246 Appendix 40 FMD and Camelids
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Appendix 40 FMD and camelids: International relevance of current research U. Wernery Central Veterinary Research Laboratory, P.O. Box 597, Dubai, U.A.E. Key words: Tylopoda, camelids, FMD Abstract Camelids regurgitate and re-chew their food and thus technically ruminate. In strict taxonomic terms, however, they are not recognized as belonging to the suborder Ruminantia. They belong to the suborder Tylopoda. Numerous differences in anatomy and physiology justify a separate classification of tylopods from ruminants. Many reports show that New World Camelids (NWC) and Old World Camelids (OWC) possess a low susceptibility to foot and mouth disease (FMD), and do not appear to be long-term carriers of the foot and mouth disease virus (FMDV). Recent preliminary results from Dubai have shown that two dromedaries infected subepidermolingually with FMD serotype 0 did not develop any clinical signs and failed to develop any lesions at the inoculation site. Infectious FMDV or FMDV RNA were not isolated and the two dromedaries failed to seroconvert. It would, therefore, appear appropriate for OIE to refine the definition of NWC and OWC by clearly stating that these animal species are not members of the suborder Ruminantia. Furthermore, these recent results suggest that dromedaries (and most probably all camelid species), which are listed in the OIE Code chapter as being susceptible to FMD similar to cattle, sheep, goats and pigs, are much less susceptible or non-susceptible to FMD. Therefore, the importance of FMD in camelids should be re- assessed. The Central Veterinary Research Laboratory (CVRL) in Dubai, U.A.E., offers to become a reference laboratory for OWC. For more than a decade, CVRL has published in excess of 150 scientific papers and three reference books on camel diseases. Classification, population and distribution Although camelids ruminate, they are not modified ruminants in a taxonomic sense. A separate evolutionary history of 35 – 40 million years divides tylopods from ruminants. Camelidae belong to the suborder Tylopoda (Fowler, 1997; Table 1). Numerous anatomical and physiological differences justify the separate classification of Tylopoda from Ruminantia. The most important differences are shown in Table 2 and some are explained in several figures. The camelid stomach system differs from that of ruminants. There are only three distinct forestomachs compared to four in ruminants. In camelids they are called compartments (C) 1, 2 and 3. The rumen equivalent is C1, which possesses cranial and caudal glandular sacs. These were once considered to represent the water store of the animal; however they mainly function as absorption and fermentation areas as well as zones of enzymatic secretion (Wilson, 1989). The second, much smaller compartment C2 is the reticulum equivalent, and the eolongated C3 is the combined omasum/abomasun equivalent, which might best be referred to as the tubular stomach due to its length. Compartments 1 and 2 are lined with non-papillary smooth epithelium (Figure 1). In camelids, the motility patterns are markedly different compared with ruminants. Another distinguished feature of all Camelidae is the unique structure of their feet (Fig. 2). The padded feet act like snowshoes allowing them to walk over soft, loose sand without sinking. Camelids walk on thick pads consisting primarily of fat. They possess two digits, and their second and third phalanges are horizontal. The reproductive physiology of camelids is of particular interest. Camels mate in a crouching position (Fig. 3) and while mating the bull exteriorises its “doula” (Fig. 4), a bright pink inflatable sac, to attract females. Camels are induced ovulators. Their gestation period lasted 13 months. A slippery surface of a third membrane surrounding the fetus eases its birth (Figure 5). Latest osteological investigations on post- cranial skeletons of Camelus dromedarius and C. bactrianus have shown that they derived from two different ancestors. Approximately twenty million OWC exist, of which two million are Bactrians (Table 3). There are four different species of NWC which inhabit the high altitudes in South America. The estimated population of NWC is shown in Table 4. Llamas and alpacas were domesticated 7.000 years ago; the dromedary and the Bactrian around 5.000 years ago. Guanacos and vicuñcas are wild and there are few wild Bactrians which roam in the Chinese and Gobi desert. There are no wild dromedaries anymore. The distribution of OWC is shown in Figure 6. 246 The knowledge of the susceptibility and resistance to infectious and parasitic diseases is of paramount importance in an area where tylopods mix with other livestock. Review of findings on FMD in camelids FMD remains the single most important animal disease, and OWC and NWC inhabit countries in North and East Africa, the Middle and Far East as well as in South America where FMD is endemic. It has been reported that dromedaries can contract the disease following experimental infection and via close contact with FMD diseased livestock, yet do not present a risk in transmitting FMD to susceptible animals (Kitching, 2002). Summarised results are presented in the following Tables 5 to 8 (Wernery and Kaaden, 2004). Only two reports exist of a natural infection. The execution of experimental infections is poor, and therefore conclusions are questionable. FMD serology and infection in Bactrian camels remains questionable, with FMD diagnosis only being made by means of clinical observations. Results of recent FMD experiments in dromedaries in Dubai with serotype 0 Two Holstein heifers of around 150 kg (6-8 months of age) and two castrated male dromedaries (Camelus dromedarius) around 400-450 kg (7-10 years of age) were each inoculated subepidermo- 7.6 lingually with 10 Tissue Culture Infectious Doses 50% (TCID50) of foot-and-mouth disease virus (FMDV) type O UAE 7/99 in a volume of 0.5 ml (Fig. 7). While the heifers developed elevated body temperatures, were drooling saliva and had typical vesicular lesions (Fig. 8) on the tongue within 24 hours, the two dromedaries did not show any clinical signs of disease and had no vesicular lesions, even at the inoculation site. Infectious FMDV and FMDV RNA were detected at relatively high levels in sera and nasal and mouth swabs from the heifers, but no infectious FMDV or FMDV RNA were isolated in similar samples from the two dromedaries (Fig. 9). Furthermore, the two dromedaries did not develop any detectable antibodies to FMDV. Based on the overall results obtained, we conclude that dromedaries (Camelus dromedarius) are not susceptible to infection with this isolate of FMDV (Wernery et al., 2005). Conclusion Camelids belong to the suborder Tylopoda; they are not ruminants. Camelids possess a low flow susceptibility to FMD, and do not appear to be long-term carriers of the FMDV. These are the main two reasons to remove them from the OIE chapter as possessing the same degree of susceptibility as cattle, sheep and goats. References Abou Zaid, A.A., 1991. Studies on some diseases of camels. PhD Thesis, Faculty of Veterinary Medicine Zagazig, Egypt Farag, M.A., Al-Sukayran, A., Mazlou, K.S, Al-Bokney, A.M., 1998. The susceptibility of camels to natural infection with foot and mouth disease virus. Assiut Veterinary Medical Journal 40, 201 – 211 Fowler, M. E. (1997), Evolutionary history and differences between camelids and ruminants, J. 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Detection of antibodies against FMDV in camels by using fluorescent antibody technique. Agricultural Research Review 64, 1079 – 1084 Moussa, A.A., Daoud, A., Tawfik, S., 1979. Susceptibility of camel and sheep to infection with foot and mouth disease virus. Agricultural Research Revision Egypt 57, 1 –19 Moussa, A., Nasser, M.I., Mowafi, L., Salah, A., 1986a. Occurrence of foot and mouth disease in different species of mammals at Sharkia province. Journal of Egypt Veterinary Medicine Association 40, 23 – 35 247 Moussa, A.A., Tantawi, H.H., Metwally, N.A., Wahba, S., Hussein, K., Osman, O.A., Saber, M.S., 1986b. Pathogenicity of foot and mouth disease virus isolated from experimentally infected camels to susceptible steers. Agricultural Research Review 64, 1071 – 1077 Moussa, A.A.M., Daoud, A., Omar, A., Meetwally, N., El-Nimr, M., McVicar, J.W. 1987. Isolation of foot and mouth disease virus from camels with ulcerative disease syndromes. Journal of Egypt Veterinary Medicine Association 47, 219 – 229 Moussa, A.A.M., 1988. The role of camels in the epizootiology of FMD in Egypt. In: FAO. The Camel: Development Research. Proceedings of Kuwait Camel Seminar, Kuwait, Oct. 20 – 23, 1986, pp. 162 – 173 Moussa, H.A.A., Youssef, N.M.A., 1998. Serological screening for some viral diseases antibodies in camel sera in Egypt. Egypt Journal of Agricultural Research 76, 867 – 873 Nasser, M., Moussa, A.A., Metwally, M.A., Saleh, R.EL.S., 1980. Secretion and persistence of foot and mouth disease virus in faeces of experimental infected camels and ram.