Unexpected bat community changes along an urban-rural gradient in the Berlin-Brandenburg metropolitan area.

Urbanization gradients are increasingly used in ecological studies to discover responses of species communities to different intensities of human-induced habitat transformation. Here, we investigated patterns of bat communities against the background of different urbanization levels using a priori defined urbanization categories based on distance classes (5 km intervals) along a linear transect from the urban core of the city of Berlin westwards into the rural outskirts of the state of Brandenburg. Using linear-mixed effects models, we found that "distance class", as a proxy for urbanization level, is a meaningful and suitable predictor of bat species richness and diversity. We observed an unexpectedly sudden increase in bat species richness and diversity and changes in species-specific activity levels relatively close to the urban center at the transition between urban and peri-urban areas. This change suggests a relevant influence of the peri-urban areas as a "buffer zone" for specific bat species not able to adapt to the heavily modified inner core of the metropolitan area. Although we could demonstrate that anthropogenic noise and artificial light have the potential to predict the variability of bat species activity along the urban-rural gradient, the actual influence on observed shifts in the bat community needs further research.

Tiger beetles produce anti-bat ultrasound and are probable Batesian moth mimics.

Echolocating bats and their eared insect prey are in an acoustic evolutionary war. Moths produce anti-bat sounds that startle bat predators, signal noxiousness, mimic unpalatable models and jam bat sonar. Tiger beetles (Cicindelidae) also purportedly produce ultrasound in response to bat attacks. Here we tested 19 tiger beetle species from seven genera and showed that they produce anti-bat signals to playback of authentic bat echolocation. The dominant frequency of beetle sounds substantially overlaps the sonar calls of sympatric bats. As tiger beetles are known to produce defensive chemicals such as benzaldehyde and hydrogen cyanide, we hypothesized that tiger beetle sounds are acoustically advertising their unpalatability. We presented captive big brown bats (Eptesicus fuscus) with seven different tiger beetle species and found that 90 out of 94 beetles were completely consumed, indicating that these tiger beetle species are not aposematically signalling. Instead, we show that the primary temporal and spectral characteristics of beetle warning sounds overlap with sympatric unpalatable tiger moth (Arctinae) sounds and that tiger beetles are probably Batesian mimics of noxious moth models. We predict that many insect taxa produce anti-bat sounds and that the acoustic mimicry rings of the night sky are hyperdiverse.

All-you-can-eat buffet: A spider-specialized bat species (Myotis emarginatus) turns into a pest fly eater around cattle.

Determining the dietary spectrum of European insectivorous bats over time is the cornerstone of their conservation, as it will aid our understanding of foraging behavior plasticity in response to plummeting insect populations. Despite the global decline in insects, a restricted number of arthropod pest species thrive. Yet past research has overlooked the potential of European bats to suppress pests harmful to woodlands or livestock, in spite of their economic relevance. Here we investigated the diet composition, its breeding season variations and pest consumption of an insectivorous bat species (Myotis emarginatus), at the northern edge of its range (Wallonia, Belgium). We also explored the prey ecology to gain insight into the hunting strategies and foraging habitats of this bat species. We used DNA metabarcoding to amplify two COI markers within 195 bat droppings collected in June, July and August, thereby identifying 512 prey taxa predominated by Diptera, Araneae and Lepidoptera. Overall, in 97% of the samples we detected at least one of the 58 potential pest taxa, 41 of which targeting trees. The June samples were marked by a diet rich in orb-weaver spiders, in accordance with the archetypal diet of M. emarginatus bats. However, during the highly energy demanding July-August parturition and lactation period, roughly 55% of the dropping samples contained two cattle fly pests (Stomoxys calcitrans and Musca domestica). Moreover, among the 88 Diptera species preyed upon by M. emarginatus in July and August, these flies accounted for around 50% of the taxa occurrences. This plasticity-the switch from a spider-rich to a fly-rich diet-seems providential considering the dramatic ongoing drop in insect populations but this involves ensuring bat-friendly cattle farming. Our results revealed that bats widely consume pest entomofauna, thereby highlighting their potential role as allies of forest managers and farmers.

Bat2Web: A Framework for Real-Time Classification of Bat Species Echolocation Signals Using Audio Sensor Data.

Bats play a pivotal role in maintaining ecological balance, and studying their behaviors offers vital insights into environmental health and aids in conservation efforts. Determining the presence of various bat species in an environment is essential for many bat studies. Specialized audio sensors can be used to record bat echolocation calls that can then be used to identify bat species. However, the complexity of bat calls presents a significant challenge, necessitating expert analysis and extensive time for accurate interpretation. Recent advances in neural networks can help identify bat species automatically from their echolocation calls. Such neural networks can be integrated into a complete end-to-end system that leverages recent internet of things (IoT) technologies with long-range, low-powered communication protocols to implement automated acoustical monitoring. This paper presents the design and implementation of such a system that uses a tiny neural network for interpreting sensor data derived from bat echolocation signals. A highly compact convolutional neural network (CNN) model was developed that demonstrated excellent performance in bat species identification, achieving an F1-score of 0.9578 and an accuracy rate of 97.5%. The neural network was deployed, and its performance was evaluated on various alternative edge devices, including the NVIDIA Jetson Nano and Google Coral.

White-nose syndrome, winter duration, and pre-hibernation climate impact abundance of reproductive female bats.

White-nose syndrome (WNS) is an infectious disease that disrupts hibernation in bats, leading to premature exhaustion of fat stores. Though we know WNS does impact reproduction in hibernating female bats, we are unsure how these impacts are exacerbated by local climate factors. We compiled data from four southeastern U.S. states and used generalized linear mixed effects models to compare effects of WNS, pre-hibernation climate variables, and winter duration on the number of reproductive females in species across the range of WNS susceptibility. We predicted we would see a decline in the number of reproductive females in WNS-susceptible species, with the effect exaggerated by longer winter durations and pre-hibernation climate variables that lead to reductions in foraging. We found that the number of reproductive females in WNS-susceptible species was positively correlated with pre-hibernation local climate conditions conducive to foraging; however, WNS-susceptible species experienced an overall decline with the presence of WNS and as winter duration increased. Our long-term dataset provides evidence that pre-hibernation climate, specifically favorable summer weather conditions for foraging, greatly influences the reproduction, regardless of WNS status.

Hierarchically embedded scales of movement shape the social networks of vampire bats.

Social structure can emerge from hierarchically embedded scales of movement, where movement at one scale is constrained within a larger scale (e.g. among branches, trees, forests). In most studies of animal social networks, some scales of movement are not observed, and the relative importance of the observed scales of movement is unclear. Here, we asked: how does individual variation in movement, at multiple nested spatial scales, influence each individual's social connectedness? Using existing data from common vampire bats (Desmodus rotundus), we created an agent-based model of how three nested scales of movement-among roosts, clusters and grooming partners-each influence a bat's grooming network centrality. In each of 10 simulations, virtual bats lacking social and spatial preferences moved at each scale at empirically derived rates that were either fixed or individually variable and either independent or correlated across scales. We found that numbers of partners groomed per bat were driven more by within-roost movements than by roost switching, highlighting that co-roosting networks do not fully capture bat social structure. Simulations revealed how individual variation in movement at nested spatial scales can cause false discovery and misidentification of preferred social relationships. Our model provides several insights into how nonsocial factors shape social networks.

The behavioural costs of overcrowding for gregarious cave-dwelling bats.

Bats are known for their gregarious social behaviour, often congregating in caves and underground habitats, where they play a pivotal role in providing various ecosystem services. Studying bat behaviour remains an underexplored aspect of bat ecology and conservation despite its ecological importance. We explored the costs and impacts of overcrowding on bat social behaviour. This study examined variations in bat behavioural patterns between two distinct groups, aggregated and non-aggregated male Rousettus amplexicaudatus, within the Monfort Bat Cave Sanctuary on Mindanao Island, Philippines. We found significant variations in the incident frequencies of various bat behavioural activities, particularly aggression and movement, between these two groups. The increase in aggregation was closely related to negative social behaviour among bats. In contrast, sexual behaviour was significantly related to the positive behaviour of individual bats and was headed in less crowded areas. The disparities in bat behaviour with an apparent decline in bat social behaviour because of overcrowding, with more aggressive behaviours emerging, align with the 'behavioural sink' hypothesis. Our study underscores the importance of considering habitat quality and resource availability in the management and conservation of bat colonies, as these factors can reduce the occurrence of aggressive and negative social behaviours in colonies with high population density by providing alternative habitats.

Physiological and behavioural adaptations by big brown bats hibernating in dry rock crevices.

Winter energy stores are finite and factors influencing patterns of activity are important for overwintering energetics and survival. Hibernation patterns (e.g., torpor bout duration and arousal frequency) often depend on microclimate, with more stable hibernacula associated with greater energy savings than less stable hibernacula. We monitored hibernation patterns of individual big brown bats (Eptesicus fuscus; Palisot de Beauvois, 1796) overwintering in rock-crevices that are smaller, drier, and less thermally stable than most known cave hibernacula. While such conditions would be predicted to increase arousal frequency in many hibernators, we did not find support for this. We found that bats were insensitive to changes in hibernacula microclimate (temperature and humidity) while torpid. We also found that the probability of arousal from torpor remained under circadian influence, likely because throughout the winter during arousals, bats commonly exit their hibernacula. We calculated that individuals spend most of their energy on maintaining a torpid body temperature a few degrees above the range of ambient temperatures during steady-state torpor, rather than during arousals as is typical of other small mammalian hibernators. Flight appears to be an important winter activity that may expedite the benefits of euthermic periods and allow for short, physiologically effective arousals. Overall, we found that big brown bats in rock crevices exhibit different hibernation patterns than conspecifics hibernating in buildings and caves.

Palaeoatmosphere facilitates a gliding transition to powered flight in the Eocene bat, Onychonycteris finneyi.

The evolutionary transition to powered flight remains controversial in bats, the only flying mammals. We applied aerodynamic modeling to reconstruct flight in the oldest complete fossil bat, the archaic Onychonycteris finneyi from the early Eocene of North America. Results indicate that Onychonycteris was capable of both gliding and powered flight either in a standard normodense aerial medium or in the hyperdense atmosphere that we estimate for the Eocene from two independent palaeogeochemical proxies. Aerodynamic continuity across a morphological gradient is further demonstrated by modeled intermediate forms with increasing aspect ratio (AR) produced by digital elongation based on chiropteran developmental data. Here a gliding performance gradient emerged of decreasing sink rate with increasing AR that eventually allowed applying available muscle power to achieve level flight using flapping, which is greatly facilitated in hyperdense air. This gradient strongly supports a gliding (trees-down) transition to powered flight in bats.

Spatial clusters, temporal behavior, and risk factors analysis of rabies in livestock in Ecuador.

Rabies, a globally distributed and highly lethal zoonotic neglected tropical disease, has a significant impact in South America. In Ecuador, animal rabies cases are primarily linked to livestock, and hematophagous bats play a crucial role in disease transmission. This study aims to identify temporal trends, spatial patterns, and risk factors for animal rabies in Ecuador between 2014 and 2019. Epidemiological survey reports from the official Animal Rabies Surveillance Program of the Phyto and Zoosanitary Regulation and Control Agency of Ecuador (AGROCALIDAD) were used. The Animal Rabies Surveillance Program from AGROCALIDAD consists of an official passive surveillance program that receives reports from farmers or individuals (both trained or untrained) who have observed animals with neurological clinical signs and lesions compatible with bat bites, or who have seen or captured bats on their farms or houses. Once this report is made, AGROCALIDAD personnel is sent for field inspection, having to confirm the suspicion of rabies based on farm conditions and compatibility of signs. AGROCALIDAD personnel collect samples from all suspicious animals, which are further processed and analyzed using the Direct Fluorescent Antibody (DFA) test for rabies confirmatory diagnosis. In this case, study data comprised 846 bovine farms (with intra-farm sample sizes ranging from 1 to 16 samples) located in different ecoregions of Ecuador; out of these, 397 (46.93%) farms tested positive for animal rabies, revealing six statistically significant spatial clusters. Among these clusters, three high-risk areas were identified in the southeast of Ecuador. Seasonality was confirmed by the Ljung-Box test for both the number of cases (p < 0.001) and the positivity rate (p < 0.001). The Pacific Coastal lowlands and Sierra regions showed a lower risk of positivity compared to Amazonia (OR = 0.529; 95% CI = 0.318 - 0.883; p = 0.015 and OR = 0.633; 95% CI = 0.410 - 0.977; p = 0.039, respectively). The breeding of non-bovine animal species demonstrated a lower risk of positivity to animal rabies when compared to bovine (OR = 0.145; 95% CI = 0.062 - 0.339; p < 0.001). Similarly, older animals exhibited a lower risk (OR = 0.974; 95% CI = 0.967 - 0.981; p < 0.001). Rainfall during the rainy season was also found to decrease the risk of positivity to animal rabies (OR = 0.996; 95% CI = 0.995 - 0.998; p < 0.001). This study underscores the significance of strengthening the national surveillance program for the prevention and control of animal rabies in Ecuador and other countries facing similar epidemiological, social, and geographical circumstances.

Resting posture drives the evolution of agonistic displays in bats.

Agonistic displays are one of the most diverse social behaviors that have important functions in animal's life history. However, their origin and driving factors have largely been unexplored. Here, we evaluated agonistic displays of 71 bat species across 10 families and classified these displays into two categories: (a) boxing displays where a bat attacks its opponent with its wrist and thumb and (b) pushing displays where a bat uses its head or body to hit a rival. We estimated the strength of the phylogenetic signal of the agonistic displays, revealed their origin, and tested the potential evolutionary relationships between agonistic behaviors and body size or resting posture (free hanging vs. contact hanging where the bat is in contact with some surface). We found that agonistic displays were phylogenetically conserved and that boxing displays are the ancestral state. Moreover, we found that bats with a free-hanging resting posture were more likely to exhibit boxing displays than pushing displays. In addition, bats with longer forearms do not have a higher propensity for boxing displays. This study expands our limited knowledge of the evolution of agonistic displays and highlights the importance of resting posture as a driving force in the diversity of agonistic displays.

Hibernating female big brown bats (Eptesicus fuscus) adjust huddling and drinking behaviour, but not arousal frequency, in response to low humidity.

Many mammals hibernate during winter, reducing energy expenditure via bouts of torpor. The majority of a hibernator's energy reserves are used to fuel brief, but costly, arousals from torpor. Although arousals likely serve multiple functions, an important one is to restore water stores depleted during torpor. Many hibernating bat species require high humidity, presumably to reduce torpid water loss, but big brown bats (Eptesicus fuscus) appear tolerant of a wide humidity range. We tested the hypothesis that hibernating female E. fuscus use behavioural flexibility during torpor and arousals to maintain water balance and reduce energy expenditure. We predicted: (1) E. fuscus hibernating in dry conditions would exhibit more compact huddles during torpor and drink more frequently than bats in high humidity conditions; and (2) the frequency and duration of torpor bouts and arousals, and thus total loss of body mass would not differ between bats in the two environments. We housed hibernating E. fuscus in temperature- and humidity-controlled incubators at 50% or 98% relative humidity (8°C, 110 days). Bats in the dry environment maintained a more compact huddle during torpor and drank more frequently during arousals. Bats in the two environments had a similar number of arousals, but arousal duration was shorter in the dry environment. However, total loss of body mass over hibernation did not differ between treatments, indicating that the two groups used similar amounts of energy. Our results suggest that behavioural flexibility allows hibernating E. fuscus to maintain water balance and reduce energy costs across a wide range of hibernation humidities.

Survival of hibernating little brown bats that are unaffected by white-nose syndrome: Using thermal cameras to understand arousal behavior.

White-nose syndrome is a fungal disease that has decimated hibernating bats from multiple North American species. In 2014, the invasive fungus arrived at a hibernaculum of little brown bats (Myotis lucifugus) inside the spillway of Tippy Dam, located near Wellston, Michigan, USA, yet surprisingly, this population has not experienced the declines seen elsewhere. Unlike a typical subterranean hibernaculum, light enters the spillway through small ventilation holes. We hypothesized that this light causes the hibernating bats to maintain a circadian rhythm, thereby saving energy via social thermoregulation during synchronous arousals. To test this idea, we used high-resolution thermal cameras to monitor arousals from October 2019 to April 2020. We found that arousals followed a circadian rhythm, peaking after sunset, and that most observed arousals (>68%) occurred within a cluster of bats allowing for social thermoregulation. These findings are consistent with the hypothesis that light-induced synchronized arousals contribute to the unprecedented absence of mass mortality from white-nose syndrome in this large population. Using light to maintain a circadian rhythm in bats should be tested as a potential tool for mitigating mortality from white-nose syndrome. More generally, studying populations that have been largely unaffected by white-nose syndrome may provide insight into mitigation strategies for protecting the remaining populations.

Vocal learning-associated convergent evolution in mammalian proteins and regulatory elements.

Vocal production learning ("vocal learning") is a convergently evolved trait in vertebrates. To identify brain genomic elements associated with mammalian vocal learning, we integrated genomic, anatomical, and neurophysiological data from the Egyptian fruit bat (Rousettus aegyptiacus) with analyses of the genomes of 215 placental mammals. First, we identified a set of proteins evolving more slowly in vocal learners. Then, we discovered a vocal motor cortical region in the Egyptian fruit bat, an emergent vocal learner, and leveraged that knowledge to identify active cis-regulatory elements in the motor cortex of vocal learners. Machine learning methods applied to motor cortex open chromatin revealed 50 enhancers robustly associated with vocal learning whose activity tended to be lower in vocal learners. Our research implicates convergent losses of motor cortex regulatory elements in mammalian vocal learning evolution.

Bat Motion can be Described by Leap Frogging.

We present models of bat motion derived from radio-tracking data collected over 14 nights. The data presents an initial dispersal period and a return to roost period. Although a simple diffusion model fits the initial dispersal motion we show that simple convection cannot provide a description of the bats returning to their roost. By extending our model to include non-autonomous parameters, or a leap frogging form of motion, where bats on the exterior move back first, we find we are able to accurately capture the bat's motion. We discuss ways of distinguishing between the two movement descriptions and, finally, consider how the different motion descriptions would impact a bat's hunting strategy.

Higher and bigger: How riparian bats react to climate change.

The altitudinal distribution of animals and changes in their body size are effective indicators of climate change. Bats are sensitive to climate change due to their dependence on temperature during critical life stages. However, long-term studies documenting responses over extended periods are rare. We present a 24-year investigation of Myotis daubentonii, a riparian bat known for altitudinal sexual segregation, along a river course in Central Italy. While males occupy the entire river course, females are confined to downstream warmer areas supporting successful reproduction due to improved foraging site productivity. In 2000, females were absent above 900 m a.s.l in our study area. We hypothesise that a) this altitude threshold is now higher, due to thermal gradient changes along the river course; and b) thermoregulatory costs for reproductive females have declined, leading to increased energy investment in offspring and subsequent generational growth in bat body size. Confirming our hypotheses, females exhibited a 175-m upward shift in altitude limit. Furthermore, we found a concurrent increase in body size (but not condition). Temperatures increased in the 24 years, likely allowing females to extend their range to higher elevations and favouring an increase in newborn body mass. Riparian vegetation remained unchanged, excluding habitat quality changes as the cause for the observed responses. The rapid female elevation rise might imply future disruption of established social structures, altering intra- and intersexual competition for roosts and food. Given the global decline in insect populations, larger bats might face future difficulties in finding food to sustain their body size, increasing mortality. However, the full impact of such changes on bat fitness remains unexplored and warrants further investigation, including other bat populations. This knowledge is crucial for informing conservation in the face of ongoing climate change and preserving the ecosystem services bats deliver in riparian ecosystems.

Bionic study of distance-azimuth discrimination of multi-scattered point objects in bat bio-sonar.

This paper presents a novel approach to enhance the discrimination capacity of multi-scattered point objects in bat bio-sonar. A broadband interferometer mathematical model is developed, incorporating both distance and azimuth information, to simulate the transmitted and received signals of bats. The Fourier transform is employed to simulate the preprocessing step of bat information for feature extraction. Furthermore, the bat bio-sonar model based on convolutional neural network (BS-CNN) is constructed to compensate for the limitations of conventional machine learning and CNN networks, including three strategies: Mix-up data enhancement, joint feature and hybrid atrous convolution module. The proposed BS-CNN model emulates the perceptual nerves of the bat brain for distance-azimuth discrimination and compares with four conventional classifiers to assess its discrimination efficacy. Experimental results demonstrate that the overall discrimination accuracy of the BS-CNN model is 93.4%, surpassing conventional CNN networks and machine learning methods by at least 5.9%. This improvement validates the efficacy of the BS-CNN bionic model in enhancing the discrimination accuracy in bat bio-sonar and offers valuable references for radar and sonar target classification.

Behavioral responses of cave-roosting bats to artificial light of different spectra and intensities: Implications for lighting management strategy.

Artificial light at night has become an emerging environmental pollutant, posing a serious threat to biodiversity. Cave-roosting animals are vulnerable to light pollution due to long-term adaptation to nocturnal niches, and the problem is especially severe in the context of cave tourism and limestone mining. Mitigating the adverse impacts of artificial light on cave-dwelling animals presents a challenge. This study aimed to assess the relative contributions of spectral parameters and light intensity to the emergence behavior of nine cave-roosting bat species: Rhinolophus macrotis, Rhinolophus pearsonii, Rhinolophus rex, Rhinolophus pusillus, Rhinolophus siamensis, Rhinolophus sinicus, Hipposideros armiger, Myotis davidii, and Miniopterus fuliginosus. We manipulated light spectra and intensities through light-emitting diode (LED) lighting and gel filters at the entrance of bat roost. We monitored nightly passes per species to quantify bat emergence under the dark control and ten lighting conditions (blue, green, yellow, red, and white light at high and low intensities) using ultrasonic recording. Our analyses showed that the number of bat passes tended to be reduced in the presence of white, green, and yellow light, independent of light intensity. In contrast, the number of bat passes showed no pronounced differences under the dark control, blue light, and red light. The number of bat passes was primarily affected by LED light's blue component, red component, peak wavelength, and half-width instead of light intensity. These results demonstrate that spectral parameters of LED light can significantly affect emergence behavior of cave-dwelling bats. Our findings highlight the importance of manipulating light colors to reduce the negative impacts of light pollution on cave-roosting bats as a function of their spectral sensitivity. We recommend the use of gel filters to manage existing artificial lighting systems at the entrance of bat-inhabited caves.

Facing the facts: adaptive trade-offs along body size ranges determine mammalian craniofacial scaling.

The mammalian cranium (skull without lower jaw) is representative of mammalian diversity and is thus of particular interest to mammalian biologists across disciplines. One widely retrieved pattern accompanying mammalian cranial diversification is referred to as 'craniofacial evolutionary allometry' (CREA). This posits that adults of larger species, in a group of closely related mammals, tend to have relatively longer faces and smaller braincases. However, no process has been officially suggested to explain this pattern, there are many apparent exceptions, and its predictions potentially conflict with well-established biomechanical principles. Understanding the mechanisms behind CREA and causes for deviations from the pattern therefore has tremendous potential to explain allometry and diversification of the mammalian cranium. Here, we propose an amended framework to characterise the CREA pattern more clearly, in that 'longer faces' can arise through several kinds of evolutionary change, including elongation of the rostrum, retraction of the jaw muscles, or a more narrow or shallow skull, which all result in a generalised gracilisation of the facial skeleton with increased size. We define a standardised workflow to test for the presence of the pattern, using allometric shape predictions derived from geometric morphometrics analysis, and apply this to 22 mammalian families including marsupials, rabbits, rodents, bats, carnivores, antelopes, and whales. Our results show that increasing facial gracility with size is common, but not necessarily as ubiquitous as previously suggested. To address the mechanistic basis for this variation, we then review cranial adaptations for harder biting. These dictate that a more gracile cranium in larger species must represent a structural sacrifice in the ability to produce or withstand harder bites, relative to size. This leads us to propose that facial gracilisation in larger species is often a product of bite force allometry and phylogenetic niche conservatism, where more closely related species tend to exhibit more similar feeding ecology and biting behaviours and, therefore, absolute (size-independent) bite force requirements. Since larger species can produce the same absolute bite forces as smaller species with less effort, we propose that relaxed bite force demands can permit facial gracility in response to bone optimisation and alternative selection pressures. Thus, mammalian facial scaling represents an adaptive by-product of the shifting importance of selective pressures occurring with increased size. A reverse pattern of facial 'shortening' can accordingly also be found, and is retrieved in several cases here, where larger species incorporate novel feeding behaviours involving greater bite forces. We discuss multiple exceptions to a bite force-mediated influence on facial proportions across mammals which lead us to argue that ecomorphological specialisation of the cranium is likely to be the primary driver of facial scaling patterns, with some developmental constraints as possible secondary factors. A potential for larger species to have a wider range of cranial functions when less constrained by bite force demands might also explain why selection for larger sizes seems to be prevalent in some mammalian clades. The interplay between adaptation and constraint across size ranges thus presents an interesting consideration for a mechanistically grounded investigation of mammalian cranial allometry.