Unexpected decadal density-dependent shifts in California sea lion size, morphology, and foraging niche.

Many marine mammal populations are recovering after long eras of exploitation.1,2 To what degree density-dependent body size declines in recovering species reflect a general response to increased resource competition is unknown. We examined skull size (as a proxy for body size), skull morphology, and foraging dynamics of the top marine predator, the California sea lion (Zalophus californianus), which have been steadily increasing over the last few decades and have approached or reached their carrying capacity in southern California.3 We show that, contrary to predictions, male California sea lions increased rather than decreased their average body size over a 46-year (1962-2008) recovery period. Larger males had proportionally longer oral cavities and more powerful bite strength, and their foraging niche expanded. Females between 1983 and 2007 maintained stable skull dimensions, but their isotopic niche was broader than contemporary males. Increased male body size is compatible with an intensification of density-dependent sexual selection for larger and more competitive individuals concurrent with an expanding foraging niche. High foraging variability among females would explain their body size stability during decades of population recovery. We demonstrate that body size reduction is not the universal response to population recovery in marine mammals and show that selective ecological dynamics could contribute to protecting populations against the increased density-dependent intraspecific competition. However, prey shifts associated with climate change will likely prevent California sea lions (and other marine mammals) from attaining these ecological dynamics, augmenting their vulnerability to resource competition and diminishing their capacity to overcome it.

Sympatric otariids increase trophic segregation in response to warming ocean conditions in Peruvian Humboldt Current System.

Determining trophic habits of predator communities is essential to measure interspecific interactions and response to environmental fluctuations. South American fur seals, Arctocephalus australis (SAFS) and sea lions Otaria byronia (SASL), coexist along the coasts of Peru. Recently, ocean warming events (2014-2017) that can decrease and impoverish prey biomass have occurred in the Peruvian Humboldt Current System. In this context, our aim was to assess the effect of warming events on long-term inter- and intra-specific niche segregation. We collected whisker from SAFS (55 females and 21 males) and SASL (14 females and 22 males) in Punta San Juan, Peru. We used δ13C and δ15N values serially archived in otariid whiskers to construct a monthly time series for 2005-2019. From the same period we used sea level anomaly records to determine shifts in the predominant oceanographic conditions using a change point analysis. Ellipse areas (SIBER) estimated niche width of species-sex groups and their overlap. We detected a shift in the environmental conditions marking two distinct periods (P1: January 2005-October 2013; P2: November 2013-December 2019). Reduction in δ15N in all groups during P2 suggests impoverished baseline values with bottom-up effects, a shift towards consuming lower trophic level prey, or both. Reduced overlap between all groups in P2 lends support of a more redundant assemblage during the colder P1 to a more trophically segregated assemblage during warmer P2. SASL females show the largest variation in response to the warming scenario (P2), reducing both ellipse area and δ15N mean values. Plasticity to adapt to changing environments and feeding on a more available food source without fishing pressure can be more advantageous for female SASL, albeit temporary trophic bottom-up effects. This helps explain larger population size of SASL in Peru, in contrast to the smaller and declining SAFS population.

Sea lions could use multilateration localization for object tracking as tested with bio-inspired whisker arrays.

Previous behavioural research on live sea lions has shown that they are able to detect the direction of oncoming vortices, even when impacting contralaterally. These experiments showed that the whisker system and the animal's neural processing is seemingly able to detect the Direction of Arrival (DoA) from just one side of the heads vibrissal pads. Therefore, temporal differences between whisker stimulation is a likely method for determining the angle. Herein, a theoretical model is presented based on multilateration, and tested by experimental studies on a 2D array of bio-inspired whiskers with regular spacing, and a 3D array of bio-inspired whiskers on a model head of a sea lion, as used in our previous studies. The results show that arrays of whiskers can in principle work as antennae to determine the DoA. This detection of the DoA is achieved by cross-correlation of triplets of whiskers, and Time Difference Of Arrival based multilateration, a method similar to signal processing in modern communication systems and other source localization applications. The results on the 2D array are conclusive and clearly support the hypothesis, while increased uncertainties were found for the 3D array, which could be explained by structural shortcomings of the experimental model. Possible ways to improve the signal are discussed.

Mother-pup recognition mechanisms in Australia sea lion (Neophoca cinerea) using uni- and multi-modal approaches.

Communication is the process by which one emitter conveys information to one or several receivers to induce a response (behavioral or physiological) by the receiver. Communication plays a major role in various biological functions and may involve signals and cues from different sensory modalities. Traditionally, investigations of animal communication focused on a single sensory modality, yet communication is often multimodal. As these different processes may be quite complex and therefore difficult to disentangle, one approach is to first study each sensorial modality separately. With this refined understanding of individual senses, revealing how they interact becomes possible as the characteristics and properties of each modality can be accounted for, making a multimodal approach feasible. Using this framework, researchers undertook systematic, experimental investigations on mother-pup recognition processes in a colonial pinniped species, the Australian sea lion Neophoca cinerea. The research first assessed the abilities of mothers and pups to identify each other by their voice using playback experiments. Second, they assessed whether visual cues are used by both mothers and pups to distinguish them from conspecifics, and/or whether females discriminate the odor of their filial pup from those from non-filial pups. Finally, to understand if the information transmitted by different sensory modalities is analyzed synergistically or if there is a hierarchy among the sensory modalities, experiments were performed involving different sensory cues simultaneously. These findings are discussed with regards to the active space of each sensory cue, and of the potential enhancements that may arise by assessing information from different modalities.

Steller Sea Lion (Eumetopias jubatus) at Tuleny Island, Russia in 2018: Abundance, Composition, and Entanglement.

The Steller sea lion (SSL, Eumetopias jubatus) inhabits the North Pacific Ocean off both the North American and Asian coasts. The abundance of the species in Asia declined by more than half in the second part of the 20th century. Decline recurred in the second decade of the 21th century after a short period of restoration. In contrast with the total dynamics of SSL in Asia, the reproductive aggregation on Tuleny I. (Sea of Okhotsk) has been growing almost continuously since the beginning of its formation in the late 1980s. Long-term monitoring of SSL at Tuleny I. always covered only summer reproductive seasons. We surveyed Tuleny I. in October 2018, and counted 1058 non-pup sea lions and 396 pups. The majority of tagged animals encountered at the rookery were of local origin. About one-third of the summer non-pup sea lions' number could remain at the rookery until the middle of October, which coincides with the seasonal appearance of sea lions off the coast of Japan. The sex-age structure was characterized by total absence of adult males and reduced proportion of subadult males and juveniles. It contrasts with autumn behavior of SSLs in the rookery of the northern Sea of Okhotsk. We observed that 0.7% of the inspected sea lions were entangled in marine debris. The proportion of entangled animals is lower in comparison with that of the whole population of Tuleny I. due to reduction in the ratios of males and juveniles, which entangle in foreign objects more often than adult females.

Fur seals do, but sea lions don't - cross taxa insights into exhalation during ascent from dives.

Management of gases during diving is not well understood across marine mammal species. Prior to diving, phocid (true) seals generally exhale, a behaviour thought to assist with the prevention of decompression sickness. Otariid seals (fur seals and sea lions) have a greater reliance on their lung oxygen stores, and inhale prior to diving. One otariid, the Antarctic fur seal (Arctocephalus gazella), then exhales during the final 50-85% of the return to the surface, which may prevent another gas management issue: shallow-water blackout. Here, we compare data collected from animal-attached tags (video cameras, hydrophones and conductivity sensors) deployed on a suite of otariid seal species to examine the ubiquity of ascent exhalations for this group. We find evidence for ascent exhalations across four fur seal species, but that such exhalations are absent for three sea lion species. Fur seals and sea lions are no longer genetically separated into distinct subfamilies, but are morphologically distinguished by the thick underfur layer of fur seals. Together with their smaller size and energetic dives, we suggest their air-filled fur might underlie the need to perform these exhalations, although whether to reduce buoyancy and ascent speed, for the avoidance of shallow-water blackout or to prevent other cardiovascular management issues in their diving remains unclear. This article is part of the theme issue 'Measuring physiology in free-living animals (Part I)'.

The northernmost haulout site of South American sea lions and fur seals in the western South Atlantic.

We present estimates of the seasonal and spatial occupation by pinnipeds of the Wildlife Refuge of Ilha dos Lobos (WRIL), based on aerial photographic censuses. Twenty aerial photographic censuses were analysed between July 2010 and November 2018. To assess monthly differences in the numbers of pinnipeds in the WRIL we used a Generalized Linear Mixed Model. Spatial analysis was carried out using Kernel density analysis of the pinnipeds on a grid plotted along the WRIL. Subadult male South American sea lions (Otaria flavescens) were the most abundant pinniped in the WRIL. Potential females of this species were also recorded during half of the census. The maximum number of pinnipeds observed in the WRIL was 304 in September 2018, including an unexpected individual southern elephant seal (Mirounga leonina), and a high number of South American fur seal yearlings (Arctocephalus australis). However, there was no statistically significant difference in counts between months. In all months analysed, pinnipeds were most often found concentrated in the northern portion of the island, with the highest abundances reported in September. This study confirms the importance of the WRIL as a haulout site for pinnipeds in Brazil, recommends that land research and recreational activities occur in months when no pinnipeds are present, and encourages a regulated marine mammal-based tourism during winter and spring months.

Pinnipeds orient and control their whiskers: a study on Pacific walrus, California sea lion and Harbor seal.

Whisker touch is an active sensory system. Previous studies in Pinnipeds have adopted relatively stationary tasks to judge tactile sensitivity, which may not accurately promote natural whisker movements and behaviours. This study developed a novel feeding task, termed fish sweeping to encourage whisker movements. Head and whisker movements were tracked from video footage in Harbor seal (Phoca vitulina), California sea lion (Zalophus californianus) and Pacific walrus (Odobenus rosmarus divergens). All species oriented their head towards the moving fish target and moved their whiskers during the task. Some species also engaged in whisker control behaviours, including head-turning asymmetry in the Pacific walrus, and contact-induced asymmetry in the Pacific walrus and California sea lion: behaviours that have only previously been observed in terrestrial mammals. This study confirms that Pinnipeds should be thought of as whisker specialists, and that whisker control (movement and positioning) is an important aspect of touch sensing in these animals, especially in sea lions and walruses. That the California sea lion controls whisker movement in relation to an object, and also had large values of whisker amplitude, spread and asymmetry, suggests that California sea lions are a promising model with which to further explore active touch sensing.

Stroke effort and relative lung volume influence heart rate in diving sea lions.

The dive response, bradycardia (decreased heart rate) and peripheral vasoconstriction, is the key mechanism allowing breath-hold divers to perform long-duration dives while actively swimming and hunting prey. This response is variable and modulated by factors such as dive duration, depth, exercise and cognitive control. This study assessed the potential role of exercise and relative lung volume in the regulation of heart rate (fH) during dives of adult female California sea lions instrumented with electrocardiogram (ECG), depth and tri-axial acceleration data loggers. A positive relationship between activity (minimum specific acceleration) and fH throughout dives suggested increased muscle perfusion associated with exercise. However, apart from late ascent, fH during dives was still less than or equal to resting fH (on land). In addition, the activity-fH relationship was weaker in long, deep dives consistent with prioritization of blood oxygen conservation over blood oxygen delivery to muscle in those dives. Pulmonary stretch receptor reflexes may also contribute to fH regulation as fH profiles generally paralleled changes in relative lung volume, especially in shallower dives and during early descent and late ascent of deeper dives. Overall, these findings support the concept that both exercise and pulmonary stretch receptor reflexes may influence the dive response in sea lions.

Flowfields produced by a robotic sea lion foreflipper starting from rest.

Sea lions swim using primarily their foreflippers, which is uncommon among aquatic mammals. While a significant body of literature exists which investigates the hydrodynamics of body-caudal swimming, relatively little research has looked at sea lion propulsion. In this work, particle imaging velocimetry is used to observe the flow around a robotic model sea lion flipper. The model flipper was cast in silicone from a high-resolution scan of a sample sea lion foreflipper. The model flipper was actuated at the root, and its motion was controlled by a programmable servomotor. It was observed that the thrust-producing clapping motion of the flipper entrained significant fluid momentum on the suction side of the flipper, which developed into a shed vortex and contributed to downstream momentum (and therefore thrust). Rotating the robotic flipper more quickly produced greater downstream jet velocities, but at a lower conversion of rotational velocity, suggesting that this mechanism of propulsion can be optimized based on the system needs.

Spontaneous Yawning and its Potential Functions in South American Sea Lions (Otaria flavescens).

Spontaneous yawning is a widespread behaviour in vertebrates. However, data on marine mammals are scarce. In this study, we tested some hypotheses on the functions of yawning in a captive group of South American sea lions (Otaria flavescens). According to the Dimorphism Hypothesis, species showing low levels of sexual dimorphism in canine size do not show sex differences in yawning distribution; this was supported by our findings, since yawning did not differ between the sexes. Yawning was more frequently performed during resting/sleeping contexts, thus supporting the Drowsiness Hypothesis. Yawning and self-scratching are considered reliable indicators of short-term anxiety in sea lions, since they immediately increased after conflicts both in aggressors and victims (Social Distress Hypothesis supported). In the long-term, yawning was not correlated with individuals' dominance status, thus showing that anxiety is similarly experienced by dominants and subordinates. The last two findings can be explained by the social competition of this species, that involves individuals independently from their sex, age or ranking status. Therefore, the exposure to frequent stressful events can induce similar levels of anxiety in all the subjects (Resource Inequity Hypothesis supported). In conclusion, spontaneous yawning in sea lions seems to share similar functions with other social mammals, suggesting that this behaviour is a possible plesiomorphic trait.

Seal and Sea lion Whiskers Detect Slips of Vortices Similar as Rats Sense Textures.

Pinnipeds like seals and sea lions use their whiskers to hunt their prey in dark and turbid situations. There is currently no theoretical model or hypothesis to explain the interaction between whiskers and hydrodynamic fish trails. The current study, however, provides a theoretical and experimental insight into the mechanism behind the detection of the Strouhal frequency from a Von-Karman vortex street, similar to that of the inverted hydrodynamic fish trail. Herein the flow around a 3D printed sea lion head, with integrated whiskers of comparable geometry and material properties to a real seal lion, is investigated when exposed to vortex streets generated by cylindrical bluff bodies. The whiskers respond to the vortices with a jerky motion, analogous to the stick-slip response of rat whiskers; this motion is found to be the time derivative of the Gaussian function. Compared to the displacement response, the time-derivative of the whisker response decodes the Strouhal frequency of the Von-Karman wake, which improves the sensing efficiency in noisy environments. The study hypothesizes that the time derivative of the whisker bending moment is the best physical variable that can be used as the input to the pinnipeds neural system.

Innervation patterns of mystacial vibrissae support active touch behaviors in California sea lions (Zalophus californianus).

Vibrissae or follicle-sinus complexes (F-SCs) are highly developed mammalian sensory structures. These blood-filled sinuses are richly innervated and possess novel mechanoreceptors. Although much is known regarding the function of F-SCs in terrestrial mammals, much less is known regarding marine carnivores such as pinnipeds. Pinnipeds possess the largest, most highly innervated vibrissae of any mammal. One such pinniped is the California sea lion, which are generalist marine predators that rely heavily upon tactile discrimination capabilities. Psychophysical studies demonstrate that haptic tactile discrimination using F-SCs is exceptionally sensitive. However, our knowledge of the structure and function of F-SCs in otariids is limited. Our objectives were to investigate the innervation and microstructure of F-SCs across the mystacial vibrissal field and infer function from haptic performance studies in California sea lions. Innervation and microstructure of vibrissae differed considerably compared to similar data available for phocids. Total innervation of mystacial vibrissae was estimated to be 86,042 axons. Investigations of innervation density and investment of microvibrissae versus macrovibrissae demonstrated a significantly increased axon density per F-SC in medial microvibrissal regions compared to lateral macrovibrissae, which supports psychophysical data and somatotopic organization of the central nervous system involved with tactile discrimination capability. Innervation increased from medial microvibrissae (705 ± 125 axons/F-SC) to lateral macrovibrissae (1,447 ± 154) as well as from dorsal (541 ± 60) to ventral (1,493 ± 327) vibrissal regions. These data provide a more complete picture of the sensory ecology of this important aquatic mammalian lineage; the specialization of peripheral sensory structures, central nervous structures with demonstrated enhanced haptic capabilities behaviorally has likely led to the ecological success of California sea lions.

Do marine reserves increase prey for California sea lions and Pacific harbor seals?

Community marine reserves are geographical areas closed to fishing activities, implemented and enforced by the same fishermen that fish around them. Their main objective is to recover commercial stocks of fish and invertebrates. While marine reserves have proven successful in many parts of the world, their success near important marine predator colonies, such as the California sea lion (Zalophus californianus) and the Pacific harbor seal (Phoca vitulina richardii), is yet to be analyzed. In response to the concerns expressed by local fishermen about the impact of the presence of pinnipeds on their communities' marine reserves, we conducted underwater surveys around four islands in the Pacific west of the Baja California Peninsula: two without reserves (Todos Santos and San Roque); one with a recently established reserve (San Jeronimo); and, a fourth with reserves established eight years ago (Natividad). All these islands are subject to similar rates of exploitation by fishing cooperatives with exclusive rights. We estimated fish biomass and biodiversity in the seas around the islands, applying filters for potential California sea lion and harbor seal prey using known species from the literature. Generalized linear mixed models revealed that the age of the reserve has a significant positive effect on fish biomass, while the site (inside or outside of the reserve) did not, with a similar result found for the biomass of the prey of the California sea lion. Fish biodiversity was also higher around Natividad Island, while invertebrate biodiversity was higher around San Roque. These findings indicate that marine reserves increase overall fish diversity and biomass, despite the presence of top predators, even increasing the numbers of their potential prey. Community marine reserves may help to improve the resilience of marine mammals to climate-driven phenomena and maintain a healthy marine ecosystem for the benefit of both pinnipeds and fishermen.

Measurement of Intraocular Pressure Using Rebound Tonometry in Anesthetized Free-ranging South American Sea Lions (Otaria byronia).

Intraocular pressures (IOPs) were measured using a rebound tonometer in 56 free-ranging adult South American sea lions (Otaria byronia) from Punta San Juan, Peru. All animals were anesthetized using medetomidine, midazolam, and butorphanol and determined to be in good health. No ocular abnormalities were observed affecting the cornea or ocular adnexa. Field conditions precluded evaluation of the lens and posterior segment of the eye. Mean (SD) IOP values for males (n=37) were 31±11 mmHg (right eye) and 31±9 mmHg (left eye). The same values for females (n=19) were 24±9 mmHg in the left eye and 27±10 mmHg in the right eye. These values are similar to those previously reported for pinnipeds under professional care and higher than those generally reported for terrestrial mammals.

Vibrissa growth rate in California sea lions based on environmental and isotopic oscillations.

Pinniped vibrissae provide information on changes in diet at seasonal and annual scales; however, species-specific growth patterns must first be determined in order to interpret these data. In this study, a simple linear model was used to estimate the growth rate of vibrissae from adult female California sea lions (Zalophus californianus) from San Esteban Island in the Gulf of California, Mexico. The δ15N and δ13C values do not display a marked oscillatory pattern that would permit direct determination of the time period contained in each vibrissa; thus, time (age) was calculated in two ways: 1) based on the correlation between the observed number of peaks (Fourier series) in the δ15N profile and the length of each vibrissa, and 2) through direct comparison with the observed number of peaks in the δ15N profile. Cross-correlation confirmed that the two peaks in the δ15N profile reflected the two peaks in the chlorophyll-a concentration recorded annually around the island. The mean growth rate obtained from the correlation was 0.08 ± 0.01 mm d-1, while that calculated based on the observed number of peaks was 0.10 ± 0.05 mm d-1. Both are consistent with the rates reported for adult females of other otariid species (0.07 to 0.11 mm d-1). Vibrissa growth rates vary by individual, age, sex, and species; moreover, small differences in the growth rate can result in significant differences over the time periods represented by the isotopic signal. Thus, it is important to assess this parameter on a species-by-species basis.

Mouth gape determines the response of marine top predators to long-term fishery-induced changes in food web structure.

Here, we analyse changes throughout time in the isotopic niche of the Franciscana dolphin (Pontoporia blainvillei), the South American fur seal (Arctocephalus australis) and the South American sea lion (Otaria flavescens) from the Río de la Plata estuary and adjacent Atlantic Ocean to test the hypothesis that fishing may modify the diet of small-gape predators by reducing the average size of prey. The overall evidence, from stable isotope and stomach contents analyses, reveals major changes in resource partitioning between the three predators considered, mainly because of an increased access of Franciscana dolphins to juvenile demersal fishes. These results are consistent with the changes in the length distribution of demersal fish species resulting from fishing and suggest that Franciscana dolphin has been the most benefited species of the three marine mammal species considered because of its intermediate mouth gape. In conclusion, the impact of fishing on marine mammals goes beyond the simple reduction in prey biomass and is highly dependent on the mouth gape of the species involved.

Characterizing estrus by trans-abdominal ultrasounds, fecal estrone-3-glucuronide, and vaginal cytology in the Steller sea lion (Eumetopias jubatus).

The ability to monitor the estrus cycle in wild and captive marine species is important for identifying reproductive failures, ensuring a successful breeding program, and monitoring animal welfare. Minimally invasive sampling methods to monitor estrus in captive populations have been developed, but results suggest these tools can be species-specific in their precision and accuracy. Therefore, the minimally invasive sampling methods of trans-abdominal ultrasounds, a fecal steroid analysis (estrone-3-glucuronide, E1G), and vaginal cytology, were evaluated for their efficacy to characterize and monitor estrus in a captive breeding population of Steller sea lions (Eumetopias jubatus). Three adult females were sampled over five breeding seasons, resulting in six estrus profiles characterized by trans-abdominal ultrasounds, five by fecal E1G, and four by vaginal cytology. Animals were trained to allow trans-abdominal ultrasounds, fecal samples, and vaginal swabs to be collected approximately daily. Of the 76 trans-abdominal ultrasound sessions attempted, 8 successfully visualized both ovaries. From these scans, the chronology of ovarian changes during proestrus and estrus was estimated. The time from the detection of developing follicles to the identification of a dominate follicle occurred in 2-5 days and a corpus hemorrhagicum formed approximately 4 days later. However, because visualization of the ovaries was prevented by the gastrointestinal system in 88% of scans, this tool was overall unreliable for monitoring changes associated with estrus. To detect fine scale physiological changes associated with estrus, we analyzed changes in fecal E1G (n = 62) and vaginal cytology (n = 157) 15 days before and after each female's single copulation event (Day = 0). Changes in fecal E1G had the highest accuracy at detecting Day = 0. Fecal E1G increased leading up to estrus, peaked at Day = 0, and then declined. Although we did observe the characteristic increase in superficial cells associated with impending estrus, the type of cell which peaked closest to Day = 0 was intermediate. The uncertainty around the peak in intermediate cells, indicating estrus, was greater than the uncertainty associated with detecting estrus from fecal E1G. Collectively, these results suggest that changes in fecal E1G and vaginal cytology are viable tools to detect estrus in Steller sea lions, but require daily sampling to detect gradual changes, limiting their applicability to studies of wild populations.

Visual cues do not enhance sea lion pups' response to multimodal maternal cues.

Mammals use multiple sensory cues for mother-offspring recognition. While the role of single sensory cues has been well studied, we lack information about how multiple cues produced by mothers are integrated by their offspring. Knowing that Australian sea lion (Neophoca cinerea) pups recognise their mother's calls, we first tested whether visual cues are used by pups to discriminate between conspecifics of different age classes (adult female vs pup). We then examined if adding a visual stimulus to an acoustic cue enhances vocal responsiveness of Australian sea lion pups, by presenting wild individuals with either a visual cue (female 3D-model), an acoustic cue (mother's call), or both simultaneously, and observing their reaction. We showed that visual cues can be used by pups to distinguish adult females from other individuals, however we found no enhancement effect of these cues on the response in a multimodal scenario. Audio-only cues prompted a similar reaction to audio-visual cues that was significantly stronger than pup response to visual-only cues. Our results suggest that visual cues are dominated by acoustic cues and that pups rely on the latter in mother recognition.

A cerebellar substrate for cognition evolved multiple times independently in mammals.

Given that complex behavior evolved multiple times independently in different lineages, a crucial question is whether these independent evolutionary events coincided with modifications to common neural systems. To test this question in mammals, we investigate the lateral cerebellum, a neurobiological system that is novel to mammals, and is associated with higher cognitive functions. We map the evolutionary diversification of the mammalian cerebellum and find that relative volumetric changes of the lateral cerebellar hemispheres (independent of cerebellar size) are correlated with measures of domain-general cognition in primates, and are characterized by a combination of parallel and convergent shifts towards similar levels of expansion in distantly related mammalian lineages. Results suggest that multiple independent evolutionary occurrences of increased behavioral complexity in mammals may at least partly be explained by selection on a common neural system, the cerebellum, which may have been subject to multiple independent neurodevelopmental remodeling events during mammalian evolution.