Tropical Conservatism Hypothesis Research Articles

Global Patterns of Earwig Species Richness

One of the most investigated patterns in species diversity is the so-called latitudinal gradient, that is, a decrease in species richness from the equator to the poles. However, few studies investigated this pattern in insects at a global scale because of insufficient taxonomic and biogeographical information. Using estimates of earwig species richness at country level, their latitudinal diversity gradient was modelled globally and for the two hemispheres separately after correcting for differences in country areas. Separate analyses were also conducted for mainland and island countries. All analyses clearly indicated the existence of latitudinal gradients. The most plausible explanation for the observed pattern is the so-called tropical conservatism hypothesis, which postulates (1) a tropical origin of many extant clades, (2) a longer time for cladogenesis in tropical environments thanks to their environmental stability, and (3) a limited ability of historically tropical lineages to adapt to temperate climates. Earwigs probably evolved on Gondwana and secondarily colonized the Northern Hemisphere. This colonization was hampered by both geographical and climatic factors. The Himalayan orogenesis obstructed earwig dispersal into the Palearctic region. Additionally, earwig preferences for warm/hot and humid climates hampered the colonization of temperate regions. Pleistocene glaciation further contributed to reducing diversity at northern latitudes.

Global variation in diversification rate and species richness are unlinked in plants

Species richness varies immensely around the world. Variation in the rate of diversification (speciation minus extinction) is often hypothesized to explain this pattern, while alternative explanations invoke time or ecological carrying capacities as drivers. Focusing on seed plants, the world's most important engineers of terrestrial ecosystems, we investigated the role of diversification rate as a link between the environment and global species richness patterns. Applying structural equation modeling to a comprehensive distribution dataset and phylogenetic tree covering all circa 332,000 seed plant species and 99.9% of the world's terrestrial surface (excluding Antarctica), we test five broad hypotheses postulating that diversification serves as a mechanistic link between species richness and climate, climatic stability, seasonality, environmental heterogeneity, or the distribution of biomes. Our results show that the global patterns of species richness and diversification rate are entirely independent. Diversification rates were not highest in warm and wet climates, running counter to the Metabolic Theory of Ecology, one of the dominant explanations for global gradients in species richness. Instead, diversification rates were highest in edaphically diverse, dry areas that have experienced climate change during the Neogene. Meanwhile, we confirmed climate and environmental heterogeneity as the main drivers of species richness, but these effects did not involve diversification rates as a mechanistic link, calling for alternative explanations. We conclude that high species richness is likely driven by the antiquity of wet tropical areas (supporting the "tropical conservatism hypothesis") or the high ecological carrying capacity of warm, wet, and/or environmentally heterogeneous environments.

Fingerprints of climatic changes through the late Cenozoic in southern Asian flora: Magnolia section Michelia (Magnoliaceae).

Ongoing global warming is a challenge for humankind. A series of drastic climatic changes have been proven to have occurred throughout the Cenozoic based on a variety of geological evidence, which helps to better understand our planet's future climate. Notably, extant biomes have recorded drastic environmental shifts. The climate in southern Asia, which hosts high biodiversity, is deeply impacted by the Asian monsoon. The origins and evolutionary dynamics of biomes occurring between the tropics and sub-tropics in southern Asia have probably been deeply impacted by climatic changes; however, these aspects remain poorly studied. We tested whether the evolutionary dynamics of the above biomes have recorded the drastic, late Cenozoic environmental shifts, by focusing on Magnolia section Michelia of the family Magnoliaceae. We established a fine time-calibrated phylogeny of M. section Michelia based on complete plastid genomes and inferred its ancestral ranges. Finally, we estimated the evolutionary dynamics of this section through time, determining its diversification rate and the dispersal events that occurred between tropical and sub-tropical areas. The tropical origin of M. section Michelia was dated to the late Oligocene; however, the diversification of its core group (i.e. M. section Michelia subsection Michelia) has occurred mainly from the late Miocene onward. Two key evolutionary shifts (dated approx. 8 and approx. 3 million years ago, respectively) were identified, each of them probably in response to drastic climatic changes. Here, we inferred the underlying evolutionary dynamics of biomes in southern Asia, which probably reflect late Cenozoic climatic changes. The occurrence of modern Asian monsoons was probably fundamental for the origin of M. section Michelia; moreover, the occurrence of asymmetric dispersal events between the tropics and sub-tropics hint at an adaptation strategy of M. section Michelia to global cooling, in agreement with the tropical conservatism hypothesis.

The adaptive challenge of extreme conditions shapes evolutionary diversity of plant assemblages at continental scales

The tropical conservatism hypothesis (TCH) posits that the latitudinal gradient in biological diversity arises because most extant clades of animals and plants originated when tropical environments were more widespread and because the colonization of colder and more seasonal temperate environments is limited by the phylogenetically conserved environmental tolerances of these tropical clades. Recent studies have claimed support of the TCH, indicating that temperate plant diversity stems from a few more recently derived lineages that are nested within tropical clades, with the colonization of the temperate zone being associated with key adaptations to survive colder temperatures and regular freezing. Drought, however, is an additional physiological stress that could shape diversity gradients. Here, we evaluate patterns of evolutionary diversity in plant assemblages spanning the full extent of climatic gradients in North and South America. We find that in both hemispheres, extratropical dry biomes house the lowest evolutionary diversity, while tropical moist forests and many temperate mixed forests harbor the highest. Together, our results support a more nuanced view of the TCH, with environments that are radically different from the ancestral niche of angiosperms having limited, phylogenetically clustered diversity relative to environments that show lower levels of deviation from this niche. Thus, we argue that ongoing expansion of arid environments is likely to entail higher loss of evolutionary diversity not just in the wet tropics but in many extratropical moist regions as well.

Global biogeography and diversification of a group of brown seaweeds (Phaeophyceae) driven by clade‐specific evolutionary processes

AbstractAimHistorical processes that shaped current diversity patterns of seaweeds remain poorly understood. Using Dictyotales, a globally distributed order of brown seaweeds as a model, we test if historical biogeographical and diversification patterns are comparable across clades. Dictyotales contain some 22 genera, three of which, Dictyota, Lobophora and Padina, are exceptionally diverse. Specifically, we test whether the evolutionary processes that shaped the latitudinal diversity patterns in these clades are in line with the tropical conservatism, out‐of‐the‐tropics or diversification rate hypotheses.LocationGlobal coastal benthic marine environments.TaxonDictyotales (Phaeophyceae).MethodsSpecies diversity was inferred using DNA‐based species delineation, addressing cryptic diversity and circumventing taxonomic problems. A six‐gene time‐calibrated phylogeny, distribution data of 3,755 specimens and probabilistic modelling of geographical range evolution were used to infer historical biogeographical patterns. The phylogeny was tested against different trait‐dependent models to compare diversification rates for different geographical units as well as different thermal affinities.ResultsOur results indicate that Dictyotales originated in the Middle Jurassic and reach a current peak of species diversity in the Central Indo‐Pacific. Ancestral range estimation points to a southern hemisphere origin of Dictyotales corresponding to the tropical southern Tethys Sea. Our results demonstrate that diversification rates were generally higher in tropical regions, but increased diversification rates in different clades are driven by different processes. Our results suggest that three major clades underwent a major diversification burst in the early Cenozoic, with Dictyota and Padina expanding their distribution into temperate regions while Lobophora retained a predominantly tropical niche.Main conclusionsOur results are consistent with both the tropical conservatism hypothesis, in which clades originate and remain in the tropics (Lobophora), and the out‐of‐the‐tropics scenario, where taxa originate and expand towards the temperate regions while preserving their presence in the tropics (Dictyota, Padina).

Ecological and evolutionary constraints on regional avifauna of passerines in China.

Strong correlations between species diversity and climate have been widely observed, but the mechanism underlying this relationship is unclear. Here, we explored the causes of the richness–climate relationships among passerine birds in China by integrating tropical conservatism and diversification rate hypotheses using path models. We found that assemblages with higher species richness southwest of the Salween–Mekong–Pearl River Divide are phylogenetically overdispersed and have shorter mean root distances (MRDs), while species-rich regions northeast of this divide (e.g., north Hengduan Mountains–south Qinling Mountains) are phylogenetically clustered and have longer MRDs. The results of the path analyses showed that the direct effect of climatic factors on species richness was stronger than their indirect effects on species richness via phylogenetic relatedness, indicating that neither tropical conservatism nor diversification rate hypotheses can well explain the richness–climate relationship among passerines in China. However, when path analyses were conducted within subregions separately, we found that the tropical conservatism hypothesis was well supported in the southwestern Salween–Mekong–Pearl River Divide, while the diversification rate hypothesis could explain the richness–climate relationship well in the northeastern divide. We conclude that the diversity patterns of passerines in different subregions of the Eastern Himalayas-Mountains of Southwest China may be shaped by different evolutionary processes related to geological and climatic histories, which explains why the tropical conservatism or diversification rate hypothesis alone cannot fully explain the richness–climate relationships.

Evolution of the latitudinal diversity gradient in the hyperdiverse ant genus Pheidole

AbstractAimThe latitudinal diversity gradient is the dominant geographic pattern of life on Earth, but a consensus understanding of its origins has remained elusive. The analysis of recently diverged, hyper‐rich invertebrate groups provides an opportunity to investigate latitudinal patterns with the statistical power of large trees while minimizing potentially confounding variation in ecology and history. Here, we synthesize global phylogenetic and macroecological data on a hyperdiverse (> 1,100 species) ant radiation, Pheidole and test predictions of three general explanations for the latitudinal gradient: variation in diversification rates, tropical conservatism and ecological regulation.LocationGlobal.Time periodThe past 35 million years.Major taxa studiedThe hyperdiverse ant genus Pheidole Westwood.MethodsWe assembled geographic data for 1,499 species and morphospecies, and inferred a dated phylogeny for 449 species of Pheidole, including 167 species newly sequenced for this study. We tested for correlations between diversification rate and latitude with Bayesian analysis of macroevolutionary mixtures (BAMM), hidden state speciation and extinction (HiSSE), geographic state speciation and extinction (GeoSSE), and a non‐parametric method (FiSSE), evaluated evidence for richness steady state, and examined patterns of diversification as Pheidole spread around the globe.ResultsThere was no evidence of systematic variation of net diversification rates with latitude across any of the methods. We found that Pheidole diversification occurred in bursts when new continents were colonized, followed by a slowdown in each region, but there is no evidence richness has saturated at an equilibrium in any region. Additionally, we found latitudinal affinity is moderately conserved with a Neotropical ancestor and simulations show that phylogenetic inertia alone is sufficient to produce the gradient pattern.Main conclusionsOur results provide no evidence that diversification rates vary systematically with latitude. Richness is far from steady state in each region, contrary to the ecological regulation hypothesis, although there is evidence that ecological opportunity promotes diversification after colonization of new areas. The fact that niche conservatism is strong enough to produce the gradient pattern is in accord with the tropical conservatism hypothesis. Overall, these results shed light on the mechanisms underlying the emergence of the diversity gradient within the past 34 million years, complementing recent work on deeper time‐scales, and more generally contribute toward much‐needed invertebrate perspective on global biodiversity dynamics.

Evolutionary origin of the latitudinal diversity gradient in liverworts

A latitudinal diversity gradient towards the tropics appears as one most recurrent patterns in ecology, but the mechanisms underlying this pattern remain an area of controversy. In angiosperms, the tropical conservatism hypothesis proposes that most groups originated in the tropics and are adapted to a tropical climatic regime, and that relatively few species have evolved physiological adaptations to cold, dry or unpredictable climates. This mechanism is, however, unlikely to apply across land plants, and in particular, to liverworts, a group of about 7500 species, whose ability to withstand cold much better than their tracheophyte counterparts is at odds with the tropical conservatism hypothesis. Molecular dating, diversification rate analyses and ancestral area reconstructions were employed to explore the evolutionary mechanisms that account for the latitudinal diversity gradient in liverworts. As opposed to angiosperms, tropical liverwort genera are not older than their extra-tropical counterparts (median stem age of tropical and extra-tropical liverwort genera of 24.35 ± 39.65 Ma and 39.57 ± 49.07 Ma, respectively), weakening the ‘time for speciation hypothesis’. Models of ancestral area reconstructions with equal migration rates between tropical and extra-tropical regions outperformed models with asymmetrical migration rates in either direction. The symmetry and intensity of migrations between tropical and extra-tropical regions suggested by the lack of resolution in ancestral area reconstructions towards the deepest nodes are at odds with the tropical niche conservatism hypothesis. In turn, tropical genera exhibited significantly higher net diversification rates than extra-tropical ones, suggesting that the observed latitudinal diversity gradient results from either higher extinction rates in extra-tropical lineages or higher speciation rates in the tropics. We discuss a series of experiments to help deciphering the underlying evolutionary mechanisms.

The macroevolution of climatic niches and its role in ant diversification

1. Investigating how climatic niches change over evolutionary timescales is a necessary step to understanding the current distribution of lineages, yet few studies have addressed this issue using comprehensive datasets. In this study, the evolution of ant climatic niches is investigated at a global scale based on bioclimatic data associated with 163 481 ant occurrence records. The resulting dataset was subjected to principal component analysis, and the scores obtained were used to characterise the main axes of ant climatic niche evolution.2. Principal component axis 1 (PC1) reflected variation in average temperature and seasonality – consistent with typical tropical/temperate gradients – whereas PC2 was associated with varying levels of aridity. Evolution along these two niche axes was markedly different: differences in the amount of explained variance between PC1 (65%) and PC2 (19%) suggest that climatic niche evolution was nearly three times more pronounced along a tropical–temperate climate axis.3. There was statistically significant phylogenetic signal on PC1, with genera occupying more tropical conditions diversifying at a faster rate, yet neither of these results is significant on PC2. In addition, most of the ancient ant lineages are associated with conditions of low seasonality and high temperatures.4. These results provide partial support for the tropical conservatism hypothesis as an explanation for geographical patterns of ant species richness.

Phylogenetic patterns in the geographic distributions of birds support the tropical conservatism hypothesis

AbstractAimRecent research suggests that the latitudinal diversity gradient (LDG) in birds is unlikely to result from faster diversification in the tropics. This puts the focus on other mechanisms, such as dispersal, as the primary drivers for the LDG. We aim to distinguish two prominent models, tropical conservatism (TCH) and out of the tropics (OTM), which make distinct predictions about dispersal across latitudes and the phylogenetic clustering of assemblages in temperate regions.LocationGlobal.MethodsWe used geographic and phylogenetic data for more than 9000 bird species to reconstruct the ancestral latitudinal zone for each node in each of 100 bird phylogenetic estimates. We used methods that consider discrete latitudinal zones, as well as those that consider latitudinal position as a continuous variable. We then estimated the frequency of ancestor–descendant dispersal within and across latitudinal zones. We also quantified phylogenetic clustering in latitudinal zones separately for bird assemblages within the Old World and the New World.ResultsLatitudinal distributions are relatively conserved: 60–96% of nodes had the same inferred latitude as their immediate ancestral node. We find that dispersal events out of the tropics were less frequent (4–5%) than dispersal events into the tropics (15–21%), the opposite of what would be expected under the OTM. Nodes with inferred temperate distributions are generally younger than the Eocene–Oligocene Climate Transition, as expected under the TCH. Phylogenetic clustering shows no regular patterns of association with latitude, and is likely to be largely driven by radiations within a few large forested biomes.Main conclusionsOur results provide support for the expectations of the TCH, but are less consistent with those of the OTM. Both the deeper origins of tropical clades and the comparatively recent but infrequent dispersal events into temperate regions appear to play a role in generating the strong disparity in tropical and temperate species richness in birds.

The Gondwanan legacy in South American biogeography

AbstractThe tropical conservatism hypothesis (TCH) suggests that phylogenetic niche conservatism (the tendency for traits to be maintained during diversification) should be the main driver of latitudinal diversity gradients. For example, the tropical–temperate diversity gradient for woody angiosperms should reflect the tropical, older origin of most clades, the constrained evolution of cold tolerance, and the fact that few clades were able to persist in temperate zones after the beginning of the global cooling in the Eocene. Evidence for this explanation has been discussed primarily from a Northern Hemisphere perspective. Recently, two studies have presented diversity and clade–age relationships in South American forests that are inconsistent with the TCH, with older woody families being more frequent at higher latitudes and higher elevations in the Andes. We argue that a broader framework considering the ancient history of the Southern Hemisphere flora and the recent history of the Andean flora can help explain these discordant patterns. Here, we provide an initial discussion of this new perspective, emphasizing the historical development of a unique and rich palaeoflora of Gondwanan ancestry at mid‐ to high latitudes of South America and Antarctica. We suggest that the idea of Austral niche conservatism (ANC) of a warm‐temperate Antarctic–South American biota should be explored further to improve our understanding of biogeographical patterns in the Southern Hemisphere.

Evolutionary histories of soil fungi are reflected in their large-scale biogeography.

Although fungal communities are known to vary along latitudinal gradients, mechanisms underlying this pattern are not well-understood. We used high-throughput sequencing to examine the large-scale distributions of soil fungi and their relation to evolutionary history. We tested the Tropical Conservatism Hypothesis, which predicts that ancestral fungal groups should be more restricted to tropical latitudes and conditions than would more recently derived groups. We found support for this hypothesis in that older phyla preferred significantly lower latitudes and warmer, wetter conditions than did younger phyla. Moreover, preferences for higher latitudes and lower precipitation levels were significantly phylogenetically conserved among the six younger phyla, possibly because the older phyla possess a zoospore stage that is vulnerable to drought, whereas the younger phyla retain protective cell walls throughout their life cycle. Our study provides novel evidence that the Tropical Conservatism Hypothesis applies to microbes as well as plants and animals.

The latitudinal species richness gradient in New World woody angiosperms is consistent with the tropical conservatism hypothesis.

Plant diversity, like that of most other taxonomic groups, peaks in the tropics, where climatic conditions are warm and wet, and it declines toward the temperate and polar zones as conditions become colder and drier, with more seasonally variable temperatures. Climate and evolutionary history are often considered competing explanations for the latitudinal gradient, but they are linked by the evolutionarily conserved environmental adaptations of species and the history of Earth's climate system. The tropical conservatism hypothesis (TCH) invokes niche conservatism, climatic limitations on establishment and survival, and paleoclimatic history to explain the latitudinal diversity gradient. Here, we use latitudinal distributions for over 12,500 woody angiosperm species, a fossil-calibrated supertree, and null modeling to test predictions of the TCH. Regional assemblages in the northern and southern temperate zones are less phylogenetically diverse than expected based on their species richness, because temperate taxa are clustered into relatively few clades. Moreover, lineages with temperate affinities are generally younger and nested within older, more tropical lineages. As predicted by the TCH, the vast majority of temperate lineages have arisen since global cooling began at the Eocene-Oligocene boundary (34 Mya). By linking physiological tolerances of species to evolutionary and biogeographic processes, phylogenetic niche conservatism may provide a theoretical framework for a generalized explanation for Earth's predominant pattern of biodiversity.

WHAT CAN MULTIPLE PHYLOGENIES SAY ABOUT THE LATITUDINAL DIVERSITY GRADIENT? A NEW LOOK AT THE TROPICAL CONSERVATISM, OUT OF THE TROPICS, AND DIVERSIFICATION RATE HYPOTHESES

We reviewed published phylogenies and selected 111 phylogenetic studies representing mammals, birds, insects, and flowering plants. We then mapped the latitudinal range of all taxa to test the relative importance of the tropical conservatism, out of the tropics, and diversification rate hypotheses in generating latitudinal diversity gradients. Most clades originated in the tropics, with diversity peaking in the zone of origin. Transitions of lineages between latitudinal zones occurred at 16-22% of the tree nodes. The most common type of transition was range expansions of tropical lineages to encompass also temperate latitudes. Thus, adaptation to new climatic conditions may not represent a major obstacle for many clades. These results contradict predictions of the tropical conservatism hypothesis (i.e., few clades colonizing extratropical latitudes), but support the out-of-the-tropics model (i.e., tropical originations and subsequent latitudinal range expansions). Our results suggest no difference in diversification between tropical and temperate sister lineages; thus, diversity of tropical clades was not explained by higher diversification rates in this zone. Moreover, lineages with latitudinal stasis diversified more compared to sister lineages entering a new latitudinal zone. This preserved preexisting diversity differences between latitudinal zones and can be considered a new mechanism for why diversity tends to peak in the zone of origin.

Tropical niche conservatism as a historical narrative hypothesis for the Neotropics: a case study using the fly family Muscidae

Aim We evaluate the extent to which the tropical conservatism hypothesis can explain the evolutionary development of the Muscidae. Furthermore, we compare the geographical patterns of muscid phylogenetic structure with biogeographical regions that have been identified for Neotropical insects. Location Central and South America. Methods We modelled the geographic distributions of 658 species using Maxent and 19 environmental variables. A generic-level supertree of the Muscidae was assembled using matrix representation with parsimony and used to map the geographic pattern of mean root distance (MRD), a metric of the relative evolutionary development of assemblages. Regression models (ordinary least squares and regression trees) were used to examine temperature and other environmental correlates of MRD to explore potential environmental drivers of muscid diversification. We used the regression tree results to recognize variable intervals that best explained MRD, and these intervals were mapped to recognize and compare with biogeographical regions of Neotropical insects. Results The geographic pattern of MRD was consistent with the tropical conservatism hypothesis: species in genera that diversified relatively early, as measured by their distance from the tree root, dominate lowland tropical South America, whereas species in genera that diversified more recently occupy extra-tropical areas, sub-Antarctic areas and the Andean highlands. Temperature was the strongest correlate of MRD. Three biogeographical regions were recognized and they coincided with two regions known for insects. Main conclusions Evolutionary responses of muscid flies to post-Eocene climate change taking the form of an expansion of a tropical group into regions with colder climates may be fundamental to explaining their distribution in the Neotropics. Our biogeographical regions delimited by temperature and the phylogenetic metric, surrogates of the tropical conservatism hypothesis, were very similar to general insect patterns, supporting the ‘tropical origin and evolutionary response to climate cooling’ as a broadly based historical narrative for the Neotropics.

Global angiosperm family richness revisited: linking ecology and evolution to climate

Aim The global richness gradient of angiosperm families is correlated with current climate, and it has been claimed that historical processes are not necessary to understand patterns of plant family richness. This claim has drawn criticism, and there have been doubts about the quality of the data used to quantify the pattern. We revisit this issue using the Angiosperm Phylogeny Group (APG) III classification and revised range maps, and we incorporate an evolutionary variable, family age, to explore covariation between evolution and ecology and their links to climate via the tropical conservatism hypothesis (TCH). Location Global. Methods The richness pattern for 408 families was derived from range maps, and family ages were derived from a dated angiosperm phylogeny. Patterns were generated for all families, 143 families composed of trees, and 149 families composed of herbs. We also examined family range size patterns to test the extent to which extratropical floras are nested subsets of tropical floras. Ordinary least squares (OLS) multiple and partial regressions were used to generate climate models for richness, mean range size and mean age for each plant dataset and to evaluate the covariation between contemporary climate and clade age as correlates of family richness. Results We confirmed the strong association between contemporary climate and family richness. Age patterns predicted by TCH were also found for families comprising trees. The richness of herbaceous families, in contrast, was correlated with climate but the age pattern was not as predicted by TCH. Floras in cold and dry areas are strongly nested within richer tropical floras. Main conclusions Phylogenetic niche conservatism at the family level offers a likely explanation for the global diversity gradient of trees, but not for non-desert herbs, probably because of the faster evolutionary rates for herbs and less constrained evolutionary responses to climate change. Thus, it appears that multiple processes account for the overall angiosperm family gradient. Our analysis also demonstrates that even very strong associations of taxon richness and climate do not preclude evolutionary processes, as has been widely argued, and that climatic and evolutionary hypotheses for richness gradients are not mutually exclusive.

Energy and the tempo of evolution in amphibians

ABSTRACTAim The evolutionary speed hypothesis (ESH) attempts to explain global patterns of species richness on the basis that rates of molecular evolution and speciation in warmer climates have led to a greater accumulation of taxa at lower latitudes. A substantial alternative hypothesis to the ESH is the tropical conservatism hypothesis (TCH). However, recent tests of the TCH, using amphibians as the model taxon, have relied on the assumption that rates of molecular evolution are stable across latitudes and elevations. Here, we test for the first time for systematic variation in rates of molecular evolution across latitude and elevation among amphibians.Location The dataset is geographically diverse with samples from all continents except Antarctica and also from many of the earth's major tropical–warm temperate archipelagos.Methods We tested for substitution rate heterogeneity across climatically varying habitats with the mitochondrial RNA genes 12S and 16S. Thus, we report here on our findings for amphibians – a taxon whose phylogenetic and trophic contexts are remote from those previously tested – using genes that have also not been examined before. The study utilized paired contrasts of sister species (188 species across 18 families, including both caudates and anurans) that are spatially separated in either latitudinal or elevational dimensions.Results We found substantially faster substitution rates for species living in warmer habitats (P= 0.001–0.002) at both lower latitudes (P < 0.02) and lower elevations (P < 0.01).Main conclusions The consistency of these results with the previous studies that used quite different organisms – and in this instance also using different genes – suggests that this is a ubiquitous pattern in nature consistent with the predictions of the ESH. Recent tests of the TCH that, in estimating diversification rates, have relied on the assumption that DNA evolution occurs at a constant rate across latitudes and elevations, require reconsideration in light of the findings presented here. Our results indicate that greater caution is required when estimating dates of divergence using DNA sequence data.

Tropical niche conservatism and the species richness gradient of North American butterflies

AbstractAim We explore the potential role of the ‘tropical conservatism hypothesis’ in explaining the butterfly species richness gradient in North America. Its applicability can be derived from the tropical origin of butterflies and the presumed difficulties in evolving the cold tolerance required to permit the colonization and permanent occupation of the temperate zone.Location North America.Methods Digitized range maps for butterfly species north of Mexico were used to map richness for all species, species with distributions north of the Tropic of Capricorn (Extratropicals), and species that also occupy the tropics (Tropicals). A phylogeny resolved to subfamily was used to map the geographical pattern of mean root distance, a metric of the evolutionary development of assemblages. Regression models and general linear models examined environmental correlates of overall richness and for Extratropicals vs. Tropicals, patterns in summer vs. winter, and patterns in northern vs. southern North America.Results Species in more basal subfamilies dominate the south, whereas more derived clades occupy the north. There is also a ‘latitudinal’ richness gradient in Canada/Alaska, whereas in the conterminous USA richness primarily varies longitudinally. Overall richness is associated with broad‐ and mesoscale temperature gradients. The richness of Tropicals is strongly associated with temperature and distance from winter population sources. The richness of Extratropicals in the north is most strongly correlated with the pattern of glacial retreat since the more recent Ice Age, whereas in the south, richness is positively associated with the range of temperatures in mountains and the presence of forests but is negatively correlated with the broad‐scale temperature gradient.Main conclusions The tropical conservatism hypothesis provides a possible explanation for the complex structure of the species richness gradient. The Canada/Alaska fauna comprises temperate, boreal and tundra species that are nevertheless constrained by cold climates and limited vegetation, coupled with possible post‐Pleistocene recolonization lags. In the USA tropical species are constrained by temperature in winter as well as recolonization distances in summer, whereas temperate‐zone groups are richer in cooler climates in mountains and forests, where winter conditions are more suitable for diapause. The evolution of cold tolerance is key to both the evolutionary and ecological patterns.

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 constraints on regional faunas: whom, but not how many

The latitudinal diversity gradient has been hypothesized to reflect past evolutionary dynamics driven by climatic niche conservation during cladogenesis, i.e. the tropical conservatism hypothesis. Here we show that the species diversity of treefrogs (Hylidae) across the western hemisphere is actually independent of evolutionary niche dynamics. We evaluated three key predictions of the tropical conservatism hypothesis that relate to the relationships between climate, species richness and the phylogenetic structure of regional treefrog faunas across the continental Americas. Species composition was dependent on the inability of some lineages to evolve cold tolerance, but the actual number of species in a region was strongly predicted by precipitation, not temperature. Moreover, phylogenetic structure was independent of precipitation. Thus, species in low-richness areas were no more closely related than species in highly diverse regions. These results provide no support for the tropical conservatism hypothesis. Instead, they show that regional species composition and richness are constrained by different climatic components, demonstrating that global biodiversity gradients can be independent of niche stasis during cladogenesis.