Biomechanics of biting in loggerhead shrikes: jaw-closing force, velocity and an argument for power.

Differences in the physical and behavioral attributes of prey are likely to impose disparate demands of force and speed on the jaws of a predator. Because of biomechanical trade-offs between force and speed, this presents an interesting conundrum for predators of diverse prey types. Loggerhead shrikes (Lanius ludovicianus) are medium-sized (∼50 g) passeriform birds that dispatch and feed on a variety of arthropod and vertebrate prey, primarily using their beaks. We used high-speed video of shrikes biting a force transducer in lateral view to obtain corresponding measurements of bite force, upper and lower bill linear and angular displacements, and velocities. Our results show that upper bill depression (about the craniofacial hinge) is more highly correlated with bite force, whereas lower bill elevation is more highly correlated with jaw-closing velocity. These results suggest that the upper and lower jaws might play different roles for generating force and speed (respectively) in these and perhaps other birds as well. We hypothesize that a division of labor between the jaws may allow shrikes to capitalize on elements of force and speed without compromising performance. As expected on theoretical grounds, bite force trades-off against jaw-closing velocity during the act of biting, although peak bite force and jaw-closing velocity across individual shrikes show no clear signs of a force-velocity trade-off. As a result, shrikes appear to bite with jaw-closing velocities and forces that maximize biting power, which may be selectively advantageous for predators of diverse prey that require both jaw-closing force and speed.

Spontaneous tempo production in cockatiels (Nymphicus hollandicus) and jungle crows (Corvus macrorhynchos).

Musical and rhythmical abilities are poorly documented in non-human animals. Most of the existing studies focused on synchronisation performances to external rhythms. In humans, studies demonstrated that rhythmical processing (e. g. rhythm discrimination or synchronisation to external rhythm) is dependent of an individual measure: the individual tempo. It is assessed by asking participants to produce an endogenous isochronous rhythm (known as spontaneous motor tempo) without any specific instructions nor temporal cue. In non-human animal literature, studies describing spontaneous and endogenous production of motor tempo without any temporal clue are rare. This exploratory study aims to describe and compare the spontaneous motor tempo of cockatiels and jungle crows. Data were collected on spontaneous beak drumming behaviours of birds housed in laboratory. Inter beak strokes intervals were calculated from sound tracks of videos. The analyses revealed that inter beak strokes intervals are non-randomly distributed intervals and are isochronous. Recorded spontaneous motor tempos are significantly different among some cockatiels. Since we could only conduct statistical analysis with one corvid, we cannot conclude about this species. Our results suggest that cockatiels and jungle crows have individual tempos, thus encouraging further investigations.

Birds are better at regulating heat loss through their legs than their bills: implications for body shape evolution in response to climate.

Endotherms use their appendages-such as legs, tails, ears and bills-for thermoregulation by controlling blood flow to near-surface blood vessels, conserving heat when it is cold, and dissipating heat in hot conditions. Larger appendages allow greater heat dissipation, and appendage sizes vary latitudinally according to Allen's rule. However, little is known about the relative importance of different appendages for thermoregulation. We investigate physiological control of heat loss via bird bills and legs using infrared thermography of wild birds. Our results demonstrate that birds are less able to regulate heat loss via their bills than their legs. In cold conditions, birds lower their leg surface temperature to below that of their plumage surface, retaining heat at their core. In warm conditions, birds increase their leg surface temperature to above that of their plumage surface, expelling heat. By contrast, bill surface temperature remains approximately 2°C warmer than the plumage surface, indicating consistent heat loss under almost all conditions. Poorer physiological control of heat loss via bird bills likely entails stronger selection for shorter bills in cold climates. This could explain why bird bills show stronger latitudinal size clines than bird legs, with implications for predicting shape-shifting responses to climate change.

Upper beak depression instead of elevation dominates cranial kinesis in woodpeckers.

The value of birds' ability to move the upper beak relative to the braincase has been shown in vital tasks like feeding and singing. In woodpeckers, such cranial kinesis has been thought to hinder pecking as delivering forceful blows calls for a head functioning as a rigid unit. Here, we tested whether cranial kinesis is constrained in woodpeckers by comparing upper beak rotation during their daily activities such as food handling, calling and gaping with those from closely related species that also have a largely insectivorous diet but do not peck at wood. Both woodpeckers and non-woodpecker insectivores displayed upper beak rotations of up to 8 degrees. However, the direction of upper beak rotation differed significantly between the two groups, with woodpeckers displaying primarily depressions and non-woodpeckers displaying elevations. The divergent upper beak rotation of woodpeckers may be caused either by anatomical modifications to the craniofacial hinge that reduce elevation, by the caudal orientation of the mandible depressor muscle forcing beak depressions, or by both. Our results suggest that pecking does not result in plain rigidification at the upper beak's basis of woodpeckers, but it nevertheless significantly influences the way cranial kinesis is manifested.

Comparative morphology and soft tissue histology of the remote-touch bill-tip organ in three ibis species of differing foraging ecology.

Ibises (order: Pelecaniformes, family: Threskiornithidae) are probe-foraging birds that use 'remote-touch' to locate prey items hidden in opaque substrates. This sensory capability allows them to locate their prey using high-frequency vibrations in the substrate in the absence of other sensory cues. Remote-touch is facilitated by a specialised bill-tip organ, comprising high densities of mechanoreceptors (Herbst corpuscles) embedded in numerous foramina in the beak bones. Each foramen and its associated Herbst corpuscles make up a sensory unit, called a 'sensory pit'. These sensory pits are densely clustered in the distal portion of the beak. Previous research has indicated that interspecific differences in the extent of sensory pitting in the beak bones correlate with aquatic habitat use of ibises, and have been suggested to reflect different levels of remote-touch sensitivity. Our study investigates the interspecific differences in the bone and soft tissue histology of the bill-tip organs of three species of southern African ibises from different habitats (mainly terrestrial to mainly aquatic). We analysed the external pitting pattern on the bones, as well as internal structure of the beak using micro-CT scans and soft tissue histological sections of each species' bill-tip organs. The beaks of all three species contain remote-touch bill-tip organs and are described here in detail. Clear interspecific differences are evident between the species' bill-tip organs, both in terms of bone morphology and soft tissue histology. Glossy Ibises, which forage exclusively in wetter substrates, have a greater extent of pitting but lower numbers of Herbst corpuscles in each pit, while species foraging in drier substrates (Hadeda and Sacred Ibises) have more robust beaks, fewer pits and higher densities of Herbst corpuscles. Our data, together with previously published histological descriptions of the bill-tip organs of other remote-touch foraging bird species, indicate that species foraging in drier habitats have more sensitive bill-tip organs (based on their anatomy). The vibrations produced by prey (e.g., burrowing invertebrates) travel poorly in dry substrates compared with wetter ones (i.e., dry soil vs. mud or water), and thus we hypothesise that a more sensitive bill-tip organ may be required to successfully locate prey in dry substrates. Furthermore, our results indicate that the differences in bill-tip organ anatomy between the species reflect complex trade-offs between morphological constraints of beak shape and remote-touch sensitivity requirements, both of which are likely related to each species' foraging behaviour and substrate usage. Our study suggests that structures in the bone of the bill-tip organ could provide valuable osteological correlates for the associated soft tissues, and consequently may provide information on the sensory ecology and habitat usage of the birds in the absence of soft tissues.

Making region-specific integumentary organs in birds: evolution and modifications.

Birds are the most diversified terrestrial vertebrates due to highly diverse integumentary organs that enable robust adaptability to various eco-spaces. Here we show that this complexity is built upon multi-level regional specifications. Across-the-body (macro-) specification includes the evolution of beaks and feathers as new integumentary organs that are formed with regional specificity. Within-an-organ (micro-) specification involves further modifications of organ shapes. We review recent progress in elucidating the molecular mechanisms underlying feather diversification as an example. (1) ß-Keratin gene clusters are regulated by typical enhancers or high order chromatin looping to achieve macro- and micro-level regional specification, respectively. (2) Multi-level symmetry-breaking of feather branches confers new functional forms. (3) Complex color patterns are produced by combinations of macro-patterning and micro-patterning processes. The integration of these findings provides new insights toward the principle of making a robustly adaptive bio-interface.

Thermoregulation and heat exchange in ospreys (Pandion haliaetus).

The osprey (Pandion haliaetus) is a cosmopolitan and long-distant migrant, found at all thermal extremes ranging from polar to tropical climates. Since ospreys may have an unusually flexible thermal physiology due to their migration over, and use of, a wide range of habitats, they represent an interesting study system to explore thermoregulatory adaptations in a raptor. In this study, we investigated the efficiency of heat exchange between body and environment in ospreys using micro-computed tomography (µ-CT), infrared thermography and behavioral observations. µ-CT revealed that the osprey bill has its largest potential for heat exchange at the proximal bill region, where arteries are situated most closely under the surface. However, thermal images of 10 juvenile ospreys showed that the bill contributes to only 0.3% of the bird's total heat exchange. The long legs and protruding claws played a more prominent role as heat dissipation areas with a contribution of 6% and 7%, respectively. Operative thresholds, i.e. the ambient temperature below which heat is lost, were high (>38.5 °C) in these body parts. However, we found no indication of active regulation of heat exchange. Instead we observed multiple behavioral adaptations starting at relatively low ambient temperatures. At 26.3 °C ospreys had a 50% probability of showing panting behavior and above 27.9 °C they additionally spread their wings to enable heat dissipation from the less insulated ventral side. The thermal images revealed that at an ambient temperature of 32.1 °C ospreys had a 50% probability of developing a ≥2 °C and up to 7.5 °C colder stripe on the head, which was likely caused by cutaneous evaporation. Our observations suggest that ospreys more strongly rely on behavioral mechanisms than on active thermal windows to cope with heat stress. This study not only improves our understanding of the role of different body parts in ospreys' total heat exchange with the environment but further provides an insight about additional adaptations of this raptor to cope with heat stress.

A mobility-based classification of closed kinematic chains in biomechanics and implications for motor control.

Closed kinematic chains (CKCs), links connected to form one or more closed loops, are used as simple models of musculoskeletal systems (e.g. the four-bar linkage). Previous applications of CKCs have primarily focused on biomechanical systems with rigid links and permanently closed chains, which results in constant mobility (the total degrees of freedom of a system). However, systems with non-rigid elements (e.g. ligaments and muscles) and that alternate between open and closed chains (e.g. standing on one foot versus two) can also be treated as CKCs with changing mobility. Given that, in general, systems that have fewer degrees of freedom are easier to control, what implications might such dynamic changes in mobility have for motor control? Here, I propose a CKC classification to explain the different ways in which mobility of musculoskeletal systems can change dynamically during behavior. This classification is based on the mobility formula, taking into account the number of loops in the CKC and the nature of the constituent joint mobilities. I apply this mobility-based classification to five biomechanical systems: the human lower limbs, the operculum-lower jaw mechanism of fishes, the upper beak rotation mechanism of birds, antagonistic muscles at the human ankle joint and the human jaw processing a food item. I discuss the implications of this classification, including that mobility itself may be dynamically manipulated to simplify motor control. The principal aim of this Commentary is to provide a framework for quantifying mobility across diverse musculoskeletal systems to evaluate its potentially key role in motor control.

Origin of the avian predentary and evidence of a unique form of cranial kinesis in Cretaceous ornithuromorphs.

The avian predentary is a small skeletal structure located rostral to the paired dentaries found only in Mesozoic ornithuromorphs. The evolution and function of this enigmatic element is unknown. Skeletal tissues forming the predentary and the lower jaws in the basal ornithuromorph Yanornis martini are identified using computed-tomography, scanning electron microscopy, and histology. On the basis of these data, we propose hypotheses for the development, structure, and function of this element. The predentary is composed of trabecular bone. The convex caudal surface articulates with rostromedial concavities on the dentaries. These articular surfaces are covered by cartilage, which on the dentaries is divided into 3 discrete patches: 1 rostral articular cartilage and 2 symphyseal cartilages. The mechanobiology of avian cartilage suggests both compression and kinesis were present at the predentary-dentary joint, therefore suggesting a yet unknown form of avian cranial kinesis. Ontogenetic processes of skeletal formation occurring within extant taxa do not suggest the predentary originates within the dentaries, nor Meckel's cartilage. We hypothesize that the predentary is a biomechanically induced sesamoid that arose within the soft connective tissues located rostral to the dentaries. The mandibular canal hosting the alveolar nerve suggests that the dentary teeth and predentary of Yanornis were proprioceptive. This whole system may have increased foraging efficiency. The Mesozoic avian predentary apparently coevolved with an edentulous portion of the premaxilla, representing a unique kinetic morphotype that combined teeth with a small functional beak and persisted successfully for ∼60 million years.

Huffin' and puffin: seabirds use large bills to dissipate heat from energetically demanding flight.

Endothermic animals regulate body temperature by balancing metabolic heat production and heat exchange with the environment. Heat dissipation is especially important during and immediately after demanding activities such as flapping flight, the most energetically expensive mode of locomotion. As uninsulated appendages, bird bills present a potential avenue for efficient heat dissipation. Puffins possess large bills and are members of the bird family with the highest known flight cost. Here, we used infrared thermography to test whether wild tufted puffins (Fratercula cirrhata) use their bills to dissipate excess heat after energetically expensive flight. Both bill surface temperature and the proportion of total heat exchange occurring at the bill decreased with time since landing, suggesting that bills are used to dissipate excess metabolic heat. We propose that bill size in puffins may be shaped by opposing selective pressures that include dissipating heat after flight and conserving heat in cold air and water temperatures.

The effect of beak tissue sloughing and post-treatment beak shape on the productivity of infrared beak-treated layer pullets and hens.

Infrared beak treatment (IRBT) results in a change in beak shape; however, it is unclear what effect variations in post-treatment beak shape have on young pullets. Additionally, the impact of sloughing of the treated beak tissue is not fully understood. Two experiments were conducted to determine the effects of beak tissue sloughing and post-treatment beak shape on the productivity of infrared beak-treated Lohmann Brown (LB) and Lohmann LSL-Lite (LW) pullets and hens. Birds were treated on day of hatch and IRBT equipment settings were adjusted to create 4 specific beak shapes: shovel (SHV), step (STP), standard (STAN), and an untreated sham control (C). Experiment 1 pullets (n = 160) were housed in cages from 1 to 29 d of age and had access to water through chick founts or 360° nipple drinkers (2 replicate cages per treatment). Data collected included body weight (BW), feed intake (FI), feed efficiency (FE), and water disappearance (WD). Experiment 2 pullets (n = 640) were housed in floor pens from 1 d to 18 wk of age (2 replicate pens per treatment) then conventional cages during the laying period (6 replicate cages per treatment). Data collected included BW, FI, egg production, and egg quality. Data were analyzed using PROC MIXED (SAS® 9.4) and differences were significant when P ≤ 0.05. During early life, the IRBT treatments and sloughing had minor effects on FI, FE, and BW. At 4 wk of age, STAN pullets were lighter than C pullets; however, differences were no longer apparent after this age. Pullets with STP or STAN beak shapes had lower WD than C pullets when allowed access to water via nipple drinkers but this did not result in reduced growth. Throughout the laying period, SHV hens laid more saleable eggs than C hens, with no other effects on production. Overall, variations in beak shape and sloughing of the beak tissue had minimal impacts on the productivity of LW and LB pullets and hens.

The impact of beak tissue sloughing and beak shape variation on the behavior and welfare of infrared beak-treated layer pullets and hens.

This research examined how infrared beak treatment (IRBT), sloughing of the treated beak tissue, and the variations in beak shape that can occur post-IRBT impact the welfare and mortality of Lohmann LSL-Lite (LW) and Lohmann Brown (LB) pullets and hens. Two experiments were conducted and birds for both experiments were treated on the day of hatch. IRBT equipment settings were adjusted to create 4 specific beak shapes: shovel (SHV), step (STP), standard (STAN), and an untreated sham control (C). Experiment 1 pullets (n = 80 per strain) were reared in bioassay cages from 1 to 29 D of age (4 replicates per treatment). Data collected included time and presence of beak sloughing, pecking force, behavioral expression, and mortality. Experiment 2 pullets (n = 320 per strain) were reared in floor pens from 1 D to 18 wk of age (2 replicates per treatment) and then conventional cages from 18 to 60 wk of age (6 replicates per treatment). Data collected for Experiment 2 included behavioral expression, feather cover, comb damage, and mortality. Data were analyzed using PROC MIXED (SAS® 9.4) with Tukey's test to separate means. Differences were significant when P ≤ 0.05. IRBT and sloughing had no effect on pecking force or mortality throughout rearing. The variations in post-IRBT beak shape had minor effects on behavior. During rearing, STAN pullets were more active than C pullets but STP and STAN pullets performed less exploratory pecking. During the laying period, SHV and STP hens preened more than C hens. The IRBT treatments, regardless of beak shape, reduced feather loss, comb damage, and cannibalism-related mortality during the laying period. Overall, the results indicate that LW and LB pullets and hens can cope with the change in beak shape that occurs with IRBT, and that welfare is not negatively impacted if some variation in beak shape occurs.

Repeated evolution of drag reduction at the air-water interface in diving kingfishers.

Piscivorous birds have a unique suite of adaptations to forage under the water. One method aerial birds use to catch fish is the plunge dive, wherein birds dive from a height to overcome drag and buoyancy in the water. The kingfishers are a well-known clade that contains both terrestrially foraging and plunge-diving species, allowing us to test for morphological and performance differences between foraging guilds in an evolutionary context. Diving species have narrower bills in the dorsoventral and sagittal plane and longer bills (size-corrected data, n = 71 species, p < 0.01 for all). Although these differences are confounded by phylogeny (phylogenetically corrected ANOVA for dorsoventral p = 0.26 and length p = 0.14), beak width in the sagittal plane remains statistically different ( p < 0.001). We examined the effects of beak morphology on plunge performance by physically simulating dives with three-dimensional printed models of beaks coupled with an accelerometer, and through computational fluid dynamics (CFD). From physically simulated dives of bill models, diving species have lower peak decelerations, and thus enter the water more quickly, than terrestrial and mixed-foraging species (ANOVA p = 0.002), and this result remains unaffected by phylogeny (phylogenetically corrected ANOVA p = 0.05). CFD analyses confirm these trends in three representative species and indicate that the morphology between the beak and head is a key site for reducing drag in aquatic species.

Lidocaine is a nocebo treatment for trigeminally mediated magnetic orientation in birds.

Even though previously described iron-containing structures in the upper beak of pigeons were almost certainly macrophages, not magnetosensitive neurons, behavioural and neurobiological evidence still supports the involvement of the ophthalmic branch of the trigeminal nerve (V1) in magnetoreception. In previous behavioural studies, inactivation of putative V1-associated magnetoreceptors involved either application of the surface anaesthetic lidocaine to the upper beak or sectioning of V1. Here, we compared the effects of lidocaine treatment, V1 ablations and sham ablations on magnetic field-driven neuronal activation in V1-recipient brain regions in European robins. V1 sectioning led to significantly fewer Egr-1-expressing neurons in the trigeminal brainstem than in the sham-ablated birds, whereas lidocaine treatment had no effect on neuronal activation. Furthermore, Prussian blue staining showed that nearly all iron-containing cells in the subepidermal layer of the upper beak are nucleated and are thus not part of the trigeminal nerve, and iron-containing cells appeared in highly variable numbers at inconsistent locations between individual robins and showed no systematic colocalization with a neuronal marker. Our data suggest that lidocaine treatment has been a nocebo to the birds and a placebo for the experimenters. Currently, the nature and location of any V1-associated magnetosensor remains elusive.

Temperature modulates photoperiodic seasonal responses in the subtropical tree sparrow, Passer montanus.

We studied the effects of temperature on the photoperiodic regulation of seasonal reproduction and related events in the subtropical tree sparrow at Shillong, India. In the first experiment, one group of birds was maintained in an outdoor open aviary receiving natural photoperiod and temperature conditions, while the other group was exposed to natural photoperiod and constant temperature of 17 ± 2 °C in an outdoor closed aviary for 12 months. Although both sexes achieved peak gonadal growth at the same time (May) under the two conditions, gonadal regression and feathers molt were delayed under the temperature controlled condition. In the second experiment, the groups of birds were exposed to three different temperatures (17, 25 and 30 °C) under both long (LD-14L:10D) and short (SD-9L:15D) day lengths for 7 months. Birds showed relatively small but significant gonadal growth, darkening of bill color and feathers molt only at 30 °C under SD. However, they behaved as though they were under natural conditions and exhibited the above responses significantly at all temperatures under LD. There was delayed gonadal regression at the lower temperature (17 °C), while feathers molt delayed with increasing temperature (25, 30 °C) under LD. These results clearly indicate that temperature modulates photoperiodic seasonal responses in the tree sparrow.

Molecular basis of tactile specialization in the duck bill.

Tactile-foraging ducks are specialist birds known for their touch-dependent feeding behavior. They use dabbling, straining, and filtering to find edible matter in murky water, relying on the sense of touch in their bill. Here, we present the molecular characterization of embryonic duck bill, which we show contains a high density of mechanosensory corpuscles innervated by functional rapidly adapting trigeminal afferents. In contrast to chicken, a visually foraging bird, the majority of duck trigeminal neurons are mechanoreceptors that express the Piezo2 ion channel and produce slowly inactivating mechano-current before hatching. Furthermore, duck neurons have a significantly reduced mechano-activation threshold and elevated mechano-current amplitude. Cloning and electrophysiological characterization of duck Piezo2 in a heterologous expression system shows that duck Piezo2 is functionally similar to the mouse ortholog but with prolonged inactivation kinetics, particularly at positive potentials. Knockdown of Piezo2 in duck trigeminal neurons attenuates mechano current with intermediate and slow inactivation kinetics. This suggests that Piezo2 is capable of contributing to a larger range of mechano-activated currents in duck trigeminal ganglia than in mouse trigeminal ganglia. Our results provide insights into the molecular basis of mechanotransduction in a tactile-specialist vertebrate.

Mitochondria-targeted molecules determine the redness of the zebra finch bill.

The evolution and production mechanisms of red carotenoid-based ornaments in animals are poorly understood. Recently, it has been suggested that enzymes transforming yellow carotenoids to red pigments (ketolases) in animal cells may be positioned in the inner mitochondrial membrane (IMM) intimately linked to the electron transport chain. These enzymes may mostly synthesize coenzyme Q10 (coQ10), a key redox-cycler antioxidant molecularly similar to yellow carotenoids. It has been hypothesized that this shared pathway favours the evolution of red traits as sexually selected individual quality indices by revealing a well-adjusted oxidative metabolism. We administered mitochondria-targeted molecules to male zebra finches (Taeniopygia guttata) measuring their bill redness, a trait produced by transforming yellow carotenoids. One molecule included coQ10 (mitoquinone mesylate, MitoQ) and the other one (decyl-triphenylphosphonium; dTPP) has the same structure without the coQ10 aromatic ring. At the highest dose, the bill colour of MitoQ and dTPP birds strongly differed: MitoQ birds' bills were redder and dTPP birds showed paler bills even compared to birds injected with saline only. These results suggest that ketolases are indeed placed at the IMM and that coQ10 antioxidant properties may improve their efficiency. The implications for evolutionary theories of sexual signalling are discussed.

Heterochronic truncation of odontogenesis in theropod dinosaurs provides insight into the macroevolution of avian beaks.

Beaks are innovative structures characterizing numerous tetrapod lineages, including birds, but little is known about how developmental processes influenced the macroevolution of these important structures. Here we provide evidence of ontogenetic vestigialization of alveoli in two lineages of theropod dinosaurs and show that these are transitional phenotypes in the evolution of beaks. One of the smallest known caenagnathid oviraptorosaurs and a small specimen of the Early Cretaceous bird Sapeornis both possess shallow, empty vestiges of dentary alveoli. In both individuals, the system of vestiges connects via foramina with a dorsally closed canal homologous to alveoli. Similar morphologies are present in Limusaurus, a beaked theropod that becomes edentulous during ontogeny; and an analysis of neontological and paleontological evidence shows that ontogenetic reduction of the dentition is a relatively common phenomenon in vertebrate evolution. Based on these lines of evidence, we propose that progressively earlier postnatal and embryonic truncation of odontogenesis corresponds with expansion of rostral keratin associated with the caruncle, and these progenesis and peramorphosis heterochronies combine to drive the evolution of edentulous beaks in nonavian theropods and birds. Following initial apomorphic expansion of rostral keratinized epithelia in perinatal toothed theropods, beaks appear to inhibit odontogenesis as they grow postnatally, resulting in a sequence of common morphologies. This sequence is shifted earlier in development through phylogeny until dentition is absent at hatching, and odontogenesis is inhibited by beak formation in ovo.

Sexual dimorphism in jaw muscles of the Japanese sparrowhawk (Accipiter gularis).

Materials suitable for anatomical research of raptorial birds are rare. Bird-eating raptors show distinct inter-sexual differences in body size and parental roles. The large females catch larger prey and prepare small morsels to feed their young using their hooked beaks. Here, we investigated the architectural properties of different jaw muscles of the Japanese sparrowhawk (Accipiter gularis) and examined whether there is sexual dimorphism in their architectural design. The results showed that musculus depressor mandibulae, the opener of the lower jaw, was characterized by relatively long fascicle length, whereas musculus pterygoideus was characterized by its larger mass and physiological cross-sectional area (PCSA) in both sexes. Females have the potential capacity to produce rapid and strong bites by their significantly longer fascicle length of M. depressor mandibulae and larger mass and PCSA of M. pterygoideus. For body size-matched gender, jaw muscles of males had fibres of relatively longer length than females, enabling greater velocity and excursion. Architectural characteristics of jaw muscles, together with the absolute dimorphism (the fascicle length of M. depressor mandibulae, the muscle mass and PCSA of M. pterygoideus) and relative dimorphism in the muscle mass of M. pterygoideus, reflect dietary difference and asymmetric parental roles between the sexes.

Functional morphology of hummingbird bill tips: their function as tongue wringers.

Nectarivores are animals that have evolved adaptations to efficiently exploit floral nectar as the main source of energy in their diet. It is well known that hummingbirds can extract nectar with impressive speed from flowers. However, despite decades of study on nectar intake rates, the mechanism by which feeding is ultimately achieved - the release of nectar from the tongue so that it can pass into the throat and be ingested - has not been elucidated. By using microCT scanning and macro high-speed videography we scrutinized the morphology and function of hummingbird bill tips, looking for answers about the nectar offloading process. We found near the bill tip, in an area of strong lateral compression of internal mandibular width, that the tomia (cutting edges of the bill) are thinner, partially inrolled, and hold forward-directed serrations. Aligned with these structures, a prominent pronglike structure projects upward and forward from the internal mandibular keel. Distal to this mandibular prong, another smaller maxillary prong protrudes downwards from the keel of the palate. Four shallow basins occur at the base of the mandibular prong on the mandibular floor. Of these, two are small basins located proximally and at the sides of the mandibular prong. A third, slightly larger basin is positioned distally to the first two and directly under the maxillary prong. And the fourth basin, the largest, is found more proximally where the bill becomes thicker, as seen from the side. We documented that this group of structures is integrated into the area of the bill where tongue extrusion occurs, and we hypothesize that they function to enhance the nectar release at each lick. We suggest that this "wringer", operated by bill and tongue movements, helps to move nectar towards the throat.