Terrestrialization, Miniaturization and Rates of Diversification in African Puddle Frogs (Anura: Phrynobatrachidae)

Terrestrialization, the evolution of non-aquatic oviposition, and miniaturization, the evolution of tiny adult body size, are recurring trends in amphibian evolution, but the relationships among the traits that characterize these phenomena are not well understood. Furthermore, these traits have been identified as possible “key innovations” that are predicted to increase rates of speciation in those lineages in which they evolve. We examine terrestrialization and miniaturization in sub-Saharan puddle frogs (Phrynobatrachidae) in a phylogenetic context to investigate the relationship between adaptation and diversification through time. We use relative dating techniques to ascertain if character trait shifts are associated with increased diversification rates, and we evaluate the likelihood that a single temporal event can explain the evolution of those traits. Results indicate alternate reproductive modes evolved independently in Phrynobatrachus at least seven times, including terrestrial deposition of eggs and terrestrial, non-feeding larvae. These shifts towards alternate reproductive modes are not linked to a common temporal event. Contrary to the “key innovations” hypothesis, clades that exhibit alternate reproductive modes have lower diversification rates than those that deposit eggs aquatically. Adult habitat, pedal webbing and body size have no effect on diversification rates. Though these traits putatively identified as key innovations for Phrynobatrachus do not seem to be associated with increased speciation rates, they may still provide opportunities to extend into new niches, thus increasing overall diversity

Biogeography of species richness gradients:Linking adaptive traits, demography and diversification

Here we review how adaptive traits contribute to the emergence and maintenance of species richness gradients through their influence on demographic and diversification processes. We start by reviewing how demographic dynamics change along species richness gradients. Empirical studies show that geographical clines in population parameters and measures of demographic variability are frequent along latitudinal and altitudinal gradients. Demographic variability often increases at the extremes of regional species richness gradients and contributes to shape these gradients. Available studies suggest that adaptive traits significantly influence demographic dynamics, and set the limits of species distributions. Traits related to thermal tolerance, resource use, phenology and dispersal seem to play a significant role. For many traits affecting demography and/or diversification processes, complex mechanistic approaches linking genotype, phenotype and fitness are becoming progressively available. In several taxa, species can be distributed along adaptive trait continuums, i.e. a main axis accounting for the bulk of inter-specific variation in some correlated adaptive traits. It is shown that adaptive trait continuums can provide useful mechanistic frameworks to explain demographic dynamics and diversification in species richness gradients. Finally, we review the existence of sequences of adaptive traits in phylogenies, the interactions of adaptive traits and community context, the clinal variation of traits across geographical gradients, and the role of adaptive traits in determining the history of dispersal and diversification of clades. Overall, we show that the study of demographic and evolutionary mechanisms that shape species richness gradients clearly requires the explicit consideration of adaptive traits. To conclude, future research lines and trends in the field are briefly outlined

Integrating Environmental, Molecular, and Morphological Data to Unravel an Ice-age Radiation of Arctic-alpine Campanula in Western North America

Many arctic-alpine plant genera have undergone speciation during the Quaternary. The bases for these radiations have been ascribed to geographic isolation,abiotic and biotic differences between populations, and/or hybridization andpolyploidization. The Cordilleran Campanula L. (Campanulaceae Juss.), a monophyletic clade of mostly endemic arctic-alpine taxa from western North America, experienced a recent and rapid radiation. We set out to unravel the factors that likely influenced speciation in this group. To do so, we integrated environmental, genetic, and morphological datasets, tested biogeographic hypotheses, and analyzed the potential consequences of the various factors on the evolutionary history of the clade. We created paleodistribution models to identify potential Pleistocene refugia for the clade and estimated niche space for individual taxa using geographic and climatic data. Using 11 nuclear loci, we reconstructed a species tree and tested biogeographic hypotheses derived from the paleodistribution models. Finally, we tested 28 morphological characters, including floral, vegetative, and seed characteristics, for their capacity to differ- entiate taxa. Our results show that the combined effect of Quaternary climatic variation, isolation among differing environments in the mountains in western North America, and biotic factors influencing floral morphology contributed to speciation in this group during the mid-Pleistocene. Furthermore, our biogeographic analyses uncovered asynchronous consequences of interglacial and glacial periods for the timing of refugial isolation within the southern and northwestern mountains, respectively. These findings have broad implications for understanding the processes promoting speciation in arctic-alpine plants and the rise of numerous endemic taxa across the region

Investigating Dispersal Ability to Infer Diversification in the Birds of Madagascar

The objective of this study was to investigate whether dispersal ability, as measured by a proxy of hand-wing index 2, influenced diversification of the birds of Madagascar at two scales. Madagascar is home to several avian lineages that have diversified greatly while other lineages are only represented by a single species. a key question in evolutionary biology is why some of these lineages diversified while others did not. One way to address this is to examine what features of these lineages promotes their diversification. Recent studies have focused on the relative importance of dispersal ability to diversification at the continental and island scales. to further test this relationship, I investigated whether dispersal ability, as measured by hand-wing index 2, influenced diversification in the birds of Madagascar at a large regional scale and smaller local scale within Madagascar. to assess whether dispersal ability influenced diversification of the birds of Madagascar I compared hand-wing index 2 of Malagasy and source (closest non-Malagasy relatives) clades of five radiating and three non-radiating lineages. I treated each lineage as an independent case study in the hopes of identifying a pattern reflecting a shift in dispersal ability upon colonization of Madagascar in radiating lineages. at a smaller local scale of macrohabitats within Madagascar, I examined whether variation in dispersal ability (HWI2) within widespread Malagasy species differed between subspecies restricted to macrohabitats reflecting local adaptation and divergence across macrohabitats. My study found that Malagasy species did not shift in their dispersal ability after colonizing Madagascar and thus, dispersal ability is not critical to the diversification of Malagasy endemics from their source clade in radiating or non-radiating lineages. However, at a smaller local scale of macrohabitats, I found variation in dispersal ability was likely due to local adaptations to macrohabitats. This study adds to our knowledge of dispersal ability and diversification patterns in Malagasy avifauna. This is a leading step towards additional studies to investigate the impact of potential geographic barriers to dispersal ability in the birds of Madagascar and could provide further insights into diversification patterns

The role of visual adaptation in cichlid fish speciation

D. Shane Wright (1) , Ole Seehausen (2), Ton G.G. Groothuis (1), Martine E. Maan (1) (1) University of Groningen; GELIFES; EGDB(2) Department of Fish Ecology & Evolution, EAWAG Centre for Ecology, Evolution and Biogeochemistry, Kastanienbaum AND Institute of Ecology and Evolution, Aquatic Ecology, University of Bern.In less than 15,000 years, Lake Victoria cichlid fishes have radiated into as many as 500 different species. Ecological and sexual sel ection are thought to contribute to this ongoing speciation process, but genetic differentiation remains low. However, recent work in visual pigment genes, opsins, has shown more diversity. Unlike neighboring Lakes Malawi and Tanganyika, Lake Victoria is highly turbid, resulting in a long wavelength shift in the light spectrum with increasing depth, providing an environmental gradient for exploring divergent coevolution in sensory systems and colour signals via sensory drive. Pundamilia pundamila and Pundamilia nyererei are two sympatric species found at rocky islands across southern portions of Lake Victoria, differing in male colouration and the depth they reside. Previous work has shown species differentiation in colour discrimination, corresponding to divergent female preferences for conspecific male colouration. A mechanistic link between colour vision and preference would provide a rapid route to reproductive isolation between divergently adapting populations. This link is tested by experimental manip ulation of colour vision - raising both species and their hybrids under light conditions mimicking shallow and deep habitats. We quantify the expression of retinal opsins and test behaviours important for speciation: mate choice, habitat preference, and fo raging performance

The Roles of Ecological Opportunity and Incumbency Effects in the Macroevolution of the Luzon Island, Philippines “Old Endemic” Murine Rodents

University of Minnesota Ph.D. dissertation. May 2019. Major: Ecology, Evolution and Behavior. Advisor: Sharon Jansa. 1 computer file (PDF); xiv, 216 pages.The evolutionary theory of adaptive radiation posits that lineages that experience a breadth of available resources in the absence of competition, known as ecological opportunity, should diversify to specialize on aspects of these available resources. The rapid evolution decelerates as niches fill in a static, spatially limited system, resulting in an assemblage of ecologically distinct species. Despite evidence to support this mode of diversification, little attention has been given to how this process unfolds in systems with multiple, ecologically similar colonizing lineages. The primary-colonizing, or incumbent, lineage, through exploiting niches in the absence of competitors, may serve to depress the rates and patterns of species and ecological diversity of subsequent colonists. In this dissertation, I explored four aims that seek to test whether the evolution of two clades of rodents endemic to Luzon Island, Philippines, Chrotomyini and Phloeomyini, exhibited evolution consistent with incumbency effects held by Phloeomyini and placed on secondary-colonizing Chrotomyini. First, I determined whether the rates of lineage diversification of the two Luzon Old Endemic (LOE) clades were consistent with reduced ecological opportunity in secondarily-colonizing Chrotomyini, resulting in lower rates of species accumulation. My results instead indicate that Chrotomyini has experienced a faster rate of diversification inconsistent with incumbency effects. Second, I tested whether the mandible of the LOE rodents, as a proxy for diet, exhibits rates of evolution consistent with lower ecological opportunity for Chrotomyini as well as patterns of diversity consistent with clade-specific partitioning of morphological variation. I found that both LOE clades evolved disparate mandible shapes at a similar rate, apart from outlying genus Rhynchomys, but that the two clades occupy nearly discrete areas of morphospace. Third, I tested whether the shape of the humerus can be used to approximate locomotory niche in a similar way to linear measurements of the ulna, metacarpal, and phalanx, to determine whether the morphology associated with locomotory strategy in the two LOE clades is convergent on shared locomotory mode. I found that although the humerus predicts some aspects of locomotory strategy, a substantial proportion of shape variation is reflected by different adaptations within shared locomotory category, thus providing a complement to, rather than replacement for, distal forelimb measurements. Finally, I tested whether the observed lack of mandibular shape overlap between the two LOE clades is consistent Chrotomyini being limited by Phloeomyini in terms of the area of morphospace it could diversify into and whether the ancestral chrotomyine lineage may have exhibited morphology disparate from Phloeomyini, thus facilitating its colonization and subsequent diversification. I found that the patterns of mandibular shape variation in the two LOE clades are consistent with the establishment of a biotic filter, meaning that Chrotomyini’s success on Luzon was facilitated by persistent ecological distinction from incumbent Phloeomyini. This dissertation illustrates that subfamily-related clades can experience substantial ecological distinction both within and between each clade. This distinction can permit repeated colonization of spatially constrained systems: as long as each colonizing clade remains ecologically distinct, evolution may proceed uninhibited by inter-clade competitive effects. Incumbency effects may thus more strongly influence the community assembly of species in a system than their evolutionary rates