Werner Conradie Publications

Peer Reviewed Publications


Portillo F, Stanley EL, Branch WR, Conradie W, Rödel M-O, Penner J, et al. (2019) Evolutionary history of burrowing asps (Lamprophiidae: Atractaspidinae) with emphasis on fang evolution and prey selection. PLoS ONE 14(4): e0214889.  

Weinell, J.L., Branch, W.R., Colston, T.J., Jackman, T.R., Kuhn, A., Conradie, W., Bauer, A.M., A species-level phylogeny of Trachylepis (Scincidae: Mabuyinae) provides insight into their reproductive mode evolution, Molecular Phylogenetics and Evolution (2019), doi:

Conradie, W., Branch, W.R., and Watson, G. 2019. Type specimens in the Port Elizabeth Museum, South Africa, including the historically important Albany Museum collection. Part 2: Reptiles (Squamata). Zootaxa 4576 (1): 001 045.

Tolley, K.A., Weeber, Maritz, B., Verburgt, L., Bates, M.F., Conradie, W., Hofmeyr, M., Turner, A.T., da Silva, J. Alexander, G. 2019. No safe haven: Protection levels show imperilled South African reptiles not sufficiently safe-guarded despite low average extinction risk. Biological Conservation 233 (2019) 61–72. 

Baptista N, Conradie W. Vaz Pinto P, Branch WR 2019. The Amphibians of Angola: Early Studies and the Current State of Knowledge. Pp 243–281. In: Biodiversity of Angola. Science & Conservation: A Modern Synthe­sis. Huntley BJ, Ferrand N, Russo V, Lages F. Springer, Switzerland. 552 p. 

Branch WR, Baptista N, Vaz Pinto P, Conradie W. 2019. The Reptiles of Angola – History, updated checklists, endemism, hot spots, and future directions for re­search. Pp 283–326. In: Biodiversity of Angola. Science & Conservation: A Modern Synthe­sis. Huntley BJ, Ferrand N, Russo V, Lages F. Springer, Switzerland. 552 p. 

Zhao Z, Verdú-Ricoy J, Mohlakoana S, Jordaan A, Conradie W, Heideman N. 2019. Unexpected phylogenetic relationships within the world's largest limbless skink species (Acontias plumbeus) highlight the need for a review of the taxonomic status of Acontias poecilus. Journal of Zoological Systematics and Evolutionary Research 2019;00:1–16.

Branch, W. R., Verburgt, L., Bayliss, J., Kucharzewski, C., Rodel, M.-O., & Conradie, W. 2019. New records of the Large-eyed Green Snake, Philothamnus macrops (Boulenger 1895), from Mozambique. Herpetology Notes, 12: 19-29. [PDF

Engelbrecht, H.M., Branch, W.R., Greenbaum, E., Alexander, G.J., Jackson, K., Burger, M., Conradie, W., Kusamba, C., Zassi-Boulou, A-G. & Tolley, K.A. 2019. Diversifying into the branches: species boundaries in African green and bush snakes, Philothamnus (Serpentes: Colubridae). Molecular phylogenetics and evolution 130: 357-365


Tolley, K.A., Conradie, W., Harvey, J., Measey, J., & Blackburn, D.G. 2018 Molecular phylogenetics reveals a complex history underlying cryptic diversity in the Bush Squeaker Frog (Arthroleptis wahlbergii) in southern Africa. African Zoology, 1-16 DOI: 10.1080/15627020.2018.1517608

Nielsen, S.V., Daniels, S.R., Conradie, W, Heinicke, M.P., & Noonan, B.P. 2018. Multilocus phylogenetics in a widespread African anuran lineage (Brevicipitidae: Breviceps) reveals patterns of diversity reflecting geoclimatic change. J Biogeogr. 2018;00:1–13.

Conradie, W., Busschau, T. & Edwards, S. 2018. Two new species of Acontias form the Mpumalanga Highveld escarpment of South Africa. Zootaxa 4429(1):089-106.

Portillo, F., Branch, W.R., Conradie, W., Rödel, M-O., Penner, J., Barej, M.F., Kusamba, C., Muninga, W.M., Aristote, M.M., Bauer, A.M., Trape, J-F., Nagy, Z.T., Carlino, P., Pauwels, O.S.G., Menegon, M., Burger, M., Mazuch, T., Jackson, K., Hughes, D.F., Behangana, M., Greenbaum, E., Phylogeny and biogeography of the African burrowing snake subfamily Aparallactinae (Squamata: Lamprophiidae), Molecular Phylogenetics and Evolution 127 (2018):288-303. doi:

Broadley, D.G., Tolley, A., Conradie, W., Wishart, S., Trape, J-F., Burger, M., Kusamba, C.,  Zassi-Boulou, A-G. & Greenbaum, E. 2018.  A phylogeny and genus-level revision of the African file snakes Gonionotophis Boulenger (Squamata: Lamprophiidae), African Journal of Herpetology, 67:1, 43-60, DOI: 10.1080/21564574.2018.1423578 

Conradie, W., Bittencourt-Silva, G.B., Farooq, H.M., Loader, S.P., Menegon, M. & Tolley K.A. 2018. New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozambique inselbergs, African Journal of Herpetology, 67:1, 61-85, DOI: 10.1080/21564574.2017.1376714

Conradie, W., Verburgt, L., Portik, D.M., Ohler, A., Bwong, B.A. & Lawson, L.P. 2018. A new Reed Frog (Hyperoliidae: Hyperolius) from coastal northeastern Mozambique. Zootaxa, 4379: 177-198,

Jongsma, G.F.M. ,  Barej, M.F. Barratt, C.D., Burger, M., Conradie, W., Ernst, R., Greenbaum, E., Hirschfeld, M.,  Leaché, A.D., Penner, J.,  Portik, D.M., Zassi-Boulou, A., Rödel, MO., &  Blackburn, D.C. 2018. Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saharan Africa, Molecular Phylogenetics and Evolution, Volume 120, March 2018, Pages 274-285, ISSN 1055-7903,


Pietersen, D.W., Pietersen, E.W., & Conradie, W.  2017. Preliminary herpetological survey of the Ngonye Falls and surrounding regions in south-western Zambia. Amphibian & Reptile Conservation 11(1) [Special Section]: 24–43 (e148).[PDF]

 Conradie, W & Bills, R. 2017. Wannabe Ranid: Notes on the morphology and natural history of the Lemaire’s Toad (Bufonidae: Sclerophrys lemairii). Salamandra 53(3):439-444. [PDF]

Branch, W.R., Haacke, H., Vaz Pinto, P., Conradie, W., Baptista, N., Verburgt, V., & Verisimmo, L. 2017. Loveridge’s Angolan geckos, Afroedura karroica bogerti and Pachydactylus scutatus angolensis (Sauria, Gekkonidae): new distribution records, comments on type localities and taxonomic status. Zoosystematics and Evolution, 93: 157-166. [PDF]

Busschau, T., Conradie, W., Jordaan, A., & Daniels, S.R. 2017 Unmasking evolutionary diversity among two closely related South African legless skink species (Acontinae: Acontias) using molecular data. Zoology 121:72-82.


Conradie, W., Reeves, B., Brown, N. & Venter, J.A. 2016. Herpetofauna of the Oviston, Commando Drift and Tsolwana nature reserves in the arid interior of the Eastern Cape Province, South Africa. Indago 32:81-98 [PDF]

Ceríaco et al. 2016. Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences. Zootaxa 4196 (3): 435–445. [PDF]

Conradie, W., Bill, R., & Branch, W.R. 2016. The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern Angola.  Amphibian & Reptile Conservation 10(2) [Special Section]: 6–36 (e126). [PDF]

Conradie, W., Bittencourt-Silva, Engelbrecht, H.M., Loader, S. P., Menegon, M., Nanvonamuquitxo, C., Scott, M. & Tolley, K.A. 2016. Exploration into the hidden world of Mozambique’s sky island forests: new discoveries of reptiles and amphibians. Zoosystematics and Evolution, 92 (2): 163-180. [PDF]

Metallinou, M., Weinell, J.L., Karin, B. R., Conradie, W., Wagner, P., Schmitz, A., Jackman, T. R. & Bauer, A. 2016. A single origin of extreme matrotrophy in African mabyine skinks. Biology Letters, 12: 20160430. [PDF]

Medina, M.F, Bauer, A.M., Branch, W.R., Schmitz, A., Conradie, W., Nagy, Z.T., Hibbitts, T.J., Ernst, R., Portik, D.M., Nielsen, S.V., Colston, T.J., Kusamba, C., Behangana, M., Rödel, M-O. & Greenbaum, E. 2016. Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa, Molecular Phylogenetics and Evolution, 100:409-423. [PDF]

Poynton, J.C., Loader, S.P., Conradie, W., Rodel, M-O &Liedtke, C. 2016. Designation and description of a neotype of Sclerophrys maculata (Hallowell, 1854), and reinstatement of S. pusilla (Mertens, 1937) (Amphibia: Anura: Bufonidae). Zootaxa 4089: 73-94. [PDF]

Conradie, W; Bittencourt-Silva, G., Loader, S.P., Menegon, M., Nanvonamuquitix, X., Kotze, A., Dalton, D.L., Engelbrecht, H.M. & Tolley, K.A. 2016. Batrachochytrium dendrobatidis survey of amphibians in the northern Mozambique ‘Sky Islands’ and low lying areas. Herpetological Review 47: 42-46 [PDF]

Bittencourt-Silva, G., Conradie, W., Siu-Ting, K., Tolley, K.A., Channing, A., Cunningham, M., Farooq, H.M., Menegon, M. & Loader, S.P. 2016. The phylogenetic position and diversity of the enigmatic mongrel frog Nothophryne Poynton, 1963 (Amphibia, Anura). Molecular Phylogenies and Evolution 99: 89-102.[PDF]


Farooq, H., Liedtke, H. C., Bittencourt-Silva, G., Conradie, W., & Loader, S. 2015. The distribution of Mertensophryne anotis with a new record in Northern Mozambique. Herpetology Notes 8: 305-307. [PDF]

Farooq, H.O.M. & Conradie, W. 2015. A second record of Scolecomorphus kirkii Boulenger, 1883 (Gymnophiona: Scolecomorphidae) for Mozambique. Herpetology Notes, 8: 59-62. [PDF]

Venter, A.J. and Conradie, W. 2015. A checklist of the reptiles and amphibians found in the protected areas along the South African Wild Coast, with notes on conservation implications. Koedoe 57 (1): 1-25. 

Conradie, W. and Conradie, C. 2015. Correlation between development and increase of number of labial tooth rows in Ghost Frog tadpoles (Anura: Heleophrynidae). Acta Herpetologica 10(2): 143-148. [PDF]

Conradie, W, Branch, W. R. & Watson, G. 2015. Type specimens in the Port Elizabeth Museum, South Africa, including the historical important AlbanyMuseum collection. Part 1: Amphibians. Zootaxa 3936 (1):042-070 [PDF]


Conradie, W. 2014. The King of the Dwarves: a new cryptic species of Dainty Frog (Anura: Pyxicephalidae: Cacosternum) from the eastern Great Escarpment of South Africa. Zootaxa 3785 (3): 438-452. 


Conradie, W., Branch, W.R., and Tolley, K.A. 2013. Fifty Shades of Grey: giving colour to the poorly known Angolan Ashy reed frog (Hyperoliidae: Hyperolius cinereus), with the description of a new species. Zootaxa 3635 (3): 201-223.

Channing, A., Hillers, A., Lötters, S., Rödel, M.O., Schick, S., Conradie, W, Rödder, R., Mercurio, V., Wagner, P., Dehling, M.J., Du Preez, L.H., Kielgast, J. & Burger, M. 2013. Taxonomy of the super-cryptic Hyperolius nasutus group of Long reed frogs of Africa (Anura: Hyperoliidae), with descriptions of six new species. Zootaxa 3620 (3): 301-350.


Conradie, W., Branch, W.R., Measey, J.G. and Tolley, K.A. 2012. A new species of Hyperolius Rapp, 1842 (Anura: Hyperoliidae) from the Serra da Chela mountains, south-western Angola. Zootaxa3269: 1–17.

Conradie, W., Measey, J.G., Branch, W.R. and Tolley, K.A. 2012. Revised phylogeny of African sand lizards (Pedioplanis), with the description of two new species. African Journal of Herpetology 61: 91-112.

Conradie, W., Tweddle, D., Makinen, T. & Bills, R. 2012 Description of the Hylarana darlingi (Anura: Ranidae) from the Mulanje Area, Malawi. Zootaxa3441:56-58.


Conradie, W., Weldon, C., Smith, K.G. & Du Preez, L.H. 2011. Seasonal pattern of chytridiomycosis in common river frog (Amietia angolensis) tadpoles in the South African Grassland Biome. African Zoology 46(1): 95–102.

Conradie, W., Harvey, J., Kotze, A., Dalton, D.L. and Cunningham, M.J. 2011. Confirmed amphibian chytrid in Mount Mulanje Area, Malawi. Herpetological Review 43(3): 369-371.


Conradie, W., Branch, W.R., Braack, H. and Mason, M. 2010. Notes on the diet of recently metamorphosed Giant African Bullfrogs (Anura: Pyxicephalidae: Pyxicephalus adspersus) and growth increase during the first nine months in a semi-natural habitat. Herpetology Notes, volume 3: 215-219.


Smith, K.G, Weldon, C., Conradie, W & Du Preez, L.H. 2007. Relationships among size, development, and Batrachochytrium dendrobatidis infection in African tadpoles. Diseases of Aquatic Organisms 74: 159-164.


Shorter Communications

Nicolau, G.K., Kemp, L. & Conradie, W. 2018. Geographical Distribution: Panaspis wahlbergii Smith, 1849. African Herp News 59:27–30.

Rebelo, A.R, Zhao, Z, Jordaan, A., Jordaan. P.R. & Conradie, W. 2018. Geographical Distribution Scelotes caffer Peters, 1861. African Herp News 68:53–56.

Rebelo, A.R, Jordaan. P.R. & Conradie, W. 2018. Geographical Distribution Ptenopus garralus maculatus Gray, 1866. African Herp News 68:50–52.

Conradie, W & Herbert, D.G. 2018. The story of the snail and the gecko egg. Biodiversity Observations 9.3:1-2

Conradie, W. & Busschau, T. Geographical Distribution: Macrelaps microlepidotus. African Herp News 67:41-43.

Kemp, L. & Conradie, W. Geographical Distribution: Aspidelaps sulcatus. African Herp News, 65:42-45.

Branch, W.R  & W. Conradie. 2017 Phrynomantis affinis Maximum Size. African Herp New, 65: 12-14.

Conradie, W., Venter, J & Du Preez, B. 2017. Geographical Distribution Note: Tropidosaura montana natalensis. African Herp News, 65:40-42

Conradie, W., Branch W.R. & Herbert, H.. 2017 Tetradactylus seps Diet. African Herp News, 65:14-16

Conradie W., King, F.V., Pressley M., & Leander, A. 2017. Natural History: Gerrhosaurus typicus. African Herp News 64:28-30

Conradie, W; Bittencourt-Silva, G., Loader, S.P., Menegon, M., Nanvonamuquitix, X., Kotze, A., Dalton, D.L., Engelbrecght, H.M. & Tolley, K.A. 2017. Batrachochytrium dendrobatidis survey of amphibians in the northern Mozambique ‘Sky Islands’ and low lying areas. FrogLog 118:95-96

Branch, W.R & Conradie, W. 2013. Naja (Boulengerina) annulata annulata - Geographical Distribution Diet. African Herp News 59:50-53

Conradie, W & Bourquin, S. 2013. Geographical Distributions: Acontias kgalagadi kgalagadi (Lamb, Biswas & Bauer, 2010): African Herp News: 60:29-30.

Conradie, W. 2012. Pedioplanis husabensis: Maximum Size. Herpetological Bulletin 120: 42

Conradie, W. 2012. Pedioplanis lineoocellata lineoocellata: Maximum Size. Herpetological Bulletin 120: 42-43

Conradie, W. 2012. Varanus albigularis - Diet. African Herp News 58:15-16

Conradie, W., Venter, J. & Nicolau, J.R. 2012. Geographical Distribution – Macrelaps microlepidotus. African Herp News 58:19-21

Conradie, W. 2012. Geographical Distribution – Nucras taeniolata. African Herp News 58: 21-22

Conradie, W. & Venter Jan. 2012. Hluleka Nature Reserve: unexpected amphibian diversity. FrogLog 103:22-23.

Conradie, W.  2012. First new frog discovery for Angola in 40 years. FrogLog 103:25.

Nielsen, S.V. and Conradie, W. 2011. Psammophis: Mortality. African Herp News 55:17-18

Conradie, W. and Tarrant, J. 2011. Amatola Toad AWOL: Thirteen years of futile searches. Froglog 97: 24

Conradie, W. and Weldon, C. 2011. Monitoring of the endangered Hewitt’s ghost frog. Froglog 97: 31.

Conradie, W., Bills, R and Bok, A. 2011. Leptopelis bocagii – Maximum Size. African Herp News 53: 42-43.

Conradie, W., Du Preez, L.H., Smith, K.G., & Weldon, C. 2008. Herpetological survey: Frogs of the Vredefort Dome Conservation Area. African Herp News 44:23-25