Dr David Reid

David Reid collecting littorinids in mangroves in Costa Rica
  • Merit Researcher
  • Life Sciences department
  • Invertebrates
Natural History Museum
Cromwell Road


Employment history

1985 Post-doctoral Fellow, National Museum of Natural History, Smithsonian Institution,
          Washington, D.C.
1986 Senior Research Fellow, Dept of Zoology, NHM
1989 Principal Scientific Officer, Mollusca Research Group, Dept of Zoology, NHM
1998 Individual Merit Promotion (band 2)


1979 BA Hons, University of Cambridge, UK
1984 PhD, James Cook University of North Queensland, Australia

Professional Roles


Open scholarship to St John’s College, Cambridge (1976-1979)
Drapers’ Company scholarship for postgraduate research overseas (1980-1983)
Keith Sutherland Award for malacological research from the Australian Museum (1981)
Annual Award for an Initial Contribution to Malacology, Malacological Society of London (1985)
Scientific Medal of the Zoological Society of London (1996)
Bicentenary Medal of the Linnean Society of London (1997)


1991–94  £119,000 from NERC: Mitochondrial DNA evolution in the gastropod genus Littorina
                  (with R.H. Thomas)
1994–97  £120,000 from NERC: Origins and speciation of the northern Atlantic marine intertidal
                  fauna: a phylogeny of patellid limpets (with J.D. Taylor, R.H. Thomas, S. Hawkins)
2001–04  £174,000 from NERC: The enigma of the marine biodiversity focus in the Indo-West
                  Pacific Ocean: phylogeography of Nodilittorina (with D.T.L. Littlewood)

Editorial boards


Journal of Molluscan Studies



Current research interests involve the reconstruction of gastropod phylogenies using both morphological and molecular approaches, and the use of such hypotheses in the study of biogeography and speciation in marine invertebrates, and in the interpretation of the evolution of adaptive morphological characters. For much of this evolutionary work, the gastropod family Littorinidae has been used as a model, having a worldwide distribution, at least a partial fossil record, and showing specialized adaptations to intertidal life. A new project investigates the evolutionary history of ecological associations between gastropods (Potamididae and Littoraria) with the mangrove habitat, integrating the fossil record with molecular phylogenetics. Supervised PhD projects have included worldwide phylogenetic analyses of Patellidae, Bullidae, gibbuline Trochidae and Muricidae. Previous research interests include the ecology of molluscs on coral reefs and the interactions between colour-polymorphic snails and their predators.General taxonomic work has been done on the marine molluscs of Chile.

Read more about my projects on their dedicated research pages.


Project: Diversification and historical biogeography of cerithioidean gastropods in mangrove environments

Department of Zoology: D.G. Reid, S.T. Williams
External collaborators: M. Glaubrecht & F. Kohler (Humboldt University, Berlin), P. Lozouet (MNHN, Paris), T. Ozawa (Cyber University, Nagoya).

Almost all of the living members of the family Potamididae have an obligate association with mangrove or subtropical salt-marsh vegetation. Using molecular phylogenetics, museum records of distribution and the fossil record, we are examining global patterns of diversification in this family, with a focus on Tethyan biogeography and the history of the mangrove environment. The family first appeared in the fossil record of the Middle Eocene, shortly after the origin of modern mangrove trees. The Indo-Pacific genus Cerithidea contains specialized tree-climbing species, and its biogeography and pattern of speciation can be compared with that of the littorinid genus Littoraria with a similar habit. The systematics of the poorly known genera Cerithidea and Cerithideopsilla are being revised.

Another cerithideoidean group found on sedimentary shores is the family Batillariidae. A molecular phylogeny of all living species together with a revision of the fossil record indicate that this is a relictual family, now restricted to the northwestern Pacific and Australasia. Two tropical American species, formerly classified in this family, do not belong in the same clade.

Cerithidea decollata can reach high densities on trunks of Avicennia mangroves in Mozambique.

Cerithidea decollata can reach high densities on trunks of Avicennia mangroves in Mozambique. © David Reid



Project: Systematics and evolution of Littorinidae (Gastropoda)

Department of Zoology: D.G. Reid, S.W. Williams
External collaborators: R. Butlin, J. Hollander, C. Smadja (University of Sheffield)

Littorinid snails are abundant on intertidal hard surfaces on almost all shores of the world, and all aspects of their biology have been intensively studied. Detailed systematic research and molecular phylogenetic analyses of many of the larger genera (e.g. Littorina, Echinolittorina, Littoraria) have established this group as a model for diversification of invertebrates in global shallow marine environments. A complete phylogeny of all 160 species of the subfamily Littorininae is now within reach, permitting analysis of patterns of life history, habitat, range size, speciation rate etc.

Reinforcement (selection against viable hybrids) has been suggested as an important mechanism to drive reproductive isolation between diverging populations, as a final step in speciation. However, the process is controversial and field evidence is hard to find. Using molecular phylogenies of littorinid genera, sister-species pairs can be confidently identified, and character displacement in genital traits (penis, sperm, oviduct) compared in sympatric and allopatric pairs. If reinforcement operates, greater divergence is predicted in sympatric pairs.

Although the molecular phylogeny of the Littorinidae is now well resolved, it is a challenge to relate this to the meagre fossil record. Examination of available littorinid fossils aims to identify the earliest occurrences of the major clades in order to make inferences about their origin and spread, and to refine the dating of phylogenetic trees. Systematic studies of living genera are continuing with the completion of revisions of worldwide Echinolittorina species.

Littorina sitkana on a rocky shore at Seldovia, Alaska.

Littorina sitkana on a rocky shore at Seldovia, Alaska. © David Reid



Project: Systematics and evolution of Rapaninae and Ergalataxinae (Gastropoda: Muricidae)

Department of Zoology: M. Claremont (PhD student), S.T. Williams, D.G. Reid
External collaborator: T. Barraclough (Imperial College, London)

The Muricidae are one of the major families of shallow-water carnivorous gastropods. There are competing morphological phylogenies based on either shell or combined shell and anatomical data. Our objectives are to use molecular markers to produce phylogenies for the subfamilies Rapaninae and Ergalataxinae, to generate a new phylogenetic classification, to examine biogeographic patters in the light of both modern distributions and the fossil record, and to investigate the role of dietary specialization in the adaptive radiation of selected genera.

Thais bitubercularis feeding on oysters, with egg capsules (Langkawi, Malaysia).

Thais bitubercularis feeding on oysters, with egg capsules (Langkawi, Malaysia). © David Reid


Research Student Supervision

S. Ridgway (Queen Mary College, University of London; 1993)
K. Caley (University of Leeds; 1994)
J. Preston (University of Southampton; 2003)
M.A.E. Malaquias (Queen Mary College, University of London; 2007)
M. Claremont (Imperial College, London)



1. Taylor, J.D. & Reid, D.G. (1983) The abundance and trophic classification of molluscs upon coral reefs in the Sudanese Red Sea. Journal of Natural History 18: 175-209.

2. Reid, D.G. (1985) Habitat and zonation patterns of Littoraria species (Gastropoda: Littorinidae) in Indo-Pacific mangrove forests. Biological Journal of the Linnean Society 26: 39-68.

3. Reid, D.G. (1986a) The Littorinid Molluscs of Mangrove Forests in the Indo-Pacific Region: the genus Littoraria. British Museum (Natural History), London. pp. xv, 228.

4. Reid, D.G. (1986b) Mainwaringia Nevill, 1885, a littorinid genus from Asiatic mangrove forests, and a case of protandrous hermaphroditism. Journal of Molluscan Studies 52: 225-242.

5. Reid, D.G. (1987) Natural selection for apostasy and crypsis acting on the shell colour polymorphism of a mangrove snail, Littoraria filosa (Sowerby) (Gastropoda: Littorinidae). Biological Journal of the Linnean Society 30: 1-24.

6. Reid, D.G. (1988) The genera Bembicium and Risellopsis (Gastropoda: Littorinidae) in Australia and New Zealand. Records of the Australian Museum 40: 91-150.

7. Reid, D.G. (1989a) The comparative morphology, phylogeny and evolution of the gastropod family Littorinidae. Philosophical Transactions of the Royal Society of London, Series B 324: 1-110.

8. Reid, D.G. (1989b) Systematic revision of the Recent species of Peasiella (Gastropoda, Littorinidae), with notes on the fossil species. Nautilus 103: 43-69.

9. Reid, D.G. (1990a) Note on the discrimination of females of Littorina mariae Sacchi & Rastelli and L. obtusata (Linnaeus). Journal of Molluscan Studies 56: 113-114.

10. Reid, D.G. (1990b) A cladistic phylogeny of the genus Littorina (Gastropoda): implications for evolution of reproductive strategies and for classification. Hydrobiologia 193: 1-19.

11. Reid, D.G. (1990c) Trans-Arctic migration and speciation induced by climatic change: the biogeography of Littorina (Mollusca: Gastropoda). Bulletin of Marine Science 47: 35-49.

12. Taylor, J.D. & Reid, D.G. (1990) Shell microstructure and mineralogy of the Littorinidae: ecological and evolutionary significance. Hydrobiologia 193: 199-215.

13. Reid, D.G., Zaslavskaya, N.I. & Sergievsky, S.O. (1991) Littorina kasatka, a new species from the Kurile Islands and the Okhotsk Sea. Nautilus 105: 1-6.

14. Reid, D.G. & Golikov, A.N. (1991) Littorina naticoides, new species, with notes on the other smooth-shelled Littorina species from the Northwestern Pacific. Nautilus 105: 7-15.

15. Reid, D.G. (1992a) The evolution of spawn and larval development in the Littorinidae (Gastropoda, Prosobranchia). Proceedings of the Tenth International Malacological Congress (Tübingen, 1989): 515-516.

16. Reid, D.G. (1992b) Predation by crabs on Littoraria species (Littorinidae) in a Queensland mangrove forest. In: Proceedings of the Third International Symposium on Littorinid Biology (J. Grahame, P.J. Mill & D.G. Reid, Eds): 141-151. Malacological Society of London.

17. Reid, D.G. (1992c) The gastropod family Littorinidae in Hong Kong. In: The Marine Flora and Fauna of Hong Kong and Southern China III (B. Morton, Ed.): 187-210. Hong Kong University Press, Hong Kong.

18. Grahame, J., Mill, P.J. & Reid, D.G., Eds (1992) Proceedings of the Third International Symposium on Littorinid Biology. Malacological Society of London. pp. vi, 324.

19. Warmoes, T., Dumoulin, E. & Reid, D.G. (1992) Distribution of Littorina and Melarhaphe along the continental coasts of the English Channel. In: Proceedings of the Third International Symposium on Littorinid Biology (J. Grahame, P.J. Mill & D.G. Reid, Eds): 293-303. Malacological Society of London.

20. Reid, D.G. (1993) Barnacle-dwelling ecotypes of three British Littorina species and the status of Littorina neglecta Bean. Journal of Molluscan Studies 59: 51-62.

21. Rumbak, E., Reid, D.G. & Thomas, R.H. (1994) Reconstruction of phylogeny of 11 species of Littorina (Gastropoda: Littorinidae) using mitochondrial DNA sequence data. Nautilus, Supplement 2: 91-97.

22. Reid, D.G. (1994) Species and speciation in Littorinidae (abstract). Cahiers de Biologie Marine 35: 258-259.

23. Reid, D.G. (1996) Systematics and Evolution of Littorina. London: The Ray Society. pp x, 463.

24. Reid, D.G., Rumbak, E. & Thomas, R. H. (1996) DNA, morphology and fossils: phylogeny and evolutionary rates of the gastropod genus Littorina. Philosophical Transactions of the Royal Society, Series B 351: 877-895.

25. Reid, D.G., Sattler, K. & Crosskey, R.W. (1996) Discussion draft of the Fourth Edition of the International Code of Zoological Nomenclature: comments (1). Bulletin of Zoological Nomenclature 53: 80-81.

26. Reid, D.G. & Geller, J.B. (1997) A new ovoviviparous species of Tectarius (Gastropoda: Littorinidae) from Niue, South Pacific, with a molecular phylogeny of the genus. Journal of Molluscan Studies 63: 207-233.

27. Reid, D.G. (1998) Superfamily Littorinoidea. Family Littorinidae. In: Mollusca: The Southern Synthesis. Fauna of Australia, Volume 5. (Beesley, P.L., Ross, G.J.B. & Wells, A., Eds): 737-739. CSIRO Publishing, Melbourne.

28. Reid, D.G. & Mak, Y-M. (1998) Additions and corrections to the taxonomy of the genus Peasiella Nevill, 1885 (Gastropoda: Littorinidae). Nautilus 112: 6-33.

29. Ridgway, S.A., Reid, D.G., Taylor, J.D., Branch, G.M. & Hodgson, A.N. (1998) A cladistic phylogeny of the family Patellidae (Mollusca: Gastropoda). Philosophical Transactions of the Royal Society, Series B 353: 1645-1671.

30. Winnepenninckx, B.M.H., Reid, D.G. & Backeljau, T. (1998) Performance of 18S rRNA in littorinid phylogeny (Gastropoda: Caenogastropoda). Journal of Molecular Evolution 47: 586-596.

31. Koufopanou, V., Reid, D.G., Ridgway, S.A. & Thomas, R.H. (1999) A molecular phylogeny of the patellid limpets (Gastropoda: Patellidae) and its implications for the origins of their antitropical distribution. Molecular Phylogenetics and Evolution 11: 138-156.

32. Reid, D.G. (1999) The genus Littoraria Griffith & Pidgeon, 1834 (Gastropoda: Littorinidae) in the tropical eastern Pacific. Veliger 42: 21-53.

33. Reid, D.G. & Mak, Y-M. (1999) Indirect evidence for ecophenotypic plasticity in radular dentition of Littoraria species (Gastropoda: Littorinidae). Journal of Molluscan Studies 65: 355-370.

34. Reid, D.G. (1999) The phylogeny of Littoraria (Gastropoda: Littorinidae): an example of the practice and application of cladistic analysis. Phuket Marine Biological Center Special Publication 19: 283-322.

35. Reid, D.G. & Osorio, C. (2000) The shallow-water marine Mollusca of the Estero Elefantes and Laguna San Rafael, southern Chile. Bulletin of the Natural History Museum. Zoology Series 66: 109-146.

36. Reid, D.G. (2000) The uses of the radula in the taxonomy and phylogeny of gastropods: cautionary cases of convergence, intraspecific variation and plasticity. Phuket Marine Biological Center Special Publication 21: 329–345

37. Reid, D.G. (2000) Preservation and curation of marine molluscan specimens. Phuket Marine Biological Center Special Publication 21: 583–590.

38. Reid, D.G. & Kaiser, K.L. (2001) Three new records of Indo-West Pacific Littorinidae (Gastropoda) in the tropical eastern Pacific. Veliger 44: 381–388.

39. Reid, D.G. (2001) New data on the taxonomy and distribution of the genus Littoraria Griffith and Pidgeon, 1834 (Gastropoda: Littorinidae) in Indo-West Pacific mangrove forests. Nautilus 115: 115–139.

40. Reid, D.G. (2001) The genus Nodilittorina von Martens, 1897 (Gastropoda: Littorinidae) in the Indo-Malayan region. Phuket Marine Biological Center Special Publication 25: 433–449.

41. Reid, D.G. (2002) The genus Nodilittorina von Martens, 1897 (Gastropoda: Littorinidae) in the eastern Pacific Ocean, with a discussion of biogeographic provinces of the rocky-shore fauna. Veliger 45: 85–169.

42. Reid, D.G. (2002) Morphological review and phylogenetic analysis of Nodilittorina (Gastropoda: Littorinidae). Journal of Molluscan Studies 68: 259–281.

43. Stuckey, M. & Reid, D.G. (2002) A new Littoraria (Gastropoda: Littorinidae) from northwestern Australia. Molluscan Research 22: 1–15.

44. Williams, S.T., Reid, D.G. & Littlewood, D.T.J. (2003) A molecular phylogeny of the Littorininae (Gastropoda: Littorinidae): unequal evolutionary rates, morphological parallelism and biogeography of the Southern Ocean. Molecular Phylogenetics and Evolution 28: 60–86.

45. Reid, D.G. & Williams, S.T. (2004) The subfamily Littorininae (Gastropoda: Littorinidae) in the temperate Southern Hemisphere: the genera Nodilittorina, Austrolittorina and Afrolittorina. Records of the Australian Museum 56: 75–122.

46. Williams, S.T. & Reid, D.G. (2004) Speciation and diversity on tropical rocky shores: a global phylogeny of snails of the genus Echinolittorina. Evolution 58: 2227–2251.

47. Osorio, C. & Reid, D.G. (2004) Moluscos marinos intermareales y submareales entre la Boca del Guafo y el estero Elefantes, sur de Chile. Investigaciones Marinas 32: 71–89.

48. Osorio, C., Reid, D.G. & Ramajo, L. (2005) Moluscos en los canales del sur de Chile entre Boca del Guafo y Estero Elefantes (CIMAR 7 Fiordos). Ciencia y Tecnología del Mar 28: 91–98.

49. Reid, D.G., Lal, K., Mackenzie-Dodds, J., Kaligis, F., Littlewood, D.T.J. & Williams, S.T. (2006) Comparative phylogeography and species boundaries in Echinolittorina snails in the central Indo-West Pacific. Journal of Biogeography 33: 990–1006.

50. Reid, D.G. (2007) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the Indo-West Pacific Ocean. Zootaxa 1420: 1–161.

51. Faubel, A., Sluys, R. & Reid, D.G. (2007) A new genus and species of polyclad flatworm found in the mantle cavities of gastropod molluscs in the high-intertidal zone of the Pacific coast of Central America. Journal of the Marine Biological Association of the United Kingdom 87: 429–434.

52. Reid, D.G. (2007) Littorina. In: The Light and Smith Manual. Intertidal Invertebrates from Central California to Oregon. Fourth Edition (Carlton, J.T., ed.), 761–766. University of California Press, Berkeley.

53. Reid, D.G., Dyal, P., Lozouet, P., Glaubrecht, M. & Williams, S.T. (2008) Mudwhelks and mangroves: the evolutionary history of an ecological association (Gastropoda: Potamididae). Molecular Phylogenetics and Evolution 47: 680–699.

54. Malaquias, M.A.E. & Reid, D.G. (2008a) Functional morphology of the gonoduct of the gastropod Bulla striata Bruguière, 1792 (Opisthobranchia: Cephalaspidea): evidence for a monaulic system. Acta Zoologica 89: 205–210.

55. Malaquias, M.A.E. & Reid, D.G. (2008b) Systematic revision of the living species of Bullidae (Mollusca: Gastropoda: Cephalaspidea), with a molecular phylogenetic analysis. Zoological Journal of the Linnean Society 153: 453–543.

56. Claremont, M., Reid, D.G. & Williams, S.T. (2008) A molecular phylogeny of the Rapaninae and Ergalataxinae (Neogastropoda: Muricidae). Journal of Molluscan Studies 74: 215–221.

57. Vermeij, G.J., Dietl, G.P. & Reid, D.G. (2008) The trans-Atlantic history of diversity and body size in ecological guilds. Ecology, supplement 89: S39–S52.

58. Torres, P., Alfiado, A., Glassom, D., Jiddawi, N., Macia, A., Reid, D.G. & Paula, J. (2008) Species composition, comparative size and abundance of the genus Littoraria (Gastropoda: Littorinidae) from different mangrove strata along the East African coast. Hydrobiologia 614: 339–351.

59. Malaquias, M.A.E., Mackenzie-Dodds, J., Bouchet, P., Gosliner, T. & Reid, D.G. (2009) A molecular phylogeny of the Cephalaspidea sensu lato (Gastropoda: Euthyneura): Architectibranchia redefined and Runcinacea reinstated. Zoologica Scripta 38: 23–41.

60. Malaquias, M.A.E., Berecibar, E. & Reid, D.G. (2009) Reassessment of the trophic position of Bullidae (Gastropoda: Cephalaspidea) and the importance of diet in the evolution of cephalaspidean gastropods. Journal of Zoology 277: 88–97.

61. Ozawa, T., Köhler, F., Reid, D.G. & Glaubrecht, M. (2009). Tethyan relicts on continental coastlines of the northwestern Pacific Ocean and Australasia: molecular phylogeny and fossil record of batillariid gastropods (Caenogastropoda, Cerithioidea). Zoologica Scripta, 38: 503–525.

62. Malaquias, M.A.E. & Reid, D.G. (2009) Tethyan vicariance, relictualism and speciation: evidence from a global molecular phylogeny of the opisthobranch genus Bulla. Journal of Biogeography, 36: 1760–1777.

63. Reid, D.G. (2009) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the western Atlantic Ocean. Zootaxa, 2184: 1–103.

64. Reid, D.G., Dyal, P. & Williams, S.T. (2010) Global diversification of mangrove fauna: a molecular phylogeny of Littoraria (Gastropoda: Littorinidae). Molecular Phylogenetics and Evolution, 55: 185-201.

65. Barco, A., Claremont, M, Reid, D.G., Houart, R., Bouchet, P., Williams, S.T., Cruaud, C., Couloux, A., Oliverio, M. (2010) A molecular phylogenetic framework for the Muricidae, a diverse family of carnivorous gastropods. Molecular Phylogenetics and Evolution, 56: 1025–1039.

66. Osorio, C. & Reid, D.G. (2010) Bivalvia (Mollusca). In: Bibliografía sobre Biodiversidad Actuática de Chile (Palma, S., Báez, P. & Pequeño, eds): 101-110. Comité Oceanográfico Nacional, Valparaíso.

67. Claremont, M., Williams, S.T., Barraclough, T.G., & Reid, D.G. (2011) The geographic scale of speciation in a marine snail with high dispersal potential. Journal of Biogeography, 38: 1016–1032.

68. Ha, W.Y., Reid, D.G., Kam, W.L., Lau, Y.Y., Sham, W.C., Tam, S.Y.K., Sin, D.W.M., Mok, C.S. (2011) Genetic differentiation between fake abalone and genuine Haliotis species using the forensically informative nucleotide sequencing (FINS) method. Journal of Agricultural and Food Chemistry, 59: 5195–5203.

69. Claremont, M., Reid, D.G. & Williams, S.T. (2011) Evolution of corallivory in the gastropod genus Drupella. Coral Reefs, 30: 977–990.

70. Reid, D.G. (2011) The genus Echinolittorina Habe, 1956 (Gastropoda: Littorinidae) in the eastern Atlantic Ocean. Zootaxa, 2974: 1–65.

71. Donald, K.M., Preston, J., Williams, S.T., Reid, D.G., Winter, D., Alvarez, R., Buge, B., Hawkins, S.J., Templado, J., Spencer, H. (2012) Phylogenetic relationships elucidate colonization patterns in the intertidal grazers Osilinus Philippi, 1847 and Phorcus Risso, 1826 (Gastropoda: Trochidae) in the northeastern Atlantic Ocean and Mediterranean Sea. Molecular Phylogenetics and Evolution, 62: 35–45.

72. Reid, D.G., Dyal, P. & Williams, S.T. (2012) A global molecular phylogeny of 147 periwinkle species (Gastropoda, Littorininae). Zoologica Scripta, 41: 125–136

73. Claremont, M., Reid, D.G. & Williams, S.T. (2012) Speciation and dietary specialization in Drupa, a genus of predatory marine snails (Gastropoda: Muricidae). Zoologica Scripta, 41: 137–149.

74. Claremont, M., Vermeij, G.J., Williams, S.T. & Reid, D.G. (2013) Global phylogeny and new classification of the Rapaninae (Gastropoda: Muricidae), dominant molluscan predators on tropical rocky seashores. Molecular Phylogenetics and Evolution, 66: 91–102.

75. Claremont, M., Houart, R., Williams, S.T. & Reid, D.G. (2013) A molecular phylogenetic framework for the Ergalataxinae (Neogastropoda: Muricidae). Journal of Molluscan Studies, 79: 19–29.

76. Hollander, J., Smadja, C.M., Butlin, R.K. & Reid, D.G. (2013) Genital divergence in sympatric sister snails. Journal of Evolutionary Biology, 26: 210–215.

77. Reid, D.G., Aravind, N.A. & Madhyastha, N.A. (2013) A unique radiation of marine littorinid snails in the freshwater streams of the Western Ghats of India: the genus Cremnoconchus W.T. Blanford, 1869 (Gastropoda: Littorinidae). Zoological Journal of the Linnean Society, 167: 93–135.

78. Reid, D.G., Claremont, M., Smith, L., Shamoto, M., Glaubrecht, M.& Ozawa, T. (2013) Mosaics in the mangroves: allopatric diversification of tree-climbing mudwhelks (Gastropoda: Potamididae: Cerithidea) in the Indo-West Pacific. Biological Journal of the Linnean Society, 110: 564–580.