Botany & Zoology
Welkom by Universiteit Stellenbosch

Prof. Savel Daniels

Evolutionary Genomics Group

Research focus

One of the central focal points of my research is to understand the interplay between species diversity and historical time in southern Africa. In this respect I have focused on evolutionary relationships of a wide number of terrestrial and inland aquatic groups, primarily invertebrates. In this respect I am currently investigating the systematics of South African velvet worms and a revision of the Afrotropical freshwater crab fauna based on DNA sequence data and morphology. I have also in recent years worked extensively on limbless skinks and have recently initiated a large-scale project on tortoises. The main objectives of all these studies are to better understand and document the spectacular biological diversity of South Africa to aid the conservation of these groups.

Current collaboration

South Africa

Dr Michelle Hamer, SANBI, Pretoria (velvet worm systematics)

Prof Neil Heideman, Dean, University of the Free State, Bloemfontein (Systematics of limbless lizards)

Dr Mike Picker, Zoology Department, University of Cape Town, Rondebosch, Cape Town (comparative phylogeography of selected mountain invertebrate species)

Dr Gavin Gouws, SAIAB, Grahamstown, South Africa (aquatic biodiversity)

Prof Retha Hofmeyr, Department of Biodiversity and Conservation, University of the Western Cape, Bellville, South Africa (tortoise phylogeography and systematics)


Prof Hilke Ruhberg, Biozentrum Grindel und Zoologisches Museum, Hamburg, Germany (velvet worm systematics)


Prof Barbara Stewart, Centre of Excellence in Natural Resource Management, University of Western Australia, Australia (comparative phylogeography of selected aquatic invertebrate species)


Dr Trisha Spears, Department of Biological Sciences, Florida State University, Tallahassee, Florida, U.S.A. (Phylogenetics of the Decapod Crustacea)

Prof Keith Crandall, Brighman Young University, Department of Integrative Biology, Provo, Utah, USA
(Systematics of freshwater decapods)

Prof Neil Cumberlidge, Department of Biology, Northern Michigan University, Presque Isle Ave, Marquette, USA (Systematics of freshwater crabs)

Prof Gonzalo Giribet, Museum of Comparative Zoology, Harvard University, Cambridge, USA (velvet worm phylogenetics)



67) Giribet, G., Boyer, S., Baker, C., Fernández, R., Sharma, P., de Divort, B., Daniels, S.R., Harvey, M., S, Neethling., J. A., Griswold, C. E. A molecular phylogeny of the temperate Gondwanan family Pettalidae (Arachnida, Opiliones, Cyphophthalmi) with biogeographic and taxonomic implications. Invertebrate Systematics (submitted)

66) Phiri, E.E., Daniels, S. R. A coalescent multilocus species delimitation approach demonstrates widespread cryptic differentiation within two mountain-living freshwater crab lineages. Invertebrate Systematics (submitted)

65) Daniels, S. R., Klaus, S., Sharma, P., Dambire, C. Unmasking alpha diversity, cladogenesis and biogeographic patterning in a ancient panarthropod lineage (Onychophora: Peripatopsidae: Opisthopatus cinctipes). Cladistics (submitted)

64) Mirimin, L., Galway, N.U.I., Carna Kitchin, N., Impson, D., Clark, P., Richard, J., Daniels, S. R., Roodt-Wilding, R. Genetic and morphologic characterization of Caridina africana (Kingsley, 1882) reveals the presence of alien shrimps in the Cape Floristic Region, South Africa. Journal of Heredity (accepted).

63) Daniels, S.R., Phiri, E.E., Klaus S., Albrecht A., Cumberlidge N. (2015) Multilocus phylogeny of the Afrotropical freshwater crab fauna reveals historical drainage connectivity and transoceanic dispersal since the Eocene. Systematic Biology, 64: 549-567

62) Myburg, A., Daniels, S.R. (2015). Exploring the impact of habitat size on phylogeographic patterning in the Overberg velvet worm Peripatopsis overbergiensis (Onychophora: Peripatopsidae). Journal of Heredity, 106: 296-305.


61) Daniels, S.R., Phiri, E. E., Bayliss J. (2014). Renewed sampling of inland aquatic habitats in southern Africa yields two novel freshwater crab species (Decapoda: Potamonautidae: Potamonautes). Zoological Journal of the Linnean Society, 171: 356-369.

60) Diedericks, G and Daniels, S. R. (2014). Ain’t no mountain high enough, ain’t no valley low enough? Phylogeography of the rupicolous Cape girdled lizard (Cordylus cordulys). Molecular Phylogenetics and Evolution, 71: 234-248.

59) Phiri, E. E., Daniels S. R. (2014). Disentangling the divergence and cladogenesis in the freshwater crab species (Potamonautidae: Potamonautes perlatus sensu lato) in the Cape Fold Mountains, South Africa, with the description of two novel cryptic lineages. Zoological Journal of the Linnean Society, 170: 310-332.

58) Murien, J., Daniels, S. R., Buckley, T., Mayer, G., Giribet. G. A living fossil tale of Pangaean biogeography. Proceedings of the Royal Society of London, Biological Sciences, 281: 1775 (20132648).

57) Cumberlidge, N and Daniels, S. R. (2014). Recognition of two new species of freshwater crabs from the Seychelles based on molecular evidence (Potamoidea: Potamonautidae).  Invertebrate Systematics, 24: 384-356.


56) Phirie, E and Daniels, S. R.  (2013). Hidden in the highlands: the description and phylogenetic position of a novel endemic freshwater crab from Zimbabwe. Invertebrate Systematics, 27: 530-539.

55) Daniels, S. R., McDonald, D.E., Picker, M. (2013). Evolutionary insight into the Peripatopsis balfouri sensu lato species complex (Onychophora: Peripatopsidae) reveals novel lineages and zoogeographic patterning. Zoologica Scripta, 42: 656-674.

54) Ruhberg, H and Daniels, S. R. (2013). Morphological assessment supports the recognition of four novel species in the widely distributed velvet worm Peripatopsis moseleyi (Onychophora: Peripatopsidae). Invertebrate Systematics, 27: 131-145.

53) Engelbrecht, H., Heideman, N. J. L., van Niekerk, A and Daniels, S.R. (2013).  Tracking the impact of Pliocene/Pleistocene sea level and climatic oscillations on the cladogenesis of the Cape legless fossorial skink, Acontias meleagris meleagris. Journal of Biogeography, 40: 492-506.

52) Weldon, C., Daniels, S. R., Clusella-Trullas, S and Chown, S. L. (2013). Metabolic and water loss rates of two cryptic species in the South African velvet worm Opisthopatus cinctipes (Onycophora). Journal of Comparative Physiology –B, 183: 323-332.


51) McDoland, D. E and Daniels, S. R. (2012). Two new velvet worm species from the Western Cape, South Africa. Zootaxa, 3380: 55-68.

50) Daniels, S. R., and Bayliss, J. (2012). Neglected refugia of biodiversity: Mountainous regions in Mozambique and Malawi yield three novel freshwater crab species (Potamonautidae: Potamonautes). Zoological Journal of the Linnean Society, 164: 498-509.

49) McDonald D.E and Daniels, S. R. (2012). Phylogeography of the Cape velvet worm (Onycophora: Peripatopsis capensis) reveals the impact of Plio/Pleistocene climatic oscillations on Afromontane forest in the Western Cape, South Africa. Journal of Evolutionary Biology, 25: 824-835.

48) Daniels, S. R. (2011). Genetic variation in the Critically endangered velvet worm Opisthopatus roseus (Onycophora: Peripatopsidae). African Zoology 46: 419-424.

47) Daniels, S. R. (2011). Reconstructing the colonisation and diversification history of the endemic freshwater crab (Seychellum alluaudi) in the granitic and volcanic Seychelles. Molecular Phylogenetics and Evolution, 61: 534-542.

46) Heideman, N.J.L., Mulcahy, D., Sites. J. W., Hendricks, M.G and Daniels, S.R. (2011) Cryptic diversity and morphological convergence in threatened species of fossorial skinks in the genus Scelotes (Squamata: Scincidae) from the Western Cape Coast of South Africa: Implications for species boundaries, digit reduction and conservation. Molecular Phylogenetics and Evolution, 61: 823-833.

45) Voua Otomo, P., Owojori, Reinceke, S.A, Daniels, S., Reinecke, A. J. (2011). Using estimates of metal bioavailability in the soil and genetic variation of allozymes to investigate heavy metal tolerance in the earthworm Eisenia fetida. Ecotoxicology and Environmental Safety, 74: 2070-2074.

44) Engelbrecht, A, Daniels, S. R., Taylor, P. J and Rambau, R. V. (2011). Cryptic speciation in the southern African vlei rat, Otomys irroratus complex: evidence derived from mitochondrial cyt b and niche modeling.  Biological Journal of the Linnean Society, 104: 192-206.

43) Engelbrecht, A, Taylor, P. J, Daniels, S. R and Rambau, R. V. (2011). Chromosomal polymorphisms in the African vlei rats, Otomys irroratus detected using banding techniques and chromosome painting: inversions, centromeric shifts, and diploid number variation. Cytogenetics and Genome Research, 133: 8-15.

42) Engelbrecht, H. M., Mouton, P. le Fras N. and Daniels, S. R. (2011). Are melanistic populations of the Karoo girdled lizard, Karusasaurus polyzonus, relics or ecotypes? A molecular investigation. African Zoology, 46:146-155.


41) Allwood, J., Gleeson, D., Mayer, G., Daniels, S., Beggs, J. R., Buckley, T. R. (2010). Support for vicariant origin of the New Zealand Onychophora. Journal of Biogeography, 37: 669 – 681.

40) Vargas-Ramírez, M., Vences, M., Branch, W.R., Daniels, S.R., Glaw, F., Hofmeyr, M.D., Kuchling, G., Maran, J., Papenfuss, T.J., Široký, P., Vieites, D.R., Fritz, U (2010). Deep genealogical lineages in the widely distributed African helmeted terrapin: evidence from mitochondrial and nuclear DNA (Testudines: Pelomedusidae: Pelomedusa subrufa). Molecular Phylogenetics and Evolution, 56: 428 – 440.

39) Fritz, U., Daniels, S.R., Hofmeyr, M.D., Gonzalez. J, Barrio-Amaros, CL., Siroky, P., Hundsdorfer, A.K., Stuckas, H. (2010). Mitochondrial phylogeography and subspecies of the wide-ranging sub-Saharan leopard tortoise Stigmochelys pardalis (Reptilia: Testudines: Testudinidae) – a case study for the pitfalls of pseudogenes and GenBank sequences. Journal of Zoological Systematics and Evolutionary Research, 48: 348 -359.

38) Daniels S. R, Ruhberg H. (2010). Molecular and morphological variation in a South African velvet worm, Peripatopsis moseleyi (Onychophora, Peripatopsidae): evidence for cryptic speciation. Journal of Zoology London, 282: 171-179.

37) Gouws, G., Stewart, B. A., Daniels, S. R. (2010) Phylogeographic structure in the gilgie (Decapoda: Parastacidae: Cherax quinquecarinatus) – a south-western Australian freshwater crayfish. Biological Journal of the Linnean Society, 101: 385-402.

36) Daniels, S. R., Hofmeyr, M. D., Henen, B. T., Baard, E. H. (2010). Systematics and phylogeography of a threatened tortoise, the speckled padloper. Animal Conservation, 13: 237-246.

35) Braband, A., Podsiadlowski, L., Cameron, S.L., Daniels, S.R, Mayer, G. (2010). The mitochondrial genome of the Onychophoran Opisthopatus cinctipes (Peripatopsidae) reflects the ancestral mitochondrial gene arrangement of Panarthropoda and Ecdysozoa. Molecular Phylogenetics and Evolution, 57: 285 – 292.

34) Braband, A., Cameron, S. L., Daniels, S. R., Mayer, G, Podsiadlowski, L. (2010). The evolution of mitochondrial genome rearrangements in the taxon Onychophora (Arthropoda): Evidence for a complete mt-genome duplicaton with random loss of genes in Metaperipatus inae (Peripatopsis, Onychophora). Molecular Phylogenetics and Evolution, 57: 293-300.


33) Daniels, S. R., Picker, M., Cowlin, R. M., Hamer, M. (2009). Unravelling evolutionary lineages among the South African velvet worms (Onychophora: Peripatopsis) provides evidence for widespread cryptic speciation. Biological Journal of the Linnean Society, 97: 200-216.

32) Daniels, S. R., Heideman, N. J., Hendricks, M. G. J. (2009). Examination of evolutionary relationships in a Cape fossorial skink species complex (Acontiinae: Acontias meleagris meleagris) reveals the presence of five cryptic lineages. Zoologica Scripta, 38: 449-463.

31) Cumberlidge, N., Ng, P. K. L., Yeo, D. C .J., Magalhaes, C., Campos, M. R., Alvarez, F., Naruse, T., Daniels, S. R., Esser, L. J, Collen, B., Attipoe, F. Y. K., Clotilde-Ba, F.L., Darwall, W., McIvor, A., Ram, M (2009). Freshwater crabs and the biodiversity crisis: importance, threats, status, and conservation challenges. Biological Conservation, 142, 16655-1673.


30) Yeo, D. C. J., Ng, P. K. L., Cumberlidge, N., Magalhaes, C., Daniels, S. R., Campos, M. R. (2008). A global assessment of freshwater crab diversity (Crustacea: Decapoda: Brachyura). Hydrobiologia, 595: 275-286.

29) Cumberlidge, N., Von Sternberg, R., Daniels, S. R. (2008). A revision of the higher taxonomy of the Afrotropical freshwater crabs (Decapoda: Brachyura) with a discussion of their biogeography. Biological Journal of the Linnean Society, 93: 399-413.

28) Cumberlidge, N., Daniels, S.R. (2008). A conservation assessment of the freshwater crabs of southern Africa (Brachyura: Potamonautidae). African Journal of Ecology, 46: 74-79.

27) Mortimer, E., McGoech, M. A., Daniels, S. R., Jansen van Vuuren, B. (2008). Growth form and population genetic structure of Azorella selago on sub-Antarctic Marion Island. Antarctic Science, 20: 381-390.

26) Heideman, N. J. L., Daniels, S. R., Hendricks, M. G. J. (2008) Maximizing empirical data in delimiting species boundaries: a morphological based discriminant function analysis of the limbless skink subfamily Acontinae. African Zoology, 43: 192-201.


25) Myburgh, M., Chown, S. L., Daniels, S. R., Jansen van Vuuren, B. (2007). Population structure, propagule pressure and conservation biogeography in sub-Antarctic: lesions from indigenous and invasive springtails. Diversity and Distribution, 13: 143-154

24) Daniels, S. R., Hofmeyr, M. D., Henen, B. T., Crandall, K. A. (2007).  Living with the genetic signature of Miocene induced changes: evidence from the phylogeographic structure of the endemic angulate tortoise Chersina angulata. Molecular Phylogenetics and Evolution, 45: 915-926.


23) Daniels, S. R., Cumberlidge, N., Perez-Losada, M., Crandall, K. A. (2006). Evolution of Afrotropical freshwater crab lineages obscured by morphological convergence. Molecular Phylogenetics and Evolution, 40: 227-235.

22) Daniels, S. R., Gouws, G., Crandall, K. A. (2006). Phylogeographic patterning in a freshwater crab species (Decapoda: Potamonautidae: Potamonautes) reveals the signature of historic climatic oscillations. Journal of Biogeography, 33: 1538-1549.

21) Marijnissen, S. A., Daniels, S. R., Schram. F., Michelle, E. (2006). Rapid recent divergence of freshwater crabs (Platythelphusidae: Platythelphusa). Molecular Phylogenetics and Evolution, 40: 628-634.

 20) Daniels, S. R., Heideman, N., Hendricks, M. J. G., Crandall, K. A. (2006). Taxonomic subdivisions within the fossorial skink subfamily Acontinae (Sauria: Scincidae:) reconsidered: a multi-locus perspective. Zoolgica Scripta, 35: 353-362.

19) Gouws, G., Stewart, B. A., Daniels, S. R. (2006). Phylogeography of the koonac (Decapoda: Parastacidae: Cherax preissii) in Western Australia: multiple lineages but only one species. Marine and Freshwater Research, 57: 837-848.


18) Daniels, S. R., Heideman, N., Hendricks, M., Mokone, M. E., Crandall, K. A. (2005). Unraveling evolutionary lineages the fossorial skink group Acontias: are subspecies equivalent systematic units. Molecular Phylogenetics and Evolution, 34: 645-654.


17) Stewart, B. A., Gouws, G., Matthee, C. A., Daniels, S. R. (2004). Delimitations of cryptic sponge crab species on the genus Pseudodromia from South Africa. Zoological Scripta, 33: 45-55.

16) Gouws, G., Stewart, B. A., Daniels, S. R. (2004). Cryptic species within the freshwater isopod Mesamphisopus capensis in the Western Cape, South Africa: allozyme, morphometrics and 12S rRNA sequence data. Biological Journal of the Linnean Society, 81: 235-253.

15) Daniels S. R, Hamer M., Rogers D. C. (2004). Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca). Biological Journal of the Linnean Society, 82: 313-327.

14) Daniels, S. R., Mouton, P. le F. N., du Toit D. (2004). Molecular data suggest that melanistic ectotherms at the southwestern tip of Africa are the products of Miocene climatic events: evidence from cordylid lizards. Journal of Zoology, London, 263: 373-383.


13) Daniels, S. R, Gouws, G., Stewart, B. A., Coke, M. (2003). Molecular and morphometric evidence for the presence of cryptic lineages among freshwater crabs from the Drakensberg Mountains, South Africa. Biological Journal of the Linnean Society, 78: 129-147.

12) Daniels, S. R. (2003). Examining the genetic structure among populations of the common Cape River crab, Potamonautes perlatus from river systems in South Africa reveals hydrographic boundaries. Journal of Crustacean Biology, 24: 936-950.


11) Gouws, G., Daniels, S. R., Stewart, B. A. (2002). Allozyme electrophoresis demonstrates the presence of a species boundary in freshwater crabs (Decapoda: Potamonautidae). Journal of Natural History, 36: 1199 – 1222.

10) Daniels, S. R., Stewart, B. A., Cook, P. A. (2002). Congruent patterns of genetic variation in a burrowing freshwater crab revealed by allozyme and mt DNA sequence analysis. Hydrobiologia, 468: 171-179.

9) Daniels, S. R., Heideman, N., Hendricks, M. G. J., Wilson, B. (2002). A molecular phylogeny of the South African limb less lizard taxa of the subfamily Acontinae with special emphasis on the relationships within the genus Acontias. Molecular Phylogenetics and Evolution, 24: 314 -322.

8) Daniels, S. R., Stewart, B. A., Gouws, G., Cunningham, M., Matthee, C. A. (2002). Phylogenetic relationships of the southern African freshwater crab fauna derived from multiple data sets reveal biogeographic patterning. Molecular Phylogenetics and Evolution, 25: 511-523.


7) Daniels, S. R., Stewart, B.A, Ridgeway, T., Florance, W. (2001). Carapace dentition patterns, morphometrics and genetic differentiation amongst two toothed river crab species
(Potamonautes warreni and P. unispinus) from river systems in South Africa. Journal of Zoology, London, 255: 389-404.

6) Daniels, S. R. (2001). Allometric growth, handedness and morphological variation, with a redescription of Potamonautes warreni (Calman, 1918). Crustaceana, 74: 237-253.

5) Daniels, S. R., Stewart, B. A., Burmeister, L. (2001) Geographic patterns of genetic and morphological divergence amongst populations of a freshwater crab with the description of a species from mountain streams in the Western Cape. Zoologica Scripta, 30: 181-197.


4) Daniels, S. R., Stewart, B. A., Gibbons, M. J. (1999). Genetic differentiation amongst populations of Potamonautes perlatus in the Olifants River system. Journal of Zoology, London, 249: 137-142.

3) Daniels, S. R., Stewart, B. A., Gibbons, M. J. (1999). Allozyme variation amongst populations of Potamonautes perlatus from the Berg River system. South African Journal of Zoology, 34: 64-68.


2) Daniels, S. R., Stewart, B. A., Gibbons, M. J. (1998). Genetic and morphometric variation amongst populations of Potamonautes parvispina from two river systems in the Western Cape. Journal of Natural History, 32: 1245-1258.

1) Daniels, S. R., Stewart, B. A., Gibbons, M. J. (1998). Potamonautes granularis sp. nov, a new cryptic species of river crab from the Olifants River System, South Africa. Crustaceana, 71: 885-903.

Popular Articles:

Daniels, S. R and Stewart, B. A. (2000). Cagey crustaceans: the freshwater crabs of Southern Africa. African Wildlife 54 (3): 31-32.

Daniels, S. R and Stewart, B. A. (2000). Hermit crabs at sea shell city. African Wildlife 54 (4): 24-25.