Research project summmary

Main research fields: zoology and reproductive physiology specialization fields: reproduction, Behavioural ecology, Behavioural physiology, . My research interests focus mainly on the sociobiology of African moles (Bathyergidae) and in particular the extrinsic and intrinsic factors that have led to the evolution of in this remarkable endemic African family. My research group is fundamentally interested in elucidating the modes and mechanisms that are responsible for reproductive suppression in the non-reproductive females of the various . Research is currently being directed at the neuroendocrine and molecular levels to elucidate the extent, nature and location of GnRH suppression. We are also interested in the photic and thermic input in the control of reproduction in seasonally reproducing -rats and the potential lack of a role in aseasonally breeding bathyergids. Long term population studies on social mole-rats from mesic and xeric environments are underway. These studies are providing empirical data on the spatial distribution of colonies, longevities, factors restricting and promoting dispersal, vagility, foraging methods and lifetime reproductive success. We are interested in the genetic relatedness of colonies and also the type of paternal skew operational in the social genera. This work is being carried out in collaboration with Dr Chris Faulkes at Queen Mary and Westfield College, London.

Students currently under supervision

MSc (research)

1. Ms. Kemba Butler Neuroendocrinology of induced ovulation in the highveld mole-rat ( hottentotus pretoriae)

2. Mr. Andre Prins What makes a good helper? A behavioural study of cooperation in Damaraland mole-rats ( damarensis).

3. Mr. Josh Sarli Seasonal Reproductive Cycle and Parasite Burden of Two Small from Saudi Arabia.

Post-Doctoral

1. Dr. Sarita Maree 1. The conservation of golden moles (Afrosoricida; Chrysochloridae) in South

2. Dr. Katarina Medger The reproductive biology of two small southern African mammals, the spiny mouse, Acomys spinosissimus (Rodentia: Muridae) and the Eastern rock elephant-shrew, Elephantulus myurus (Macroscelidea: Macroscelididae) 3. Dr. M Oosthuizen Factors influencing neurogenesis and memory in three mole-rat species exhibiting a wide spectrum of social organisation.

Students currently under co-supervision

MSc (research)

1. Ms. Liz Archer The effects of the environment and sociality on parasite transmission and MHC selection in common mole-rats (Cryptomys hottentotus hottentotus)

2. Mr. Josh Sarli Seasonal Reproductive Cycle and Parasite Burden of Two Small Mammals from Saudi Arabia.

PhD

1. Mr. Low de Vries Ecology of the aardwolf, Proteles cristatus with special reference to diet, home range and prey abundance

2. Mrs. Jestina Katandukila A multidisplinary analysis on the status of the north-east African mole-rat, Tachyoryctes splendens (Rodentia: ) from Tanzania: systematics, ecology, physiology and epidemiology

3. Mr. James van Sandwyk Molecular epidemiology and evolution of Cardioviruses (Picornaviridae) in endemic South African murid

Recent Publications

2013

1. Lutermann, H., Young, A.J., Bennett, N.C. (2013) Reproductive status and testosterone among females in cooperative mole-rat societies. General and Comparative Endocrinology. 187:60-65

2. Lutermann, H., Bennett, N.C., Speakman, J.R., Scantlebury, M. (2013) Energetic benefits of sociality offset the costs of parasitism in a cooperative . PLoS One. 8:57969-57969 3. van Sandwyk, J., Bennett, N.C., Swanepoel, R., Bastos, A.D.S. (2013) Retrospective genetic characterisation of encephalomyocarditis viruses from African elephant and swine recovers two distinct lineages in South Africa. Veterinary Microbiology. 162(1):23-31

2012

1. Lutermann, H., Bodenstein, C., Bennett, N.C. (2012) Natural parasite infection affects the tolerance but not the response to a simulated secondary parasite infection. PLoS One. 7:52077-52077

2. Kotze, R., Bennett, N., Cameron, E.Z., de Vries, J.L., Marneweck, D.G., Pirk, C.W.W., Dalerum, F. (2012) Temporal patterns of den use suggest polygamous mating patterns in an obligate monogamous mammal. Animal Behaviour. 84:1573-1578

3. Medger, K., Chimimba, C.T., Bennett, N.C. (2012) Seasonal reproduction in the eastern rock elephant-shrew: Influenced by rainfall and ambient temperature? Journal of Zoology. 288:283-293

4. Ganswindt, A., Muilwijk, C., Engelkes, M., Muenscher, S., Bertschinger, H., Paris, M., Palme, R., Cameron, E., Bennett, N., Dalerum, F. (2012) Validation of Noninvasive Monitoring of Adrenocortical Endocrine Activity in Ground-Feeding Aardwolves (Proteles cristata): Exemplifying the Influence of Consumption of Inorganic Material for Fecal Steroid Analysis. Physiological and Biochemical Zoology. 85(2):194-199

5. Lutermann, H., Bennett, N.C. (2012) Determinants of Helminth Infection in a Subterranean Rodent, the Cape Dune Mole-Rat (Bathyergus suillus). Journal of Parasitology. 98:686-689

6. Brettschneider, H., Bennett, N.C., Chimimba, C.T., Bastos, A.D.S. (2012) Bartonellae of the Namaqua rock mouse, Micaelamys namaquensis (Rodentia: Muridae), from South Africa. Veterinary Microbiology. 157:132-136

7. Medger, K., Chimimba, C.T., Bennett, N.C. (2012) Seasonal changes in reproductive development in male spiny mice (Acomys spinosissimus) from South Africa. Mammalian Biology. 77:153-159

8. Medger, K., Chimimba, C.T., Bennett, N.C. (2012) Reproductive photoresponsiveness in male spiny mice from South Africa. Journal of Zoology. 286:243-249

2011

1. Viljoen, H., Bennett, N.C., Ueckermann, E.A., Lutermann, H. (2011) The role of host traits, season and group size on parasite burdens in a cooperative mammal. PLoS One. 6:-

2. Viljoen, H., Bennett, N.C., Lutermann, H. (2011) Life-history traits, but not season, affect the febrile response to a lipopolysaccharide challenge in highveld mole-rats. Journal of Zoology. 285:222-229 2010

1. Faulkes, C.G., Sichilima, A.M., van Sandwyk, J., Lutermann, H., Bennett, N.C. (2010) Control of ovulation in female giant mole-rats Fukomys mechowii(Rodentia: Bathyergidae), and phylogenetic trends within the family. Journal of Zoology. 282:64-74

2. Bennett, N.C., van Sandwyk, J., Lutermann, H. (2010) The pattern of ovulation in Ansell�s mole-rat, Fukomys anselli: phylogenetic or ecological constraints? Journal of Zoology. 281:66-73

3. Young, A.J., Oosthuizen, M.K., Lutermann, H., Bennett, N.C. (2010) Physiological suppression eases in Damaraland mole-rat societies when ecological constraints on dispersal are relaxed. Hormones and Behavior. 57:177-183

2009

1. Lutermann, H., Bennett, N.C. (2009) Parasites - a cost of sociality? Journal of the South African Veterinary Association. 80:131-131

2. Costanzo, M.S., Bennett, N.C., Lutermann, H. (2009) Spatial learning and memory in African mole-rats: the role of sociality and sex. Physiology and Behaviour. 96:128-134

3. Taylor, P.J., Maree, S., van Sandwyk, J., Kerbis Peterhans, J.C., Stanley, W.T., Verheyen, E., Kaliba, P., Verheyen, W., Kaleme, P., Bennett, N.C. (2009) mirrors geomorphology and palaecclimatic history in African laminate-toothed rats (Muridae; Otomyini) of the Otomys denti and O. lacustris species-complexes in the. Biological Journal of the Linnean Society. 96:913-941

4. Bennett, N.C., Faulkes, C.G., Hart, L., Jarvis, J.U.M. (2009) The Cape mole-rat, Bathyergus suillus. Mammalian Species. 828:1-7

2008

1. de Vries, L., Oosthuizen, M.K., Sichilima, A.M., Bennett, N.C. (2008) Circadian rhythms of locomotor activity in Ansell's mole-rat: are mole-rat's clocks ticking? Journal of Zoology. 276:343-349

2. Kotze, J., Bennett, N.C., Scantlebury, M. (2008) The energetics of huddling in two species of mole-rat (Rodentia: Bathyergidae). Physiology and Behaviour. 93:215-221

3. Bennett, N.C. (2008) The Mammal Research Institute 1966-2006. Transactions of the Royal Society of South Africa . 63(1):53-60

4. Manjerovic, M.B., Kinahan, A.A., Waterman, J.M., Bennett, N.C., Bateman, P.W. (2008) Structure and allometry of genitalia in males and females of a social African ground squirrel with high polygynandry. Journal of Zoology. 275:375-380 5. Oosthuizen, M.K., Bennett, N.C., Lutermann, H., Coen, C.W. (2008) Reproductive suppression and the seasonality of reproduction in the social Natal mole-rat (Cryptomys hottentotus natalensis). General and Comparative Endocrinology. 159:236-240

6. Sichilima, A.M., Bennett, N.C., Faulkes, C.G., Le Comber, S.C. (2008) Evolution of African mole-rats sociality: burrow architecture, rainfall and foraging in colonies of the cooperatively breeding Fukomys mechowii. Journal of Zoology. 275:276-282

7. Scantlebury, M., Waterman, J.M., Bennett, N. (2008) Alternative reproductive tactics in male Cape ground squirrels Xerus inauris. Physiology and Behaviour. 94:359-367

8. Scantlebury, M., Lovegrove, B.G., Jackson, C.R., Bennett, N.C., Lutermann, H. (2008) Hibernation and non-shivering thermogenesis in the Hottentot golden mole (Amblysomus hottentottus. J Comp Physiol B. 178:887-897

9. Cameron, E.Z., Lemons, P.R., Bateman, P.W., Bennett, N.C. (2008) Experimental alteration of litter sex ratios in a mammal. Proceedings of the Royal Society London, B. 275:323-327

10. Lutermann, H., Bennett, N.C. (2008) Strong immune function: a benefit promoting the evolution of sociality?. Journal of Zoology. 275(1):26-32

2007

1. Jackson, T.P., Waterman, J.M., Bennett, N.C. (2007) LH responses to single doses of exogenous GnRH in female social ground squirrels exhibiting low reproductive skew. Journal of Zoology. 273:8-13

2. Malherbe, G.P., Bennett, N.C. (2007) An assessment of behavioural dominance in a social subterranean rodent mole Cryptomys hottentotus pretoriae using the cardinal dominance index method. African Zoology. 42(2):187-198

3. Scantlebury, M., Waterman, J.M., Hillegass, M., Speakman, J.R., Bennett, N. (2007) Energetic costs of parasitism in the Cape ground squirrel Xerus inauris. Proceedings of the Royal Society - B. 274:2169-2177

4. Hart, L., Chimimba, C.T., Jarvis, J.U.M., O'Raian, J., Bennett, N.C. (2007) Craniometric sexual dimorphism and age variation in the solitary South African endemic Cape dune mole-rat, Bathyergus suillus (Rodentia: Muridae). Journal of Mammalogy. 88:657-666

5. Yeakel, J.D., Bennett, N.C., Koch, P.L., Dominy, N.J. (2007) The isotopic ecology of African mole-rats informs hypotheses on the evolution of human diet. Proceedings of the Royal Society, London. 274:1723-1730

6. Dalerum, F., Bennett, N.C., Clutton-Brock, T.H. (2007) Longitudinal differences in 15N between mothers and offspring during and after weaning in a small cooperative mammal, the (Suricata suricatta). Rapid Communications in Mass Spectrometry. 21:1889- 1892 7. Scantlebury, M., Waterman, J.M., Hillegrass, M., Speakman, J.R., Bennett, N.C. (2007) Effects of age and parasite status on the foraging energetics of female Cape ground squirrels, Xerus inauris. Proceedings of the Royal Society - B. 274:2169-2177

8. Kinahan, A.A., Bennett, N.C., O'Riain, J., Hart, L., Bateman, P.W. (2007) Size matters: genital allometry in an African mole-rat (Bathyergidae) . Evolutionary Ecology. 21:201-213

2006

1. Deuve, J.L., Bennett, N.C., O'Brien, P.C.M., Ferguson-Smith, M., Faulkes, C.G., Britton- Davidan, J., Robinson, T.J. (2006) Complex evolution of X and Y autosomal translocations in the Giant mole-rat, Cryptomys mechowi (Bathyergidae). Research. 14:681-691

2. Hart, L., O'Riain, M.J., Jarvis, J.U.M., Bennett, N.C. (2006) Is the Cape dune mole-rat Bathyergus suillus a seasonal or aseasonal breeder? Journal of Mammalogy. 87:1078-1085

3. Herbst, M., Bennett, N.C. (2006) The burrow architecture and burrowing dynamics of the endangered Namaqua dune mole-rat, Bathyergus janetta (Rodentia: Bathyergidae). Journal of Zoology. 270:420-428

4. Du Toit, L., Bennett, N.C., Gutjahr, G.H., Coen, C.W. (2006) Reproductive suppression in subordinate female highveld mole-rats, Cryptomys hottentotus pretoriae: No apparent role for endogenous opioid peptides. Physiology and Behaviour. 87:897-902

5. Snyman, P.C., Jackson, C.R., Bennett, N.C. (2006) Do dispersing female Damaraland mole- rats, Cryptomys damarensis (Rodentia: Bathyergidae) exhibit spontaneous ot induced ovulation? Physiology and Behaviour. 87:88-94

6. Scantlebury, M., Bennett, N.C., Speakman, J.R., Pillay, N., Schradin, C. (2006) The energetics of huddling in group living African four striped field mice Rhabdomys pumilio. Functional Ecology. 20:166-173

7. Broekman, M., Bennett, N.C., Jackson, C.R., Weber, R. (2006) Does altitude difference modulate the respiratory properties in subterranean rodent Cryptomys hottentotus mahali blood? Physiology and Behaviour. 88:77-81

8. Scantlebury, M., Speakman, J.R., Bennett, N.C. (2006) The energy costs of sexual dimorphism in mole-rats are morphological not behavioural. Proceedings of the Royal Society - B. 273:57-63

9. Bouche, L., Bennett, N.C., Jackson, T.P., Waterman, J.M. (2006) Is the Cape ground squirrel, Xerus inauris an induced or spontaneous ovulator? Journal of Mammalogy. 87:60-66

10. Bennett, N.C., Maree, S., Faulkes, C.G. (2006) The Cape , Georychus capensis. Mammalian Species. 799:1-4

11. Du Toit, L., Bennett, N.C., Kallo, I., Katz, A., Coen, C.W. (2006) A neuroanatomical and neuroendocrinological study into the relation between social status and the GnRH system in two cooperatively breeding species of African mole-rats Cryptomys h. hottentotus and C. h. pretoriae. Journal of Comparative Neurology. 494:303-313

12. Muteka, S.P., Chimimba, C.T., Bennett, N.C. (2006) Reproductive seasonality in the Tete veld rat, Aethomys ineptus (Rodentia: Muridae) from southern Africa. Journal of Zoology (London). 270:314-322

13. Muteka, S.P., Chimimba, C.T., Bennett, N.C. (2006) Reproductive seasonality in Aethomys namaquensis (Rodentia: Muridae) from southern Africa. Journal of Mammalogy. 87:67-74

14. Muteka, S.P., Chimimba, C.T., Bennett, N.C. (2006) Reproductive photoresponsiveness in Aethomys ineptus and A. namaquensis (Rodentia: Muridae) from southern Africa. Journal of Zoology (London). 268:225-231

15. Young, A.J., Monfort, S.L., Carlson, A.A., Russell, A.F., Bennett, N.C., Clutton-Brock, T.H. (2006) Stress and the suppression of subordinate reproduction in cooperatively breeding . Proceedings of the National Academy of Sciences. 103:12005-12010

16. Clutton-Brock, T.H., Hodge, S.J., Spong, G., Russell, A.F., Jordan, N.R., Bennett, N.C., Sharp, L., Manser, M.B. (2006) Intrsexual competition and sexual selection in cooperative mammals. Nature. (444):1065-1068

17. Scantlebury, M., Speakman, J.R., Oosthuizen, M.K., Roper, T.J., Bennett, N.C. (2006) Energetics reveals physiologically distinct castes in a eusocial mammal. Nature. 440:795-797

18. Bateman, P.W., Bennett, N.C. (2006) The biology of human sexuality: evolution, ecology and physiology. Verbum et Ecclesia. 27:245-264

19. Parag, A., Bennett, N.C., Faulkes, C.G., Bateman, P.W. (2006) Penile morphology of African mole-rats (Bathyergidae): structural modification in relation to mode of ovulation and degree of sociality. Journal of Zoology. 270:323-329

2005

1. Vasicek, C., Malpaux, B., Fleming, P.A., Bennett, N.C. (2005) Melatonin secretion in the Mashona mole-rat, Cryptomys darlingi -influence of light on rhythmicity. Physiology and Behaviour. 83:689-697

2. Vasicek, C.A., Oosthuizen, M.K., Cooper, H.M., Bennett, N.C. (2005) Endogenous circadian rhythms of locomotor activity in the subterranean Mashona, Cryptomys darlingi: the effects of light on entrainment. Physiology and Behaviour. 84:181-191

3. Scantlebury, M., Oosthuizen, M.K., Speakman, J.R., Bennett, N.C. (2005) Seasonal field energetics of the Hottentot golden mole at high altitude: an ancient African Afrotherian. Physiology and Behaviour. 84:739-745 4. Oosthuizen, M.K., Bennett, N.C., Cooper, H.M. (2005) C-fos photic induction in two species of African mole-rat (Rodentia: Bathyergidae). Neuroscience. 133:555-560

5. van Sandwyk, J., Bennett, N.C. (2005) Do solitary, seismic signalling Cape mole-rats, Georychus capensis demonstrate spontaneous or induced ovulation? Journal of Zoology. 267:75-80

6. Maswanganye, K.A., Malherbe, G.P. ., Bennett, N.C. (2005) Ultrastructure of epididymal spermatozoa in three social African mole-rats of the genus Cryptomys. African Zoology. 40:293-300

7. Schumann, D.M., Cooper, H.M., Hofmeyr, M.D., Bennett, N.C. (2005) Circadian rhythms of locomotor activity in the four striped field mouse, Rhabdomys pumilio: a diurnal or crepuscular African rodent. Physiology and Behaviour. 85:231-239

8. Jackson, C.R., Bennett, N.C. (2005) Is the Natal mole-rat (Cryptomys hottentotus natalensis) a spontaneous or induced ovulator? Journal of Mammalogy. 86:1-6

9. Chimimba, C.T., Bennett, N.C. (2005) Order Rodentia. In: The Mammals of the Southern African Subregion (JD Skinner & CT Chimimba, Revisers). Cambridge University Press, Cambridge. :77-209

2004

1. Hart, L., Bennett, N.C., Malpaux, B., Chimimba, C.T., Oosthuizen, M.K. (2004) The chronobiology of the Natal mole-rat, Cryptomys hottentotus natalensis. Physiology & Bahaviour. 82:563-569