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Studies of vertebrate brain evolution have mainly focused on measures of brain size, particularly relative mass and its allometric scaling across lineages, commonly with the goal of identifying the substrates that underly differences in cognition. However, recent studies on birds and mammals have demonstrated that brain size is an imperfect proxy for neuronal parameters that underly function, such as the number of neurons that make up a given brain region. Here we present estimates of neuron numbers and density in two species of lizard, Anolis cristatellus and A. evermanni, representing the first such data from squamate species, and explore its implications for differences in cognitive performance and vertebrate brain evolution. The isotropic fractionator protocol outlined in this article is optimized for the unique challenges that arise when using this technique with lineages having nucleated erythrocytes and relatively small brains. The number and density of neurons and other cells we find in Anolis for the telencephalon, cerebellum, and the rest of the brain (ROB) follow similar patterns as published data from other vertebrate species. Anolis cristatellus and A. evermanni exhibited differences in their performance in a motor task frequently used to evaluate behavioral flexibility, which was not mirrored by differences in the number, density, or proportion of neurons in either the cerebellum, telencephalon, or ROB. However, the brain of A. evermanni had a significantly higher number of nonneurons and a higher nonneuron to neuron ratio across the whole brain, which could contribute to the observed differences in problem solving between A. cristatellus and A. evermanni. Although limited to two species, our findings suggest that neuron number and density in lizard brains scale similarly to endothermic vertebrates in contrast to the differences observed in brain to body mass relationships. Data from a wider range of species are necessary before we can fully understand vertebrate brain evolution at the neuronal level.

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Across vertebrates, there is a broad correlation between neuroanatomy and the type of habitat preferred by a species. In general, species occupying habitats classified as more structurally complex have relatively larger brains and exaggerated structures related to navigating and exploiting those habitats. We empirically measured the structural habitat complexity of six species of Puerto Rican Anolis lizards, which have traditionally been classified as occupying three distinct habitat types. We also measured the volume of the whole brain as well as six structures putatively related to exploiting complex habitats in these species. We found a significant interspecific variation in structural habitat complexity, including a substantial variation between those belonging to the same ecomorph category. Despite this, we found no evidence to support the hypothesis that interspecific differences in habitat preferences, particularly differences in the relative structural complexity of those habitats, can favor a divergence in neuroanatomy. However, our findings indicate that, at a finer scale, species preferences for structural habitats vary greatly between Anolis species belonging to the same ecomorph category. This variation might contribute to the community structure of anoles by allowing multiple sympatric species of the same ecomorph category to occupy what, at a coarse scale, appears to be the same structural niche. We propose that, in the case of arboreal species, differences in the complexity of arboreal habitats alone may not be sufficient to favor divergent brain evolution.

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In many forensic cases, the job of forensic pathologists and anthropologists is to determine whether pediatric death is due to an abusive act or an accidental fall. The goal of this study was to compare the cranial fracture patterns generated on the parietal bone of a developing, infant porcine (pig, Sus scrofa) model by a controlled energy head drop onto a plate versus previous data generated by blunt force impact at the same energy onto the head constrained to a plate. The results showed that blunt force impacts on a head constrained to a rigid plate produces more fracture, but the same general pattern, as that for a head dropped onto the plate with the same level of impact energy. The study suggests that head constraint may be an important factor to consider in the evaluation of death causation for blunt force impacts to the pediatric skull.

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Patterns of brain evolution have been widely studied across vertebrates, with the bulk of studies using mammals and/or birds as model systems. Within these groups, species occupying different habitats have been shown to have divergent neuroanatomy, particularly with regard to differences in the relative size of different brain structures, correlated with differences in habitat complexity. We examined the pattern of allometric scaling across the telencephalon, dorsal cortex, dorsomedial cortex, medial cortex, dorsal ventricular ridge, medulla and cerebellum in six species of Puerto Rican Anolis lizards, which are grouped in three distinct ecomorphs (i.e. ecological types) according to interspecific differences in preferred habitat type. The differences in habitat preferences are accompanied by morphological and behavioral adaptations for effective use of each habitat type. Our results challenge this trend and demonstrate a lack of convergence in the relative size of different brain structures between species belonging to the same ecomorph type. Overall brain volume explained between 92.5 and 99.8% of the variance in the volume of each of the brain regions measured and 93.8 and 98.5% of the variance in the volume of each component measured within the telencephalon. This pattern of brain allometry is consistent with concerted brain evolution. However, in the case of the cerebellum, interspecific differences in volume exhibit a trend in accordance with mosaic brain evolution. This suggests that both concerted and mosaic brain evolution have shaped the anole brain, with the former playing a dominant role. Concerted brain evolution is the primary mechanism shaping the brain in mammals and cartilaginous fishes, and its presence in Anolis lizards provides additional evidence supporting the hypothesis that concerted brain evolution might result from a conserved pattern of brain development common to all vertebrates. More generally, our findings highlight the necessity of further studies of brain evolution in reptiles as they can provide valuable insights into the mechanisms underlying vertebrate brain evolution.

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The role of behavioural flexibility in responding to new or changing environmental challenges is a central theme in cognitive ecology. Studies of behavioural flexibility have focused mostly on mammals and birds because theory predicts that behavioural flexibility is favoured in species or clades that exploit a diversity of habitats or food sources and/or have complex social structure, attributes not associated with ectothermic vertebrates. Here, we present the results of a series of experiments designed to test cognitive abilities across multiple cognitive modules in a tropical arboreal lizard: Anolis evermanni. This lizard shows behavioural flexibility across multiple cognitive tasks, including solving a novel motor task using multiple strategies and reversal learning, as well as rapid associative learning. This flexibility was unexpected because lizards are commonly believed to have limited cognitive abilities and highly stereotyped behaviour. Our findings indicate that the cognitive abilities of A. evermanni are comparable with those of some endothermic species that are recognized to be highly flexible, and strongly suggest a re-thinking of our understanding of the cognitive abilities of ectothermic tetrapods and of the factors favouring the evolution of behavioural flexibility.

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The objective of this study was to document patterns of fracture on infant porcine skulls aged 2-28 days (n = 57) because of a single, high energy blunt impact to the parietal bone with rigid (nondeformable) and compliant (deformable) interfaces. Fracture patterns were mapped using Geographic Information System software. For the same generated impact force, the rigid interface produced more fractures than the compliant interface for all ages. This study also showed that this increased level of impact energy versus an earlier study using a lower energy resulted in new sites of fracture initiation and also caused previously defined fractures that propagate into an adjacent bone. Several unique characteristics of bone and diastatic fracture were documented as a function of specimen age, impact energy, and interface. These data describe some baseline characteristics of skull fracture using an animal model that may help guide future studies from forensic case files.

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This study documents skull fracture characteristics on infant porcine specimens under known impact conditions with respect to age and interface. A single impact causing fracture was conducted on the skull of porcine specimens aged 2-28 days (n = 76). Paired rigid and compliant impacts at the same energy were conducted at each specimen age. Impact force, impact duration, and fracture length were recorded. Energy required to initiate skull fracture increased with specimen age. For a given energy, impact of the skull with a compliant interface caused more fracture damage than with a rigid interface for specimens aged under 17 days, but less damage for specimens aged 24-28 days. The documentation of energy required to cause fracture and resulting fracture propagation with respect to impact interface and age may be of critical importance in forensic investigations of infant skull trauma.

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An infant less than 18 months of age with a skull fracture has a one in three chance of abuse. Injury biomechanics are often used in the investigation of these cases. In addition to case-based investigations, computer modeling, and test dummies, animal model studies can aid in these investigations. This study documents age effects on the mechanical properties of parietal bone and coronal suture in porcine infants and correlates the bending properties of the bone to existing human infant data. Three beam specimens were cut from porcine specimens aged 3 days, 7 days, 10 days, 14 days, 18 days, and 21 days: one across the coronal suture and two from the parietal bone, one parallel to and one perpendicular to the coronal suture. An actuator-mounted probe applied four-point bending in displacement control at 25 mm/s until failure. Bending stiffness of bone specimens increased with age; bone-suture-bone specimens showed no change up to 14 days but increased from 14 days to 18 days. All three specimen types showed decreases in ultimate stress with age. Ultimate strain for the bone-suture-bone specimens was significantly higher than that for the bone specimens up to 14 days with no differences thereafter. There was no change in the bending modulus with age for any specimen type. Bone-suture-bone bending modulus was lower than that of the bone specimens up to 14 days with no differences thereafter. There was no change in strain energy to failure with age for the bone specimens; bone-suture-bone specimens showed no change up to 14 days but decreased from 14 days to 18 days. There was an increase in specimen porosity with age. Correlation analysis revealed a weak (-0.39) but significant and negative correlation between ultimate stress and porosity. While the mechanical properties of parietal bone and coronal suture did not change significantly with age, bone specimens showed an increase in bending stiffness with age. Bone-suture-bone specimens showed an increase in bending stiffness only between 14 days and 18 days of age. Correlation analyses using existing and new data to compute the bending rigidity of infant parietal bone specimens suggested that days of pig age may correlate with months of human age during the most common time frame of childhood abuse cases.

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