Comparative morphology of shark pectoral fins

Sharks vary greatly in morphology, physiology, and ecology. Differences in whole body shape, swimming style, and physiological parameters have previously been linked to varied habitat uses. Pectoral fin morphology has been used to taxonomically classify species and hypotheses on the functional differences in shape are noted throughout the literature; however, there are limited comparative datasets that quantify external and skeletal morphology. Further, fins were previously categorized into two discrete groups based on the amount of skeletal support present: (a) aplesodic, where less than half of the fin is supported and (b) plesodic where greater than half of the fin is supported. These discrete classifications have been used to phylogenetically place species, though the methodology of classification is infrequently described. In this study, we sampled fins from 18 species, 6 families, and 3 orders, which were also grouped into five ecomorphotype classifications. We examined the external morphology, extent of skeletal support, and cross-sectional shape of individual cartilaginous elements. Using phylogenetic comparative methods, we show that fin shape does not differ significantly between ecomorphotypes, suggesting there may be some mechanical constraint. However, we find that the internal anatomy of the fin does vary significantly between ecomorphotypes, especially the extent and distribution of calcification of skeletal support, suggesting that the superficial similarity of fin shapes across ecomorphotypes may belie differences in function. Finally, we find that a number of morphological variables such as number of radials, radial calcification and shape, and fin taper all correlate with the extent of skeletal support. Within these morphospaces, we also describe that some orders/families tend to occupy certain areas with limited overlap. While we demonstrate that there is some mechanical constraint limiting external variations in shark pectoral fin morphology, there are compounding differences in skeletal anatomy that occur within ecomorphotypes which we propose may affect function.     

Steady diversification of derived liverworts under Tertiary climatic fluctuations

Tropical forests contain the majority of extant plant diversity and their role as a cradle and/or museum of biodiversity is an important issue in our attempts to assess the long-term consequences of global climate change for terrestrial biomes. Highly diverse groups of liverworts are an often ignored but extremely common element in rainforests, and thus their evolution may shed light on the ecological robustness of rainforest biomes to climate fluctuations. We record a remarkable constant accumulation of diversity through time for the most species-rich family of liverworts, Lejeuneaceae, inferred by divergence time estimates. The observed pattern supports the recently developed concept of a dual role of the tropics as both a museum and a cradle of biodiversity.     

The effect of community on the territorial behavior of the threespot damselfish

Synopsis I examined the hypothesis that animals alter their behavior in response to changes at the community scale of organization, rather than simply reacting to a sequential set of independent interactions with other organisms. The focal species was the threespot damselfish, Stegastes planifrons,a territorial coral reef fish that experiences a high degree of variation in community composition. Field observations of individual threespot damselfish showed a significant increase in the amount of time spent on active territorial defense when the community contained greater proportions of food competitors. Territorial behavior also increased concurrently with higher species diversity in the community. However, no behavioral change was observed in response to the total number of fish entering the defended territory, suggesting that threespot damselfish are responding to more complex environmental cues associated with community structure rather than simple density.     

The influence of size on body shape diversification across Indo‐Pacific shore fishes*

Understanding the causes of body shape variability across the tree of life is one of the central issues surrounding the origins of biodiversity. One potential mechanism driving observed patterns of shape disparity is a strongly conserved relationship between size and shape. Conserved allometry has been shown to account for as much as 80% of shape variation in some vertebrate groups. Here, we quantify the amount of body shape disparity attributable to changes in body size across nearly 800 species of Indo‐Pacific shore fishes using a phylogenetic framework to analyze 17 geometric landmarks positioned to capture general body shape and functionally significant features. In marked contrast to other vertebrate lineages, we find that changes in body size only explain 2.9% of the body shape variation across fishes, ranging from 3% to 50% within our 11 sampled families. We also find a slight but significant trend of decreasing rates of shape evolution with increasing size. Our results suggest that the influence of size on fish shape has largely been overwhelmed by lineage‐specific patterns of diversification that have produced the modern landscape of highly diverse forms that we currently observe in nature.     

Primate diversification inferred from phylogenies and fossils

Biodiversity arises from the balance between speciation and extinction. Fossils record the origins and disappearance of organisms, and the branching patterns of molecular phylogenies allow estimation of speciation and extinction rates, but the patterns of diversification are frequently incongruent between these two data sources. I tested two hypotheses about the diversification of primates based on ～600 fossil species and 90% complete phylogenies of living species: (1) diversification rates increased through time; (2) a significant extinction event occurred in the Oligocene. Consistent with the first hypothesis, analyses of phylogenies supported increasing speciation rates and negligible extinction rates. In contrast, fossils showed that while speciation rates increased, speciation and extinction rates tended to be nearly equal, resulting in zero net diversification. Partially supporting the second hypothesis, the fossil data recorded a clear pattern of diversity decline in the Oligocene, although diversification rates were near zero. The phylogeny supported increased extinction ～34 Ma, but also elevated extinction ～10 Ma, coinciding with diversity declines in some fossil clades. The results demonstrated that estimates of speciation and extinction ignoring fossils are insufficient to infer diversification and information on extinct lineages should be incorporated into phylogenetic analyses.     

Food capture kinematics of the suction feeding horn shark, Heterodontus francisci

The goal of this study was to examine the feeding kinematics of the horn shark, Heterodontus francisci, a member of the most basal clade of galeomorph sharks, the Heterodontiformes. The accessibility of the food was manipulated to determine if the horn shark modulated capture. Three different methods of presenting food were used to mimic the different positions of prey items found in the natural diet of the horn shark. Food was presented unattached to the substrate, securely attached, or fitted snugly in a tube. Using high-speed video kinematic analysis, capture events were examined. Heterodontus francisci uses inertial suction facilitated by rapid mandible depression and labial cartilage protrusion to capture food. The horn shark conforms to a capture kinematic profile characteristic of both basal and derived inertial suction feeding sharks. Unusual post-capture behaviors include body leveraging, use of the mouth to form a seal over food, and chisel-like palatoquadrate protrusion. When presented with food of different accessibility, Heterodontus francisci used one consistent kinematic pattern for capture that was not modulated. Only post-capture behaviors varied according to food accessibility.     

Genomic signatures of host‐associated divergence and adaptation in a coral‐eating snail,Coralliophila violacea (Kiener, 1836)

The fluid nature of the ocean, combined with planktonic dispersal of marine larvae, lowers physical barriers to gene flow. However, divergence can still occur despite gene flow if strong selection acts on populations occupying different ecological niches. Here, we examined the population genomics of an ectoparasitic snail, Coralliophila violacea (Kiener 1836), that specializes on Porites corals in the Indo‐Pacific. Previous genetic analyses revealed two sympatric lineages associated with different coral hosts. In this study, we examined the mechanisms promoting and maintaining the snails’ adaptation to their coral hosts. Genome‐wide single nucleotide polymorphism (SNP) data from type II restriction site‐associated DNA (2b‐RAD) sequencing revealed two differentiated clusters of C. violacea that were largely concordant with coral host, consistent with previous genetic results. However, the presence of some admixed genotypes indicates gene flow from one lineage to the other. Combined, these results suggest that differentiation between host‐associated lineages of C. violacea is occurring in the face of ongoing gene flow, requiring strong selection. Indeed, 2.7% of all SNP loci were outlier loci (73/2,718), indicative of divergence with gene flow, driven by adaptation of each C. violacea lineage to their specific coral hosts.     

Coral reefs promote the evolution of morphological diversity and ecological novelty in labrid fishes

Although coral reefs are renowned biodiversity hotspots it is not known whether they also promote the evolution of exceptional ecomorphological diversity. We investigated this question by analysing a large functional morphological dataset of trophic characters within Labridae, a highly diverse group of fishes. Using an analysis that accounts for species relationships, the time available for diversification and model uncertainty we show that coral reef species have evolved functional morphological diversity twice as fast as non-reef species. In addition, coral reef species occupy 68.6% more trophic morphospace than non-reef species. Our results suggest that coral reef habitats promote the evolution of both trophic novelty and morphological diversity within fishes. Thus, the preservation of coral reefs is necessary, not only to safeguard current biological diversity but also to conserve the underlying mechanisms that can produce functional diversity in future.     

Composition and structure of reef fish communities in Paraíba State,north‐eastern Brazil

The composition and trophic structure of reef fish communities in two natural and two artificial reefs along the coast of Paraíba State in north‐eastern Brazil were investigated. A total of 114 species of fish belonging to 47 families were recorded during 120 stationary visual surveys, slightly less than half (46·55%) of which were recorded at all four surveyed localities. Most species are widely distributed on the western Atlantic coast, but several are endemic to Brazil. The greatest diversity and equitability indexes were recorded at the reefs of Sapatas and Cabeço dos Cangulos, whereas the greatest richness and abundance were found at the Queimado shipwreck. The Alvarenga shipwreck reef had the least richness, diversity and equitability. The four localities studied had very similar ichthyofaunas, especially in relation to species composition. The reefs along the Paraíba coast are considered priority conservation areas by the Brazilian Ministry of the Environment, and the information generated by this study will be useful for comparison with other reefs in the region and can be directly applied to programmes seeking to protect and manage these environments.     

High‐resolution imaging sheds new light on a multi‐tier symbiotic partnership between a “walking” solitary coral,a sipunculan,and a bivalve from East Africa

Marine symbioses are integral to the persistence of ecosystem functioning in coral reefs. Solitary corals of the species Heteropsammia cochlea and Heterocyathus aequicostatus have been observed to live in symbiosis with the sipunculan worm Aspidosiphon muelleri muelleri, which inhabits a cavity within the coral, in Zanzibar (Tanzania). The symbiosis of these photosymbiotic corals enables the coral holobiont to move, in fine to coarse unconsolidated substrata, a process termed as “walking.” This allows the coral to escape sediment cover in turbid conditions which is crucial for these light‐dependent species. An additional commensalistic symbiosis of this coral‐worm holobiont is found between the Aspidosiphon worm and the cryptoendolithic bivalve Jousseaumiella sp., which resides within the cavity of the coral skeleton. To understand the morphological alterations caused by these symbioses, interspecific relationships, with respect to the carbonate structures between these three organisms, are documented using high‐resolution imaging techniques (scanning electron microscopy and µCT scanning). Documenting multi‐layered symbioses can shed light on how morphological plasticity interacts with environmental conditions to contribute to species persistence.     

Larval connectivity across temperature gradients and its potential effect on heat tolerance in coral populations

Coral reefs are increasingly exposed to elevated temperatures that can cause coral bleaching and high levels of mortality of corals and associated organisms. The temperature threshold for coral bleaching depends on the acclimation and adaptation of corals to the local maximum temperature regime. However, because of larval dispersal, coral populations can receive larvae from corals that are adapted to very different temperature regimes. We combine an offline particle tracking routine with output from a high‐resolution physical oceanographic model to investigate whether connectivity of coral larvae between reefs of different thermal regimes could alter the thermal stress threshold of corals. Our results suggest that larval transport between reefs of widely varying temperatures is likely in the Coral Triangle and that accounting for this connectivity may be important in bleaching predictions. This has important implications in conservation planning, because connectivity may allow some reefs to have an inherited heat tolerance that is higher or lower than predicted based on local conditions alone.     

The Evolution of Calcification in Reef-Building Corals

Corals build the structural foundation of coral reefs, one of the most diverse and productive ecosystems on our planet. Although the process of coral calcification that allows corals to build these immense structures has been extensively investigated, we still know little about the evolutionary processes that allowed the soft-bodied ancestor of corals to become the ecosystem builders they are today. Using a combination of phylogenomics, proteomics, and immunohistochemistry, we show that scleractinian corals likely acquired the ability to calcify sometime between ∼308 and ∼265 Ma through a combination of lineage-specific gene duplications and the co-option of existing genes to the calcification process. Our results suggest that coral calcification did not require extensive evolutionary changes, but rather few coral-specific gene duplications and a series of small, gradual optimizations of ancestral proteins and their co-option to the calcification process.     

CO2 partial pressure controls the calcification rate of a coral community

Previous studies have demonstrated that coral and algal calcification is tightly regulated by the calcium carbonate saturation state of seawater. This parameter is likely to decrease in response to the increase of dissolved CO₂ resulting from the global increase of the partial pressure of atmospheric CO₂. We have investigated the response of a coral reef community dominated by scleractinian corals, but also including other calcifying organisms such as calcareous algae, crustaceans, gastropods and echinoderms, and kept in an open‐top mesocosm. Seawater pCO₂ was modified by manipulating the pCO₂ of air used to bubble the mesocosm. The aragonite saturation state (Ω_arag) of the seawater in the mesocosm varied between 1.3 and 5.4. Community calcification decreased as a function of increasing pCO₂ and decreasing Ω_arag. This result is in agreement with previous data collected on scleractinian corals, coralline algae and in a reef mesocosm, even though some of these studies did not manipulate CO₂ directly. Our data suggest that the rate of calcification during the last glacial maximum might have been 114% of the preindustrial rate. Moreover, using the average emission scenario (IS92a) of the Intergovernmental Panel on Climate Change, we predict that the calcification rate of scleractinian‐dominated communities may decrease by 21% between the pre‐industrial period (year 1880) and the time at which pCO₂ will double (year 2065).     

Coral reef mesopredators switch prey,shortening food chains,in response to habitat degradation

Diet specificity is likely to be the key predictor of a predator's vulnerability to changing habitat and prey conditions. Understanding the degree to which predatory coral reef fishes adjust or maintain prey choice, in response to declines in coral cover and changes in prey availability, is critical for predicting how they may respond to reef habitat degradation. Here, we use stable isotope analyses to characterize the trophic structure of predator–prey interactions on coral reefs of the Keppel Island Group on the southern Great Barrier Reef, Australia. These reefs, previously typified by exceptionally high coral cover, have recently lost much of their coral cover due to coral bleaching and frequent inundation by sediment‐laden, freshwater flood plumes associated with increased rainfall patterns. Long‐term monitoring of these reefs demonstrates that, as coral cover declined, there has been a decrease in prey biomass, and a shift in dominant prey species from pelagic plankton‐feeding damselfishes to territorial benthic algal‐feeding damselfishes, resulting in differences in the principal carbon pathways in the food web. Using isotopes, we tested whether this changing prey availability could be detected in the diet of a mesopredator (coral grouper, Plectropomus maculatus). The δ¹³C signature in grouper tissue in the Keppel Islands shifted from a more pelagic to a more benthic signal, demonstrating a change in carbon sources aligning with the change in prey availability due to habitat degradation. Grouper with a more benthic carbon signature were also feeding at a lower trophic level, indicating a shortening in food chains. Further, we found a decline in the coral grouper population accompanying a decrease in total available prey biomass. Thus, while the ability to adapt diets could ameliorate the short‐term impacts of habitat degradation on mesopredators, long‐term effects may negatively impact mesopredator populations and alter the trophic structure of coral reef food webs.     

The evolution of South American endemic canids: a history of rapid diversification and morphological parallelism

The origin of endemic South American canid fauna has been traditionally linked with the rise of the Isthmus of Panama, suggesting that diversification of the dog fauna on this continent occurred very rapidly. Nevertheless, despite its obvious biogeographic appeal, the tempo of Canid evolution in South America has never been studied thoroughly. This issue can be suitably tackled with the inference of a molecular timescale. In this study, using a relaxed molecular clock method, we estimated that the most recent common ancestor of South American canids lived around 4 Ma, whereas all other splits within the clade occurred after the rise of the Panamanian land bridge. We suggest that the early diversification of the ancestors of the two main lineages of South American canids may have occurred in North America, before the Great American Interchange. Moreover, a concatenated morphological and molecular analysis put some extinct canid species well within the South American radiation, and shows that the dental adaptations to hypercarnivory evolved only once in the South American clade.     

Symbiont footprints highlight the diversity of scleractinian‐associated Zanclea hydrozoans (Cnidaria,Hydrozoa)

Hydrozoans of the genus Zanclea have been acknowledged only recently as a fundamental component of the highly diverse fauna associated with reef‐building scleractinian corals. Although widely distributed in coral reefs and demonstrated to be important in protecting corals from predation and diseases, the biodiversity of these hydrozoans remains enigmatic due to the paucity of available morphological characters, incomplete morphological characterisations and the possible existence of cryptic species. Recently, molecular techniques have revealed the existence of multiple hidden genetic lineages not yet supported by diagnostic morphological characters. In this work, we further explore the morpho‐diversity of three genetic lineages, namely Zanclea associated with the coral genera Goniastrea (clade I), Porites (clade II) and Pavona (clade VI). Aside from providing a complete classical characterisation of the polyp and medusa stage of each clade, we searched for new potential taxonomic indicators either on symbiotic hydroids or on host corals. On the hydroids, statistical analyses on almost 7,000 nematocyst capsules revealed a significant difference in terms of nematocyst size among the three Zanclea clades investigated. On each host coral genus, we identified peculiar skeletal modifications related to the presence of Zanclea symbionts. Lastly, we discussed the potential diagnostic value of these footprints in the characterisation of Zanclea–scleractinian associations.     

DO REEFS DRIVE DIVERSIFICATION IN MARINE TELEOSTS? EVIDENCE FROM THE PUFFERFISH AND THEIR ALLIES (ORDER TETRAODONTIFORMES)

A major challenge in evolutionary biology lies in explaining patterns of high species numbers found in biodiversity hot spots. Tropical coral reefs underlie most marine hot spots and reef-associated fish faunas represent some of the most diverse assemblages of vertebrates on the planet. Although the standing diversity of modern reef fish clades is usually attributed to their ecological association with corals, untangling temporal patterns of codiversification has traditionally proved difficult. In addition, owing to uncertainty in higher-level relationships among acanthomorph fish, there have been few opportunities to test the assumption that reef-association itself leads to higher rates of diversification compared to other habitats. Here we use relaxed-clock methods in conjunction with statistical measures of species accumulation and phylogenetic comparative methods to clarify the temporal pattern of diversification in reef and nonreef-associated lineages of tetraodontiforms, a morphologically diverse order of teleost fish. We incorporate 11 fossil calibrations distributed across the tetraodontiform tree to infer divergence times and compare results from models of autocorrelated and uncorrelated evolutionary rates. All major tetraodontiform reef crown groups have significantly higher rates of diversification than the order as a whole. None of the nonreef-associated families show this pattern with the exception of the aracanid boxfish. Independent contrasts analysis also reveals a significantly positive relationship between diversification rate and proportion of reef-associated species within each family when aracanids are excluded. Reef association appears to have increased diversification rate within tetraodontiforms. We suggest that both intrinsic factors of reef habitat and extrinsic factors relating to the provincialization and regionalization of the marine biota during the Miocene (about 23-5 MY) played a role in shaping these patterns of diversity.     

Comparative feeding ecology of the yellow ray Urobatis jamaicensis (Urotrygonidae) from The Bahamas

Stomach contents were collected from 117 yellow rays Urobatis jamaicensis from three locations in south Eleuthera, The Bahamas and compared with ambient infauna via sediment surveys. Diets were relatively limited with a total of 535 prey items recovered, representing five taxonomic groups and dominated by polychaetes and decapod crustaceans (87% of total diet), while environmental sampling reported 5249 individual taxa represented by 62 taxonomic groups. Regardless of gravidity, sex or density of prey items among sites, no significant differences were detected. Foraging strategy plots suggested preferential prey is rare within the environment and the Manly‐Chesson index validates polychaetes were consumed with high selectivity. This is the most comprehensive and updated assessment of comparative feeding in this species, particularly for The Bahamas, allowing insight into invertebrate community richness and diversity in ecologically sensitive coastal and nearshore habitats.