Using movement data from electronic tags in fisheries stock assessment: A review of models, technology and experimental design

Fisheries Research

Volumn 163, Issue , 2015, Pages 152-160

  Sippel, Tim ^a   Paige Eveson, J ^b   Galuardi, Benjamin ^c   Lam, Chi ^c   Hoyle, Simon ^d   Maunder, Mark ^e   Kleiber, Pierre ^f   Carvalho, Felipe ^g   Tsontos, Vardis ^h   Teo, Steven L H ^a   Aires da Silva, Alexandre ^e   Nicol, Simon ^d  

^a NOAA SOUTHWEST FISHERIES SCIENCE CENTER   (United States)

^b CSIRO   (Australia)

^c Large Pelagics Research Center   (United States)

^d SECRETARIAT OF THE PACIFIC COMMUNITY   (New Caledonia)

^e INTER AMERICAN TROPICAL TUNA COMMISSION   (United States)

^f PACIFIC ISLANDS FISHERIES SCIENCE CENTER   (United States)

^g UNIVERSITY OF FLORIDA   (United States)

^h CALIFORNIA INSTITUTE OF TECHNOLOGY   (United States)

Author keywords

Electronic tags; Experimental design; Fish movement; Spatial stock assessment

Indexed keywords

EID: 84920914472     PISSN: 01657836     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.fishres.2014.04.006     Document Type: Article

References (107)

1. Adam S., Sibert J. Population dynamics and movements of skipjack tuna (Katsuwonus pelamis) in the Maldivian fishery: analysis of tagging data from an advection-diffusion-reaction model. Aquat. Living Resour. 2002, 15:13-23.
2. Adam S., Sibert J., Itano D., Holland K. Dynamics of bigeye (Thunnus obsesus) and yellowfin (T. albacares) tuna in Hawaii's pelagic fisheries: analysis of tagging data with a bulk-transfer model incorporating size-specific attrition. Fish. Bull. 2003, 101:215-228.
3. Aires-da-Silva A.M., Maunder M.N., Gallucci V.F., Kohler N.E., Hoey J.J. A spatially structured tagging model to estimate movement and fishing mortality rates for the blue shark (Prionace glauca) in the North Atlantic Ocean. Mar. Freshw. Res. 2009, 60:1029-1043.
4. Anganuzzi A., Hilborn R., Skalski J.R. Estimation of size selectivity and movement rates from mark-recovery data. Can. J. Fish. Aquat. Sci. 1994, 51:734-742.
5. Bayliff W.H. Migrations of yellowfin tuna in the eastern Pacific Ocean as determined from tagging experiments initiated during 1968-1974. Inter-Am. Trop. Tuna Comm. Bull. 1979, 17:445-506.
6. Besbeas P., Lebreton J.-D., Morgan B.J.T. The efficient integration of abundance and demographic data. J. R. Stat. Soc. Ser. C (Appl. Stat.) 2003, 52:95-102.
7. Bestley S., Patterson T.A., Hindell M.A., Gunn J.S. Feeding ecology of wild migratory tunas revealed by archival tag records of visceral warming. J. Anim. Ecol. 2008, 77:1223-1233.
8. Beverton R.J.H., Holt S.J. On the Dynamics of Exploited Fish Populations 1957, Springer, London.
9. Block B.A., Dewar H., Farwell C., Prince E.D. A new satellite technology for tracking the movements of Atlantic bluefin tuna. Proc. Natl. Acad. Sci. U. S. A. 1998, 95:9384.
10. Block B.A., Jonsen I.D., Jorgensen S.J., Winship A.J., Shaffer S.A., Bograd S.J., Hazen E.L., Foley D.G., Breed G.A., Harrison A.-L., Ganong J.E., Swithenbank A., Castleton M., Dewar H., Mate B.R., Shillinger G.L., Schaefer K.M., Benson S.R., Weise M.J., Henry R.W., Costa D.P. Tracking apex marine predator movements in a dynamic ocean. Nature 2011, 475:86-90.
11. Block B.A., Teo S.L.H., Walli A., Boustany A., Stokesbury M.J.W., Farwell C.J., Weng K.C., Dewar H., Williams T.D. Electronic tagging and population structure of Atlantic bluefin tuna. Nature 2005, 434:1121-1127.
12. Breed G.A., Costa D.P., Goebel M.E., Robinson P.W. Electronic tracking tag programming is critical to data collection for behavioral time-series analysis. Ecosphere 2011, 2:art10.
13. Brownie C., Anderson D.R., Burnham K.P., Robson D.S. Statistical Inference from Band Recovery Data: A Handbook 1985, United States Fish and Wildlife Service.
14. Bull B., Francis R., Dunn A., McKenzie A., Gilbert D., Smith M., Bian R. CASAL (C++ Algorithmic Stock Assessment Laboratory): CASAL User Manual v2 2005.
15. Cadrin S.X., Secor D.H. Accounting for spatial population structure in stock assessment: past, present, and future. The Future of Fisheries Science in North America 2009, 405-426. Springer Netherlands, Dordrecht. R.J. Beamish, B.J. Rothschild (Eds.).
16. Carruthers T., McAllister M. Spatial surplus production modelling of Atlantic tunas and billfish. Collect. Vol. Sci. Pap. ICCAT 2011, 66:2182-2201.
17. Carruthers T.R., McAllister M.K. Quantifying tag reporting rates for Atlantic tuna fleets using coincidental tag returns. Aquat. Living Resour. 2010, 23:343-352.
18. Carruthers T.R., McAllister M.K., Taylor N.G. Spatial surplus production modeling of Atlantic tunas and billfish. Ecol. Appl. 2011, 21:2734-2755.
19. Dagorn L., Holland K.N., Itano D.G. Behavior of yellowfin (Thunnus albacares) and bigeye (T. obesus) tuna in a network of fish aggregating devices (FADs). Mar. Biol. 2007, 151:595-606.
20. Domeier M.L., Dewar H., Nasby-Lucas N. Mortality rate of striped marlin (Tetrapturus audax) caught with recreational tackle. Mar. Freshw. Res. 2003, 54:435-446.
21. Evans K., Abascal F., Kolody D., Sippel T., Holdsworth J., Maru P. The horizontal and vertical dynamics of swordfish in the South Pacific Ocean. J. Exp. Mar. Biol. Ecol. 2014, 450:55-67.
22. Eveson J.P., Basson M., Hobday A.J., Jech J.M. Using electronic tag data to improve mortality and movement estimates in a tag-based spatial fisheries assessment model. Can. J. Fish. Aquat. Sci. 2012, 69:869-883.
23. Eveson J.P., Laslett G.M., Polacheck T. A spatial model for estimating mortality rates, abundance and movement probabilities from fishery tag-recovery data. Modeling Demographic Processes in Marked Populations, Environmental and Ecological Statistics 2009, 987-1010. Springer, USA. D.L. Thomson, E.G. Cooch, M.J. Conroy, G.P. Patil, G.P. Patil, T.G. Gregoire, A.B. Lawson, B.D. Nussbaum (Eds.).
24. Fedak M.A. Marine animals as platforms for oceanographic sampling: a win-win situation for biology and operational oceanography. Mem. Natl. Inst. Polar Res. 2004, 58:133-147.
25. Fournier D.A., Hampton J., Sibert J.R. MULTIFAN-CL: a length-based, age-structured model for fisheries stock assessment, with application to South Pacific albacore, Thunnus alalunga. Can. J. Fish. Aquat. Sci. 1998, 55:2105-2116.
26. Fournier D.A., Skaug H.J., Ancheta J., Ianelli J., Magnusson A., Maunder M.N., Nielsen A., Sibert J. AD Model Builder: using automatic differentiation for statistical inference of highly parameterized complex nonlinear models. Optim. Methods Softw. 2012, 27:233-249.
27. Fromentin J.-M. Descriptive analysis of the ICCAT bluefin tuna tagging database. Collect. Vol. Sci. Pap. ICCAT 2002, 54:353-362.
28. Galuardi B., Lam C.H. Chapter nineteen - telemetry analysis of highly migratory species. Stock Identification Methods 2014, 447-476. Academic Press, San Diego. 2d ed. S.X. Cadrin, L.A. Kerr, S. Mariani (Eds.).
29. Galuardi B., Lutcavage M. Dispersal routes and habitat utilization of juvenile Atlantic bluefin tuna, Thunnus thynnus, tracked with mini PSAT and archival tags. PLoS ONE 2012, 7:e37829.
30. Galuardi B., Royer F., Golet W., Logan J., Neilson J., Lutcavage M. Complex migration routes of Atlantic bluefin tuna (Thunnus thynnus) question current population structure paradigm. Can. J. Fish. Aquat. Sci. 2010, 67:966-976.
31. Gillanders B.M., Ferrell D.J., Andrew N.L. Estimates of movement and life-history parameters of yellowtail kingfish (Seriola lalandi): how useful are data from a cooperative tagging programme?. Mar. Freshw. Res. 2001, 52:179-192.
32. Goethel D.R., Quinn T.J., Cadrin S.X. Incorporating spatial structure in stock assessment: movement modeling in marine fish population dynamics. Rev. Fish. Sci. 2011, 19:119-136.
33. Gunn J., Polacheck T., Davis T., Sherlock M., Betlehem A. The development and use of archival tags for studying the migration, behaviour and physiology of southern bluefin tuna, with an assessment of the potential for transfer of the technology to groundfish research. Proc. ICES Symp. Fish Migration 1994, 1-23.
34. Hampton J. Effect of tagging on the condition of southern bluefin tuna, Thunnus maccoyii. Aust. J. Mar. Freshw. Res. 1986, 38:699-705.
35. Hampton J., Fournier D.A. A spatially disaggregated, length-based, age-structured population model of yellowfin tuna (Thunnus albacares) in the western and central Pacific Ocean. Mar. Freshw. Res. 2001, 52:937-963.
36. Hartog J.R., Patterson T.A., Hartmann K., Jumppanen P., Cooper S., Bradford R. Developing integrated database systems for the management of electronic tagging data. Tagging and Tracking of Marine Animals with Electronic Devices 2009, 367-380.
37. Hearn W.S., Polacheck T., Pollock K.H., Whitelaw W. Estimation of tag reporting rates in agestructured multicomponent fisheries where one component has observers. Can. J. Fish. Aquat. Sci. 1999, 56:1255-1265.
38. Hilborn R. Determination of fish movement patterns from tag recoveries using maximum likelihood estimators. Can. J. Fish. Aquat. Sci. 1990, 47:635-643.
39. Hilborn R., Walters C. Quantitative Fisheries Stock Assessment: Choice, Dynamics and Uncertainty 1992, Chapman & Hall, London.
40. Hill R.D. Theory of geolocation by light levels. Elephant Seals. Population Ecology, Behavior, and Physiology 1994, 227-236.
41. Hill R.D., Braun M.J. Geolocation by light level. Electronic Tagging and Tracking in Marine Fisheries: Proceedings of the Symposium on Tagging and Tracking Marine Fish with Electronic Devices 2001, 315. East-West Center, University of Hawaii.
42. Hoenig J.M., Barrowman N.J., Pollock K.H., Brooks E.N., Hearn W.S., Polacheck T. Models for tagging data that allow for incomplete mixing of newly tagged animals. Can. J. Fish. Aquat. Sci. 1998, 55:1477-1483.
43. Holdsworth J., Sippel T., Block B. Near real time satellite tracking of striped marlin (Kajikia audax) movements in the Pacific Ocean. Mar. Biol. 2009, 156:505-514.
44. Hoolihan J.P., Luo J., Abascal F.J., Campana S.E., De Metrio G., Dewar H., Domeier M.L., Howey L.A., Lutcavage M.E., Musyl M.K., et al. Evaluating post-release behaviour modification in large pelagic fish deployed with pop-up satellite archival tags. ICES J. Mar. Sci. 2011, 68:880-889.
45. Ishii T. Attempt to estimate migration of fish population with survival parameters from tagging experiment data by the simulation method. Investig. Pesquera 1979, 43:301-317.
46. Kaltongga B. Regional Tuna Tagging Project: data summary. Oceanic Fisheries Program Secretariat of the Pacific Community, Technical Report No. 35 1988, 70.
47. Kell L.T., Mosqueira I., Grosjean P., Fromentin J.-M., Garcia D., Hillary R., Jardim E., Mardle S., Pastoors M.A., Poos J.J., Scott F., Scott R.D. FLR: an open-source framework for the evaluation and development of management strategies. ICES J. Mar. Sci. 2007, 64:640-646.
48. Kerstetter D.W., Graves J.E. Post-release survival of white marlin released from commercial pelagic longline gear in the western North Atlantic. Fish. Bull. 2006, 104:434-444.
49. Kleiber P., Argue A.W., Kearney R.E. Assessment of Pacific skipjack tuna (Katsuwonus pelamis) resources by estimating standing stock and components of population turnover from tagging data. Can. J. Fish. Aquat. Sci. 1987, 44:1122-1134.
50. Kleiber P., Hampton J. Modeling effects of FADs and Islands on movements of skipjack tuna (Katsuwonus pelamis): estimating parameters from tagging data. Can. J. Fish. Aquat. Sci. 1994, 51:2642-2653.
51. Kolody D., Campbell R., Davies N. A MULTIFAN-CL Stock Assessment of South-West Pacific Swordfish 1952-2007 (No. WCPFC-SC-4/SA-WP-6) 2008.
52. Kranstauber B., Cameron A., Weinzerl R., Fountain T., Tilak S., Wikelski M., Kays R. The Movebank data model for animal tracking. Environ. Model. Softw. 2011, 26:834-835.
53. Kranstauber B., Kays R., LaPoint S.D., Wikelski M., Safi K. A dynamic Brownian bridge movement model to estimate utilization distributions for heterogeneous animal movement. J. Anim. Ecol. 2012, 81:738-746.
54. Kurota H., McAllister M.K., Lawson G.L., Nogueira J.I., Teo S.L.H., Block B.A. A sequential Bayesian methodology to estimate movement and exploitation rates using electronic and conventional tag data: application to Atlantic bluefin tuna (Thunnus thynnus). Can. J. Fish. Aquat. Sci. 2009, 66:321-342.
55. Lam C.H., Nielsen A., Sibert J.R. Improving light and temperature based geolocation by unscented Kalman filtering. Fish. Res. 2008, 91:15-25.
56. Lam C.H., Tsontos V.M. Integrated management and visualization of electronic tag data with tagbase. PLoS ONE 2011, 6:e21810.
57. Langley A., Million J. Determining an appropriate tag mixing period for the Indian Ocean yellowfin tuna stock assessment. IOTC-2012-WPTT-14-31 2012, 53.
58. Latour R.J., Hoenig J.M., Olney J.E., Pollock K.H. A simple test for nonmixing in multiyear tagging studies: application to striped bass tagged in the Rappahannock River, Virginia. Trans. Am. Fish. Soc. 2001, 130:848-856.
59. Leroy B., Nicol S., Lewis A., Hampton J., Kolody D., Caillot S., Hoyle S. Lessons learned from implementing three, large-scale tuna tagging programmes in the western and central Pacific Ocean. Fish. Res. 2013, (in this volume).
60. Lutcavage M.E., Brill R.W., Skomal G.B., Chase B.C., Howey P.W. Results of pop-up satellite tagging of spawning size class fish in the Gulf of Maine: do North Atlantic bluefin tuna spawn in the mid-Atlantic?. Can. J. Fish. Aquat. Sci. 1999, 56:173-177.
61. Maunder M. Integrated tagging and catch-at-age analysis (ITCAAN): model development and simulation testing. Spatial Processes and Management of Marine Populations 2001, 123-146. University of Alaska Sea Grant, Fairbanks.
62. Maunder M.N. Integration of Tagging and Population Dynamics Models in Fisheries Stock Assessment 1998, (Ph.D. thesis), University of Washington, 306 pp.
63. Maunder M.N. Is it time to discard the Schaefer model from the stock assessment scientist's toolbox?. Fish. Res. 2003, 61:145-149.
64. Maunder M.N., Langley A.D. Integrating the standardization of catch-per-unit-of-effort into stock assessment models: testing a population dynamics model and using multiple data types. Fish. Res. 2004, 70:389-395.
65. Maunder M.N., Punt A.E. Standardizing catch and effort data: a review of recent approaches. Fish. Res. 2004, 70:141-159.
66. Maunder M.N., Punt A.E. A review of integrated analysis in fisheries stock assessment. Fish. Res. 2013, 142:61-74.
67. Methot R.D., Wetzel C.R. Stock synthesis: a biological and statistical framework for fish stock assessment and fishery management. Fish. Res. 2013, 142:86-99.
68. Miller T.J., Andersen P.K. A finite-state continuous-time approach for inferring regional migration and mortality rates from archival tagging and conventional tag-recovery experiments. Biometrics 2008, 64:1196-1206.
69. Morales J.M., Moorcroft P.R., Matthiopoulos J., Frair J.L., Kie J.G., Powell R.A., Merrill E.H., Haydon D.T. Building the bridge between animal movement and population dynamics. Philos. Trans. R. Soc. B Biol. Sci. 2010, 365:2289-2301.
70. Musyl M.K., Brill R.W., Curran D.S., Fragoso N.M., McNaughton L.M., Nielsen A., Kikkawa B.S., Moyes C.D. Postrelease survival, vertical and horizontal movements, and thermal habitats of five species of pelagic sharks in the central Pacific Ocean. Fish. Bull. 2011, 109:341-368.
71. Nathan R., Getz W.M., Revilla E., Holyoak M., Kadmon R., Saltz D., Smouse P.E. A movement ecology paradigm for unifying organismal movement research. Proc. Natl. Acad. Sci. 2008, 105:19052.
72. Nielsen A., Sibert J.R. State-space model for light-based tracking of marine animals. Can. J. Fish. Aquat. Sci. 2007, 64:1055-1068.
73. Nielsen A., Sibert J.R., Ancheta J., Galuardi B., Lam C.H. trackit: Track Tagged Individuals from Light Measurements, R Package Version 0 2012, 2-6.
74. Tagging and Tracking of Marine Animals with Electronic Devices 2009, Springer, Dordrecht. J.L. Nielsen, H. Arrizabalaga, N. Fragoso, A. Hobday, M. Lutcavage, J. Sibert (Eds.).
75. O'Dor R.K., Stokesbury M.J.W. The ocean tracking network - adding marine animal movements to the global ocean observing system. Tagging Tracking Mar. Anim. Electron. Devices 2009, 91-100.
76. Patterson T.A., Evans K., Carter T.I., Gunn J.S. Movement and behaviour of large southern bluefin tuna (Thunnus maccoyii) in the Australian region determined using pop-up satellite archival tags. Fish. Oceanogr. 2008, 17:352-367.
77. Patterson T.A., Hartmann K. Designing satellite tagging studies: estimating and optimizing data recovery. Fish. Oceanogr. 2011, 20:449-461.
78. Patterson T.A., McConnell B.J., Fedak M.A., Bravington M.V., Hindell M.A. Using GPS data to evaluate the accuracy of state-space methods for correction of Argos satellite telemetry error. Ecology 2010, 91:273-285.
79. Patterson T.A., Thomas L., Wilcox C., Ovaskainen O., Matthiopoulos J. State-space models of individual animal movement. Trends Ecol. Evol. 2008, 23:87-94.
80. Pedersen M.W., Weng K.C. Estimating individual animal movement from observation networks. Methods Ecol. Evol. 2013, 4:920-929.
81. Polacheck T., Eveson J.P., Laslett G.M., Pollock K.H., Hearn W.S. Integrating catch-at-age and multiyear tagging data: a combined Brownie and Petersen estimation approach in a fishery context. Can. J. Fish. Aquat. Sci. 2006, 63:534-548.
82. Polacheck T., Paige Eveson J., Laslett G.M. Classifying tagging experiments for commercial fisheries into three fundamental types based on design, data requirements and estimable population parameters. Fish Fish. 2010, 11:133-148.
83. Porch C., Restrepo V., Turner S., Scott G. Virtual population analyses of Atlantic bluefin tuna incorporating movement and tagging data. ICCAT Collective Volume of Scientific Papers. ICCAT 1995, 44:183-208.
84. Porch C.E., Turner S.C., Powers J.E. Virtual population analyses of Atlantic bluefin tuna with alternative models of transatlantic migration: 1970-1997. Collect. Vol. Sci. Pap. ICCAT 2001, 52:1022-1045.
85. Punt A., Butterworth D. Use of tagging data within a VPA formalism to estimate migration rates of bluefin tuna across the North Atlantic. Collect. Vol. Sci. Pap. ICCAT 1995, 44:166-182.
86. R Core Development Team R: A Language and Environment for Statistical Computing. Vienna, Austria 2012.
87. Schaefer K.M., Fuller D.W. Horizontal movements of bigeye tuna (Thunnus obesus) in the eastern Pacific Ocean, as determined from conventional and archival tagging experiments initiated during 2000-2005. Inter-Am. Trop. Tuna Comm. Bull. 2009, 24:191-247.
88. Schaefer K.M., Fuller D.W., Block B.A. Movements, behavior, and habitat utilization of yellowfin tuna (Thunnus albacares) in the Pacific Ocean off Baja California, Mexico, determined from archival tag data analyses, including unscented Kalman filtering. Fish. Res. 2011, 112:22-37.
89. Schnute J.T., Richards L.J. Analytical models for fishery reference points. Can. J. Fish. Aquat. Sci. 1998, 55:515-528.
90. Seber G. A review of estimating animal abundance. Biometrics 1986, 42:267-292.
91. Senina I., Royer F., Lehodey P., Hampton J., Nicol S., Ogura M., Kiyofuji H., Sibert J. Integrating conventional and electronic tagging data into SEAPODYM. Pelagic Fish. Res. Progr. Newslett. 2012, 16:9-14.
92. Sibert J.R., Hampton J., Fournier D.A., Bills P.J. An advection-diffusion-reaction model for the estimation of fish movement parameters from tagging data, with application to skipjack tuna (Katsuwonus pelamis). Can. J. Fish. Aquat. Sci. 1999, 56:925-938.
93. Sibert J.R., Lutcavage M.E., Nielsen A., Brill R.W., Wilson S.G. Interannual variation in large-scale movement of Atlantic bluefin tuna (Thunnus thynnus) determined from pop-up satellite archival tags. Can. J. Fish. Aquat. Sci. 2006, 63:2154-2166.
94. Sibert J.R., Musyl M.K., Brill R.W. Horizontal movements of bigeye tuna (Thunnus obesus) near Hawaii determined by Kalman filter analysis of archival tagging data. Fish. Oceanogr. 2003, 12:141-151.
95. Sippel T., Holdsworth J., Dennis T., Montgomery J. Investigating behaviour and population dynamics of striped marlin (Kajikia audax) from the Southwest Pacific Ocean with satellite tags. PLoS ONE 2011, 6:e21087.
96. Skaug H., Fournier D., Nielsen A., Magnusson A., Bolker B. glmmADMB: generalized linear mixed models using AD model builder. R package version 0.6 2011, 4.
97. Skellam J.G. Random dispersal in theoretical populations. Biometrika 1951, 38:196-218.
98. Skomal G. Evaluating the physiological and physical consequences of capture on post-release survivorship in large pelagic fishes. Fish. Manage. Ecol. 2007, 14:81-89.
99. Smith P. Determining Fish Movement from an 'Archival' Tag: Precision of Geographical Positions Made from a Time Series of Swimming Temperature and Depth. NOAA Technical Memorandum 1986.
100. Sumner M. The Tag Location Problem 2011, (Ph.D. thesis), University of Tasmania, Hobart.
101. Taylor I., Stewart I., Hicks A., Garrison T., Punt A., Wallace J., Wetzel C. r4ss: R code for Stock Synthesis. R package version 1.16 2011.
102. Taylor N.G., McAllister M.K., Lawson G.L., Carruthers T., Block B.A. Atlantic bluefin tuna: a novel multistock spatial model for assessing population biomass. PLoS ONE 2011, 6:e27693.
103. Vecchi G.A., Harrison M.J. An observing system simulation experiment for the Indian Ocean. J. Climate 2007, 20:3300-3319.
104. Welch D.W., Boehlert G.W., Ward B.R. POST - the Pacific Ocean salmon tracking project. Oceanol. Acta 2002, 25:243-253.
105. Whitlock R.E., McAllister M.K., Block B.A. Estimating fishing and natural mortality rates for Pacific bluefin tuna (Thunnus orientalis) using electronic tagging data. Fish. Res. 2012, 119-120:115-127.
106. Wilson S.G., Stewart B.S., Polovina J.J., Meekan M.G., Stevens J.D., Galuardi B. Accuracy and precision of archival tag data: a multiple-tagging study conducted on a whale shark (Rhincodon typus) in the Indian Ocean. Fish. Oceanogr. 2007, 16:547-554.
107. Xiao Y. An individual-based approach to evaluating experimental designs for estimating rates of fish movement from tag recoveries. Ecol. Model. 2000, 128:149-163.