South American biota. However, there is little understanding of the patterns of species diversity across latitudinal and altitudinal gradients. In this paper, we examine the diversity of small mammals along the South Central Dry Andes (SCDA) within the framework of two contrasting hypotheses: (a) species richness decreases with increasing elevation and latitude; and (b) species richness peaks at altitudinal midpoints (mid‐domain). We explore the composition of the species pool, the impact of species–area relationships and the Rapoport effect (i.e. size of geographic ranges) along latitudinal and elevational gradients. First, we constructed a database of SCDA small mammals. Then, species richness patterns were analysed through generalized models, and species–area relationships were assessed by log–log regressions; the curvilinear method (c = S/Az) was use to compute richness corrected by area size. Lastly, the Rapoport effect was evaluated using the midpoint method. Our results show: (1) a richness of 67 small mammals along the SCDA, of which 36 are endemic; (2) a hump‐shaped pattern in species richness along elevation and latitudinal gradients; (3) a species–area relationship for both gradients; (4) endemic species corrected by area present a strong and positive relationship with elevation; (5) a Rapoport effect for the latitudinal ranges, but no effect across the elevational gradient; and (6) a major species turnover between 28° and 30° south latitude. This is the first study quantifying the diversity of small mammals encompassing the central Andean region. Overall, our macrogeographic analysis supports the previously postulated role of the Andes in the diversification of small mammals (i.e. in situ cladogenesis) and highlights some basic attributes (i.e. anatomy of geographic ranges; species–area relationships) when considering the consequences of climate change on biodiversity conservation of mountain ecosystems.     

Latitudinal variation in the shape of the species body size distribution: an analysis using freshwater fishes

Many taxonomic and ecological assemblages of species exhibit a right-skewed body size-frequency distribution when characterized at a regional scale. Although this distribution has been frequently described, factors influencing geographic variation in the distribution are not well understood, nor are mechanisms responsible for distribution shape. In this study, variation in the species body size-frequency distributions of 344 regional communities of North American freshwater fishes is examined in relation to latitude, species richness, and taxonomic composition. Although the distribution of all species of North American fishes is right-skewed, a negative correlation exists between latitude and regional community size distribution skewness, with size distributions becoming left-skewed at high latitudes. This relationship is not an artifact of the confounding relationship between latitude and species richness in North American fishes. The negative correlation between latitude and regional community size distribution skewness is partially due to the geographic distribution of families of fishes and apparently enhanced by a nonrandom geographic distribution of species within families. These results are discussed in the context of previous explanations of factors responsible for the generation of species size-frequency distributions related to the fractal nature of the environment, energetics, and evolutionary patterns of body size in North American fishes.     

What determines conformity to Bergmann's rule?

Aim  Bergmann's rule, the tendency of body size within species in bird and mammal populations to be positively correlated with latitude, is among the best known biogeographical generalizations. The factors behind such clines, however, are not well understood. Here we use a large data base of 79 mammalian carnivore species to examine the factors affecting latitudinal size clines.
Location  Worldwide.
Methods  We measured the skulls and teeth of carnivores in natural history museums, and calculated the amount of variation in size explained by latitude, supplementing our measurements with published data. We examined the effects of a number of variables on the tendency to show latitudinal clines.
Results  We found that geographical range and latitudinal extent are strongly related to size clines. Minimum temperatures across the range, net primary productivity and habitat diversity also have some, albeit much less, influence.
Main conclusions  We suggest that species with large geographical ranges are likely to encounter significant heterogeneity in those factors that influence body size, and are thus likely to exhibit size clines. However, the key factors that determine body size may not always operate along a latitudinal (or other geographical) cline, but be spatially linked to patches in the species range. One such important factor is likely to be food availability, which we show is a strong predictor of size in the brown bear ( Ursus arctos ) but is not associated with a latitudinal cline. We argue that the spatial distribution of key resources within the species range constitutes a significant predictor of carnivore body size.     

Latitudinal gradients of species richness: a test of the geographic area hypothesis at two ecological scales

The geographic area hypothesis advances area as the primary cause of latitudinal gradients in diversity. The greater area of tropical zones, it suggests, stimulates speciation, inhibits extinction, and leads to increased species richness compared to the situation in smaller temperate and boreal zones. Because bats exhibit exceptionally strong latitudinal gradients of richness at multiple spatial scales in the New World, they are an appropriate system with which to test the geographic area hypothesis. We used range maps for 250 species of New World bats to estimate species richness in biogeographic zones at two hierarchical spatial scales: biome types and provinces. We then conducted a series of regression analyses to evaluate the ability of area to account for latitudinal gradients in species richness. However, spillover (zonal bleeding) of tropical species into extra-tropical zones may mask the species-area relationship and alter perceptions of the latitudinal gradient. To address this issue, we conducted additional analyses excluding tropical species, using a series of increasingly inclusive definitions of tropical ranges. Ecogeographic zones of the New World are not larger at tropical versus extra-tropical latitudes. Moreover, spillover of tropical species into ecogeographic zones within extra-tropical regions generally does not diminish the association between richness and area. Nonetheless, the latitudinal gradient of species richness is strong and significant at both ecogeographic scales. Clearly, area does not drive the latitudinal gradient of bat species richness in the New World. In fact, area represents a source of noise rather than a dominant signal at the focal scale of biome types and provinces in the Western Hemisphere.     

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.     

Beyond Rapoport's rule: evaluating range size patterns of New World birds in a two-dimensional framework

Aim To evaluate Rapoport's rule for New World birds in two‐dimensional geographical space. We specifically test for a topography × climate interaction that predicts little difference in range sizes between lowlands and mountains in cold climates, whereas in the tropics, montane species have narrow ranges and lowland species have broad ranges. Location The western hemisphere. Methods We used digitized range maps of breeding birds to generate mean range sizes in grids of 27.5 × 27.5 km and 110 × 110 km across North and South America. We examined the geographical pattern with respect to range in elevation, mean temperature in the coldest month, their interaction, biome size and continental width, using model II analysis of variance, multiple regression and simple correlation. Results In northern latitudes species have broad ranges in both mountainous and flat areas. However, range sizes in the mountains and lowlands diverge southwards, with the most extreme differences in the tropics. Further, there are minimal differences in range sizes across latitudes in lowlands. The smallest mean ranges occur in the tropical Andes. Mean range sizes in north‐central Canada, Central America and Argentina/Chile are also small, reflecting the narrowing of the continents in these areas. The best regression model explained 51% of the variation in mean range size. Main conclusions The two‐dimensional range size pattern indicates that neither winter temperature nor annual variability in temperature strongly influences the distribution of range sizes directly; rather, climate influences bird range sizes indirectly via effects on habitat size. Also, macroclimate interacts with topographic relief across latitudes, generating sharp mesoscale habitat gradients in tropical mountains but not in high latitude mountains or in lowlands at any latitude. Birds respond to these habitat gradients, resulting in ‘latitudinal’ range size gradients in topographically complex landscapes but not in simple landscapes.     

Wing morphology and migration status,but not body size,habitat or Rapoport's rule predict range size in North‐American dragonflies (Odonata: Libellulidae)

Understanding why species range sizes vary is important for predicting the impact of environmental change on biodiversity. Here we use a multi‐variable approach in a phylogenetic comparative context to understand how four morphological, two ecological, and two eco‐geographical variables are associated with range size, latitudinal range and longitudinal range in 81 species of North‐American libellulid dragonflies. Our results show that: 1) migratory species and species with a more expanded basal hindwing lobe have a larger range size; 2) opposite to Rapoport's rule, latitudinal range is negatively correlated with mid‐range latitude; 3) longitudinal range is predicted by wing morphology and migration; 4) body size and larval habitat are not correlated with range size, latitudinal range or longitudinal range. These results suggest that dispersal‐related traits, such as wing shape and migratory status, are important factors in predicting the range size of libellulid dragonflies. In addition, the reverse Rapoport's rule suggests that more northern‐centred species might be more specialized than more southern‐centred species. We suggest that the variables predicting range size are likely imposed by taxon‐specific morphological, ecological, physiological and behavioural traits. Taxon‐specific knowledge is thus necessary to understand the dynamics of range sizes and is important to implement successful restoration and conservation plans of threatened species.     

Latitudinal gradients in butterfly body sizes: is there a general pattern?

We tested for the existence of latitudinal gradients in the body sizes of butterflies in North America, Europe, Australia and the Afrotropics. We initially compared body sizes (measured as male forewing length) of all butterflies found in 5° latitudinal bands in each region, and then evaluated the relationship between body size and latitude statistically using the latitudinal midpoint of each species' distribution. Trends were examined for species in all butterfly families together and for each family separately. We found that gradients in body sizes were inconsistent in different geographical regions and butterfly families; in some cases species were larger towards the tropics, in some they were smaller, and in other cases there were no relationships. Most of the gradients, when they existed, reflected between-family effects arising from changes in the relative numbers of species in each family across regions. We conclude that general ecological explanations for geographical trends in butterfly body sizes are inappropriate, and gradients largely reflect historical patterns of speciation within and between taxa in each biogeographical realm. Thus, the robustness of body size gradients found in other insect groups should be confirmed in future studies by including more than one geographical region whenever possible.     

Sensitivity of intraspecific latitudinal clines of body size for tetrapods to sampling, latitude and body size

Recent studies have shown that most tetrapod groups (mammals,birds, chelonians, amphibians) show general intraspecific tendenciesfor increasing body size with latitude, whereas squamates (lizardsand snakes) show an intraspecific tendency towards decreasingbody size with latitude. Here I evaluate whether these sizetrends are general by using independent contrasts analysis toinvestigate the dependence of intraspecific size-latitude relationships(r), and the magnitude alone of size-latitude relationships([r]), for tetrapod vertebrates, on sample size, range of latitudessampled, average latitude sampled, and body size. Range of latitudessampled, average latitude sampled, and body size did not influencebody size-latitude relationships (r) or the magnitude aloneof body size-latitude relationship ([r]). Sample size did notinfluence size-latitude relationships (r), but did influencethe magnitude alone of size-latitude relationships ([r]), possiblyindicating increased precision of estimating size-latitude relationshipswith increased sampling. In short, intraspecific size-latituderelationships are similar for species of different sizes, occurringat different latitudes, sampled over different latitudinal ranges,and differing in number of populations sampled (though magnitudealone is influenced by sample size). These results suggest thatintraspecific size-latitude trends are general, and biologicallysignificant (i.e., are not artifacts of sampling), thus deservingexplanation.     

The 29°N latitudinal line: an important division in the Hengduan Mountains, a biodiversity hotspot in southwest China

This paper aimed to explore the division of the southern and northern Hengduan Mountains based on gradients in species similarity and richness, and to analyze species richness in each sub-region. The Hengduan Mountain region was divided into nine latitudinal belts using one degree of latitude to define the belt after which distribution of seed plants within each latitudinal belt was recorded. Latitudinal patterns of species similarity were measured using the Jaccard similarity index for each pair of adjacent latitudinal belts. Non-metric multidimentional scaling (NMDS) was also used to analyze the similarity in species composition among the nine latitudinal belts. The latitudinal pattern of species similarity and the NMDS ordination both showed a great change in species composition across the 29°N latitudinal line, essentially dividing the Hengduan Mountain region into southern and northern sub-regions. Species richness, shown by the c-value of the species–area power function, and species–area ratio along a latitudinal gradient both showed a sharp decrease across the latitudinal belt from 29°0' to 29°59'N. The southern sub-region occupied 40% of the total area of the Hengduan Mountain region, but contained more than 80% of all the seed plants in the region. The higher species richness and endemism in the southern sub-region showed it to be the core of the Hengduan biodiversity hotspot, a result not unexpected because of the greater extremes of topography and wider diversity of habitats in the southern portion.