Sun et al.: Reproductive biology of Makaira nigricans 
431 
parameters could be explored further by using data 
collected over a broader spatial domain and hence an 
exploration could be conducted to determine whether 
this variation is related to abiotic or biotic factors. We 
recommend that the methods used in this study be 
applied to estimate reproductive parameters for billfish 
and other pelagic fish. 
Acknowledgments 
The authors express sincere gratitude to A. E. Punt, 
School of Aquatic and Fishery Sciences, University of 
Washington, for his valuable comments and comprehen- 
sive editing of an earlier version of this manuscript. The 
authors also thank the four anonymous reviewers for 
their constructive comments. This study was partially 
financially supported by the National Science Coun- 
cil and the Fisheries Agency, Council of Agriculture, 
Taiwan, through grants NSC89-2316-B-002-038 and 
90AS-1.4.5-FA-F2(3) to Chi-Lu Sun. 
Literature cited 
Arocha, F. 
2002. Oocyte development and maturity classification 
of swordfish from the north-western Atlantic. J. Fish 
Biol. 60:13-27. 
Arocha, F., and Barrio. 
2009. Sex ratios, spawning seasonality, sexual maturity, 
and fecundity of white marlin ( Tetrapturus albidus ) from 
the western central Atlantic. Fish. Res. 95:98-111. 
Arocha, F., and L. Marcano. 
2008. Life history characteristics of blue marlin, white 
marlin, and sailfish from the eastern Caribbean Sea 
and adjacent waters. In Proceedings of the fourth 
world fisheries congress: reconciling fisheries with 
conservation (J. Nielsen, J. Dodson, K. Friedland, T. 
Hamon, N. Hughes, J. Musick, and E. Verspoor, eds), p. 
1481-1491. Am. Fish. Soc., Symposium 49, Vancouver, 
British Columbia, Canada, 2-6 May, 2004. 
Begg G. A., G. A. Hopper. 
1997. Feeding patterns of school mackerel ( Scomb - 
eromorus queenslandicus) and spotted mackerel (S. 
munroi) in Queensland east-coast waters. Aust. J. 
Mar. Freshw. Res. 48:565-571 
Buonaccorsi, V. R, K. S. Reece, L. W. Morgan, and J. E. Graves. 
1999. Geographic distribution of molecular variance 
within the blue marlin (Makaira nigricans): A hier- 
archical analysis of allozyme, single-copy nuclear 
DNA, and mitochondrial DNA markers. Evolution 
53:568-579. 
Chen, B. J. 
2001. Age and growth of the blue marlin, Makaira 
mazara, in the western Pacific Ocean. Master’s thesis, 
76 p. Institute of Oceaography, National Taiwan Uni- 
versity, Taipei, Taiwan. 
Chiang, W. C., C. L. Sun, S. Z. Yeh, W. C. Su, and D. C. Liu. 
2006a. Spawning frequency and batch fecundity of 
the sailfish ( Istiophorus platypterus ) (Istiophoridae) 
in waters off Eastern Taiwan. Zool. Stud. 45:483- 
491. 
Chiang, W. C„ C. L. Sun, S. Z. Yeh, W. C. Su, D. C. Liu, and 
W. Y. Chen. 
2006b. Sex ratios, size at sexual maturity, and spawn- 
ing seasonality of sailfish Istiophorus platypterus from 
eastern Taiwan. Bull. Mar. Sci. 79:727-737. 
Cyr, E. C. 
1987. Age, growth and reproduction of blue marlin, 
Makaira nigricans, from South Carolina tourmament 
collections. Master’s thesis, 41 p. Univ. South Caro- 
lina, Columbia, SC. 
DeMartini, E. E., J. H. Uchiyama, and H. A. Williams. 
2000. Sexual maturity, sex ratio, and size composition of 
swordfish, Xiphias gladius, caught by the Hawaii-based 
pelagic longline fishery. Fish. Bull. 98:489-506. 
deSylva, D. P. 
1974. A review of world sport fishery for billfishes 
(Istiophoridae and Xiphidae). In Proceedings of the 
international billfish symposium (R. S. Shomura, and 
F. Williams, eds.), p. 12-33. NOAA Technical Report 
NMFS SSRF-675, Kailua-Kona, Hawaii, 9-12 August, 
1972. 
deSylva, D. P, and P. R. Breder. 
1997. Reproduction, gonad histology, and spawning cycles 
of North Atlantic billfishes (Istiophoridae). Bull. Mar. 
Sci. 60:668-697. 
Erdman, D. S. 
1968. Spawning cycle sex ratio and weights of blue marlin 
off Puerto Rico and Virgin Islands. Trans. Am. Fish. 
Soc. 97:131-137. 
Fournier, D. A. 
2000. An introduction to AD model builder, vers. 4: for 
use in nonlinear modelling and statistics, 127 p. Otter 
Research Ltd, Sidney, Canada. 
Grier, H. J. 
1981. Cellular-organization of the testis and spermato- 
genesis in fishes. Am. Zool. 21:345-357. 
Hinton, M. G. 
2001. Status of blue marlin in the Pacific Ocean. IATTC 
Stock Assess. Rep. 1:284-318. 
Hopper, C. N. 
1990. Patterns of Pacific blue marlin reproduction in 
Hawaiian waters. In Proceedings of the second inter- 
national billfish symposium; 1—5 August 1988 (R. H. 
Stroud, ed.), p. 29-39. Kailua-Kona, HI. 
Hunter, J. R., N. C. H. Lo, and R. J. Leong 
1985. Batch fecundity in multiple spawning fishes, /n An 
egg production method for estimating spawning biomass 
of pelagic fish: Application to the northern anchovy, 
Engraulis mordax (R. Lakser, ed.), p. 67—78. NOAA 
Tech. Rep. NMFS 36. 
Hunter, J. R., and B. J. Macewicz. 
2003. Improving the accuracy and precision of reproduc- 
tive information used in fisheries. In Report of the 
working group on modern approaches to assess maturity 
and fecundity of warm- and cold-water fish and squids, 
4-7 September, 2001 (O. S. Kjesbu, J. R. Hunter, and P. 
R. Witthames, eds), p. 57—68. Bergen, Norway. 
Hunter, J. R., and B. J. Macewicz. 
1985. Measurement of spawning frequency in multiple 
spawning fishes. In An egg production method for esti- 
mating spawning biomass of pelagic fish: application 
to the northern anchovy, Engraulis mordax (R. Lakser, 
ed.), p. 79-94. NOAA Tech. Rep. NMFS 36. 
King, M. 
1995. Fisheries biology assessment and management, 
341 p. Fishing News Books, Oxford, England. 
