LOLLDAIGA HILLS RESEARCH PROGRAMME NEWSLETTER

Tom Butynski & Yvonne de Jong

May–June 2018 (Issue 19)

Cover photograph: Adult Maned (or Crested) Rat Lophiomys imhausi, Lolldaiga Hills Ranch, Kenya. Photograph by Stephanie Higgins.

New to lolldaiga.com • Blog: Research on Lolldaiga Hills’ poisonous ---the Maned Rat • New publication: Primates of East Africa Pocket Identification Guide • New publication: Geographic range, , and conservation of the Mount Kilimanjaro Guereza Colobus Monkey (Primates: Cercopithecidae: Colobus) • New publication: Sleep patterns, daytime , and the evolution of diurnal sleep site selection in lorisiforms

Blog Research on Lolldaiga Hills’ poisonous rodent---the Maned Rat Sara B. Weinstein, Department of Biology, University of Utah and National Zoological Park, Smithsonian Institution

Many defend against predation by co-opting the chemical defenses of their food. Although this poison sequestration behaviour has been most studied in butterflies, similar behaviours occur in other , amphibians and even birds (Savitzky et al. 2012, Nishida 2014). Remarkably, although many consume poisonous vegetation (Dearing et al. 2005), the defensive use of sequestered toxins is rare among mammals. The European Hedgehog Erinaceus europaeus coats its quills with noxious chemicals (Brodie 1977) and the Maned (or Crested) Rat Lophiomys imhausi anoints its fur with compounds from the Poison Arrow Tree (Kingdon et al. 2012).

Adult Maned (or Crested) Rats Lophiomys imhausi. Photograph by Stephanie Higgins.

The Maned Rat defends itself with cardiac glycosides from the Poison Arrow Tree. These large (adult body weight = 600–1200 g) chew the bark and leaves of the Poison Arrow Tree, releasing ouabain, acovenoside, and other cardiac glycosides which they then lick onto specialized hairs on their sides. When disturbed, they expose these areas to reveal warning coloration and the toxin-anointed hairs (Kingdon et al. 2012). The ecology, behaviour and distribution of the Maned Rat remain poorly known (Kingdon 1974, Happold 2013). These rodents are, however, repeatedly observed around Mount Kenya, including numerous sightings on Lolldaiga Hills Ranch (see past blog post on www.lolldaiga.com/maned-rat-lolldaiga). Lolldaiga’s Maned Rat population is being monitored, sampled, and studied in order to better understand the behaviour and ecology of

this enigmatic rodent. 1

Adult Maned Rat displaying black and white warning colours and presenting its poison laden hairs on Lolldaiga Hills Ranch, central Kenya. Photograph by S. B. Weinstein.

Trapping of Maned Rats has focused on the riverine forest along the Timau River on the south end of the Ranch. Trapped rats are examined for poison levels, parasite loads, and behaviour. All tested animals had cardiac glycosides in their specialized hairs, although poison levels varied among individuals.

Team ‘Poison Rat’ sets and monitors trap lines along the Timau River, south Lolldaiga Hills Ranch, central Kenya. Photograph by Stephanie Higgins.

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In most animals, feeding on even a small amount of the Poison Arrow Tree leads to death, as all parts of the plant (except the ripe fruit) are highly toxic. We have filmed Maned Rats chewing and applying masticated Poison Arrow Tree bark and leaves to their fur, thereby confirming previous observations of this behaviour [see link to Kingdon (2012): https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3248729/]. Using motion activated infra-red cameras, we have also observed other behaviours, including mating, grooming and social interactions.

Above: Specialized hairs collected from a wild Maned Rat. The unusual porous structure of these hairs wicks and retains toxins, which stain the hairs brown. Magnification 40x. Photograph by S. B. Weinstein.

Above right: Cardiac glycosides are extracted with water. The difference between ’poison’ (’SBW1’) and ’regular’ (’SBW2’) hair is visible, even before chemical analyses. Chemical composition and concentration in hairs and faeces is analyzed using liquid chromatography mass spectrometry at the International Centre for Physiology and Ecology (ICIPE) in Nairobi. Photograph by S. B. Weinstein.

Louse collected from a Maned Rat. Magnified 40x. Photograph by S. B. Weinstein.

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Link to chewing video: https://www.youtube.com/watch?v=kbYvPVNmdWw Link to feeding video: https://youtu.be/D1h_NBejN1Y

Sampled Maned Rats yielded lice and fleas, suggesting that these ectoparasites, like their hosts, have unique adaptations for managing toxin exposure and its effects. Interestingly, ticks have not been observed on Maned Rats.

This research project will continue to study the Maned Rat on Lolldaiga Hills Ranch, while expanding sampling to other areas to further examine the natural history and mechanisms that facilitate toxin sequestration in this unusual rodent.

Adult Maned Rat being released at its site of capture following sampling and behavioural observations on Lolldaiga Hills Ranch. Photograph by S. B. Weinstein.

References: Brodie, E. D. 1977. Hedgehogs use toad venom in their own defence. Nature 268(5621): 627––628. Dearing, M. D., Foley, W. J. & McLean, S. 2005. The influence of plant secondary metabolites on the nutritional ecology of herbivorous terrestrial vertebrates. Annual Review of Ecology, Evolution, and Systematics 36(1): 169–189. Happold, D. C. D. 2013. Lophiomys imhausi Maned Rat (Crested Rat). In: Mammals of Africa. Volume III: Rodents, Hares and Rabbits, Happold, D. C. D., ed., pp. 214–215. Bloomsbury, London. Kingdon, J. 1974. East African Mammals. Volume IIB: Hares and Rodents. University of Chicago Press, Chicago. Kingdon, J., Agwanda, B., Kinnaird, M., O'Brien, T., Holland, C., Gheysens, T., Boulet-Audet, M. & Vollrath, F. 2012. A poisonous surprise under the coat of the African crested rat. Proceedings of the Royal Society of London B: Biological Sciences 279(1729): 675–680. Nishida, R. 2014. Chemical ecology of insect–plant interactions: ecological significance of plant secondary metabolites. Bioscience, Biotechnology, and Biochemistry 78(1): 1–13. Savitzky, A. H., Mori, A., Hutchinson, D. A., Saporito, R. A., Burghardt, G. M., Lillywhite, H. B. & Meinwald, J. 2012. Sequestered defensive toxins in tetrapod vertebrates: principles, patterns, and prospects for future studies. Chemoecology 22(3): 141–158.

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Global Wildlife Conservation Primates of East Africa Pocket Identification Guide Yvonne de Jong & Tom Butynski Illustrations & Design Stephen Nash 2018

70 taxa • 32 distribution maps • 73 drawings

Get a copy at the IPS Congress (Nairobi) or go to www.wildsolutions.nl/pg. Soon available at www.nhbs.com

Eastern Africa Primate Diversity and Conservation Program, www.wildsolutions.nl Lolldaiga Hills Research Programme, www.lolldaiga.com

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Request for information Hyaena Distribution Mapping Project 2018 IUCN/SSC Hyaena Specialist Group Range maps for the Spotted Hyaena, Striped Hyaena, Brown Hyaena, and Aardwolf have not been updated since 1998! Over the next 7 months, we will attempt to compile all verifiable observations of hyaenas (and their spoor) since 2010, across their ranges. These new range maps will be the first step towards a new status survey and conservation action plan for the Hyaenidae.

Guiding research objectives for this project are:

1) to provide an updated global distribution map for all four hyaena species 2) to compare the distribution maps to the 1998 Action Plan 3) to collate all known density information 4) to identify key threats for each species and/or geographic region

Adult Spotted Hyaena Crocuta crocuta, El Karama Ranch, Laikipia, Kenya. Photograph by Yvonne de Jong.

You can help by:

Submitting data

• If you have 50 of fewer observations, you can complete the attached data spreadsheet and email it to Andrew Jacobson: [email protected]. • Alternatively, you can post observations at iNaturalist.org, and they will be automatically added to our project there (https://www.inaturalist.org/projects/hyaena-distribution-mapping-project-2018). There are now over 1000 observations included in the project. • Finally, if you have a large number of potential data points from a research project or camera trapping survey please email Andrew and we can determine the best way to handle your data.

Providing additional contacts If you know of people and/or organizations that might have useful data, and you are willing to share their contact information with us, we would be very grateful. We are especially interested in gaining contacts for observations outside of protected areas, and in countries outside of eastern and southern Africa. Potential contacts can be sent to Steph Dloniak: [email protected] (please note that we have sent out a request to all SSC members working in hyaena range countries).

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Publications and Reports

Publication

• Butynski, T. M. & De Jong, Y. A. Geographic range, taxonomy, and conservation of the Mount Kilimanjaro Guereza Colobus Monkey (Primates: Cercopithecidae: Colobus). Hystrix. http://www.italian-journal-of- mammalogy.it/Geographic-range-taxonomy-and-conservation-of-the-Mount-Kilimanjaro-guereza- colobus,82059,0,2.html • De Jong, Y. A. & Butynski, T. M. Primates of East Africa. Pocket Identification Guide. Global Wildlife Conservation, Tropical Pocket Guide Series. http://www.wildsolutions.nl/pg/ • De Jong, Y. A., d’Huart, J. P. & Butynski, T. M. Biogeography of the Desert Warthog Phacochoerus aethiopicus (Pallas, 1766) and Common Warthog Phacochoerus africanus (Gmelin, 1788) in the Horn of Africa. Poster: 12th International Symposium on Wild Boar and Other Suids Lázně Bělohrad, Czech Republic, 4-7 September 2018. • Svensson, M. S, Nekaris, K. A. I., Bearder, S. K., Bettridge, C., Butynski, T. M., Cheyne, S. M., Das, N., De Jong, Y. A., Luhrs, A. M., Luncz, L., Maddock, S. T., Perin, A., Pimley, E., Poindexter, S., Reinhardt, K. D., Spaan, D., Stark, D. J., Starr, C. & Nijman, V. Sleep patterns, daytime predation, and the evolution of diurnal sleep site selection in lorisiforms. American Journal of Physical Anthropology 166: 563–577. https://www.researchgate.net/publication/326300424_Sleep_patterns_daytime_predation_and_the_evolution_of_di urnal_sleep_site_selection_in_lorisiforms

In press

• Butynski, T. M. & De Jong, Y. A. Primates of Africa’s coastal deltas and their conservation. In: Primates in Flooded Habitats: Ecology and Conservation. Barnett, A. A., Matsuda, I. & Nowak, K., eds. Cambridge University Press, Cambridge, UK. • Cunneyworth, P., De Jong, Y. A., Butynski, T. M. & Perkin, A. W. IUCN/SSC Red List assessment for Peter's Angolan Colobus Colobus angolensis palliatus. The IUCN Red List of Threatened Species 2018. IUCN/SSC, Gland, Switzerland. • De Jong, Y. A. & Butynski, T. M. IUCN/SSC Red List assessments for 55 taxa of African primates. The IUCN Red List of Threatened Species 2018. IUCN/SSC, Gland, Switzerland. • Rovero, F., Davenport, T. R. B., De Jong, Y. A. & Butynski, T. M. IUCN/SSC Red List assessment for Sharpe's Angolan Colobus Colobus angolensis palliatus. The IUCN Red List of Threatened Species 2018. IUCN/SSC, Gland, Switzerland.

Submitted

• Kenworthy, S. P., Butynski, T. M., De Jong, Y. A., de Kort, S. & Bettridge, C. M. Intraspecific structural differences in loud calls of the Small-eared Greater Galago (Otolemur garnettii). • Kivai, S. M., Butynski, T. M., De Jong, Y. A., King, J., Loyola, L. C., Mbora, D. N. M. & Ting, N. Tana River Red Colobus P. rufomitratus / Critically Endangered. IUCN/SSC Red Colobus Action Plan. IUCN/SSC, Gland, Switzerland. • Svensson, M. S., Butynski T. M., De Jong, Y. A., Bearder, S. K. & Nijman, V. Geographic variation in the loud call of the Northern Lesser Galago (Galago senegalensis).

In preparation

• Butynski, T. M., De Jong, Y. A. & Brugieri, D. African colobine conservation. In: The Colobines: Natural History, Behaviour and Ecological Diversity. Matsuda, I, Grueter, C. C. & Teichroeb, J. A., eds. Cambridge University Press, Cambridge, UK. • De Jong, Y. A. & Butynski, T. M. Biogeography, taxonomy and phenotypic clines of Olive Baboon Papio anubis and Yellow Baboon Papio cynocephalus in Kenya and Tanzania. In: Baboons. Wallis, J., ed. Cambridge University Press, Cambridge, UK. • De Jong, Y. A. & Butynski, T. M. Vocal pattern of the advertisement call of the Somali Lesser Galago Galago gallarum Thomas, 1901. • De Jong, Y. A., d’Huart, J. P. & Butynski, T. M. Biogeography of the Desert Warthog Phacochoerus aethiopicus (Pallas, 1766) and Common Warthog Phacochoerus africanus (Gmelin, 1788) in the Horn of Africa.

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New publication Geographic range, taxonomy, and conservation of the Mount Kilimanjaro Guereza Colobus Monkey (Primates: Cercopithecidae: Colobus) Thomas M. Butynski & Yvonne A. de Jong, Lolldaiga Hills Research Programme

Abstract The Mount Kilimanjaro Guereza Colobus Monkey is endemic to northern Tanzania and southern Kenya, occurring on and near Mount Kilimanjaro/Mount Meru. Currently referred to as “Colobus guereza caudatus Thomas 1885”, this monkey is geographically very isolated, and phenotypically distinct from all other taxa of guereza monkeys. As such, application of the “Phylogenetic Species Concept” resurrects the Mount Kilimanjaro guereza to specific rank as Colobus caudatus. The geographic range of C. caudatus is small (ca. 4,030 km2) and in decline, as is the number of individuals and area of habitat. Colobus caudatus qualifies as an IUCN Red List globally “Endangered” species, as a nationally “Endangered” species in Tanzania, and as a nationally “Critically Endangered” species in Kenya. Colobus caudatus is Kenya’s most threatened species of primate. Recommendations for research and conservation actions are provided.

Full publication on: http://www.wildsolutions.nl/colobuscaudatus/

Left: Adult male Mount Kilimanjaro Guereza Colobus Monkey Colobus caudatus, Usa River, central north Tanzania. Photograph by Yvonne de Jong and Tom Butynski.

Adult Mount Kilimanjaro Guereza Colobus Monkey Colobus caudatus, Kitobo Forest Reserve, southeastern Kenya. Photograph by Yvonne de Jong and Tom Butynski.

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New publication Sleep patterns, daytime predation, and the evolution of diurnal sleep site selection in lorisiforms Svensson, M. S, Nekaris, K. A. I., Bearder, S. K., Bettridge, C., Butynski, T. M., Cheyne, S. M., Das, N., De Jong, Y. A., Luhrs, A. M., Luncz, L., Maddock, S. T., Perin, A., Pimley, E., Poindexter, S., Reinhardt, K. D., Spaan, D., Stark, D. J., Starr, C. & Nijman, V.

Abstract Objectives: Synthesize information on sleep patterns, sleep site use, and daytime predation at sleep sites in lorisiforms of Asia and Africa (10 genera, 36 species), and infer patterns of evolution of sleep site selection. Materials and methods: We conducted fieldwork in 12 African and six Asian countries, collecting data on sleep sites, timing of sleep, and predation during daytime. We obtained additional information from literature and through correspondence. Using a phylogenetic approach, we established ancestral states of sleep site selection in lorisiforms and traced their evolution.

Adult Somali Lesser Galago Galago gallarum, Meru National Park, central Kenya. Photograph by Yvonne de Jong and Tom Butynski.

Results: The ancestral lorisiform was a fur-clinger and used dense tangles and branches/forks as sleep sites. Use of tree holes and nests as sleep sites emerged ~22 Mya (range 17–26 Mya) in Africa, and use of bamboo emerged ~11 (7–14) Mya in Asia and later in Africa. Fur clinging and some sleep sites (e.g., tree holes, nests, but not bamboo or dense tangles) show strong phylogenetic signal. Nests are used by Galagoides, Paragalago, Galago and Otolemur; tree holes by Galago, Paragalago, Sciurocheirus and Perodicticus; tangles by Nycticebus, Loris, Galagoides, Galago, Euoticus, Otolemur, Perodicticus and Arctocebus; all but Sciurocheirus and Otolemur additionally sleep on branches/forks. Daytime predation may affect sleep site selection and sleep patterns in some species of Nycticebus, Galago, Galagoides, Otolemur and Perodicticus. Most lorisiforms enter their sleep sites around sunrise and leave around sunset; several are active during twilight or, briefly, during daytime. Conclusion: Variations in sleep behavior, sleep patterns, and vulnerability to daytime predation provide a window into the variation that was present in sleep in early primates. Overall, lorisiforms use the daytime for sleeping and no species can be classified as cathemeral or polycyclic.

Full publication here: http://www.wildsolutions.nl/sleep-lorisiforms/

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Rainfall on Lolldaiga Hills Ranch Rainfall data kindly provided by Peter Karani.

Monthly rainfall (mm) on Lolldaiga Hills Ranch Rainfall (mm) at four sites on Lolldaiga Hills during 2018 Ranch during May and June 2018 (mean of four sites) 250 300 214 262 200 250 161 150 200 103 100 160 91 96 95 100 85 150 99 100 84 50

50 0 0 0 South West Central North 0 January February March April May June May June

Species totals as of end of June 2018

Mammals Lolldaiga Hills Conservation Landscape……..106 species

Mammals Lolldaiga Hills Ranch.………………………………….99 species Birds Lolldaiga Hills Ranch………………………………………..406 species Birds proposed IBA/KBA……………………………….………….633 species Reptiles Lolldaiga Hills Ranch………………..…….………….….33 species Amphibians Lolldaiga Hills Ranch...... 12 species Butterflies Lolldaiga Hills Ranch………………….……………135 species Moths Lolldaiga Hills Ranch...... >200 species

Except for moths, species list for the above taxonomic groups can be viewed at www.lolldaiga.com

Adult female Smith’s Dik-dik Madoqua smithii, Lolldaiga Hills Ranch, Kenya. Photograph by Per Aronsson.

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Best Zoological Society of London/Lolldaiga Hills Research Programme Camera Trap Project photographs on Lolldaiga Hills Ranch (May–June 2018)

Lion Panthera leo Lion Panthera leo

Savanna Elephant Loxodonta africana Savanna Elephant Loxodonta africana

Steenbuck Raphicerus campestris Common Duiker Sylvicapra grimmia

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African Buffalo Syncerus caffer Striped Hyaena Hyaena hyaena

Impala Aepyceros melampus White-tailed Mongoose Ichneumia albicauda

Common Warthog Phacochoerus africanus Common Warthog Phacochoerus africanus

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