14) Immunology
No studies are known to us, other than serologic surveys. Thus, Anderson & Rowe (1998) found antibodies to Rift Valley Fever (RVF) virus in black rhinos of Zimbabwe . Fischer-Tenhagen et al. (2000) found antibodies to a variety of other viral agents, but not against RVF virus in 281 animals from South Africa , Kenya and Namibia.
15) Pathological features
A major problem for black rhinoceroses is the propensity to develop hemolytic anemia (Chaplin, et al., 1986; Miller & Boever, 1982; Paglia et al., 1986; Paglia, 1993). The major enzymatic causes of this phenomenon were well described by Paglia (1993), and hematologic parameters studied by Chaplin et al. (1986). Veterinary problems, diseases and numerous procedures, were summarized by Silberman & Fulton (1979), and a complete veterinary bibliography was compiled by Miller (1983). It includes references to reproductive laboratory data as well.
Dermal problems and infections appear to be the commonest ailments, including pox (Schaller & Pilaski, 1979; Griner, 1983). Skin ulcers, studied by Kock & Kock (1990) in free-living animals from Zimbabwe were mostly or only due to filarial nematodes ( Stephanofilaria dinniki ). Other parasites have been described; thus, Keirans (1993) found Ixodid ticks in white and black rhinos; babesiasis was reported in three animals from Tanzania (Nijhof et al., 2003); trypanosomiasis was reported by Mihok et al. (1992) and Knapp et al. (1997) collected various arthropods and helminths from black rhinos. Neiffer et al. (2001) described leptospirosis in two black rhinos from Pittsburgh Zoo, presumably acquired from raccoons. Salmonellosis killed two rhinos in Denver (Kenny et al., 1997) and a survey of US zoos (Kenny, 1999) indicated an approximately 10% infection rate.
Three captive black rhinos showed severe encephalomalacia at death whose etiology remains unknown (Miller et al., 1990). Jones (1979) saw ulcerative stomatitis in black rhinos, a lesion later also described by Ott et al. (1982). Pessier et al. (2004) later associated this to eosinophilic granulomas occurring in rhinos but distinguished it from the necrolytic dermatitis described by Munson et al. (1998). Such skin lesions led Grant et al. (2002) to study fatty acids of the foods eaten; diets in the wild were significantly different in their composition. Following prolonged antibiotic therapy of hoof abscesses, a black rhino died from pulmonary aspergillosis (Woods et al., 1999). Acute lymphoblastic leukemia in a 21 months-old animal was the cause of death, perhaps also related to cardiotoxicity of the therapeutic agents (Radcliffe et al., 2000; Paglia & Radcliffe, 2000). That case study suggested excess iron in black rhinos, a notion that was more recently confirmed by additional studies (Paglia et al., 2001). These studies suggested greater prevalence of iron storage in females and suggested a link to the encephalopathy. Stegmann et al. (2001) described the fascinating story of anesthesia and operation of a necrotic rectal prolapse in a free-ranging black rhino. The animal died and was then found to have old pelvic fractures that may have been contributory (Olivier et al., 2001).
16) Physiologic data
Whatever relevant physiologic data are available have been provided in review form by Silberman & Fulton (1979), and as bibliography by Miller (1983). Schaurte (1966) described in detail the various dental formulas and depicts teeth of all rhinocerotidae. Schaffer et al. (1998) studied the reasons for the difficulty of obtaining semen by using sonography. They demonstrated the urethral filling in conditioned animals and the benefits of sonographic usage. O'Brien & Roth (2000) provided a protocol for the conservation and recovery of semen from Sumatran and black rhinos. Meiswinkel (1987) described mosquitoes grown from elephant and rhino dung. Schaffer et al. (2001) found the anatomy and histology of reproductive tracts similar to other species. Maluf (91991; 1994) described the anatomy of the kidney with its 60 lobes. Dierenfeld et al. (1988) found that plasma vitamin E levels of 31 wild animals were significantly higher than in eleven animals of the captive population. In a subsequent study, these results were not verified, presumably because of vitamin supplementation in captive animals (Clauss et al., 2002). Hematologic parameters, especially following the frequent translocations occurring in Africa, were delineated by Kock et al. (1999). The episodic occurrences of hemolysis have led to study particulars of enzymes etc. in rhino red cells. Thus, Weber et al. (2004) found significantly higher tyrosine levels in wild animals compared to captive specimens. For the same reasons, the hemoglobins were studied in several animals and related species (Fairbanks & Miller, 1990). They found considerable polymorphism of black rhino hemoglobins and proposed a genetic mechanism for these proteins.
17) Other resources
Cell strains of four species of rhinoceros are available from the “Frozen zoo” at the Zoological Society of San Diego by contacting Dr. Oliver Ryder at: oryder@ucsd.edu . None are available of the severely endangered Javan rhinoceros. An International Rhino Conference was held at the San Diego Zoo in 1991. It reviews many topics; a copy of the review of this conference can also be obtained from Dr. O. Ryder.
18) Other remarks – What additional Information is needed?
There are no studies on early implantation; information on the structure and length of umbilical cord is needed.
Acknowledgement
The animal photograph in this chapter comes from the Zoological Society of San Diego. The B&W photographs come from R. Montali at the National Zoo, Washington, DC.
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One female D. bicomis individual, (#360), was found to have a diploid number of 2n 85 because of an additional large submetacentric chromosome. Analysis of the 0-bands revealed a trisomy X chromosomal complement. Presumably the additional X did not have any deleterious effects because of X inactivation (LYON , 1962). This animal died at age 2.5 years. Autopsy findings gave no indications of anomalies or untoward effects of trisomy X. Diffuse encephalopathy was deemed to be the result of an unknown toxin or nutritional deficiency (Eric Miller, personal communication). The dam, (#267), of this individual was sampled and revealed a normal 2n 84 karyotype. The sire was not sampled directly, but karyotype analyses of his paternal grandparents, (#124 and #125), revealed normal chromosomal complements.
Two other female D. bicomis individuals, that were wild-caught and presumably unrelated, had normal diploid numbers, but showed very large heterochromatic areas in the p-arm of one chromosome. In one animal, (#293), the additional heterochromatin resulted in a submetacentric element similar in size to the X chromosomes. Due to the quality of the G-banding it was not possible to determine which chromosome was affected. In the other animal, (#188), a metacentric chromosome was created by the addition of a large heterochromatic arm. Q-banding showed a metacentric chromosome in which the q-arm paired with that of a subtelocentric autosome.
This study illustrates the importance of chromosomal analyses in support of animal health and conservation management programs. Further comparative studies need to be conducted on chromosomal polymorphisms which may distinguish different geographic populations of D. bicomis . In addition, the numerical polymorphism noted in the critically endangered northern white rhinoceros ( C. s. cottoni ) should be monitored through additional study of captive specimens and, where feasible, of wild populations.
Acknowledgements
The authors wish to thank Ken Kelley, Grace Magee, Suellen Charter and Reneê Cabrera for their assistance and cooperation. Appreciation is extended to Steve Kingswood, Dr. Valentine L. Vance and Dr. E. Ann 0akenfull for their critical review of the manuscript and to Dr. Mike H. Jurke for translation of the summary. Clonetics Corporation generously provided the necessary growth medium for the fibroblast cultures.
Summary
Cytogenetics of the Rhinocerotidae
Cytogenetic studies were conducted on 113 rhinoceroses, representing four of the five extant species. Karyotype analyses revealed a modal diploid number of 82 chromosomes for Ceratotherium simum, Rhinoceros unicomis, Dicerorhinus sumatrensis and 84 chromosomes for Diceros bicomis . Comparison of the autosomes of the four species revealed differences in centromere location, ranging from all acrocentric elements in D. sumatrensis to mostly submetacentric elements in D. bicomis. The X chromosome was identified as a large submetacentric element in all four species. The X chromosome of D. sumatrensis was distinct from the other three species because of an addition of heterochromatic material in the long arm.
Abnormal chromosomal complements were identified in three C. simum and three D. bicomis including one trisomy X individual. The diploid number of D. sumatrensis is presented for the first time, as well as the first banding studies of D. sumatrensis, R. unicornis and D. bicornis.
Zusammenfassung CYTOGENETIK DER RHINOCEROTIDAE
Wir berichten über cytogenetische Untersuchungen an 113 Nashömem von vier der fünf bestehenden Nashomarten. Die Chromosomenzahl (“diploide Zahl" “2n”) beträgt 82 Chromosomen in den folgenden drei Arten: Ceratotherium simum, Rhinoceros unicornis und Dicerorhinus sumatrensis. Hingegen hat Diceros bicomis 84 Chromosomen. Wenn man die Autosomen dieser Arten vergleicht, so findet man eine unterschiedliche Lokalisation der Zentromeren, von ausschliesslich akrozentrischen Autosomen in D. sumatrensis zu hauptsächlich submetazentrischen E!ementen im D. bicomis. Das X-Chromosom ist ein grosses submetazentrisches Element in allen Arten. Das X-Chromosom von D. sumatrensis unterscheidet sich allerdings von den anderen Arten durch ein zusätzliches Stück von Heterochromatin im langen Arm. Chromosomenabnormalitäten wurden in je drei Nashömem von D. simum und von D. bicornis gefunden, einschliesslich einer Trisomie des X-Chromosoms. Dieser Bericht stellt die erste Beschreibung der diploiden Chromosomenzahl des D. sumatrensis dar; ebenso präsentieren wir die ersten gebandeten Chromosomen von D. sumatrensis, R. unicomis und von D. bicomis.
Résumé
La cytogénetique des rhinocérotidae
Nous avons effectué des examens cytogénétiques sur 113 rhinoceros ayant represente quatre des cinq espèces existantes. Le nombre de chromosomes (“chiffre diploide" "2n”) se chiffre a 82 chromosomes pour les trois espèces qui suivent: Ceratotherium simum, Rhinoceros unicornis et Dicerorhinus sumatrensis . Diceros bicomis par contre, a 84 XX chromosomes. La comparaison des autosomes de ces espèces révèle une localisation differente des centromères , a partir d'autosomes exclusivement acrocentriques dans l'espèce D. sumatrensis jusqu'aux elements essentiellement submétacentriques dans D. bicomis . Le chromosome X est un grand élément submetacentrique dans toutes les espèces. Cependant, le chromosome X de D. sumatrensis se distingue des autres espèces par un élement additionnel d'héterochromatine au bras long. Nous avons observe des anomalies de chromosomes dans trois specimens de C. simum et dans trois de D. bicornis ainsi qu'une trisomie du chromosome X. Ce rapport présente pour Ia premiere fois une description du chiffre diploide des chromosomes de D. sumatrensis et fait etat des bandes chromosomales de D. sumatrensis, de R. unicomis et de D. bicomis.
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Address of authors: Marlys L. Houck, Center for Reproduction of Endangered Species, Zoological Society of San Diego , P. 0. Box 551 , San Diego , C.A. 92112-0551 (USA).
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