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Notamacropus rufogriseus (red-necked wallaby) are in the family Macropodidae, which is the second largest family of marsupials after the family Didelphidae. This study was conducted with the aim of providing a detailed description of the origin and distribution of the brachial plexus in N. rufogriseus. Two-year-old male and 3-year-old female red-necked wallabies were used for the study. The brachial plexus was formed by ventral rami of C4, C5, C6, C7, C8, and T1 spinal nerves. It is composed of three trunks that give rise to 12 principal nerves. The cranial trunk is formed by the combination of the rami C4-C7; the middle trunk is formed by the combination of the rami C6 and C7; and the caudal trunk is formed by the combination of the rami C8 and T1. Differences between left and right side of the plexus brachialis were not observed. C6 ventral spinal rami contribute the most to brachial plexus nerve formation, while C4 contributes the least. The formation and distribution of the plexus in N. rufogriseus exhibited more resemblance to the patterns observed in marsupial animals rather than placental mammals. Marsupial mammals demonstrate the involvement of C4 in the development of the brachial plexus. The formation and branching of the brachial plexus sequentially adapt in accordance with changes in their thoracic limb activities and innervation points. Anatomical data from brachial plexus studies optimizes thoracic limb clinical and surgical treatments. This work can provide baseline data for future marsupial brachial plexus studies and fill gaps in the scarce literature., (© 2024 American Association for Anatomy.)

Dental disease is common in captive-managed macropods, including Bennett's wallabies, and is a significant cause of morbidity and mortality. Dental extractions and debridement of diseased tissue is often necessary for those undergoing treatment for severe dental disease. Regional anesthesia of the dentition is considered standard of care for domestic animals undergoing orofacial surgery, however, it is not routinely performed in macropods due to limited information on dental anatomy and block approaches. Regional block descriptions for the infraorbital, maxillary, inferior alveolar, and mental blocks in domestic dogs and cats were evaluated and adapted for use in Bennett's wallabies based on descriptions of their anatomy and examination of 2 skulls. These approaches were then performed on cadaver heads with iohexol and methylene blue dye, and block placement was assessed on computed tomography scans and by gross dissection. All block approaches described in this study resulted in appropriate placement of regional anesthesia of the dentition in Bennett's wallabies. They can thus be used by clinicians to improve the intra and postoperative pain control of patients and provide a high level of veterinary care., Competing Interests: Declaration of Conflicting InterestsThe authors declared no potential conflicts of interest with respect to the research, authorship, and/or publication of this article.

Objective: Tooth growth and wear are commonly used tools for determining the age of mammals. The most speciose order of marsupials, Diprotodontia, is characterised by a pair of procumbent incisors within the lower jaw. This study examines the growth and wear of these incisors to understand their relationship with age and sex., Design: Measurements of mandibular incisor crown and root length were made for two sister species of macropodid (kangaroos and wallabies); Macropus giganteus and Macropus fuliginosus. Histological analysis examined patterns of dentine and cementum deposition within these teeth. Broader generalisability within Diprotodontia was tested using dentally reduced Tarsipes rostratus - a species disparate in body size and incisor function to the studied macropodids., Results: In the macropodid sample it is demonstrated that the hypsodont nature of these incisors makes measurements of their growth (root length) and wear (crown length) accurate indicators of age and sex. Model fitting finds that root growth proceeds according to a logarithmic function across the lifespan, while crown wear follows a pattern of exponential reduction for both macropodid species. Histological results find that secondary dentine deposition and cementum layering are further indicators of age. Incisor measurements are shown to correlate with age in the sample of T. rostratus., Conclusions: The diprotodontian incisor is a useful tool for examining chronological age and sex, both morphologically and microstructurally. This finding has implications for population ecology, palaeontology and marsupial evolution., Competing Interests: Declaration of Competing Interest None., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Using finite element analysis on the astragali of five macropodine kangaroos (extant and extinct hoppers) and three sthenurine kangaroos (extinct proposed bipedal striders) we investigate how the stresses experienced by the ankle in similarly sized kangaroos of different hypothesized/known locomotor strategy compare under different simulation scenarios, intended to represent the moment of midstance at different gaits. These tests showed a clear difference between the performance of sthenurines and macropodines with the former group experiencing lower stress in simulated bipedal strides in all species compared with hopping simulations, supporting the hypothesis that sthenurines may have utilized this gait. The Pleistocene macropodine Protemnodon also performed differently from all other species studied, showing high stresses in all simulations except for bounding. This may support the hypothesis of Protemnodon being a quadrupedal bounder., (© 2024 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

The purpose of this prospective and anatomic study was to describe the ultrasonographic anatomy of the kidneys, urinary bladder, adrenal glands, spleen, liver, gall bladder, and gastrointestinal tract in healthy juvenile eastern grey kangaroos (Macropus giganteus). As ultrasonographic descriptions are lacking in marsupial species, it was also conducted to develop a systematic approach for abdominal ultrasonographic evaluation in the kangaroo and to provide preliminary quantitative and qualitative references. Ten macropod cadavers (eight eastern grey kangaroos and two swamp wallabies (Wallabia bicolor)) were used for initial dissections and preliminary ultrasonographic examinations. Seven eastern grey kangaroos (four females and three males; mean mass 18 kg (±4.5)) were ultrasonographically examined under heavy sedation in lateral recumbency. The gaseous forestomach occupied a large proportion of the entire abdomen ultrasonographically; therefore, the majority of cranial landmarks were based on an intercostal approach comparable to a deep-chested dog. Compared to domestic species, ultrasonographic differences in anatomy include the forestomach, hindstomach, liver orientation, distinguishable adrenal glands, splenic branching, and epipubic bones, all of which were described. The study was limited by the small sample size (7) and weight range (14-25 kg). The systematic approach and description of the normal ultrasonographic anatomy of the abdominal organs in the eastern grey kangaroo should provide a foundation for the ultrasonographic diagnosis and interpretation of abdominal disease in this species., (© 2023 The Authors. Veterinary Radiology & Ultrasound published by Wiley Periodicals LLC on behalf of American College of Veterinary Radiology.)

Tree-kangaroos of the genus Dendrolagus occupy forest habitats of New Guinea and extreme northeastern Australia, but their evolutionary history is poorly known. Descriptions in the 2000s of near-complete Pleistocene skeletons belonging to larger-bodied species in the now-extinct genus Bohra broadened our understanding of morphological variation in the group and have since helped us to identify unassigned fossils in museum collections, as well as to reassign species previously placed in other genera. Here we describe these fossils and analyse tree-kangaroo systematics via comparative osteology. Including B. planei sp. nov., B. bandharr comb. nov. and B. bila comb. nov., we recognise the existence of at least seven late Cenozoic species of Bohra, with a maximum of three in any one assemblage. All tree-kangaroos (Dendrolagina subtribe nov.) exhibit skeletal adaptations reflective of greater joint flexibility and manoeuvrability, particularly in the hindlimb, compared with other macropodids. The Pliocene species of Bohra retained the stepped calcaneocuboid articulation characteristic of ground-dwelling macropodids, but this became smoothed to allow greater hindfoot rotation in the later species of Bohra and in Dendrolagus. Tree-kangaroo diversification may have been tied to the expansion of forest habitats in the early Pliocene. Following the onset of late Pliocene aridity, some tree-kangaroo species took advantage of the consequent spread of more open habitats, becoming among the largest late Cenozoic tree-dwellers on the continent. Arboreal Old World primates and late Quaternary lemurs may be the closest ecological analogues to the species of Bohra.

The extinct sthenurine (giant, short-faced) kangaroos have been proposed to have a different type of locomotor behavior to extant (macropodine) kangaroos, based both on physical limitations (the size of many exceeds the proposed limit for hopping) and anatomical features (features of the hind limb anatomy suggestive of weight-bearing on one leg at a time). Here, we use micro computerised tomography (micro-CT) scans of the pedal bones of six kangaroos, three sthenurine, and three macropodine, ranging from ~50 to 150 kg, to investigate possible differences in bone resistances to bending and cortical bone distribution that might relate to differences in locomotion. Using second moment of area analysis, we show differences in resistance to bending between the two subfamilies. Distribution of cortical bone shows that sthenurines had less resistant calcaneal tubers, implying a different foot posture during locomotion, and the long foot bones were more resistant to the medial bending stresses. These differences were the most pronounced between Pleistocene monodactyl sthenurines (Sthenurus stirlingi and Procoptodon browneorum) and the two species of Macropus (the extant M. giganteus and the extinct M. cf. M. titan) and support the hypothesis that these derived sthenurines employed bipedal striding. The Miocene sthenurine Hadronomas retains some more macropodine-like features of bone resistance to bending, perhaps reflecting its retention of the fifth pedal digit. The Pleistocene macropodine Protemnodon has a number of unique features, possibly indicative of a type of locomotion unlike the other kangaroos., (© 2022 The Authors. Journal of Morphology published by Wiley Periodicals LLC.)

Human ribs are thought to be less affected by mechanical strain at the microscopic level than limb bones, implying that rib remodelling better reflects bone physiological homeostasis. Here, we test the hypothesis that rib tissue will be well vascularized and thus enhance susceptibility to metabolic influence. An intra-skeletal comparison of bone vascular canal density was conducted using a macropod animal model adapted to bipedal habitual hopping. The right humerus, ulna, radius, femur, tibia, fibula, a mid-thoracic and upper-thoracic rib of an eastern grey kangaroo (Macropus giganteus) were sectioned at the midshaft, from which histological sections were prepared. Bone vascularity from a maximum of 12 mm 2  of sub-periosteal parallel-fibred and lamellar bone was recorded, resulting in a total of 2047 counted vessels. Vascular canal density data were corrected by cortical width, maximum length, and midshaft circumference robusticity indices computed for each bone. The fibula consistently had the highest vascular canal density, even when corrected for maximum length, cortical width and midshaft circumference robusticities. This was followed by the mid- and upper-thoracic ribs. Vascularity differences between bones were relatively consistent whether vascular canal density was controlled for by cortical width or midshaft circumference robusticities. Vascular canal density and robusticity indices were also positively and negatively correlated (p < 0.05). Results confirm that the ribs are well vascularized, which facilitates bone metabolic processes such as remodelling, but the fibula also appears to be a well vascularized bone. Future research investigating human bone metabolism will benefit from examining thoracic rib or fibula samples.

The upright posture of the kangaroo suggests that the spine of the kangaroo could be a possible substitute model for biomechanical studies of the human spine. A prerequisite for this should be the agreement of anatomy in humans and kangaroos. The purpose of this study was to investigate the anatomical parameters of the kangaroo spine from C4 to S4 and compare them with existing anatomical data of the human spine. Eight complete spines of the red giant kangaroo were obtained and 21 anatomical parameters were measured from the vertebral bodies, spinal canal, endplate, pedicles, intervertebral discs, transverse, and spinous processes. Most similarities between kangaroo and human spines were found for the vertebral bodies in the cervical and the lumbar spine. The largest differences were evident for the spinous processes. Although both species are somehow upright, these differences may be explained by the way how they move. Jumping probably requires more muscle strength than walking on two legs., (© 2020 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

This study assesses the functional morphology of the ankle extensor muscle-tendon units of the springhare Pedetes capensis, an African bipedal hopping rodent, to test for convergent evolution with the Australian bipedal hopping macropods. We dissect and measure the gastrocnemius, soleus, plantaris, and flexor digitorum longus in 10 adult springhares and compare them against similar-sized macropods using phylogenetically informed scaling analyses. We show that springhares align reasonably well with macropod predictions, being statistically indistinguishable with respect to the ankle extensor mean weighted muscle moment arm (1.63 vs. 1.65 cm, respectively), total muscle mass (41.1 vs. 29.2 g), total muscle physiological cross-sectional area (22.9 vs. 19.3 cm 2 ), mean peak tendon stress (26.2 vs. 35.2 MPa), mean tendon safety factor (4.7 vs. 3.6), and total tendon strain energy return capacity (1.81 vs. 1.82 J). However, total tendon cross-sectional area is significantly larger in springhares than predicted for a similar-sized macropod (0.26 vs. 0.17 cm 2 , respectively), primarily due to a greater plantaris tendon thickness (0.084 vs. 0.048 cm 2 ), and secondarily because the soleus muscle-tendon unit is present in springhares but is vestigial in macropods. The overall similarities between springhares and macropods indicate that evolution has favored comparable lower hindlimb body plans for bipedal hopping locomotion in the two groups of mammals that last shared a common ancestor ~160 million years ago. The springhare's relatively thick plantaris tendon may facilitate rapid transfer of force from muscle to skeleton, enabling fast and accelerative hopping, which could help to outpace and outmaneuver predators., (© 2020 Anatomical Society.)

The Sthenurinae were a diverse subfamily of short-faced kangaroos that arose in the Miocene and diversified during the Pliocene and Pleistocene. Many species possessed skull morphologies that were relatively structurally reinforced with bone, suggesting that they were adapted to incorporate particularly resistant foods into their diets. However, the functional roles of many unique, robust features of the sthenurine cranium are not yet clearly defined. Here, the finite element method is applied to conduct a comprehensive analysis of unilateral biting along the cheek tooth battery of a well-represented sthenurine, Simosthenurus occidentalis. The results are compared with those of an extant species considered to be of most similar ecology and cranial proportions to this species, the koala (Phascolarctos cinereus). The simulations reveal that the cranium of S. occidentalis could produce and withstand comparatively high forces during unilateral biting. Its greatly expanded zygomatic arches potentially housed enlarged zygomaticomandibularis muscles, shown here to reduce the risk of dislocation of the temporomandibular joint during biting with the rear of a broad, extensive cheek tooth row. This may also be a function of the zygomaticomandibularis in the giant panda (Ailuropoda melanoleuca), another species known to exhibit an enlarged zygomatic arch and hypertrophy of this muscle. Furthermore, the expanded frontal plates of the S. occidentalis cranium form broad arches of bone with the braincase and deepened maxillae that each extend from the anterior tooth rows to their opposing jaw joints. These arches are demonstrated here to be a key feature in resisting high torsional forces during unilateral premolar biting on large, resistant food items. This supports the notion that S. occidentalis fed thick, lignified vegetation directly to the cheek teeth in a similar manner to that described for the giant panda when crushing mature bamboo culms., Competing Interests: The author has declared that no competing interests exist.

Combined "total evidence" analysis of molecular and morphological data offers the opportunity to objectively merge fossils into the tree of life, and challenges the primacy of solely DNA based phylogenetic and dating inference, even among modern taxa. To investigate the relative utility of DNA, morphology, and total evidence for evolutionary inference, we sequenced the first near-complete mitochondrial genomes from extinct Australian megafauna: a 40-50 thousand year old giant short-faced kangaroo (Simosthenurus occidentalis) and giant wallaby (Protemnodon anak). We analyzed the ancient DNA and fossil data alongside comparable data from extant species to infer phylogeny, divergence times, and ancestral body mass among macropods (kangaroos and wallabies). Our results confirm a close relationship between Protemnodon and the iconic kangaroo genus complex "Macropus", and unite the giant Simothenurus with the hare-sized Lagostrophus fasciatus (banded hare-wallaby), suggesting that the latter is the closest living link to the once diverse sthenurine kangaroo radiation. We find that large body size evolved multiple times among kangaroos, coincident with expansion of open woodland habitats beginning in the Late Miocene. In addition, our results suggest that morphological data mislead macropod phylogeny reconstruction and in turn can distort total evidence estimation of divergence dates. However, a novel result with potentially broad application is that the accuracy and precision of reconstructing ancestral body mass was improved by tracing body mass on morphological branch lengths. This is likely due to positive allometric correlation between morphological and body size variation-a relationship that may be masked or even misleadingly inverted with the temporal or molecular branch lengths that typically underpin ancestral body size reconstruction. Our study supports complementary roles for DNA and morphology in evolutionary inference, and opens a new window into the evolution of Australia's unique marsupial fauna., (© The Author(s) 2018. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Background: The kangaroo pericardium might be considered to be a good candidate material for use in the manufacture of the leaflets of percutaneous heart valves based upon the unique lifestyle. The diet consists of herbs, forbs and strubs. The kangaroo pericardium holds an undulated structure of collagen., Material and Method: A Red Kangaroo was obtained after a traffic fatality and the pericardium was dissected. Four compasses were cut from four different sites: auricular (AUR), atrial (ATR), sternoperitoneal (SPL) and phrenopericardial (PPL). They were investigated by means of scanning electron microscopy, light microscopy and transmission electron microscopy., Results: All the samples showed dense and wavy collagen bundles without vascularisation from both the epicardium and the parietal pericardium. The AUR and the ATR were 150±25μm thick whereas the SPL and the PPL were thinner at 120±20μm. The surface of the epicardium was smooth and glistening. The filaments of collagen were well individualized without any aggregation, but the banding was poorly defined and somewhat blurry., Conclusion: This detailed morphological analysis of the kangaroo pericardium illustrated a surface resistant to thrombosis and physical characteristics resistant to fatigue. The morphological characteristics of the kangaroo pericardium indicate that it represents an outstanding alternative to the current sources e.g., bovine and porcine. However, procurement of tissues from the wild raises supply and sanitary issues. Health concerns based upon sanitary uncertainty and reliability of supply of wild animals remain real problems., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

The mammalian penis is a complex hydraulic organ of cavernous (spongy) tissue supported by both smooth and skeletal muscle structures. In placental mammals, the paired ‎Musculus ischiocavernosi anchor the corpora cavernosa to the pelvis (at the ischium), and the paired M. bulbospongiosi converge as they envelop the base of the corpus spongiosum. Male marsupials have a dramatically different anatomy, however, in which both sets of paired muscles remain separate, have a bulbous, globular shape and do not have any direct connection to the pelvis. Here we provide the first detailed anatomical investigation of the muscles of the penis in the western grey kangaroo (Macropus fuliginosus) incorporating dissection, histology, vascular casting and computed tomography. The M. ischiocavernosus and M. bulbospongiosus form massive, multipennate bodies of skeletal muscle surrounding the paired roots of the corpus cavernosum and corpus spongiosum, respectively. Bilateral vascular supply is via both the artery of the penis and the ventral perineal artery. Histological examination reveals cavernous tissues with substantial smooth muscle supported by fibroelastic trabeculae, surrounded by the thick collagenous tunica albuginea. The M. ischiocavernosus and M. bulbospongiosus are known to function during erection of the penis and ejaculation via muscular contraction increasing blood pressure within cavernous vascular tissues. The thick muscular anatomy of the kangaroo would be well suited to this function. The absence of any connection to the bony pelvis in marsupials suggests the possibility of different mechanisms of action of these muscles with regard to reduction of venous return, eversion from the cloaca, or movements such as penile flips, which have been described in some placental mammals. This highlights a greater diversity in form and function in the evolution of the mammalian penis than has been previously considered., (© 2019 Anatomical Society.)

Differentiating between ancient and younger, more rapidly evolved clades is important for determining paleoenvironmental drivers of diversification. Australia possesses many aridity-adapted lineages, the origins of which have been closely linked to late Miocene continental aridification. Using dental macrowear and molar crown height measurements, spanning the past 25 million years, we show that the most iconic Australian terrestrial mammals, "true" kangaroos (Macropodini), adaptively radiated in response to mid-Pliocene grassland expansion rather than Miocene aridity. In contrast, low-crowned, short-faced kangaroos radiated into predominantly browsing niches as the late Cenozoic became more arid, contradicting the view that this was an interval of global browser decline. Our results implicate warm-to-cool climatic oscillations as a trigger for adaptive radiation and refute arguments attributing Pleistocene megafaunal extinction to aridity-forced dietary change., (Copyright © 2018 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Increasing body size is accompanied by facial elongation across a number of mammalian taxa. This trend forms the basis of a proposed evolutionary rule, cranial evolutionary allometry (CREA). However, facial length has also been widely associated with the varying mechanical resistance of foods. Here, we combine geometric morphometrics and computational biomechanical analyses to determine whether evolutionary allometry or feeding ecology have been dominant influences on facial elongation across 16 species of kangaroos and relatives (Macropodiformes). We found no support for an allometric trend. Nor was craniofacial morphology strictly defined by dietary categories, but rather associated with a combination of the mechanical properties of vegetation types and cropping behaviours used to access them. Among species examined here, shorter muzzles coincided with known diets of tough, resistant plant tissues, accessed via active slicing by the anterior dentition. This morphology consistently resulted in increased mechanical efficiency and decreased bone deformation during incisor biting. Longer muzzles, by contrast, aligned with softer foods or feeding behaviours invoking cervical musculature that circumvent the need for hard biting. These findings point to a potential for craniofacial morphology to predict feeding ecology in macropodiforms, which may be useful for species management planning and for inferring palaeoecology., (© 2018 The Author(s).)

Traffic collisions between kangaroos and motorists are on the rise on Australian roads. According to a recent report, it was estimated that there were more than 20,000 kangaroo vehicle collisions that occurred only during the year 2015 in Australia. In this work, we are proposing a vehicle-based framework for kangaroo detection in urban and highway traffic environment that could be used for collision warning systems. Our proposed framework is based on region-based convolutional neural networks (RCNN). Given the scarcity of labeled data of kangaroos in traffic environments, we utilized our state-of-the-art data generation pipeline to generate 17,000 synthetic depth images of traffic scenes with kangaroo instances annotated in them. We trained our proposed RCNN-based framework on a subset of the generated synthetic depth images dataset. The proposed framework achieved a higher average precision (AP) score of 92% over all the testing synthetic depth image datasets. We compared our proposed framework against other baseline approaches and we outperformed it with more than 37% in AP score over all the testing datasets. Additionally, we evaluated the generalization performance of the proposed framework on real live data and we achieved a resilient detection accuracy without any further fine-tuning of our proposed RCNN-based framework.

A hallmark of mammalian brain evolution is the emergence of the neocortex, which has expanded in all mammalian infraclasses (Eutheria, Marsupialia, Monotremata). In eutherians, neocortical neurons derive from distinct neural stem and progenitor cells (NPCs). However, precise data on the presence and abundance of the NPCs, especially of basal radial glia (bRG), in the neocortex of marsupials are lacking. This study characterized and quantified the NPCs in the developing neocortex of a marsupial, the tammar wallaby (Macropus eugenii). Our data demonstrate that its neocortex is characterized by high NPC diversity. Importantly, we show that bRG exist at high relative abundance in the tammar indicating that this cell type is not specific to the eutherian neocortex and that similar mechanisms may underlie the formation of an expanded neocortex in eutherian and marsupial mammals. We also show that bRG are likely to have been present in the therian ancestor, so did not emerge independently in the eutherian and marsupial lineages. Moreover, our data support the concept that changes in multiple parameters contribute to neocortex expansion and demonstrate the importance of bRG and other NPCs for the development and expansion of the mammalian neocortex., (© The Author 2016. Published by Oxford University Press. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

Bipedal hopping is used by macropods, including rat-kangaroos, wallabies and kangaroos (superfamily Macropodoidea). Interspecific scaling of the ankle extensor muscle-tendon units in the lower hindlimbs of these hopping bipeds shows that peak tendon stress increases disproportionately with body size. Consequently, large kangaroos store and recover more strain energy in their tendons, making hopping more efficient, but their tendons are at greater risk of rupture. This is the first intraspecific scaling analysis on the functional morphology of the ankle extensor muscle-tendon units (gastrocnemius, plantaris and flexor digitorum longus) in one of the largest extant species of hopping mammal, the western grey kangaroo Macropus fuliginosus (5.8-70.5 kg post-pouch body mass). The effective mechanical advantage of the ankle extensors does not vary with post-pouch body mass, scaling with an exponent not significantly different from 0.0. Therefore, larger kangaroos balance rotational moments around the ankle by generating muscle forces proportional to weight-related gravitational forces. Maximum force is dependent upon the physiological cross-sectional area of the muscle, which we found scales geometrically with a mean exponent of only 0.67, rather than 1.0. Therefore, larger kangaroos are limited in their capacity to oppose large external forces around the ankle, potentially compromising fast or accelerative hopping. The strain energy return capacity of the ankle extensor tendons increases with a mean exponent of ~1.0, which is much shallower than the exponent derived from interspecific analyses of hopping mammals (~1.4-1.9). Tendon safety factor (ratio of rupture stress to estimated peak hopping stress) is lowest in the gastrocnemius (< 2), and it decreases with body mass with an exponent of -0.15, extrapolating to a predicted rupture at 160 kg. Extinct giant kangaroos weighing 250 kg could therefore not have engaged in fast hopping using 'scaled-up' lower hindlimb morphology of extant western grey kangaroos., (© 2017 Anatomical Society.)

The kangaroo knee is, as in other species, a complex diarthrodial joint dependent on interacting osseous, cartilaginous and ligamentous components for its stability. While principal load bearing occurs through the femorotibial articulation, additional lateral articulations involving the fibula and lateral fabella also contribute to the functional arrangement. Several fibrocartilage and ligamentous structures in this joint remain unexplained or have been misunderstood in previous studies. In this study, we review the existing literature on the structure of the kangaroo 'knee' before providing a new description of the gross anatomical and histological structures. In particular, we present strong evidence that the previously described 'femorofibular disc' is best described as a fibular meniscus on the basis of its gross and histological anatomy. Further, we found it to be joined by a distinct tendinous tract connecting one belly of the m. gastrocnemius with the lateral meniscus, via a hyaline cartilage cornu of the enlarged lateral fabella. The complex of ligaments connecting the fibular meniscus to the surrounding connective tissues and muscles appears to provide a strong resistance to external rotation of the tibia, via the restriction of independent movement of the proximal fibula. We suggest this may be an adaptation to resist the rotational torque applied across the joint during bipedal saltatory locomotion in kangaroos., (© 2017 Anatomical Society.)

Purpose: Based on the structural anatomy, loading condition and range of motion (ROM), no quadruped animal has been shown to accurately mimic the structure and biomechanical function of the human spine. The objective of this study is to quantify the thoracic vertebrae geometry of the kangaroo, and compare with adult human, pig, sheep, and deer., Methods: The thoracic vertebrae (T1-T12) from whole body CT scans of ten juvenile kangaroos (ages 11-14 months) were digitally reconstructed and geometric dimensions of the vertebral bodies, endplates, pedicles, spinal canal, processes, facets and intervertebral discs were recorded. Similar data available in the literature on the adult human, pig, sheep, and deer were compared to the kangaroo. A non-parametric trend analysis was performed., Results: Thoracic vertebral dimensions of the juvenile kangaroo were found to be generally smaller than those of the adult human and quadruped animals. The most significant (p < 0.001) correlations (Rho) found between the human and kangaroo were in vertebrae and endplate dimensions (0.951 ≤ Rho ≤ 0.963), pedicles (0.851 ≤ Rho ≤ 0.951), and inter-facet heights (0.891 ≤ Rho ≤ 0.967). The deer displayed the least similar trends across vertebral levels., Conclusions: Similarities in thoracic spine vertebral geometry, particularly of the vertebrae, pedicles and facets may render the kangaroo a more clinically relevant human surrogate for testing spinal implants. The pseudo-biped kangaroo may also be a more suitable model for the human thoracic spine for simulating spine deformities, based on previously published similarities in biomechanical loading, posture and ROM.

In mammals, embryonic diapause, or suspension of embryonic development, occurs when embryos at the blastocyst stage are arrested in growth and metabolism. In the tammar wallaby (Macropus eugenii), there are two separate uteri, only one of which becomes gravid with the single conceptus at a post-partum oestrus, so changes during pregnancy can be compared between the gravid and non-gravid uterus within the same individual. Maintenance of the viable blastocyst and inhibition of further conceptus growth during diapause in the tammar is completely dependent on the uterine environment. Although the specific endocrine and seasonal signals are well established, much less is known about the cellular changes required to create this environment. Here we present the first detailed study of uterine morphology during diapause and early pregnancy of the tammar wallaby. We combined transmission electron microscopy and light microscopy to describe the histological and ultrastructural changes to luminal and glandular epithelial cells. At entry into diapause after the post-partum oestrus and formation of the new conceptus, there was an increase in abundance of organelles associated with respiration in the endometrial cells of the newly gravid uterus, particularly in the endoplasmic reticulum and mitochondria, as well as an increase in secretory activity. Organelle changes and active secretion then ceased in these cells as they became quiescent and remained so for the duration of diapause. In contrast, cells of the non-gravid, post-partum, contralateral uterus underwent sloughing and remodelling during this time and some organelle changes in glandular epithelial cells continued throughout diapause, suggesting these cells are not completely quiescent during diapause, although no active secretion occurred. These findings demonstrate that diapause, like pregnancy, is under unilateral endocrine control in the tammar, and that preparation for and maintenance of diapause requires substantial changes to uterine endometrial cell ultrastructure and activity., (© 2016 Anatomical Society.)

A study concerning the upper (UE) and lower (LE) eyelids, lacrimal gland (LG), superficial gland of the third eyelid (SGTE) and third eyelid (TE) was conducted on 4 sexually mature red kangaroos (2 males and 2 females). Gross anatomical, histological, histometrical, histochemical and ultrastructural (TEM) components were compared. Tissue sections were stained with haematoxylin and eosin, azan-trichrome, van Gieson, Masson-Goldner trichrome, methyl green-pyronin Y, periodic acid-Schiff, alcian blue pH 2.5, aldehyde fuchsin and Hale's dialysed iron. The location of the LG, SGTE, TE, UE and LE was similar to that of other mammals. Organized lymphoid follicles were also found in the LE. The TE resembled the letter T and was composed of cartilage (hyaline tissue). The LG was relatively larger than the SGTE. The LG and SGTE were multilobar tubuloacinar glands. The LG had more plasma cells than the SGTE. The SGTE and LG secretions were mucoserous in composition. The TEM study showed that the secretory cells of the LG and SGTE have a similar ultrastructural appearance. Two types of secretory vesicles were detected in the cytoplasm in acini and one type of secretory vesicle was found in the tubules of these glands.

The heart and left ventricle of the marsupial western grey kangaroo Macropus fuliginosus exhibit biphasic allometric growth, whereby a negative shift in the trajectory of cardiac growth occurs at pouch exit. In this study, we used transmission electron microscopy to examine the scaling of left ventricle cardiomyocyte ultrastructure across development in the western grey kangaroo over a 190-fold body mass range (0.355-67.5 kg). The volume-density (%) of myofibrils, mitochondria, sarcoplasmic reticuli and T-tubules increase significantly during in-pouch growth, such that the absolute volume (ml) of these organelles scales with body mass (Mb; kg) with steep hyperallometry: 1.41Mb (1.38), 0.64Mb (1.29), 0.066Mb (1.45) and 0.035Mb (1.87), respectively. Maturation of the left ventricle ultrastructure coincides with pouch vacation, as organelle volume-densities scale independent of body mass across post-pouch development, such that absolute organelle volumes scale in parallel and with relatively shallow hypoallometry: 4.65Mb (0.79), 1.75Mb (0.77), 0.21Mb (0.79) and 0.35Mb (0.79), respectively. The steep hyperallometry of organelle volumes and volume-densities across in-pouch growth is consistent with the improved contractile performance of isolated cardiac muscle during fetal development in placental mammals, and is probably critical in augmenting cardiac output to levels necessary for endothermy and independent locomotion in the young kangaroo as it prepares for pouch exit. The shallow hypoallometry of organelle volumes during post-pouch growth suggests a decrease in relative cardiac requirements as body mass increases in free-roaming kangaroos, which is possibly because the energy required for hopping is independent of speed, and the capacity for energy storage during hopping could increase as the kangaroo grows., (© 2015. Published by The Company of Biologists Ltd.)

Exceptionally well-preserved skulls and postcranial elements of a new species of the plesiomorphic stem macropodiform Balbaroo have been recovered from middle Miocene freshwater limestone deposits in the Riversleigh World Heritage Area of northwestern Queensland, Australia. This constitutes the richest intraspecific sample for any currently known basal "kangaroo", and, along with additional material referred to Balbaroo fangaroo, provides new insights into structural variability within the most prolific archaic macropodiform clade--Balbaridae. Qualitative and metric evaluations of taxonomic boundaries demonstrate that the previously distinct species Nambaroo bullockensis is a junior synonym of B. camfieldensis. Furthermore, coupled Maximum Parsimony and Bayesian phylogenetic analyses reveal that our new Balbaroo remains represent the most derived member of the Balbaroo lineage, and are closely related to the middle Miocene B. camfieldensis, which like most named balbarid species is identifiable only from isolated jaws. The postcranial elements of Balbaroo concur with earlier finds of the stratigraphically oldest balbarid skeleton, Nambaroo gillespieae, and suggest that quadrupedal progression was a primary gait mode as opposed to bipedal saltation. All Balbaroo spp. have low-crowned bilophodont molars, which are typical for browsing herbivores inhabiting the densely forested environments envisaged for middle Miocene northeastern Australia.

Sthenurine kangaroos (Marsupialia, Diprotodontia, Macropodoidea) were an extinct subfamily within the family Macropodidae (kangaroos and rat-kangaroos). These "short-faced browsers" first appeared in the middle Miocene, and radiated in the Plio-Pleistocene into a diversity of mostly large-bodied forms, more robust than extant forms in their build. The largest (Procoptodon goliah) had an estimated body mass of 240 kg, almost three times the size of the largest living kangaroos, and there is speculation whether a kangaroo of this size would be biomechanically capable of hopping locomotion. Previously described aspects of sthenurine anatomy (specialized forelimbs, rigid lumbar spine) would limit their ability to perform the characteristic kangaroo pentapedal walking (using the tail as a fifth limb), an essential gait at slower speeds as slow hopping is energetically unfeasible. Analysis of limb bone measurements of sthenurines in comparison with extant macropodoids shows a number of anatomical differences, especially in the large species. The scaling of long bone robusticity indicates that sthenurines are following the "normal" allometric trend for macropodoids, while the large extant kangaroos are relatively gracile. Other morphological differences are indicative of adaptations for a novel type of locomotor behavior in sthenurines: they lacked many specialized features for rapid hopping, and they also had anatomy indicative of supporting their body with an upright trunk (e.g., dorsally tipped ischiae), and of supporting their weight on one leg at a time (e.g., larger hips and knees, stabilized ankle joint). We propose that sthenurines adopted a bipedal striding gait (a gait occasionally observed in extant tree-kangaroos): in the smaller and earlier forms, this gait may have been employed as an alternative to pentapedal locomotion at slower speeds, while in the larger Pleistocene forms this gait may have enabled them to evolve to body sizes where hopping was no longer a feasible form of more rapid locomotion.

Introduction: The tammar wallaby has a short gestation (26.5 days) and vascular modifications to expedite transport during that brief pregnancy. Here we examine trophoblast structural attributes that would facilitate materno-fetal exchange., Materials and Methods: Four specimens of Macropus eugenii between days 23 and 26 gestation were examined using electron microscopy and 24 lectins to characterise glycosylated secretions and their internalisation., Results: Two trophoblast phenotypes were found, flattened cells generally in contact with the underlying uterine epithelium and giant cells associated with histiotrophe. The latter appeared to penetrate uterine clefts, occasionally detach and become necrotic. Lectin histochemistry and ultrastructure indicated the presence of many lysosomes and residual bodies especially in trophoblast giant cells; these contained glycans, mainly apically, which were also detected in secretions and cell debris. Trophoblast basal membranes bore extensive filopodia. Giant cells were less common in vascular trilaminar areas and here the trophoblast barrier became thinner near term., Discussion: Loss of Maackia amurensis agglutinin binding suggested cleavage of terminal sialic acid residues as an early post-internalisation event in the trophoblast. Lectin staining indicated degradation occurred in an apical-basal direction, and the heavily glycosylated basal membrane appeared specialised for transport out of the cell., Conclusion: Granules seen ultrastructurally and histochemically, particularly in giant trophoblast cells of the bilaminar area, suggest that internalised histiotrophe is broken down here and nutrients transferred to the embryo via the specialised basal plasma membrane. The trilaminar vascular area contained mostly flattened trophoblast cells, supporting the suggestion that gaseous exchange is its primary function., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

Introduction: Blood vessel glycosylation at the fetomaternal interface of four near-term specimens of tammar wallaby, Macropus eugenii, has been examined at days 23-26 of the 26.5 day pregnancy and compared with that of other species., Methods: A panel of 23 lectins was used to compare vasculature in tammar with non-mammalian (shark, skink) and eutherian species at early and late gestation (camel, horse and alpaca), and term/near-term (cat, lion, dog, mink and elephant)., Results: Strikingly low levels of all the glycans tested, apart from sialic acids, were found in capillary endothelium of both the trilaminar omphalopleure and underlying surface endometrium of the tammar, though deeper endometrial vessels showed normally high levels of glycosylation. Only maternal vasculature of the mink placenta showed a comparable lack of expression., Discussion: One reason for a reduced endothelial glycocalyx may be to facilitate diffusion of gases and nutrients as the tammar trophoblast lacks the indentation by overlying vessels that is seen in the other near-term placentae. Early epitheliochorial placentae of other species with equal diffusion distances to the tammar, showed normal vascular glycosylation. However, their pregnancies are much longer., Conclusion: The hypoglycosylation of tammar vessels at the fetomaternal interface may allow continued transfer of nutrients and gaseous exchange during the extremely rapid period of organogenesis which occurs during the short 26.5 day pregnancy of this marsupial. Given the short gestation period of the tammar, we suggest that a thinner endothelial glycocalyx has evolved to facilitate diffusion of gases and nutrients between the maternal and fetal compartments., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Rensch's rule, which states that the magnitude of sexual size dimorphism tends to increase with increasing body size, has evolved independently in three lineages of large herbivorous mammals: bovids (antelopes), cervids (deer), and macropodids (kangaroos). This pattern can be explained by a model that combines allometry, life-history theory, and energetics. The key features are that female group size increases with increasing body size and that males have evolved under sexual selection to grow large enough to control these groups of females. The model predicts relationships among body size and female group size, male and female age at first breeding, death and growth rates, and energy allocation of males to produce body mass and weapons. Model predictions are well supported by data for these megaherbivores. The model suggests hypotheses for why some other sexually dimorphic taxa, such as primates and pinnipeds (seals and sea lions), do or do not conform to Rensh's rule.

The mammalian cerebellar cortex is apparently uniform in composition, but a complex heterogeneous pattern can be revealed by using biochemical markers such as zebrin II/aldolase C, which is expressed by a subset of Purkinje cells that form a highly reproducible array of transverse zones and parasagittal stripes. The architecture revealed by zebrin II expression is conserved among many taxa of birds and mammals. In this report zebrin II immunohistochemistry has been used in both section and whole-mount preparations to analyze the cerebellar architecture of the Australian tammar wallaby (Macropus eugenii). The gross appearance of the wallaby cerebellum is remarkable, with unusually elaborate cerebellar lobules with multiple sublobules and fissures. However, despite the morphological complexity, the underlying zone and stripe architecture is conserved and the typical mammalian organization is present., (Copyright © 2012 S. Karger AG, Basel.)

Tree-kangaroos are a unique group of arboreal marsupials that evolved from terrestrial ancestors. The recent discovery of well-preserved specimens of extinct tree-kangaroo species (genus Bohra) within Pleistocene cave deposits of south-central Australia provides a unique opportunity to examine adaptive evolution of tree-kangaroos. Here, we provide the first detailed description of the functional anatomy of the forelimb, a central component of the locomotor complex, in the extant Dendrolagus lumholtzi, and compare its structure and function with representatives of other extant marsupial families. Several features were interpreted as adaptations for coping with a discontinuous, uneven and three-dimensional arboreal substrate through enhanced muscular strength and dexterity for propulsion, grasping, and gripping with the forelimbs. The forelimb musculoskeletal anatomy of Dendrolagus differed from terrestrial kangaroos in the following principal ways: a stronger emphasis on the development of muscles groups responsible for adduction, grasping, and gripping; the enlargement of muscles that retract the humerus; and modified shape of the scapula and bony articulations of the forelimb bones to allow improved mobility. Many of these attributes are convergent with other arboreal marsupials. Tree-kangaroos, however, still retain the characteristic bauplan of their terrestrial ancestors, particularly with regard to skeletal morphology, and the muscular anatomy of the forelimb highlights a basic conservatism within the group. In many instances, the skeletal remains of Bohra have similar features to Dendrolagus that suggest adaptations to an arboreal habit. Despite the irony of their retrieval from deposits of the Nullarbor "Treeless" Plain, forelimb morphology clearly shows that the species of Bohra were well adapted to an arboreal habitat., (Copyright © 2011 Wiley-Liss, Inc.)

Hypospadias is increasingly common, and requires surgery to repair, but its aetiology is poorly understood. The marsupial tammar wallaby provides a unique opportunity to study hypospadias because penile differentiation occurs postnatally. Androgens are responsible for penile development in the tammar, but the majority of differentiation, in particular formation and closure of the urethral groove forming the penile urethra in males, occurs when there is no measurable sex difference in the concentrations of testosterone or dihydrotestosterone in either the gonads or the circulation [corrected]. Phalluses were examined morphologically from the sexually indifferent period (when androgens are high) to well after the time that the phallus becomes sexually dimorphic. We show that penile development and critical changes in the positioning of the urethra occur in the male phallus begin during an early window of time when androgens are high. Remodelling of the urethra in the male occurs between days 20-60. The critical period of time for the establishment urethral closure occurs during the earliest phases of penile development. This study suggests that there is an early window of time before day 60 when androgen imprinting must occur for normal penile development and closure of the urethral groove., (Copyright © 2011 S. Karger AG, Basel.)

Objective: To perform electroretinography on normal anesthetized western gray kangaroos (Macropus fuliginosus). Animals studied  Six captive western gray kangaroos., Procedures: The kangaroos were anesthetized using a combination of ketamine and medetomidine via a remote drug delivery system, then were maintained on isoflurane after endotracheal intubation and reversal of the medetomidine with atipamazole. After a minimum of 20 min of dark adaptation, electroretinograms were obtained using a handheld electroretinography (ERG) machine using a single flash protocol at three light intensities: 10 mcd.s/m(2), 3000 mcd.s/m(2), 10 000 mcd.s/m(2)., Results: At 10 mcd.s/m(2) the mean b-wave amplitude and implicit time was 102.0 μV (SD ± 41.3 and 95% CI 68.9-135.1) and 78.4 ms (SD ± 8.3 and 95% CI 71.8-85.0). At 3000 mcd.s/m(2) the mean a-wave amplitude and implicit time was 69.9 μV (SD ± 20.5 and 95% CI 53.5-86.3) and 17.6 ms (SD ± 1.5 and 95% CI 16.4-18.8) and the mean b-wave amplitude and implicit time was 175.4 μV (SD ± 35.9 and 95% CI 146.7-204.1) and 74.1 ms (SD ± 3.5 and 95% CI 71.2-76.9). At 10 000 mcd.s/m(2) the mean a-wave amplitude and implicit time was 89.1 μV (SD ± 27.1 and 95% CI 67.5-110.8) and 16.8 ms (SD ± 1.0 and 95% CI 16.0-17.0) and the mean b-wave amplitude and implicit time was 203.7 μV (SD ± 41.4 and 95% CI 170.6-236.8) and 75.4 ms (SD ± 3.3 and 95% CI 72.8-78.1)., Conclusion: Electroretinography outside of the typical clinical setting is feasible using a portable ERG system and allows for quick analysis of retinal function in exotic species.

CART (cocaine- and amphetamine-regulated transcript) is a neuromodulator involved in feeding, drug reward, stress and cardiovascular function. We have immunohistochemically studied the distribution of the CART peptide in the brains of two adult marsupial species: the brown antechinus (Antechinus stuartii) as a representative of polyprotodont marsupials and the tammar wallaby (Macropus eugenii) as a representative of diprotodont marsupials. We have also examined the distribution of CART during postnatal development in the tammar wallaby. There were similarities and differences both between the two marsupial species and between the marsupials and eutherians in CART distribution. Both marsupials showed immunoreactivity to CART in the olfactory bulb, piriform cortex, extended amygdala, the supraoptic, paraventricular and arcuate nuclei of the hypothalamus, somatosensory and auditory nuclei of the brainstem, vagal/solitary complex, raphe obscurus and raphe pallidus and presumptive presympathetic neurons of the ventrolateral medulla, as has been seen in eutherians. On the other hand, immunoreactivity to CART was weak in or absent from isocortical areas, and immunoreactivity to CART was poor or minimal in the ventral tegmental area and nucleus accumbens of both species; regions where immunoreactivity to CART is very strong in the brains of eutherians. During development, CART was present at birth (P0) in the lateral trigeminal ganglion, spinal trigeminal tract and the vagal sensorimotor complex, but did not appear in mid- or forebrain regions until much later (from P37). These anatomical findings indicate that although CART is likely to serve very similar functions in both eutherians and marsupials, there are potentially functionally significant differences between the two mammalian groups., (Copyright © 2010 S. Karger AG, Basel.)

In kangaroos and wallabies at birth the highly altricial newborn young climbs unassisted from the urogenital opening to the teat. Negative geotropism is important for the initial climb to the pouch opening, but nothing is known of the signals that then direct the neonate downwards to the teat. Here we show that the newborn tammar wallaby (Macropus eugenii) has the olfactory apparatus to detect smell. Both the main olfactory system and vomeronasal organ (VNO) are developed at the time of birth. Receptor cells of the main olfactory epithelium immunopositive for G(oalpha)-protein project to the three layered main olfactory bulb (MOB). The receptor epithelium of the VNO contains G-protein immunopositive cells and has olfactory knob-like structures. The VNO is connected to an area between the two MOBs. Next, using a functional test, we show that neonates can respond to odours from their mother's pouch. When neonatal young are presented with a choice of a pouch-odour-soaked swab or a saline swab, they choose the swab with their mother's pouch secretions significantly more often (P<0.05) than the saline swab. We conclude that both olfactory systems are capable of receiving odour signals at birth, a function that must be a critical adaptation for the survival of an altricial marsupial neonate such as the tammar for its journey to the pouch.

Kangaroos are the world's most diverse group of herbivorous marsupials. Following late-Miocene intensification of aridity and seasonality, they radiated across Australia, becoming the continent's ecological equivalents of the artiodactyl ungulates elsewhere. Their diversity peaked during the Pleistocene, but by approximately 45,000 years ago, 90% of larger kangaroos were extinct, along with a range of other giant species. Resolving whether climate change or human arrival was the principal extinction cause remains highly contentious. Here we combine craniodental morphology, stable-isotopic, and dental microwear data to reveal that the largest-ever kangaroo, Procoptodon goliah, was a chenopod browse specialist, which may have had a preference for Atriplex (saltbushes), one of a few dicots using the C(4) photosynthetic pathway. Furthermore, oxygen isotope signatures of P. goliah tooth enamel show that it drank more in low-rainfall areas than its grazing contemporaries, similar to modern saltbush feeders. Saltbushes and chenopod shrublands in general are poorly flammable, so landscape burning by humans is unlikely to have caused a reduction in fodder driving the species to extinction. Aridity is discounted as a primary cause because P. goliah evolved in response to increased aridity and disappeared during an interval wetter than many it survived earlier. Hunting by humans, who were also bound to water, may have been a more decisive factor in the extinction of this giant marsupial.

The jaw muscles were studied in seven genera of macropodoid marsupials with diets ranging from mainly fungi in Potorous to grass in Macropus. Relative size, attachments, and lamination within the jaw adductor muscles varied between macropodoid species. Among macropodine species, the jaw adductor muscle proportions vary with feeding type. The relative mass of the masseter is roughly consistent, but grazers and mixed-feeders (Macropus and Lagostrophus) had relatively larger medial pterygoids and smaller temporalis muscles than the browsers (Dendrolagus, Dorcopsulus, and Setonix). Grazing macropods show similar jaw muscle proportions to "ungulate-grinding" type placental mammals. The internal architecture of the jaw muscles also varies between grazing and browsing macropods, most significantly, the anatomy of the medial pterygoid muscle. Potoroines have distinctly different jaw muscle proportions to macropodines. The masseter muscle group, in particular, the superficial masseter is enlarged, while the temporalis group is relatively reduced. Lagostrophus fasciatus is anatomically distinct from other macropods with respect to its masticatory muscle anatomy, including enlarged superficial medial pterygoid and deep temporalis muscles, an anteriorly inflected masseteric process, and the shape of the mandibular condyle. The enlarged triangular pterygoid process of the sphenoid bone, in particular, is distinctive of Lagsotrophus., ((c) 2009 Wiley-Liss, Inc.)

Macropods are herbivorous foregut-fermenters that have adapted to a wide variety of habitats. Anatomic adaptations such as dentition reflect differences in the diet consumed in the wild. Several key differences exist in anatomy and digestive physiology between macropods and other foregut-fermenters such as ruminants. The diet fed to macropods in captivity should be formulated with species-specific wild diets as a model. Hand-rearing orphaned macropods should be done with the unique physiology of macropod lactation in mind to ensure success. This article provides a summary of anatomic, physiologic, metabolic, and behavioral peculiarities of kangaroos and wallabies, with recommendations on captive management based on these traits.

Fragments of bone shafts that lack diagnostic features can be difficult to identify as human or non-human-an important task in forensic science and archaeology. Some workers have found the thickness of cortical bone in the shaft to be a useful distinction, although the sparse literature in the field is contradictory in how this may be applied. The aim of the present study was to determine whether any difference is discernible between humans, kangaroos and sheep (mammals whose remains are commonly confused with those of humans in Australia) at the femoral midshaft, with a view to a larger-scale analysis if differences were discovered. Cross-sections at the midpoint of the shaft were measured to determine the diameter of the whole shaft and the medullary cavity on each bone; an index describing cortical thickness relative to shaft diameter was calculated. Statistically significant differences were found between all three groups, with humans showing the thickest cortical bone, and sheep the thinnest. These differences may be linked to a higher load on the human femur, due to a larger body mass carried on two legs, as opposed to the sheep's four. Further work now needs to be carried out to determine if differences are present when comparing multiple sites on the skeleton, and between non-human mammals of different sizes.

The vomeronasal organ is the primary olfactory organ that detects sexual pheromones in mammals. We investigated the anatomy of the vomeronasal organ of the tammar wallaby (Macropus eugenii), a small macropodid marsupial. Pheromones may be important for activation of the hypothalamo-pituitary axis of tammar males at the start of the breeding season because plasma testosterone and luteinizing hormone concentration in males rise concurrently with pregnancy and the post-partum ovulation in females. The gross anatomy and the connection to the brain of the vomeronasal organ were examined by light and electron microscopy in adult male and female tammars. The vomeronasal organ was well developed in both sexes. The vomeronasal organ is a tubular organ connected at the rostral end via the nasopalatine duct (incisive duct) to the mouth and nasal cavity. At the rostral end the lumen of the vomeronasal organ was crescent shaped, changing to a narrow oval shape caudally. Glandular tissue associated with the vomeronasal organ increased towards the blind end of the organ. The tammar has the typical pattern of mammalian vomeronasal organs with electron-dense supporting cells and electron-lucent receptor cells. Microvilli were present on the surface of both epithelia while cilia were only found on the surface of the non-receptor epithelium. Some non-receptor epithelial cells appeared to secrete mucus into the vomeronasal organ lumen. The vomeronasal organ shows a high degree of structural conservation compared with eutherian mammals. The degree of vomeronasal organ development makes it likely that, as in other mammals, pheromones are important in the reproduction of the tammar.

The aim of this study was to examine hind limb scaling of the musculoskeletal system in the Macropodoidea, the superfamily containing wallabies and kangaroos, to re-examine the effect of size on the locomotor mechanics and physiology of marsupial hopping. Morphometric musculoskeletal analyses were conducted of 15 species and skeletal specimens of 21 species spanning a size range from 0.8 to 80 kg that included representatives of 12 of the 16 extant genera of macropodoids. We found that unlike other groups, macropodoids are able to match force demands associated with increasing body size primarily through a combination of positive allometry in muscle area and muscle moment arms. Isometric scaling of primary hind limb bones suggests, however, that larger species experience relatively greater bone stresses. Muscle to tendon area ratios of the ankle extensors scale with strong positive allometry, indicating that peak tendon stresses also increase with increasing body size but to a lesser degree than previously reported. Consistent with previous morphological and experimental studies, large macropodoids are therefore better suited for elastic strain energy recovery but operate at lower safety factors, which likely poses an upper limit to body size. Scaling patterns for extant macropodoids suggest that extinct giant kangaroos (approximately 250 kg) were likely limited in locomotor capacity.

We used carbocyanine dye tracing techniques in conjunction with hematoxylin and eosin staining, immunohistochemistry for GAP-43, and tritiated thymidine autoradiography to examine the development of the olfactory pathways in early pouch young tammar wallabies (Macropus eugenii). The overarching aim was to test the hypothesis that the olfactory system of newborn tammars is sufficiently mature at birth to contribute to the guidance of the pouch young to the nipple. Although GAP-43 immunoreactive fibers emerge from the olfactory epithelium and enter the olfactory bulb at birth, all other components of the olfactory pathway in newborn tammars are very immature at birth, postnatal day (P0). In particular, maturation of the vomeronasal organ and its projections to the accessory olfactory bulb appears to be delayed until P5 and the olfactory bulb is poorly differentiated until P12, with glomerular formation delayed until P25. The lateral olfactory tract is also very immature at birth with pioneer axons having penetrated only the most rostral portion of the piriform lobe. Interestingly, there were some early (P0) projections from the olfactory epithelium to the medial septal region and lamina terminalis (by the terminal nerve) and to olfactory tubercle and basal forebrain. The former of these is presumably serving the transfer of LHRH(+) neurons to the forebrain, as seen in eutherians, but neither of these very early pathways is sufficiently robust or connected to the more caudal neuraxis to play a role in nipple finding. Tritiated thymidine autoradiography confirmed that most piriform cortex pyramidal neurons are generated in the first week of life and are unlikely to be able to contribute to circuitry guiding the climb to the pouch. Our findings lead us to reject the hypothesis that olfactory projections contribute to guidance of the newborn tammar to the pouch and nipple. It appears far more likely that the trigeminal pathways play a significant role in this behavior because the central projections of the trigeminal nerve are more mature at birth in this marsupial., (Copyright 2008 S. Karger AG, Basel.)

We have studied the early development of the vestibular apparatus and its central connections in the tammar wallaby (Macropus eugenii) in order to determine whether the vestibular system anatomy is sufficiently mature at birth to assist in climbing to the pouch. Structural development was studied with the aid of hematoxylin and eosin stained sections and immunoreactivity for GAP-43, whereas the development of vestibular system connections was examined by carbocyanine dye tracing. At the time of birth, the otocyst has distinct utricle, saccule and semicircular canals with immature sensory regions receiving innervation by GAP-43 immunoreactive fibers. Vestibular nerve fibers can be traced into the brainstem to the developing vestibular nuclei, which are not yet cytoarchitectonically distinct. The vestibular nuclei do not contribute direct projections to the lower cervical spinal cord at birth; most bulbospinal projections in the newborn appear to be derived bilaterally from the gigantocellular, lateral paragigantocellular reticular and ventral medullary nuclei. A substantial bilateral projection to the vestibular ganglion and apparatus from the region of the gigantocellular and lateral paragigantocellular nuclei was seen at birth, but not in subsequent ages. This is similar to a projection seen in newborn Ameridelphians. By postnatal day (P) 5, the vestibular apparatus had extensive projections to all vestibular nuclei and neurons projecting in the lateral vestibulospinal tract could be identified in the lateral vestibular nucleus. Cytoarchitectonic differentiation of the vestibular nuclei proceeded over the next 3 to 4 weeks with the emergence of discrete parvicellular and magnocellular components of the medial vestibular nucleus by P19. GAP-43 immunoreactivity stayed high in the lateral vestibulospinal tract for several months after birth, suggesting that the development of this tract followed a prolonged timecourse. Our findings indicate that central and peripheral connections of the vestibular ganglion are present at birth, but that there is no direct projection from the vestibular nuclei to the cervical spinal cord until P5. Nevertheless, the possibility remains that an indirect projection between the vestibular nuclei and the medial reticular formation is present at birth and mediates control of the climb., ((c) 2008 S. Karger AG, Basel)

This study describes pubertal changes in testes and epididymides and seasonal changes in the adult male reproductive organs and plasma androgen concentrations of the swamp wallaby (Wallabia bicolor). Pre-pubescent males had testes with solid seminiferous cords and spermatogenesis only to the stage of gonocytes. Their epididymides had empty lumina along their entire length. The testes of three males undergoing puberty had some lumen formation and mitotic activity. Their epididymides were similar in appearance to those of adult males but were entirely devoid of any cells within the lumen of the duct. Three other pubescent males showed full lumen formation in the testes and spermatogenesis up to the elongating spermatid stage. Their epididymides were similar in appearance to those of adult males but with no spermatozoa in the duct. However, cells of testicular origin were found in the lumen of the duct in all regions suggesting that testicular fluids and immature germ cells shed into the rete testes flow through the seminiferous tubules into the epididymis before the release of mature testicular spermatozoa. The weights of testes and epididymides of adult males showed no change throughout the year but prostate weight and plasma androgen concentrations varied significantly with season, with maximums in spring and summer and minimums in winter. The volume fraction of Leydig cells and seminiferous tubules was significantly lower in winter than in summer; but, despite this, maturing spermatozoa were found in the testes throughout the year. Females in the area conceived year-round, suggesting that seasonal changes in the male reproductive tract did not prevent at least some males from breeding throughout the year.

A morphological and morphometric study of the lung of the newborn quokka wallaby (Setonix brachyurus) was undertaken to assess its morphofunctional status at birth. Additionally, skin structure and morphometry were investigated to assess the possibility of cutaneous gas exchange. The lung was at canalicular stage and comprised a few conducting airways and a parenchyma of thick-walled tubules lined by stretches of cuboidal pneumocytes alternating with squamous epithelium, with occasional portions of thin blood-gas barrier. The tubules were separated by abundant intertubular mesenchyme, aggregations of developing capillaries and mesenchymal cells. Conversion of the cuboidal pneumocytes to type I cells occurred through cell broadening and lamellar body extrusion. Superfluous cuboidal cells were lost through apoptosis and subsequent clearance by alveolar macrophages. The establishment of the thin blood-gas barrier was established through apposition of the incipient capillaries to the formative thin squamous epithelium. The absolute volume of the lung was 0.02 +/- 0.001 cm(3) with an air space surface area of 4.85 +/- 0.43 cm(2). Differentiated type I pneumocytes covered 78% of the tubular surface, the rest 22% going to long stretches of type II cells, their precursors or low cuboidal transitory cells with sparse lamellar bodies. The body weight-related diffusion capacity was 2.52 +/- 0.56 mL O(2) min(-1) kg(-1). The epidermis was poorly developed, and measured 29.97 +/- 4.88 microm in thickness, 13% of which was taken by a thin layer of stratum corneum, measuring 4.87 +/- 0.98 microm thick. Superficial capillaries were closely associated with the epidermis, showing the possibility that the skin also participated in some gaseous exchange. Qualitatively, the neonate quokka lung had the basic constituents for gas exchange but was quantitatively inadequate, implying the significance of percutaneous gas exchange.

Neuronal activity is implicated as a driving force in the development of sensory systems. In order for it to play a developmental role, however, the pathways involved must be capable of transmitting this activity. The relationship between afferent arrival, synapse formation and the onset of chemical neurotransmission has been examined using the advantageous model of a marsupial mammal, the wallaby (Macropus eugenii), to determine at what stage activity has the capacity to influence cortical development. It is known that thalamocortical afferents arrive in the somatosensory cortex on postnatal day (P)15 and that their growth cones reach to the base of the compact cell zone of the cortical plate. However, electronmicroscopy showed that thalamocortical synapses were absent at this stage. Glutamatergic responses were recorded in the cortex following stimulation of the thalamus in slices at this time but only in magnesium-free conditions. The responses were mediated entirely by N-methyl-d-aspartate (NMDA) receptors. From P28, responses could be recorded in normal magnesium and comprised a dominant NMDA-mediated component and a non-NMDA mediated component. At this time thalamocortical synapses were first identified and they were in the cortical plate. By P63 the non-NMDA-mediated component had increased relative to the NMDA-mediated component, and by P70 layer IV began to emerge and contained thalamocortical synapses. By P76 a fast non-NMDA-mediated peak dominated the response. This coincides with the appearance of cortical whisker-related patches and the onset in vivo of responses to peripheral stimulation of the whiskers.

The great-gray kangaroo (Macropus giganteus) belongs to the Diprotodontia suborder (herbivorous marsupials of Australia) of the order of marsupials. We dissected the masticatory muscles in the great-gray kangaroo and classified them based on their innervation. Three (two male and one female) adult great-gray kangaroos (M. giganteus), fixed with 10% formalin, were examined. The masseter muscle of the great-gray kangaroo was classified into four layers (superficial layers 1, 2, 3, and a deep layer), all innervated by masseteric nerves. Layer 1 of the masseter muscle was well developed and the deep layer inserted into the masseteric canal. The zygomaticomandibular muscle, which belongs to both the masseter and temporalis muscles, was innervated by both the masseteric nerve and posterior deep temporal nerve, and the temporalis muscle was innervated by the anterior and posterior deep temporal nerves. The medial pterygoid muscle, which was innervated by the medial pterygoid nerve, was divided into superficial and deep portions. The lateral pterygoid muscle was divided into superior and inferior heads by the buccal nerve. We propose that the relationship of the masticatory muscles in the kangaroo has evolved by passive anterior invasion of the deep layer of the masseter by the medial pterygoid muscle via the masseteric canal, associated with the development of an anteroposterior mode of mastication.

Plasticity within the visual system was assessed in the quokka wallaby following unilateral superior collicular (SC) ablation at postnatal days (P) 8-10, prior to the arrival of retinal ganglion cell (RGC) axons. At maturity (P100), projections were traced from the eye opposite the ablation, and total RGC numbers were estimated for both eyes. Ablations were partial (28-89% of SC remaining) or complete (0-5% of SC remaining). Projections to the visual centers showed significant bilateral (P < 0.05) increases in absolute volume. Minor anomalous projections also formed within the deep, surviving non-retino-recipient layers of the ablated SC and via a small bundle of RGC axons recrossing the midline to innervate discrete patches in the SC contralateral to the lesion. Total absolute volume of projections did not differ between partial and complete ablations; moreover, values did not differ from normal (P > 0.05). Compared with normal, total RGC numbers were significantly (P < 0.05) reduced in the eye opposite the ablation but increased (P < 0.05) in the other eye. Consequently, the sum of the two RGC populations did not differ from normal (P > 0.05). As in rodents, the visual system in quokka compensates following injury by maintaining a set volume of arborization but does so by forming only minor anomalous projections. Furthermore, increased RGC numbers in the eye ipsilateral to the lesion indicate that compensation occurs transneuronally, thus maintaining total numbers of projecting neurons. The implication is that the visual system acts in concert following unilateral injury to maintain set values for RGC terminal arbors as well as their cell bodies.

The biochemical composition of uterine and fetal fluids during pregnancy of the grey short-tailed opossum was compared with new and published data on the tammar wallaby. In the grey short-tailed opossum, there are three main phases of embryonic nourishment. During the first phase, the embryo obtains nutrients from uterine secretion transferred into the yolk sac. The amount of uterine secretion declines during the second phase up to the time of shell coat rupture. As a result, the protein concentration in yolk sac fluid also declines. During phase three, which begins with shell coat rupture, nutrients are predominantly available from the maternal blood. In the grey short-tailed opossum that lacks a vesicular, fluid-filled allantois, waste products such as urea are apparently stored in the yolk sac and from there pass into the maternal circulation across the invasive yolk sac placenta. In contrast, in the tammar wallaby, the main source of nutrients available to the late term fetus is glandular secretion that is complemented by substances from the maternal circulation via the chorio-vitelline placenta, and waste products are stored in the large, fluid-filled allantois.