zebra stripe

herder and sheep
herder and sheep
zebra herder and sheep
Department of Ecology and Evolutionary Biology | Princeton University



Crawford, .C.L., Volenec, Z.M., Sisanya, M., Kibet, R., Rubenstein, D.I. (2019). Behavioral and ecological implications of bunched, rotational cattle grazing in east African savanna ecosystem. Rangeland Ecology & Management, 72: 1, 204-209. DOI/ 10.1016/j.rama.2018.07.016 Open Access

Ng’weno, C., Buskirk, S., Georgiadis, N., Gituku, B., Kibungei, A., Porensky, L., Rubenstein, D. I. & Goheen, J. Apparent Competition, Lion Predation, and Managed Livestock Grazing: Can Conservation Value Be Enhanced? Frontiers in Ecology and Evolution. 7, 123. DOI/10.3389/fevo.2019.00123 (PDF)  Open Access

Nunez, C. M., & Rubenstein, D. I. (2019). Mother-infant communication in feral horses (Equus caballus): what are they saying, why are they saying it, and what might it tell us about the mammalian juvenile stage? Integrative and Comparative Biology (Vol. 59, pp. E171-E171).

O'Bryan, L.R., Abaid, N, Nakayama, N., Dey, T., King, A. J., Cowlishaw, G., Garnier, S., Rubenstein, D. I. (2019). Contact Calls Facilitate Group Contraction in Free-ranging Goats (Capra aegagrus hircus). Frontiers in Ecology and Evolution, 7, 73. DOI/10.3389/fevo.2019.00073 (PDF)  Open Access

O’Connor, D., Stacy-Dawes, J., Muenuza, A., Rubenstein, D.I., et al. (2019) Updated geographic range maps for giraffe, Giraffa spp., throughout sub‐Saharan Africa, and implications of changing distributions for conservation. Mammal Review. DOI/10.1111/mam.12165 (PDF) Open Access

Rubenstein, D. I. (2019). On assessing the importance of demographic change for social structure: a comment on Shizuka and Johnson. Behavioral Ecology .doi.org/10.1093/beheco/arz164 Open Access

Tombak, K. J., Wikberg, E. C., Rubenstein, D. I., Chapman, C. A. (2019). Reciprocity and rotating social advantage among females in egalitarian primate societies. Animal Behaviour. doi.org/10.1016/j.anbehav.2019.09.010. (PDF)  Open Access


Cabal, C. & Rubenstein, D.I.  (2018) Above- and Below-Ground Biomass Allocation and Functional Trait Response to Soil Water Inputs and Evaporation of two Common Savanna Grasses. Journal of Arid Environments Vol.157 10/2018 1-12 DOI/10.1016/j.jaridenv.2018.06.008 Open Access

Caputo., M., Bouveroux, T.,  & Rubenstein, D.I.  (2017)Striping patterns may not influence social interactions and mating in zebra: evidence from melanic zebra in South Africa.   African Journal of Ecology. DOI: 10.1111/aje.12463 Open Access

Costelloe, B.R. & Rubenstein, D.I. (2018) Temporal structuring of vigilance behavior by female Thomson’s gazelles with hidden fawns.  Animal Behavior. 145, 87-97. DOI/10.1016/j.anbehav.2018.09.007 Open Access

Greenbaum, G., Renan, S., Templeton, A.R., Bouskila, A., Saltz, D., Rubenstein, D.I., Bar-David, S. (2017) Revealing life-history characteristics and their impacts on genetic drift by contrasting genetic estimation with theoretical predictions of effective population size. Conservation Biology. DOI/10.1111/cobi.13068 Open Access

Henry, L., Tokita, C. Misra, M. & Rubenstein, D.I. (2018)  Mutualistic Acacia Ants Exhibit Lower Defensive Behavior and Higher Off-Tree Movement Near Termite Mounds. Biotropica. DOI/10.1111/btp.12572 Open Access

Kendall, C., Rubenstein, D.I., Slater, P., Mondajem, A. (2017) An assessment of tree availability as a possible cause in the population decline of scavenging raptors. Journal of Avian Biology.   DOI: 10.1111/jav.01497 Open Access

Kulachi, I., Ghazanfar, A., & Rubenstein, D.I. (2018) Consistent individual variation across interaction networks indicates social personalities in lemurs. Animal Behavior. DOI/10.1016/j.anbehav.2017.11.012 Open Access

Kulachi, I., Ghazanfar, A., & Rubenstein, D.I. Knowledgeable lemurs become more central in social networks. Current Biology 28 (8), 1306-1310. e2. DOI/10.1016/j.cub.2018.02.079 Open Access

Monk, C., Romanczuk, R., Watson, J., Arlinghaus, R., Levin, S., & Rubenstein, D.I.  (2018) How ecology shapes exploitation: a framework to predict the behavioural response of natural resource users along exploration-exploitation tradeoffs. Ecology Letters. 21 (6), 779-793 DOI/10.1111/ele.12949 Open Access

O’Brien, T.G., Kinnaird, M.F., Ekwanga, S., Williams, T., Alayne, O.C., Rubenstein, D.I., Frank, L., (2018) Resolving a conservation dilemma: vulnerable lions eating endangered zebras. PloS one, 13 (8). DOI/10.1371/journal.pone.0201983 Open Access

Odadi, W., Riginos, C. & Rubenstein, D.I. (2018) Tightly bunched herding improves cattle performance in an African savanna rangeland. Rangeland Ecology & Management 71 (4), 481-491 DOI/10.1016/j.rama.2018.03.008 Open Access

Parham, J., Stewart, C., Crall, J., Rubenstein, D.I., Holmberg, J., & Berger-Wolf, T. (2018) An Animal Detection Pipeline for Indentification. 2018 IEEE Winter Conference on Applications of Computer Vision (WACV), 1075-1083 DOI/10.1109/WACV.2018.00123 Open Access

Rubenstein, D.I. (2018) Moving beyond structure to function. Animal Behavior. 136. 173-174. DOI/10.1016/j.anbehav.2018.01.001

Schuttler, S., Rubenstein, D.I., et al (2018). Citizen science in schools: Students collect valuable mammal data for science, conservation, and community engagement. BioScience. DOI/10.1093/biosci/biy141 Open Access

Tucker, M.A., Böhning-Gaese, K., Fagan, W.F., Fryxell, J.M., Moorter, B., Rubenstein, D.I. et al.  Moving in the Anthropocene: global reductions in terrestrial mammalian movements. Science. 26 Jan 2018: 466-469. DOI: 10.1126/science.aam9712 Open Access


Berger-Wolf, T.,  Rubenstein, D.I., Stewart, C.V., Holmberg, J.A., Parham, J., Meron, S., Crall, J., van    Oast, J., Kiciman, E. & L. Joppa. (2017) Wildbook:  Crowdsourcing, computer vision and data science for conservaton.  arXiv:1710.08880 [cs.CY] Open Access

Gersick, A. S., & Rubenstein, D. I. (2017). Physiology modulates social flexibility and collective behaviour in equids and other large ungulates. Phil. Trans. R. Soc. B, 372(1727), DOI/10.1098/rstb.2016.0241 Open Access

Hauck, S., & Rubenstein, D. I. (2017). Pastoralist societies in flux: A conceptual framework analysis of herding and land use among the Mukugodo Maasai of Kenya. Pastoralism, 7(1), 18. DOI/10.1186/s13570-017-0090-4 Open Access

Nuñez, C. M., Adelman, J. S., Carr, H. A., Alvarez, C. M., & Rubenstein, D. I. (2017). Lingering effects of contraception management on feral mare (Equus caballus) fertility and social behavior. Conservation Physiology, 5(1), DOI/10.1093/conphys/cox018 Open Access

Nuñez, C. M., Adelman, J. S. A., Carr, H.A.,Knight, C.,& Rubenstein, D.I.. (2017) “Prolonged Effects of Contraception Management on Feral Horse (Equus Caballus) Reproductive Physiology and Behavior.Integrative and Comparative Biology, vol. 57, pp. E366-E366, DOI: 10.1093/icb/icx002.

Odaddi, W, Fragione, J. and Rubenstein, D.I. (2017) Vegetation, wildlife and livestock responses to planned grazing management in an African pastoral landscape.  Land Degradation & Development. DOI/10.1002/ldr.2725  Open Access

J. Parham, J. Crall, C. Stewart, T. Berger-Wolf, and D.I. Rubenstein, “Animal Population Censusing at Scale with Citizen Science and Photographic Identification,” presented at the AAAI 2017 Spring Symposium on AI for Social Good, Stanford, CA, 2017 (PDF)

Schieltz,  J., Okanga, S., Allan, B., Rubenstein. D.I. (2017) GPS tracking cattle as a monitoring tool for conservation and management, African Journal of Range and Forage Science. 34:3 DOIabs/10.2989/10220119.2017.1387175


Barnier, F., Duncan, P., Fritz, H., Blanchard, P., Rubenstein, D. I., & Pays, O. (2016). Between-gender differences in vigilance do not necessarily lead to differences in foraging-vigilance tradeoffs. Oecologia, 1-12. DOI/10.1007/s00442-016-3614-5

Kahumbu, P., Martins, D., Schieltz, J. & Rubenstein, D.I. (2016) Transforming the next generation through citizen science in Kenya: the kids twiga tally. Swara, April-June, 52-56. (PDF)

Kulahci, I. G., Rubenstein, D. I., Bugnyar, T., Hoppitt, W. J. E., Mikus, N., & Schwab, C. (2016). Social networks predict selective observation and information spread in ravens. Royal Society Open Science. 3(7), 160256.  DOI/10.1098/rsos.160256 Open Access

Lalampaa, P. K., Wasonga, O. V., Rubenstein, D. I., & Njoka, J. T. (2016). Effects of holistic grazing management on milk production, weight gain, and visitation to grazing areas by livestock and wildlife in Laikipia County, Kenya. Ecological Processes, 5(1), 17. DOI/10.1186/s13717-016-0061-5 Open Access

Pappano, D. J., Bergman, T. J., Beehner, J. C., Berger-Wolf, T., & Rubenstein, D. I. (2016). Seasonal influences on gelada social networks. American Journal of Physical Anthropology. 159, 247-247. (link)

Rubenstein, D.I.  (2016). Anthropogenic Impacts on Behavior:  The Pros and Cons of Plasticity.  In “Conservation Behavior”. Eds. Saltz, D. and O. Berger-tal.  Pp. 121-131. Cambridge Univ. Press. DOI/10.1017/CBO9781139627078

Rubenstein, D.I., Low Mackey, B., Davidson, ZD, Kebede, F. & King, S.R.B. (2016).  Equus grevyi. The IUCN Red List of Threatened Species 2016. (link)

Rubenstein, D.I., Cao, Q.  and J. Chui.  (2016). Equids and ecological niches:  behavioral and life history variations on a common theme. In “Wild Equids: Ecology, Management, and Conservation”, Eds. Ransom, J.R. & P. Kaczencky. Pp. 58-68.  Johns Hopkins University Press. (link)

Rubenstein, D. R., & Rubenstein, D. I. (2016). From Pleistocene to trophic rewilding: A wolf in sheep’s clothing. Proceedings of the National Academy of Sciences, 201521757. DOI/10.1073/pnas.1521757113 Open Access

Schieltz, J. M., & Rubenstein, D. I. (2016). Evidence-based review: positive versus negative effects of livestock grazing on wildlife. What do we really know?. Environmental Research Letters, 11(11), 113003. DOI/10.1088/1748-9326/11/11/113003 Open Access

Uyehara, I. K., Sisanya, M., Hemp, C., & Rubenstein, D. I. (2016). Effects of traditional pastoralism on grasshopper (Caelifera) assemblages in East Africa. African Journal of Ecology, 54(2), 167-173. DOI/10.1111/aje.12279


Barale, C., Rubenstein., D.I., Beehner, J. (2015) Juvenile social relationships reflect adult patterns of behavior in wild geladas. American Journal of Primatology. 77(10), 1086-1096. DOI/10.1002/ajp.22443 

Costelloe, B.R. and Rubenstein, D.I. (2015) Coping with transition: offspring risk and maternal behavioral changes at the end of the hiding phase. Animal Behaviour 109: 217-2 DOI/10.1016/j.anbehav.2015.08.022 Open Access

de Froment, A.J., Rubenstein, D.I., and S.A. Levin. (2015) An extra dimension to decision-making in animals: The three-way trade-off between speed, effort per-unit-time and accuracy, PLoS Computational Biology. 10(12), e1003937 PDF

Kartzinel, T., Chen, P., Coverdale, T., Erickson, D., Kress, J., Rubenstein, D.I., Wang, W., Pringle, R. (2015) DNA metabarcoding illuminates dietary niche partitioning by large African herbivores. Proceedings of the National Academy of Sciences. vol. 112 no. 26, 8019–8024, (link) Open Access

Kulachi, I., Rubenstein, D.I., and Ghazanfar, A. (2015) Lemurs groom-at-a-distance through vocal networks. Animal Behaviour.110. 179-186. DOI/10.1016/j.anbehav.2015.09.016

Larison, B., Harrigan, R.J., Rubenstein, D.I., & Smith, T.B. (2015) Concordance on zebra stripes is not black and white. Royal Society Open Science. 2(1), 140452. DOI/10.1098/rsos.150359 Open Access

Larison, B., Harrigan, R. J., Thomassen, H. A., Rubenstein, D. I., Chan-Golston, A. M., Li, E., & Smith, T. B. (2015). How the zebra got its stripes: a problem with too many solutions. Royal Society Open Science, 2(1), 140452. Open Access

Nuñez, C., Adelman, J. and Rubenstein, D. (2015). Sociality increases juvenile survival after a catastrophic event in the feral horse (Equus caballus). Behavioral Ecology 26 (1): 138-147. (link)

Odadi, W. O., & Rubenstein, D. I. (2015). Herd Size-Dependent Effects of Restricted Foraging Time Allowance on Cattle Behavior, Nutrition, and Performance. Rangeland Ecology & Management. 68:4, pp. 341-348. (link)

Riginos, C., M.A. Karande, D.I. Rubenstein, and T.M. Palmer. (2015). Disruption of a protective ant-plant mutualism by an invasive ant increases elephant damage to savanna trees. Ecology 96(3), 2015, pp. 654–661 (link)

Rubenstein D 2015. Networks of terrestrial ungulates: linking form and function. In: Animal social networks (Krause J, James R, Franks DW & Croft DP, eds). Oxford University Press, Oxford. pp. 184-196. (link)

Rubenstein, D.I. , Berger-Wolf, T., Tantipathananandh, C., Sundaresan, S.,& Fischoff, I. & (2015) Similar but Different: Dynamic Social Network Analysis Highlights Fundamental Differences between the Fission-Fusion Societies of Two Equid Species, the Onager and Grevy's Zebra. PLoS One. DOI/10.1371/journal.pone.0138645 Open Access

Rubenstein, Daniel I, Kammen, Daniel M. (2015). The launch of Environmental Research Reviews. Environmental Research Letters, 10 (12), 120402 - 120402. DOI/10.1088/1748-9326/10/12/120402/meta Open Access

Schieltz, J. M., & Rubenstein, D. I. (2015). Caught between two worlds: genes and environment influence behaviour of plains× Grevy's zebra hybrids in central Kenya. Animal Behaviour, 106, 17-26. (link)

Searchinger, T. D., Estes, L., Thornton, P. K., Beringer, T., Notenbaert, A., Rubenstein, D., Heimlich, R., Licker, R. & Herrero, M. (2015). High carbon and biodiversity costs from converting Africa/'s wet savannahs to cropland. Nature Climate Change. doi:10.1038/nclimate2584 (link)

Tong, W. Shapiro, B. and Rubenstein, D.I. (2015) Genetic relatedness in two-tiered plains zebra societies suggests that females choose to associate with kin. Behaviour. DOI: 10.1163/1568539X-00003314 (PDF)

Zhang, Y., Cao Q.S., Rubenstein D.I., Zang, S., Songer, M., Leimgruber, P., Chu, H.,  Cao, J., Li, K.,  and Hu, D.. Water use patterns of sympatric Przewalski's horse and Kulan: Interspecific comparison reveals niche differences. PLOS ONE.  DOI: 10.1371/journal.pone.0132094. Open Access


Kendall, Corinne J., Virani, Munir Z., Hopcraft, J. Grant C., Bildstein, Keith L., Rubenstein, D.I. (2014). African Vultures Don’t Follow Migratory Herds: Scavenger Habitat Use Is Not Mediated by Prey Abundance. PLoS ONE, DOI: 10.1371/journal.pone.0083470 Open Access

Kulahci, I. G., Drea, C. M., Rubenstein, D. I. and Asif A. Ghazanfar, A. A. (2014)Individual recognition through olfactory–auditory matching in lemurs. Proceedings of the Royal Society Vol. 281 No. 1784 Open Access

Lekerpes, S. S., Jung'a, J. O., Badamana, M. S., & Rubenstein, D. I. (2014). Genetic polymorphism of beta-lactoglobulin in Kenyan small east African goat breed using PCR-RFLP and sequencing. Scientific Journal of Animal Science, 3(9), 233-239. (link)

Nowzari, H., Hemami, M., Karami, M., Zarkesh, M.M.K., Riazi, B. and Rubenstein, D.I.(2013). Habitat use by the Persian Onager Equus hemionus onager (Perissodactyla: Equidae) in Qatrouyeh National Park, Fars, Iran. Journal of Natural History 47:43-44, 2795-2814, DOI: 10.1080/00222933.2013.802040 PDF (link)

Nunez, C.M.V, Adelman, J.S. and Rubenstein, D.I. (2013). A Free-Ranging, Feral Mare Equus caballus Affords Similar Maternal Care to Her Genetic and Adopted Offspring. The American Naturalist November 2013, Vol. 182, No. 5, pp. 674-681 (link) Open Access

Nunez, C.M.V, Adelman, J.S., Smith, J., Gesquiere, L.R., Rubenstein, D.I. (2014) Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels. General and Comparative Endocrinology, Vol. 196, pp. 26–33 (link)

Roberts, B.A. and Rubenstein, D.I. (2014) Maternal tactics for mitigating neonate predation risk during the postpartum period in Thomson’s gazelle. Behavior 151: 1229-1248 (link)


Asa, C., Beever, E., Coughenour, M., Eggert, L., Garrott, R., Huntsinger, L., Kalof, L., Krausman, P., Oli, M., Palmer, G., Petersen, S., Powell, D., Rubenstein, D., Thain, D. (2013). National Research Council. Using science to improve the BLM Wild Horse and Burro Program: A way forward. National Academies Press. 630 pp. (link)

Crall, J.P., Stewart, C.V., Berger-Wolf, T.Y., Rubenstein, D. I., and Sundaresan, S.R. (2013). HotSpotter - Pattern Species Instance Recognition. IEEE Workshop on Applications of Computer Vision. (link)

Giuggioli, L, Potts, J.R., Rubenstein, D.I. and Levin, S.A. (2013). Stigmergy, collective actions, and animal social spacing. Proceedings of the National Academy of Sciences. (link)

Nowzari, H., Hemami, M., Karami, M., Zarkesh, M.M.K., Riazi, B., Rubenstein, D. (2013). Habitat associations of persian wild ass (Equus Hermionus Onager) in QA - Trouyeh National Park, Iran, pp. 25-30 PDF

Rubenstein, D. I., and Hack Mace A. (2013). Migration. In: Levin S.A. (ed.) Encyclopedia of Biodiversity, second edition, Volume 5, pp. 309-320. Waltham, MA: Academic Press (link)

Rubenstein D. I., and Rubenstein D. R. (2013). Social Behavior. In: Levin S.A. (ed.) Encyclopedia of Biodiversity, second edition, Volume 6, pp. 571-579. Waltham, MA: Academic Press. (link)

Ward, A., Herbert-Read, J., Jordan, L., James, R., Krauser, J., Ma, Q., Rubenstein, D. I., Sumpter, D., Morrell, L.J. (2013). Initiators, Leaders, and Recruitment Mechanisms in the Collective Movements of Damselfish. The American Naturalist, Vol. 181, No. 6, pp. 748-760 (link) Open Access


Chandler, M., Bebber, D.P., Castro, S., Lowman. M.D., Muoria, P., Oguge, N., Rubenstein, D. I. (2012). International citizen science: making the local global. Frontiers in Ecology and the Environment. 10: 328-331 (link)

Dixit, A., Levin, S.A., and Rubenstein, D. I. (2012). Reciprocal insurance among Kenyan pastoralists. Theoretical Ecology: Volume 6, pp. 173-187.DOI: 10.1007/s12080-012-0169-x. (link)


Crofoot, M. C., Rubenstein, D. I., Maiya. A.S., Berger-Wolf, T.Y.  (2011). Aggression, grooming and group-level cooperation in white-faced capuchins (Cebus capucinus): Insights from social networks. American Journal of Primatology. 73: 821-833. 
DOI: 10.1002/ajp.20959 (pdf)

Didier, K. A., Cotterill, A., Douglas-Hamilton, I., Frank, L., Georgiadis, N.J., Graham, M., Ihwagi, F., King, J., Malleret-King, D., Rubenstein, D. I., Wilkie, D. & Woodroffe, R.. (2011). Conserving Wildlife in African Landscapes: Kenyas Ewaso Ecosystem. In: Smithsonian Contributions to Zoology, 632: 105-123. N. J. Georgiadis, ed. Smithsonian Institution Scholarly Press. (link) (pdf)

Lahiri, M., Tantipathananandh, C., Warungu, R., Rubenstein, D. I. & Berger-Wolf, T.Y.. (2011). Biometric Animal Databases from Field Photographs: Identification of Individual Zebra in the Wild. Proceedings of the ACM International Conference on Multimedia Retrieval (ICMR 2011), Trento, Italy.  (pdf)

Nuñez, C., Asa, C. S. & Rubenstein, D. I.. (2011). Zebra Reproduction: Plains Zebra (Equus burchelli), Mountain Zebra (Equus zebra), and Grevy’s Zebra (Equus grevyi). Pp. 2851-2865. In: Equine Reproduction, 2nd edition. A. O. McKinnon, E. L Squires, W. E. Vaala & D. D. Varner, eds. Wiley-Blackwell, Ames, Iowa. (pdf)

Odadi, W. O., Jain, M., Van Wieren, S.E., Prins, H.H.T. & Rubenstein, D. I.. (2011) Facilitation between bovids and equids in and African savanna. Evolutionary Ecology Research 13: 237-252. (pdf)

Rubenstein, D. I. (2011). Family equidae (Horses and relatives). Pp. 106-143. In: Handbook of Mammals of the World, Vol. 2, Hoofed Mammals, D. E. Wilson and R. A. Mittermeier, eds. Lynx Edicions, Barcelona, Spain. (PDF)


Fischhoff, I. R., Sundaresan, S.R., Larkin, H.M., Cordingley, J.E., Sellier, M-J. & Rubenstein, D. I. (2010). A rare fight between two female plains zebra (Equus burchelli). Journal of Ethology 28: 201-205. DOI 10.1007/s10164-009-0183-7 (pdf)

Jordan, L. A., Avolio, C., Herbert-Read, J.E., Krause, J., Rubenstein D. I. & Ward, A.J.W.(2010). Group structure in a restricted entry system is mediated by both resident and joiner preferences. Behavioral Ecology and Sociobiology.64(7): 1099-1106. DOI: 10.1007/s00265-010-0924-1 (pdf)

Madosky, J. M., Rubenstein, D. I., Howard, J.J. & Stuska, S.. (2010). The effects of immuno-contraception on harem fidelity in a feral horse (Equus caballus) population. Applied Animal Behavior Science 128: 50-56. DOI: 10.1016/j.applanim.2010.09.013 (pdf)

Nuñez, C.M.V., Adelman, J.S. & Rubenstein, D. I.. (2010). Immunocontraception in wild horses (Equus caballus) extends reproductive cycling beyond the normal breeding season. PLoS ONE. 5(10): e13635. doi:10.1371/journal.pone.0013635 (pdf)

Rubenstein, D. I. (2010). Ecology, social behavior, and conservation in zebras. Pp. 231­258. In: Advances in the Study Behavior: Behavioral Ecology of Tropical Animals, Vol. 42. R. Macedo, ed. Elsevier Press, Oxford, UK. (pdf)

Rubenstein, D. I. (2010). The secret lives of horses. Odyssey Magazine. 19(5): 24-27. (pdf)


Cordingley, J. E., S. R. Sundaresan, I. R. Fischhoff, B. Shapiro, J. Ruskey & D. I. Rubenstein. (2009). Is the endangered Grevy¹s zebra (Equus grevyi) threatened by hybridization? Animal Conservation 12: 505-513. DOI: 10.1111/j.1469-1795.2009.00294.x (pdf)

Cordingley, J. E., S. R. Sundaresan, B. J. Larison, B. Shapiro & D. I. Rubenstein. (2009). Response: Grevy¹s zebra conservation: overcoming threats of isolation, genetic hybridization and demographic instability. Animal Conservation 12: 520-521. (pdf)

Didier, K. A., D. Wilkie, I. Douglas-Hamilton, L. Frank, N. Georgiadis, M. Graham, F. Ihwagi, A. King, A. Cotterill, D. Rubenstein & R. Woodroffe. (2009). Conservation planning on a budget: a "resource light" method for mapping priorities at a landscape scale? Biodiversity and Conservation 18(7): 1979-2000. (pdf)

Fischhoff I. R., J. Dushoff, S. R. Sundaresan, J. E. Cordingley, D. I. Rubenstein. (2009). Reproductive status influences group size and persistence of bonds in male plains zebra (Equus burchelli). Behavioral Ecology and Sociobiology 63:1035-1043. (pdf)

Low B., S. R. Sundaresan I. R. Fischhoff & D. I. Rubenstein (2009).  Partnering with local communities to identify conservation priorities for endangered Grevy's zebra. Biological Conservation 142(7): 1548-1555. (pdf)

Nuñez, C.M.V., J. S. Adelman, C. Mason & D. I. Rubenstein. (2009). Immunocontraception decreases group fidelity in a feral horse population during the non-breeding season. Applied Animal Behaviour Science 117(1-2): 74-83. (pdf)

Rubenstein, D. I. (2009). Social behavior and sociobiology. Pp. 237-255. In: Evolution: The First Four Billion Years. M. Ruse and J. Travis (eds). Harvard University Press, Cambridge, MA. (pdf) (link)

Rubenstein, D. I., C. Nuñez. (2009). Sociality and reproductive skew in horses and zebras. Pp. 196-226. In: Reproductive Skew in Vertebrates: Proximate and Ultimate Causes. R. Hager, C. B. Jones (eds).  Cambridge University Press. (pdf) (link)

Franceschini, M. D., D. I. Rubenstein, B. Low & L. M. Romero. (2008). Fecal glucocorticoid metabolite analysis as an indicator of stress during translocation and acclimation in a, endangered large mammal, the Grevy’s zebra. Animal Conservation 11: 263-269. (pdf)

Moehlman, P.D., Rubenstein, D.I. & Kebede, F. 2008. Equus grevyi. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2.(link)

Sundaresan, S. R., I. R. Fischhoff, H. M. Hartung, P. Akilong & D. I. Rubenstein. (2008). Habitat choice of Grevy's zebras (Equus grevyi) in Laikipia, Kenya. African Journal of Ecology 46(3): 359-364. (pdf)

Fischhoff I. R., S. R. Sundaresan, J. Cordingley, H. M. Larkin, M-J. Sellier & D. I. Rubenstein. (2007). Social relationships and reproductive state influence leadership roles in movements of plains zebra (Equus burchelli). Animal Behaviour 73: 825-831. (pdf)

Fischhoff, I. R., S. R. Sundaresan, J. Cordingley & D. I. Rubenstein. (2007). Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions. Behavioral Ecology 18(4): 725-729. (pdf)

Pringle, R. M., T. P. Young, D. I. Rubenstein & D. J. McCauley. (2007). Herbivore-initiated interaction cascades and their modulation by productivity in an African savanna. Proceedings of the National Academy of Sciences, USA 104(1): 193-197. (pdf)

Rubenstein, D. I., S. Sundaresan, I. Fischhoff & D. Saltz (2007). Social networks in wild asses: Comparing patterns and processes among populations. Pp: 159-176. In: Exploration into the Biological Resources of Mongolia, Vol. 10. A. Stubbe, P. Kaczensky, K. Wesche, R. Samjaa, M. Stubbe (eds.). Martin-Luther-University Halle-Wittenberg, Halle (Saale). (pdf)

Sundaresan, S. R., I. R. Fischhoff, J. Dushoff & D. I. Rubenstein. (2007). Network metrics reveal differences in social organization between two fission-fusion species, Grevy’s zebra and onager. Oecologia, 151(1): 140-149. (pdf)

Sundaresan, S. R., I. R. Fischhoff & D. I. Rubenstein. (2007). Male harassment influences female movements and associations in Grevy’s zebra (Equus grevyi). Behavioral Ecology 18(5): 860-865. doi:10.1093/beheco/arm055 (pdf)

Rubenstein, D. R., D. I. Rubenstein, P. W. Sherman & T. A. Gavin (2006). Pleistocene Park: Does re-wilding North America represent sound conservation for the 21st century? Biological Conservation 132: 232-238. (pdf)
Saltz, D., D. I. Rubenstein & G. C. White (2006). The impact of increased environmental stochasticity, due to climate change on the dynamics of Asiatic wild ass. Conservation Biology 20(5): 1402-1409. (pdf)

Muoria, P. K., P. Muruthi, D. Rubenstein, N. O. Oguge & E. Munene (2005). Cross-sectional survey of gastro-intestinal parasites of Grevy’s zebras in southern Samburu, Kenya. African Journal of Ecology 43: 392-395. (pdf)

Weinstock, J., E. Willerslev, A. Sher, W. Tong, S. Y.W. Ho, D. Rubenstein, J. Storer, J. Burns, L. Martin, C. Bravi, A. Preito, D. Froese, E. Scott, L. Xulong & A. Cooper (2005). Evolution, systematics, and phylogeography of Pleistocene horses in the New World: A molecular perspective. PloS Biology 3(8): 1373-1379. (pdf)
Zhang, P., C.M. Sadler, T. Liu, I. Fischhoff, Martonosi, M., S. A, Lyons & D. I. Rubenstein. (2005). Habitat monitoring with ZebraNet: Design and experiences. Pp: 235-257. In: Wireless Sensor Networks: A Systems Perspective. N. Bulusu and S. Jha (eds.). Artech House, Norwood, MA. (pdf)

Rubenstein, D. I. (2004). Zebra sociality: Different stripes for different types. Balliol College Record. Pp 18-23. (pdf)

Rubenstein, D. I. (2004). Herd dynamics: Why aggregations vary in size and complexity. Pp 994-1000. In: Encyclopedia of Animal Behavior, M. Bekoff, (ed.). Greenwood Press, Westport, CT. (pdf)

Rothley, K. D., C. N. Berger, C. Gonzalez, E. M. Webster, & D. I. Rubenstein. (2004). Combining strategies to identify reserves in fragmented landscapes. Conservation Biology 18: 1121-1131. (pdf)

Tien, J. H., D. I. Rubenstein & S. A. Levin (2004). A field study on the proximate mechanisms of fish shoaling. Evolutionary Ecology Research 6: 555-565. (pdf)

Rubenstein, D. I. & M. Hack (2004) Natural and sexual selection and the evolution of multi-level societies: insights from zebras with comparisons to primates. Pp. 266-279. In: Sexual Selection in Primates: New and Comparative Perspectives. P. Kappeler and C. P. van Schaik (eds.). Cambridge University Press. (pdf)

Hack, M. A., R. East & D. I. Rubenstein (2002). Status and action plan for the Plains zebra (Equus burchelli). Pp 43-57. In Equids: Zebras, Asses, and Horses. Status Survey and Conservation Action Plan. P. D. Moehlman (ed.). IUCN/SSC Equid Specialist Group. IUCN, Gland, Switzerland. (pdf) (link)

Juang, P., H. Oki, Y. Want, M. Martonosi, L.-S. Peh & D. Rubenstein (2002). Energy-efficient computing for wildlife tracking: Design tradeoffs and early experiences with ZebraNet. In: Architectural Support for Programming Languages and Operating Systems. (Proceedings of ASPLOS-X, San Jose, CA, October 2002). (pdf)

Rubenstein, D. I. (2001). Horses, asses, and zebras. Pp. 482-487. In: Encyclopedia of Mammals. D. W. MacDonald (ed.). Oxford University Press. (pdf)

Hack, M. A. & D. I. Rubenstein (2001). Migration. Pp. 221-234. In: Encyclopedia of Biodiversity, Volume 4. S. A. Levin (ed.). Academic Press. (pdf)

Rubenstein, D. I. (2001). Social behavior. Pp. 295-304. In: Encyclopedia of Biodiversity, Volume 5. S. A. Levin (ed.). Academic Press. (pdf)

Saltz, D., M. Rowen & D. I. Rubenstein (2000). The effect of space-use patterns of reintroduced Asiatic wild ass on effective population size. Conservation Biology 14: 1852-1861. (pdf)

Krause, J., D. J. Hoare, S. Krause, C. K. Hemelrijk & D. I. Rubenstein (2000). Leader-ship in fish shoals. Fish and Fisheries 1: 82-89. (pdf)


Rubenstein, D. I. (1999). Reforming Science & Mathematics Education: Making Inquiry Based Science Self-Sustaining. January 21, 1999.  Online publication.
Merck Institute for Science Education/NSF at their Local Systemic Change Meeting.

Hack, M. & D. I. Rubenstein (1998). Zebra zones. Natural History 107 (2): 26-29. (pdf)

Rubenstein, D. I. (1998). Behavioral ecology and conservation policy: On balancing science, applications, and advocacy. Pp 527-553. In: Behavioral Ecology & Conservation Biology. T. Caro (ed.). Oxford University Press. (pdf)

Krause J., G. D. Ruxton & D. I. Rubenstein (1998). Is there an influence of group size on predator hunting success? Journal of Fish Biology 52: 494-501. (pdf)

Krause, J., G. J. Godin, & D. I. Rubenstein (1998). Group choice as a function of group size difference and assessment time in fish: the influence of species vulnerability to predation. Ethology 104: 68-74. (pdf)

Bumann D., J. Krause & D. I. Rubenstein (1997). Mortality risk of spatial positions in animal groups: the danger of being in the front. Behaviour 134: 1063-1076. (pdf)

Krause J., D. I. Rubenstein & D. Brown (1997). Shoal choice behaviour in fish: the relationship between assessment time and assessment quality. Behaviour 134: 1051-1062. (pdf)

Gueron, S., S. A. Levin & D. I. Rubenstein (1996). The dynamics of mammalian herds: From individuals to aggregations. Journal of Theoretical Biology 182: 85-98. (pdf)

Saltz, D. & D. I. Rubenstein (1995). Population dynamics of a reintroduced Asiatic Wild Ass (Equus hemionus) herd. Ecological Applications 5: 327-335. (pdf)

Rubenstein, D. I. (1994). Princeton’s science advisors: A neural network. Pp. 167-169. In: Scientists, Educators, and National Standards Action at the Local Level, Forum Proceedings. Sigma Xi, The Scientific Research Society.

Rubenstein, D. I. (1994). The ecology of female social behavior in horses, zebras, and asses. Pp. 13-28. In: Animal Societies: Individuals, Interactions, and Organization. P. Jarman and A. Rossiter (eds.). Kyoto University Press. (pdf)

Rubenstein, D. I. (1993). Science and the pursuit of a sustainable world. Ecological Applications 3: 585-587. (pdf)

Rubenstein, D. I. (1993). On the evolution of juvenile lifestyles in mammals. Pp. 38-57. In: Juvenile Primates: Life history, Development, and Behavior. M. E. Pereira and L. A. Fairbanks (eds.). Oxford University Press. (pdf)

Rubenstein, D. I. & M. Hack (1992). Horse signals: the sounds and scents of fury. Evolutionary Ecology 6: 254-260. (pdf)

Rubenstein, D. I. (1991). The greenhouse effect and changes in animal behavior: effects on social structure and life-history strategies. Pp. 180-192. In: Consequences of Global Warming for Biodiversity. R. Peters (ed.). Yale University Press. (pdf)

Ginsburg, J.R., D.I. Rubenstein (1990). Sperm competition and zebra mating behaviour. Behavioral Ecology and Sociobiology 26: 427-434. (pdf)

Rubenstein, D. I. (1990). Critical thinking and biology teaching. Pp. 11-23. In: Continuing Dialogue. R. H. Dollase (ed.). CETE Report, Middlebury College.


Dobson, A., A. Jolly & D. I. Rubenstein (1989). The greenhouse effect and biological diversity. Trends in Ecology and Evolution 4: 64-78. (pdf)

Rubenstein, D. I. & M. E. Hohmann (1989). Parasites and social behavior of island feral horses. Oikos 55: 300-312. (pdf)

Rubenstein, D. I. (1989). Life history and social organization in arid adapted ungulates. Journal of Arid Environments 17: 145-156. (pdf)

Rubenstein, D. I. (1989). Sperm competition in the water strider Gerris remigis. Animal Behaviour 38: 631-636. (pdf)

Rubenstein, D. I. (1987). Alternative reproductive tactics in the spider Meta segmentata. Behavioral Ecology and Sociobiology 20: 229-237. (pdf)

Wrangham, R. W. & D. I. Rubenstein (1986). Social evolution in birds and mammals. Pp. 452-470. In: Ecological Aspects of Social Evolution. D. I. Rubenstein & R. W. Wrangham, (eds.). Princeton University Press, Princeton, NJ. (pdf)

Rubenstein, D. I. & R. W. Wrangham (1986). Socioecology: Origins and trends. Pp. 3-20. In: Ecological Aspects of Social Evolution. D. I. Rubenstein & R. W. Wrangham, (eds.). Princeton University Press, Princeton, NJ. (pdf)

Rubenstein, D. I. (1986). Ecology and sociality in horses and zebras. Pp. 282-302. In: Ecological Aspects of Social Evolution. D. I. Rubenstein & R. W. Wrangham, (eds.). Princeton University Press, Princeton, NJ. (pdf)

Bischoff, R. J., J. L. Gould & D. I. Rubenstein (1985). Tail size and female choice in the guppy (Poecilia reticulata). Behavioral Ecology and Sociobiology 17: 253-255. (pdf)

May, R. M. & D. I Rubenstein (1985). Reproductive strategies. Pp. 1-23. In: Reproduction in Mammals, Volume 4. C. R. Austin and R. V. Short (eds.). Cambridge University Press, Cambridge, UK. (pdf)

Rubenstein, D. I. (1984). Resource acquisition and alternative mating strategies in water striders. American Zoologist 24: 345-353. (pdf)
Horn, H. S., & D. I. Rubenstein (1984). Behavioural adaptations and life history. Pp. 279-300. In: Behavioural Ecology. J. R. Krebs and N. B. Davies (eds.). Blackwell Scientific Publications. Oxford. (pdf)

Schulman, S. R. & D. I. Rubenstein (1983). Kinship, need and the distribution of altruism. American Naturalist 121: 776-788. (pdf)

Rubenstein, D. I. (1982). Reproductive value and behavioural strategies: Coming of age in monkeys and horses. Perspectives in Ethology 5: 469-487. (pdf)

Rubenstein, D. I. (1982). Complexity in evolutionary processes. Pp. 87-89. In: Current Problems in Sociobiology. King's Sociobiology Group (eds.). Cambridge University Press. Cambridge, UK. (pdf)

Rubenstein, D. I. (1982). Risk, uncertainty, and evolutionary strategies. Pp. 91-111. In: Current Problems in Sociobiology. Bertram, B. C. R., Clutton-Brock, T. H., Dunbar, R. I. M., Rubenstein, D. I. & Wrangham, R. W. (eds.). Cambridge University Press, UK. (pdf)

Parker, G. A. & D. I. Rubenstein (1981). Role assessment, reserve strategy, and acquisition of information in asymmetric animal contests. Animal Behaviour 29: 221-40. (pdf)

Rubenstein, D. I. (1981). Combat and communication in the Everglades pygmy sunfish. Animal Behaviour 29: 249-258. (pdf)

Rubenstein, D. I. (1981). Population density, resource patterning, and territoriality in the Everglades pygmy sunfish. Animal Behaviour 29: 155-172. (pdf)

Rubenstein, D. I. (1981). Individual variation and competition in the Everglades pygmy sunfish. Journal of Animal Ecology 50: 337-350. (pdf)

Rubenstein, D. I. (1981). Behavioural ecology of island feral horses. Equine Veterinarian J. 13: 27-34. (pdf)

Rubenstein, D. I. & R. W. Wrangham (1980). Why is altruism toward kin so rare? Zeitschrift fur Tierpsychology 54: 381-387. (pdf)

Rubenstein, D. I. (1980). On the evolution of alternative mating strategies. Pp. 65-100. In: Limits to Action: The Allocation of Individual Behaviour. J. E. R. Stadden (ed.). Academic Press, N.Y. (pdf)

1974 - 1979

Wilbur, H. M., D. I. Rubenstein & L. Fairchild (1978). Sexual selection in toads: the role of female choice and male body size. Evolution 32: 264-270. (pdf)

Rubenstein, D. I. (1978). On predation, competition, and the advantages of group living. Perspectives in Ethology 3: 205-231. (pdf)

Fairchild, L., D. I. Rubenstein, S. T. Patti & P. H. Klopfer (1977). Seasonal changes in the feeding strategies of mixed and single species flocks. Ibis 119: 85-87. (pdf)

Klopfer, P. H. & D. I. Rubenstein (1977). The concept "Privacy" and its biological basis. Journal of Social Issues 33: 52-65. (pdf)

Rubenstein, D. I., R. J. Barnett, R. S. Ridgely & P. H. Klopfer (1977). Adaptive advantages of mixed species feeding flocks in Costa Rica. Ibis 119: 10-21. (pdf)

Hazlett, B. A., D. I. Rubenstein & D. Rittschoff (1975). Starvation, aggression and energy reserves in the crayfish (Orconectes virilis). Crustaceana 28: 11-26. (pdf)

Patti, S. T., D. I. Rubenstein & N. Rubenstein (1974). Distributional notes on the birds of Cayman Brac. Florida Scientist 37: 155-156. (pdf)

Rubenstein, D. I. & B. A. Hazlett (1974). Examination of the antagonistic behavior of the crayfish Orconectes virilis by character analysis. Behavior 50: 193-216. (pdf)

Klopfer, P. H., D. I. Rubenstein, R. S. Ridgely & R. J. Barnett (1974). Migration and species diversity in the tropics. Proceedings of the American Academy of Science, USA 71(2): 339-340. (pdf)

Hazlett, B. A., D. Rittschoff & D. I. Rubenstein (1974). Behavioral biology of the crayfish Orconectes virilis 1. Home ranges. American Midland Naturalist 92: 301-319. (pdf)

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