Howard Choat, Recipient of the First Bleeker Award in Ecology
By Kendall Clements
We are honouring Professor Howard Choat as one of the recipients of the 1stBleeker Award for his contributions to Indo-Pacific ichthyology. In fact his scientificcontributions extend beyond the field of fish biology to include kelp dynamics,
cephalopod demography, and marine ecology in general. It is probably fair to say that
Howard is the finest Australasian marine ecologist, in the sense that he has made anenormous contribution to marine ecology on both sides of the Tasman Sea. This broadcontribution has been acknowledged by a number of previous awards, including anAustralian Research Council Special Investigator Award in 1993 and the Jubilee Awardfor Outstanding Marine Scientist by the Australian Marine Sciences Society in 1998. It is,however, extremely fitting that Howard should now be honoured for his major researchcontribution, which has been his work on the biology of reef fishes.
Howard’s scientific career started in the 1960s in New Zealand with a BSc andMSc at Victoria University of Wellington, and he went on to complete his PhD at theUniversity of Queensland in 1969. After postdoctoral experience in California, Howard
took up a position at the University of Auckland, where he was extremely active indeveloping subtidal research at the Leigh Marine Laboratory. In 1986 Howard took up
the post of head of the Department of Marine Biology at James Cook University, andretired from administrative duties in 2001 to take up the research position of adjunctprofessor.
Howard has operated largely by developing broadly-based research programs inassociation with graduate students. Howard’s particular contributions to theichthyological literature have been in the fields of herbivory and coral reef fish ecology,
with a recent focus on age-based dynamics. However, his contribution to ichthyology
cannot be measured simply by his impressive publication output. He has been
instrumental in the development of reef fish research both as an inspiring supervisor ofmany graduate students from different parts of the world and through his service on
numerous science management committees. Howard’s unselfish and diligent efforts overmany years towards developing reef fish biology and biologists make him an extremely
worthy recipient of the 1st Bleeker Award.

Publications of Howard Choat
1. Choat JH (1965)

Sexual dimorphism in the labrid fish Pseudolabrus celidotus.

Pacific Science 19: 451-457.
2. Choat JH (1968) The status of Pseudolabrus psittaculus with notes on other speciesof the genus Transactions of the Royal Society of New Zealand (Zoology)
10:151-157.
3. Robertson DR, Choat JH (1974) Protogynous hermaphroditism and social systems
in labrid fish Proceedings of the Second International Coral Reef
Symposium Vol 1: 217-228
4. Choat JH, Robertson DR (1975). Protogynous hermaphroditism in fishes of thefamily Scaridae. In Intersexuality in the Animal Kingdom (Reinboth R, ed.),pp.
263–283. Berlin: Springer-Verlag.
5. Choat J H (1977) Influence of sessile organisms on population biology of threespecies of acmaeid limpets. Journal of Experimental Marine Biology andEcology 26 1-26 : 25-50
6. Choat J H, Black R (1979) Life histories of limpets and the limpet-laminarianrelationship. Journal of Experimental Marine Biology and Ecology 41 :25-50
7. Schiel DR, Choat JH (1980) Effects of density on monospecific stands ofmarine-algae.

Nature 285 (5763): 324-326

8. Choat JH, Schiel DR (1980) Population-structure of Placostylus hongii on the PoorKnights islands. New Zealand Journal of Zoology 7 (2): 199-205 1980
9. Leum LL, Choat JH (1980) Density and distribution patterns
marine fishof the temperate

Cheilodactylus spectabilis in a reef environment. Marine

Biology 57: 327-337
10. Randall JE, Choat JH (1980) Two new parrotfishes of the genus Scarus from thecentral and south Pacific with further examples of sexual dichromatism
Zoological Journal of the Linnean Society 70: 383-419
11. Kingett PD, Choat JH (1981) Analysis of the density and distribution patterns ofChrysophrys auratus within a reef environment: an experimental approach.
Marine Ecology Progress Series 5: 283-294.
12. Andrew NL, Choat JH (1982) The influence of predation and conspecific adults on
the abundance of juvenile Evechinus chloroticus. Oecologia 54: 80-87
13. Choat JH (1982)

Fish feeding and the structure of benthic communities intemperate waters. Annual Review of Ecology and Systematics 13: 423-449

14. Choat JH, Kingett PD (1982) The influence of fish predation on the abundancecyclesof an algal turf invertebrate fauna. Oecologia 54: 88-95

15. Choat JH, Schiel DR (1982) Patterns of distribution and abundance of large brownalgae and invertebrate herbivores in the subtidal of northern New Zealand.
Journal of Experimental Marine Biology and Ecology 60:129-162
16. Choat JH, Bellwood DR (1985) Interactions amongst herbivorous fishes on a coral
reef: influence of spatial variation. Marine Biology 89: 221-234
17. Kingsford MJ, Choat JH (1985) The fauna associated with drift algae captured with aplankton-mesh purse seine net. Limnology and Oceanography 30: 618-630
18. Andrew NL Choat JH (1985) Habitat related differences in the survivorship andgrowth of juvenile sea urchins. Marine Ecology Progress Series 27: 155-161
19. Choat JH, Andrew NL (1986) Interactions amongst species in a guild of subtidal
benthic herbivores. Oecologia 68: 387-394
20. Kingsford MJ, Choat JH (1986) Influence of surface slicks on the distribution andmovements of small fish. Marine Biology 91: 161-171
21. Schiel DR, Kingsford MJ, Choat JH (1986) Depth distribution and abundance ofbenthic organisms and fishes in the subtropical Kermadec Islands. New
Zealand Journal of Marine and Fresh Water Research. 20: 521-535
22. Choat JH, Randall JE (1986)

A review of the parrotfishes (Family Scaridae) of theGreat Barrier Reef Australia with a description of a new species Records of
the Australian Museum 38: 175-228
23. McCormick MI, Choat JH (1987) Estimating total abundance of a large temperate
reef fish. Marine Biology. 96: 469-478
24. Choat JH, Ayling AM (1987).

The influence of habitat structure on reef fish

communities in north-eastern New Zealand.

Journal of Experimental

Marine Biology and Ecology. 110: 257-272.
25. Choat JH, Ayling AM, Schiel DR (1988).

Temporal and spatial variation in an island

fish fauna. Journal of Experimental Marine Biology and Ecology. 121:91-112.
26. Kingsford MJ, Choat JH (1989) Horizontal distribution patterns of presettlement reef
fish: are they influenced by the proximity of reefs? Marine Biology 101:285-297
27. Bellwood DR, Choat JH (1989) A description of the juvenile colour phase patterns of
24 parrotfish species (family Scaridae) from the Great Barrier Reef Australia.
Records of the Australian Museum 41: 1-42

28. Choat JH, Barnes DJ, Borowitzka MA, Coll JC , Davies PJ, Flood P, Hatcher BG,Hopley, D

Hutchings PA, Kinsey D, Orme GR, Pichon M, Sale PF,Sammarco PW, Wallace CC, Wilkinson CR, Wolanski E, Bellwood O (eds),(1989).

Proceedings of Sixth International Coral Reef Symposium,

Townsville, Australia., vols 1-3: 1733 pp
29. Bellwood DR, Choat JH (1990) A functional analysis of grazing in parrotfishes family
Scaridae the ecological implications. Environmental Biology of Fishes
28:189-214
30. Kingsford MJ, Wolanski E, Choat JH (1991) Influence of tidally induced fronts andlangmuir circulations on distribution and movements of presettlement fishes
around a coral reef Marine Biology 109: 167-180
31. Choat, JH (1991). The biology of herbivorous fishes on coral reefs. In: Sale, P.F.
(ed) The Ecology of Coral Reef Fishes. Academic Press, 120-155.
32. Choat, JH, Bellwood, DR (1991). Reef fishes: their history and evolution. In: Sale,P.F.(ed) The Ecology of Coral Reef Fishes. Academic Press, 39-60.
33. Choat JH, Clements KD. (1992)

Diet in odacid and aplodactylid fishes fromAustralia and New Zealand. Australian Journal of Marine and Freshwater
Research 43:1451-1459.
34. Jackson GD, Choat JH (1992) Growth in tropical cephalopods: an analysis based onstatolith microstructure. Canadian Journal of Fisheries and Aquatic Sciences
49: 218-228.
35. Lewis AR, Choat JH (1993) Spawning mode and reproductive output of the tropicalcephalopod Idiosepius pygmaeus. Canadian Journal of Fisheries andAquatic Sciences. 50: 20-28.
36. Choat, JH, Doherty PJ, Kerrigan BA, Leis JM (1993). A comparison of towed nets,
purse seine and light aggregation devices for sampling larval and pelagicjuveniles of coral reef fishes. Fishery Bulletin 91:195-209.
37. Choat JH, Clements KD (1993). Daily feeding rates in herbivorous labroid fishes.
Marine Biology 117, 205-211.
38. Clements KD, Choat JH (1993). Influence of season, ontogeny and tide on the diet of
the temperate marine herbivorous fish Odax pullus (Odacidae). Marine Biology
117(2): 213-220.
39. Clements KD, Choat JH (1995). Fermentation in tropical marine herbivorous fishes.
Physiological Zoology 68(3): 355-378.
40. Choat JH, Axe LM (1996). Growth and longevity in acanthurid fishes; an analysis ofotolith increments. Marine Ecology Progress Series 134: 15-26.

41. Choat JH, Axe LM, Lou DC (1996) Growth and longevity in fishes of the family
Scaridae. Marine Ecology Progress Series 145: 33-41.
42. Clements KD, Choat JH (1997). Mechanisms of herbivory in marine fish: acomparison of the closely-related genera Girella and Kyphosus. MarineBiology 127: 579-586.
43. Meekan MG, Choat JH (1997) Latitudinal variation in abundance of herbivorous fishes:a comparison of temperate and tropical reefs. Marine Biology 128: 373-384.
44. Craig PC, Choat JH, Axe LM, Saucerman S (1997). Population biology and harvestof the coral reef surgeonfish, Acanthurus lineatus, in American Samoa.
Fishery Bulletin 95: 680-693.
45. Choat, JH (1998). The question of spatial scale in reef fish studies. In: Jones GP,Doherty PJ, Mapstone, BD, Howlett L. (Eds.) ReeFish ’95: Recruitment andPopulation Dynamics of Coral Reef Fishes. CRC Reef Research Centre,Townsville, Australia. pp 83-86.
46. Choat JH, Bellwood DR (1998). Wrasses and Parrotfishes. In: Paxton JR ,Eschmeyer WN (eds) Encyclopedia of Animals: Fishes 2nd Edition. WeldonOwen. pp 209-213.
47. Choat JH, Clements KD (1998). Vertebrate herbivores in marine and terrestrial
environments: a nutritional ecology perspective. Annual Review of Ecologyand Systematics 29, 375-403.
48. Jackson GD, Alford RA, Choat, JH (2000). Can squid populations be analysed using
standard fishery length-based techniques? ICES Journal of Marine Science
57: 948-954
49. Gust N, Choat JH, McCormick MI (2001). Spatial variability in reef fish distribution,abundance, size and biomass: a multi-scale analysis. Marine Ecology
Progress Series 214: 237-251.
50. Crossman DJ, Choat JH, Clements KD, Hardy T, McConochie J (2001) Detritus asfood for grazing fishes on coral reefs. Limnology and Oceanography 46:
1596-1605.
51. Choat JH, Clements KD, Robbins WD (2002). The trophic status of herbivorous fishes
on coral reefs. I Dietary analyses. Marine Biology 140: 613-623
52. Gust N, Choat JH, Ackerman J (2002) Demographic plasticity in tropical reef fishes.
Marine Biology 140: 1039-1051.
53. Choat JH, Robertson DR (2002). Age based studies. In: (PF Sale Ed) Coral Reef
Fishes : Diversity and dynamics in a complex system. Academic Press: 57-80.

54. van Herwerden L, Davies CR, Choat JH (2002) Phylogenetic and evolutionary
perspectives of the Indo-Pacific grouper Plectropomus species on the Great
Barrier Reef, Australia. Journal of Fish Biology 60: 1591-1596.
55. Zemke-White L, Choat JH, Clements, KD (2002) A re-evaluation of the diel feeding
hypothesis for marine herbivorous fishes. Marine Biology 141: 571-579.
56. Clements KD, Gray RD, Choat JH (2003) Rapid evolutionary divergences in reeffishes of the family Acanthuridae (Perciformes; Teleostei) MolecularPhylogenetics and Evolution. 26: 190-201
57. Choat JH, Robertson DR, Ackerman J, Posada J (2003) An age-based demographic
analysis of the Caribbean stoplight parrotfish Sparisoma viride. Marine
Ecology Progress Series 246: 265-277
58. Grutter AS, Murphy JM, Choat JH (2003) Cleaner fish drives local fish diversity oncoral reefs. Current Biology 13: 64-67
59. Bellwood DR, Hoey AS, Choat JH (2003) Limited functional redundancy in highdiversity systems: resilience and ecosystem function on coral reefs. Ecology
Letters 6 :281-285
60. Klanten SO, van Herwerden L, Choat JH (2003) Acquiring DNA sequences fromformalin fixed museum specimens: application for reef fish.

Bulletin of Marine

Science 73 (3): 771-776
61. Wilson SK, Bellwood DR, Choat JH, Furnas MJ (2003) Detritus in the epilithic matrix
and its use by coral reef fishes. Oceanography and Marine Biology Annual Review
41: 279-309
62. Bay L, Choat JH, Van Herwerden L, Robertson DR (2003) High genetic diversities
and complex genetic structure in an Indo-Pacific tropical reef fish (Chlorurussordidus): evidence of an unstable evolutionary past?

Marine Biology 144:757-767
63. Klanten S, van Herwerden L, Choat JH, Blair D (2004) Patterns of lineage
diversification in the genus Naso (Acanthuridae). Molecular Phylogenetics andEvolution. 32: 221-235
64. Choat JH, Robbins WD, Clements KD (2004) The trophic status of herbivorous fishes oncoral reefs II: Food processing modes and trophodynamics Marine Biology 145:445-454
65. Munday PL, Hodges AL, Choat JH, Gust N (2004 ) Sex-specific growth effects inprotogynous hermaphrodites. Canadian Journal of Fisheries and Aquatic Sciences325-329
66. Crossman DJ, Choat JH, Clements KD (2005) Nutritional ecology of nominallyherbivorous fishes on coral reefs Marine Ecology Progress Series 296:129-142

61:

67. Robertson DR, Ackerman JL, Choat JH, Posada J, Pitt J (2005) Latitude, habitat
and the geography of demography
Marine Ecology Progress Series

in the ocean surgeonfish, Acanthurus bahianus.
295: 229-244

68. Robertson DR, Ackerman JL, Choat JH, Posada J, Pitt J (2005) Fishing and thelongevity, size and abundance of the west Atlantic surgeonfish, Acanthurusbahianus. Marine Ecology Progress Series 295: 245-256
69. Pears R, Choat JH, Mapstone BM, Begg G (2005) Demography of a large grouper,
Epinephelus fuscoguttatus, from the Great Barrier Reef: implications for harvest
limits and size regulations. Marine Ecology Progress Series (in press)
70. Choat JH, Davies CR, Ackerman JL, Mapstone BM (2005) Demography of a large
teleost, Cheilinus undulatus with a review of size distribution in labrid fishes. MarineEcology Progress Series (in review)