Daniel S. Moen

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.

Why does diversification slow down

Studies of phylogenetic diversification often show evidence for slowdowns in diversification rates over the history of clades. Recent studies seeking biological explanations for this pattern have emphasized the role of niche differentiation, as in hypotheses of adaptive radiation and ecological limits to diversity. Yet many other biological explanations might underlie diversification slowdowns. In this paper, we focus on the geographic context of diversification, environment-driven bursts of speciation, failure of clades to keep pace with a changing environment, and protracted speciation. We argue that, despite being currently underemphasized, these alternatives represent biologically plausible explanations that should be considered along with niche differentiation. Testing the importance of these alternative hypotheses might yield fundamentally different explanations for what influences species richness within clades through time.

Phylogenetic origins of local-scale diversity patterns and the causes of Amazonian megadiversity

What explains the striking variation in local species richness across the globe and the remarkable diversity of rainforest sites in Amazonia? Here, we apply a novel phylogenetic approach to these questions, using treefrogs (Hylidae) as a model system. Hylids show dramatic variation in local richness globally and incredible local diversity in Amazonia. We find that variation in local richness is not explained primarily by climatic factors, rates of diversification (speciation and extinction) nor morphological variation. Instead, local richness patterns are explained predominantly by the timing of colonization of each region, and Amazonian megadiversity is linked to the long-term sympatry of multiple clades in that region. Our results also suggest intriguing interactions between clade diversification, trait evolution and the accumulation of local richness. Specifically, sympatry between clades seems to slow diversification and trait evolution, but prevents neither the accumulation of local richness over time nor the co-occurrence of similar species.

An expanded phylogeny of treefrogs (Hylidae) based on nuclear and mitochondrial sequence data

The treefrogs (Hylidae) make up one of the most species-rich families of amphibians. With 885 species currently described, they contain >13% of all amphibian species. In recent years, there has been considerable progress in resolving hylid phylogeny. However, the most comprehensive phylogeny to date (Wiens et al., 2006) included only 292 species, was based only on parsimony, provided only poor support for most higher-level relationships, and conflicted with previous hypotheses in several parts (including the monophyly and relationships of major clades of Hylinae). Here, we present an expanded phylogeny for hylid frogs, including data for 362 hylid taxa for up to 11 genes (4 mitochondrial, 7 nuclear), including 70 additional taxa and >270 sequences not included in the previously most comprehensive analysis. The new tree from maximum likelihood analysis is more well-resolved, strongly supported, and concordant with previous hypotheses, and provides a framework for future systematic, biogeographic, ecological, and evolutionary studies.

PHYLOGENETIC EVIDENCE FOR COMPETITIVELY DRIVEN DIVERGENCE: BODY‐SIZE EVOLUTION IN CARIBBEAN TREEFROGS (HYLIDAE: OSTEOPILUS)

Understanding the role of competition in explaining phenotypic diversity is a challenging problem, given that the most divergent species may no longer compete today. However, convergent evolution of extreme body sizes across communities may offer evidence of past competition. For example, many treefrog assemblages around the world have convergently evolved species with very large and small body sizes. To better understand this global pattern, we studied body-size diversification within the small, endemic radiation of Caribbean treefrogs (Osteopilus). We introduce a suite of analyses designed to help reveal the signature of past competition. Diet analyses show that Osteopilus are generalist predators and that prey size is strongly associated with body size, suggesting that body-size divergence facilitates resource partitioning. Community assembly models indicate that treefrog body-size distributions in Jamaica and Hispaniola are consistent with expectations from competition. Phylogenetic analyses show that similar body-size extremes in Jamaica and Hispaniola have originated through parallel evolution on each island, and the rate of body-size evolution in Osteopilus is accelerated relative to mainland treefrogs. Together, these results suggest that competition may have driven the rapid diversification of body sizes in Caribbean treefrogs to the extremes seen in treefrog communities around the world.

Evolutionary conservatism and convergence both lead to striking similarity in ecology, morphology and performance across continents in frogs

Many clades contain ecologically and phenotypically similar species across continents, yet the processes generating this similarity are largely unstudied, leaving fundamental questions unanswered. Is similarity in morphology and performance across assemblages caused by evolutionary convergence or by biogeographic dispersal of evolutionarily conserved ecotypes? Does convergence to new ecological conditions erase evidence of past adaptation? Here, we analyse ecology, morphology and performance in frog assemblages from three continents (Asia, Australia and South America), assessing the importance of dispersal and convergent evolution in explaining similarity across regions. We find three striking results. First, species using the same microhabitat type are highly similar in morphology and performance across both clades and continents. Second, some species on different continents owe their similarity to dispersal and evolutionary conservatism (rather than evolutionary convergence), even over vast temporal and spatial scales. Third, in one case, an ecologically specialized ancestor radiated into diverse ecotypes that have converged with those on other continents, largely erasing traces of past adaptation to their ancestral ecology. Overall, our study highlights the roles of both evolutionary conservatism and convergence in explaining similarity in species traits over large spatial and temporal scales and demonstrates a statistical framework for addressing these questions in other systems.

Community Assembly Through Evolutionary Diversification and Dispersal in Middle American Treefrogs

How are ecologically diverse organisms added to local assemblages to create the community structure we see today? In general, within a given region or community, a given trait (character state) may either evolve in situ or be added through dispersal after having evolved elsewhere. Here, we develop simple metrics to quantify the relative importance of these processes and then apply them to a case study in Middle American treefrogs. We examined two ecologically important characters (larval habitat and body size) among 39 communities, using phylogenetic and ecological information from 278 species both inside and outside the region. For each character, variation among communities reflects complex patterns of evolution and dispersal. Our results support several general hypotheses about community assembly, which may apply to many other systems: (1) elevation can play an important role in creating patterns of community structure within a region, (2) contrary to expectations, species can invade communities in which species with similar ecological traits are already present, (3) dispersal events tend to occur between areas with similar climatic regimes, and (4) the first lineage to invade a region diversifies the most ecologically, whereas later invasions show limited change.

Cope's rule in cryptodiran turtles: do the body sizes of extant species reflect a trend of phyletic size increase?

Copes rule of phyletic size increase is questioned as a general pattern of body size evolution. Most studies of Copes rule have examined trends in the paleontological record. However, neontological approaches are now possible due to the development of model‐based comparative methods, as well as the availability of an abundance of phylogenetic data. I examined whether the phylogenetic distribution of body sizes in extant cryptodiran turtles is consistent with Copes rule. To do this, I examined body size evolution in each of six major clades of cryptodiran turtles and also across the whole tree of cryptodirans (n = 201 taxa). Extant cryptodiran turtles do not appear to follow Copes rule, as no clade showed a significant phyletic body size trend. Previous analyses in other extant vertebrates have also found no evidence for phyletic size increase, which is in contrast to the paleontological data that support the rule in a number of extinct vertebrate taxa.

Testing Convergence Versus History: Convergence Dominates Phenotypic Evolution for over 150 Million Years in Frogs.

Striking evolutionary convergence can lead to similar sets of species in different locations, such as in cichlid fishes and Anolis lizards, and suggests that evolution can be repeatable and predictable across clades. Yet, most examples of convergence involve relatively small temporal and/or spatial scales. Some authors have speculated that at larger scales (e.g., across continents), differing evolutionary histories will prevent convergence. However, few studies have compared the contrasting roles of convergence and history, and none have done so at large scales. Here we develop a two-part approach to test the scale over which convergence can occur, comparing the relative importance of convergence and history in macroevolution using phylogenetic models of adaptive evolution. We apply this approach to data from morphology, ecology, and phylogeny from 167 species of anuran amphibians (frogs) from 10 local sites across the world, spanning ~160 myr of evolution. Mapping ecology on the phylogeny revealed that similar microhabitat specialists (e.g., aquatic, arboreal) have evolved repeatedly across clades and regions, producing many evolutionary replicates for testing for morphological convergence. By comparing morphological optima for clades and microhabitat types (our first test), we find that convergence associated with microhabitat use dominates frog morphological evolution, producing recurrent ecomorphs that together encompass all sampled species in each community in each region. However, our second test, which examines whether and how much species differ from their inferred optima, shows that convergence is incomplete: that is, phenotypes of most species are still somewhat distant from the estimated optimum for each microhabitat, seemingly because of insufficient time for more complete adaptation (an effect of history). Yet, these effects of history are related to past ecologies, and not clade membership. Overall, our study elucidates the dominant drivers of morphological evolution across a major vertebrate clade and shows that evolution can be repeatable at much greater temporal and spatial scales than commonly thought. It also provides an analytical framework for testing other potential examples of large-scale convergence.

Microhabitat and Climatic Niche Change Explain Patterns of Diversification among Frog Families

A major goal of ecology and evolutionary biology is to explain patterns of species richness among clades. Differences in rates of net diversification (speciation minus extinction over time) may often explain these patterns, but the factors that drive variation in diversification rates remain uncertain. Three important candidates are climatic niche position (e.g., whether clades are primarily temperate or tropical), rates of climatic niche change among species within clades, and microhabitat (e.g., aquatic, terrestrial, arboreal). The first two factors have been tested separately in several studies, but the relative importance of all three is largely unknown. Here we explore the correlates of diversification among families of frogs, which collectively represent ∼88% of amphibian species. We assemble and analyze data on phylogeny, climate, and microhabitat for thousands of species. We find that the best-fitting phylogenetic multiple regression model includes all three types of variables: microhabitat, rates of climatic niche change, and climatic niche position. This model explains 67% of the variation in diversification rates among frog families, with arboreal microhabitat explaining ∼31%, niche rates ∼25%, and climatic niche position ∼11%. Surprisingly, we show that microhabitat can have a much stronger influence on diversification than climatic niche position or rates of climatic niche change.