Trait decoupling promotes evolutionary diversification of the trophic and acoustic system of damselfishes.

Trait decoupling, wherein evolutionary release of constraints permits specialization of formerly integrated structures, represents a major conceptual framework for interpreting patterns of organismal diversity. However, few empirical tests of this hypothesis exist. A central prediction, that the tempo of morphological evolution and ecological diversification should increase following decoupling events, remains inadequately tested. In damselfishes (Pomacentridae), a ceratomandibular ligament links the hyoid bar and lower jaws, coupling two main morphofunctional units directly involved in both feeding and sound production. Here, we test the decoupling hypothesis by examining the evolutionary consequences of the loss of the ceratomandibular ligament in multiple damselfish lineages. As predicted, we find that rates of morphological evolution of trophic structures increased following the loss of the ligament. However, this increase in evolutionary rate is not associated with an increase in trophic breadth, but rather with morphofunctional specialization for the capture of zooplanktonic prey. Lineages lacking the ceratomandibular ligament also shows different acoustic signals (i.e. higher variation of pulse periods) from others, resulting in an increase of the acoustic diversity across the family. Our results support the idea that trait decoupling can increase morphological and behavioural diversity through increased specialization rather than the generation of novel ecotypes.

Using Pareto optimality to explore the topology and dynamics of the human connectome.

Graph theory has provided a key mathematical framework to analyse the architecture of human brain networks. This architecture embodies an inherently complex relationship between connection topology, the spatial arrangement of network elements, and the resulting network cost and functional performance. An exploration of these interacting factors and driving forces may reveal salient network features that are critically important for shaping and constraining the brain's topological organization and its evolvability. Several studies have pointed to an economic balance between network cost and network efficiency with networks organized in an 'economical' small-world favouring high communication efficiency at a low wiring cost. In this study, we define and explore a network morphospace in order to characterize different aspects of communication efficiency in human brain networks. Using a multi-objective evolutionary approach that approximates a Pareto-optimal set within the morphospace, we investigate the capacity of anatomical brain networks to evolve towards topologies that exhibit optimal information processing features while preserving network cost. This approach allows us to investigate network topologies that emerge under specific selection pressures, thus providing some insight into the selectional forces that may have shaped the network architecture of existing human brains.

The Evolution of Modularity in the Mammalian Skull I: Morphological Integration Patterns and Magnitudes

Morphological integration refers to the modular structuring of inter-trait relationships in an organism, which could bias the direction and rate of morphological change, either constraining or facilitating evolution along certain dimensions of the morphospace. Therefore, the description of patterns and magnitudes of morphological integration and the analysis of their evolutionary consequences are central to understand the evolution of complex traits. Here we analyze morphological integration in the skull of several mammalian orders, addressing the following questions: are there common patterns of inter-trait relationships? Are these patterns compatible with hypotheses based on shared development and function? Do morphological integration patterns and magnitudes vary in the same way across groups? We digitized more than 3,500 specimens spanning 15 mammalian orders, estimated the correspondent pooled within-group correlation and variance/covariance matrices for 35 skull traits and compared those matrices among the orders. We also compared observed patterns of integration to theoretical expectations based on common development and function. Our results point to a largely shared pattern of inter-trait correlations, implying that mammalian skull diversity has been produced upon a common covariance structure that remained similar for at least 65 million years. Comparisons with a rodent genetic variance/covariance matrix suggest that this broad similarity extends also to the genetic factors underlying phenotypic variation. In contrast to the relative constancy of inter-trait correlation/covariance patterns, magnitudes varied markedly across groups. Several morphological modules hypothesized from shared development and function were detected in the mammalian taxa studied. Our data provide evidence that mammalian skull evolution can be viewed as a history of inter-module parcellation, with the modules themselves being more clearly marked in those lineages with lower overall magnitude of integration. The implication of these findings is that the main evolutionary trend in the mammalian skull was one of decreasing the constraints to evolution by promoting a more modular architecture.

The evolution of behavioural systems: a study of grooming in rodents

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Morphological patterns and community organization in leaf-litter ant assemblages

We present a new approach to determine the number and composition of guilds, using the hyperdiverse leaf-litter ant fauna as a model, based on appropriate morphological variables and species co-occurrence null models to describe the complex assemblages of interacting Species Community structure at the 1-m(2) scale. We obtained 18 linear morphometric measures from 949 workers of 171 leaf-litter ant species (18762 measurements) surveyed in four Atlantic Forest localities to test whether the assemblages are morphologically structured; the morphological characters were selected to indicate diet and foraging habits. Principal components analysis was used to characterize the morphospace and to describe the guild structure (number of species and composition). The guild proportionality assembly rule (significant tendency toward constant proportion of species in guilds) was assessed at the 1-m(2) scale. Our analysis indicates that the division of leaf-litter ants into guilds is based mainly on microhabitat distribution in the leaf-litter, body size and shape, eye size, and phylogeny. The same guild scheme applied to four more sites shows that different Atlantic Forest areas have the same leaf-fitter ant guilds. The guild proportionality assembly rule was confirmed for most guilds, Suggesting that there are guild-specific limitations on species coexistence within assemblages; on the other hand, in a few cases the variance in guild proportion was greater than expected under the null assumptions. Other studies on ant functional group classification are partially supported by our quantitative morphological analysis. Our results, however, imply that there are more compartments than indicated in previous models, particularly among cryptic species (confined to soil and litter) and tropical climate specialists. We argue that a general null model for the analysis of species association based oil morphology can reveal objectively defined groups and may thus contribute to a robust theory to explain community structure in general and have important consequences on studies of litter ant community ecology in particular.

Biomechanics on the Half Shell: Functional Performance Influences Patterns of Morphological Variation in the Emydid Turtle carapace

This study uses the carapace of emydid turtles to address hypothesized differences between terrestrial and aquatic species. Geometric morphometrics are used to quantify shell shape, and performance is estimated for two shell functions: shell strength and hydrodynamics. Aquatic turtle shells differ in shape from terrestrial turtle shells and are characterized by lower frontal areas and presumably lower drag. Terrestrial turtle shells are stronger than those of aquatic turtles; many-to-one mapping of morphology to function does not entirely mitigate a functional trade-off between mechanical strength and hydrodynamic performance. Furthermore, areas of morphospace characterized by exceptionally poor performance in either of the functions are not occupied by any emydid species. Though aquatic and terrestrial species show no significant differences in the rate of morphological evolution, aquatic species show a higher lineage density, indicative of a greater amount of convergence in their evolutionary history. The techniques employed in this study, including the modeling of theoretical shapes to assess performance in unoccupied areas of morphospace, suggest a framework for future studies of morphological variation.

Morphological Diversity and the Roles of Contingency, Chance and Determinism in African Cichlid Radiations

Background: Deterministic evolution, phylogenetic contingency and evolutionary chance each can influence patterns of morphological diversification during adaptive radiation. In comparative studies of replicate radiations, convergence in a common morphospace implicates determinism, whereas non-convergence suggests the importance of contingency or chance. Methodology/Principal Findings: The endemic cichlid fish assemblages of the three African great lakes have evolved similar sets of ecomorphs but show evidence of non-convergence when compared in a common morphospace, suggesting the importance of contingency and/or chance. We then analyzed the morphological diversity of each assemblage independently and compared their axes of diversification in the unconstrained global morphospace. We find that despite differences in phylogenetic composition, invasion history, and ecological setting, the three assemblages are diversifying along parallel axes through morphospace and have nearly identical variance-covariance structures among morphological elements. Conclusions/Significance: By demonstrating that replicate adaptive radiations are diverging along parallel axes, we have shown that non-convergence in the common morphospace is associated with convergence in the global morphospace. Applying these complimentary analyses to future comparative studies will improve our understanding of the relationship between morphological convergence and non-convergence, and the roles of contingency, chance and determinism in driving morphological diversification.

Relaxed trait covariance in interspecific cichlid hybrids predicts morphological diversity in adaptive radiations

The process of adaptive radiation involves multiple events of speciation in short succession, associated with ecological diversification. Understanding this process requires identifying the origins of heritable phenotypic variation that allows adaptive radiation to progress. Hybridization is one source of genetic and morphological variation that may spur adaptive radiation. We experimentally explored the potential role of hybridization in facilitating the onset of adaptive radiation. We generated first- and second-generation hybrids of four species of African cichlid fish, extant relatives of the putative ancestors of the adaptive radiations of Lakes Victoria and Malawi. We com- pared patterns in hybrid morphological variation with the variation in the lake radiations. We show that significant fractions of the interspecific mor- phological variation and the major trajectories in morphospace that charac- terize whole radiations can be generated in second-generation hybrids. Furthermore, we show that covariation between traits is relaxed in second- generation hybrids, which may facilitate adaptive diversification. These results support the idea that hybridization can provide the heritable pheno- typic diversity necessary to initiate adaptive radiation.

Phenotypic evolution in a fossil gastropod species lineage: evidence for adaptive radiation?

Detecting speciation in the fossil record is a particularly challenging matter. Palaeontologists are usually confronted with poor preservation and limited knowledge on the palaeoenvironment. Even in the contrary case of adequate preservation and information, the linkage of pattern to process is often obscured by insufficient temporal resolution. Consequently, reliable documentations of speciation in fossils with discussions on underlying mechanisms are rare. Here we present a well-resolved pattern of morphological evolution in a fossil species lineage of the gastropod Melanopsis in the long-lived Lake Pannon. These developments are related to environmental changes, documented by isotope and stratigraphical data. After a long period of stasis, the ancestral species experiences a phenotypic change expressed as shift and expansion of the morphospace. The appearance of several new phenotypes along with changes in the environment is interpreted as adaptive radiation. Lake-level high stands affect distribution and availability of habitats and, as a result of varied functional demands on shell geometry, the distribution of phenotypes. The on-going divergence of the morphospace into two branches argues for increasing reproductive isolation, consistent with the model of ecological speciation. In the latest phase, however, progressively unstable conditions restrict availability of niches, allowing survival of one branch only.

Morphometrics data on Holocene Melanopsidae of Lake Petea

The dataset includes measurements of Microcolpia parreyssii parreyssii (Philippi, 1847) and Microcolpia parreyssii sikorai (Brusina, 1903) from Holocene deposits of Lake Petea near Oradea, Romania. Additionally, the tps-files generated with the program TpsDig2 and containing pairwise x,y-coordinates describing the outlines of the digitized images are supplied. Finally, the matrix of Fourier coefficients resulting from the Fast Fourier Transform is provided.

Abundance patterns and isotopic signals of morphotypes of Globigerinoides ruber

The chemical composition of shells of the planktonic foraminifer Globigerinoides ruber (white) is frequently used to determine past sea surface conditions. Recently, it has been shown that arbitrarily defined morphotypes within this species exhibit different chemical and isotopic signatures. Here, we investigate the occurrence through time and in space of morphological types of G. ruber (white) in late Quaternary and Holocene sediments of the central and the eastern Mediterranean Sea. In 115 samples representing two distinct time intervals (MIS 1-2 and MIS 9-12) at ODP Site 964 and the piston core GeoTü-SL96, we have defined three morphological types within this species and determined their relative abundances and stable isotopic composition. A quantitative analysis of morphological variation within G. ruber (white) in four samples revealed that the subjectively defined morphotypes occupy separate segments of a continuous and homogenous morphospace. We further show that the abundance of the morphotypes changes significantly between glacials and interglacials and that the three morphotypes of G. ruber show significant offsets in their stable isotopic composition. These offsets are consistent within glacial and interglacial stages but their sign is systematically reversed between the two Sites. Since the isotopic shifts among the three G. ruber morphotypes are systematic and often exceed 1per mil, their understanding is essential for the interpretation of all G. ruber-based proxy records for the paleoceanographic development of the Mediterranean during the late Quaternary.

Raw-data to morphometric investigations about the Neogene planktonic foraminifer Globorotalia menardii and related forms from ODP Hole 154-925B (Céara Rise, western tropical Atlantic)

Evolutionary prospection is the study of morphological evolution and speciation in calcareous plankton from selected time-slices and key sites in the world oceans. In this context, the Neogene menardiform globorotalids serve as study objects for morphological speciation in planktic foraminifera. A downcore investigation of test morphology of the lineage of G. menardii-limbata-multicamerata during the past 8 million years was carried out in the western tropical Atlantic ODP Hole 925B. A total of 4669 specimens were measured and analyzed from 38 stratigraphic levels and compared to previous studies from DSDP Sites 502 and 503. Collection of digital images and morphometric measurements from digitized outlines were achieved using a microfossil orientation and imaging robot called AMOR and software, which was especially developed for this purpose. Most attention was given to the evolution of spiral height versus axial length of tests in keel view, but other parameters were investigated as well. The variability of morphological parameters in G. menardii, G. limbata, and G. multicamerata through time are visualized by volume density diagrams. At Hole 925B results show gradual test size increase in G. menardii until about 3.2 Ma. The combination of taxonomic determination in the light microscope with morphometric investigations shows strong morphological overlap and evolutionary continuity from ancestral to extant G. menardii (4-6 chambers in the final whorl) to the descendent but extinct G. limbata (seven chambers in the final whorl) and to G. multicamerata (>=8 chambers in the final whorl). In the morphospace defined by spiral height (dX) and axial length (dY) Globorotalia limbata and G. multicamerata strongly overlap with G. menardii. Distinction of G. limbata from G. menardii is only possible by slight differences in the number of chambers of the final whorl, nuances in spiral convexity, upper keel angles, radii of osculating circles, or by differences in reflectance of their tests. Globorotalia multicamerata can be distinguished from the other two forms by more than eight chambers in the final whorl. It appeared as two stratigraphically separate clusters during the Pliocene. Between 2.88 and 2.3 Ma G. menardii was severely restricted in size and abundance. Thereafter, it showed a rapid and prominent expansion of the upper test size extremes between 2.3 and 1.95 Ma persisting until present. The size-frequency distributions at Hole 925B are surprisingly similar to trends of menardiform globorotalids from Caribbean DSDP Site 502. There, the observations were explained as an adaptation to changes in the upper water column due to the emergence of the Isthmus of Panama. In light of more recent paleontological and geological investigations about the completion of the permanent land connection between North and South America since about 3 Ma the present study gives reason to suspect the sudden test size increase of G. menardii to reflect immigration of extra-large G. menardii from the Indian Ocean or the Pacific. It is hypothesized that during the Late Pliocene dispersal of large G. menardii into the southern to tropical Atlantic occurred during an intermittent episode of intense Agulhas Current leakage around the Cape of Good Hope and from there via warm eddy transport to the tropical Atlantic (Agulhas dispersal hypothesis).

Morphological variability of Globorotalia menardii in DSDP Sites 68-502 and 68-503

Variability in the test of Globorotalia menardii during the past 8 million years has been investigated at DSDP Site 502A (Caribbean Sea) and DSDP Site 503A (Eastern Equatorial Pacific). Measurements were made of spire height (delta x), maximum diameter (delta y), the tangent angles of the upper and lower peripheral keels (phi 1, phi 2, respectively), the number of chambers in the final whorl, and the area of the silhouette in keel view. Four morphotypes alpha, beta, gamma, and delta were distinguished. Morphotype alpha was found in strata ranging in age from the Late Miocene through the Holocene. It shows a continuous increase in delta x and delta y until the Late Pleistocene. During and after the final closure of the ancient Central American Seaway (between 2.4 Ma and 1.8 Ma) there was a rapid increase in the area of the test in keel view. At the Caribbean Sea site, morphotype beta evolved during the past 0.22 Ma. It is less inflated than alpha and has a more delicate test. In the morphospace of delta x vs. delta y, morphotypes alpha and beta can be distinguished by a separation line delta y = 3.2 * delta x - 160 (delta x and delta y in µm). Plots of morphotype alpha are below that line, those of beta are above it. Morphotype alpha is taken to be Globorotalia menardii menardii Parker, Jones & Brady (1865) and includes G. menardii 'A' Bolli (1970). Morphotype beta is identified as G. menardii cultrata (d'Orbigny). Morphotypes gamma and delta are extinct Upper Miocene to Pliocene forms which evolved from morphotype alpha. They have a narrower phi 1 angle and more chambers (>=7) than morphotype alpha commonly with 5 to 6 chambers (7 in transitional forms). In contemporaneous samples morphotype delta can be distinguished from gamma by a smaller value of phi 1 and 8 or more chambers in the final whorl. Morphotype gamma is taken to be G. limbata (Fornasini, 1902) and includes the junior synonym G. menardii 'B' Bolli (1970). Morphotype delta is G. multicamerata Cushman & Jarvis (1930). With the exception of the Late Pleistocene development of G. menardii cultrataonly in the Caribbean the morphological changes of G. menardii at DSDP Sites 502A and 503A are similar. The development from the ancestral G. menardii menardii of the G. limbata - G. multicamerata lineage during the Pliocene and of G. menardii cultrata during the Late Pleistocene suggests responses at the two sites to a changing palaeoceanography during and after the formation of the Isthmus of Panama.