Polar bear energetic and behavioral strategies on land with implications for surviving the ice-free period.

Declining Arctic sea ice is increasing polar bear land use. Polar bears on land are thought to minimize activity to conserve energy. Here, we measure the daily energy expenditure (DEE), diet, behavior, movement, and body composition changes of 20 different polar bears on land over 19-23 days from August to September (2019-2022) in Manitoba, Canada. Polar bears on land exhibited a 5.2-fold range in DEE and 19-fold range in activity, from hibernation-like DEEs to levels approaching active bears on the sea ice, including three individuals that made energetically demanding swims totaling 54-175 km. Bears consumed berries, vegetation, birds, bones, antlers, seal, and beluga. Beyond compensating for elevated DEE, there was little benefit from terrestrial foraging toward prolonging the predicted time to starvation, as 19 of 20 bears lost mass (0.4-1.7 kg•day-1). Although polar bears on land exhibit remarkable behavioral plasticity, our findings reinforce the risk of starvation, particularly in subadults, with forecasted increases in the onshore period.

Cadaveric dissection of connection between accessory hemiazygos vein and left brachiocephalic vein.

PURPOSE: The azygos system of veins has many anatomical variations that can impact mediastinal and vascular interventions. While radiological reports on these are of great clinical value, this study is among the first to present a high-quality cadaveric dissection of a rare anatomical variant to supplement previously published radiologic studies. The azygos venous system consists of the azygos vein (AV), hemiazygos vein (HAV), and the accessory hemiazygos vein (AHAV), which develop from the last portion of the posterior cardinal veins. The normal anatomical configuration includes drainage of the posterior intercostal veins, vertebral vein, esophageal veins, HAV, and AHAV to an unpaired right-side AV at the level of the 8th/9th thoracic vertebra. The reported incidence of AHAV draining directly into the left brachiocephalic vein is 1-2%. METHODS: An adult formalin-fixed 70-year-old female cadaver was dissected as part of a medical gross anatomy elective course. RESULTS: Gross documentation of a direct connection of the HAV to the AHAV with the AHAV draining into the left brachiocephalic vein. CONCLUSION: It is important to note the variations of the azygos system to avoid confusion with a potential pathology such as mediastinal masses. Understanding of the rare variant reported here could be useful in the prevention of iatrogenic bleeding from the misplacement of venous catheters and help facilitate radiological diagnosis in the incidence of venous clot formation.

A body composition model with multiple storage compartments for polar bears (Ursus maritimus).

Climate warming is rapidly altering Arctic ecosystems. Polar bears (Ursus maritimus) need sea ice as a platform from which to hunt seals, but increased sea-ice loss is lengthening periods when bears are without access to primary hunting habitat. During periods of food scarcity, survival depends on the energy that a bear has stored in body reserves, termed storage energy, making this a key metric in predictive models assessing climate change impacts on polar bears. Here, we developed a body composition model for polar bears that estimates storage energy while accounting for changes in storage tissue composition. We used data of dissected polar bears (n = 31) to link routinely collected field measures of total body mass and straight-line body length to the body composition of individual bears, described in terms of structural mass and two storage compartments, adipose and muscle. We then estimated the masses of metabolizable proteins and lipids within these storage compartments, giving total storage energy. We tested this multi-storage model by using it to predict changes in the lipid stores from an independent dataset of wild polar bears (n = 36) that were recaptured 8-200 days later. Using length and mass measurements, our model successfully predicted direct measurements of lipid changes via isotopic dilutions (root mean squared error of 14.5 kg). Separating storage into two compartments, and allowing the molecular composition of storage to vary, provides new avenues for quantifying energy stores of individuals across their life cycle. The multi-storage body composition model thus provides a basis for further exploring energetic costs of physiological processes that contribute to individual survival and reproductive success. Given bioenergetic models are increasingly used as a tool to predict individual fitness and population dynamics, our approach for estimating individual energy stores could be applicable to a wide range of species.

The acute physiological response of polar bears to helicopter capture.

Many wildlife species are live captured, sampled, and released; for polar bears (Ursus maritimus) capture often requires chemical immobilization via helicopter darting. Polar bears reduce their activity for approximately 4 days after capture, likely reflecting stress recovery. To better understand this stress, we quantified polar bear activity (via collar-mounted accelerometers) and body temperature (via loggers in the body core [Tabd] and periphery [Tper]) during 2-6 months of natural behavior, and during helicopter recapture and immobilization. Recapture induced bouts of peak activity higher than those that occurred during natural behavior for 2 of 5 bears, greater peak Tper for 3 of 6 bears, and greater peak Tabd for 1 of 6 bears. High body temperature (>39.0°C) occurred in Tper for 3 of 6 individuals during recapture and 6 of 6 individuals during natural behavior, and in Tabd for 2 of 6 individuals during recapture and 3 of 6 individuals during natural behavior. Measurements of Tabd and Tper correlated with rectal temperatures measured after immobilization, supporting the use of rectal temperatures for monitoring bear response to capture. Using a larger dataset (n = 66 captures), modeling of blood biochemistry revealed that maximum ambient temperature during recapture was associated with a stress leukogram (7-26% decline in percent lymphocytes, 12-21% increase in percent neutrophils) and maximum duration of helicopter operations had a similar but smaller effect. We conclude that polar bear activity and body temperature during helicopter capture are similar to that which occurs during the most intense events of natural behavior; high body temperature, especially in warm capture conditions, is a key concern; additional study of stress leukograms in polar bears is needed; and additional data collection regarding capture operations would be useful.

History of nasopharyngeal study: A 5,000-year odyssey into our evolving understanding of a small but vital region.

The nasopharynx has been understudied relative to neighboring anatomical regions. It is a highly complex, integrated space whose function, development, and evolution remains unclear after nearly 5,000 years of study. Historically, most work on the nasopharynx was done with a focus on adjacent structures. It has most often been mentioned in relation to the middle ear (via the Eustachian tube) in ancient texts and has only later been given a designation as one of three portions of a tripartite pharynx among adult humans. As human dissection became practiced more widely in Renaissance Europe, understanding of the nasopharyngeal boundaries improved. With further advancements in the study of nasopharyngeal development, evolution, and anatomical variation from the 19th century up until the present, this region has been shown to be functionally vital and still complicated to define.

The nasopharynx as a window to half a billion years of evolutionary change to the upper respiratory tract.

The nasopharynx is a region at the nexus of several vital physiological systems, including the nasal cavity, oral cavity, braincase, middle ear, and cervical vertebrae. It has undergone pronounced morphological change over the course of tetrapod, mammalian, and human evolution. However, despite its place in evolutionary history, the nasopharynx has received relatively little attention. This special issue focuses on "the evolution, development, and functional morphology of the nasopharynx and its boundaries." Topics covered here include evolutionary developmental biology (or evo-devo), nasopharyngeal adaptions in bats, the importance of the nasopharynx and adjacent structures over the course of human evolution, normal development, middle ear morphology, clinical importance, and the study of the nasopharynx throughout history. Contributions to this special issue range among reviews and syntheses, descriptive analyses, phylogenetic analysis, traditional morphometrics, three-dimensional geometric morphometrics, and computational fluid dynamics. Here, we discuss the central importance of the nasopharynx as can be seen through vertebrate paleontology and comparative morphology. It is via the composite evolutionary history of the nasopharyngeal boundaries that our origins may be better understood, starting with the derivation of the choanae from the median olfactory pit of jawless fish nearly half a billion years ago to the basicranial flexion and facial reduction that distinguish Homo sapiens from all other living mammals. Indeed, the nasopharynx must be acknowledged for its importance in the processes of encephalization and acquisition of speech that have become the hallmark of our species.

Nasopharyngeal morphology contributes to understanding the "muddle in the middle" of the Pleistocene hominin fossil record.

The late archeologist Glynn Isaac first applied the term "muddle in the middle" to a poorly understood period in the Middle Pleistocene human fossil record. This study uses the nasopharyngeal boundaries as a source of traits that may inform this unclear period of human evolution. The nasopharynx lies at the nexus of several vital physiological systems, yet relatively little is known about its importance in human evolution. We analyzed a geographically diverse contemporary Homo sapiens growth series (n = 180 adults, 237 nonadults), Homo neanderthalensis (La Chapelle aux Saints 1, La Ferrassie 1, Forbes Quarry 1, Monte Circeo 1, and Saccopastore 1), mid-Pleistocene Homo (Atapuerca 5, Kabwe 1, Petralona 1, and Steinheim 1), and two Homo erectus sensu lato (KNM-ER 3733 and Sangiran 17). Methods include traditional (Analysis 1) and 3D geometric morphometric analysis (Analysis 2). H. erectus exhibited tall, narrow nasopharyngeal shape, a robust, ancestral morphology. Kabwe 1 and Petralona 1 plotted among H. sapiens in Analysis 2, exhibiting relatively shorter and vertical cartilaginous Eustachian tubes and vertical medial pterygoid plates. Atapuerca 5 and Steinheim 1 exhibited horizontal vomeral orientation similar to H. neanderthalensis, indicating greater relative soft palate length and anteroposterior nasopharynx expansion. They may exhibit synapomorphies with H. neanderthalensis, supporting the accretionary hypothesis. Species-level differences were found among H. sapiens and H. neanderthalensis, including relatively longer dilator tubae muscles and extreme facial airorhynchy among Neanderthals. Furthermore, H. neanderthalensis were autapomorphic in exhibiting horizontal pterygoid plate orientation similar to human infants, suggesting that they may have had inferiorly low placement of the torus tubarius and Eustachian tube orifice on the lateral nasopharyngeal wall in life. This study supports use of osseous nasopharyngeal boundaries both for morphological characters and understanding evolution of otitis media susceptibility in living humans.

The cranial base and related internal anatomical features in Homo neanderthalensis and Homo sapiens.

The cranial anatomy of Homo neanderthalensis and Homo sapiens is well documented in the paleoanthropological and medical literature. However, there are few high-quality visual guides of their comparative morphology. We give here a detailed description of the anatomy of two important fossil specimens, La Chapelle-aux-Saints 1 and abri Pataud 1, based on high-resolution imaging data with each specimen representing the respective morphologies of H. neanderthalensis and H. sapiens. We describe the comparative morphology of external, endocranial, and internal characteristics of the cranium, with a focus on the petrous and tympanic portions of the temporal bone. This descriptive approach shows differences between our specimens, including in positions of cerebral components relative to cranial structures and patterns of dural sinus drainage. Numerous external and internal differences in the shape of the petrous temporal are also described, including its articulation with the tympanic bone and the orientation of the petrotympanic crest. The presence of a large protuberance between the osseous Eustachian tube orifice and carotid foramen in H. neanderthalensis suggests that the levator veli palatini muscle took origin more laterally than the dilator tubae arm of the tensor veli palatini muscle, a feature shared with H. sapiens. The overall pattern that emerges is one in which two species have undergone large-scale evolutionary changes in a functionally critical region. Such differences necessitate high-quality visualization and consideration of both internal and external morphology.

Physiological consequences of Arctic sea ice loss on large marine carnivores: unique responses by polar bears and narwhals.

Rapid environmental changes in the Arctic are threatening the survival of marine species that rely on the predictable presence of the sea ice. Two Arctic marine mammal specialists, the polar bear (Ursus maritimus) and narwhal (Monodon monoceros), appear especially vulnerable to the speed and capriciousness of sea ice deterioration as a consequence of their unique hunting behaviors and diet, as well as their physiological adaptations for slow-aerobic exercise. These intrinsic characteristics limit the ability of these species to respond to extrinsic threats associated with environmental change and increased industrial activity in a warming Arctic. In assessing how sea ice loss may differentially affect polar bears that hunt on the ice surface and narwhals that hunt at extreme depths below, we found that major ice loss translated into elevated locomotor costs that range from 3- to 4-fold greater than expected for both species. For polar bears this instigates an energy imbalance from the combined effects of reduced caloric intake and increased energy expenditure. For narwhals, high locomotor costs during diving increase the risk of ice entrapment due to the unreliability of breathing holes. These species-specific physiological constraints and extreme reliance on the polar sea ice conspire to make these two marine mammal specialists sentinels of climate change within the Arctic marine ecosystem that may foreshadow rapid changes to the marine ecosystem.

Identification of critical windows in early development of human upper respiratory tract and middle ear disease.

Otitis media (OM) or middle ear disease is a prevalent pediatric condition generally related to early growth of the cartilaginous Eustachian tube (CET). This study used a developmental series of dry crania to reconstruct CET and dilator tubae (DT, the muscle opening the CET) morphology. Timing and directionality of CET and upper respiratory tract (URT) growth were investigated. Traditional and 3D geometric morphometrics (GM) were used to assess bony landmarks on the crania. The series was divided using dental eruption into seven growth stages ranging from before eruption of deciduous dentition (approximately the first 6 postnatal months) to eruption of the first permanent maxillary molar (after approximately 6 years). Bony endpoints of the CET and DT were used to calculate their morphology. GM analysis showed substantial shape differences between newborns, early infants, and all later developmental stages. Univariate measures showed the largest growth change between birth and 6 months. Subsequently, CET morphology changed little in the latter half of year 1, instead maturing gradually until approximately 3 years whereas DT relative length and orientation finish growth by the end of year 1. Incongruence in slower CET growth and faster DT growth could impact CET function between 6 and 12 months and be a contributing factor of OM. Tubal aeration may improve after this time when both CET and DT morphology mature, coinciding with clinically reported drop-off in ear infections.

Identifying reliable indicators of fitness in polar bears.

Animal structural body size and condition are often measured to evaluate individual health, identify responses to environmental change and food availability, and relate food availability to effects on reproduction and survival. A variety of condition metrics have been developed but relationships between these metrics and vital rates are rarely validated. Identifying an optimal approach to estimate the body condition of polar bears is needed to improve monitoring of their response to decline in sea ice habitat. Therefore, we examined relationships between several commonly used condition indices (CI), body mass, and size with female reproductive success and cub survival among polar bears (Ursus maritimus) measured in two subpopulations over three decades. To improve measurement and application of morphometrics and CIs, we also examined whether CIs are independent of age and structural size-an important assumption for monitoring temporal trends-and factors affecting measurement precision and accuracy. Maternal CIs and mass measured the fall prior to denning were related to cub production. Similarly, maternal CIs, mass, and length were related to the mass of cubs or yearlings that accompanied her. However, maternal body mass, but not CIs, measured in the spring was related to cub production and only maternal mass and length were related to the probability of cub survival. These results suggest that CIs may not be better indicators of fitness than body mass in part because CIs remove variation associated with body size that is important in affecting fitness. Further, CIs exhibited variable relationships with age for growing bears and were lower for longer bears despite body length being related to cub survival and female reproductive success. These results are consistent with findings from other species indicating that body mass is a useful metric to link environmental conditions and population dynamics.

The seasonal energetic landscape of an apex marine carnivore, the polar bear.

Divergent movement strategies have enabled wildlife populations to adapt to environmental change. In recent decades, the Southern Beaufort Sea subpopulation of polar bears (Ursus maritimus) has developed a divergent movement strategy in response to diminishing sea ice where the majority of the subpopulation (73-85%) stays on the sea ice in summer and the remaining bears move to land. Although declines in sea ice are generally considered a challenge to energy balance in polar bears residing in some regions of the Arctic, little quantitative data exists concerning the seasonal energy expenditures of this apex marine carnivore. We used GPS satellite collars with tri-axial accelerometers and conductivity sensors to measure the location, behavior, and energy expenditure of five adult female polar bears in the southern Beaufort Sea across seasons of sea ice breakup and minimum extent. Using a Bayesian mixed-effects model, we found that energy expenditure was influenced by month, ocean depth, and habitat type (sea ice or land). Total energy expenditure from May through September ranged from 37.7 to 47.2 mJ/kg for individual bears. Bears that moved to land expended 7% more energy on average from May through September than bears that remained on the receding sea ice. In August, when bears were moving from the sea ice to land or moving north with the receding pack ice, bears that moved to land spent 7% more time swimming and expended 22% more energy. This means the immediate cost of moving to land exceeded the cost of remaining on the receding summer pack ice. These findings suggest a physiological reason why the majority of the Southern Beaufort Sea subpopulation continues to inhabit a diminishing summer ice platform. However, bears that moved to land spent 29% more time in preferred hunting habitats over the continental shelf than bears that remained on the sea ice. Bears on land also had access to subsistence-harvested bowhead whale carcasses. Hence, our findings indicate there may be a greater overall energetic benefit to move to land in this region, which suggests that the use of the diminishing summer sea ice may be functioning as an ecological trap.

Reconstructing the Neanderthal Eustachian Tube: New Insights on Disease Susceptibility, Fitness Cost, and Extinction.

Neanderthals are among the best studied and yet most enigmatic fossil human groups with aspects of their anatomy and functional morphology remaining poorly understood. We present the first anatomical reconstruction of the Neanderthal cartilaginous Eustachian tube (CET), a vital component of the upper respiratory tract and nexus for the middle ear and postnasal airway. The Eustachian (auditory, pharyngotympanic) tube, comprised of a bony and cartilaginous (CET) portion, is integral to normal physiological functions such as middle ear aeration and pressure equilibration. Findings indicate that Neanderthal tubal morphology may have predisposed them to high rates of middle ear disease (otitis media [OM]). In living humans, mechanical CET dysfunction underlies OM in infants and young children, with sequelae including hearing loss, meningitis, and pneumonia. Despite proven linkage of CET malfunction with OM, the role of CET morphology in Neanderthal health and disease remains unstudied. We reconstructed Neanderthal CET morphology, comparing their crania to a modern human growth series. Methods included geometric morphometrics and univariate measures among Procrustes-fitted coordinates. Results showed Neanderthal adults exhibiting primitively tall and narrow nasopharynges with infant-like horizontal CET and choanal orientation. As horizontal CET orientation is associated with increased OM incidence in infants and children until around age six, its appearance in Neanderthal adults strongly indicates persistence of high OM susceptibility at this time. This could have compromised fitness and disease load relative to sympatric modern humans, affecting Neanderthals' ability to compete within their ecological niche, and potentially contributing to their rapid extinction. Anat Rec, 302:2109-2125, 2019. © 2019 American Association for Anatomy.

Estimating the energy expenditure of free-ranging polar bears using tri-axial accelerometers: A validation with doubly labeled water.

Measures of energy expenditure can be used to inform animal conservation and management, but methods for measuring the energy expenditure of free-ranging animals have a variety of limitations. Advancements in biologging technologies have enabled the use of dynamic body acceleration derived from accelerometers as a proxy for energy expenditure. Although dynamic body acceleration has been shown to strongly correlate with oxygen consumption in captive animals, it has been validated in only a few studies on free-ranging animals. Here, we use relationships between oxygen consumption and overall dynamic body acceleration in resting and walking polar bears Ursus maritimus and published values for the costs of swimming in polar bears to estimate the total energy expenditure of 6 free-ranging polar bears that were primarily using the sea ice of the Beaufort Sea. Energetic models based on accelerometry were compared to models of energy expenditure on the same individuals derived from doubly labeled water methods. Accelerometer-based estimates of energy expenditure on average predicted total energy expenditure to be 30% less than estimates derived from doubly labeled water. Nevertheless, accelerometer-based measures of energy expenditure strongly correlated (r 2 = 0.70) with measures derived from doubly labeled water. Our findings highlight the strengths and limitations in dynamic body acceleration as a measure of total energy expenditure while also further supporting its use as a proxy for instantaneous, detailed energy expenditure in free-ranging animals.

The Nasal Complex of a Semiaquatic Artiodactyl, the Moose (Alces alces): Is it a Good Evolutionary Model for the Ancestors of Cetaceans?

Among Cetartiodactyla, cetaceans are the only obligate aquatic dwellers. Given morphological similarities between cetacean relatives such as Indohyus (the best represented Eocene raoellid artiodactyl) with other, later artiodactyls, any crown artiodactyl that engages in aquatic behaviors is of interest as an evolutionary model for the adaptations that accompanied the origins of cetaceans. The American moose (Alces alces) is the only non-cetacean artiodactyl to engage in aquatic foraging and, other than Hippopotamus, is distinctive in its diving behaviors. This study surveyed the soft and hard tissue nasal morphology of Alces alces to assess phylogenetic polarity and the presence of adaptations for diving and feeding in fresh water habitats. A fresh dissection of the facial musculature and nasal cavity was performed on one subadult male individual and osteological analyses were also performed on dry crania. This species was analyzed alongside fossil crania of Cervalces (its presumed ancestor), other cervids (e.g., Odocoileus virginianus, the white tail deer; Dama dama, the fallow deer), a bovid (Bos taurus, domestic cattle), and a carnivoran (Ursus americanus, the American black bear). A fresh dissection of the facial musculature and nasal anatomy of one fallow deer specimen was also performed for comparison with the moose. Results indicate that Alces alces exhibited a primitive configuration of maxillolabial muscles and, like Dama, exhibited a series of subcutaneous fibrous tissues connecting these muscles to skin. Alces and Dama, however, both exhibited autapomorphies in the soft tissue anatomy of the external nares. The former possessed a series of muscles that act to constrict the anterior nares, likely during diving. Extremely large fibrofatty pads that were perforated by muscle tendon supported their alar fold. Internally, a double-scrolled maxilloturbinal occupied nearly the entire volume of the anterior nasal cavity and protruded beyond the rim of the piriform aperture in dry crania. Dama had long, thin muscles taking origin on their nasal conchae and inserting onto the alar fold. Yet, despite these anterior nasal autapomorphies, the ethmoturbinal patterns of all observed cervids and the one bovid all appeared primitive with a posteroinferiorly oriented array of ethmoturbinals in close contact with a relatively straight cribriform plate, a macrosmatic condition. These differed from the curved cribriform plate of Ursus whose posterior nasal anatomy appeared hyper-macrosmatic. Indohyus exhibits no skeletal sign of a fleshy proboscis such as an enlarged piriform aperture or shortened nasal bones. Thus, there is little evidence that the early ancestors of cetaceans engaged in prolonged bouts of diving for aquatic foods but more probably were surface swimmers traveling between terrestrial food sources or fleeing predators. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc. Anat Rec, 302:667-692, 2019. © 2018 Wiley Periodicals, Inc.

Energetic costs of locomotion in bears: is plantigrade locomotion energetically economical?

Ursids are the largest mammals to retain a plantigrade posture. This primitive posture has been proposed to result in reduced locomotor speed and economy relative to digitigrade and unguligrade species, particularly at high speeds. Previous energetics research on polar bears (Ursus maritimus) found locomotor costs were more than double predictions for similarly sized quadrupedal mammals, which could be a result of their plantigrade posture or due to adaptations to their Arctic marine existence. To evaluate whether polar bears are representative of terrestrial ursids or distinctly uneconomical walkers, this study measured the mass-specific metabolism, overall dynamic body acceleration, and gait kinematics of polar bears and grizzly bears (Ursus arctos) trained to rest and walk on a treadmill. At routine walking speeds, we found polar bears and grizzly bears exhibited similar costs of locomotion and gait kinematics, but differing measures of overall dynamic body acceleration. Minimum cost of transport while walking in the two species (2.21 J kg-1 m-1) was comparable to predictions for similarly sized quadrupedal mammals, but these costs doubled (4.42 J kg-1 m-1) at speeds ≥5.4 km h-1 Similar to humans, another large plantigrade mammal, bears appear to exhibit a greater economy while moving at slow speeds.

Spring fasting behavior in a marine apex predator provides an index of ecosystem productivity.

The effects of declining Arctic sea ice on local ecosystem productivity are not well understood but have been shown to vary inter-specifically, spatially, and temporally. Because marine mammals occupy upper trophic levels in Arctic food webs, they may be useful indicators for understanding variation in ecosystem productivity. Polar bears (Ursus maritimus) are apex predators that primarily consume benthic and pelagic-feeding ice-associated seals. As such, their productivity integrates sea ice conditions and the ecosystem supporting them. Declining sea ice availability has been linked to negative population effects for polar bears but does not fully explain observed population changes. We examined relationships between spring foraging success of polar bears and sea ice conditions, prey productivity, and general patterns of ecosystem productivity in the Beaufort and Chukchi Seas (CSs). Fasting status (≥7 days) was estimated using serum urea and creatinine levels of 1,448 samples collected from 1,177 adult and subadult bears across three subpopulations. Fasting increased in the Beaufort Sea between 1983-1999 and 2000-2016 and was related to an index of ringed seal body condition. This change was concurrent with declines in body condition of polar bears and observed changes in the diet, condition and/or reproduction of four other vertebrate consumers within the food chain. In contrast, fasting declined in CS polar bears between periods and was less common than in the two Beaufort Sea subpopulations consistent with studies demonstrating higher primary productivity and maintenance or improved body condition in polar bears, ringed seals, and bearded seals despite recent sea ice loss in this region. Consistency between regional and temporal variation in spring polar bear fasting and food web productivity suggests that polar bears may be a useful indicator species. Furthermore, our results suggest that spatial and temporal ecological variation is important in affecting upper trophic-level productivity in these marine ecosystems.

Cranial Indicators Identified for Peak Incidence of Otitis Media.

Acute otitis media (AOM) is one of the most common pediatric conditions worldwide. Peak age of occurrence for AOM has been identified within the first postnatal year and it remains frequent until approximately six postnatal years. Morphological differences between adults and infants in the cartilaginous Eustachian tube (CET) and associated structures may be responsible for development of this disease yet few have investigated normal growth trajectories. We tested hypotheses on coincidence of skeletal growth changes and known ages of peak AOM occurrence. Growth was divided into five dental eruption stages ranging from edentulous neonates (Stage 1) to adults with erupted third maxillary molars (Stage 5). A total of 32 three-dimensional landmarks were used and Generalized Procrustes Analysis was performed. Next, we performed principal components analysis and calculated univariate measures. It was found that growth change in Stage 1 was the most rapid and comprised the largest amount of overall growth in upper respiratory tract proportions (where time is represented by the natural logarithmic transformation of centroid size). The analysis of univariate measures showed that Stage 1 humans did indeed possess the relatively shortest and most horizontally oriented CET's with the greatest amount of growth change occurring at the transition to Stage 2 (eruption of deciduous dentition at five postnatal months, commencing peak AOM incidence) and ceasing by Stage 3 (approximately six postnatal years). Skeletal indicators appear related to peak ages of AOM incidence and may contribute to understanding of a nearly ubiquitous human disease. Anat Rec, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 300:1721-1740, 2017. © 2017 Wiley Periodicals, Inc.

Habitat degradation affects the summer activity of polar bears.

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species' distributions, few studies have examined associated changes in behavior. Polar bear (Ursus maritimus) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50-75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.

Toward Understanding the Mammalian Zygoma: Insights From Comparative Anatomy, Growth and Development, and Morphometric Analysis.

The zygoma, or jugum, is a cranial element that was present in Mesozoic tetrapods, well before the appearance of mammals. Although as an entity the zygoma is a primitive retention among mammals, it has assumed myriad configurations as this group diversified. As the zygoma is located at the intersection of the visual, respiratory, and masticatory apparatuses, it is potentially of great importance in systematic, phylogenetic, and functional studies focused on this region. For example, the facial component of the zygoma and its contribution to a postorbital bar (POB) appear to be relevant to the systematics of a number of mammalian subclades, and the formation of a bony postorbital septum (POS) that separates the orbit from the infratemporal fossa is unique to, and thus potentially phylogenetically significant for uniting anthropoid primates, while the zygoma itself appears to serve to resist tension and bending forces during mastication. In order to better understand the zygoma in the context of its contributions to the circumorbital region, we documented its morphological expression in specimens representing 10 orders of mammals. Since the presence of a POB and of a POS has long been used to justify uniting extant primates and anthropoid primates as respective clades, and because postorbital closure (POC) is morphologically more complex than a POB, we provide detail necessary to address these claims. Our taxically broad overview also allowed us to provide for the first time definitions of configurations that can be applied to future studies. Using a different, but also taxically broad sample of mammals, and of primates in particular, we performed two geometric morphometric analyses that were geared toward testing long-held interpretations of the functional role of the zygoma, especially with regard to mastication and in the context of orbital frontation (to which the zygoma contributes). Further, overall, zygomatic morphology tends not to scale with allometry, sexual dimorphism, or angle of orbital convergence, but it does contribute to unique patterns of intraspecies variation. Anat Rec, 300:76-151, 2017. © 2016 Wiley Periodicals, Inc.