DIVERSITY‐DEPENDENT CLADOGENESIS AND TRAIT EVOLUTION IN THE ADAPTIVE RADIATION OF THE AUKS (AVES: ALCIDAE)

Through the course of an adaptive radiation, the evolutionary speed of cladogenesis and ecologically relevant trait evolution are expected to slow as species diversity increases, niches become occupied, and ecological opportunity declines. We develop new likelihood‐based models to test diversity‐dependent evolution in the auks, one of only a few families of seabirds adapted to underwater “flight,” and which exhibit a large variety of bill sizes and shapes. Consistent with the expectations of adaptive radiation, we find both a decline in rates of cladogenesis (a sixfold decline) and bill shape (a 64‐fold decline) evolution as diversity increased. Bill shape diverged into two clades at the basal cladogenesis event with one clade possessing mostly long, narrow bills used to forage primarily on fish, and the other with short thick bills used to forage primarily on plankton. Following this initial divergence in bill shape, size, a known correlate of both prey size and maximum diving depth, diverged rapidly within each of these clades. These results suggest that adaptive radiation in foraging traits underwent initial divergence in bill shape to occupy different food resources, followed by size differentiation to subdivide each niche along the depth axis of the water column.     

Evolution and development of the primate limb skeleton

The order Primates is composed of many closely related lineages, each having a relatively well established phylogeny supported by both the fossil record and molecular data. 1 Primate evolution is characterized by a series of adaptive radiations beginning early in the Cenozoic era. Studies of these radiations have uncovered two major trends. One is that substantial amounts of morphological diversity have been produced over short periods of evolutionary time. 2 The other is that consistent and repeated patterns (variational tendencies 3 ) are detected. Taxa within clades, such as the strepsirrhines of Madagascar and the platyrrhines of the Neotropics, have diversified in body size, substrate preference, and diet. 2 , 4 - 6 The diversification of adaptive strategies within such clades is accompanied by repeated patterns of change in cheiridial proportions 7 , 8 (Fig. 1) and tooth‐cusp morphology. 9 There are obvious adaptive, natural‐selection based explanations for these patterns. The hands and feet are in direct contact with a substrate, so their form would be expected to reflect substrate preference, whereas tooth shape is related directly to the functional demands of masticating foods having different mechanical properties. What remains unclear, however, is the role of developmental and genetic processes that underlie the evolutionary diversity of the primate body plan. Are variational tendencies a signature of constraints in developmental pathways? What is the genetic basis for similar morphological transformations among closely related species? These are a sampling of the types of questions we believe can be addressed by future research integrating evidence from paleontology, comparative morphology, and developmental genetics.     

Cats and Dogs Down the Tree: The Tempo and Mode of Evolution in the Lower Carnassial of Fossil and Living Carnivora

Teeth are usually considered the primer to understand mammalian evolution. Their morphology is the result of the complex interplay between adaptation to food processing and somewhat fixed developmental programs. We analyse the morphometry of the first lower molar (m1 = lower carnassial) in 199 carnivore species (64 extant and 135 extinct) in order to clarify the tempo and mode of evolution of two different and possibly related traits, tooth size and tooth angular height. We applied a set of comparative methods and produced disparity through time plots by using four alternative phylogenetic trees. Three of them include fossil species, and were used in order to estimate what an effect the inclusion of fossil species has on the patterns we found for extant species. Our analyses on living species show that m1 size exhibits a weak phylogenetic signal, and that its variance occurs more within than among clades. The opposite applies to m1 angular height, which evolved in an adaptive radiation-like fashion, showing very strong phylogenetic signal and great among-clade variation. These results are thoroughly confirmed after the inclusion of fossil data. Major differences in m1 shape in Carnivora were clearly established early in their evolutionary history, probably as a result of rapid morphospace occupation after the clade radiation. Successive variation in tooth size allowed species to avoid potential overlap in feeding morphologies as a mechanism to limit interspecific competitive interaction within subclades.     

Eating with a saw for a jaw: Functional morphology of the jaws and tooth‐whorl in Helicoprion davisii

The recent reexamination of a tooth‐whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth‐whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth‐whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite‐forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft‐bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth‐whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft‐bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth‐whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft‐bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior‐posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth‐whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion. J. Morphol. 276:47–64, 2015. © 2014 Wiley Periodicals, Inc.     

Mandibular shape correlates of tooth fracture in extant Carnivora: implications to inferring feeding behaviour of Pleistocene predators

Percentages of tooth fracture and mandible shape are robust predictors of feeding habits in Carnivora. If these parameters co‐vary above the species level, more robust palaeobiological inferences could be made on fossil species. A test of association is presented between mandible shape and tooth fracture in a subset of extant carnivorans together with large Pleistocene fossil predators from Rancho La Brea (Canis dirus, Panthera atrox, and Smilodon fatalis). Partial least square (PLS) and comparative methods are employed to validate co‐variation of these two parameters in extant carnivorans. Association between mandible shape and percentage of tooth fracture is strongly supported, even if both blocks of data exhibit a phylogenetic signal to a different degree. Dietary adaptations drive shape/fracture co‐variation in extant species, although no significant differences occur in the PLS scores between carnivores and bone/hard food consumers. The fossil species project into PLS morphospace as outliers. Their position suggests a unique feeding behaviour. The increase in the size of prey, together with consumption of skin and hair from carcasses in a cold environment, might have generated unusual tooth breakage patterns in large predators from Rancho La Brea. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 70–80.     

Fast adaptive responses in the oral jaw of Lake Victoria cichlids

Rapid morphological changes in response to fluctuating natural environments are a common phenomenon in species that undergo adaptive radiation. The dramatic ecological changes in Lake Victoria provide a unique opportunity to study environmental effects on cichlid morphology. This study shows how four haplochromine cichlids adapted their premaxilla to a changed diet over the past 30 years. Directly after the diet change toward larger and faster prey in the late 1980s, the premaxilla (upper jaw) changed in a way that is in agreement with a more food manipulating feeding style. During the 2000s, two zooplanktivorous species showed a reversal of morphological changes after returning to their original diet, whereas two other species showed no reversal of diet and morphology. These rapid changes indicate a potential for extremely fast adaptive responses to environmental fluctuations, which are likely inflicted by competition release and increase, and might have a bearing on the ability of haplochromines to cope with environmental changes. These responses could be due to rapid genetic change or phenotypic plasticity, for which there is ample evidence in cichlid fish structures associated with food capture and processing. These versatile adaptive responses are likely to have contributed to the fast adaptive radiation of haplochromines.     

Ecological opportunity alters the timing and shape of adaptive radiation

The uneven distribution of diversity is a conspicuous phenomenon across the tree of life. Ecological opportunity is a prominent catalyst of adaptive radiation and therefore may alter patterns of diversification. We evaluated the distribution of shifts in diversification rates across the cichlid phylogeny and the distribution of major clades across phylogenetic space. We also tested if ecological opportunity influenced these patterns. Colonization‐associated ecological opportunity altered the tempo and mode of diversification during the adaptive radiation of cichlid fishes. Clades that arose following colonization events diversified faster than other clades. Speciation rate shifts were nonrandomly distributed across the phylogeny such that they were disproportionally concentrated around nodes that corresponded with colonization events (i.e., of continents, river basins, or lakes). Young clades tend to expand faster than older clades; however, colonization‐associated ecological opportunity accentuated this pattern. There was an interaction between clade age and ecological opportunity that explained the trajectory of clades through phylogenetic space over time. Our results indicate that ecological opportunities afforded by continental and ecosystem‐scale colonization events explain the dramatic speciation rate heterogeneity and phylogenetic imbalance that arose during the evolutionary history of cichlid fishes.     

Major issues in the origins of ray‐finned fish (Actinopterygii) biodiversity

Ray‐finned fishes (Actinopterygii) dominate modern aquatic ecosystems and are represented by over 32000 extant species. The vast majority of living actinopterygians are teleosts; their success is often attributed to a genome duplication event or morphological novelties. The remainder are ‘living fossils’ belonging to a few depauperate lineages with long‐retained ecomorphologies: Polypteriformes (bichirs), Holostei (bowfin and gar) and Chondrostei (paddlefish and sturgeon). Despite over a century of systematic work, the circumstances surrounding the origins of these clades, as well as their basic interrelationships and diagnoses, have been largely mired in uncertainty. Here, I review the systematics and characteristics of these major ray‐finned fish clades, and the early fossil record of Actinopterygii, in order to gauge the sources of doubt. Recent relaxed molecular clock studies have pushed the origins of actinopterygian crown clades to the mid‐late Palaeozoic [Silurian–Carboniferous; 420 to 298 million years ago (Ma)], despite a diagnostic body fossil record extending only to the later Mesozoic (251 to 66 Ma). This disjunct, recently termed the ‘Teleost Gap’ (although it affects all crown lineages), is based partly on calibrations from potential Palaeozoic stem‐taxa and thus has been attributed to poor fossil sampling. Actinopterygian fossils of appropriate ages are usually abundant and well preserved, yet long‐term neglect of this record in both taxonomic and systematic studies has exacerbated the gaps and obscured potential synapomorphies. At the moment, it is possible that later Palaeozoic‐age teleost, holostean, chondrostean and/or polypteriform crown taxa sit unrecognized in museum drawers. However, it is equally likely that the ‘Teleost Gap’ is an artifact of incorrect attributions to extant lineages, overwriting both a post‐Palaeozoic crown actinopterygian radiation and the ecomorphological diversity of stem‐taxa.     

Life in the arena: infaunal gastropods and the late Phanerozoic expansion of marine ecosystems into sand

Marine ecosystems have expanded into the infaunal realm below the surface of soft sediments throughout the Phanerozoic eon. During the Palaeozoic era, this expansion largely involved sedentary animals living in permanent resting places. Active sand‐burrowing animals colonized the infaunal environment later, but when this happened and when specialization for infaunal life evolved remain open questions. Here, phylogenetic evidence, fossil occurrences and previously established criteria for recognizing the sand‐burrowing habit in marine gastropods are used to determine how many gastropod clades became infaunal and when the transitions from surface‐dwelling to infaunal life occurred. At least 20, and as many as 35, clades (all but one of post‐Palaeozoic age) contain actively infaunal species. The overwhelming majority (15 of 20 clades) became infaunal during the Cenozoic, and clades with hundreds of infaunal species in the living fauna diversified beginning in the Early Miocene. The repeated evolution of, and specialization to, the sand‐burrowing habit by gastropods and other animals was enabled by increased habitat availability and higher marine productivity, and was necessitated by intensifying predation. As a result, the infaunal realm was transformed from a marine refuge to an integrated part of the marine biosphere in which high performance in locomotion and defence has become the norm.     

Adaptive morphological shifts to novel habitats in marine sculpin fishes

Sculpin fishes of the North American Pacific Coast provide an ideal opportunity to examine whether adaptive morphological character shifts have facilitated occupation of novel habitat types because of their well‐described phylogeny and ecology. In this group, the basal‐rooted species primarily occupy the subtidal habitat, whereas the species in the most distal clades are found in the intertidal. We tested multiple evolutionary models to determine whether changes in body size and changes in number of scales are adaptive for habitat use in sculpins. Based on a statistically robust, highly resolved molecular phylogeny of 26 species of sculpins, in combination with morphometric and habitat affinity data, our analyses show that an adaptive model based on habitat use best explains changes in body size and number of scales. The habitat model was statistically supported over models of neutral evolution, stabilizing selection across all habitats, and three clade‐based models. We suggest that loss of scales and reduction of body size in the intertidal may facilitate cutaneous breathing in air when tidepools become hypoxic during low tides. This study demonstrates how the combined use of phylogenetic, ecological and statistical approaches helps to identify traits that are likely adaptive to novel habitats.     

A molecular phylogenetic framework for the subfamily Ocenebrinae (Gastropoda,Muricidae)

The Ocenebrinae is a subfamily of marine predatory gastropods known as oyster and mussel drills. Their current phylogenetic framework is traditionally based on shell and radular characters, but a consensus on relationships among genera is still lacking. We investigated the molecular phylogeny of Ocenebrinae using 50 species and DNA data from one nuclear (28S) and two mitochondrial (COI and 16S) genes, the largest data set so far assembled for this subfamily. We found support for the monophyly of the Ocenebrinae, and species were divided into four major lineages. Within groups, genera had similar geographic distributions, suggesting that except in a few cases, species diversification within clades occurred without range expansions. We discuss the phylogenetic distribution of a labral tooth and a sealed siphonal canal, two characteristic ocenebrine features. We also show that Ocinebrina species in the north‐eastern Pacific are not monophyletic with north‐eastern Atlantic and Mediterranean species, and that the Ocinebrina edwardsii species complex belongs to Ocenebra.     

Dynamics of evolutionary patterns of clades in a food web system model

The evolutionary patterns of animal species clades in an evolving food web system were examined by computer simulation. In this system, each animal species fed on other species according to feeding preference. The food web system evolved via the appearance and extinction of species. The model succeeded in reproducing evolutionary patterns of diversity similar to those seen in the fossil record. This result indicates that the model reproduced the temporal changes of the rates of colonization and extinction of species in the system, which have been decided a priori in the previous stochastic models. In the food web system, the numbers of both predatory and prey species influenced the temporal diversity patterns in each clade in the system. The number of prey species fluctuated strongly, whereas the number of predatory species gradually increased with time. Therefore, temporal diversity patterns were influenced mainly by the number of predatory species. As a result of the gradual increase of the number of predatory species, it was difficult for each clade to maintain its species diversity for a long time. Slight changes of interspecific interaction can sometimes decide the destiny of a clade. When a clade is faced with extinction, if one predatory species of the clade becomes extinct and one or two prey species of the clade appear, the species diversity in the clade increases again. This result indicates that slight changes of interspecific interaction sometimes decide the destiny of a clade.     

Key innovations and the ecology of macroevolution

The origin or evolutionary `success' of taxa is often attributed to key innovations-aspects of organismal phenotype that promote diversification. Different ways of delimiting taxa and measuring `success' (i.e. number or longevity of species, morphological variety or differential control of energy) give rise to different ideas of how key innovations might operate. Key innovations may enhance competitive ability, relax adaptive trade-offs or permit exploitation of a new productive resource base. Recent key innovation studies comparing species richness in extant sister clades may miss important observations possible only with consideration of the fossil record, traditional higher taxa and phenotypic diversity.     

Interactive effects of productivity and predation on zooplankton diversity

Recent studies suggest the necessity of understanding the interactive effects of predation and productivity on species coexistence and prey diversity. Models predict that coexistence of prey species with different competitive abilities can be achieved if inferior resource competitors are less susceptible to predation and if productivity and/or predation pressure are at intermediate levels. Hence, predator effects on prey diversity are predicted to be highly context dependent: enhancing diversity from low to intermediate levels of productivity or predation and reducing diversity of prey at high levels of productivity or predation. While several studies have examined the interactive effects of herbivory and productivity on primary producer diversity, experimental studies of such effects in predator‐prey systems are rare. We tested these predictions using an aquatic field mesocosm experiment in which initial density of the zooplankton predator Notonecta undulata and productivity were manipulated to test their interactive effects on diversity of seven zooplankton, cladoceran species that were common in surrounding ponds. Two productivity levels were imposed via phosphorus enrichment at levels comparable to low and intermediate levels found within neighboring natural ponds. We used open systems to allow for natural dispersal and behaviorally‐mediated numerical responses by the flight‐capable predator. Effects of predators on zooplankton diversity depended on productivity level. At low and high productivity, prey species richness declined while at high productivity it showed a unimodal relationship with increasing the predator density. Effects of treatments were weaker when using Pielou's evenness index or the inverse Simpson index as measures of prey diversity. Our findings are generally consistent with model predictions in which predators can facilitate prey coexistence and diversity at intermediate levels of productivity and predation intensity. Our work also shows that the functional form of the relationship between prey diversity and predation intensity can be complex and highly dependent on environmental context.     

'First' appearances in the Cenozoic land-mammal record of the Greater Antilles: significance and comparison with South American and Antarctic records

Aim Through analysis of fossil records, the aim of this paper is to show that fossil representatives of at least three land‐mammal clades (pitheciine atelid primates, heteropsomyine echimyid rodents, and megalonychid phyllophagan xenarthrans) that once lived in the Greater Antilles are as old as, if not older than, ‘first’ occurrences of these same groups on the South American mainland. Location Greater Antilles, South America, Antarctic Peninsula. Methods Analysis of Cenozoic land‐mammal fossil records for the three areas. Results Comparison reveals an interesting similarity to the Tertiary vertebrate palaeontological record for the Antarctic Peninsula (Seymour Island), in the sense that the latter also includes early (Eocene) representatives of some typical ‘South American’ groups (e.g. meridiungulates, sloths, certain marsupial groups). Conclusions Given how limited the Antillean and Antarctic records are in quantity and quality, it seems unlikely that these ‘first’ appearances have much bearing on real origins (basal divergences). Rather, it suggests that the fossil basis for interpreting the origin and earliest diversification of ‘South American’ clades during the latest Cretaceous/early Cenozoic is probably even scantier than generally realized. In particular, the Antillean record strengthens arguments that some crown‐group continental lineages are considerably older than fossil evidence currently allows – a point increasingly (if unevenly) supported by molecular studies of many of the same clades.     

Performance of shark teeth during puncture and draw: implications for the mechanics of cutting

The performance of an organism's feeding apparatus has obvious implications for its fitness and survival. However, the majority of studies that focus on chondrichthyan feeding have largely ignored the role of teeth. Studying the functional morphology of shark teeth not only elucidates the biological role that teeth play in feeding, but also provides insight specifically into the evolution of shark feeding because teeth are often the only structures available in the fossil record. In the present study, we investigate the puncture and draw performance of three general categories of extant teeth, tearing‐type, cutting‐type, and cutting–clutching type, as well as three fossil morphologies, utilizing a universal testing system. Differences in puncturing performance occurred among different prey items, indicating that not all ‘soft’ prey items are alike. The majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types; however, few patterns emerged. In some cases, broader triangular teeth were less effective at puncturing than narrow‐cusped teeth. There were no differences between the maximum draw forces and maximum puncture forces. Many of the shark teeth in the present study were not only able to perform draw and puncture equally well, but also many tooth morphologies were functionally equivalent to each other. The findings obtained in the present study lend little support to the belief that shark tooth morphology is a good predictor of biological role. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 271–286.     

Morphological integration and adaptation in the snake feeding system: a comparative phylogenetic study

A long-standing hypothesis for the adaptive radiation of macrostomatan snakes is that their enlarged gape--compared to both lizards and basal snakes--enables them to consume "large" prey. At first glance, this hypothesis seems plausible, or even likely, given the wealth of studies showing a tight match between maximum consumed prey mass and head size in snakes. However, this hypothesis has never been tested within a comparative framework. We address this issue here by testing this hypothesis in 12 monophyletic clades of macrostomatan snakes using recently published phylogenies, published maximum consumed prey mass data and morphological measurements taken from a large sample of museum specimens. Our nonphylogenetically corrected analysis shows that head width--independent of body size--is significantly related to mean maximum consumed prey mass among these clades, and this relationship becomes even more significant when phylogeny is taken into account. Therefore, these data do support the hypothesis that head shape is adapted to prey size in snakes. Additionally, we calculated a phylogenetically corrected morphological variance-covariance matrix to examine the role of morphological integration during head shape evolution in snakes. This matrix shows that head width strongly covaries with both jaw length and out-lever length of the lower jaw. As a result, selection on head width will likely be associated with concomitant changes in jaw length and lower jaw out-lever length in snakes.     

Longer in the tooth,shorter in the record? The evolutionary correlates of hypsodonty in Neogene ruminants

The acquisition of hypsodont molars is often regarded as a key innovation in the history of ruminant ungulates. Hypsodont ruminants diversified rapidly during the later Neogene, circa 15-2 Myr ago, and came to dominate the ruminant fossil record in terms of species diversity. Here we show that hypsodont clades had higher speciation and diversification rates than other clades. Hypsodont species had, on average, shorter stratigraphic durations, smaller range size and lower occupancy than non-hypsodont species. Within hypsodont clades, some species were very common and acquired large geographical ranges, whereas others were quite rare and geographically limited. We argue that hypsodont clades diversified in an adaptive radiation-like fashion, with species often splitting cladogenetically while still in the expansive phase of their occupancy history.     

TESTING FOR ANCIENT ADAPTIVE RADIATIONS IN NEOTROPICAL CICHLID FISHES

Most contemporary studies of adaptive radiation focus on relatively recent and geographically restricted clades. It is less clear whether diversification of ancient clades spanning entire continents is consistent with adaptive radiation. We used novel fossil calibrations to generate a chronogram of Neotropical cichlid fishes and to test whether patterns of lineage and morphological diversification are congruent with hypothesized adaptive radiations in South and Central America. We found that diversification in the Neotropical cichlid clade and the highly diverse tribe Geophagini was consistent with diversity‐dependent, early bursts of divergence followed by decreased rates of lineage accumulation. South American Geophagini underwent early rapid differentiation in body shape, expanding into novel morphological space characterized by elongate‐bodied predators. Divergence in head shape attributes associated with trophic specialization evolved under strong adaptive constraints in all Neotropical cichlid clades. The South American Cichlasomatini followed patterns consistent with constant rates of morphological divergence. Although morphological diversification in South American Heroini was limited, Eocene invasion of Central American habitats was followed by convergent diversification mirroring variation observed in Geophagini. Diversification in Neotropical cichlids was influenced by the early adaptive radiation of Geophagini, which potentially limited differentiation in other cichlid clades.     

A review of the impacts of corvids on bird productivity and abundance

Corvids are often viewed as efficient predators capable of limiting prey species populations. Despite this widely held belief, a comprehensive review quantifying the effect of corvids on the demography of prey species is lacking. We examine the impacts of crows, ravens Corvus spp. and Eurasian Magpies Pica pica on the population parameters of other bird species. We summarize results from 42 studies, which included 326 explicit evaluations of relationships between a corvid and a potential prey species. Population parameters of studied prey species were categorized as abundance‐related (numbers, nest density) or productivity‐related (nest success, brood size). Information from both experimental removal studies and correlative studies was examined. Combining all studies, no negative influence of corvids on either abundance or productivity of prey species was found in 81% of cases. Negative impacts were significantly more likely in cases examining productivity rather than abundance (46 vs. 10%). Experimental studies that removed only corvid species were significantly less likely to show a positive impact on productivity than those removing corvids alongside other predators (16 vs. 60%). This suggests that the impact of corvids is smaller than that of other predators, or that compensatory predation occurs. The impact of corvids was similar between diverse avian groups (such as gamebirds, passerines and waders; or ground‐nesting and other species). Crows were found to be significantly more likely to have a negative impact on prey species productivity than were Magpies (62 vs. 12%), but no differences were found in relation to prey abundance. We conclude that while corvids can have a negative impact on bird species, their impact is small overall, and nearly five times more frequent for productivity than for abundance. These results suggest that in most cases bird populations are unlikely to be limited by corvid predation and that conservation measures may generally be better targeted at other limiting factors. However, negative impacts were found in a minority of cases, and those may require further investigation to develop management tools to mitigate such impacts where they are of economic or conservation concern.