High Tropical Diversity Research Articles

Niche conservatism limits the distribution ofMedicagoin the tropics

The increase in species number from poles to the equator is one of the most fundamental patterns in ecology. Although several hypotheses have been proposed, there is a lack of consensus on the mechanisms underlying this pattern. While most hypotheses provide plausible explanation for high tropical diversity of tropical clades, it is unclear if similar mechanisms drive the diversity of extra‐tropical clades. Here, we investigated the environmental drivers influencing the diversity pattern of a Mediterranean plant genusMedicagoand dissect their effects across continents and biomes. We compiled a comprehensive dataset on the distribution of allMedicagospecies and mapped their distribution at the spatial resolution of 100 × 100 km2. We used generalized linear models to quantify the relative effects of environmental factors on the richness patterns ofMedicagoand its two life forms. Next, using geographically weighted regression, we explored the variation in richness–environment relationship across latitude. We found that Quaternary climate change and environmental energy were important explanatory variables, and their effects were consistent at global, continental and biome scales. However, we found contrasting effect of environmental energy in temperate and tropical regions, with species richness increasing with energy in the temperate and decreasing in the tropics. We also found strong affinity ofMedicagospecies to its ancestral Mediterranean climatic niche. Our results suggest that the spatial variation inMedicagorichness patterns may be strongly influenced by heat tolerance as opposed to the tropical clades that depend upon cold tolerance. This dependency on heat tolerance can be attributed to strong niche conservatism exhibited byMedicagospecies. Our findings provide new insights into the richness‐energy hypothesis and suggest that high environmental energy may not necessarily yield high species diversity but may also lower species diversity, especially of extra‐tropical clades towards the tropics.

Local adaptation to herbivory within tropical tree species along a rainfall gradient.

In tropical forests, insect herbivores exert significant pressure on plant populations. Adaptation to such pressure is hypothesized to be a driver of high tropical diversity, but direct evidence for local adaptation to herbivory in tropical forests is sparse. At the same time, herbivore pressure has been hypothesized to increase with rainfall in the tropics, which could lead to differences among sites in the degree of local adaptation. To assess the presence of local adaptation and its interaction with rainfall, we compared herbivore damage on seedlings of local vs. nonlocal populations at sites differing in moisture availability in a reciprocal transplant experiment spanning a rainfall gradient in Panama. For 13 native tree species, seeds collected from multiple populations along the rainfall gradient were germinated in a shadehouse and then transplanted to experimental sites within the species range. We tracked the proportion of seedlings attacked over 1.5yr and quantified the percentage of leaf area damaged at the end of the study. Seedlings originating from local populations were less likely to be attacked and experienced lower amounts of herbivore damage than those from nonlocal populations, but only on the wetter end of the rainfall gradient. However, overall herbivore damage was higher at the drier site compared to wetter sites, contrary to expectation. Taken together, these findings support the idea that herbivory can result in local adaptation within tropical tree species; however, the likelihood of local adaptation varies among sites because of environmentally driven differences in investment in defense or herbivore specialization or both.

Patterns of niche diversification in south-east Asian crocodile newts

Niche conservatism underlies the process of speciation in amphibians on the global scale. However, high tropical diversity is partly attributable to niche shifts within evolutionary lineages. Crocodile newts (genera Echinotriton and Tylototriton) are a monophyletic group endemic to south-east Asia, and are one of the few lineages of urodeles that appear in tropical regions. Here, we examined the patterns of ecological diversification within the crocodile newts based on 207 records of 23 species. The niche space was described according to topographic, climatic variables (temperature, rainfall, and seasonality) and forest cover. Our analysis showed that the occurrence of most of these species was strongly associated with the precipitation regime and seasonality. The diversification patterns of the crocodile newts are complex, showing both conservatism and divergence. These patterns can be related to the Tibetan uplift phase during the Neogene, resulting in the emergence of new montane habitats and subsequent changes in the palaeo-drainage system and climate. The range of ecological diversification within a lineage is possibly the result of several interacting processes, including environmental heterogeneity, the regional climatic history, and physiological constraints on species.

Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity

Many groups show higher species richness in tropical regions but the underlying causes remain unclear. Despite many competing hypotheses to explain latitudinal diversity gradients, only three processes can directly change species richness across regions: speciation, extinction and dispersal. These processes can be addressed most powerfully using large-scale phylogenetic approaches, but most previous studies have focused on small groups and recent time scales, or did not separate speciation and extinction rates. We investigate the origins of high tropical diversity in amphibians, applying new phylogenetic comparative methods to a tree of 2871 species. Our results show that high tropical diversity is explained by higher speciation in the tropics, higher extinction in temperate regions and limited dispersal out of the tropics compared with colonization of the tropics from temperate regions. These patterns are strongly associated with climate-related variables such as temperature, precipitation and ecosystem energy. Results from models of diversity dependence in speciation rate suggest that temperate clades may have lower carrying capacities and may be more saturated (closer to carrying capacity) than tropical clades. Furthermore, we estimate strikingly low tropical extinction rates over geological time scales, in stark contrast to the dramatic losses of diversity occurring in tropical regions presently.

Specialization of Mutualistic Interaction Networks Decreases toward Tropical Latitudes

Species-rich tropical communities are expected to be more specialized than their temperate counterparts. Several studies have reported increasing biotic specialization toward the tropics, whereas others have not found latitudinal trends once accounting for sampling bias or differences in plant diversity. Thus, the direction of the latitudinal specialization gradient remains contentious. With an unprecedented global data set, we investigated how biotic specialization between plants and animal pollinators or seed dispersers is associated with latitude, past and contemporary climate, and plant diversity. We show that in contrast to expectation, biotic specialization of mutualistic networks is significantly lower at tropical than at temperate latitudes. Specialization was more closely related to contemporary climate than to past climate stability, suggesting that current conditions have a stronger effect on biotic specialization than historical community stability. Biotic specialization decreased with increasing local and regional plant diversity. This suggests that high specialization of mutualistic interactions is a response of pollinators and seed dispersers to low plant diversity. This could explain why the latitudinal specialization gradient is reversed relative to the latitudinal diversity gradient. Low mutualistic network specialization in the tropics suggests higher tolerance against extinctions in tropical than in temperate communities.

DNA Barcode Libraries Provide Insight into Continental Patterns of Avian Diversification

BackgroundThe causes for the higher biodiversity in the Neotropics as compared to the Nearctic and the factors promoting species diversification in each region have been much debated. The refuge hypothesis posits that high tropical diversity reflects high speciation rates during the Pleistocene, but this conclusion has been challenged. The present study investigates this matter by examining continental patterns of avian diversification through the analysis of large-scale DNA barcode libraries.Methodology and Principal FindingsStandardized COI datasets from the avifaunas of Argentina, the Nearctic, and the Palearctic were analyzed. Average genetic distances between closest congeners and sister species were higher in Argentina than in North America reflecting a much higher percentage of recently diverged species in the latter region. In the Palearctic genetic distances between closely related species appeared to be more similar to those of the southern Neotropics. Average intraspecific variation was similar in Argentina and North America, while the Palearctic fauna had a higher value due to a higher percentage of variable species. Geographic patterning of intraspecific structure was more complex in the southern Neotropics than in the Nearctic, while the Palearctic showed an intermediate level of complexity.Conclusions and SignificanceDNA barcodes can reveal continental patterns of diversification. Our analysis suggests that avian species are older in Argentina than in the Nearctic, supporting the idea that the greater diversity of the Neotropical avifauna is not caused by higher recent speciation rates. Species in the Palearctic also appear to be older than those in the Nearctic. These results, combined with the patterns of geographic structuring found in each region, suggest a major impact of Pleistocene glaciations in the Nearctic, a lesser effect in the Palearctic and a mild effect in the southern Neotropics.

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

ABSTRACTAim 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’.

EVOLUTIONARY AND BIOGEOGRAPHIC ORIGINS OF HIGH TROPICAL DIVERSITY IN OLD WORLD FROGS (RANIDAE)

Differences in species richness between regions are ultimately explained by patterns of speciation, extinction, and biogeographic dispersal. Yet, few studies have considered the role of all three processes in generating the high biodiversity of tropical regions. A recent study of a speciose group of predominately New World frogs (Hylidae) showed that their low diversity in temperate regions was associated with relatively recent colonization of these regions, rather than latitudinal differences in diversification rates (rates of speciation-extinction). Here, we perform parallel analyses on the most species-rich group of Old World frogs (Ranidae; approximately 1300 species) to determine if similar processes drive the latitudinal diversity gradient. We estimate a time-calibrated phylogeny for 390 ranid species and use this phylogeny to analyze patterns of biogeography and diversification rates. As in hylids, we find a strong relationship between the timing of colonization of each region and its current diversity, with recent colonization of temperate regions from tropical regions. Diversification rates are similar in tropical and temperate clades, suggesting that neither accelerated tropical speciation rates nor greater temperate extinction rates explain high tropical diversity in this group. Instead, these results show the importance of historical biogeography in explaining high species richness in both the New World and Old World tropics.

Marine longitudinal biodiversity: causes and conservation

ABSTRACTThe horizontal temperature zones of the earth tend to restrict the latitudinal ranges of species but allow the possibility of exceedingly broad longitudinal dispersals. In the Tropical Zone, biodiversity on the continental shelves is not homogeneous but is concentrated in two conspicuous peaks, one in the Indo‐Pacific Ocean and the other in the Atlantic. The Indo‐Pacific biodiversity peak is located within a relatively small area called the East Indies Triangle. The Atlantic peak is located in the southern Caribbean Sea. Evidence that has been accumulated over the years indicates that each area functions as a centre of origin and evolutionary radiation. What are the causes of these concentrations and their present functions? A newly published model indicates a positive relationship between environmental temperature and the rate of speciation. While this helps to explain the generally high tropical diversity, and the negative relationship between diversity and latitude, it does not provide a reason for the longitudinal concentrations. But, other new research serves to substantiate previous indications of a positive relationship between speciation rate and species diversity. The existence of this positive feedback, together with some contributory factors, provides the reason why concentrations occur. The evolutionary radiation probably begins when the build‐up of species diversity reaches a critical level. The warm‐temperate biotas are derived from the tropics. Their northern longitudinal relationships tend to be minor but, in the southern hemisphere, the West Wind Drift is an important dispersal mechanism for both warm‐temperate and cold‐temperate species. The cold‐temperate biotas peaked in two areas, the North Pacific and the Antarctic; each has developed into a centre of origin. The continuous dispersal of well‐adapted species from the centres helps peripheral communities maintain diversity.

The Latitudinal Gradient in Recent Speciation and Extinction Rates of Birds and Mammals

Although the tropics harbor greater numbers of species than do temperate zones, it is not known whether the rates of speciation and extinction also follow a latitudinal gradient. By sampling birds and mammals, we found that the distribution of the evolutionary ages of sister species-pairs of species in which each is the other's closest relative-adheres to a latitudinal gradient. The time to divergence for sister species is shorter at high latitudes and longer in the tropics. Birth-death models fitting these data estimate that the highest recent speciation and extinction rates occur at high latitudes and decline toward the tropics. These results conflict with the prevailing view that links high tropical diversity to elevated tropical speciation rates. Instead, our findings suggest that faster turnover at high latitudes contributes to the latitudinal diversity gradient.

Evolutionary and Ecological Causes of the Latitudinal Diversity Gradient in Hylid Frogs: Treefrog Trees Unearth the Roots of High Tropical Diversity

Why are there more species in the tropics than in temperate regions? In recent years, this long-standing question has been addressed primarily by seeking environmental correlates of diversity. But to understand the ultimate causes of diversity patterns, we must also examine the evolutionary and biogeographic processes that directly change species numbers (i.e., speciation, extinction, and dispersal). With this perspective, we dissect the latitudinal diversity gradient in hylid frogs. We reconstruct a phylogeny for 124 hylid species, estimate divergence times and diversification rates for major clades, reconstruct biogeographic changes, and use ecological niche modeling to identify climatic variables that potentially limit dispersal. We find that hylids originated in tropical South America and spread to temperate regions only recently (leaving limited time for speciation). There is a strong relationship between the species richness of each region and when that region was colonized but not between the latitudinal positions of clades and their rates of diversification. Temperature seasonality seemingly limits dispersal of many tropical clades into temperate regions and shows significant phylogenetic conservatism. Overall, our study illustrates how two general principles (niche conservatism and the time-for-speciation effect) may help explain the latitudinal diversity gradient as well as many other diversity patterns across taxa and regions.

Evolutionary and Ecological Causes of the Latitudinal Diversity Gradient in Hylid Frogs: Treefrog Trees Unearth the Roots of High Tropical Diversity

Evolutionary and Ecological Causes of the Latitudinal Diversity Gradient in Hylid Frogs: Treefrog Trees Unearth the Roots of High Tropical Diversity

TESTING FOR LATITUDINAL BIAS IN DIVERSIFICATION RATES: AN EXAMPLE USING NEW WORLD BIRDS

Study of the latitudinal diversity gradient to date has focused largely on pattern description, with relatively little work on the possible mechanisms underlying the pattern. One proximate mechanism is a latitudinal bias in the discrepancy between speciation and extinction rates, leading to higher rates of species diversification toward lower latitudes. Despite being central to many explanations for high tropical diversity, this mechanism is tested very rarely. We discuss some of the problems involved in testing for latitudinal bias in diversification rates and present an example phylogenetic analysis for endemic bird genera of the New World. The results provide evidence for higher diversification rates in clades inhabiting lower latitudes, both when genera are considered independent and when phylogeny is controlled for using independent contrasts. High rates of diversification are also associated with larger geographic area and higher net primary productivity, although these do not fully account for the latitudinal effect. The latitudinal pattern is stronger in younger clades, supporting the prediction of a simple model in which the signal of latitudinal bias in diversification rates diminishes as clades age and become saturated with species. Our study demonstrates that a clade-based approach can help answer important questions that a geographic approach cannot, but large phylogenies and geographic databases are needed to cope with the large amount of noise inherent in this type of analysis.

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

AbstractAre 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.