Elephants

Elephants constitute the largest living land animals today and their ancestors have held that status going back to the mammoths and mastodons (Rohland et al 2010). There are actually three species of extant elephants. One is found in India (Elephas maximus indicus) and two live in Africa. The two African species feature one that primarily lives in dense jungle/woodlands mostly in equatorial and west Africa  (Loxodonta Africa cyclotis) and one that sprawls across the savannahs of eastern and southern Africa (Loxodonta Africana Africana). There is a small zone of hybridization in central Africa, although this is not as common in the wild as it is in zoos.

Elephants in Asia

Elephants now number a few hundred thousand in Asia, where perhaps as many as a million used to roam. They are more common in India than any other state although Bangladesh and all the Southeast Asian countries have significant populations.

A particular feature of the lives of some elephants in Asia, as opposed to Africa, is that at one time a  significant proportion of them were used in hauling operations in remote areas (generally in the lumber industry) and some still are. (They have largely been supplanted by trucks, tractors, and specialized mechanized equipment).

A much smaller percentage today are housed in temple quarters owing, in part, to their connection with Hindu religious traditions, and for use in ceremonies and festival parades. (This is not to say they live a cosseted life, owing to factors that will be discussed shortly).  The majority that remain in India and Sri Lanka, at least, live in forest preserves, where some attempts are made to conserve them (Badola et al 2010, Baskaran, Udhayan & Desai 2010, Govindaraj 2010, Gubbi 2011).

While both Asian and forest elephants in Africa routinely prefer leaves and twigs from trees and bushes, they can switch to grasses opportunistically. One of the favorites of Asian elephants  (and forest elephants in Africa) are tree fruit, which are a major cash crop, which puts them in conflict with farmers, although ironically in Sri Lanka at least, farmers are, in consequence,  turning to citrus production, in part because elephants there do not prefer it. However, by far the  the largest cash crop in all Asia, rice, is often raided by elephant bands at harvest time (Chelliah et al 2010, Jotikapukkana,  Berg, &  Pattanavibool 2010, Santiappillai & Reed 2010, Santiapillai et al 2010). Deaths to both elephants and humans are not infrequent  in trying to scare off the elephants and preserve the harvest (Doyle et al 2010).

Elephants in Africa

While a tiny percentage of African elephants can be found in  zoos and circuses, they are overwhelmingly wild, undomesticated animals, whose  present population numbers are  in the hundreds of thousands,  down from the millions as recently as the early twentieth century.

The forest type are significantly more threatened (Bechem & Luhunu 2010, Bouche et al 2010, Lindsell, Klop & Siaka 2011, Lindsell, Klop, &  Siaka 2011, Rood, Ganie  & Nijman 2010 ) than the savannah species.

While there are some truly gigantic game parks (large relative to European and even most North American equivalents), a significant number of African elephants of both types are still living close to the park borders  or in unrestricted areas where deadly clashes with ivory poachers (Martin & Martin 2010, Messer 2010) and  natives hunter-gatherer tribespeople and indigenous farmers are a sorry fact of life (Graham et al 2010, Monney,  Dakwa, & Wiafe 2010, Ouattara, Soulemane, Nandjui & Tondoh 2010). Elephants are scarcely alone in their fatal encounters with humans and their crops. To the surprise of many who think them solely placid water plant eaters, hippos also frequently raid crops and attack and kill humans who try to fend them off (Kendall 2011).

The Diet & Water Needs of Elephants Actively Determine the Landscape

Because they are so large, elephants (particularly African Savannah elephants) are seen as much as shapers of their surroundings rather than as being shaped by their surroundings.  They do this largely through the long term effects of  their dietary  habits and water-seeking (Buitenwerf, Swemmer & Peel 2011, de Knegt et al 2011, O’Connor 2010).

Probably no other animal in the wild has its diet and travel ways  so thoroughly analyzed through the seemingly continuous analysis of their poop: largely because the droppings are  enormous and  easily located  with  fresh new specimens available to researchers at least once daily from individual elephants  so changes in thier diet can be  monitored (Codron et al 2011).  

When African savannah elephants enter a field which contains a mixture of trees, shrubs, and grasses, they may choose any one of them for forage to obtain the 400 lbs. or so of vegetation that they typically ingest every day. However the choice most likely to be adverse to the local ecosystems occurs when they forage on  the youngest trees or the mid-level shrubs.

Why would they do this? At certain times of the year (mostly during dry spells) ordinarily grass-loving  savannah elephants are likely to find  greater nutritional density in the browse from trees and bushes than from the available grasses.  The impact of savannah elephant browsing in terms of lost shade and diminished topsoil retention  can be immediate , and recovery can take quite a while, particularly if an area has already been logged commercially (Bonnell, Reyna-Hurtado, & Chapman 2011).

However, during the rainy season almost all savannah elephants switch back almost exclusively to tall lush (and presumably more nutrient-dense)  grasses, sparing the damage to trees and mid-level shrubs (Codron et al 2011).

Interestingly a number of trees and mid-level shrubs have evolved mutualisms with insects, which do not appreciably harm the trees,  whereby the insects will instead bite/harass the elephants who attempt to browse on them (Goheen & Palmer 2010).

This savannah elephant tree and mid-level brush browsing would seem to be entirely detrimental to the landscape save for perhaps three offsetting behaviors.

First, elephant dung is a tremendous source of food to burrowing insects, such as termites and dung beetles,  whose ferocious level of industry further mixes  the animal waste into the substrate (60% of which is actually relatively undigested  and serves as an excellent mulch) actually enriching the soil (Aarrestad et al 2011, Banks et al 2010, Levick 2010) and supporting  more green growth.

Second, savannah elephants are also among the first to return to areas of the veldt that have been burned via lightning storms, as they can better tolerate the coarse remnants digestively, and their subsequent droppings actually speed the recovery of the local ecosystem in terms of small grazers returning to well fertilized tender growth (Sensenig, Demment & Laca 2010).

 

Third, elephants are actually rather good at finding underground water and will use their tusks to dig up what become temporary water holes that a wide variety of other animals can subsequently share (Augustine 2010,  Franz  et al 2011). These water holes may become more permanent when the rains come, changing the landscape for the better on the largely arid savannahs. This behavior is perhaps even more altruistic than might be expected since it is almost exclusively only during water hunting forays and "well digging"  that the distracted elephants experience the otherwise rarely  successful attacks of lions on their drought-weakened young (Joubert 2006, Loveridge et al 2006). 

However, elephant overpopulation may not be entirely offset by the good the herds  do, and even abundant rainfall may not provide enough vegetation to sustain densely crowded elephant  populations (Hayward &  Zawadzka 2010).

Elephant Sensory Capacity, Intelligence & Sociability

Elephants have enormous and exceptionally well developed brains (Fowler &  Mikota2006, Hakeem et al. 2009, Hart & Pinter-Woman 2008, Shoshani, Kupsky & Marchant 2006). They possess self-awareness, recognize themselves in mirrors (Plotnik, de Waal, & Reiss 2006, Plotnik et al 2010), and demonstrate innate intelligence at least on a par with primates and are possibly as bright as dolphins (Byrne, Bates &  Moss 2009).

They have excellent hearing and communicate with a wide variety of vocalizations that seem to convey not only specific information but also mood states (Nair et al 2009, Soltis, Blowers, & Savage 2011).

While they have a keen sense of smell (Santiapillai &  Read 2010) they have terrible eyesight (Baproda et al 2010, Pettigrew et al 2010).

They are highly social and live for the most part in matriarchal groups and practice a great deal of caring for one another’s offspring  (Bates 2008, Bradshaw &  Schore 2007, De Waal 1998, De Waal 2006,  Lee 1987).

They appear to show sadness at the death of important individuals from their group, especially matriarchs (Douglas-Hamilton et al 2006).

There is much that we still do not understand about an elephant’s interior life  (Irie &  Hasegawa 2009) but we do know that it seems built for cooperativity with its own kind (Schulte 2000) and that elephants actually suffer in captivity when kept alone and isolated from other elephants (Nolen 2011).

Perhaps the most intriguing insight into elephant intelligence and willingness to cooperate is documented by Plotnik et al (2011). In an experiment conducted with the cooperation of mahouts (handlers who maintain a special lifelong welfare oversight and training  relationship with a particular elephant ) conducted in a Thai elephant refuge center, a  series of elephants were trained to pull a rope, for which they were rewarded with corn in a bowl.

The table on which the corn was placed was later rigged with a special rope with two ends, both ends of which had to be pulled in order for the table to slide within reach, and for the bowl then to be emptied of its corn.

No elephant acting on his or her own could pull the rope and make the table slide forward. Instead, the rope merely pulled out from the table.

If, in a pair of elephants one of them pulled at the rope but the other did not, the table would not slide forward, and no reward was forthcoming either.

Eventually, the elephants determined that they would somehow have to do this rope-pulling task together without any prompting from their mahouts, and did so.

Their techniques varied. Some pulled from their side using their trunk and their mouth. Others used only their trunk.

One particularly clever female managed to have her male elephant partner do all the real work, for she simply trapped the rope on her side under her foot, making her partner have to pull twice as hard on his end  to get the table to slide forward.

Elephants who saw no rope around the table eventually did not bother going forward to manage in some other way to obtain the corn. They sensed (correctly as it turned out) that without the rope there was no use wasting their time.  

However, the elephants developed a sense of patience. Some elephants who knew that another elephant might be nearby would advance  to the  usual location of the rope on  his or her side alone, and wait for another elephant to show up on the other side.

 If none showed up, the dsappointed elephant  moved away. But they had an unusually long period in which they were content to stay in hope: 45 seconds. This is much longer  than chimps and a variety of other animals tested along similar lines.

Interestingly, their intent to seek cooperative opportunities was not extinguished if no one showed up. They would try again later, and as test showed, they often succeeded.

What is the future of the elephant?

Despite the fact that encroaching human population and elephant habitat loss are very real threats, there is a significant shift among many citizens in countries where elephants are native. They have come to see that elephant conservation and resultant eco-tourism may well be a better way to make a living than subsistence farming or ivory poaching (Fischer, Muchapondwa & Sterner 2011), and in addition, that such preserves as are being set up allow  for as much space as possible about which elephants might forage and roam, especially including protection of incoming sources of water (Jotikapukkana,  Berg,  & Pattanavibool 2010), and that temple and zoo animals be allowed to live in groups where they could develop a social life or elephant culture (Vanitha, Thiyagesan, & Baskaran 2011).

Tony Stankus FSLA [email protected] Life Sciences Librarian, Science Coordinator & Professor

Editor-in-Chief Science & Technology Libraries

University of Arkansas Libraries MULN 223 E

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 Anonymous.  2011. Elephants can lend a helping trunk. Science 331 (6022): 1248-.

Aarrestad, P. A., G. S. Masunga, H. Hytteborn, M. L. Pitlagano, W. Marokane, and C. Skarpe. 2011. Influence of soil, tree cover and large herbivores on field layer vegetation along a savanna landscape gradient in Northern Botswana. Journal of Arid Environments 75 (3): 290-7.

Augustine, David J. 2010. Response of native ungulates to drought in semi-arid Kenyan rangeland. African Journal of Ecology 48 (4) (DEC 2010): 1009-20.

Badola, Ruchi, Syed Ainul Hussain, Bidyut Kumar Mishra, Bidyarani Konthoujam, Sneha Thapliyal, and Parag Madhukar Dhakate. 2010. An assessment of ecosystem services of Corbett Tiger Reserve, India. Environmentalist 30 (4).

Banks, John E., Colin Jackson, Lisa M. Hannon, Christopher M. Thomas, Albert Baya, and Laban Njoroge. 2010. The cascading effects of elephant presence/absence on arthropods and an Afrotropical thrush in Arabuko-Sokoke forest, Kenya. African Journal of Ecology 48 (4) (DEC 2010): 1030-8.

Bapodra, Priya, Tim Bouts, Paul Mahoney, Sally Turner, Ayona Silva-Fletcher, and Michael Waters. 2010. Ultrasonographic anatomy of the Asian elephant (Elephas maximus) eye. Journal of Zoo and Wildlife Medicine 41 (3) (SEP 2010): 409-17.

Baskaran, N., A. Udhayan, and Ajay A. Desai. 2010. Status of the Asian elephant population in Mudumalai Wildlife Sanctuary, Southern India. Gajah 32: 6-13.

Bates, L.A. et al. 2008. Do elephants show empathy? Journal of Conscious Studies. 15: 204-25.

Bechem, M., and S. Luhunu. 2010. Elephant surveys in Central African forests. Pachyderm (47): 111-4.

Bonnell, T. R., R. Reyna-Hurtado, and C. A. Chapman. 2011. Post-logging recovery time is longer than expected in an East African tropical forest. Forest Ecology and Management 261 (4): 855-64.

Bouche, Philippe, Pierre-Cyril Renaud, Philippe Lejeune, Cedric Vermeulen, Jean-Marc Froment, Alfred Bangara, Okclefort Fiongai, Antoine Abdoulaye, Raymond Abakar, and Mike Fay. 2010. Has the final countdown to wildlife extinction in northern Central African Republic begun? African Journal of Ecology 48 (4) (DEC 2010): 994-1003.

Bradshaw, G.A., and Schore, A.N. 2007. How elephants are opening doors: Developmental neuroethology, attachment and social context. Ethology 113: 426-36.

Byrne, R.W., Bates, L.A., and Moss, C.J. 2009. Elephant cognition in primate perspective. Comp. Cog. Behav. Rev. 4: 1-15.

Buitenwerf, Robert, Anthony M. Swemmer, and Mike J. S. Peel. 2011. Long-term dynamics of herbaceous vegetation structure and composition in two African savanna reserves. Journal of Applied Ecology 48 (1) (FEB 2011): 238-46.

Chelliah, K., G. Kannan, S. Kundu, N. Abilash, A. Madhusudan, N. Baskaran, and R. Sukumar. 2010. Testing the efficacy of a chili-tobacco rope fence as a deterrent against crop-raiding elephants. Current Science (Bangalore) 99 (9) (NOV 10 2010): 1239-43.

Codron, Jacqueline, Daryl Codron, Julia A. Lee-Thorp, Matt Sponheimer, Kevin Kirkman, Kevin J. Duffy, and Judith Sealy. 2011. Landscape-scale feeding patterns of African elephant inferred from carbon isotope analysis of feces. Oecologia (Berlin) 165 (1) (JAN 2011): 89-99.

de Knegt, H. J., F. van Langevelde, A. K. Skidmore, A. Delsink, R. Slotow, S. Henley, G. Bucini, et al. 2011. The spatial scaling of habitat selection by African elephants. Journal of Animal Ecology 80 (1): 270-81.

DeFries, Ruth, Krithi K. Karanth, and Sajid Pareeth. 2010. Interactions between protected areas and their surroundings in human-dominated tropical landscapes. Biological Conservation 143 (12, Sp. Iss. SI) (DEC 2010): 2870-80.

De Waal, F.B.M. 2008. Putting the altruism back into altruism: The evolution of empathy. Annu. Rev. Psychol. 59(4): 1-4.22.

De Waal, F.B.M. 1996. Good natured. Cambridge, MA: Harvard University Press.

Douglas-Hamilton, I., Bhalla, S., Wittemyer, G., and Vollrath, F. 2006. Behavioral reactions of elephants towards a dying or deceased matriarch. Appl. Anim. Behav. Sci. 100: 87-102.

Dow, T. L., W. Roudebush, F. N. Parker, and J. L. Brown. 2011. Influence of age and gender on secretion of anti-Mullerian hormone in Asian (Elephas maximus) and African (Loxodonta africana) elephants. Theriogenology 75 (4): 620-7.

Doyle, Sarah, Melissa Groo, Christie Sampson, Melissa Songer, Marshall Jones, and Peter Leimgruber. 2010. Human-elephant conflict - What can we learn from the news? Gajah 32 : 14-20.

Fischer, C., E. Muchapondwa, and T. Sterner. 2011. A bio-economic model of community incentives for wildlife management under CAMPFIRE. Environmental & Resource Economics 48 (2): 303-19.

Fowler, M.E., and Mikota, S.K. Eds. 2006. Biology, medicine and surgery of elephants. Ames, IA: Wiley-Blackwell.

Franz, Mathias, Stephanie Kramer-Schadt, Werner Kilian, Christian Wissel, and Juergen Groeneveld. 2010. Understanding the effects of rainfall on elephant-vegetation interactions around waterholes. Ecological Modelling 221 (24) (DEC 15 2010): 2909-17.

Goheen, Jacob R., and Todd M. Palmer. 2010. Defensive plant-ants stabilize megaherbivore-driven landscape change in an African savanna. Current Biology 20 (19) (OCT 12 2010): 1768-72.

Gordon, J.A. 1966. Elephants do think. African Wildlife 20: 75-9.

Govindaraj, Kannan. 2010. Solitary behaviour of female elephants in Idukki Wildlife Sanctuary, Kerala. Gajah 32 : 40-1.

Graham, Maximilian D., Benedikt Notter, William M. Adams, Phyllis C. Lee, and Tobias Nyumba Ochieng. 2010. Patterns of crop-raiding by elephants, Loxodonta africana, in Laikipia, Kenya, and the management of human-elephant conflict. Systematics and Biodiversity 8 (4) (2010): 435-45.

Gubbi, S. 2011. India proposes new strategies to conserve elephants. Oryx 45 (1): 14-5.

Hakeem, A.Y. et al. 2009. Von economo neurons in the elephant brain. Anat. Rec. 292: 242-48.

Hart, B.L., Hart, L.A. and Pinter-Wollman, N. 2008. Large brains and cognition: Where do elephants fit in? Neurosci. Biobehav. Rev. 32: 86-98.

Hayward, Matt W., and Bernadetta Zawadzka. 2010. Increasing elephant Loxodonta africana density is a more important driver of change in vegetation condition than rainfall. Acta Theriologica 55 (4) (OCT 2010): 289-99.

Hock, K., K. L. Ng, and N. H. Fefferman. 2010. Systems approach to studying animal sociality: Individual position versus group organization in dynamic social network models. Plos One 5 (12).

Irie, N. and Hasegawa, T. 2009. Elephant psychology: What we know and what we would like to know. Jpn Psychology Res. 51: 177-81.

Jotikapukkana, Sukanda, Ake Berg, and Anak Pattanavibool. 2010. Wildlife and human use of buffer-zone areas in a wildlife sanctuary. Wildlife Research 37 (6) (2010): 466-74.

Joubert, D. 2006. Hunting behavior of lions (Panthera leo) on elephants (Loxodonta Africana) in the Chobe National Park, Botswana). African Journal of Ecology 44: 279-281.

Kendall, C. I. 2011. The spatial and agricultural basis of crop raiding by the vulnerable common Hippopotamus hippopotamus amphibius around Ruaha National Park, Tanzania. Oryx 45 (1): 28-34.

Kuntner, Matjaz, Laura J. May-Collado, and Ingi Agnarsson. 2011. Phylogeny and conservation priorities of Afrotherian mammals (Afrotheria, mammalia). Zoologica Scripta 40 (1) (JAN 2011): 1-15.

Lair, R.C. 1997. Gone Astray: The care and management of the Asian elephant in domesticity. Rome: Food and Agricultural Organization of the United Nations.

Lee, P.C. 1987. Allomothering among African elephants. Animal Behavior. 35: 278-91.

Levick, Shaun R., Gregory P. Asner, Ty Kennedy-Bowdoin, and David E. Knapp. 2010. The spatial extent of termite influences on herbivore browsing in an African savanna. Biological Conservation 143 (11) (NOV 2010): 2462-7.

Lindsell, J. A., E. Klop, and A. M. Siaka. 2011. The impact of civil war on forest wildlife in West Africa: Mammals in Gola Forest, Sierra Leone. Oryx 45 (1): 69-77.

Loveridge, A. J.E. Hunt, F. Murindagomo & D.W. Macdonald. 2006. Influence of drought on predation of elephant calves by lions. Journal of Zoology 270 (3): 523-530.

Martin, E., and C. Martin. 2010. Russia's mammoth ivory industry expands: What effect on elephants? Pachyderm (47): 26-35.

Messer, Kent D. 2010. Protecting endangered species: When are shoot-on-sight policies the only viable option to stop poaching? Ecological Economics 69 (12) (OCT 15 2010): 2334-40.

Monney, Kweku Ansah, Kwaku Brako Dakwa, and Edward Debrah Wiafe. 2010. Assessment of crop raiding situation by elephants (Loxodonta africana cyclotis) in farms around Kakum Conservation Area, Ghana. International Journal of Biodiversity and Conservation 2 (9): 243-9.

Moss, C. 1988. Elephant memories: Thirteen years in the life of an elephant family. New York: Fawcett Columbine.

Murphree, R., J. V. Warkentin, J. R. Dunn, W. Schaffner, and T. F. Jones. 2011. Elephant-to-human transmission of tuberculosis, 2009. Emerging Infectious Diseases 17 (3): 366-71.

Nair, S., Balakrishnan, R., Seelamantula, C.S., and Sukumar, R. 2009. Vocalization of wild Asian elephants (Elephas maximus): Structural classification and social context. Journal of Acoust. Soc. Am. 126: 2768-78.

Noë, R. 2006. Cooperation experiments: Coordination through communication versus acting apart together. Animal Behavior 71: 1-18.

Nolen, R. S. 2011. Captive elephant welfare study under way. JAVMA-Journal of the American Veterinary Medical Association 238 (3): 270-.

O'Connor, Timothy G. 2010. Transformation of riparian forest to woodland in Mapungubwe National Park, South Africa, between 1990 and 2007. Austral Ecology 35 (7) (NOV 2010): 778-86.

Ouattara, F. A., O. Soulemane, A. Nandjui, and E. J. Tondoh. 2010. State of marauding and damage to crops with elephants in the west sector Djouroutou in south-west of the Parc National De Tai (Cote d'ivoire). Pachyderm(47): 36-44.

Pettigrew, John D., Adhil Bhagwandin, Mark Haagensen, and Paul R. Manger. 2010. Visual acuity and heterogeneities of retinal ganglion cell densities and the Tapetum lucidum of the African elephant (Loxodonta africana). Brain Behavior and Evolution 75 (4) (2010): 251-61.

Plotnik, J. M., de Waal, F.B.M., Moore, D. 3^rd., and Reiss, D. 2010. Self-recognition in the Asian elephant and future directions for cognitive research with elephants in zoological settings. Zoo Biol. 29: 179-91.

Plotnik, J. M., de Waal, F.B.M., and Reiss, D. 2006. Self-recognition in an Asian elephant. Proc. Natl. Acad. Sci. USA 103: 17053-57.

Plotnik, J.M., R. Lair, W. Suphachoksahkun & F.B.M. de Waal. 2011. Elephants know when they need a helping trunk in a cooperative task. Proc. Natl. Acad. Sci. USA 108 (12):  5116-5121.

Poilecot, Pierre, Etienne Bemadjim N'Gakoutou, and Nicolas Taloua. 2010. Evolution of large mammal populations and distribution in Zakouma National Park (Chad) between 2002 and 2008. Mammalia 74 (3) (SEP 2010): 235-46.

Poole, J.H., and Granli, P.K.  2011. Signals, gestures and behaviors of African elephants. The Amboseli elephants: A long-term perspective on a long-lived mammal, Eds Moss, C.J. and Croze, H.J. Chicago: University of Chicago Press.

Poole, J. H. and Moss, C. 2008. Elephants and ethics, Eds Wemmer, C., and Christen, C.A. Baltimore: Johns Hopkins University Press: 69-98.

Poole, J. 1996. Coming of age with elephants. New York: Hyperion

Rohland, Nadin, David Reich, Swapan Mallick, Matthias Meyer, Richard E. Green, Nicholas J. Georgiadis, Alfred L. Roca, and Michael Hofreiter. 2010. Genomic DNA sequences from mastodon and woolly mammoth reveal deep speciation of forest and savanna elephants. PLoS Biology 8 (12) (DEC 2010): ArteNo.:e1000564.

Rood, Ente, Abdullah A. Ganie, and Vincent Nijman. 2010. Using presence-only modelling to predict Asian elephant habitat use in a tropical forest landscape: Implications for conservation. Diversity and Distributions 16 (6) (NOV 2010): 975-84.

Santiapillai, Charles, and Bruce Read. 2010. Would masking the smell of ripening paddy-fields help mitigate human-elephant conflict in Sri Lanka? Oryx 44 (4) (OCT 2010): 509-11.

Santiapillai, Charles, S. Wijeyamohan, Ganga Bandara, Rukmali Athurupana, Naveen Dissanayake, and Bruce Read. 2010. An assessment of the human-elephant conflict in Sri Lanka. Ceylon Journal of Science Biological Sciences 39 (1): 21-33.

Sarker, A. H. M. Raihan, and Eivin Roskaft. 2010. Human attitudes towards conservation of Asian Elephants (Elephas maximus) in Bangladesh. International Journal of Biodiversity and Conservation 2 (10): 316-27.

Schulte, B.A. 2000. Social structure and helping behavior in captive elephants. Zoo Biol. 19: 447-59.

Seed, A., and R. Byrne. 2010. Animal tool-use. Current Biology 20 (23): R1032-9.

Seidensticker, J. and Lumpkin, S. Eds. 1990. Asian elephants: John F. Eisenberg, George M. McKay, and John Seidensticker. Washington D.C: Friends Natl Zoo and the Zoolog Park.

Sensenig, Ryan L., Montague W. Demment, and Emilio A. Laca. 2010. Allometric scaling predicts preferences for burned patches in a guild of East African grazers. Ecology (Washington D C) 91 (10) (OCT 2010): 2898-907.

Shannon, Graeme, Bruce R. Page, Kevin J. Duffy, and Rob Slotow. 2010. The ranging behaviour of a large sexually dimorphic herbivore in response to seasonal and annual environmental variation. Austral Ecology 35 (7) (NOV 2010): 731-42.

Shoshani, J. Kupsky, W.J. and Marchant, G.H. 2006. Elephant brain, Part 1: Gross morphology, functions, comparative anatomy and evolution. Brain Res. Bull. 70: 124-57.

 Shoshani, J. 1998. Understanding proboscidean evolution: A formidable task. Trends Ecol. Evol. 13: 480-87.

Soltis, J., T. E. Blowers, and A. Savage. 2011. Measuring positive and negative affect in the voiced sounds of African elephants (Loxodonta africana). Journal of the Acoustical Society of America 129 (2): 1059-66.

Sukumar, R. 1989. The Asian elephant: Ecology and management. Cambridge, UK: Cambridge University Press.

Suwattana, D., J. Prasupphachok, S. Kanchanapangka, and W. Koykul. 2011. Tetranucleotide microsatellite markers for molecular testing in Thai domestic elephants (Elephas maximus indicus). Strategic Management Journal 32 (3): 405-9.

Teren, G., and N. Owen-Smith. 2010. Elephants and riparian woodland changes in the Linyanti Region, Northern Botswana. Pachyderm(47): 18-25.

Theuerkauf, J., and R. Gula. 2010. Towards standardisation of population estimates: Defecation rates of elephants should be assessed using a rainfall model. Annales Zoologici Fennici 47 (6): 398-402.

Valeix, M., H. Fritz, R. Sabatier, F. Murindagomo, D. Cumming, and P. Duncan. 2011. Elephant-induced structural changes in the vegetation and habitat selection by large herbivores in an African savanna. Biological Conservation 144 (2): 902-12.

Vanak, Abi Tamim, Maria Thaker, and Rob Slotow. 2010. Do fences create an edge-effect on the movement patterns of a highly mobile mega-herbivore? Biological Conservation 143 (11) (NOV 2010): 2631-7.

Vanitha, V., K. Thiyagesan, and N. Baskaran. 2011. Social life of captive Asian elephants (Elephas maximus) in Southern India: Implications for elephant welfare. Journal of Applied Animal Welfare Science 14 (1): 42-58.

Williams, J.H. 1950. Elephant Bill. London: Rupert Hart-Davis.

Wrege, Peter H., Elizabeth D. Rowland, Bruce G. Thompson, and Nikolas Batruch. 2010. Use of acoustic tools to reveal otherwise cryptic responses of forest elephants to oil exploration. Conservation Biology 24 (6) (DEC 2010): 1578-85.

Young, K. D., and R. J. Van Aarde. 2011. Science and elephant management decisions in South Africa. Biological Conservation 144 (2): 876-85.

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