G. E. Weissengruber

Hyoid apparatus and pharynx in the lion (Panthera leo), jaguar (Panthera onca), tiger (Panthera tigris), cheetah (Acinonyx jubatus) and domestic cat (Felis silvestris f. catus)

Structures of the hyoid apparatus, the pharynx and their topographical positions in the lion, tiger, jaguar, cheetah and domestic cat were described in order to determine morphological differences between species or subfamilies of the Felidae. In the lion, tiger and jaguar (species of the subfamily Pantherinae) the Epihyoideum is an elastic ligament lying between the lateral pharyngeal muscles and the Musculus (M.) thyroglossus rather than a bony element like in the cheetah or the domestic cat. The M. thyroglossus was only present in the species of the Pantherinae studied. In the lion and the jaguar the Thyrohyoideum and the thyroid cartilage are connected by an elastic ligament, whereas in the tiger there is a synovial articulation. In adult individuals of the lion, tiger and jaguar the ventral end of the tympanohyal cartilage is rotated and therefore the ventral end of the attached Stylohyoideum lies caudal to the Tympanohyoideum and the cranial base. In newborn jaguars the Apparatus hyoideus shows a similar topographical position as in adult cheetahs or domestic cats. In adult Pantherinae, the Basihyoideum and the attached larynx occupy a descended position: they are situated near the cranial thoracic aperture, the pharyngeal wall and the soft palate are caudally elongated accordingly. In the Pantherinae examined the caudal end of the soft palate lies dorsal to the glottis. Differences in these morphological features between the subfamilies of the Felidae have an influence on specific structural characters of their vocalizations.

The structure of the cushions in the feet of African elephants (Loxodonta africana).

The uniquely designed limbs of the African elephant, Loxodonta africana, support the weight of the largest terrestrial animal. Besides other morphological peculiarities, the feet are equipped with large subcutaneous cushions which play an important role in distributing forces during weight bearing and in storing or absorbing mechanical forces. Although the cushions have been discussed in the literature and captive elephants, in particular, are frequently affected by foot disorders, precise morphological data are sparse. The cushions in the feet of African elephants were examined by means of standard anatomical and histological techniques, computed tomography (CT) and magnetic resonance imaging (MRI). In both the forelimb and the hindlimb a 6th ray, the prepollex or prehallux, is present. These cartilaginous rods support the metacarpal or metatarsal compartment of the cushions. None of the rays touches the ground directly. The cushions consist of sheets or strands of fibrous connective tissue forming larger metacarpal/metatarsal and digital compartments and smaller chambers which were filled with adipose tissue. The compartments are situated between tarsal, metatarsal, metacarpal bones, proximal phalanges or other structures of the locomotor apparatus covering the bones palmarly/plantarly and the thick sole skin. Within the cushions, collagen, reticulin and elastic fibres are found. In the main parts, vascular supply is good and numerous nerves course within the entire cushion. Vater–Pacinian corpuscles are embedded within the collagenous tissue of the cushions and within the dermis. Meissner corpuscles are found in the dermal papillae of the foot skin. The micromorphology of elephant feet cushions resembles that of digital cushions in cattle or of the foot pads in humans but not that of digital cushions in horses. Besides their important mechanical properties, foot cushions in elephants seem to be very sensitive structures.

Anatomical Description of the Muscles of the Pelvic Limb in the Ostrich (Struthio camelus)

Dissections of 12 formalin‐fixed ostriches were performed to give anatomical descriptions of the muscles and tendons of the pelvic, femoral, tibiotarsal, tarsometatarsal and digital regions. In the pelvic limb of the ostrich, 36 muscles can be determined. The ostrich lacks those muscles to the first and second toes (with exception of the M. flexor hallucis longus), which can be found in birds with four toes. The Mm. iliotrochantericus medius, plantaris, extensor proprius digiti IV and adductor digiti IV, which are present in other birds, are also absent, whereas the Mm. pectineus and femorotibialis accessorius additionally occur in the ostrich. The Pars supramedialis is a tendineous part of the M. gastrocnemius, on which the Mm. flexor cruris lateralis and flexor cruris medialis insert by means of a fascial sheet. The caudal part of the M. iliofibularis terminates within the caudal aspect of the superficial fascia cruris. The caudal heads of the Mm. flexor perforatus digiti III and flexor perforatus digiti IV as well as the M. flexor hallucis longus have a common origin on the Fossa poplitea of the femur. The lateral head of the M. flexor perforatus digiti IV and the femoral head of the M. flexor perforans et perforatus digiti III originate on the tendon of origin of the Caput laterale of the M. flexor perforatus digiti III. Furthermore, the last named tendon fuses with the tendon of insertion of the M. ambiens. The M. extensor proprius digiti III originates on a plate‐like fascial sheet part of the dorsal joint capsule of the intertarsal joint.

Nordic rattle: the hoarse vocalization and the inflatable laryngeal air sac of reindeer (Rangifer tarandus)

Laryngeal air sacs have evolved convergently in diverse mammalian lineages including insectivores, bats, rodents, pinnipeds, ungulates and primates, but their precise function has remained elusive. Among cervids, the vocal tract of reindeer has evolved an unpaired inflatable ventrorostral laryngeal air sac. This air sac is not present at birth but emerges during ontogenetic development. It protrudes from the laryngeal vestibulum via a short duct between the epiglottis and the thyroid cartilage. In the female the growth of the air sac stops at the age of 2–3 years, whereas in males it continues to grow up to the age of about 6 years, leading to a pronounced sexual dimorphism of the air sac. In adult females it is of moderate size (about 100 cm3), whereas in adult males it is large (3000–4000 cm3) and becomes asymmetric extending either to the left or to the right side of the neck. In both adult females and males the ventral air sac walls touch the integument. In the adult male the air sac is laterally covered by the mandibular portion of the sternocephalic muscle and the skin. Both sexes of reindeer have a double stylohyoid muscle and a thyroepiglottic muscle. Possibly these muscles assist in inflation of the air sac. Head‐and‐neck specimens were subjected to macroscopic anatomical dissection, computer tomographic analysis and skeletonization. In addition, isolated larynges were studied for comparison. Acoustic recordings were made during an autumn round‐up of semi‐domestic reindeer in Finland and in a small zoo herd. Male reindeer adopt a specific posture when emitting their serial hoarse rutting calls. Head and neck are kept low and the throat region is extended. In the ventral neck region, roughly corresponding to the position of the large air sac, there is a mane of longer hairs. Neck swelling and mane spreading during vocalization may act as an optical signal to other males and females. The air sac, as a side branch of the vocal tract, can be considered as an additional acoustic filter. Individual acoustic recognition may have been the primary function in the evolution of a size‐variable air sac, and this function is retained in mother–young communication. In males sexual selection seems to have favoured a considerable size increase of the air sac and a switch to call series instead of single calls. Vocalization became restricted to the rutting period serving the attraction of females. We propose two possibilities for the acoustic function of the air sac in vocalization that do not exclude each other. The first assumes a coupling between air sac and the environment, resulting in an acoustic output that is a combination of the vocal tract resonance frequencies emitted via mouth and nostrils and the resonance frequencies of the air sac transmitted via the neck skin. The second assumes a weak coupling so that resonance frequencies of the air sac are lost to surrounding tissues by dissipation. In this case the resonance frequencies of the air sac solely influence the signal that is further filtered by the remaining vocal tract. According to our results one acoustic effect of the air sac in adult reindeer might be to mask formants of the vocal tract proper. In other cervid species, however, formants of rutting calls convey essential information on the quality of the sender, related to its potential reproductive success, to conspecifics. Further studies are required to solve this inconsistency.

Glottal opening and closing events investigated by electroglottography and super-high-speed video recordings

Previous research has suggested that the peaks in the first derivative (dEGG) of the electroglottographic (EGG) signal are good approximate indicators of the events of glottal opening and closing. These findings were based on high-speed video (HSV) recordings with frame rates 10 times lower than the sampling frequencies of the corresponding EGG data. The present study attempts to corroborate these previous findings, utilizing super-HSV recordings. The HSV and EGG recordings (sampled at 27 and 44 kHz, respectively) of an excised canine larynx phonation were synchronized by an external TTL signal to within 0.037 ms. Data were analyzed by means of glottovibrograms, digital kymograms, the glottal area waveform and the vocal fold contact length (VFCL), a new parameter representing the time-varying degree of ‘zippering’ closure along the anterior–posterior (A–P) glottal axis. The temporal offsets between glottal events (depicted in the HSV recordings) and dEGG peaks in the opening and closing phase of glottal vibration ranged from 0.02 to 0.61 ms, amounting to 0.24–10.88% of the respective glottal cycle durations. All dEGG double peaks coincided with vibratory A–P phase differences. In two out of the three analyzed video sequences, peaks in the first derivative of the VFCL coincided with dEGG peaks, again co-occurring with A–P phase differences. The findings suggest that dEGG peaks do not always coincide with the events of glottal closure and initial opening. Vocal fold contacting and de-contacting do not occur at infinitesimally small instants of time, but extend over a certain interval, particularly under the influence of A–P phase differences.

The elephant knee joint: morphological and biomechanical considerations

Elephant limbs display unique morphological features which are related mainly to supporting the enormous body weight of the animal. In elephants, the knee joint plays important roles in weight bearing and locomotion, but anatomical data are sparse and lacking in functional analyses. In addition, the knee joint is affected frequently by arthrosis. Here we examined structures of the knee joint by means of standard anatomical techniques in eight African (Loxodonta africana) and three Asian elephants (Elephas maximus). Furthermore, we performed radiography in five African and two Asian elephants and magnetic resonance imaging (MRI) in one African elephant. Macerated bones of 11 individuals (four African, seven Asian elephants) were measured with a pair of callipers to give standardized measurements of the articular parts. In one Asian and three African elephants, kinematic and functional analyses were carried out using a digitizer and according to the helical axis concept. Some peculiarities of healthy and arthrotic knee joints of elephants were compared with human knees. In contrast to those of other quadruped mammals, the knee joint of elephants displays an extended resting position. The femorotibial joint of elephants shows a high grade of congruency and the menisci are extremely narrow and thin. The four‐bar mechanism of the cruciate ligaments exists also in the elephant. The main motion of the knee joint is extension–flexion with a range of motion of 142°. In elephants, arthrotic alterations of the knee joint can lead to injury or loss of the cranial (anterior) cruciate ligament.

Musculature of the crus and pes of the African elephant (Loxodonta africana): insight into semiplantigrade limb architecture

The limbs of elephants are designed to support the weight of the largest terrestrial animal, and they display unique morphological peculiarities among mammals. In this article we provide a new and detailed anatomical description of the muscles of the lower hindlimb in African elephants (Loxodonta africana), and we place our observations into a comparative anatomical as well as a functional morphological context. At the cranial aspect of the shank (crus) and the foot (pes), the flexors of the tarsal joint and the extensors of the toes form a flat muscular plate covering the skeletal elements. Caudal to the tibia and the fibula the Musculus (M.) soleus is strongly developed, whereas the M. gastrocnemius and the M. flexor digitorum superficialis are thin. Small flexors, adductors, and abductors of the toes are present. The M. tibialis caudalis as well as the Mm. fibularis longus and brevis mainly support the tarsal joint. The design of the muscular structures matches the specific requirements of heavy-weight bearing as well as of proboscidean limb posture and locomotion patterns.

Structure and innervation of the tusk pulp in the African elephant (Loxodonta africana)

African elephants (Loxodonta africana) use their tusks for digging, carrying and behavioural display. Their healing ability following traumatic injury is enormous. Pain experience caused by dentin or pulp damage of tusks seems to be negligible in elephants. In this study we examined the pulp tissue and the nerve distribution using histology, electron microscopy and immunhistochemistry. The results demonstrate that the pulp comprises two differently structured regions. Randomly orientated collagen fibres characterize a cone‐like part lying rostral to the foramen apicis dentis. Numerous nerve fibres and Ruffini endings are found within this cone. Rostral to the cone, delicate collagen fibres and large vessels are orientated longitudinally. The rostral two‐thirds of the pulp are highly vascularized, whereas nerve fibres are sparse. Vessel and nerve fibre distribution and the structure of connective tissue possibly play important roles in healing and in the obviously limited pain experience after tusk injuries and pulp alteration. The presence of Ruffini endings is most likely related to the use of tusks as tools.

Occurrence and structure of epipharyngeal pouches in bears (Ursidae)

The infrequent mention of epipharyngeal pouches occurring in some species of bears indicates the scarcity of morphological and functional knowledge about these structures. In order to provide precise morphological data on the structure of these remarkable formations and to verify their taxonomic utility, the pharyngeal regions of 1 spectacled bear and 3 brown bears were examined. All these individuals possessed epipharyngeal pouches, which are tubular, blind‐ending outpouchings of the caudodorsal pharyngeal wall equipped with respiratory epithelium and a thick layer of elastic fibres. While the spectacled bear and Ursus arctos syriacus possessed a single pouch on the caudodorsal wall of the nasopharynx, in Ursus arctos and Ursus arctos beringianus 2 unequally sized pouches were present. Two additional sacs of smaller size, representing outpouchings of the lateral pharyngeal wall, occurred in the spectacled bear. These findings prove epipharyngeal pouches to be constant and unique morphological features of the family Ursidae, the anatomical features suggesting involvement in the respiratory system most probably in important aspects of ursid phonation. This is the first description of epipharyngeal pouches in the spectacled bear.

Gross Anatomy of the Female Genital Organs of the Domestic Donkey (Equus asinus Linné, 1758)

Although donkeys play an important role as companion or pack and draught animals, theriogenological studies and anatomical data on the genital organs of the jenny are sparse. To provide anatomical descriptions and morphometric data, the organa genitalia feminina, their arteries and the ligamentum latum uteri of 10 adult but maiden jennies were examined by means of gross anatomical and morphometric techniques. In comparison with anatomical data of horses obtained from literature the genital organs of jennies appear to be more voluminous in relation to the body mass and the position of the ovaries is slightly further cranial than in mares. In asses, the ovaries contain large follicles reaching a diameter of up to 40 mm. The mesosalpinx is much wider than in the horse forming a considerably spacious bursa ovarica. The asinine ligamentum teres uteri reveals a very prominent cranial end, the ‘appendix’. Tortuous mucosal folds occur in the wall of the jenny’s cervical channel. The vascularization of the female genital organs of asses is very similar to that of horses. One of the examined specimens reveals a large mucosal fold dividing the cranial part of the vagina into a left and right compartment.