Staff Directory

Prof Daryl Codron
Zoology and Entomology
Dept Zoology and Entomology
IB 49
Biology Building 24

Short CV

Contact Me:

UFS Animal Ecology Research group

Previous Positions

2013 - 2019: Specialist Scientist, National Museum, Bloemfontein, South Africa

2011 - 2013: DeutscheForschungsGemeinschaft Post-doctoral Fellow, University of Zürich, Switzerland 

2009 - 2011: EU Marie-Curie Post-doctoral Fellow, University of Zürich, Switzerland 

2007 - 2009: Lecturer, University of KwaZulu-Natal, Pietermaritzburg, South Africa

2006 - 2007: Post-doctoral Fellow, National Museum, Bloemfontein, South Africa



2006: PhD (Ecology), University of Cape Town, South Africa 

2021: BSc Hons (Applied Statistics), University of the Free State, South Africa 

2003: MSc (Quaternary Science), University of Cape Town, South Africa 

2001: BSc Hons (Biodiversity and Conservation), Rand Afrikaans University (now the University of Johannesburg), South Africa

1998: BSc (Biological Sciences), Rand Afrikaans University (now the University of Johannesburg), South Africa


Research Interests

Trophic interactions and stability  of multispecies communities

Ecological and environmental determinants of mammalian resource use 

Life history evolution of mammals in relation to diet niches

Stable isotope ecology and its applications in ecology and evolution


Modules Taught

ZLGY3734 - Conservation Ecology (Level 3)

ZLGY/ENTO6852 - Quantitative Ecology (Honours)

ZLGY6824 - Conservation Ecology (Honours)

BTNY6894 - Methods in Palaeoecology (Honours)




Journal Articles

Clauss M, Codron D, Hummel J (2023) Equid nutritional physiology and behavior: an evolutionary perspective. Journal of Equine Veterinary Sciences 124:104265. doi:10.1016/j.jevs.2023.104265

Codron D, Bousman CB, Buschke F, Clauss M, Lewis C, Van der Merwe R, van der Mescht A, Brink JS (2023) Competition drives the evolution of emergent neutrality in the dietary niches of mammalian herbivores. Quaternary International 650:28-39. doi:10.1016/j.quaint.2021.11.002

Martin LF, Codron D, Winkler DE, Tütken T, Hatt J-M, Clauss M (2023) Macroscopic dental measures in guinea pigs (Cavia porcellus) fed natural and pelleted diets of different abrasiveness: implications for wear and compensatory growth in a hypselodont species. Journal of the Royal Society Interface 20:20230012. doi:10.1098/rsif.2023.0012

Clauss M, Tschuor A, Codron D, Hummel J (2022) Reticular contraction frequency and ruminal gas dome development in goats do not differ between grass and browse diets. Journal of Animal Physiology and Animal Nutrition 106:1208-1215. doi:10.1111/jpn.13663

Codron D, Holt S, Wilson B, Horwitz LK (2022) Skeletal allometries in the leopard tortoise (Stigmochelys pardalis): Predicting chelonian body size and mass distributions in archaeozoological assemblages. Quaternary International 614:59-72. doi:10.1016/j.quaint.2021.07.021

Duque-Correa MJ, Clauss M, Hoppe MI, Buyse K, Codron D, Meloro C, Edwards MS (2022) Diet, habitat and flight characteristics correlate with intestine length in birds. Proc Biol Sci 289:20220675. doi:10.1098/rspb.2022.0675

Martin LF, Ackermans NL, Richter H, Kircher PR, Hummel J, Codron D, Clauss M, Hatt J-M (2022) Macrowear effects of external quartz abrasives of different size and concentration in rabbits (Oryctolagus cuniculus). Journal of Experimental Zoology B 338:586-597. doi:10.1002/jez.b.23104

Martin LF, Winkler DE, Ackermans NL, Müller J, Tütken T, Kaiser T, Codron D, Schulz-Kornas E, Hatt J-M, Clauss M (2022) Dental microwear texture analysis correlations in guinea pigs (Cavia porcellus) and sheep (Ovis aries) suggest that dental microwear texture signal consistency is species-specific. Frontiers in Ecology and Evolution 10:958576. doi:10.3389/fevo.2022.958576

Mohale NE, Codron D, Horwitz LK (2022) Stable isotope evidence for mid-Pleistocene paleoenvironmental conditions at the site of Kathu Pan 1 (central interior, South Africa). Quaternary International 614:37-49. doi:10.1016/j.quaint.2021.02.027

Valerio SO, Hummel J, Codron D, Hatt J-M, Clauss M (2022) The ruminant sorting mechanism protects teeth from abrasives. Proceedings of the National Academy of Sciences of the United States of America Biological Sciences 119:e2212447119. doi:org/10.1073/pnas.2212447119

van der Mescht AC, Codron D (2022) Mountains and their ecotones increase landscape heterogeneity and maintain a unique assemblage of grasshoppers in the southern Kalahari. Ecol Entomol. doi:

van der Mescht AC, Lewis C, van der Merwe R, Codron D (2022) Hurry up and sing: early onset of diel calling behaviour and ecological drivers of calling behaviour of Acanthoplus discoidalis. Bioacoustics 32:284-300. doi:10.1080/09524622.2022.2112289

Venter C, Haddad CR, Codron D (2022) A novel approach to determine the surface area of buckspoor spider webs and other irregular-shaped two-dimensional objects. MethodsX 9:101904. doi:org/10.1016/j.mex.2022.101904

Winter N, Clauss M, Codron D, Hummel J, Müller J, Richter H, Kircher PR, Hatt J-M, Martin LF (2022) Sand accumulation in the digestive tract of rabbits (Oryctolagus cuniculus) and guinea pigs (Cavia porcellus). Journal of Morphology 283:5-15. doi:10.1002/jmor.21423

Ackermans NL, Martin LF, Codron D, Kircher PR, Clauss M, Hatt J-M (2021) Confirmation of a wear-compensation mechanism in dental roots of ruminants. Anatomical Record 304:425-436. doi:10.1002/ar.24402

Clauss M, Zerbe P, Bingaman Lackey L, Codron D, Müller DWH (2021) Basic considerations on seasonal breeding in mammals including their testing by comparing natural habitats and zoos. Mammalian Biology 101:373-386. doi:10.1007/s42991-020-00078-y

Duque-Correa MJ, Codron D, Meloro C, McGrosky A, Schiffmann C, Edwards MS, Clauss M (2021) Mammalian intestinal allometry, phylogeny, trophic level and climate. Proceedings of the Royal Society B 288:20202888. doi:10.1098/rspb.2020.2888

Hatt J-M, Codron D, Richter H, Kircher PR, Hummel J, Clauss M (2021) Preliminary evidence for a forestomach washing mechanism in llamas (Lama glama). Mammalian Biology - Zeitschrift für Säugetierkunde 101:941-948. doi:10.1007/s42991-021-00142-1

Holt S, Horwitz LK, Wilson B, Codron D (2021) Leopard tortoise Stigmochelys pardalis (Bell, 1928) mortality caused by electrified fences in central South Africa and its impact on tortoise demography. African Journal of Herpetology 70:32-52. doi:10.1080/21564574.2020.1860140

Hoppe MI, Meloro C, Edwards ME, Codron D, Clauss M, Duque-Correa MJ (2021) Less need for differentiation? Intestinal length of reptiles as compared to mammals. PLOS ONE 16:e0253182. doi:10.1371/journal.pone.0253182.t001

Martin LF, Ackermans NL, Tollefson TN, Kircher PR, Richter H, Hummel J, Codron D, Hatt J-M, Clauss M (2021) Tooth wear, growth, and height in rabbits (Oryctolagus cuniculus) fed pelleted or extruded diets with added sand. Journal of Animal Physiology and Animal Nutrition 106:630-641. doi:10.1111/jpn.13565

Winkler DE, Clauss M, Roelle M, Schulz-Kornas E, Codron D, Kaiser TM, Tütken T (2021) Dental microwear texture gradients in guinea pigs reveal that material properties of the diet affect chewing behaviour. Journal of Experimental Biology 224:242446. doi:10.1242/jeb.242446

Ackermans NL, Martin LF, Codron D, Hummel J, Kircher PR, Richter H, Kaiser TM, Clauss M, Hatt J-M (2020) Mesowear represents a lifetime signal in sheep (Ovis aries) within a long-term feeding experiment. Palaeogeography Palaeoclimatology Palaeoecology 553:109793. doi:10.1016/j.palaeo.2020.109793

De Cuyper A, Meloro C, Abraham AJ, Müller DWH, Codron D, Janssens GPJ, Clauss M (2020) The uneven weight distribution between predators and prey: comparing gut fill between terrestrial herbivores and carnivores. Comparative Biochemistry and Physiology - Part A: Molecular & Integrative Physiology 243:110683. doi:10.1016/j.cbpa.2020.110683

Hatt J-M, Codron D, Ackermans NL, Martin LF, Richter H, Gerspach C, Hummel J, Clauss M (2020) Differences in the rumen washing mechanism in sheep fed diets with abrasives of various concentrations and sizes. Palaeogeography, Palaeoclimatology, Palaeoecology 550:109728. doi:10.1016/j.palaeo.2020.109728

Hohl CJM, Codron D, Kaiser TM, Martin LF, Müller DWH, Hatt J-M, Clauss M (2020) Chewing, dental morphology and wear in tapirs (Tapirus spp.) and a comparison of freeranging and captive specimens. PLOS ONE 15:e0234826. doi:10.1371/journal.pone.0234826

Malindie S, Buschke F, Codron D (2020) Carbon isotope differences between grassland and savanna herbivores reveal environmentally driven rather than phylogenetically conserved niches. Journal of Zoology (London) 311:116-125. doi:10.1111/jzo.12762

Marston CG, Wilkinson DM, Sponheimer M, Codron D, Codron J, O`Regan HJ (2020) ‘Remote’ behavioural ecology: Do megaherbivores consume vegetation in proportion to its presence in the landscape? PeerJ 8:e8622. doi:10.7717/peerj.8622

Martin LF, Krause L, Ulbricht A, Winkler DE, Codron D, Kaiser TM, Müller J, Hummel J, Clauss M, Hatt J-M, Schulz-Kornas E (2020) Dental wear at macro- and microscopic scale in rabbits fed diets of different abrasiveness: A pilot investigation. Palaeogeography Palaeoclimatology Palaeoecology 556:109886. doi:org/10.1016/j.palaeo.2020.109886

van Aardt AC, Codron D, Theron EJ, du Preez PJ (2020) Plant community structure and possible vegetation changes after drought on a granite catena in the Kruger National Park, South Africa. Koedoe 62:a1585. doi:10.4102/koedoe.v62i2.1585

Winkler DE, Schulz-Kornas E, Kaiser TM, Codron D, Leichliter J, Hummel J, Martin LF, Clauss M, Tütken T (2020) The turnover of dental microwear texture: Testing the ”last supper” effect in small mammals in a controlled feeding experiment. Palaeogeography Palaeoclimatology Palaeoecology 557:109930. doi:10.1016/j.palaeo.2020.109930

Clauss M, Müller DWH, Codron D (2019) Within-niche pace of life acceleration as a fundamental evolutionary principle: a mammal pilot test case. Evolutionary Ecology Research 20:385-401.

De Cuyper A, Clauss M, Carbone C, Codron D, Cools A, Hesta M, Janssens G (2019) Predator size and prey size-gut capacity ratios determine kill frequency and carcass production in terrestrial carnivorous mammals. Oikos 128:13-22. doi:10.1111/oik.05488

Ehrlich C, Codron D, Hofmann RR, Hummel J, Clauss M (2019) Comparative omasum anatomy in ruminants: relationships with natural diet, digestive physiology, and general considerations on allometric investigations. Journal of Morphology 280:259-277. doi:10.1002/jmor.20942

Hatt J-M, Codron D, Müller DWH, Ackermans NL, Martin LF, Kircher PR, Hummel J, Clauss M (2019) The rumen washes off abrasives before heavy-duty chewing in ruminants. Mammalian Biology - Zeitschrift für Säugetierkunde 97:104-111. doi:10.1016/j.mambio.2019.06.001

Henry AG, Hutschenreuther A, Paine OO, Leichliter J, Codron D, Codron J, Loudon J, Adolph S, Sponheimer M (2019) Influences on plant nutritional variation and their potential effects on hominin diet selection. Review of Palaeobotany and Palynology 261:18-30. doi:10.1016/j.revpalbo.2018.11.001

Holt S, Horwitz LK, Hoffman J, Codron D (2019) Structural density of the leopard tortoise (Stigmochelys pardalis) shell and its implications for taphonomic research. Journal of Archaeological Science Reports 26:101819. doi:10.1016/j.jasrep.2019.04.008

Martin LF, Winkler DE, Tütken T, Codron D, De Cuyper A, Hatt J-M, Clauss M (2019) The way wear goes – phytolith-based wear on the dentine-enamel system in guinea pigs (Cavia porcellus). Proceedings of the Royal Society B 286:20191921. doi:10.1098/rspb.2019.1921

McGrosky A, Codron D, Müller DWH, Navarrete A, Isler K, Hofmann RR, Clauss M (2019) Gross intestinal morphometry and allometry in ruminants. Journal of Morphology 280:1254-1266. doi:10.1002/jmor.21028

Paine O, Henry A, Codron D, Lambert JB, Sponheimer M, Koppa A, Leichliter J, Codron J (2019) Seasonal and habitat effects on the nutritional properties of savanna vegetation: potential implications for early hominin dietary ecology. Journal of Human Evolution 133:99-107. doi:10.1016/j.jhevol.2019.01.003

Paine OCC, Leichliter JN, Avenant N, Codron D, Lawrence A, Sponheimer M (2019) The ecomorphology of southern African rodent incisors: Potential applications to the hominin fossil record. PLOS ONE 14:e0205476. doi:10.1371/journal.pone.0205476

Przybylo M, Hummel J, Ortmann S, Codron D, Kohlschein G-M, Kilga D, Smithyman J, Przybylo U, Swierk S, Hammer S, Hatt J-M, Górka P, Clauss M (2019) Digesta passage in nondomestic ruminants: separation mechanisms in ‘moose-type’ and ‘cattle-type’ species, and seemingly atypical browsers. Comparative Biochemistry and Physiology - Part A: Molecular & Integrative Physiology 235:180-192. doi:10.1016/j.cbpa.2019.06.010

Schiffmann C, Clauss M, Hoby S, Codron D, Hatt J-M (2019) Body Condition Scores (BCS) in European zoo elephants (Loxodonta africana and Elephas maximus) lifetimes – a longitudinal analysis. Journal of Zoo and Aquarium Research 7:74-86. doi:10.19227/jzar.v7i2.375

Schiffmann C, Hatt J-M, Hoby S, Codron D, Clauss M (2019) Elephant body mass cyclicity suggests effect of molar progression on chewing efficiency. Mammalian Biology - Zeitschrift für Säugetierkunde 96:81-86. doi:10.1016/j.mambio.2018.12.004

Sponheimer M, Clauss M, Codron D (2019) Dietary evolution: the panda paradox. Current Biology 29:R417-R419. doi:10.1016/j.cub.2019.04.045

Codron D, Clauss M, Codron J, Tütken T (2018) Within trophic level shifts in collagen–carbonate stable carbon isotope spacing are propagated by diet and digestive physiology in large mammal herbivores. Ecology and Evolution 8:3983-3995. doi:10.1002/ece3.3786

Codron D, Radloff FGT, Codron J, Kerley GIH, Tambling CJ (2018) Meso-carnivore niche expansion in response to an apex predator`s reintroduction - a stable isotope approach. African Journal of Wildlife Research 48: 10.3957/056./048.013004. doi:10.3957/056.048.013004

Codron J, Avenant NL, Wigley-Coetsee C, Codron D (2018) Carnivore stable carbon isotope niches reflect predator-prey size relationships in African savannas. Integrative Zoology 13:166-179. doi:10.1111/1749-4877.12290

Holt S, Codron D, Horwitz LK (2018) Bone mineral density in the leopard tortoise: implications for inter-taxon variation and bone survivorship in an archaeozoological assemblage. Quaternary International 495:64-78. doi:10.1016/j.quaint.2018.04.020

Mutirwara R, Radloff FGT, Codron D (2018) Growth rate and stable carbon and nitrogen isotope trophic discrimination factors of lion and leopard whiskers. Rapid Communications in Mass Spectrometry 32:33-47. doi:10.1002/rcm.8003

Paine OCC, Koppa A, Henry AG, Leichliter JN, Codron D, Codron J, Lambert JE, Sponheimer M (2018) Grass leaves as potential hominin dietary resources. Journal of Human Evolution 117:44-52. doi:10.1016/j.jhevol.2017.10.013

Scott L, Codron D, Horwitz LK (2018) Sub-Saharan Archaeology, Zooarchaeology and Paleoenvironments - Papers in Honour of James Simpson Brink on his 60th Birthday. Quaternary International 495:1-3. doi:10.1016/j.quaint.2018.10.034

Clauss M, Fritz J, Tschuor A, Braun U, Hummel J, Codron D (2017) Dry matter and digesta particle size gradients along the goat digestive tract on grass and browse diets. Journal of Animal Physiology and Animal Nutrition 101:61-69. doi:10.1111/jpn.12505

Codron J, Botha-Brink J, Codron D, Huttenlocker A, Angielczyk KD (2017) Predator–prey interactions amongst Permo-Triassic terrestrial vertebrates as a deterministic factor influencing faunal collapse and turnover. Journal of Evolutionary Biology 30:40-54. doi:10.1111/jeb.12983

Gillis-Germitsch N, Vybiral P-R, Codron D, Clauss M, Kotze A, Mitchell EP (2017) Intrinsic factors, adrenal gland morphology, and disease burden in captive cheetahs (Acinonyx jubatus) in South Africa. Zoo Biology 36:40-49. doi:10.1002/zoo.21341

Leichliter J, Sandberg P, Passey B, Codron D, Avenant N, Paine O, Codron J, de Ruiter D, Sponheimer M (2017) Stable carbon isotope ecology of modern small mammals from the Sterkfontein Valley: implications for habitat reconstruction in mosaic environments. Palaeogeography Palaeoclimatology Palaeoecology 485:57-67. doi:10.1016/j.palaeo.2017.06.003

Richard OK, Codron D, Hagen KB, Südekum K-H, Clauss M (2017) Little differences in digestive efficiency for protein and fat in mammals of different trophic guilds and digestive strategies: data constraints or fundamental functional similarity? Journal of Animal Physiology and Animal Nutrition 101 (suppl 1):127-141. doi:10.1111/jpn.12657

Botha-Brink J, Codron D, Huttenlocker AK, Angielczyk KD, Ruta M (2016) Breeding young as a survival strategy during Earth`s greatest mass extinction. Scientific Reports 6:24053. doi:10.1038/srep24053

Codron D, Codron J, Sponheimer M, Clauss M (2016) Within-population isotopic niche variability in savanna mammals: disparity between carnivores and herbivores. Frontiers in Ecology and Evolution 4:15. doi:10.3389/fevo.2016.00015

Djagoun CAMS, Codron D, Sealy J, Mensah GA, Sinsin B (2016) Isotopic niche structure of a mammalian herbivore assemblage from a West African savanna: Body mass and seasonality effect. Mammalian Biology 81:644-650. doi:10.1016/j.mambio.2016.09.001

Leichliter JN, Sponheimer M, Avenant NL, Sandberg PA, Paine OCC, Codron D, Codron J, Passey BH (2016) Small mammal insectivore stable carbon isotope compositions as habitat proxies in a South African savanna ecosystem. Journal of Archaeological Science Reports 8:335-345. doi:10.1016/j.jasrep.2016.05.013

Codron J, Duffy KJ, Avenant NL, Sponheimer M, Leichliter J, Paine O, Sandberg P, Codron D (2015) Stable isotope evidence for trophic niche partitioning in a South African savanna rodent community. Current Zoology 61:397-411. doi:10.1093/czoolo/61.3.397

Müller J, Clauss M, Codron D, Schulz E, Hummel J, Kircher P, Hatt J-M (2015) Tooth length and incisal wear and growth in guinea pigs (Cavia porcellus) fed diets of different abrasiveness. Journal of Animal Physiology and Animal Nutrition 99:591-604. doi:10.1111/jpn.12226

Carbone C, Codron D, Scofield C, Clauss M, Bielby J (2014) Geometric factors influencing the diet of vertebrate predators in marine and terrestrial environments. Ecology Letters 17:1553-1559. doi:10.1111/ele.12375

Clauss M, Dittmann MT, Müller DWH, Zerbe P, Codron D (2014) Low scaling of a life history variable: analysing eutherian gestation periods with and without phylogeny-informed statistics. Mammalian Biology 79:9-16. doi:10.1016/j.mambio.2013.01.002

Clauss M, Schiele K, Ortmann S, Fritz J, Codron D, Hummel J, Kienzle E (2014) The effect of very low food intake on digestive physiology and forage digestibility in horses. Journal of Animal Physiology and Animal Nutrition 98:107-118. doi:10.1111/jpn.12053

Gattiker C, Espie I, Kotze A, Lane EP, Codron D, Clauss M (2014) Diet and diet-related disorders in captive ruminants at the National Zoological Gardens of South Africa. Zoo Biology 33:426-432. doi:10.1002/zoo.21150

Lane EP, Clauss M, Kock ND, Hill FWG, Majok AA, Kotze A, Codron D (2014) Body condition and ruminal morphology responses of free-ranging impala (Aepyceros melampus) to changes in diet. European Journal of Wildlife Research 60:599-612. doi:10.1007/s10344-014-0824-1

Müller J, Clauss M, Codron D, Schulz E, Hummel J, Fortelius M, Kircher P, Hatt J-M (2014) Growth and wear of incisor and cheek teeth in domestic rabbits (Oryctolagus cuniculus) fed diets of different abrasiveness. Journal of Experimental Zoology 321A:283-298. doi:10.1002/jez.1864

Ritz J, Codron D, Wenger S, REnsch EE, Hatt J-M, Braun U, Clauss M (2014) Ruminal pH in cattle (Bos primigenius f. taurus) and moose (Alces alces) under different feeding conditions: a pilot investigation. Journal of Zoo and Aquarium Research 2:44-51. doi:10.5167/uzh-95706

Taylor LA, Müller DWH, Schwitzer C, Kaiser TM, Codron D, Schulz E, Clauss M (2014) Tooth wear in captive rhinos differs from that of free-ranging conspecifics. Contributions to Zoology 83:107-117. doi:10.5167/uzh-94211

Clauss M, Dittmann MT, Müller DWH, Meloro C, Codron D (2013) Bergmann’s rule in mammals: a cross-species interspecific pattern. Oikos 122:1465-1472. doi:10.1111/j.1600-0706.2013.00463.x

Clauss M, Müller DWH, Codron D (2013) Source references and the scientist’s mind-map: Harvard vs. Vancouver style. Journal of Scholarly Publishing 44:274-282. doi:10.3138/jsp.44.3.005

Clauss M, Steuer P, Müller DWH, Codron D, Hummel J (2013) Herbivory and body size: allometries of diet quality and gastrointestinal physiology, and implications for herbivore ecology and dinosaur gigantism. PLOS ONE 8:e68714. doi:10.1371/journal.pone.0068714

Codron D, Carbone C, Clauss M (2013) Ecological interactions in dinosaur communities: Influences of small offspring and complex ontogenetic life histories. PLoS ONE 8:e77110. doi:10.1371/journal.pone.0077110

Codron D, Carbone C, Müller DWH, Clauss M (2013) Ecological modelling, size distributions and taphonomic size bias in dinosaur faunas: reply to Brown et al. Biology Letters doi: 10.1098/rsbl.2012.0922.

Codron J, Kirkman K, Duffy KJ, Sponheimer M, Lee-Thorp JA, Ganswindt A, Clauss M, Codron D (2013) Stable isotope turnover and variability in tail hairs of captive and free-ranging African elephants (Loxodonta africana) reveal dietary niche differences within populations. Canadian Journal of Zoology 91:124-134. doi:10.1139/cjz-2012-0155

Codron J, Lee-Thorp JA, Sponheimer M, Codron D (2013) Plant stable isotope composition across habitat gradients in a semi-arid savanna: implications for environmental reconstruction. Journal of Quaternary Science 28:301-310. doi:10.1002/jqs.2614

Djagoun CAMS, Codron D, Sealy J, Mensah GA, Sinsin B (2013) Stable carbon isotope analysis of the diets of West African bovids in Pendjari Biosphere Reserve, Northern Benin. South African Journal of Wildlife Research 43:33-43. doi:10.3957/056.043.0107

Kaiser DP, Müller DWH, Fortelius M, Schulze E, Codron D, Clauss M (2013) Hypsodonty and tooth facet development in relation to diet and habitat in herbivorous ungulates: implications for understanding tooth wear. Mammal Review 43:34-46. doi:10.1111/j.1365-2907.2011.00203.x

Lauper M, Lechner I, Barboza PS, Collins WB, Hummel J, Codron D, Clauss M (2013) Rumination of different-sized particles in muskoxen (Ovibos moschatus) and moose (Alces alces) on grass and browse diets, and implications for rumination in different ruminant feeding types. Mammalian Biology 78:142-152. doi:10.1016/j.mambio.2012.06.001

Müller DWH, Codron D, Meloro C, Munn A, Schwarm A, Hummel J, Clauss M (2013) Assessing the Jarman–Bell Principle: scaling of intake, digestibility, retention time and gut fill with body mass in mammalian herbivores. Comparative Biochemistry and Physiology A 164:129-140. doi:10.1016/j.cbpa.2012.09.018

Ritz J, Hofer K, Hofer E, Hackländer K, Immekus D, Codron D, Clauss M (2013) Forestomach pH in hunted roe deer (Capreolus capreolus) in relation to forestomach region, time of measurement, supplemental feeding, and a comparison among wild ruminant species. European Journal of Wildlife Research 59:505-517. doi:10.1007/s10344-013-0698-7

Taylor LA, Kaiser TM, Schwitzer C, Müller DWH, Codron D, Clauss M, Schulz E (2013) Detecting inter-cusp and inter-tooth weat patterns in rhinocerotids. PLoS ONE 8:e80921. doi:10.1371/journal.pone.0080921

Wernick MB, Steinmetz HW, Martin-Jurado O, Howard J, Vogt R, Codron D, Hatt J-M (2013) Comparison of fluid types for resuscitation in acute hemorrhagic shock and evaluation of gastric luminal and transcutaneous PCO2 in Leghorn Chickens. Journal of Avian Medicine and Surgery 27:109-119. doi:10.1647/2012-018

Codron D, Carbone C, Müller DWH, Clauss M (2012) Ontogenetic niche shifts in dinosaurs influenced size, diversity and extinction in terrestrial vertebrates. Biology Letters 8:620-623. doi:10.1098/rsbl.2012.0240

Codron D, Sponheimer M, Codron J, Hammer S, Tschuor A, Braun U, Bernasconi SM, Clauss M (2012) Tracking the fate of digesta 13C and 15N compositions along the ruminant gastrointestinal tract: does digestion influence the relationship between diet and faeces? European Journal of Wildlife Research 58:303-313. doi:10.1007/s10344-011-0581-3

Codron D, Sponheimer M, Codron J, Newton I, Lanham JL, Clauss M (2012) The confounding effects of source isotopic heterogeneity on consumer-diet and tissue-tissue stable isotope relationships. Oecologia 169:939-953. doi:10.1007/s00442-012-2274-3

Codron J, Codron D, Sponheimer M, Kirkman K, Duffy KJ, Raubenheimer EJ, Mélice JL, Grant R, Clauss M, Lee-Thorp JA (2012) Stable isotope series from elephant ivory reveal lifetime histories of a true dietary generalist. Proceedings of the Royal Society B 279:2433-2441. doi:10.1098/rspb.2011.2472

Müller DWH, Codron D, Werner J, Fritz J, Hummel J, Griebeler EM, Clauss M (2012) Dichotomy of eutherian reproduction and metabolism. Oikos 121:102-115. doi:10.1111/j.1600-0706.2011.19505.x

Zerbe P, Clauss M, Codron D, Lackey LB, Rensch E, Streich WJ, Hatt J-M, Müller DWH (2012) Reproductive seasonality in captive wild ruminants: implications for biogeographical adaptation, photoperiodic control, and life history. Biological Reviews (Cambridge) 87:965-990. doi:10.1111/j.1469-185X.2012.00238.x

Clauss M, Lechner I, Barboza P, Collins W, Tervoort TA, Südekum K-H, Codron D, Hummel J (2011) The effect of size and density on the mean retention time of particles in the reticulorumen of cattle (Bos primigenius f. taurus), muskoxen (Ovibos moschatus) and moose (Alces alces). British Journal of Nutrition 105:634-644. doi:10.1017/S0007114510004101

Clauss M, Lunt N, Ortmann S, Plowman A, Codron D, Hummel J (2011) Fluid and particle passage in three duiker species. European Journal of Wildlife Research 57:143-148. doi:10.1007/s10344-010-0407-8

Codron D, Codron J, Sponheimer M, Bernasconi SM, Clauss M (2011) When animals are not quite what they eat: Diet digestibility influences 13C-incorporation rates and apparent discrimination in a mixed-feeding herbivore. Canadian Journal of Zoology 89:453-465. doi:10.1139/Z11-010

Codron D, Hull J, Brink JS, Codron J, Ward D, Clauss M (2011) Effect of competition on niche dynamics of syntopic grazing ungulates: contrasting the predictions of habitat selection models using stable isotope analysis. Evolutionary Ecology Research 13:217-235. doi:10.5167/uzh-49216

Codron J, Codron D, Lee-Thorp JA, Sponheimer M, Kirkman K, Duffy KJ, Sealy J (2011) Landscape-scale feeding patterns of African elephant inferred from carbon isotope analysis of feces. Oecologia 165:89-99. doi:10.1007/s00442-010-1835-6

Copeland SR, Sponheimer M, de Ruiter DJ, Lee-Thorp JA, Codron D, le Roux PJ, Grimes V, Richards MP (2011) Strontium isotope evidence for landscape use by early hominins. Nature 474:76-79. doi:10.1038/nature10149

Hummel J, Findeisen E, Südekum K-H, Ruf I, Kaiser T, Bucher M, M. C, Codron D (2011) Another one bites the dust: faecal silica levels in large herbivores correlate with high-crowned teeth. Proceedings of the Royal Society B 278:1742-1747. doi:10.1098/rspb.2010.1939

Müller DWH, Caton J, Codron D, Schwarm A, Lentle R, Streich WJ, Hummel J, Clauss M (2011) Phylogenetic constraints on digesta separation: Variation in fluid throughput in the digestive tract in mammalian herbivores. Comparative Biochemistry and Physiology A 160:207-220. doi:10.1016/j.cbpa.2011.06.004

Müller DWH, Zerbe P, Codron D, Clauss M, Hatt J-M (2011) A long life among ruminants: giraffids and other special cases. Schweizer Archiv fuer Tierheilkunde 153:515-519. doi:10.1024/0036-7281/a000263

Pirovino M, Heistermann M, Zimmermann N, Zingg R, Clauss M, Codron D, Kaup F-J, Steinmetz HW (2011) Fecal glucocorticoid measurements and their relation to rearing, behavior, and environmental factors in the population of pileated gibbons (Hylobates pileatus) held in European zoos. International journal of primatology 32:1161-1178. doi:10.1007/s10764-011-9532-9

Tschudin A, Clauss M, Codron D, Hatt J-M (2011) Preference of rabbits for drinking from open dishes versus nipple drinkers. Veterinary Record 168:190-191. doi:10.1136/vr.c6150

Tschudin A, Clauss M, Codron D, Hatt J-M (2011) Water intake in domestic rabbits (Oryctolagus cuniculus) from open dishes and nipple drinkers under different water and feeding regimes. Journal of Animal Physiology and Animal Nutrition 95:499-511. doi:10.1111/j.1439-0396.2010.01077.x

Codron D, Clauss M (2010) Rumen physiology constrains diet niche: linking digestive physiology and food selection across wild ruminant species. Canadian Journal of Zoology 88:1129-1138. doi:10.1139/Z10-077

Copeland SR, Sponheimer M, Lee-Thorp JA, de Ruiter DJ, le Roux PJ, Grimes V, Codron D, Codron J, Berger L, Richards MP (2010) Using strontium isotopes to study site accumulation processes. Journal of Taphonomy 8:115-127. doi:10.5167/uzh-38862

Codron D, Codron J (2009) Reliability of d13C and d15N in faeces for reconstructing savanna herbivore diet. Mammalian Biology 74:36–48. doi:10.1016/j.mambio.2007.12.005

Codron D, Codron J, Lee-Thorp JA, Sponheimer M, Grant CC, Brink JS (2009) Stable isotope evidence for nutritional stress, competition, and loss of functional habitat as factors limiting recovery of rare antelope in southern Africa. Journal of Arid Environments 73:449–457. doi:10.1016/j.jaridenv.2008.12.003

Copeland SR, Sponheimer M, Spinage CA, Lee-Thorp JA, Codron D, Reed KE (2009) Stable isotope evidence for impala Aepyceros melampus diets at Akagera National Park, Rwanda. African Journal of Ecology 47:490-501. doi:10.1111/j.1365-2028.2008.00969.x

Sponheimer M, Codron D, Passey BH, Lee-Thorp JA, De Ruiter D, Cerling TE (2009) Using carbon isotopes to track dietary change in modern, historical, and ancient primates: a look at the methods and a discussion of fallback foods in australopith diets. American Journal of Physical Anthropology 140:661-670. doi:10.1002/ajpa.21111

Codron D, Brink JS, Rossouw L, Clauss M (2008) The evolution of ecological specialization in southern African ungulates: competition or physical environmental turnover? Oikos 117:344-353. doi:10.1111/j.2007.0030-1299.16387.x

Codron D, Brink JS, Rossouw L, Clauss M, Codron J, Lee-Thorp JA, Sponheimer M (2008) Functional differentiation of African grazing ruminants: an example of specialized adaptations to very small changes in diet. Biological Journal of the Linnaean Society 94:755–764. doi:10.1111/j.1095-8312.2008.01028.x

Codron D, Lee-Thorp JA, Sponheimer M, de Ruiter D, Codron J (2008) What insights can baboon feeding ecology provide for early hominin niche differentiation? International journal of primatology 29:757-772. doi:10.1007/s10764-008-9261-x

Codron D, Brink JS (2007) Trophic ecology of two savanna grazers, blue wildebeest Connochaetes taurinus and black wildebeest Connochaetes gnou. European Journal of Wildlife Research 53:90-99. doi:10.1007/s10344-006-0070-2

Codron D, Codron J, Lee-Thorp JA, Sponheimer M, de Ruiter D, Brink JS (2007) Stable isotope characterization of mammalian predator–prey relationships in a South African savanna. European Journal of Wildlife Research 53:161-170. doi:10.1007/s10344-006-0075-x

Codron D, Codron J, Lee-Thorp JA, Sponheimer M, de Ruiter D, Sealy J, Grant R, Fourie N (2007) Diets of savanna ungulates from stable carbon isotope composition of faeces. Journal of Zoology London 273:21-29. doi:10.1111/j.1469-7998.2007.00292.x

Codron D, Lee-Thorp JA, Sponheimer M, Codron J (2007) Nutritional content of savanna plant foods: implications for browser/grazer models of ungulate diversification. European Journal of Wildlife Research 53:100-111. doi:10.1007/s10344-006-0071-1

Codron D, Lee-Thorp JA, Sponheimer M, Codron J (2007) Stable carbon isotope reconstruction of ungulate diet changes through the seasonal cycle. South African Journal of Wildlife Research 37:117-125. doi:10.3957/0379-4369-37.2.117

Codron D, Lee-Thorp JA, Sponheimer M, Codron J, de Ruiter D, Brink JS (2007) Significance of diet type and diet quality for ecological diversity of African ungulates. Journal of Animal Ecology 76:526-537. doi:10.1111/j.1365-2656.2007.01222.x

Codron D, Codron J, Lee-Thorp JA, Sponheimer M, de Ruiter D, Brink JS (2006) Dietary variation in impala Aepyceros melampus recorded by carbon isotope composition of feces. Acta Zoologica Sinica 52:1015-1025. doi:10.5167/uzh-25360

Codron D, Lee-Thorp JA, Sponheimer M, de Ruiter D, Codron J (2006) Inter- and intra-habitat dietary variability of chacma baboons (Papio ursinus) in South African savannas based on fecal d13C, d15N and %N. American Journal of Physical Anthropology 129:204-214. doi:10.1002/ajpa.20253

Codron J, Lee-Thorp JA, Sponheimer M, Codron D, Grant RC, De Ruiter DJ (2006) Elephant (Loxodonta africana) diets in Kruger National Park, South Africa: spatial and landscape differences. Journal of Mammalogy 87:27-34. doi:10.1644/05-MAMM-A-017R1.1

Sponheimer M, Loudon JE, Codron D, Howells ME, Pruetz JD, Codron J, De Ruiter D (2006) Do savanna chimpanzees consume C4 resources? Journal of Human Evolution 51:128-133. doi:10.1016/j.jhevol.2006.02.002

Codron D, Codron J, Lee-Thorp JA, Sponheimer M, de Ruiter D (2005) Animal diets in the Waterberg based on stable isotopic composition of faeces. South African Journal of Wildlife Research 35:43-52. doi:10.5167/uzh-25362

Codron D, Codron J, Sponheimer M, Lee-Thorp JA, Robinson T, Grant CC, de Ruiter D (2005) Assessing diet in savanna herbivores using stable carbon isotope ratios of faeces. Koedoe 48:115-124. doi:10.4102/koedoe.v48i1.170

Codron D, Luyt J, Lee-Thorp JA, Sponheimer M, de Ruiter DJ, Codron J (2005) Reply to Thackeray. South African Journal of Science 101:484.

Codron D, Luyt J, Lee-Thorp JA, Sponheimer M, de Ruiter DJ, Codron J (2005) Utilization of savanna-based resources by Plio-Pleistocene baboons. South African Journal of Science 101:245-248. doi:10.5167/uzh-25352

Codron J, Codron D, Lee-Thorp JA, Sponheimer M, Bond WJ, De Ruiter D, Grant CC (2005) Taxonomic, anatomical, and spatio-temporal variations in the stable carbon and nitrogen isotopic compositions of plants from an African savanna. Journal of Archaeological Science 32:1757-1772. doi:10.1016/j.jas.2005.06.006

Sponheimer M, Lee-Thorp JA, De Ruiter D, Codron D, Codron J, Baugh A, Thackeray JF (2005) Hominins, sedges, and termites: new carbon isotope data from the Sterkfontein Valley and Kruger National Park. Journal of Human Evolution 48:301-312. doi:10.1016/j.jhevol.2004.11.008

Sponheimer M, Grant CC, De Ruiter D, Lee-Thorp JA, Codron D, Codron J (2003) Diets of impala from Kruger National Park: evidence from stable carbon isotopes. Koedoe 46:101-106. doi:10.4102/koedoe.v46i1.43

Book Chapters

Bousman B, Codron D, Gowlett J, Herries AIR, Rossouw L, Toffolo M (2023) Cornelia-Uitzoek, South Africa. In: Beyin A, Wright DK, Wilkins J,Olszewski DI (eds.) Handbook of Pleistocene Archaeology of Africa : Hominin behavior, geography, and chronology. p1327-1347. Springer International Publishing, Cham. doi:10.1007/978-3-031-20290-2_86

Brink JS, Gowlett JAJ, Herries AIR, Hancox J, Moggi-Cecchi J, Codron D, Rossouw L, Trower G, Stone OML, Bousman B, Grün R, Souron A (2022) Cornelia-Uitzoek: Paleoecology, Archeology, and Geochronology. In: Bobe R,Reynolds SC (eds.) African Paleoecology and Human Evolution. p120-134. Cambridge University Press, Cambridge. doi:10.1017/9781139696470.012

Codron D (2020) Evolution of large mammal herbivores in savannas. In: Scogings PF,Sankaran M (eds.) Savanna woody plants and large herbivores. p213-234. John Wiley & Sons, West Sussex, UK.

Codron D, Hofmann RR, Clauss M (2019) Morphological and physiological adaptations for browsing and grazing. In: Gordon IJ,Prins HHT (eds.) The impact of browsing and grazing. p81-125. Springer, Berlin, Germany.

Codron D (2013) African megaherbivores. In: Grzimek B, MacLeod N, Archibald JD, Levin PS,Blanchfield DS (eds.) Grzimek`s animal life encyclopedia Extinction No. ISBN: 9781414490700. p385-395. Gale/Cengage Learning, Detroit.

Sponheimer M, Lee-Thorp JA, Codron D (2013) A brief update on developments in early hominin biogeochemistry. In: Armitage RA,Burton JH (eds.) Archaeological Chemistry VIII ACS Symposium Series. p295-307. American Chemical Society, Washington DC.



Courses Presented




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