420 Abstract—Demographic parameters Demographic characteristics of exploited from seven exploited coral reef lutja- nid species were compared as a case tropical lutjanids: a comparative analysis study of the implications of intra- family variation in life histories for 1 2 multispecies harvest management. Michelle R. Heupel (contact author) Ann Penny Modal lengths varied by 4 cm among Ashley J. Williams2,3 Jacob P Kritzer6 four species (Lutjanus fulviflamma, L. David J. Welch2,4 Ross J. Marriott7 vitta, L. carponotatus, L. adetii), which were at least 6 cm smaller than the Campbell R. Davies5 Bruce D. Mapstone5 modal lengths of the largest species Email address for contact author: [email protected] (L. gibbus, Symphorus nematophorus, 1 5 Aprion virescens). Modal ages, indicat- School of Earth and Environmental Sciences CSIRO Marine and Atmospheric Research ing ages of full selection to fishing James Cook University GPO Box 1538 gear, were 10 years or less for all spe- Townsville, Queensland, 4811, Australia Hobart, Tasmania, 7001, Australia cies, but maximum ages ranged from 2 Fishing and Fisheries Research Centre 6 Environmental Defense Fund 12 (L. gibbus) to 36 years (S. nema- School of Earth and Environmental Sciences 18 Tremont Street, Suite 850 tophorus). Each species had a unique James Cook University Boston, Massachusetts 02108 growth pattern, with differences in Townsville, Queensland, 4811, Australia 7 Department of Fisheries WA length-at-age and mean asymptotic 3 Oceanic Fisheries Program Western Australian Fisheries and fork length (L∞), but smaller species Secretariat of the Pacific Community Marine Research Laboratories generally grew fast during the first BP 5, 98848 Noumea, New Caledonia PO Box 20 1–2 years of life and larger species 4 Queensland Primary Industries and Fisheries North Beach, Western Australia 6920 grew more slowly over a longer period. PO Box 1085 Australia Total mortality rates varied among Oonoonba, Queensland 4811, Australia species; L. gibbus had the highest mortality and L. fulviflamma, the lowest mortality. The variability in life history strategies of these tropical lutjanids makes generalizations about lutjanid life histories difficult, but the fact that all seven had characteristics that would make them particularly Knowledge of the life history charac- it is inappropriate to apply the same vulnerable to fishing indicates that teristics of an exploited species can management strategy to all those harvest of tropical lutjanids should help elucidate inherent resilience to species. Here we consider the lutja- be managed with caution. anthropogenic effects such as fishing nids as a case study. (Parent and Schriml, 1995; Jennings The Lutjanidae comprises approxi- et al., 1998; Musick et al., 2000). mately 103 species with a wide ar- Interspecific variation in life history ray of size and body form, making it characteristics may indicate that one of the largest and most diverse some species are more susceptible to families of fish (Carpenter and Niem, overfishing than others. Life history 2001). Lutjanids are found in tropi- characteristics vary widely among cal waters around the globe and are families of coral reef fish (Sale, 1991; often associated with reef habitats Gust et al., 2002)—a feature particu- (Carpenter and Niem, 2001). Lutja- larly evident within the Lutjanidae. nids are of high commercial value Some lutjanid species grow to sizes throughout the world and are taken in excess of 1 m (e.g., cubera snapper regularly in artisanal, recreational, [Lutjanus cyanopterus], red snapper and commercial fisheries (Newman et [L. sebae]) and others reach smaller al., 1996; Kaunda-Arara and Ntiba, maximum sizes of less than 30 cm 1997; Marriott et al., 2007; Amezcua (e.g., two-spot banded snapper [L. et al., 2006). The propensity of lut- biguttatus], bluestriped snapper [L. janids to aggregate, particularly for notatus]). Variable life spans and spawning, and their predictably reef- spatial distributions also have been associated distribution, make them Manuscript submitted 5 November 2009. recorded within the family. Lutja- potentially susceptible to overfishing. Manuscript accepted 7 July 2010. Fish. Bull. 108:420–432 (2010). nids are typically gonochoristic (i.e., Concerns have been raised about the do not change sex) and often exhibit level of harvest and sustainability The views and opinions expressed sex specific variation in life history of fishing for lutjanid populations in or implied in this article are those traits (Polovina and Ralston, 1987). some regions (Kamukuru et al., 2005; of the author (or authors) and do not necessarily reflect the position A high degree of variation in life his- Amezcua et al., 2006). of the National Marine Fisheries tory strategies among reef fish spe- Little is known about the specific Service, NOAA. cies within a family would suggest spatial distribution and demograph- Heupel et al.: Demographic characteristics of exploited tropical lutjanids 421 ic characteristics of many lutjanid species (but see Newman et al., 1996, 2000a; Kaun- da-Arara and Ntiba, 1997; Kritzer, 2002, 2004; Amezcua et al., 2006; Marriott et al., 2007). Many species remain unstudied and comparisons among species are few, even though research has covered much of the family’s geographic range. including Mexico Lizard Island (Arreguín-Sánchez and Manickchand-Heile- Coral Sea man, 1998; Amezcua et al., 2006), Kenya (Kaunda-Arara and Ntiba, 1997), the United States (Wilson and Nieland, 2001; Meyer et al., 2007), Japan (Shimose and Tachihara, Townsville 2005), the Great Barrier Reef (Newman et al., 1996, 2000a; Marriott et al., 2007), Ara- Mackay bian Gulf (Grandcourt et al., 2006), and the Indian Ocean (Newman and Dunk, 2003; Storm Cay Pilling et al., 2000). The few data that do exist indicate that at least some lutjanid species are long lived with life spans over 30 years (Newman et al., 1996; Wilson and Nieland, 2001; Newman and Dunk, 2003; Australia Marriott et al., 2007). In this study we compared the demo- graphic characteristics of seven lutjanid species on mid and outer shelf reefs of the Great Barrier Reef (GBR), Australia, as a case study of the implications for intra- Figure 1 family demographic diversity for managing Map of the Great Barrier Reef region off the east coast of Australia multispecies fisheries. The species exam- showing the four sampling regions where lutjanids were collected for ined included the following: chinamanfish use in this study. [Symphorus nematophorus]; green jobfish [Aprion virescens]; dory snapper [Lutjanus fulviflamma]; yellow-banded snapper [L. adetii]; hump- Materials and methods back red snapper [L. gibbus]; brownstripe red snapper [L. vitta]; and spanish flag snapper [L. carponotatus]. Sample collection These species are among the most common lutjanids caught on the GBR after the highly valued red snap- Individuals of the seven lutjanids considered here were pers (L. erythropterus, L. malabaricus, L. sebae) (New- collected during the effects of line fishing (ELF) experi- man et al., 2000b). ment1 (Campbell et al., 2001) between 1995 and 2005 Demographic characteristics examined for each from four regions of the GBR spanning 7° of latitude: species included size, age, growth, mortality, and Lizard Island, 14°S; Townsville, 18°S; Mackay, 20°S; and sex-specific patterns. Demographic parameters were Storm Cay, 21°S (Fig. 1). All fishing was done by hand- compared to quantify differences in life history strat- line with the same gear as that used in the operational egies among species and to infer the likely resilience commercial fishery, but survey effort was stratified to of each species to fishing impacts. This information ensure roughly equal distribution around each reef and was used to assess whether common management over two depth strata (above 12 m and below 15 m). Oto- measures would be appropriate across species within liths (sagittae) and gonads of fish were either dissected the lutjanid family. Such descriptive field studies also on the day of capture and processed as described below can be useful for providing the data sets to be used or fish heads were removed from filleted fish carcasses in in meta-analyses for further developing our collective the field, frozen, and otoliths were dissected upon return understanding of life history theory (e.g., Roff, 1984; to the laboratory. Otoliths of five of the seven species Molloy et al., 2007). In this study we describe differ- were examined for age analysis (A. virescens, S. nema- ent aspects of biology across these closely related spe- cies. Samples from the same shallow coral reef habi- 1 Mapstone, B.D., C. R. Davies, L. R. Little, A. E. Punt, A. D. tats allowed us to control for taxonomic, geographic, M. Smith, F. Pantus, D. C. Lou, A. J. Williams, A. Jones, A. and environmental effects to provide insight into the M. Ayling, G. R. Russ, and A. D. McDonald. 2004. The effects of line fishing on the Great Barrier Reef and evalu- similarities or differences in life history strategies ations of alternative potential management strategies. CRC and thus theoretical trade-offs in evolved traits for Reef Research Centre Technical Report no 52, 205 p. CRC moderate-size gonochoristic teleosts. Reef Research Centre, Townsville, Australia. 422 Fishery Bulletin 108(4) tophorus, L. carponotatus, L. gibbus, L. fulviflamma). examined to define the sex and maturity of individuals, Sufficient specimens were collected for comparisons but low sample sizes prevented meaningful analysis of among regions for only one species (L. carponotatus) detailed histological staging or gonadosomatic index and specimens for all other species were pooled across analyses. Gonads were dissected and either frozen or regions and years for analyses. preserved in FAACC (formaldehyde 4%, acetic acid 5%, calcium chloride 1.3%) or 10% buffered formalin imme- Fish processing diately after removal. Tissues were preserved in FAACC before May 1999 and 10% buffered formalin thereafter. All fish were measured to the nearest millimeter fork The sex and maturity of L. carponotatus was estab- length (FL) and weighed to the nearest gram before lished by macroscopic methods (Kritzer, 2004).