Descended and mobile larynx, vocal tract elongation and rutting roars in male goitred gazelles (Gazella subgutturosa Güldenstaedt, 1780)

Similar to male humans, Homo sapiens, the males of a few polygynous ruminants - red deer Cervus elaphus, fallow deer Dama dama and Mongolian gazelle Procapra gutturosa- have a more or less enlarged, low-resting larynx and are capable of additional dynamic vocal tract elongation by larynx retraction during their rutting calls. The vocal correlates of a large larynx and an elongated vocal tract, a low fundamental frequency and low vocal tract resonance frequencies, deter rival males and attract receptive females. The males of the polygynous goitred gazelle, Gazella subgutturosa, provide another, independently evolved, example of an enlarged and low-resting larynx of high mobility. Relevant aspects of the rutting behaviour of territorial wild male goitred gazelles are described. Video and audio recordings served to study the acoustic effects of the enlarged larynx and vocal tract elongation on male rutting calls. Three call types were discriminated: roars, growls and grunts. In addition, the adult male vocal anatomy during the emission of rutting calls is described and functionally discussed using a 2D-model of larynx retraction. The combined morphological, behavioural and acoustic data are discussed in relation to the hypothesis of sexual selection for male-specific deep voices, resulting in convergent features of vocal anatomy in a few polygynous ruminants and in human males.     

Savannah roars: The vocal anatomy and the impressive rutting calls of male impala (Aepyceros melampus) – highlighting the acoustic correlates of a mobile larynx

A retractable larynx and adaptations of the vocal folds in the males of several polygynous ruminants serve for the production of rutting calls that acoustically announce larger than actual body size to both rival males and potential female mates. Here, such features of the vocal tract and of the sound source are documented in another species. We investigated the vocal anatomy and laryngeal mobility including its acoustical effects during the rutting vocal display of free-ranging male impala (Aepyceros melampus melampus) in Namibia. Male impala produced bouts of rutting calls (consisting of oral roars and interspersed explosive nasal snorts) in a low-stretch posture while guarding a rutting territory or harem. For the duration of the roars, male impala retracted the larynx from its high resting position to a low mid-neck position involving an extensible pharynx and a resilient connection between the hyoid apparatus and the larynx. Maximal larynx retraction was 108 mm based on estimates in video single frames. This was in good concordance with 91-mm vocal tract elongation calculated on the basis of differences in formant dispersion between roar portions produced with the larynx still ascended and those produced with maximally retracted larynx. Judged by their morphological traits, the larynx-retracting muscles of male impala are homologous to those of other larynx-retracting ruminants. In contrast, the large and massive vocal keels are evolutionary novelties arising by fusion and linear arrangement of the arytenoid cartilage and the canonical vocal fold. These bulky and histologically complex vocal keels produced a low fundamental frequency of 50 Hz. Impala is another ruminant species in which the males are capable of larynx retraction. In addition, male impala vocal folds are spectacularly specialized compared with domestic bovids, allowing the production of impressive, low-frequency roaring vocalizations as a significant part of their rutting behaviour. Our study expands knowledge on the evolutionary variation of vocal fold morphology in mammals, suggesting that the structure of the mammalian sound source is not always human-like and should be considered in acoustic analysis and modelling.     

Vocal anatomy, tongue protrusion behaviour and the acoustics of rutting roars in free-ranging Iberian red deer stags (Cervus elaphus hispanicus)

Roaring in rutting Iberian red deer stags Cervus elaphus hispanicus is unusual compared to other subspecies of red deer, which radiated from the Iberian refugium after the last glacial maximum. In all red deer stags, the larynx occupies a permanent low mid‐neck resting position and is momentarily retracted almost down to the rostral end of the sternum during the production of rutting calls. Simultaneous with the retraction of the larynx, male Iberian red deer pronouncedly protrude the tongue during most of their rutting roars. This poses a mechanical challenge for the vocal tract (vt) and for the hyoid apparatus, as tongue and larynx are strongly pulled in opposite directions. This study (i) examines the vocal anatomy and the acoustics of the rutting roars in free‐ranging male C. e. hispanicus; (ii) establishes a potential mechanism of simultaneous tongue protrusion and larynx retraction by applying a two‐dimensional model based on graphic reconstructions in single video frames of unrestrained animals; and (iii) advances a hypothesis of evaporative cooling by tongue protrusion in the males of a subspecies of red deer constrained to perform all of the exhausting rutting activities, including acoustic display, in a hot and arid season.     

A nose that roars: anatomical specializations and behavioural features of rutting male saiga

The involvement of the unique saiga nose in vocal production has been neglected so far. Rutting male saigas produce loud nasal roars. Prior to roaring, they tense and extend their noses in a highly stereotypic manner. This change of nose configuration includes dorsal folding and convex curving of the nasal vestibulum and is maintained until the roar ends. Red and fallow deer males that orally roar achieve a temporary increase of vocal tract length (vtl) by larynx retraction. Saiga males attain a similar effect by pulling their flexible nasal vestibulum rostrally, allowing for a temporary elongation of the nasal vocal tract by about 20%. Decrease of formant frequencies and formant dispersion, as acoustic effects of an increase of vtl, are assumed to convey important information on the quality of a dominant male to conspecifics, e.g. on body size and fighting ability. Nasal roaring in saiga may equally serve to deter rival males and to attract females. Anatomical constraints might have set a limit to the rostral pulling of the nasal vestibulum. It seems likely that the sexual dimorphism of the saiga nose was induced by sexual selection. Adult males of many mammalian species, after sniffing or licking female urine or genital secretions, raise their head and strongly retract their upper lip and small nasal vestibulum while inhalating orally. This flehmen behaviour is assumed to promote transport of non-volatile substances via the incisive ducts into the vomeronasal organs for pheromone detection. The flehmen aspect in saiga involves the extensive flexible walls of the greatly enlarged nasal vestibulum and is characterized by a distinctly concave configuration of the nose region, the reverse of that observed in nasal roaring. A step-by-step model for the gradual evolution of the saiga nose is presented here.     

Cytogenetic comparison between Vietnamese sika deer and cattle: R-banded karyotypes and FISH mapping

R-banded chromosomes of Vietnamese sika (VS) deer (Cervus nippon pseudaxis, 2N = 66), a threatened subspecies of sika deer endemic to Vietnam, are presented for the first time and were compared with bovine R-banded chromosomes to define a standard karyotype. Nineteen VS deer autosomes (CNP) were identified on the basis of the banding pattern relative to bovine chromosomes (BTA), while hypotheses for the remaining thirteen were proposed from comparisons with the published deer genetic map, BTA 1, 2, 5, 6, 8 and 9 each equivalent to two separate acrocentric CNP chromosomes and BTA 26 and 28 associated in a tandem fusion. To confirm these hypotheses, probes for the twenty-nine Texas nomenclature type I markers specific for each cattle autosome, sixteen other type I and fourteen microsatellite markers were used in FISH experiments on VS deer chromosomes. CNP7 presented the most complex rearrangement as compared with cattle chromosomes. A complete correspondence between VS deer and cattle chromosomes was established and it was extended with a comparison with the human karyotype to transfer human map information to this species of scientific and economic interest. Moreover, this work anchors the deer genetic linkage map to chromosome-specific markers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Acoustical and Anatomical Determination of Sound Production and Transmission in West Indian (Trichechus manatus) and Amazonian (T. inunguis) Manatees

West Indian (Trichechus manatus) and Amazonian (T. inunguis) manatees are vocal mammals, with most sounds produced for communication between mothers and calves. While their hearing and vocalizations have been well studied, the actual mechanism of sound production is unknown. Acoustical recordings and anatomical examination were used to determine the source of sound generation. Recordings were performed on live captive manatees from Puerto Rico, Cuba and Colombia (T. manatus) and from Peru (T. inunguis) to determine focal points of sound production. The manatees were recorded using two directional hydrophones placed on the throat and nasal region and an Edirol‐R44 digital recorder. The average sound intensity level was analyzed to evaluate the sound source with a T test: paired two sample for means. Anatomical examinations were conducted on six T. manatus carcasses from Florida and Puerto Rico. During necropsies, the larynx, trachea, and nasal areas were dissected, with particular focus on identifying musculature and soft tissues capable of vibrating or constricting the airway. From the recordings we found that the acoustical intensity was significant (P < 0.0001) for both the individuals and the pooled manatees in the ventral throat region compared to the nasal region. From the dissection we found two raised areas of tissue in the lateral walls of the manatee's laryngeal lumen that are consistent with mammalian vocal folds. They oppose each other and may be able to regulate airflow between them when they are adducted or abducted by muscular control of arytenoid cartilages. Acoustic and anatomical evidence taken together suggest vocal folds as the mechanism for sound production in manatees. Anat Rec, 297:1896–1907, 2014. © 2014 Wiley Periodicals, Inc.     

Blood biochemistry values in some wild ruminants in captivity

Analyses of blood chemical parameters were carried out in eight axis deer (Cervus axis), 12 fallow deer (Cervus dama), 16 red deer (Cervus elaphus hippelaphus), three sambar (Cervus unicolor), nine Père David deer (Elaphurus davidianus), 20 European bison (Bison bonasus), seven nilgai (Boselaphus tragocamelus), eight mouflon (Ovis musimon), four white-bearded gnu (Connochaetes taurinus) and six barbary sheep (Ammotragus lervia). The following parameters were determined: glucose, urea, uric acid, cholesterol, triglycerides, creatinine, total bilirubin, aspartate aminotransferase, alanine aminotransferase, alkaline phosphatase, creatinine phosphokinase, lactate dehydrogenase, γ-glutamyl transpeptidase, total serum proteins, albumin, globulins, albumin/globulin ratio, sodium, potassium, chloride, calcium, magnesium, total phosphorus and serum osmolality. Some trends and statistically significant differences were observed between the species and the young and adult groups.     

Identification by R-banding and FISH of chromosome arms involved in Robertsonian translocations in several deer species

We constructed and analyzed the RBG-banded karyotype of five deer species: Chital (Axis axis), White-lipped deer (Cervus albirostris), Rusa deer (Cervus timorensis russa), Sambar deer (Cervus unicolor) and Eld's deer (Cervus eldi siamensis). Among these five species, only Eld's deer had been previously karyotyped using R-banding. In order to identify all the chromosome correspondences with cattle and precisely which chromosome arms are involved in Robertsonian translocations, we compared the karyotypes of these five species with those of the closely related and well-characterized species, cattle (Bos taurus) and Vietnamese Sika deer (Cervus nippon pseudaxis). Among these six deer species (the five above plus the Vietnamese Sika deer), we found thirteen different Robertsonian translocations involving nineteen different chromosome arms. Thirteen chromosome arms were identified by comparison of R-banding patterns only and the remaining six were either confirmed or identified by FISH-mapping of bovine or caprine probes previously localized in cattle. Finally, we observed that five of the thirteen Robertsonian translocations are shared by at least two species and that some chromosome arms are more frequently involved in Robertsonian translocations than others. This revised version was published online in July 2006 with corrections to the Cover Date.     

Some Haematological Values of Farmed Fallow Deer (>Dama dama)

Haematological examinations have been carried out on 62 healthy farmed male and female fallow deer after they had been shot during a routine cull. Comparisons were made between the sexes, between young (up to 2 years) and older (>2 years) animals, and between age groups according to sex. The following parameters (means and SDs) were examined on all samples: erythrocyte count, haemoglobin concentration, packed cell volume, mean cell volume, mean cell haemoglobin, mean cell haemoglobin concentration, total leucocyte count, and numbers of neutrophils, eosinophils, basophils, lymphocytes, band neutrophils and monocytes. Only minor differences in red cell and white cell parameters between different groups of animals were found. However, significant differences in mean cell volume, mean cell haemoglobin concentration, eosinophils and band neutrophils were discovered in different age and sex groups. The values provide useful information for diagnostic purposes in fallow deer. Correspondence and offprint requests to: G. Vengušt, Institute for Breeding and Health Care of Wild Animals, Fishes and Bees, University of Ljubljana, Veterinary Faculty, Gerbičeva 60, 1000 Ljubljana, Slovenia. e-mail: vengusgo@vf.uni-lj.si     

Zur Wirtsspezifität von Babesia motasi und Babesia ovis (Piroplasmidea)

Zusammenfassung 1. Babesia motasi wurde auf ein Hausrind, eine Ziege, einen Mufflon, einen Rothirsch, zwei Damhirsche, ein Reh und eine Sömmering-Gazelle übertragen. Mit Ausnahme des Mufflons und eines Damhirsches waren die Tiere entmilzt. Als empfänglich erwiesen sich Ziege, Mufflon, Rothirsch und beide Damhirsche.2. Babesia ovis wurde auf zwei Damhirsche, einen Rothirsch und eine Sömmering-Gazelle übertragen. Die Tiere waren bis auf einen Damhirsch entmilzt. Es kam nur zu kurzen, wahrscheinlich latenten Infektionen bei den Damhirschen und dem Rothirsch. Die Sömmering-Gazelle war unempfänglich für B. ovis.

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Summary 1. Babesia motasi has been inoculated by intravenous injection of infected sheep-blood into a calf (Bos taurus L.), a goat (Capra hircus L.), a moufflon (Ovis aries musimon Pallas), a red deer (Cervus elaphus L.), two fallow deer (Dama dama L.), a roe deer (Capreolus capreolus) and a Soemmering's gazelle (Gazella soemmeringi). All of these ruminants except the moufflon and one of the fallow deer had been splenectomized previously. The goat, the moufflon, the red deer and the two fallow deer proved to be susceptible for B. motasi.2. B. ovis has been inoculated by intravenous injection of sheep-blood into a red deer, a fallow deer and a Soemmering's gazelle and by feeding infected ticks (Rhipicephalus bursa) on another fallow deer. All of these ruminants except the fallow deer which has been infected with infected ticks had been splenectomized previously. A slight parasitemia of short duration could be detected only in the red deer and in the two fallow deer.

     

Serum Biochemistry of Persian Fallow Deer (Dama Mesopotamica)

In order to study serum biochemistry of Persian fallow deer (Dama mesopotamica), sampling was conducted from eight clinicaly normal Persian fallow deer (three females and five males) aged between 6 months and 10 years and the following results were obtained: total protein 68.4 t 5 g/l; albumin 28.1 t 8.9 g/l; globulin 42.7 t 7.5 g/l; A: G 0.63 t 0.25; glucose 6.61 t 3.41 mmol/l; triglyceride 0.37 t 0.22 mmol/l; cholesterol 2.86 t 1.46 mmol/l; blood urea nitrogen 8.91 t 1.21 mmol/l; creatinin 146.75 t 53.92 mmol/l; total bilirubin 12.99 t 2.05 mmol/l; direct bilirubin 7.01 t 1.88 mmol/l; aspartate aminotransferase (AST) 20.25 t 7.36 IU/l; alanine aminotransferase (ALT) 26.38 t 8.94 IU/l; alkaline phosphatase (AP) 118.25 t 70.79 IU/l; creatine kinase (CK) 219.13 t 151.81 IU/l; Na 147.5 t 9.09 mmol/l; K 6.82 t 1.92 mmol/l; Cl 103.88 t 5.44 mmol/l; Ca 2.79 t 0.43 mmol/l; P 1.99 t0.47 mmol/l; Mg 1.12 t 0.23 mmol/l. Significant differences were seen for Na and P between age groups, with higher levels in deer aged less than 2 years (p<0.05). Glucose had higher values in does than in stags (p<0.05). Correlations between measured parameters were also determined.     

The first detection of nematodes Ashworthius sidemi in elk Alces alces (L.) in Poland and remarks of ashworthiosis foci limitations

Ashworthius sidemi, a nematode belonging to the family of Trichostrongylidae, is a primary parasite of the Asian deer, mainly sika deer (Cervus nippon), with which it was introduced to Ukraine, as well as Slovakia, the Czech Republic and France. Migrating red deer carried this parasite from neighboring countries to Polish territory. Until now, in Poland, this parasite has been recorded in European bison, red deer, roe deer and fallow deer. As a result of post-mortem examinations of 10 elk, 2 of them from the Augustów Forest and Biebrza Marshes, A. sidemi were found in abomasa for the first time in Poland. The intensity of the invasions was 120 and 7 specimens, respectively. This finding of Ashworthius sidemi in elk indicates a further expansion of the focus of ashworthiosis in Bia?owie?a towards the north into the Biebrza Marshes and the Augustowska Forest. The growth of the elk population and their tendency for long distance migrations can contribute to the spread of the parasitosis in much greater distances than deer. On the basis of our own research and data from the literature, the current spread of ashworthiosis in Poland is discussed.     

Selection on vocal output affects laryngeal morphology in rats

Although laryngeal morphology often reflects adaptations for vocalization, the structural consequences of selection for particular aspects of vocal behavior remain poorly understood. In this study, we investigated the effects of increased ultrasonic calling in pups on the adult larynx morphology in selectively bred rat lines. Laryngeal morphology was assessed using multiple techniques: mineralized cartilage volumes were compared in 3D-models derived from microCT scans, internal structure was compared using clearing and staining procedures combined with microscopy, cellular structure was compared using histology and microscopy, and element composition was assessed with scanning energy dispersive X-ray spectroscopy. Our results show that adult rats from lines bred to produce ultrasonic calls at higher rates as pups have shorter vocal folds and a more mineralized thyroid cartilage compared to rats bred to produce ultrasonic calls at lower rates. The change in vocal fold length appears to account for differences in low-frequency calls in these two rat lines. We suggest that the observed increases in mineralization of the thyroid cartilage in the high-ultrasound lineage provide increased reinforcement of the laryngeal structure during ultrasonic call production. Our findings therefore demonstrate an effect of selection for vocal behavior on laryngeal morphology, with acoustic consequences.     

Status and origin of Haemonchinae (Nematoda: Trichostrongylidae) in deer: a survey conducted in France from 1985 to 1998

During our investigations on helminthofauna in roe deer (Capreolus capreolus) and red deer (Cervus elaphus) in France (1985–1998) we isolated nematodes not only of the genus Haemonchus Cobb, 1898 but also of the genus Ashworthius Le Roux 1930, both of which belong to the same subfamily of Haemonchinae. The prevalence of Ashworthius was 22% (65/294) in roe deer and 40% (10/25) in red deer. H. contortus was not found in red deer, whereas its prevalence was only 3% (9/294) in roe deer. These data contrast with those observed in other European countries. The presence of A. sidemi in red and roe deers in France suggests a case of parasitism imported via sika deer. A hypothesis of their spread is proposed. Received: 19 November 1999 / Accepted: 23 December 1999     

A Computational Study of the Effect of False Vocal Folds on Glottal Flow and Vocal Fold Vibration During Phonation

The false vocal folds are believed to be components of the acoustic filter that is responsible for shaping the voice. However, the effects of false vocal folds on the vocal fold vibration and the glottal aerodynamic during phonation remain unclear. This effect has implications for computational modeling of phonation as well as for understanding laryngeal pathologies such as glottal incompetence resulting from unilateral vocal fold paralysis. In this study, a high fidelity, two-dimensional computational model, which combines an immersed boundary method for the airflow and a continuum, finite-element method for the vocal folds, is used to examine the effect of the false vocal folds on flow-induced vibration (FIV) of the true vocal folds and the dynamics of the glottal jet. The model is notionally based on a laryngeal CT scan and employs realistic flow conditions and tissue properties. Results show that the false vocal folds potentially have a significant impact on phonation. The false vocal folds reduce the glottal flow impedance and increase the amplitude as well as the mean glottal jet velocity. The false vocal folds also enhance the intensity of the monopole acoustic sources in the glottis. A mechanism for reduction in flow impedance due to the false vocal folds is proposed.     

活体犬声带控制振动的声门图研究——Ⅱ、振动模式与声门图特征变化

本文以定义并由多功能声门图仪提取的一组特征参数为基础,对声门气流、喉上神经与喉返神经电刺激变化,活体犬声带控制振动模式的影响进行了定量研究。实验结果表明:(1)对相同发声方式,犬喉与人体声带振动模式是类似的,但也因声带尺寸与组织学上的差异而引起特征参数差别;(2)采用多功能声门图仪与活体犬模型,能对声带振动进行深入的研究;(3)本文的研究方法和结果为喉病临床诊断,发展新方法提供重要的基础。     

The giant liver fluke <Emphasis Type="Italic">Fascioloides magna</Emphasis> (Bassi 1875) in cervids in the Czech Republic and potential of its spreading to Germany

The giant liver fluke Fascioloides magna is an important parasite of cervids in Europe. From September 2003 to December 2005, faecal samples and livers of red deer (Cervus elaphus) and fallow deer (Dama dama) were investigated to determine the current distribution of the fluke in the Czech Republic. Faecal samples were collected from 20 different areas, and livers of hunted deer were dissected from each locality to confirm F. magna infection. The prevalence of F. magna in examined areas determined by coprological examination varied from 4% to 95%. Moreover, new foci of F. magna infection were discovered in all localities in the Šumava mountains where F. magna was observed; this has epizootiological importance due to the possibility of the spread of F. magna into the German territory (Bavaria).     

A morphological study of the human prepubertal and pubertal larynx

Measurements were made from ten Caucasian male and ten Caucasian female cadaveric larynges ranging in age from 9 to 18 years. The sample was divided into prepubertal and pubertal groups of each sex, respectively. Each specimen was dissected and measured according to a specified protocol so that data on linear and angular dimensions of the laryngeal cartilages and the weight of each cartilage were obtained. Results from this study highlighted differences in the developmental morphology of the circumpubertal larynx. With few exceptions, pubertal laryngeal measurements were significantly larger than prepubertal counterparts in both sexes. No clear sexual dimorphism was found between male and female prepubertal larynges though measurements of the prepubertal female larynx were closer to adult counterparts in size and weight than were those in the male. Thus the female larynx requires less growth per unit time to reach maturity than the male. By puberty, clear sexual dimorphism was evident in the larynx. Linear and weight measurements of the pubertal male larynx were significantly larger than in the female. Though the angle of the thyroid laminae was not significantly different in pubertal male and female cartilages, the thyroid eminence was clearly more prominent in the male. The vocal folds in both sexes reached essentially their adult length by puberty; however, the absolute length of the male vocal folds had increased by over two times that of the female.     

Identity of rumen fluke in deer

As evidence is growing that in many temperate areas paramphistome infections are becoming more common and widespread, this study was undertaken to determine the role of deer as reservoirs for rumen fluke infections in livestock. A total of 144 deer faecal samples (88 from fallow deer, 32 from red deer and 24 samples from sika, sika/red deer hybrids) were screened for the presence of fluke eggs. Based on the ITS-2 rDNA locus plus flanking 5.8S and 28S sequences (ITS-2+), fluke eggs were identified to species level. Our results indicate that, of the 3 deer species, fallow deer had the highest fluke infection rates. Two rumen fluke species, Calicophoron daubneyi and Paramphistomum leydeni, with morphologically distinct eggs, were identified. Concurrent infections of the two paramphistome species and liver fluke, Fasciola hepatica, were common. Considering the comparatively low egg burdens observed in this study, it is unlikely that deer represent a significant source of infection for Irish livestock.