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* indicates equal contribution • underline indicates high school/undergraduate student mentee

X) Burtner A, Grossnickle D, Santana S, & Law CJ. In review. Gliding towards an understanding of the origin of flight in bats. Submitted to PeerJ. BioRxiv 2022.09.26.509622

X) Law CJ, Linden T, & Flynn JJ. In review. Skull evolution and lineage diversification in endemic Malagasy carnivorans. Target journal: Royal Society Open Science. BioRxiv 2024.03.25.586658

X) Law CJ, Hlusko LJ, & Tseng ZJ. In review. The carnivoran adaptive landscape reveals trade-offs among functional traits in the skull, appendicular, and axial skeleton. Target journal: Proceedings of Royal Society B. BioRxiv 2024.05.06.592620

X) López-Torres S, Bertrand OC, Fostowicz-Frelik Ł, Lang MM, Law CJ, Martin-Flores GS, Schillaci MA, Silcox MT. In revision. The allometry of brain size in Euarchontoglires and an exploration of group-specific encephalization. Invited to resubmit to Journal of Mammalogy

34) Law CJ, Tinker MT, Fujii, JA, Nicholson T, Staedler M, Tomoleoni J, Young C, & Mehta RS. 2024. Tool use increases foraging success in southern sea otters. Science. 384:798-802.

Press: NPR: Morning Edition, The Guardian, The Hill, ReuterSüdwestrundfunk, Science News

Goal: To investigate the mechanical benefits of tool use behavior and test if tool use leads to a higher rate of energy intake in environments depleted with preferred prey

Findings: Tool use facilitates access to harder or larger prey items that are otherwise inaccessible, especially for female otters

Findings: Tool use reduces tooth damage overall but exhibits a non-linear relationship with dentine damage

Findings: Tool use facilitates two alternative strategies in maintaining energetic intake when preferred prey are depleted: 1) the “high risk, high payoff” strategy: low frequency tool users feed on hard but energetically rich prey like clams, resulting in increased energetic intake. However, these prey are not abundant and are some of the hardest, resulting in some dentine damage. 2) the “low risk, low payoff” strategy: high frequency tool users feed on hard but very abundant prey like snails, resulting in sufficient energetic intake. However, these prey are energetically poor, making the daily total energetic demands difficult to reach 

Conclusions: Tool use allows individuals to maintain energetic requirements through the processing of alternative hard prey that are typically inaccessible with biting alone, suggesting that this behavior is a necessity for the survival of some otters in environments with limited resources.

33) Grossnickle DM, Brightly WH, Pevsner SK, Polly PD, Roston RA, Stanchak KE, Stayton T, Weaver LN, & Law CJ. In press. Challenges and advances in methods for measuring phenotypic convergence. Evolution.

Goal: To investigate the performance of current methods of quantifying phenotypic convergence including C-measures and Ornstein–Uhlenbeck models

Findings: Both measure types are influenced by the position of putatively convergent taxa in morphospace such that morphological outliers are statistically more likely to exhibit convergence by chance

Findings: some methods will misidentify divergent and parallel lineages as convergent

Findings: To help address these problems, we developed a new distance-based method (Ct-measure) that incorporates distances between lineages through time and minimizes the possibility of divergent taxa being misidentified as convergent

Conclusions: Nevertheless, we emphasize that all available convergence measures are imperfect, and researchers should recognize the limitations of all methods and use multiple lines of evidence to test convergence hypotheses.

32) Jiangzuo Q, Wang X, Law CJ, Jia Y, Li S, Fu J, Wang S, Deng T, & Ji X. In press. Presence of Cernictis and Lutravus (Ictonychinae, Mustelidae, Carnivora) in eastern Asia and the dispersal of Ictonychinae during the Late Miocene. Journal of Systematic Paleontology.

31) Law CJ, Hlusko LJ, & Tseng ZJ. 2024. Uncovering the mosaic evolution of the carnivoran skeletal system. Biology Letters. 20:20230526.

Goal: To test if ecological and phylogenetic effects led to distinct modes of evolution among the cranial, appendicular and vertebral regions in extant carnivoran skeletons

Findings: The carnivoran skeleton exhibits mosaic evolution, where only the mandible, hindlimb and posterior (i.e. last thoracic and lumbar) vertebrae showed evidence of adaptation towards ecological regimes

Findings: The remaining skeletal components (i.e., cranium, forelimb, and anterior (cervical and thoracic) vertebrae exhibit clade-specific evolutionary shifts that corresponded well to carnivoran clades.

Findings: Decoupled evolution of individual skeletal components may have led to the origination of distinct adaptive zones and morphologies among extant carnivoran families that reflect phylogenetic hierarchies.

Conclusions: Our work highlights the importance of examining multiple skeletal components simultaneously in ecomorphological analyses. Ongoing work integrating the fossil record will further clarify deep-time drivers that govern the carnivoran diversity we see today and reveal the complexity of evolutionary processes in multicomponent systems.

30) Kemp ME, Boville AE*, Carneiro CM*, Jacisin III JJ*, Law CJ*, Ledesma DT*, Meza A*, Shields-Estrada A*, Xu T*. 2023. Looking back for the future: the ecology of terrestrial communities through the lens of conservation paleobiology. Annual Review of Ecology, Evolution, and Systematics. 54:obad017.

Goal: A review to investigate how conservation paleobiology provides deep-time perspectives on terrestrial organisms, populations, communities, and ecosystems impacted by anthropogenic processes. We also highlight how the colonial legacies of conservation biology confound our understanding of present-day biodiversity, ecosystem processes, and conservation outlooks, and we make recommendations for more inclusive and ethical practices moving forward.

29) Rickman J*, Burtner A*, Linden T, Santana S, & Law CJ*. 2023. Size and locomotor ecology have differing effects on the external and internal morphologies of squirrel (Rodentia: Sciuridae) limb bones. Integrative Organismal Biology. 5:obad017.

Goal: To test the effects of locomotor mode and scaling on the external shape and structure of the two major limb bones in squirrels, the humerus and femur

Findings: Shapes of the humerus and femur (to a lesser extent) are best explained by locomotor ecology rather than by size. Gliders tend to exhibit gracile, elongate limbs; ground squirrels tend to exhibit robust limbs; & chipmunks & tree squirrels exhibit intermediate limbs

Findings: Bone structures are best explained by interactions between locomotor ecology and scaling. As limb size ⬆️ across all squirrels; however, scaling patterns in bone structure variables of the humerus and femur differ among the ecotypes, serving as adaptations to different ecological specialization

Findings: The statistical relationships between limb morphologies and ecotype were lost when accounting for phylogenetic relationships among species under Brownian motion. 

Conclusions: our results suggest that humeral and femoral variation in squirrels partitioned early between clades and their ecomorphologies were maintained to the present 

28) Linden T*, Burtner A, Rickman J, McFeely A, Santana S, & Law CJ*. 2023. Scaling patterns of body plans differ between squirrel ecotypes. PeerJ. e14800.

Goal: To test scaling patterns of body shape, body size, & limb lengths among squirrel ecotypes and describe the underlying morphological components that contribute to body shape

Findings: Body shape scale with body size, but these allometric patterns differ among squirrel ecotypes: chipmunks & gliding squirrels exhibited more elongate bodies with increasing body sizes whereas ground squirrels exhibited more robust bodies with increasing body size 

Findings: Only ground squirrels exhibit a relationship between forelimb length and body shape, where more elongate species exhibit relatively shorter forelimbs

Findings: Relative length of the ribs and elongation or shortening of the thoracic region contributes the most to body shape evolution across squirrels

Conclusions: This work contributes to the growing understanding of mammalian body shape evolution and how it is influenced by body size and locomotor ecology, in this case from robust subterranean to gracile gliding squirrels

27) Law CJ, Blackwell E, Curtis AA, Dickinson E, Hartstone-Rose A, and Santana SE. 2022. Decoupled evolution of crania and mandibles in carnivoran mammals. Evolution. 76:2959-2974.

Goal: To examine the evolutionary patterns of the cranium and mandible in terrestrial carnivoran mammals

Findings: Cranial shape follows clade-based evolutionary shifts, suggesting that the complexity and variation of carnivoran cranial adaptations cannot be captured effectively by dietary categories

Findings: Mandibular shape evolution is linked to broad dietary regimes

Findings: The evolution of both cranial and mandibular size in hypercarnivores is associated with relative prey size, demonstrating that dietary diversity can be loosely structured by craniomandibular size within some guilds

Conclusions: Cranial and mandibular shape exhibit decoupled evolution; carnivoran crania exhibit hierarchical evolution whereas mandibles exhibit greater evolutionary lability with respect to diet

26) Lang MM, Bertrand OC, Flores GSM, Law CJ, Abdul-Sater JAD, Spakowski S, Silcox MT. 2022. Scaling patterns of cerebellar petrosal lobules in Euarchontoglires: impacts of ecology and phylogeny. Anatomical Records. 305:3472-3503.

Goal: To test whether ecology influences relative petrosal lobule size

Findings: Lobule size exhibited strong positive relationships with both endocranial volume and body mass

Findings: Relative petrosal lobule size differed between clades where lagomorphs exhibited relatively larger petrosal lobules

Findings: Fossorial species exhibited relatively smaller petrosal lobules compare  to other locomotor groups; there were no other differences between other ecological groups

Conclusions: Ecological effects on petrosal lobule scaling patterns remain difficult to disentangle across large clades

25) Law CJ. 2022. Different evolutionary pathways lead to incomplete convergence of elongate body shapes in carnivoran mammals. Systematic Biology. 71:788–796.

Goal: To test whether “elongate,” “slender,” and “long” carnivorans exhibit convergence in body shape 

Findings: Qualitatively-described elongate carnivorans exhibited shared adaptive peaks and incomplete convergence towards elongate bodies compared to other terrestrial carnivorans

Findings: In contrast, the morphological components underlying body shape variation do not exhibit convergence despite evidence that these components are more elongate in elongate carnivorans compared to non-elongate carnivorans

Findings: Short but different phylogenetic half-lives associated with each component indicated that they are strongly pulled towards distinct peaks over different adaptive slopes or rates

Conclusions: Different evolutionary pathways from underlying morphological components can lead to incomplete convergence of elongate body shapes in carnivoran mammals

24) Shrestha MB, Shrestha G, Reule S, Oli S, Ghartimagar TB, Singh G, Tripathi DM, Law CJ, Shah KB, & Savage M. 2021. First evidence of Eurasian otter in Nepal in three decades. IUCN-SSC Otter Specialist Group Bulletin. 38:279-291.

Goal: To assess evidence of Eurasian otter presence in Nepal 

Findings: Camera trap images from the Barekot River and the Tubang River confirm the presence of Eurasian otters in Nepal

Findings: Craniomandibular measurements are more similar to Eurasian otter skulls compared to sympatric smooth-coated otters 

Conclusions: Further study would elucidate the distribution of Eurasian otters in this region

23) Higham TE, Ferry LA, Schmitz L, Irschick DJ, Starko S, Anderson PSL, Bergmann PJ, Jamniczky HA, Monteiro LR, Navon D, Messier J, Carrington E, Farina SC, Feilich KL, Hernandez LP, Johnson MA, Kawano SM, Law CJ, Longo SJ, Martin CH, Martone PT, Rico-Guevara A, Santana SE, and Niklas KJ. 2021. Linking biomechanical models and functional traits to investigate phenotypic diversity. Trends in Ecology and Evolution. 36:860-873.

Goal: To create an ecomechanical framework to understanding patterns of species distributions, organism–environment interactions, developmental patterns, and evolutionary processes. 

Findings: We must expand functional trait databases to include biomechanically-meaningful traits, standardize the collection of these biomechanical traits, and increase the access to models using these traits via freely-available online platforms

Findings: Integrating functional organismal traits with the ecological variables will inform our ability to predict species shifts in survival and distribution and provides critical insights into phenotypic diversity.

22) Law CJ. 2021. Ecological drivers of carnivoran body shape evolution. The American Naturalist. 198:406–420. Presentation 🖥 

Goal: To assess whether locomotor mode, hunting behavior, and dietary ecologies influenced body shape evolution in carnivorans

Findings: Carnivorans exhibit clade-based evolutionary shifts in body shape and tis underlying components, indicating species with shared locomotor, hunting, or dietary ecologies do not evolve towards similar morphological optima

Findings: The water to land boundary appears to drive opposing body shape allometry in terrestrial and aquatic carnivorans, highlighting that allometry may be more important than specific ecological traits in influencing body shape evolution

Conclusions: The multidimensionality of body shape may conceal one-to-one mapping of morphology onto ecology, making it difficult to disentangle the complex relationship between morphological evolution, ecological diversity, and phylogeny across macroevolutionary scales

21) Law CJ. 2021. Evolutionary and morphological patterns underlying carnivoran body shape diversity. Evolution. 75:365-375. SICB Presentation 🖥 

Goal: To investigate how the cranial and axial components contribute to the diversity of carnivoran body shapes

Findings: Clade age, but not rates of body shape evolution or species richness, predicted the variation of body shapes across carnivoran families, indicating that older families simply had more evolutionary time to accumulate more disparate body shapes than younger families

Findings: Body shape scaled with negative allometry with respect to body size, where smaller species exhibited more elongate bodies and larger species exhibited more robust bodies

Findings: Evolutionary changes in carnivoran body shape was primarily driven through changes of the thoracic and lumbar regions and rib length, albeit pathways differ between different families

Conclusion: Pathways to body shape variation are diverse but provide a strong morphological foundation for future research investigating the evo-devo and evolutionary ecology of mammalian body shapes.

20) Law CJ. 2020. Sex-specific ontogenetic patterns of cranial morphology, theoretical bite force, and underlying jaw musculature in fishers and American martens. Journal of Anatomy. 237:727-740.

Goal: To examine sex‐specific ontogenetic changes in cranial morphology, musculature, and theoretical bite force in the fisher (Pekania pennanti ) and American marten (Martes americana) 

Findings: Cranial shape development drives ontogenetic increases in relative bite force and jaw adductor muscle size by broadening the zygomatic arches and enlargement of the sagittal crest

Findings: In contrast, sexual dimorphism in cranial shape does not translate to dimorphism in either relative bite forces or jaw adductor muscle size

Conclusion: Morphological variation can have different influences on bite performance depending on the level of intraspecific variation that is examined (i.e. ontogenetic versus sexual dimorphism)

19) Law CJ. 2019. Solitary meat-eaters: solitary, carnivorous carnivorans exhibit the highest degree of sexual size dimorphism. Scientific Reports. 9:15344.

Goal: To determine how social system (as a proxy for sexual selection) and diet (as a proxy for natural selection) influenced the evolution of SSD in terrestrial carnivorans

Findings: Territorial solitary and carnivorous carnivorans exhibited selection towards increased degree of male-biased sexual size dimorphism compared to other carnivorans with alternative social systems and diets

Findings: Most carnivoran families do not follow predictions of Rensch’s rule of increased male-biased sexual size dimorphism with increasing body size

Conclusion: Both sexual selection and niche divergence work simultaneously to reinforce or increase sexual size dimorphism in extant terrestrial carnivorans

18) Law CJ & Mehta RS. 2019. Dry versus wet and gross: Comparisons between the dry skull method and gross dissection-based modeling in bite force estimations. Journal of Morphology. 280:1706–1713.

Goal:  To test how accurately skeletomuscular traits estimated from the dry skull method predicts traits derived from dissection-based biomechanical modeling

Findings: The dry skull method overestimates PCSA of the masseter but underestimates PCSA of the temporalis

Findings: Sexual dimorphism in skull shape affects temporalis PCSA estimations; the dry skull method predicted female temporalis PCSA well but underestimates male temporalis PCSA

Conclusion: The dry skull method provides an underestimation of theoretical bite force over ontogeny, and the underlying anatomical components driving bite force may be misrepresented

17) Law CJ. 2019. Evolutionary shifts in extant mustelid (Mustelidae: Carnivora) cranial shape, body size, and body shape coincides with the Mid-Miocene Climate Transition. Biology Letters. 15:20190155.

Goal: To examine the timing of evolutionary shifts in cranial shape, body size and body shape within extant mustelids (martens, otters, polecats and weasels) during the climatic and environmental changes of the Cenozoic

Findings: Extant mustelid subclades evolved more elongate body plans followed by concurrent shifts towards smaller body sizes and more robust crania during the Late Miocene

Findings: These cranial and body adaptations allowed mustelids to exploit new resources, which in turn facilitated significant increases in species richness

Conclusion: Future work incorporating the fossil record is needed to understand whether selection for smaller, more elongate bodies and stronger jaws occurred within crown-ward mustelids or if extinct ancestors already exhibited those characteristics

16) Law CJ, Slater GJ, & Mehta RS. 2018. Shared extremes by ectotherms and endotherms: Body elongation in mustelids is associated with small size and reduced limbs. Evolution. 73:735–749.

Goal: To tested if mustelid subclades exhibited evolutionary shifts towards more elongate bodies compared to the remaining clade

Findings: Weasels are elongate! Specifically, mustelids exhibit an evolutionary transition toward more elongate bodies driven by the elongation of the head and various regions of the axial skeleton

Findings: The elongation of the body is associated with the evolutionary reduction of body size and a reduction in forelimb length but not hindlimb length

Conclusion: The relationship between body elongation and forelimb length has not previously been quantitatively established for mammals but is consistent with trends exhibited by ectothermic vertebrates and suggests a common pattern of trait covariance associated with body shape evolution

15) Law CJ, Duran E, Hung N, Richards E, Santillan I, & Mehta RS. 2018. Effects of diet on cranial morphology and biting ability in musteloid mammals. Journal of Evolutionary Biology. 31:1918-1931. 

Goal: To examine relationships between cranial size and shape, estimated bite force, and dietary ecologies across Musteloidea

Findings: Cranial size but not cranial shape predicts estimated bite forces in musteloids; many-to-one mapping of form to function may explain this pattern as the cranium is complex, and a variety of morphological shape differences contribute to similar relative biting performance outcomes

Findings: Musteloids with different diets exhibit different cranial shapes; however, they have similar estimated bite forces suggesting that other feeding performance metrics and potentially nonfeeding traits are also important contributors to cranial evolution

14) Higgins BA, Law CJ, & Mehta RS. 2018. Eat whole and less often: ontogenetic shift reveals size specialization on kelp bass by the California moray eel, Gymnothorax mordax. Oecologia. 188:875–887. 

Findings: California morays are dietary specialists, feeding primarily on kelp bass despite not being the most commonly occurring fish species in the environment

Findings: California morays exhibit an ontogenetic shift for kelp bass where maximum prey size increases with moray size but smaller prey are dropped from the diet

Findings: Moray bite forces increased disproportionately with increasing head size, suggesting that larger morays can consume prey that exceed their head lengths; however, there was no significant relationship between prey size and moray bite force, indicating that prey size selection is not limited by moray bite force

13) Law CJ & Mehta RS. 2018. Carnivory maintains cranial dimorphism between males and females: Evidence for niche divergence in extant Musteloidea. Evolution. 72:1950–1961.

Findings: Niche divergence rather than sexual selection maintains the evolution of sexual dimorphism in cranial size and bite force across Musteloidea

Findings: Hypercarnivorous musteloids exhibit the greatest degree of cranial size and bite force dimorphism; in contrast, neither dietary regime nor social system influenced the evolution of sexual dimorphism in cranial shape

Conclusion: These results demonstrate the importance of diet in reducing intraspecific competition for resources as an important mechanism that maintains the evolution of sexual dimorphism in extant musteloids.

12) Law CJ, Slater G, & Mehta RS. 2018. Lineage diversity and size disparity in Musteloidea: testing patterns of adaptive radiation using molecular and fossil-based methods. Systematic Biology. 67:127–144.

Findings: Musteloids do not exhibit an adaptive radiation as previously hypothesized; however, a subclade of “elongate” mustelids exhibit increased lineage carrying capacity and increased rates of body length evolution but not body mass evolution

Findings: Lack of correspondence in rates of body mass and length evolution suggest that phenotypic evolutionary rates under a single morphological metric may not capture the evolution of diversity in clades that exhibit elongate body shapes

Conclusion: These results suggest that body elongation led to the radiation of some musteloids

11) Jones K† & Law CJ†. 2018. Differentiation of craniomandibular morphology in two sympatric Peromyscus mice (Cricetidae: Rodentia). Mammal Research. 63:277–283. †Joint first authors 

Findings: Granivorous Pinyon mice (Peromyscus truei) exhibit skulls that facilitate relatively larger jaw adductor muscles compared to insectivorous California mice (Peromyscus californicus)

Findings: Pinyon mice also exhibit higher mechanical advantage of the masseter jaw muscle

Conclusion: These traits are consistent with the dietary differences exhibited by the two sympatric species

10) Law CJ. 2018. Mustela sibirica (Carnivora: Mustelidae). Mammalian Species. 50:109-118.

Goal: Provide a comprehensive literature review of the Siberian weasel

9) Kienle SS, Law CJ, Costa DP, Berta A, & Mehta RS. 2017. Revisiting the behavioural framework of feeding in predatory aquatic mammals. Proceedings of the Royal Society B. 284:20171035.

An aquatic tetrapod feeding cycle with 5 stages—ingestion, intraoral transport, processing, water removal and swallowing—allows aquatic mammals to be examined in the same framework as other tetrapods and provides flexibility to accommodate behaviorally diverse lineages

Evolution of aquatic tetrapod feeding is not a progression of linear events and should be modeled as a tree-like process

8) Law CJ, Baliga VB, Tinker MT, & Mehta RS. 2017. Asynchrony in craniomandibular development and growth in Enhydra lutris nereis (Carnivora: Mustelidae): Are southern sea otters born to bite? Biological Journal of Linnean Society. 121:420-438

Findings: The majority of sea otter skull growth and development occurs during the pup stage; however, the skull does not fully mature until adulthood and therefore may constrain feeding on hard-shelled prey

Findings: Sexual dimorphism in adult skulls arose through differences in developmental and growth rates and duration

Findings: Male crania mature faster to presumably reach adult biting ability sooner, gaining a competitive advantage in obtaining food and in male–male agonistic interactions

7) Hung N & Law CJ. 2016. Lutra lutra (Carnivora: Mustelidae). Mammalian Species. 48:109–122.

Goal: Provide a comprehensive literature review of the Eurasian otter

6) Law CJ, Young C, and Mehta RS. 2016. Ontogenetic scaling of theoretical bite forces in southern sea otters (Enhydra lutris nereis). Physiological and Biochemical Zoology 89:347-363.

Findings: Sea otters exhibit positive allometric increases in theoretical bite force, which is primarily driven by allometric increases in temporalis muscle mass

Findings: Adult males exhibited greater bite forces as a result of their larger sizes

Findings: However, scaling patterns of theoretical bite force and morphological traits do not differ between the sexes

5) Law CJ, Venkatram V, & Mehta RS. 2016. Sexual dimorphism of craniomandibular morphology in southern sea otters (Enhydra lutris nereis). Journal of Mammalogy. 97:1764-1773.

Findings: Male sea otters exhibit larger skulls compared to females

Findings: Sea otters also exhibit significant shape differences driven by 3 craniomandibular traits that increase the surface area of jaw muscle attachment sites

Findings: These morphological differences indicate males and females may also exhibit differences in biting abilities

4) Baliga VB & Law CJ. 2016. Cleaners amongst wrasses: phylogenetics and evolutionary patterns of cleaning behavior within Labridae. Molecular Phylogenetics and Evolution. 94A:424–435.

3) Law CJ, Dorgan KM, & Rouse GW. 2014. Relating divergence in polychaete musculature to different burrowing behaviors: A study using Opheliidae (Annelida). Journal of Morphology. 571:548–571.

2) Dorgan KM, Law CJ, & Rouse GW. 2013. Meandering worms: Mechanics of undulatory burrowing in muds. Proceedings of the Royal Society B. 280:20122948.

1) Law CJ, Dorgan KM, & Rouse GW. 2013. Validation of three sympatric Thoracophelia species (Annelida: Opheliidae) from Dillon Beach, California using mitochondrial and nuclear DNA sequence data. Zootaxa. 3608:67–74.

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