A brain-wide analysis maps structural evolution to distinct anatomical module.

The vertebrate brain is highly conserved topologically, but less is known about neuroanatomical variation between individual brain regions. Neuroanatomical variation at the regional level is hypothesized to provide functional expansion, building upon ancestral anatomy needed for basic functions. Classically, animal models used to study evolution have lacked tools for detailed anatomical analysis that are widely used in zebrafish and mice, presenting a barrier to studying brain evolution at fine scales. In this study, we sought to investigate the evolution of brain anatomy using a single species of fish consisting of divergent surface and cave morphs, that permits functional genetic testing of regional volume and shape across the entire brain. We generated a high-resolution brain atlas for the blind Mexican cavefish Astyanax mexicanus and coupled the atlas with automated computational tools to directly assess variability in brain region shape and volume across all populations. We measured the volume and shape of every grossly defined neuroanatomical region of the brain and assessed correlations between anatomical regions in surface fish, cavefish, and surface × cave F2 hybrids, whose phenotypes span the range of surface to cave. We find that dorsal regions of the brain are contracted, while ventral regions have expanded, with F2 hybrid data providing support for developmental constraint along the dorsal-ventral axis. Furthermore, these dorsal-ventral relationships in anatomical variation show similar patterns for both volume and shape, suggesting that the anatomical evolution captured by these two parameters could be driven by similar developmental mechanisms. Together, these data demonstrate that A. mexicanus is a powerful system for functionally determining basic principles of brain evolution and will permit testing how genes influence early patterning events to drive brain-wide anatomical evolution.

Covariation of brain and skull shapes as a model to understand the role of crosstalk in development and evolution.

Covariation among discrete phenotypes can arise due to selection for shared functions, and/or shared genetic and developmental underpinnings. The consequences of such phenotypic integration are far-reaching and can act to either facilitate or limit morphological variation. The vertebrate brain is known to act as an "organizer" of craniofacial development, secreting morphogens that can affect the shape of the growing neurocranium, consistent with roles for pleiotropy in brain-neurocranium covariation. Here, we test this hypothesis in cichlid fishes by first examining the degree of shape integration between the brain and the neurocranium using three-dimensional geometric morphometrics in an F5 hybrid population, and then genetically mapping trait covariation using quantitative trait loci (QTL) analysis. We observe shape associations between the brain and the neurocranium, a pattern that holds even when we assess associations between the brain and constituent parts of the neurocranium: the rostrum and braincase. We also recover robust genetic signals for both hard- and soft-tissue traits and identify a genomic region where QTL for the brain and braincase overlap, implicating a role for pleiotropy in patterning trait covariation. Fine mapping of the overlapping genomic region identifies a candidate gene, notch1a, which is known to be involved in patterning skeletal and neural tissues during development. Taken together, these data offer a genetic hypothesis for brain-neurocranium covariation, as well as a potential mechanism by which behavioral shifts may simultaneously drive rapid change in neuroanatomy and craniofacial morphology.

Breaking constraints: The development and evolution of extreme fin morphology in the Bramidae.

The developmental process establishes the foundation upon which natural selection may act. In that same sense, it is inundated with numerous constraints that work to limit the directions in which a phenotype may respond to selective pressures. Extreme phenotypes have been used in the past to identify tradeoffs and constraints and may aid in recognizing how alterations to the Baupläne can influence the trajectories of lineages. The Bramidae, a family of Scombriformes consisting of 20 extant species, are unique in that five species greatly deviate from the stout, ovaloid bodies that typify the bramids. The Ptericlinae, or fanfishes, are instead characterized by relatively elongated body plans and extreme modifications to their medial fins. Here, we explore the development of Bramidae morphologies and examine them through a phylogenetic lens to investigate the concepts of developmental and evolutionary constraints. Contrary to our predictions that the fanfishes had been constrained by inherited properties of an ancestral state, we find that the fanfishes exhibit both increased rates of trait evolution and differ substantially from the other bramids in their developmental trajectories. Conversely, the remaining bramid genera differ little, both among one another and in comparison, to the sister family Caristiidae. In all, our data suggest that the fanfishes have broken constraints, thereby allowing them to mitigate trade-offs on distinctive aspects of morphology.

The influence of divergent reproductive strategies in shaping modularity and morphological evolution in mammalian jaws.

Marsupial neonates are born at an earlier developmental stage than placental mammals, but the rapid development of their forelimbs and cranial skeleton allows them to climb to the pouch, begin suckling and complete their development ex utero. The mechanical environment in which marsupial neonates develop is vastly different from that of placental neonates, which exhibit a more protracted development of oral muscles and bones. This difference in reproductive strategy has been theorized to constrain morphological evolution in the oral region of marsupials. Here, we use 3D morphometrics to characterize one of these oral bones, the lower jaw (dentary), and assess modularity (pattern of covariation among traits), morphological disparity and rates of morphological evolution in two clades of carnivorous mammals: the marsupial Dasyuromorphia and placental fissiped Carnivora. We find that dasyuromorph dentaries have fewer modules than carnivorans and exhibit tight covariation between the angular and coronoid processes, the primary attachment sites for jaw-closing muscles. This pattern of modularity may result from the uniform action of muscles on the developing mandible during suckling. Carnivorans are free from this constraint and exhibit a pattern of modularity that more strongly reflects genetic and developmental signals of trait covariation. Alongside differences in modularity, carnivorans exhibit greater disparity and faster rates of morphological evolution compared with dasyuromorphs. Taken together, this suggests dasyuromorphs have retained a signal of trait covariation that reflects the outsized influence of muscular force during early development, a feature that may have impacted the ability of marsupial carnivores to explore specialized regions of morphospace.

The cichlid oral and pharyngeal jaws are evolutionarily and genetically coupled.

Evolutionary constraints may significantly bias phenotypic change, while "breaking" from such constraints can lead to expanded ecological opportunity. Ray-finned fishes have broken functional constraints by developing two jaws (oral-pharyngeal), decoupling prey capture (oral jaw) from processing (pharyngeal jaw). It is hypothesized that the oral and pharyngeal jaws represent independent evolutionary modules and this facilitated diversification in feeding architectures. Here we test this hypothesis in African cichlids. Contrary to our expectation, we find integration between jaws at multiple evolutionary levels. Next, we document integration at the genetic level, and identify a candidate gene, smad7, within a pleiotropic locus for oral and pharyngeal jaw shape that exhibits correlated expression between the two tissues. Collectively, our data show that African cichlid evolutionary success has occurred within the context of a coupled jaw system, an attribute that may be driving adaptive evolution in this iconic group by facilitating rapid shifts between foraging habitats, providing an advantage in a stochastic environment such as the East African Rift-Valley.

Extreme Morphology, Functional Trade-offs, and Evolutionary Dynamics in a Clade of Open-Ocean Fishes (Perciformes: Bramidae).

When novel or extreme morphologies arise, they are oft met with the burden of functional trade-offs in other aspects of anatomy, which may limit phenotypic diversification and make particular adaptive peaks inaccessible. Bramids (Perciformes: Bramidae) comprise a small family of 20 extant species of fishes, which are distributed throughout pelagic waters worldwide. Within the Bramidae, the fanfishes (Pteraclis and Pterycombus) differ morphologically from the generally stout, laterally compressed species that typify the family. Instead, Pteraclis and Pterycombus exhibit extreme anterior positioning of the dorsal fin onto the craniofacial skeleton. Consequently, they possess fin and skull anatomies that are radically different from other bramid species. Here, we investigate the anatomy, development, and evolution of the Bramidae to test the hypothesis that morphological innovations come at functional (proximate) and evolutionary (ultimate) costs. Addressing proximate effects, we find that the development of an exaggerated dorsal fin is associated with neurocrania modified to accommodate an anterior expansion of the dorsal fin. This occurs via reduced development of the supraoccipital crest (SOC), providing a broad surface area on the skull for insertion of the dorsal fin musculature. While these anatomical shifts are presumably associated with enhanced maneuverability in fanfishes, they are also predicted to result in compromised suction feeding, possibly limiting the mechanisms of feeding in this group. Phylogenetic analyses suggest craniofacial and fin morphologies of fanfishes evolved rapidly and are evolutionarily correlated across bramids. Furthermore, fanfishes exhibit a similar rate of lineage diversification as the rest of the Bramidae, lending little support for the prediction that exaggerated medial fins are associated with phylogenetic constraint. Our phylogeny places fanfishes at the base of the Bramidae and suggests that nonfanfish bramids have reduced medial fins and re-evolved SOCs. These observations suggest that the evolution of novel fin morphologies in basal species has led to the phylogenetic coupling of head and fin shape, possibly predisposing the entire family to a limited range of feeding. Thus, the evolution of extreme morphologies may have carryover effects, even after the morphology is lost, limiting ecological diversification of lineages.

新規または極端な形態が発生すると、解剖学的構造の他の側面で機能的なトレードオフの負担に直面することが多く、表現型の多様化が制限され、特定の適応ピークにアクセスできなくなる可能性があります。ブラミド（スズキ目：シマガツオ科）は、20種の現存する魚の小さな家族で構成されており、世界中の遠洋水域に分布しています。ブラミダエ内では、シマガツオ（PteraclisとPterycombus）は、家族を代表する一般的に頑丈な横方向に圧縮された種と形態学的に異なります。代わりに、PteraclisとPterycombusは、頭蓋顔面骨格への背びれの極端な前方位置を示します。その結果、それらは他のブラミド種とは根本的に異なるひれと頭蓋骨の解剖学的構造を持っています。ここでは、形態学的革新が機能的（近接）および進化的（究極）コストでもたらされるという仮説をテストするために、ブラミダエの解剖学、発達、および進化を調査します。近接効果に対処すると、誇張された背びれの発達は、背びれの前方拡張に対応するように修正された脳頭蓋に関連していることがわかります。これは、後頭上頂（SOC）の発達の低下を介して発生し、背びれの筋肉組織を挿入するための頭蓋骨の広い表面積を提供します。これらの解剖学的変化はおそらくファンフィッシュの操作性の向上に関連していると思われますが、吸引餌の低下をもたらし、このグループの餌のメカニズムを制限する可能性もあると予測されています。系統発生分析は、シマガツオの頭蓋顔面およびヒレの形態が急速に進化し、ブラミド間で進化的に相関していることを示唆しています。さらに、ファンフィッシュは他のブラミダエと同様の系統多様化率を示し、誇張された内側のひれが系統発生の制約に関連しているという予測をほとんど支持していません。私たちの系統発生は、シマガツオをシマガツオ科の根元に配置し、シマガツオ以外のシマガツオが内側のひれを減らし、SOCを再進化させたことを示唆しています。これらの観察結果は、基底種における新しいヒレの形態の進化が、頭とヒレの形状の系統発生的結合をもたらし、おそらく家族全員が限られた範囲の摂食にかかりやすくなっていることを示唆しています。したがって、極端な形態の進化は、形態が失われた後でも持ち越し効果をもたらす可能性があり、系統の生態学的多様化を制限します。.

Cuando surgen morfologías nuevas o extremas, a menudo se encuentran con la carga de compensaciones funcionales en otros aspectos de la anatomía, lo que puede limitar la diversificación fenotípica y hacer inaccesibles los picos adaptativos particulares. Las bramidas (Perciformes: Bramidae) comprenden una pequeña familia de 20 especies de peces existentes, que se distribuyen en las aguas pelágicas de todo el mundo. Dentro de los Bramidae, los fanfishes (Pteraclis y Pterycombus) difieren morfológicamente de las especies generalmente robustas y comprimidas lateralmente que caracterizan a la familia. En cambio, Pteraclis y Pterycombus exhiben una posición anterior extrema de la aleta dorsal sobre el esqueleto craneofacial. En consecuencia, poseen anatomías de aletas y cráneo que son radicalmente diferentes de otras especies de bramidas. Aquí, investigamos la anatomía, el desarrollo y la evolución de Bramidae para probar la hipótesis de que las innovaciones morfológicas tienen un costo funcional (próximo) y evolutivo (último). Al abordar los efectos inmediatos, encontramos que el desarrollo de una aleta dorsal exagerada se asocia con neurocráneo modificado para adaptarse a una expansión anterior de la aleta dorsal. Esto ocurre a través del desarrollo reducido de la cresta supraoccipital (SOC), proporcionando una amplia área de superficie en el cráneo para la inserción de la musculatura de la aleta dorsal. Si bien estos cambios anatómicos presumiblemente están asociados con una mayor maniobrabilidad en los peces fanfishes, también se predice que darán como resultado una alimentación por succión comprometida, lo que posiblemente limite los mecanismos de alimentación en este grupo. Los análisis filogenéticos sugieren que las morfologías craneofaciales y de aletas de los fanfishes evolucionaron rápidamente y están correlacionadas evolutivamente entre las bramidas. Además, los fanfishes exhiben una tasa similar de diversificación de linajes que el resto de los Bramidae, lo que brinda poco apoyo a la predicción de que las aletas mediales exageradas están asociadas con restricciones filogenéticas. Nuestra filogenia coloca a los peces abanico en la base de las Bramidae y sugiere que las bramidas que no son peces abanico tienen aletas mediales reducidas y SOC reevolucionado. Estas observaciones sugieren que la evolución de nuevas morfologías de aletas en especies basales ha llevado al acoplamiento filogenético de la forma de la cabeza y la aleta, lo que posiblemente predisponga a toda la familia a un rango limitado de alimentación. Por lo tanto, la evolución de morfologías extremas puede tener efectos de arrastre, incluso después de que se pierde la morfología, lo que limita la diversificación ecológica de los linajes.

Quando surgem morfologias novas ou extremas, muitas vezes enfrentam o fardo de compensações funcionais em outros aspectos da anatomia, que podem limitar a diversificação fenotípica e tornar determinados picos adaptativos inacessíveis. Bramids (Perciformes: Bramidae) compreendem uma pequena família de 20 espécies existentes de peixes, que estão distribuídos em águas pelágicas em todo o mundo. Dentro dos Bramidae, os fanfishes (Pteraclis e Pterycombus) diferem morfologicamente das espécies geralmente robustas e comprimidas lateralmente que tipificam a família. Em vez disso, Pteraclis e Pterycombus exibem posicionamento anterior extremo da nadadeira dorsal no esqueleto craniofacial. Conseqüentemente, eles possuem anatomias de barbatana e crânio que são radicalmente diferentes de outras espécies de bramida. Aqui, investigamos a anatomia, o desenvolvimento e a evolução dos Bramidae para testar a hipótese de que as inovações morfológicas têm custos funcionais (proximais) e evolutivos (finais). Abordando os efeitos imediatos, descobrimos que o desenvolvimento de uma nadadeira dorsal exagerada está associado a neurocrania modificada para acomodar uma expansão anterior da nadadeira dorsal. Isso ocorre por meio do desenvolvimento reduzido da crista supraoccipital (SOC), proporcionando uma ampla área de superfície no crânio para a inserção da musculatura da nadadeira dorsal. Embora essas mudanças anatômicas estejam presumivelmente associadas a maior capacidade de manobra em peixes-leque, também se prevê que resultem em alimentação de sucção comprometida, possivelmente limitando os mecanismos de alimentação neste grupo. As análises filogenéticas sugerem que as morfologias craniofaciais e das nadadeiras de fanfishes evoluíram rapidamente e estão evolutivamente correlacionadas entre as bramidas. Além disso, fanfishes exibem uma taxa semelhante de diversificação de linhagem como o resto dos Bramidae, emprestando pouco suporte para a previsão de que nadadeiras mediais exageradas estão associadas a restrições filogenéticas. Nossa filogenia coloca fanfishes na base dos Bramidae e sugere que bramids não fanfish possuem nadadeiras mediais reduzidas e SOCs re-evoluídos. Essas observações sugerem que a evolução de novas morfologias de nadadeiras em espécies basais levou ao acoplamento filogenético da forma da cabeça e da nadadeira, possivelmente predispondo toda a família a uma faixa limitada de alimentação. Assim, a evolução de morfologias extremas pode ter efeitos de transporte, mesmo após a perda da morfologia, limitando a diversificação ecológica das linhagens.

Diversity in rest-activity patterns among Lake Malawi cichlid fishes suggests a novel axis of habitat partitioning.

Animals display remarkable diversity in rest and activity patterns that are regulated by endogenous foraging strategies, social behaviors and predator avoidance. Alteration in the circadian timing of activity or the duration of rest-wake cycles provide a central mechanism for animals to exploit novel niches. The diversity of the >3000 cichlid species throughout the world provides a unique opportunity to examine variation in locomotor activity and rest. Lake Malawi alone is home to over 500 species of cichlids that display divergent behaviors and inhabit well-defined niches throughout the lake. These species are presumed to be diurnal, though this has never been tested systematically. Here, we measured locomotor activity across the circadian cycle in 11 Lake Malawi cichlid species. We documented surprising variability in the circadian time of locomotor activity and the duration of rest. In particular, we identified a single species, Tropheops sp. 'red cheek', that is nocturnal. Nocturnal behavior was maintained when fish were provided shelter, but not under constant darkness, suggesting that it results from acute response to light rather than an endogenous circadian rhythm. Finally, we showed that nocturnality is associated with increased eye size after correcting for evolutionary history, suggesting a link between visual processing and nighttime activity. Together, these findings identify diversity of locomotor behavior in Lake Malawi cichlids and provide a system for investigating the molecular and neural basis underlying variation in nocturnal activity.

Weak genetic signal for phenotypic integration implicates developmental processes as major regulators of trait covariation.

Phenotypic integration is an important metric that describes the degree of covariation among traits in a population, and is hypothesized to arise due to selection for shared functional processes. Our ability to identify the genetic and/or developmental underpinnings of integration is marred by temporally overlapping cell-, tissue- and structure-level processes that serve to continually 'overwrite' the structure of covariation among traits through ontogeny. Here, we examine whether traits that are integrated at the phenotypic level also exhibit a shared genetic basis (e.g. pleiotropy). We micro-CT scanned two hard tissue traits, and two soft tissue traits (mandible, pectoral girdle, atrium and ventricle, respectively) from an F5 hybrid population of Lake Malawi cichlids, and used geometric morphometrics to extract 3D shape information from each trait. Given the large degree of asymmetric variation that may reflect developmental instability, we separated symmetric from asymmetric components of shape variation. We then performed quantitative trait loci (QTL) analysis to determine the degree of genetic overlap between shapes. While we found ubiquitous associations among traits at the phenotypic level, except for a handful of notable exceptions, our QTL analysis revealed few overlapping genetic regions. Taken together, this indicates developmental interactions can play a large role in determining the degree of phenotypic integration among traits, and likely obfuscate the genotype to phenotype map, limiting our ability to gain a comprehensive picture of the genetic contributors responsible for phenotypic divergence.

Ecomorphological divergence and habitat lability in the context of robust patterns of modularity in the cichlid feeding apparatus.

BACKGROUND: Adaptive radiations are characterized by extreme and/or iterative phenotypic divergence; however, such variation does not accumulate evenly across an organism. Instead, it is often partitioned into sub-units, or modules, which can differentially respond to selection. While it is recognized that changing the pattern of modularity or the strength of covariation (integration) can influence the range or rate of morphological evolution, the relationship between shape variation and covariation remains unclear. For example, it is possible that rapid phenotypic change requires concomitant changes to the underlying covariance structure. Alternatively, repeated shifts between phenotypic states may be facilitated by a conserved covariance structure. Distinguishing between these scenarios will contribute to a better understanding of the factors that shape biodiversity. Here, we explore these questions using a diverse Lake Malawi cichlid species complex, Tropheops, that appears to partition habitat by depth. RESULTS: We construct a phylogeny of Tropheops populations and use 3D geometric morphometrics to assess the shape of four bones involved in feeding (mandible, pharyngeal jaw, maxilla, pre-maxilla) in populations that inhabit deep versus shallow habitats. We next test numerous modularity hypotheses to understand whether fish at different depths are characterized by conserved or divergent patterns of modularity. We further examine rates of morphological evolution and disparity between habitats and among modules. Finally, we raise a single Tropheops species in environments mimicking deep or shallow habitats to discover whether plasticity can replicate the pattern of morphology, disparity, or modularity observed in natural populations. CONCLUSIONS: Our data support the hypothesis that conserved patterns of modularity permit the evolution of divergent morphologies and may facilitate the repeated transitions between habitats. In addition, we find the lab-reared populations replicate many trends in the natural populations, which suggests that plasticity may be an important force in initiating depth transitions, priming the feeding apparatus for evolutionary change.

Evolution of a soft-tissue foraging adaptation in African cichlids: Roles for novelty, convergence, and constraint.

Understanding the origins of biodiversity demands consideration of both extrinsic (e.g., ecological opportunity) and intrinsic (e.g., developmental constraint) factors. Here, we use a combination of phylogenetic and genetic tools to address the origin of novelty in African cichlids. In particular, we focus on an extreme hypertrophied snout that is structurally integrated with the upper jaw. We show that this bizarre trait has evolved independently in at least two distinct and ecologically successful cichlid clades. We find that snout dimensions are decoupled both phenotypically and genetically, which has enabled it to evolve independently in multiple directions. Further, patterns of variation among species and within a genetic mapping pedigree suggest that relative to snout length, depth is under greater genetic and/or developmental constraint. Models of evolution suggest that snout shape is under selection for feeding behavior, with snout depth being important for algae scraping and snout length for sand sifting. Indeed, the deep snout of some algivores is achieved via an expansion of the intermaxillary ligament, which is important for jaw stability and may increase feeding performance. Overall, our data imply that the evolution of exaggerated snout depth required overcoming a genetic/developmental constraint, which led to expanded ecological opportunity via foraging adaptation.

Muscle-induced loading as an important source of variation in craniofacial skeletal shape.

The shape of the craniofacial skeleton is constantly changing through ontogeny and reflects a balance between developmental patterning and mechanical-load-induced remodeling. Muscles are a major contributor to producing the mechanical environment that is crucial for "normal" skull development. Here, we use an F5 hybrid population of Lake Malawi cichlids to characterize the strength and types of associations between craniofacial bones and muscles. We focus on four bones/bone complexes, with different developmental origins, alongside four muscles with distinct functions. We used micro-computed tomography to extract 3D information on bones and muscles. 3D geometric morphometrics and volumetric measurements were used to characterize bone and muscle shape, respectively. Linear regressions were performed to test for associations between bone shape and muscle volume. We identified three types of associations between muscles and bones: weak, strong direct (i.e., muscles insert directly onto bone), and strong indirect (i.e., bone is influenced by muscles without a direct connection). In addition, we show that although the shape of some bones is relatively robust to muscle-induced mechanical stimulus, others appear to be highly sensitive to muscular input. Our results imply that the roles for muscular input on skeletal shape extend beyond specific points of origin or insertion and hold significant potential to influence broader patterns of craniofacial geometry. Thus, changes in the loading environment, either as a normal course of ontogeny or if an organism is exposed to a novel environment, may have pronounced effects on skeletal shape via near and far-ranging effects of muscular loading.

Genetic and developmental origins of a unique foraging adaptation in a Lake Malawi cichlid genus.

Phenotypic novelties are an important but poorly understood category of morphological diversity. They can provide insights into the origins of phenotypic variation, but we know relatively little about their genetic origins. Cichlid fishes display remarkable diversity in craniofacial anatomy, including several novelties. One aspect of this variation is a conspicuous, exaggerated snout that has evolved in a single Malawi cichlid lineage and is associated with foraging specialization and increased ecological success. We examined the developmental and genetic origins for this phenotype and found that the snout is composed of two hypertrophied tissues: the intermaxillary ligament (IML), which connects the right and left sides of the upper jaw, and the overlying loose connective tissue. The IML is present in all cichlids, but in its exaggerated form it interdigitates with the more superficial connective tissue and anchors to the epithelium, forming a unique ligament-epithelial complex. We examined the Transforming growth factor ß (Tgfß) → Scleraxis (Scx) candidate pathway and confirmed a role for these factors in snout development. We demonstrate further that experimental up-regulation of Tgfß is sufficient to produce an expansion of scx expression and concomitant changes in snout morphology. Genetic and genomic mapping show that core members of canonical Tgfß signaling segregate with quantitative trait loci (QTL) for snout variation. These data also implicate a candidate for ligament development, adam12, which we confirm using the zebrafish model. Collectively, these data provide insights into ligament morphogenesis, as well as how an ecologically relevant novelty can arise at the molecular level.

The Influence of Climatic Variability on Morphological Integration, Evolutionary Rates, and Disparity in the Carnivora.

Biodiversity is unevenly distributed in space and time. One possible explanation for this is the influence of climate on the ecology, evolution, and morphology of taxa. Here we investigated the link between climatic variability and phenotypic integration, rates of morphological evolution, and disparity (morphological diversity) in three carnivoran clades (Canidae, Felidae, and Mustelidae). We gathered landmark data from the lower jaw and extracted current temperature and precipitation data from range maps. We found a significant negative relationship between climatic variability and integration for canids and felids. Among canids, variability in temperature was the key climatic variable, while in felids it was a combination of variability in temperature and precipitation. In both cases, relatively variable climates were associated with low phenotypic integration. We also found evidence for a negative association between climatic variability and both disparity and rates of morphological evolution in canids and mustelids. Selection can drive the evolution of jaw shape along lines of least resistance defined by patterns of integration, and this study suggests that climate may be a predictor of phenotypic integration. As a result, taxa in more variable regions (e.g., temperate, montane) may be more evolvable and more able to respond to fluctuating environmental conditions over a period of generations.

The evolution of jaw protrusion mechanics is tightly coupled to bentho-pelagic divergence in damselfishes (Pomacentridae).

Most species-rich lineages of aquatic organisms have undergone divergence between forms that feed from the substrate (benthic feeding) and forms that feed from the water column (pelagic feeding). Changes in trophic niche are frequently accompanied by changes in skull mechanics, and multiple fish lineages have evolved highly specialized biomechanical configurations that allow them to protrude their upper jaws toward the prey during feeding. Damselfishes (family Pomacentridae) are an example of a species-rich lineage with multiple trophic morphologies and feeding ecologies. We sought to determine whether bentho-pelagic divergence in the damselfishes is tightly coupled to changes in jaw protrusion ability. Using high-speed video recordings and kinematic analysis, we examined feeding performance in 10 species that include three examples of convergence on herbivory, three examples of convergence on omnivory and two examples of convergence on planktivory. We also utilized morphometrics to characterize the feeding morphology of an additional 40 species that represent all 29 damselfish genera. Comparative phylogenetic analyses were then used to examine the evolution of trophic morphology and biomechanical performance. We find that pelagic-feeding damselfishes (planktivores) are strongly differentiated from extensively benthic-feeding species (omnivores and herbivores) by their jaw protrusion ability, upper jaw morphology and the functional integration of upper jaw protrusion with lower jaw abduction. Most aspects of cranial form and function that separate these two ecological groups have evolved in correlation with each other and the evolution of the functional morphology of feeding in damselfishes has involved repeated convergence in form, function and ecology.

The functional significance of morphological changes in the dentitions of early mammals.

The Mesozoic marked a time of experimentation in the tooth morphology of early mammals. One particular experiment involved the movement of three points, or cusps, on the surface of a molar tooth from a line into a triangle. This transition is exemplified by two extinct insectivorous mammals, Morganucodon (cusps in a line) and Kuehneotherium (cusps in a triangle). Here we test whether this difference in cusp arrangement, alongside cusp heights and angles between cusps, is associated with differences in the ability of the teeth to fracture proxy-insect prey. We gathered measurements from molar teeth of both species and used them to create physical models. We then measured the force, time and energy at fracture and peak force, and the amount of damage inflicted by the models on hard and soft gels encased in a tough film that mimicked the material properties of insects. The Morganucodon model required less force and energy to fracture hard gels and reach peak force compared with KuehneotheriumKuehneotherium required a similar time, force and energy to fracture soft gels but reduced the time, force and energy to reach peak force. More importantly, Kuehneotherium also inflicted more damage to both the hard and the soft gels. These results suggest that changes in dental morphology in some early mammals was driven primarily by selection for maximizing damage, and secondarily for maximizing biomechanical efficiency for a given food material property.