The World Conservation

SSC Logo

In comparison to many other species, hippos have been the subject of little formal research.  Much of what we know about hippos comes from research performed during mass culling programs in Uganda in the late 1950's and early 1960's.  However, more recent discoveries, some of which have occurred quite unexpectedly, have researchers taking a new look at both common and pygmy hippos.  

To learn more, please take a look at the following:


- A -

Abaturov, B.D., Kolesnikov, M.P., Nikonova, O.A., and Pozdnyakova, M.K. (2003 Jan) Experience of quantitative investigation of nutrition in free-ranging mammals in natural habitat [Russian] Zoologichesky Zhurnal, 82(1); pp. 104-114.

Alix * (1879) Sur la Glande lachrymale de l'Hippopotame, Bull, Soc. Zool. Fr, iv; 117-*.

Alix * (1879) Sur la Glande lacrymale de l'Hippopotame, Bull. Soc. Philom. Paris, 7(iii); pp. 146 -*.

Alix * (1879*) Sur le Glande Sympathique de l'Hippopotame, Bull. Soc. Philom. Paris, 7(iii); p. 168.

Alix * (1879*) Sur le Plexus cervicale, Bull Soc. Philom. Paris, 7(iii); p. 167.

Allbrook, D. (1962) The morphology of the subdermal glands of Hippopotamus amphibius, Proceedings of the Zoological Society of London, 139; pp. 67-73.

Allbrook, D.B., Luck, C.P. & Wright, P.G. (1959) The cutaneous glands in hippopotamus, Proc. Anat. Soc. in J. Anat. London, 93; pp. 587-588.

Alving, T. (1932) Flusspferdgeburt und andere Ereignisse im Zoo Kopenhagen, Zool. Gart. Lpz. N.P., 5; pp. 34-37.

Amedee-Pichot, P. (1919) Les immersions de l'Hippopotame, Bull. Soc. Nat. Acclim. France, Lxvi; pp. 297-300.

Amorosa, E.C., Hancock, N.A. & Kellas, L. (1958) The foetal membranes and placenta of the hippopotamus (Hippopotamus amphibius (Linnaeus)), Proceedings of the Zoological Society of London, 130; pp. 437-447, 7 plates.

Anghi, C. (1972) Data on lung capacity of the hippopotamus, Allattani Kozl, 59(1-4); pp. 29-35.

Anon. (1993) Hippos return to Sundays River, African Wildlife, 47; p. 31.

Ansell, W.F.H. (1960) Mammals of Northern Rhodesia. Government. Printer, Lusaka.

Ansell, W.F.H. (1965) Feeding habits of Hippopotamus amphibius L., The Puku, 3; p. 171.

Ansell, W.F.H. (1965) Hippo census on the Luangwa River 1963-1964, The Puku, 3; p. 15-27.

Ansell, W.F.H. (1965) Measurements and weights of adult hippo from Zambia, Puku, 3; p. 181.

Ansell, W.F.H. (1971) Order Artiodactyla (In) Meester, J & Setzer, H.W. The mammals of Africa: An identification manual. Smithsonian Institution Press. Washington.

Ansell, W.F.H. (1973) A hippo count on the upper Luangwa River, The Puku, 7; pp. 193-195.

Ansell, W.H. (1978) The mammals of Zambia. Chilanga, Zambia.

Anstey, S. (1991) Large mammal distribution in Liberia. WWF-International Gland.

Anthony, J. (1948) Etude de moulages endocraniens d'hippopotames disparus Memoires du Museum National D'Histoire Naturelle, N.S. 26 (2); pp. 31-56.

Appleton, C.C. & Bruton, M.N. (1979) The epidemiology of schistosomiasis in the vicinity of Lake Sibaya with a note on other areas of Tongaland Natal South Africa, Ann Trop Med Parasitol, 73 (6); pp. 547-562.

Arman, P. & Field, C.R. (1973) Digestion in the hippopotamus, East African Wildlife Journal, 11; pp. 9-17.

Asdell, S.A. (1946) Patterns of Mammalian Reproduction, New York, Comstock.

Attwell, R.I.G. (1963) Surveying Luangwa hippo, Puku, 1.; pp 29-49.

Attwell, R.I.G. (1966) Oxpeckers and their associations with mammals in Zambia, Puku, 4; pp. 17-48.

- B -

Barklow, W. (1997). Hippo talk. Natural History 5/95:54.

Barklow, W.E. (1997) Some underwater sounds of the hippopotamus (Hippopotamus amphibius), Marine and Freshwater Behaviour and Physiology, 29(1-4); pp. 237-249.

Barrett, J. (1991) Tanzania tinkles with the ivory trade, The Guardian, 27th September.

Bartlett, A.D. (1871) Notes on the birth of a hippopotamus in the Society's Gardens, Proc. Zool. Soc. London; pp. 255-257.

Bartlett, A.D. (1872) Additional notes on the breeding of hippopotamus in the Society's Gardens, Proc. Zool. Society London; pp. 819-821.

Beckwitt, R., Shea, J., Osborne, D., Krueger, S., and Barklow, W. (2002 Oct) A PCR-based method for sex identification in Hippopotamus amphibius, African Zoology, 37(2); pp. 127-130.

Beddard, F.E. (1910) A contribution to the anatomy of Hippopotamus amphibius, Proceedings of the Zoological Society of London; pp. 220-232.

Bellon, P. (1551) L'histoire naturelle des estranges poisons marins avec la vraie peincture et description du dauiphin, et de plusiers autres de son espece, Paris: Regnaud Chaudiere; pp 47-52.

Bennett, D., Green, N., and Basuglo, B. (2000 Dec) The abundance of Hippopotamus amphibius in the Black Volta River at Bui National Park, Ghana., African Journal of Ecology, 38(4); pp. 372-373.

Benson, C.W. (1961) Jacanas and other birds perching on Hippopotamus, Bull. Brit. Orn. Club, 81; pp. 85-86.

Benson, C.W. (1962) Jacanas and other birds perching on Hippopotamus, Bull. Brit. Orn. Club, 82; pp.21-22.

Bere, R. M. (1959) Queen Elizabeth National Park, Uganda. The hippopotamus problem and experiment, Oryx, 5; pp. 116-124.

Bere, R.M. (1957) Fauna research. Proposal to reduce hippopotamus in the Queen Elizabeth National Park, Afr. wild Life, 11(3); pp. 213-217.

Berry, P.S.M. (19**) A hippo count on the upper Luangwa River, Puku **; 193-195.

Bhima, R. (1996 Mar) Census of hippopotamus (Hippopotamus amphibius (l)) in the Upper Shire River, Malawi, African Journal of Ecology, 34(1); pp. 83-85.

Blaszkiewitz, B. (1980) Flusspferde (Hippopotamus amphibious) in Zoo Berlin, Bongo, Berlin. 4, pp. 17-22.

Boisserie, J.R., Lihoreau, F., & Brunet, M. (2005) The position of Hippopotamidae within Cetartiodactyla, Proceedings of the National Academy of Science; 102 (5); pp. 1537-1541.

Bonelli, S.U. (1834) Ueber einem im Museum zu Turin sich befindenden Hippotamus, Isis von Oken, Leipzig, 1; p. 76.

Bourliere, F. & Verschuren, J. (1960) Introduction a l'ecologie des ongules du Parc National Albert. Explor. Parc natn. Albert. Inst. Parcs. Nat. Congo Belge, Bruxelles.

Braunitzer, G., Wright, P.G., Stangl, A., Schrank, B. & Krombach, C. (1983) Amino acid sequence of haemoglobin of hippopotamus (Hippopotamus amphibius, Artiodactyla), South African Journal of Science, 79; pp. 411-412.

Brett, M.R. (1986) The hippopotamus, Natura, No.9; 2pp.

Briand Petersen, J.C. & Casebeer, R.L. (1971) A bibliography relating to the ecology and energetics of East African large mammals, East African Wildlife Journal, 9; pp. 1-23.

Brown, C.E. (1924) Rearing hippopotamuses in captivity, Journal of Mammalogy, 5; pp. 243-246.

Bruton, M.N. (1980) An outline of the ecology of Lake Sibaya, with emphasis on the vertebrate communities. In: Studies on the Ecology of Maputaland, edited by M.N. Bruton & K.H. Cooper. pp. 382-407. Rhodes University , Grahamstown.

Bruton, M.N. (1978) Recent mammal records from eastern Tongaland in KwaZulu, with notes on hippopotamus in Lake Sibaya, Lammergeyer 24; pp. 19-27.

Bruton, M.N. (1979) The amphibians, reptiles, birds and mammals of Lake Sibaya. In. Lake Sibaya. Allanson, B.R. (ed). Junk: The Hague.

Bruton, M.N. (1979) The utilization and conservation of Lake Sibaya. In. Lake Sibaya. Allanson, B.R. (ed). Junk: The Hague.

Buck, A.C., Leivestad, H., Meya, A., Nganwa-Baguma, A.B., Nkugwa, J., Rayner, D.V., Richardson, H.D., Tejani, D. & Wright, P.G. (1969) Some observations on the blood of the hippopotamus (H. amphibius), Makerere Medical Journal, 13; pp. 25-28.

Buffon, G.L.L. (1775) De Natuurlyke historie van den Hippopotamus of het Rivierpaard. Amsterdam: J.H. Schneider

Bungartz, M.A.H. (1928) "Kiboko" das achte junge Flusspferd unseres Zoologischen Gartens, Hamburg Zool. Ztg.; pp. 2-12.

Burton, M. (1962) Systematic Dictionary of Mammals of the World, London Museum Press.

- C -

Cave, A.J.E. & Blackwood, H.J.J. (1965) Traumatic deformity of hippopotamus tusks, Proc. Zool. Soc.Lond., 146; pp. 447-450.

Cena, K. (1964) Thermoregulation in the hippopotamus, International Journal of Biometeorology 8(1); pp. 57-60.

Chapman, H.C. (1881) Observations upon the Hippopotamus, P. Acad. Nat. Sci. Philad. 1881; pp. 126-148.

Child, D. (1962) Hippo hunt in the Sahara, Afr. Wildl. 16(4); pp. 337-339.

Chilvers, H.A. (1931) Huberta goes south, CNA, South Africa.

Clark, J. (1872) Notes on the visceral anatomy of the Hippopotamus, Proc. Zool. Soc. London (1872); pp. 185-195.

Clarke, J.R. (1953) The hippopotamus in Gambia, West Africa, Journal of Mammalogy, 34(3); pp. 299-315.

Clauss, M., Frey, R., Kiefer, B., Lechner-Doll, M., Loehlein, W., Polster, C., Rossner, G.E., and Streich, W.J. (2003 Jun) The maximum attainable body size of herbivorous mammals: morphophysiological constraints on foregut, and adaptations of hindgut fermenters [Review], Oecologia, 136(1); pp. 14-27.

Clemens, E.T. & Maloiy, G.M.O. (1982) The digestive physiology of three East African herbivores: the elephant, rhinoceros and hippopotamus, J.Zool., Lond., 198; pp. 141-156.

Clough, G. (1966) Reproduction in the hippopotamus (Hippopotamus amphibius, Linn.). Unpublished Ph.D. thesis, University of Cambridge.

Clough, G. (1970) A record of "Testis cords" in the ovary of a mature hippopotamus (Hippopotamus amphibius, Linn.), Anat. Rec., 166; pp. 47-50.

Cmelik, S.H.W. & Ley, H. (1976) Composition of the lipids from the adrenal gland of Hippopotamus amphibius, Comp Biochem Physiol B Comp Biochem, 54(3); pp. 347-350.

Cohen, M. (1970) Game digest for hippopotamus Hippopotamus amphibius Linn., Unpublished document, Department of Zoology, University of Pretoria. 83pp.

Cole, M. (1992) Zimbabwe's hippo threatened by drought, Time, April 8.

Cole, M. (1992) Zimbabwe's hippos threatened by drought., New Scientist, 134(1817); p. 9.

Colyer, J.F. (1936) Variations and diseases of the teeth of animals, London, John Bale & Danielsson.

Coryndon, S.C.(1977) The taxonomy and nomenclature of the Hippopotamidae (Mammalia, Artiodactyla) and a description of two new fossil species, Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, B, 80; pp. 61-88.

Coryndon, S.C. (1978) Hippopotamidae. In Evolution of African Mammals (Ed. V.J. Maglio & H.B.S. Cooke) Harvard University Press: Cambridge, Mass. pp 483-495.

Coulson, I. (1981) Annual hippo census, South-eastern lowveld. Progress report no. 4 project no. GNP/B4/4a/2, Department of National Parks and Wildlife Management, Zimbabwe.

Cowan, D.F., Thurlbeck, W.M. & Laws, R.M .(1967) Some diseases of Hippopotamus in Uganda, Path. Vet., 4; pp. 553-567.

Crisp, E. (1867) On some points connected with the anatomy of the hippopotamus (Hippopotamus amphibius), Proceedings of the Zoological Society of London, 39; pp. 601-612.

Crisp, E. (1867) On the form, size and structure of the viscera of the Hippopotamus, as compared with the same parts in the members of the pachyderm family and in some other animals, Proceedings of the Zoological Society of London, 39; pp. 689-695.

Crowther, R. & Whyte, I. (1989) Hippos interfere, Custos, 18(1); p.10.

Cumming, D.H.M. (1982) The influence of large herbivores on savanna structure in Africa. Ecology of Tropical Savannas. Ed. B.J. Huntley and B.H. Walker. Springer-Verlag, Berlin.

Curry-Lindahl, K. (1961) Contribution a l'etude des vertebres terrestres en Afrique tropicale, Inst.Parcs Nationaux du Congo et du Ruanda. Bruxelles.

Cuvier, G. (1804) Sur l'Hippotame et sur son osteologie, Annales du Museum National D'Histoire Naturelle, Paris, 4; pp. 299-328.

- D -

Dauth, J., Dreyer, M.J., Raubenheimer, E.J., Lemmer, L.B. & De Vos, D. (1988). Blood lead concentrations in hippopotami (Hippopotamus amphibius) in the Kruger National Park. Journal of the South African Veterinary Association 59(3):153-154.

De Lalouviere, P.la.H. (1989). Record of a hippopotamus Hippopotamus amphibius killing livestock. Lammergeyer, 40, p.6-7,

De Vos, V. & De Klerk, B.D. (1980). Tetanus in a free living hippopotamus, Hippopotamus amphibius capensis, from the Kruger National Park. Koedoe 23: 171-175.

Dean, W.R.J. & MacDonald, I.A.W. (1981). A review of African birds feeding in association with mammals. Ostrich 52 (3): 135-155.

Delvingt W (1978). Ecologie de l'Hippopotame (Hippopotamus amphibius L.) au Parc National des Virunga (Zaire). PhD thesis, U of Gambloux, Belgium, 333pp.

Demuth, K. (1921) Die Structure des hodens und der accessorischen Geschlechtsdrussen bei Hippopotamus amphibious, Inaugural Dissertation, Berlin.

Denis, A. (1964). The hippos of Lake Baringo. Oryx 7 (5): 251-252.

Deocampo, D.M. (2002 Apr) Sedimentary structures generated by Hippopotamus amphibius in a lake-margin wetland, Ngorongoro Crater, Tanzania, Palaios, 17(2); pp. 212-217.

Deraniyagala, P.E.P. (1951). Some hippopotamuses of Ceylon, Western Asia, East Africa and England. Spolia Zeylanica 26 (2): 125-132.

Deraniyagala, P.E.P. (1955). The extinct Hippopotamus antiquns of Europe. Spolia Zeylanica, Bull. Nat. Mus. Ceylon. 27(2): 219-222.

Dietrich, W.O. (1928). Pleistocene deutschostafrikanische Hippopotamus-Reste. Wiss. Erg. Oldoway Exp., new series 3: 1-41.

Dinnik, J.A. & Dinnik, N.N. (1961). On the morphology and life history of Fasciola nyanzae Leiper, 1910 from the hippopotamus. J. Helminthology (R.T. Leiper Suppl.) 53-62.

Dittrich, L. (1976). Age of sexual maturity in the hippopotamus. International Zoo Yearbook 16: 171-173.

Dorst, J. & Dandelot, P. (1970). A field guide to the larger mammals of Africa. Collins: London.

Du Plessis, S.F. (1969). The past and present geographical distribution of the perissodactyla and artiodactyla in southern Africa. University of Pretoria: Unpublished M.Sc. thesis.

Dudley, J.P. (1996) Record of carnivory, scavenging and predation for Hippopotamus amphibius in Hwange National Park, Zimbabwe, Mammalia, 60(3); pp. 486-488.

Dudley, J.P. (1998 Jun) Reports of carnivory by the common hippo hippopotamus amphibius, South African Journal of Wildlife Research, 28(2); pp. 58-59.

Dudley, J.P. (1998) Strange but true - first records of carnivory for hippos in southern and eastern Africa. African wildlife, Vol 52, Issue 5, Sep-Oct. p.12-13.

Dugmore, H. (2001) Seeking four female hippos. Africa geographic, Vol 9, Issue 9, Oct. p.41.

Duncan, W.R.H. & Garton, G.A. (1968). The fatty acid composition and intramolecular structure of triglycerides from adipose tissue of the hippopotamus and the African elephant. Comp. Biochem. Physiol. 25: 319-325.


Eksteen, Jacob Johannes. (1994). The determination of acceptable hippopotamus (Hippopotamus amphibius, Linn.) densities in the Crocodile River, outside the Kruger National Park Unpublished MSc thesis, University of the Witwatersrand.

Ellerman, J.R., Morrison-Scott, T.C.S. & Hayman, R.W. (1953). Southern African Mammals 1758-1951: A reclassification. British Museum: London.

Ellery, W.N., Dahlberg, A.C., Strydom, R., Neal, M.J., and Jackson, J. (2003 May) Diversion of water flow from a floodplain wetland stream: an analysis of geomorphological setting and hydrological and ecological consequences, Journal of Environmental Management, 68(1), pp. 51-71.

Elsner, R. (1966). Diving bradycardia in the unrestrained hippopotamus. Nature, 212: 408.

Eltringham, S.K. (1974) Community changes following removal of the hippopotamus. Journal of Applied Ecology. 11(3):855-865.

Eltringham, S.K. (1974). Changes in the large mammal communities of Mweya Peninsula, Rwenzori National Park, Uganda, following removal of hippopotamus. Journal of Applied Ecology 11 (3): 855-865.

Eltringham, S. K. (1993): The common hippopotamus (Hippopotamus amphibius): In; W.L.R.Oliver (ed): Pigs, Peccaries and Hippos: Status Survey and Conservation Action Plan. IUCN, Gland, Switzerland; pp: 43-55.

Eltringham, S.K. (1999). The hippopotamus. Academic Press

Engelbrechht, E. (2000). Seekoeijag. Man Magnum, Vol 25, Issue 7, Jul. p.29-31.

- F -

Feldhake, G. (2005) Hippos: Natural History and Conservation; Voyageur Press

Field, C. R. (1970). A study of the feeding habits of the hippopotamus in the Queen Elizabeth National Park, Uganda, with some management implications. Zoologica Africana 5:71-86.

Field, C.R. (1968). A comparative study of the food habits of some wild ungulates in the Queen Elizabeth National Park, Uganda. Symp. Zool. 50c. London. 21 : 135-151.

Field, C.R. (1970). A study of the feeding habits of the hippopotamus (Hippopotamus amphibius Linn.) in the Queen Elizabeth National Park, Uganda, with some management implications. Zoologica Africana 5 (1): 71-86.

Flach, E. J. ; Furrokh, I. K. ; Thornton, S. M. ; Smith, J. (1998). Caesarean section in a pygmy hippopotamus (Choeropsis liberiensis) and the management of the wound. The Veterinary Record : Journal of the British Veterinary Association. 143, no. 22,: 611.

Flower, W.H. (1885) An introduction to the osteology of the Mammalia, Macmillan: London.

Forbes-Watson, A.D. & Turner, D.A. (1977). Little Grebe Tachybaptus ruficollis associating with hippo. Scopus 1 (1): 21.

Fradrich, H. (1967). Das verhalten der Schweine und Flusspferde. Handb. Zool. 8 (10): 1-44.

Fripp, P.J. (1981). Non-specific esterase isoenzymes of adult schistosomes from the hippopotamus (Hippopotamus amphibius). Onderstepoort J. Vet. Res. 48(4): 257.

Fry, Stephen. (1995*). The hippopotamus. London : Arrow, 356 p. *

Fuller, G. (1975). Hippopotamus amphibius along the Omo River. Walia 6: 4-7.

- G -

Garnham, P.C.C. (1960). East Afr. med. J. 37: 495-*

Garrod, A.H. (1872). Note on the placenta of the hippopotamus. Proceedings of the Zoological Society of London 821-822.

Garrod, A.H. (1879). On the brain and other parts of the hippopotamus (H. amphibius). Trans. Zool. Soc. Lond. 11: 11-17. 2 Plates.

Geoffroy St Hilaire, I.S (1858). Note sur la naissance d'un jeune hippopotame a la Menagerie du Museum de l'Histoire naturelle. in: Comptes rendus hebd. des Sc. de l'Acd. des.Sc. T. XLVI, 19: 879.

Gerneke, W.H. (1965). The chromosomes and neutrophil nuclear appendages of Hippopotamus amphibius Linnaeus 1758. Onderstepoort J. Vet. Res. 32 (1): 181-186.

Ges in Regensburg, 241

Gijzen, A. (1964) Finalement des Hippopotames nains de nouveau presents au zoo; Zoo, 29; pp. 99-101.

Goddard, J. (1971). Hippopotamus populations on the Luangwa river. Report on UN (SF)/FAO Luangwa valley conservation and development project.

Goss, D.J. (1960). Breeding notes on the hippopotamus and giraffe at Cleveland Zoo. Int. Zoo Yb. 2: 90.

Graham, L.H., Reid, K., Webster, T., Richards, M., and Joseph, S. (2002) Endocrine patterns associated with reproduction in the Nile hippopotamus (Hippopotamus amphibius) as assessed by fecal progestagen analysis, General and Comparative Endocrinology, 128(1); pp. 74-81.

Graham, L.H., Webster, T., Richards, M., Reid, K., and Joseph, S. (2002) Ovarian function in the Nile hippopotamus and the effects of Depo-Provera(TM) administration, Reproduction, (Suppl 60); pp. 65-70.

Graham-Jones, O., Senior, M. & Tong, E.H. (1961). Tranquillisation of a pregnant hippopotamus (Hippopotamus amphibius). Int. Zoo. Yearbook 3: 113-116.

Gratiolet, L.P. (1860). Recherches sur le systeme vasculaire sanguin de l'Hippopotame. C.R. seances Acad. Sc. Paris, 51: 524-528.

Gratiolet, L.P. (1867). Recherches sur l'anatomie de l'hippopotame. (ed) Alix* Paris: Victor Manson et Fils.

Gray, R.F. (1959). Hippopotamus milk analysis. Int. Zoo. Yearbook 1: 46.

Greasley, W.H. (1973). Protective maternal behaviour of hippopotamus. Puku 7: 196.

Grey, J., and Harper, D.M. (2002 Dec) Using stable isotope analyses to identify allochthonous inputs to Lake Naivasha mediated via the hippopotamus gut, Isotopes in Environmental & Health Studies, 38(4); pp. 245-250.

Griffin, M. & Grobler, H.J.W. (1991). Wetland-associated mammals of Namibia - a national review. Madoqua, Vol 17, Issue 2. p.233-237. ISSN: 1010-2299.

Guggisberg, C. (1965). The mystery of the hippo. Africana 2 (3): 6-9.

Guilbride, P.D.L., Coyle, T.J., McAnulty, E.G., Barber, L. & Lomax, G.D. (1962). Some pathogenic agents found in hippopotamus in Uganda. J. Comp. Path. 72: 137-141.

- H -

Hancock, P.G. (1978). Dynamics of an overutilized area in the Ndumu Game Reserve, Tongaland. University of Natal, Unpublished M.Sc. thesis.

Hanks, J. & Albl, P. (1973). Observations on the amniotic nodules of the hippopotamus (Hippopotamus amphibius Linn.) Puku 7: 133-136.

Harthoorn, A.M. & Lock, J.A. (1961). Advances in the use of muscle relaxing drugs for immobilization and handling of larger land mammals. Journal of Small Animal Practice. 2: 163-169.

Harthoorn, A.M. (1963). Techniques of handling large animals such as elephant, rhinoceros and hippopotamus etc. by the use of drugs. Proc. 17th World Vet Congress, Hanover 2: 183-187.

Harthoorn, A.M. (1973). The drug immobilization of large wild herbivores other than the antelopes. In: Younge, E. (ed.) The Capture and care of wild animals. Human & Rousseau, Cape Town.

Hasegawa, M. and Adachi, J. (1996 May) Phylogenetic position of cetaceans relative to artiodactyls - reanalysis of mitochondrial and nuclear sequences, Molecular Biology & Evolution, 13(5); pp. 710-717.

Hatfield, J.R., Samuelson, D.A., Lewis, P.A., and Chisholm, M.(2003 Mar) Structure and presumptive function of the iridocorneal angle of the West Indian manatee (Trichechus manatus), short-finned pilot whale (Globicephala macrorhynchus), hippopotamus (Hippopotamus amphibius), and African elephant (Loxodonta africana), Veterinary Ophthalmology, 6(1); pp. 35-43.

Havinga, J. & Beintema, J.J. (1980). Pancreatic RNAse of mammals with ruminant-like digestion amino-acid sequences of hippopotamus and sloth. Eur J Biochem, 110 (1): 131-142.

Hediger, H. (1951). Observations sur la psychologie animale dans les Parcs Nationaux du Congo Belge. Inst. Parcs. Nationaux du Congo Belge, Bruxelles.

Hediger, H. (1953). Ein symbioseartiges Verhaltnis zwischen Flusspferd und Fisch. Saugetierk. Mitt. 1: 75-76.

Hediger, H. (1955). In Traite de Zoologie. Grasse, P. Vol. 17: 510-521 Paris: Masson.

Hediger, H. (1977). The hippo takes refuge. Jet Tales Magazine NR3/77 INTA Advertising, Inc., New York.

Hegner, B. & Pilleri, G. (1968). Note on the morphology of the eye of Hippopotamus amphibius and Choeropsis liberiensis (Mammalia Artiodactyla Hippopotamidae) with special consideration of the tapetum lucidum fibrosum and the Schlemms canal. Acta. Anat. 69 (3): 458-461.

Henwood, R.R. & Keep, M.E. (1989). The capture and translocation of hippopotamus by means of chemical immobilization. Lammergeyer, Issue 40, p.30-38.

Herring, S.W. (1973). Adaptations for gape in the hippopotamus. AM Zool. 13 (4): 1343.

Herring, S.W. (1975). Adaptations for gape in the hippopotamus and its relatives. Forma et Functio 8: 85-100.

Herring, S.W. (1980). Functional design of cranial muscles comparative and physiological studies in pigs. AM Zool 20 (1): 283-294.

Hill, A. & Wyatt, J. (1976). Non-aggressive tactile interactions of Hippopotamus amphibius Linn. with Syncerus caffer (Sparrman). Mammalia 40 (1): 161-162.

Hillaby, J. (1962). Harvesting the hippo. New Scientist. 291 : 588-589.

Hoier, R. (1950). Hippopotames (Hippopotamus amphibius Linne). In: A travers plaines et volcans au Parc National Albert, Explorations du Parc National Albert. Institut des Parcs Nationaux du Congo Belge, Brussels, 22-30.

Hoier, R. (1950). Les hippopotames au Parc Albert. Zooleo No.5: 15-21.

Hoier, R. (1955) A Travers Plaines et Volcans au Parc National Albert. Brussels : Institut des Parcs Nationaux du Congo Belge.

Hoier, R. (1955). L'hippopotame In Le Grande Livre de la Faune Africaine et da sa Chasse. Kisten, Geneva 41-51.

Hofmann, L. (1923) Zur Anatomie des männlichen elephanten - tapir und hippopotamus Genitale, Zoologische Jahrbücher: Abteilung fur Anatomie und Ontogenie der Tiere, 45; pp. 161-212.

Honacki, J.H., Kinman, K.E. & Koeppl, J.W. (eds) (1982). Mammal species of the world - a taxonomic and geographic reference. Allen Press. p317.

Hooijer, D.A. (1942). On the supposed hexaprotodont milk dentition of Hippopotamus amphibius L. Zool. Med. Museum Leiden 24: 187-196.

Hooijer, D.A. (1950). The fossil Hippopotamidae of Asia, with notes on the recent species. Zool. Verh. Rijksmus 8: 1-124.

Hooijer, D.A. (1958). Pleistocene remains of hippopotamus from the Orange Free State. Researches of the Nasionale Museum, 1 (2): 259-266.

Houtekamer, J.L. & Sondaar, P.Y. (1979). Osteology of the fore limb of the Pleistocene dwarf hippopotamus Phanourios minor from Cyprus with special reference to phylogeny and function 1. Proc. K Ned. Akad. Wet. Ser. B Palaeontol. Geol. Phys. Chem. 82 (4): 411-429.

Houtekamer, J.L. & Sondaar, P.Y. (1979). Osteology of the fore limb of the Pleistocene dwarf hippopotamus Phanourios minor from Cyprus with special reference to phylogeny and function 2. Proc. K Ned. Akad. Wet. Ser. B Palaeontol. Geol. Phys. Chem. 82 (4): 430-448.

- I -

Ito, A ; Miyazaki, Y ; Imal, S. (2001). Descriptions of new Parentodinium ciliates in the family Parentodiniidae n. fam. from Hippopotamus amphibius in comparison with some entodiniomorphs from horses and cattle. European Journal of Protistology. 37, no. 4, 405 (22 pages). ISSN0932-4739.

- J -

Jacobsen, N. H. G., and C. J. Kleynhaus. (1993). The importance of weirs as refugia for hippopotamus and crocodiles in the Limpopo River, South Africa. Water South Africa 19(4):301-306.

Jewell, P.A. (1963). Wild life research in East and Central Africa. Oryx 7 (2 & 3): 77-87.

Jobaert, A.J. (1954). L'hippopotame. Zooleo 29: 525-537. Leopoldville.

Joleand, L. (1921). Considerations sur le systeme dentaire des Hippopotames. Bull. Soc. Zool. France 46 (1-7): 17-22.

- K -

Kain, D.N.E. (1956). South Africa's hippo herds on the march. Veld & Vlei 1 (44): 9-26.

Karstad, E.L. (1984). The ecology of hippotami (Hippopotamus amphibius ) in southwestern Kenya. M.Sc. thesis, University of Alberta.

Karstad, E.L. & Hudson, R.J. (1984). Numbers and distribution of hippopotami of the Mara River, southwestern Kenya. African Journal of Ecology 22;143-147.

Karstad, E.L. & Hudson, R.J. (1986). Social organization and communication of riverine hippopotami in southwestern Kenya. Mammalia 50(2):153-164.

Kenneth, J.H. & Ritchie, G.R. (1953). Gestation periods. A table and bibliography. Technical communication No. 5. Commonwealth Bureau of Animal Breeding and Genetics. Edinburgh.

Kerbert, C. (1922). Over dracht, geboorte, puberteit en levensduur van Hippopotamus amphibius L. Bijdragen tot de Dierkunde, Festnummer, M. Webber, 22: 185-191.

Kettlitz, W.K. (1962). The distribution of some of the larger game mammals in the Transvaal (excluding the Kruger National Park). Annals of the Cape Provincial Museums 2: 118-137.

Kilham, P. (1982). The effect of hippopotamuses on potassium and phosphate ion concentrations in an African lake. The American Midland Naturalist. 108 (1): 202-205.

Kirchshofer, Rosl. (1970). Notiz uber eine steissgeburt beim flusspferd (Hippopotamus amphibius Linne). Einige Bemerkungen zur Vermehrungsrate des Frankfurter Flusspferdpaares. [A short account on a pelvic presentation in a hippopotamus Hippopotamus amphibius some remarks on the multiplication rate of the pair of hippopotamuses in the Frankfurt zoo.] Zeitschrift fur saugetierkunde 35 (1): 27-34.

Klingel H. (1979). Das Kriegsgemetzel in Uganda. Das Tier 10, 10-13.

Klingel H. (1979). Soziale Organisation des Flusspferdes Hippopotamus amphibius. Verhandlungen der Deutschen Zoologischen Gesellschaft. in Regensburg, 241.

Klingel H. (1981). Großes Maul, aber friedlich. Sielmanns Tierwelt 11, 46-51.

Klingel H. (1981). Wie Flusspferde ihre Umwelt gestalten. Sielmanns Tierwelt 2, 30-37

Klingel H. (1983). Life with the gentle giants. Swara 6, 24-27.

Klingel H. (1988). Flusspferd. in Grzimek B (ed): Grzimeks Enzyklopädie 5, 60-61, 64-79, Kindler.

Klingel H. (1989). Hippopotamuses. in Grzimek B (ed): Grzimek's Encyclopedia of Mammals 5, 60-79, McGraw Hill.

Klingel H. (1991). Hippos. Anima Japan 9, 32-37.

Klingel H. (1991). Sizing up a Heavyweight. Int.Wildlife 21, 4-10.

Klingel H. (1991). The social organization and behaviour of Hippopotamus amphibius. In Kayanja IB & Edroma EL (eds): African Wildlife: Research and Management 73-75, ICSU, Paris.

Klingel H. (1993). Marking behaviour in Hippopotamus amphibius. XXIII International Ethological Conference 90.

Klingel H. (1995). Fluctuating Fortunes of the River Horse. Natural History 5/95, 46-57.

Klingel H. (1995). Zoff in Hippo Hills. BBC Wildlife 8, 6-15.

Kock, von D. (1970). Die verbreitungs geschichte des Flusspferdes, Hippopotamus amphibius. Sauget Mitt. 18 (1): 12-15.

Krahnert, R. (1942). Zur Anatomie des Flusspferdeherzens (Hippopotamus amphibius und Choeropsis liberiensis). Zeits. Wiss. Zool. CLV, 4: 317-342.

Krueger, S.K. (1998). When things go as planned: Introducing an unrelated ten month old hippo to an established adult pair of over forty years. Proceedings of the 1998 American Zoo and Aquarium Association Regional Conferences, pp. 470-475.

Krueger, S. K. (2000). Investigating echolocation in the Hippopotamus (Hippopotamus amphibius). M.S. Thesis. Bowling Green State University, Bowling Green, Ohio.

Krueger, S., Barklow, W., Schlagheck, L. (1998). Developing a hippo skin tissue biopsy dart. In: Proceedings of the 1998 American Association of Zoo Veterinarians and American Association of Wildlife Veterinarians Joint Conference, pp. 512-515.

Kruger, F.J. (1990). Host cellular components adhering to the tegument of schistosomes from cattle, buffalo, hippopotamus and lechwe. Onderstepoort Journal of Veterinary Research, Vol 57, Issue 2,. p.137-9.

Kruger, F.J. & Hamilton-Attwell, V.L. (1988). The tegument of Schistosoma hippopotami from Hippopotamus amphibius in the Kruger National Park. Onderstepoort Journal of Veterinary Research, Vol 55, Issue 3, Pages: p.153-5.

Kyriacou, X.S. (1980). Decomposition studies at Lake Bhangazi, Zululand. University of Natal: Unpublished B.Sc. (Hons) thesis.


Lakshminarasimhan, A & Seshadri, V.K. (1959). Observations on the anatomy of a hippopotamus foetus (Hippopotamus amphibius). Indian Veterinary Journal 36: 584-591.

Langer, P. (1974). Stomach evolution in the Artiodactyla. Mammalia 38 (2): 295-314.

Langer, P. (1975). Macroscopic anatomy of the stomach of the Hippopotamidae. Gray, 1821. Zbl. Vet. Med. C. 4: 334-359.

Langer, P. (1976). Functional anatomy of the Stomach of Hippopotamus amphibius L. 1758. South African Journal of Science 72: 12-16.

Langer, P. (1979). Phylogenetic adaptation of the stomach of the macropodidae to food. Z Saeugetierkd 44 (6): 321-333.

Laurie, A & Olivier, R. (1977). Pachyderms in peril. New Scientist 15 September 1977:658-660.

Laws, R.M. & Clough, G. (1965). Observation of reproduction in the hippopotamus (Hippopotamus amphibius). Journal of Reproduction and Fertility 9: 369-370.

Laws, R.M. & Clough, G. (1966). Observations on reproduction in the hippopotamus Hippopotamus amphibius Linn. in Rowlands, I.W. (ed) Comparative biology of reproduction in mammals. Symposium of the Zoological Society of London. 15: 117-140.

Laws, R.M. (1968). Dentition and ageing of the hippopotamus. East African Wildlife Journal 6: 19-52.

Laws, R.M. (1968). Interactions between elephant and hippopotamus populations and their environment. Bull. Epizoot. Dis. Afr. 16 (2): 238.

Laws, R.M. (1968). Interactions between elephant and hippopotamus populations and their environments. Proc. of the symposium on wildlife, Nairobi East African Agricultural and Forestry Journal XXXIII: 140-147.

Laws, R.M. (1989) Development of research on large mammals in East Africa. Symposium of the Zoological Society of London. No.61:267-298.

Laws, R.M., Parker, I.S.C. & Johnstone, R.C.B. (1975). Elephants and their habitats. The ecology of elephants in North Bunyoro, Uganda. Clarendene Press, Oxford.

Leidy, J. (1952). On the osteology of the head of Hippopotamus and a description of the osteological characters of a new genus of Hippopotamidae. Jour. Acad. Nat. Sci. n.s. 2(3): 207-234.

Leiper, R.T. (1910). The entozoa of the hippopotamus. Proc. Zool. Soc. Lond. 1:233-251.

Leivestad, H., Richardson, D. & Wright, P.G. (1973). The respiratory properties of the blood of the hippopotamus. Respiration Physiology 19: 19-25.

Leuthold, W. (1977). African Ungulates: A comparative view of their Ethology and Behavioural Ecology. Springer Verlag, Berlin.

Lewison, R. (1998 Sep) Infanticide in the hippopotamus - evidence for polygynous ungulates, Ethology Ecology & Evolution, 10(3); pp. 277-286.

Lewison, R. (2007) Population responses to environmental and human-mediated distrubances: Assessing the vulnerability of the common hippopotamus (Hippopotamus amphibius), African Journal of Ecology (in Press).

Lewison, R. and Carter, J. (2004 Jan 1) Exploring behavior of an unusual megaherbivore: a spatially explicit foraging model of the hippopotamus, Ecological Modelling, 171(1-2); pp. 127-138.

Leyhausen, P. (1976). Erosion and the hippos. Oryx XIII (3): 303-304.

Lochte, T.H. (1951). Untersuchungen an Haaren eines neugeborenen Nilpferdes und eins Zwergflusspferdes. Zool. Garten, XVIII (3-4): 119-124.

Lock, J. M. (1972). The effect of hippopotamus grazing on grasslands. Journal of Ecology 60(2):445-467.

Lock, J.M. & Field, C.R. (1969). Effects of hippopotamus grazing on grasslands. J Anim. Ecol. 38 (3): 24p.

Longhurst, W. M. (1958). Wildlife research in Uganda. Trustees of the Ugandan National Parks, Progress report No. 3.

Luck, C.P. & Wright, P.G. (1959). Aqueous humour of the hippopotamus. Nature 183 : 1595-1596.

Luck, C.P. & Wright, P.G. (1959). The body temperature of the hippopotamus. J. Physiol 147: 53-54.

Luck, C.P. & Wright, P.G. (1962). Aspects of the body temperature and habitat of large animals. (in) Biometeorology p 334 Oxford: Pergamon Press.

Luck, C.P. & Wright, P.G. (1964). Aspects of the anatomy and physiology of the skin of the hippopotamus H. amphibius. Quarterly Journal of Experimental Physiology and Cognate Medical Sciences, 49 (1): 1-14.

Lucke, R. (1929). Der Kehlkopf des Hippopotamus amphibius. Anat. Anz. LXVII: 241-264.

Lydekker, (1915).

Lyle, A. (1931). Hippopotamus in: T.F.Dreyer and A.Lyle, New fossil mammals and man from South Africa. Bloemfontein (Nasionale Pers Bpk.) 13-20.

- M -

Macdonald, A.A. and Hartman, W. (1983) Comparative and functional morphology of the stomach in the adult and newborn Pigmy Hippopotamus (Choeropsis liberiensis), Journal of Morphology 177; pp. 269-276.

Macdonald, A.A. and Bosma, A.A. (1985) Notes on placentation in the Suina. Placenta, 6; pp. 83-92.

Macdonald, A.A. and Fowden, A.L. (1997) Microscopic anatomy of the ungulate placenta. Equine Veterinary Journal, Supplement, 24; pp. 7-13.

MacDonald, I.A.W. & Brooks, P.M. (1983) Monitoring for detection of mammal overabundance in small areas. Management of Large Mammals in African Conservation Areas, ed. N. Owen-Smith. Haum Publishers, Pretoria.

Mackie, C. (1976). Feeding habits of the hippopotamus on the Lundi River, Rhodesia. Arnoldia Rhodesia 7 (34): 1-16.

Mackie, C.S. (1973). Interactions between the hippopotamus (Hippopotamus amphibius L.) and its environment on the Lundi River. Unpublished thesis, Department of Botany and Zoology. University of Rhodesia.

Maluf, N. S. R. (1994) Renal Anatomy of the Pigmy Hippopotamus (Choeropsis liberiensis): An Overview. Zentralblatt für Veterinärmedizin. Reihe C, Anatomia, histologia, embryologia : Journal der Weltvereinigung der Veterinäranatomen. 23, no. 3,: 189.

Manlius, N. (2000 Jan) Historical ecology and biogeography of the hippopotamus in Egypt [French], Belgian Journal of Zoology, 130(1); pp. 59-66.

Marshall, P. (1972). A hippopotamus (Hippopotamus amphibius L.) population : its structure, dynamics and management. Unpublished B.Sc. Thesis, University of Edinborough. 72pp.

Marshall, P.J. & Sayer, J.A. (1976) Population ecology and response to cropping of a hippopotomus population in eastern Zambia. Journal of Applied Ecology 13 (2): 391-404.

Matsuzaki, M. & Nakagawa, S. (1973). The copulation of hippopotamus on land. J. Mamm. Soc. Japan 5 (6): 212.

Maung, M. (1975). An ascaridoid nematode from a hippopotamus. Int J Parasitol 5 (4): 431-440.

McCarthy, T.S., Ellery, W.N., and Bloem, A. (1998 Mar) Some observations on the geomorphological impact of hippopotamus (Hippopotamus amphibius l) in the Okavango Delta, Botswana, African Journal of Ecology, 36(1); pp. 44-56.

McCully, R.M., Van Niekerk, J. & Kruger, S.P. (1965). The pathology of bilharziasis in Hippopotamus amphibius. South African Medical Journal. 39: 1026.

McCully, R.M., Van Niekerk, J.W. & Kruger, S.P. (1967). Observations on the pathology of Bilharziasis and other parasitic infestations of Hippopotamus amphibius Linnaeus 1758, from the Kruger National Park. Onderstepoort J. Vet. Res. 34 (2): 563-618.

Members of the R.V.C. East African Expedition 1962 (1963). The use of Thiambutene/Phencylidine/Hyoscine mixture for the immobilisation of topi (Damaliscus korrigum) and the hippopotamus (Hippopotamus amphibius). Vet. Rec. 75(24): 630-633.

Middleton, I. (1972). A study of a low density hippopotamus population in the mid-Zambezi valley, Rhodesia. Unpublished thesis, University of Rhodesia.

Mkanda, F.X. (1994 Mar) Conflicts between hippopotamus (Hippopotamus-amphibius (l)) and man in Malawi, African Journal of Ecology, 32(1); pp. 75-79.

Mkanda, F X ; Kumchedwa, B. (1997). Relationship between crop damage by hippopotamus (Hippopotamus amphibius L.) and farmer complaints in the Elephant Marsh. Journal of African zoology. 111, no. 1,: 27 (12 pages)

Moffet, J. P. (1958). Handbook of Tanganyika, 2nd edition. Government of Tanzania, Dar es Salaam, Tanzania.

Moloi, S.K., Kutuza, S.B. & Boreham, P.F.L. (1980). Studies on Glossina pallidipes, Glossina fuscipes fuscipes and Glossina brevipalpis in terms of the epidemiology and epizootiology of trypanosomiases in Southern-eastern Uganda. Ann.Trop. Med. Parasitol 74 (2): 219-238.

Morton, S.G. (1844). On a supposed new species of hippopotamus. Proc. Acad. nat. Sci. Philad. 2: 14-17.

Morton, S.G. (1849). Additional observations on a new living species of hippopotamus. J. Acad. nat. Sci. Philad. (2)1:231-239.

Mouquet, A. (1919). Gestations d'une femelle d'Hippopotame. Bull. Soc. Nat. d'Accl. France 167-180.

Msuya, P. (1963). A chemical examination of the red dye secreted in the sweat of the hippopotamus. Biochem. Journal. 89 (2): 80.

Mugangu T.E. & Hunter M. L. (1992). Aquatic foraging by hippopotamus in Zaire - Response to a food shortage. Mammalia 56 (3):345-349.

Muwanika, V.B., Siegismund, H.R., Bosco, J., Okello, A., Masembe, C., Arctander, P., and Nyakaana, S. (2003 Aug) A recent bottleneck in the warthog and elephant populations of Queen Elizabeth National Park, revealed by a comparative study of four mammalian species in Uganda national parks, Animal Conservation, 6(3); pp. 237-245.

Mwase, M., Almli, B., Sivertsen, T., Musonda, M.M., and Flaoyen, A. (2002 Sep) Hepatic and renal concentrations of copper and other trace elements in hippopotami (Hippopotamus amphibius L) living in and adjacent to the Kafue and Luangwa Rivers in Zambia, Onderstepoort Journal of Veterinary Research, 69(3); pp. 207-214.

- N -

Nakagawa, S. (1971). A case of precocious maturity in a female hippopotamus of Ueno Zoological Garden. J. mammal Soc. Japan 5: 139-141.

Naylor, J.N. Caughley, G.J. Abel, N.D.J. & Liberg, O. (1973). Luangwa valley conservation and development project, Zambia. Game management and habitat manipulation. Rome: FAO.

Ngog Nje, J. (1988). Contribution à l'etude de la population des hippopotames (Hippopotamus amphibius L.) au Parc national de la Benoue (Cameroun). Mammalia 52(2):149-158.

Nomura, O., Lin, Z.H., Muladno, Wada, Y., and Yasue, H. (1998 Jul) A sine species from hippopotamus and its distribution among animal species, Mammalian Genome, 9(7); pp. 550-555.

Nomura, O. and Yasue, H. (1999 May) Genetic relationships among hippopotamus, whales, and bovine based on SINE insertion analysis, Mammalian Genome, 10(5); pp. 526-527.

Norton, P.M. (1988). Hippopotamus numbers in the Luangwa Valley, Zambia, in 1981. African Journal of Ecology 26:337-339.

- O -

O'Connor, T.G. & Campbell, B.M. (1986). Hippopotamus habitat relationships on the Lundi River, Ghonarezhou National Park, Zimbabwe. African Journal of Ecology 24:7-26.

Ogden, C.G. (1967). A little-known nematode Cobboldina vivipara from the hippopotamus re-described. J Natur. Hist. 1 (3): 389-392.

Olivier, R. (1973). The enigmatic hippopotamus. African Wildlife Leadership Foundation News 8 (3): 14-17.

Olivier, R.C.D. & Laurie, W.A. (1974). Birds associating with hippopotamuses. Auk, 91:169-170.

Olivier, R.C.D. & Laurie, W.A. (1974). Habitat utilization by hippopotamus in the Mara River. East African Wildlife Journal 12: 249-271.

Owen, R. (1840-1845). Odontography 2: P1 142. London: Bailliere.

Owen-Smith, R.N. (1989). Megaherbivores: The influence of very large body size on ecology. Cambridge University Press, Cambridge.

- P -

Parker, G.H. (1932). The normal period of submergence for the Hippopotamus. Amer J. Physiol., 99, 577-578.

Parker, G.H. (1935). The breathing rate of the hippopotamus as indicated by its submergence periods. Journal of Mammalogy, XVI, 115-117.

Passey, B.H., and Cerling, T.E. (2002 Sep) Tooth enamel mineralization in ungulates: Implications for recovering a primary isotopic time-series, Geochimica et Cosmochimica Acta, 66(18); pp. 3225-3234.

Peters, *. (1852). Reise nach Mossambique Saugetiere 1852: 180-*.

Petrides, G.A. & Swank, W.G. (1965) Population densities and the range-carrying capacity for large mammals in Queen Elizabeth National Park, Uganda. Zoologica Africana 1 (1): 209-225.

Phillipson, J, & Pomeroy, D.E. (1972). Hippopotamus defending carcass? East African Natural History Society Bulletin 10: 173-174.

Physick, R. (1972). Hippopotamus behaviour. Lammergeyer 15: 78.

Pienaar, U. de V. (1963). The large mammals of the Kruger National Park - Their distribution and present-day status. Koedoe 6: 1-37.

Pienaar, U. de V. (1967). The field-immobilization and capture of hippopotami (Hippopotamus amphibius Linnaeus) in their aquatic element. Koedoe 10: 149-157.

Pienaar, U. de V. (1968). Recent advances in the field immobilization and restraint of wild ungulates in South African National Parks. Acta Zoologica et Pathologica Antverpiensia 46: 17-38.

Pienaar, U. de V. (1969). Capture and immobilizing techniques currently employed in South African National Parks and Reserves. In. Golley, F.R. & Buechner, H.K. (eds.) A practical guide to the study of the productivity of large herbivores. Oxford: Blackwell.

Pienaar, U. de V. (1969). The use of drugs in the field immobilization and restraint of large wild mammals in South African National Parks. Acta Zoologica Antverpiensia 48: 163-177.

Pienaar, U. de V., van Niekerk, J.W., Young, E. & van Wyk, P. (1966). Neuroleptic narcosis of large wild herbivores in South African National Parks with the new potent morphine analogues M-99 and M-183. Journal of South African Veterinary Medical Association 37(3): 277-291.

Pienaar, U. de V., Van Wyk, P. & Fairall, N. (1966). An experimental cropping scheme of hippopotami in the Letaba river of the Kruger National Park. Koedoe 9: 1-33.

Pilliet, A. & Boulart, R. (1886). Sur l'estomae de l'hippopotame, du kanguroo de Bennet et du paresseaux Ai. J. Ant. Physiol. norm. pathol. de l'Homme et des Animaux 22, 402-423.

Pitchford, R.J. (1966). Some African schistosomes: genus Schistosoma. S. Afr. J. Sci. 62: 202-205.

Pitchford, R.J. Visser, P. (1981). Schistosoma, Weinland, 1858 from Hippopotamus amphibius, Linnaeus, 1758 in the Kruger National Park. Onderstepoort Journal of Veterinary Research. 48: *.

Pitman, C.R.S. (1961). Birds perching on hippopotamus. Bull. Brit. Orn. Club 81: 148.

Pitman, C.R.S. (1962). Albino hippopotamus. Proceedings of the Zoological Society of London 139(4): 531-534.

Plowright, W., Laws, R.M. & Rampton, C.S. (1964). Serological evidence for the susceptibility of the hippopotamus (Hippopotamus amphibius Linnaeus) to natural infection with rinderpest virus. J. Hyg. Camb. 62: 329-336.

Pocock, R.I. (1918). The hippopotamus and duration of submergence. The Field, CXXXII, 348.

Pooley, A.C. (1967). Bird/crocodile and bird/hippopotamus commensalism in Zululand. Ostrich Vol* 11-12.

Pycraft, W.P. (1938). Pygmies and giants: Two species of hippopotamus. Illus. London News 193 (5182): 273.

- R -

Ramsay, E.C., Loomis, M.R., Mehren, K.G., Boardman, W.S.J., Jensen, J., and Geiser, D. (1998 Mar) Chemical restraint of the Nile hippopotamus (hippopotamus amphibius) in captivity, Journal of Zoo & Wildlife Medicine, 29(1); pp. 45-49.

Rautenbach, I.L. (1982). Mammals of the Transvaal. Ecoplan monograph No. 1: Pretoria.

Reed, G.T. (1979). Immobilization of 2 captive adult Nile hippopotami (Hippopotamus amphibius). AM Assoc. Zoo. Vet. Ann. Proc. 150-153.

Rice, D.W. (1963) Birds associating with elephants and hippopotamuses. Auk 80: 196-197.

Roberts, A. (1951). The mammals of South Africa. Central News Agency: South Africa.

Rollinson, D.H.L. & Lomax, G.D. (1959). Paper electrophoresis of hippopotamus haemoglobin. Nature 184: 1653-1654.

Ruggiero, R.G. (1996 Apr) Interspecific feeding associations - mutualism and semi-parasitism between hippopotami (Hippopotamus amphibius) and African Jacanas (Actophilornis Africanus), Ibis, 138(2); pp. 346-348.

- S -

Saban, R, (1971). La musculature peauciere de la tete et du con chez Hippopotamus amphibius L. 1758 Gegenbaurs morph. Jb. 116: 303-321.

Saikawa, Y., Hashimoto, K., Nakata, M., Yoshihara, M., Nagai, K., Ida, M., and Komiya, T. (2004 May 27) Pigment chemistry: The red sweat of the hippopotamus, Nature, 429(6990); p. 363.

Sailer, O. (1950). Aus der Nilpferd-Aufzucht des Dresdener Zoo. Nene Ergebn und Prob. Zool., Zool. Anz. Erganz 165: 835-839.

Sayer, J.A. & Rakha, A.M. (1974). The age of puberty of the hippopotamus (Hippopotamus amphibius Linn.) in the Luangwa River in Eastern Zambia. East African Wildlife Journal 12: 227-232.

Schmidt, F. (1935). Das erste im Halle geborene Flusspferd. Mitt. Zool. Garten Halle, XXX.

Schumacher, S.V. (1918). Histologische Untersuchung der ausseren Haut eines neugeborenen Hippopotamus amphibius. Denkschr. Akad. Wiss. Wien. 94: 1-*.

Schumachers, S. (1918). Mucous glands in the skin of a hippopotamus. Denkschr. Akad. Wiss. Wien. 94: 1-53.

Schutte, G.W. (1973). Hippopotamus behaviour after shooting an adult female. Lammergeyer 19: 36.

Schwartz, D. (1996). Snatching scientific secrets from the hippo's gaping jaws. Smithsonian 26(12): 90-102.

Sclater, W.L. (1900). The mammals of South Africa. London, R.H. Porter.

Scotcher, J.S.B. (1973). Observations on the breeding behaviour of hippopotamus. Lammergeyer 19: 35-36.

Scotcher, J.S.B. (1974). A quantitative assessment of the food preferences of Hippopotamus amphibius L. in the Ndumu Game Reserve, Tongaland. University of Natal: Unpublished M.Sc. thesis.

Scotcher, J.S.B. (1978). Hippopotamus numbers and movements in Ndumu Game Reserve. Lammergeyer 24: 5-12.

Scotcher, J.S.B., Stewart, D.R.M. & Breen, C.M. (1978). The diet of the hippopotamus in Ndumu Game Reserve, Natal, as determined by faecal analysis. South African Journal of Wildlife Research 8(1): 1-11.

Seefeld, A. (1889). Nilpferdzucht in Zool. Garten Zu St Petersburg. Der Zool Garten Jahrg xxx.

Senior, M. & Tong, E.H. (1963). Parturition in a hippopotamus. Proceedings of the Zoological Society of London. 140(1): 57-59.

Sey, O. & Graber, M. (1980). Nilocotyle duplicisphinctris new-species trematoda paramphistomidae from Hippopotamus amphibius. Dep. Zool. Coll. Educ. Pecs. Hung. J Helminthol 54 (2): 123-128.

Shimoyama, M., Morimoto, S., and Ozaki, Y. (2004) Non-destructive analysis of the two subspecies of African elephants, mammoth, hippopotamus, and sperm whale ivories by visible and short-wave near infrared spectroscopy and chemometrics, Analyst, 129(6); pp. 559-563.

Shortridge, G.C. (1934). The mammals of South West Africa. Heinemann: London.

Sidney, J. (1965). The past and present distribution of some African ungulates. Transactions of the Zoological Society of London 30: 1-397.

Sikes, S.K. (1968). Observations on the internal structure of the aorta in East African mammals, with particular reference to the klipspringer, gerenuk and hippopotamus. Zeits. fur Sauget. 33 (1):27-42.

Skead, C.J. (1980). Historical mammal incidence in the Cape Province. 1:The Western and Northern Cape 392-432.

Skinner, J.A., Scorer, J.A. & Millar, R.P. (1975). Observations on the reproductive physiological status of mature herd bulls, bachelor bulls and young bulls in the hippopotamus. Hippopotamus amphibius Linnaeus General and Comparative Endocrinology 26: 92-95.

Sleeper, B. (1977). Introduction of a young male to an adult female Hippopotamus amphibius. In: Crockett, C & Hutchins, M. (eds). Applied behavioral research at the Woodland Park Zoological Gardens, Seattle, Washington: 331-369.

Sleeper, B. (1982). Hippos: river horses of Africa. Swara 5(3): 25-27.

Smart, A.C. (1990) The density of Hippopotamus amphibius, Linnaeus at Lake Naivasha Kenya. Tropical Freshwater Biology. 2(2). 1990.

Smith, N.S. (1970). Appraisal of condition estimation methods for East African Ungulates. E.Afr. Wildl. J. 8: 123-129.

Smithers, R.H.N. & Lobao Tello, J.L.P. (1976). Checklist and atlas of the mammals of Mocambique. Museum memoir No. 8. National Museums Monuments of Rhodesia: Salisbury.

Smithers, R.H.N. & Wilson, V.J. (1979). Checklist and atlas of the mammals of Zimbabwe, Rhodesia. Museum memoir No. 9. National Museums and Monuments: Salisbury.

Smithers, R.H.N. (1966). The mammals of Rhodesia, Zambia and Malawi. Collins: London.

Smithers, R.H.N. (1971). The mammals of Botswana. Museum Memoir No. 4. National Museums of Rhodesia: Salisbury.

Smuts, G.L. & Whyte, I.J. (1981). Relationships between reproduction and environment in the hippopotamus Hippopotamus amphibius in the Kruger National Park. Koedoe 24: 169-185.

Smuts, G.L. (1978). Reproduction and population data for hippopotami from Kruger National Park. National Parks Board: Unpublished report.

Sokolowsky, A. (1915). Die Fortpflanzung de Flusspferdes. in: Promethus, Leipzig, Otto Spamer. XXVI (1326): 410-411.

Spinage, C.A. (1959). Hippos galore. The Kazinga Channel and its hippo problem. African Wildlife 13 (2): 117-121.

Stone, S.S. & Heuchele, W.P. (1965). The role of the hippopotamus in the epizootiology of African swine fever. Bull. Epiz. Dis. Afr. 13: 23-28.

Stoskopf, M.K. & Bishop, L. (1978). Immobilization of two captive adult Nile hippo (Hippopotamus amphibius). J. Zoo. Anim. Med. 9(3): 103-107.

Stunkard, H.L. (1924). A new trematode, Oculotrema hippopotami n.g n.sp. from the eye of the hippopotamus. Parisitology 16: 436-440.

Swart, P.J. (1961). Nilocotyle hepaticae n. sp. from Hippopotamus amphibius. Onderstepoort J. Vet. Research 28: 551-556.

Swart, P.J. (1966). A redescription of Nylocotyle (Nilocotyle) praesphinctris Nasmark, 1937 (Trematoda: Paramphistomidae from Hippopotamus amphibius) Onderstepoort J. Vet. Res. 33(1): 73-76.


Tanzania Wildlife Conservation Monitoring (TWCM) (1991). Wildlife census: Katavi- Rukwa, November 1991.

Taylor, R.H. (1976). Hippopotamuses at Lake St Lucia. In: Heydorn, A.E.F. St Lucia Scientific Advisory Council Workshop Meeting - Charter's Creek February 1976. Natal Parks Board, Pietermaritzburg.

Taylor, R.H. (1980). A land capability study for hippopotamuses at Lake St Lucia, Zululand. Unpublished M.Sc. thesis, University of Natal.

Tembo, A. (1987). Population status of the hippopotamus on the Luangwa River, Zambia. African Journal of Ecology 25:71-77.

Thal, J. & Ngog Nje, J. (1987). Rapport concernant les mortalites d'hippopotames au Parc National de la Bernoue. Ecole de Faune, Garoua, Cameroun. 6 p.

Thompson, R.B., Borden, D., Tarr, G.E. & Margoliash, E. (1978). Heterogeneity of amino acid sequence in hippopotamus cytochrome C. J. Biol. Chem. 253 (24): 8957-8961.

Thomson, A.L. (1964). Birds associated with elephants and hippopotamuses. Auk 81 (3): 436.

Thornton, D. D. (1971). The effect of complete removal of hippopotamus on grassland in the Queen Elizabeth National Park, Uganda. East African Wildlife Journal 9:47-55.

Thurlbeck, W.M. (1965). Artiosclerosis in hippopotami. In: Comparative artiosclerosis: 15 6 Roberts, J.C. and Straus, S. (eds). New York: Harper & Row.

Thurston, J.P. & Grain, J. (1971). Holotrich ciliates from the stomach of Hippopotamus amphibius, with descriptions of two new genera and four new species. Journal of Protozoology 18(1): 133-141.

Thurston, J.P. & Noirot-Timothee, C. (1973). Entodiniomorph ciliates from the stomach of Hippopotamus amphibius with descriptions of two new genera and three new species. J. Protozol. 20 (5): 562-565.

Thurston, J.P. (1963). Schistosomes from Hippopotamus amphibius L. I. The morphology of Schistosoma hippopotami sp. nov. Parasitology 53: 49-54.

Thurston, J.P. (1964). Schistosomes from Hippopotamus amphibius L. II. The morphology of Schistosoma edwardiense sp. nov. Parasitology 54:67-72.

Thurston, J.P. Noirot-Timothee, C. & Arman, P. (1968). Fermentative digestion in the stomach of Hippopotamus amphibius (Artiodactyla: Suiformes) and associated ciliate protozoa. Nature 218: 882-883.

Tinley, K.L. (1976). The Ecology of Tongaland. Wildlife Society of Southern Africa, Natal Branch, Durban.

Tischendorf, F. (1951). Quantitativ-vergleichende Untersuchungen zur funktionellen Morphologie des Hippopotamidengebisses. Anat. Anz.98:317-323.

Toldt Jun, K. (1915). Aeusserliche Untersuchung eines neugebornen Hippopotamus amphibius L. mit besonderer Berucksichtigung des Integuments und Bemerkungen uber die fetalen Formen der Zehenspitsenbekleidung bei Saugetieren. in: Denkschr des Kaiserl. Akad. d. Wiss. in Wien. Bd. 92: 653-*.

Tomes, J. (1850). On the blood-coloured exudation from the skin of the hippopotamus. Proc. Zool. Soc. Lond. (1850): 160-162.

Turnbull et al (1991).

- U -

Ursing, B.M. and Arnason, U. (1998 Dec 7) Analyses of mitochondrial genomes strongly support a hippopotamus-whale clade, Proceedings of the Royal Society of London - Series B: Biological Sciences, 265(1412); pp. 2251-2255.

- V-

Van Doorn, C. (1968). Einiges uber Flusspferde-ihre Zucht und Haltung im Rotterdamer Zoo. Freunde des Kolner Zoo. 11(3): 85-90.

Van Foreest, A.W., Greenwood, A.G. & Taylor, D.C. (1979). Hand rearing of an infant nile hippopotamus (Hippopotamus amphibius). AM Assoc. Zoo Vet. Ann. Proc. 170-178.

Van Hoven, W. (1974) Ciliate protozoa and aspects of the nutrition of the hippopotamus in the Kruger National Park. South African Journal of Science 70: 107-109.

Van Hoven, W. (1977). Stomach fermentation rate in the hippopotamus. South African Journal of Science 73: 216.

Van Hoven, W. (1978). Digestion physiology in the stomach complex and hindgut of the hippopotamus (Hippopotamus amphibius). South African Journal of Wildlife Research 8 (2): 59-64.

Van Niekerk, J.W. & Pienaar, U. de V. (1962). Adaptations of the immobilizing technique to the capture, marking and translocation of game animals in the Kruger National Park. Koedoe 5: 137-143.

Van Riet, W. (1969). A study of hippo populations. Afr. Wildl. 23 (3): 236-242.

Vega, I. (1995). The hippo, threatened due to ivory trade. Quercus III, Mayo.

Verheyen (1951)

Verheyen, R. (1954). Monographie ethologique de l'hippopotame. Exploration du Parc National Albert, Institut des Parcs Nationaux du Congo Belge, Bruxelles.

Vevers, G.M. (1926). Some notes on the recent birth of hippopotamus in the Gardens. Proc. Zool. Soc. London, 1097-1100.

Viljoen, P. C. (1995). Changes in number and distribution of hippopotamus (Hippopotamus amphibius) in the Sabie River, Kruger National Park, during the 1992 drought. Koedoe 38(2):115-121.

Viljoen, P. C., & H. C. Biggs. (1998). Population trends of hippopotami in the rivers of Kruger National Park, South Africa. Pages 251-279 in Behavior and Ecology of Riparian Mammals, N. Dunstone and M. L. Gorman, editors. Symposia of the Zoological Society of London, Cambridge Press, London, UK.

Viljoen, P.C. (1980). Distribution and numbers of the hippopotamus in the Olifants and Blyde Rivers. South African Journal of Wildlife Research 10: 129-132.

Vincent, J. (1962). The distribution of ungulates in Natal. Annals of the Cape Provincial Museums 2: 110-117.

Viner, A.B. (1975). The supply of minerals to tropical rivers and lakes (Uganda). In: coupling of land and water systems, ed. Hasler, A.D. Ecological studies 10. Berlin: Springer-Verlag.

Vosseler, J. (1923). Zur Fortpflanzung und Aufzucht der Nilpferde. Zoologica Palaearctica I (3): 145-155.

- W -

Walker, O. (1967). The hippo poacher. London: Cassell.

Weber, E. (1970). Hand-rearing a hippopotamus Hippopotamus amphibius at Melbourne Zoo. Int. Zoo Yb. 10: 153.

Weber, M. (1886). Studien uber Saugethiere: 3-8. Beitrage zur Anatomie von Hippopotamus amphibius. Jena: Gustav Fisher.

Weber, M. (1886). Uber den sogenannten blutigen Schweiss des Hippopotamus. in: Studien uber Saugetiere. Jena, Gustav Fischer, 9.

Weiler, P., T. De-Meulenaer, and A. Vanden-Block. (1994). Recent trends in the international trade of hippopotamus ivory. Traffic Bulletin- IUCN Wildlife Trade Monitoring Unit 15(1): 47-49.

Weinbren, B.M. (1960). Serum proteins of Hippopotamus amphibius (Linnaeus). J. Comp. Path. 70: 217-221.

Westerman, G.F. (1863). Geboorte van een Nijlpaard. Ned. Tijdschrift Dierkunde, deel (I): 1-5.

Weston, E. M. (2000). A new species of hippopotamus Hexaprotodon lothagamensis (Mammalia: Hippopotamidae) from the late Miocene of Kenya. Journal of Vertebrate Paleontology 20, no. 1: 177-185.

Weston, E.M. (2003 Dec) Evolution of ontogeny in the hippopotamus skull: using allometry to dissect developmental change, Biological Journal of the Linnean Society, 80(4); pp. 625-638.

Wetzel, H. (1973). On the nasal bot fly of the hippopotamus Rhinoestrus hippopotami (Diptera : Oestridae). Z Angew Zool. 60 (1): 5-14.

Wetzel, Von H. (1973). Zur kenntnis des Rachenfliege des Flusspferdes, Rhinoestrus hippopotami Grunberg (Diptera : Oestridae). Zeitscrift fur angewandte Zoologie 60(1): 5-14.

Wilson, J. & Littlewood, A. (1978). First year of a hand-reared hippopotamus Hippopotamus amphibius at Portland Zoo. International Zoo Yearbook, Zoological Society of London 18: 211-213.

Wright, P.G. (1964). Wild animals in the tropics. Symp. Zool. Soc. Lond. 13: 17-28.

Wright, P.G. (1972). Considerations on the hippopotamus as a diving animal. J. Physiol. (Lond) 222 (1): 56.

Wright, P.G. (1973). Observations on the adaptation of the hippopotamus to an aquatic environment. S. Afr. J. Lab. Clin. Med. 19 (1): 30-31.

Wright, P.G. (1973). Observations on the adaptation of the hippopotamus to an aquatic environment. South African Medical Journal 47 (22): 968-969.

Wright, P.G. (1975). Thermoregulation on land, and the hippopotamus. South African Medical Journal 49: 275.

Wright, P.G. (1987). Thermoregulation in the hippopotamus on land. South African Journal of Zoology 22(3):237-242.

Wright, P.G. & Luck, C.P. (1959). The skin secretion of the hippopotamus. J. Physiol. 148: 22p.

- Y -

Young, E. (1966). Nutrition of the hippopotamus (Hippopotamus amphibius ). African Wildlife 20 (2): 165-167.

- Z -

Zapico, T.A. (1999) First documentation of flehmen in a common hippopotamus (Hippopotamus amphibius), Zoo Biology, 18(5); pp. 415-420.

Zapico, T.A. (2000). The behavior of captive common hippos (Hippopotamus amphibius): the effects of environment and observations of sociosexual investigatory behaviors. M.S. Thesis. University of Denver, Denver, Colorado.

Zschokke, S. (2002) Distorted sex ratio at birth in the captive pygmy hippopotamus, Journal of Mammalogy, 83(3); pp. 674-681. (e-mail: samuel.zschokke@alumni.ethz.ch)