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Elephants playing together in jungle – stock photo, courtesy of Getty Images. Taken in a private game reserve bordering the Kruger National Park, South Africa.

Evidence suggests that the Cape south coast of South Africa supported a considerable elephant population during the Pleistocene.  Although at times during that period, the enormous Palaeo-Agulhas Plain may have been inundated with water, it was often an open sandy environment, filled with dunes and beaches.  The recent paper by Charles Helm, Martin Lockley, Lizette Moolman, Hayley Cawthra, Jan De Vynck, Mark Dixon, Willo Stear and Guy Thesen describes sites containing 400,000- to 80,000-year-old elephant footprints.  Those trace fossils have survived on the significantly smaller remnants of that ancient plain today.  

Figure 1. (colour online) Map of the Cape south coast of South Africa, indicating the main study area and the study site. With permission from Cambridge University Press.

Other proboscidean species such as mammoths and mastodons were alive in that same timeframe in other parts of the world, but no fossil evidence of any earlier type of elephant has been found in South Africa younger than 400,000 years.  The authors, as mentioned in part one of this two-part series, believe the fossil footprints were made by the African savanna elephant (Loxodonta africana).  

But that’s not the end of the story.  

As wonderful and exciting as the many trace fossils are, the plight of the elephants in that area is—in direct contrast—profoundly alarming.  The entire continent may have been home to about 20 million elephants before Europeans colonized it, perhaps 3000 of them roaming the Cape south coast.  In less than 400 years, their numbers dropped precipitously. A mere 30-50 elephants existed at the beginning of the 20th century.  In 1925, there were just 12 left.  And today? 

Only one.

For an intelligent, social species known to live in herds, a single elephant living in the wild is absolutely heartbreaking.  She is 42 years old, and like all other “Knysna elephants,” she lives within the Knysna forest.  She tends to avoid open areas.

Screenshot of a tweet from South African National Parks, featuring Oupoot, the last known Knysna elephant.

Screenshot from Google Maps indicating a part of South Africa that includes the town of Knysna (and its surrounding forests).

She is not an African forest elephant (Loxodonta cyclotis) despite her habitat.  The authors contend that the Knysna elephants were forced into the local forests to avoid human contact.  In a 2019 paper, lead author and wildlife ecology scientist at South Africa National Parks, Lizette Moolman and her colleagues describe records of humans shooting elephants in the southern Cape as far back as 1755.  South Africa as a whole lost most of its elephant population due to increases in the number of humans and their farms, as well as the ivory trade.  

Once decimated, the Knysna elephants struggled to survive.  They were given protection in 1908, and in 1994, new elephants from other areas were brought in, but despite all efforts, nothing worked.  Various theories have been proposed—from a suboptimal forest diet, to reproductive problems, to the increased vulnerability of a dwindling population—but the authors suggest we have much more to learn before we understand the reasons behind it.

“So the question is,” Lizette explained in an email, “would elephants have preferred the current Knysna elephant range if there were no anthropogenic threats and barriers? And is the current range the type of environment that will enhance a population or limit its fitness? We do not have the answer to [these questions] but can make assumptions based on historic distributions of elephant. What we concluded in our research is that there is not enough evidence or data to support any hypothesis fully (or in other words, to take any of the hypotheses off the table) and we theorised that these elephants probably declined due to synergistic (a mix of) workings of sub-optimal habitat quality, killings (people shooting them) and demographic stochasticity (i.e. the population was so tiny by the time they were declared protected that any one chance event would have caused their demise).”

Understanding why this elephant population fell so drastically and why it couldn’t recover will help scientists decide how to proceed in the near future.

“The fact that we do not know for sure why the population declined, together with the fact that the elephant range is not fenced off and surrounded by farmland and close to towns and that the elephant range falls across 4 different landowners land, makes this a complex issue,” she continued.

South African National Parks (SANParks) and Nelson Mandela University (NMU) are working with everyone who might be impacted by “free roaming elephants in the Knysna forest.” Their proposed solutions keep the lone surviving elephant’s health and best interests in mind, but they face a unique challenge. The Knysna population is one elephant. Do they have time to find a workable answer? One hopes that the intense amount of effort, care and study conducted by those in South Africa will do just that.

A sloping aeolianite surface in the Goukamma Nature Reserve, showing elephant tracks and two elongated grooves interpreted as possible trunk-drag impressions from ancient African elephants in the Cape south coast, courtesy of Dr. Charles Helm and with permission from Cambridge University Press.

Another enormous thank you to Lizette Moolman, Mpilo Nxumalo, Hayley Cawthra and Charles Helm for their insight, their time, and their help! It was a great pleasure connecting with all of you!


  1. Chapman Poulsen, Zoë. Forests of South Africa. Botanical Society of South Africa. July 27, 2020.
  2. Elephant Voices. Elephants are socially complex. Elephantvoices.org.
  3. Grobler, Riaan. Meet Oupoot, the only elephant left in the entire Knysna forest. News24. February 15, 2019.
  4. Helm, C., Lockley, M., Moolman, L., Cawthra, H., De Vynck, J., Dixon, M., . . . Thesen, G. (2021). Morphology of Pleistocene elephant tracks on South Africa’s Cape south coast and probable elephant trunk-drag impressionsQuaternary Research, 1-15. doi:10.1017/qua.2021.32
  5. Moolman L, Ferreira SM, Gaylard A, Zimmerman D, Kerley GIH. The decline of the Knysna elephants: Pattern and hypotheses. S Afr J Sci. 2019;115(3/4), Art. #4805, 7 pages. https://doi.org/10.17159/ sajs.2019/4805
  6. SABC News. Knysna elephant alive and well. February 9, 2019.
  7. South African National Biodiversity Institute. Fynbos Biome. PlantZAfrica.