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Von Bertalanffy growth parameters of non-fish marine organisms. Palomares, Maria Lourdes D.; Pauly, Daniel 2008

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ISSN 1198-6727  Fisheries Centre Research Reports  2008 Volume 16 Number 10  Von Bertalanffy Growth Parameters of Non-Fish Marine Organisms  Fisheries Centre, University of British Columbia, Canada  VON BERTALANFFY GROWTH PARAMETERS OF NON-FISH MARINE ORGANISMS  Edited by Maria Lourdes D. Palomares and Daniel Pauly  Fisheries Centre Research Reports 16(10) 137 pages © published 2008 by The Fisheries Centre, University of British Columbia 2202 Main Mall Vancouver, B.C., Canada, V6T 1Z4  ISSN 1198-6727  Fisheries Centre Research Reports 16(10) 2008 VON BERTALANFFY GROWTH PARAMETERS OF NON-FISH MARINE ORGANISMS Edited by  Maria Lourdes D. Palomares and Daniel Pauly  CONTENTS  PAGE  DIRECTOR’S FOREWORD ................................................................................................................................. 1 Growth of marine mammals M.L. Deng Palomares, Patricia M.E. Sorongon, Andrea Hunter and Daniel Pauly .............................. 2 Life-history patterns in marine birds Vasiliki S. Karpouzi and Daniel Pauly .................................................................................................... 27 Growth of marine reptiles M.L. Deng Palomares, Christine Dar, Gary Fry ..................................................................................... 32 Growth of leatherback sea turtles (Dermochelys coriacea) in captivity with inferences on growth in the wild T. Todd Jones, Mervin Hastings, Brian Bostrom, Daniel Pauly and David R. Jones ........................... 82 Length-weight relationships and additional growth parameters for sea turtles Colette Wabnitz and Daniel Pauly .......................................................................................................... 98 A preliminary compilation of life-history data for Mediterranean marine invertebrates Charalampos A. Apostolidis and Konstantinos I. Stergiou.................................................................... 102 Growth estimates of the spiny lobster, Panulirus longipes (A. Milne-Edwards, 1868) in captivity Len R. Garces .......................................................................................................................................... 122 Development and growth of edible oysters (Ostreidae) in Papua New Guinea J. L. Maclean and M.L. Deng Palomares................................................................................................ 127  A Research Report from the Fisheries Centre at UBC 137 pages © Fisheries Centre, University of British Columbia, 2008  FISHERIES CENTRE RESEARCH REPORTS ARE ABSTRACTED IN THE FAO AQUATIC SCIENCES AND FISHERIES ABSTRACTS (ASFA)  ISSN 1198-6727  Von Bertalanffy Growth Parameters of Non-Fish Marine Organisms, Palomares, M.L.D., Pauly, D.  1  DIRECTOR’S FOREWORD The report presented here is ‘only’ a compilation of growth and related parameters for marine, non-fish vertebrates and invertebrates, with only brief analyses, mainly to compare, and thus partly validate, the datasets. One could ask, what is the point? Why such compilation? In the late 1970s, I undertook a similar compilation of growth parameters in fish [Pauly, D. 1978. A preliminary compilation of fish length growth parameters. Berichte des Institut für Meereskunde an der Universität Kiel, No. 55, 200 p.], one of the first of its kind. Its purpose, among other things, was to provide colleagues with a baseline against which to compare progress in the estimation of new growth parameters, whose absence was then perceived as a major impediment to the management of tropical fisheries. Within ten years, this compilation had morphed into the beginnings of FishBase. FishBase at first existed on various diskettes and CD-ROMs, then became an online database of fish - the only such database which not only presents the names and pretty picture of the species it covers, but also important features of their biology, such as, for example, their growth parameters (see www.fishbase.org). The 7 papers of this report, which present growth parameters for marine mammals, seabirds, marine reptiles and many of the invertebrate tribes, however, will not lead to the creation of another database. This is so because a database and website have recently been created for non-fish marine metazoans. It is SeaLifeBase (www.sealifebase.org), which is patterned after FishBase, and which can therefore accommodate, in addition to the names of marine animals, any amount of biological information, notably growth and related parameters (length-weight relationships, size at first maturity, longevity, etc.). Hence this report, in addition to its direct utility to readers, will serve as documentation for a large set of the growth parameters incorporated into SeaLifeBase. These parameters will be of interest to theoreticians, i.e., biologists who want to compare life history strategies in a wide range of taxa, and especially to ecosystem modelers, who need to populate their models with growth parameters and/or parameters derived from these, such as production/biomass ratios, an indication of productivity. This report, therefore, endeavors to cover as wide a range of morphologies and ecological niches as possible. This was particularly successful for the non-fish vertebrates, of which all major groups are represented. For example, in the case of the reptilians, all marine families are represented, and most of the species, except for the very speciose sea snakes (Family Hydrophidae), for which, however, a very good sample of representative species is available. The invertebrates, obviously, are the group for which we have the smallest sample relative to the number of extant species. The compilation that we have here, covering mainly commercial species from the Mediterranean is a good start, however, as are the two papers from the Western Central Pacific, with growth parameters for lobsters and oysters, respectively. Jointly with the growth parameters that were already included in SeaLifeBase, this should provide a starting point for most analyses.  2  Growth of marine mammals, Palomares, M.L.D., et al.  GROWTH OF MARINE MAMMALS 1 M.L. Deng Palomares  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email: m.palomares@fisheries.ubc.ca  Patricia M.E. Sorongon  The SeaLifeBase Project, WorldFish Center, Khush Hall, IRRI, Los Baños, Laguna, Philippines; Email: p.sorongon@cgiar.org Andrea Hunter  Andrea Hunter  Golder Associates Ltd., 2640 Douglas Street, Victoria, BC, Canada V8T 4M1; Email: hunter@zoology.ubc.ca  Daniel Pauly  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email: d.pauly@fisheries.ubc.ca  ABSTRACT Growth and length-weight data were obtained from the literature for 187 populations of 61 species of marine mammals ranging from sea otters to pygmy blue whales. Length-weight parameter estimates yielded a mean b value of 2.86. Estimates of the von Bertalanffy growth function indicate that smaller marine mammals, i.e., seals, sea lions, walruses and dolphins, tend to have growth performance indices between 3.5 to 4.5 and that larger marine mammals, i.e., male fur and elephant seals and whales, tend to have indices higher than 4.5. However, the auximetric plot of log K vs log W∞ shows a decreasing trend in growth performance, similar to that shown for fishes, seabirds and aquatic reptiles.  INTRODUCTION Interest in marine mammals, primarily harvesting and use of products derived from them (e.g., fur/hide, oil and meat), can be traced back to ancient times (Cotté & Guinet, 2007; Allen & Keay, 2006; Christensen, 2006; Tillman & Donovan, 1983). This interest evolved through time, graduating from the need to know of their seasonal whereabouts for obvious reasons connected to the hunt (Christensen, 2006), to a need to know how much fish they consume, i.e., the extent of their competition with fisheries (Kaschner & Pauly, 2005; Kastelein & Vaughan, 1989; Goode, 1884). For some rare species, interest is also growing as to the effect of climate change on their populations (see, e.g., Laidre et al., 2006; Cotté & Guinet, 2007; Newsome et al., 2007). Studying animals living in aquatic environments has always been a challenge because of their inaccessibility to us, their observers. This inconvenience is compounded when the subject are marine mammals, many of which are highly migratory, or which can, on rare occasions, pose a threat to their human observers, as is the case with polar bears. Studying marine mammals is more difficult now that many have become in danger of extinction and, in most parts of the world, are protected species. Traditional life-history studies involve field sampling, and usually sacrificing a subset of the population being studied (see, e.g., True, 1885), or laboratory experiments following the life stages and growth of individual specimens. Nowadays, field sampling of marine mammal populations is done in the context of 1 Cite as: Palomares, M.L.D., Sorongon, P.M.E., Hunter, A., Pauly, D., 2008. Growth of marine mammals. In: Palomares, M.L.D., Pauly, D. (eds.), Von Bertalanffy Growth Parameters of Non-Fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 2-26.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  3  ‘scientific whaling’ (see, e.g., Tamura & Konishi, 2006; Amano & Miyazaki, 2004; Bryden & Harrison, 1986) or when they are caught as by-catch (see, e.g., Miller et al., 1998; Yoshida et al., 1994; Bryden & Harrison, 1986), or stranded. Marine mammal field studies, some dating back to the early 1970s (see, e.g., Stirling, 2002; Burns & Harbo, 1972), employed expensive field observation methods, e.g., helicopter observations or tagging and recapture methods, and were aimed primarily at collecting biogeographical data. There was no known method of determining ages of cetaceans and pinnipeds until the 1950s, and thus, the data required for growth analyses could not be obtained (Gaskin & Blair, 1977). This has changed, however, and length-atage data are available and may be obtained from studies of: bones; GLG’s (growth layer groups) of dentine, e.g., in odontocete cetaceans (Scheffer & Myrick, 1980); weight of eye lenses (Gaskin & Blair, 1977); track width measurements, e.g., in harbor seals (Reijnders 1976); amino acid racemisation (Bada et al., 1980); counts of ovarian corpora albicantia (Kleinenberg & Klevezal, 1962); counts of periosteal bones (van Bree et al., 1986; Brodie, 1969; Kleinenberg & Klevezal, 1962; Laws, 1960); and skeletal and external morphology (Stuart & Morejohn, 1980), e.g., in sea otters (Schneider, 1973). Such data were used to describe the growth of marine mammals using the Gompertz equation (Laird, 1969), logistic equations and, occasionally, the von Bertalanffy growth function (VBGF). This paper assembles growth parameters for marine mammals, estimated using a variety of methods, and standardizes them using the VBGF, along with length-weight relationships. These life-history parameters are available through SeaLifeBase (www.sealifeabase.org), an information system on non-fish marine organisms patterned after the successful model for fish, FishBase (www.fishbase.org). Thus, a preliminary comparison of the growth performance of marine mammal can be presented.  MATERIALS AND METHODS Growth parameter estimation Growth parameters of marine mammal populations were obtained from published literature, and cover the following: (i) the parameters of growth equations other than the VBGF, notably the logistic and Gompertz curves; (ii) age-at-length or growth increment data; and (iii) time series of size frequency distributions. The parameters of the VBGF were recalculated from age-at-length data generated from (i), and all age-at-length and growth increment data were fitted to the VBGF (see von Bertalanffy, 1957) of the form:  Lt = L∞ (1 - e -K·(t-t0))  … (1)  where Lt is the length at age t, L∞ is the asymptotic length, i.e., the mean length the animal would reach if it could grow forever, K is a coefficient of dimension t-1, and t0 is a parameter setting the origin of the curve on the age-axis. Size frequency distributions were fitted to the Powell-Wetherall Plot (PW-Plot; see Pauly, 1998; Wetherall, 1986; Powell, 1979) to estimate L∞, based on the assumption that the resulting distribution is representative of the population. Plotting of successive mean lengths (Lmean), computed from successive cut-off lengths (Li+1), minus the Li (i.e., Lmean- Li) against Li. The downward trend of the points were then fitted with a linear regression of the form Y = a + bX, with L∞ = a/-b) and Z/K = (1+b)/(-b), where Z is the instantaneous rate of total mortality (Pauly, 1998). This method allows the estimation of L∞ and Z/K, i.e., exploited populations, where Z is the instantaneous rate of total mortality. Z/K is equivalent to M/K in unexploited populations. In cases where only L∞ estimates are available, e.g., results of the PW-Plot, values of K were obtained using the growth performance index (Ф’) defined by Pauly & Munro (1984) as Ф’ = log10 K + 2·log10L∞, and mean values of Ф’, available from L∞ and K pairs for: (a) the same species in different localities; (b) other species in the same genus; (c) other species in the same family. Growth parameters obtained through this method are marked as such in SeaLifeBase.  4  Growth of marine mammals, Palomares, M.L.D., et al.  Asymptotic weight estimation Asymptotic weight, W∞, was estimated using the length-weight relationship of the form  W = a · Lb where a is a multiplicative term equivalent to the y-intercept of the loglog transformed linear regression, L the length, and b the exponent, equivalent to the slope of the regression. In many cases, sufficient length-weight data pairs were not available for linear regression analyses. Thus, condition factors (c.f.) using individual lengthweight pairs were estimated with c.f. = W · 100/L3, where W is the weight in grams, and L the length in centimeters (Pauly, 1984). The value of the length-weight parameter a was then obtained as a = c.f./100, assuming that b=3.  … (2) Table 1. Summary of marine mammal species and populations for which data on growth, length-weight relationships (L/W) and condition factors (c.f.) were obtained from the literature. Order Carnivora  Family Mustelidae Odobenidae Otariidae Phocidae Ursidae Balaenidae Balaenopteridae Delphinidae Eschrichtiidae Iniidae Monodontidae Phocoenidae Physeteridae Ziphiidae  Cetacea  Species 1 1 10 16 1 1 8 14 1 1 2 3 1 1  c.f.  L/W -  2 2 17 28 2 2 32 12 4 2 2 4 11 -  8 7 13 14 1 3 3 3 1  VBGF 12 11 24 88 2 1 9 19 1 4 2 11 3 -  RESULTS AND DISCUSSION 18 16 14 12 Frequency  Our literature search, which relied heavily on Internet sources and electronic or ‘soft’ reprints, resulted in 173 length-weight relationships covering 61 species (Table A1), 187 asymptotic size estimates for 47 species and 179 L∞ and K pairs for 46 species (Table A2). Table 1 summarizes the results obtained from this exercise. Note that only two estimates of Z/K were obtained (see Table A2 for values of Z/K calculated through the PowellWetherall Plot for the killer whale, Orcinus orca (Linnaeus, 1758)). The over-representation of phocids and otariids may be due to the fact that their populations remain on- or nearshore and are thus accessible for research. Among cetacean families, delphinids and balaenopterids are best represented. This may be a product of improved ageing techniques, but may also be a by product of whaling and fisheries by-catch. Few data are available for the oceanic Ziphiidae (Baird's beaked whale).  10 8 6 4  Sperm whale  2  Pygmy blue whale  0 2.50  2.75  3.00  3.25  3.50  3.75  4.00  L/W relationship coefficient 'b'  Figure 1. Frequency distribution of the length-weight relationship coefficient b for 53 populations of marine mammals with lengthweight data pairs (see Table A1 for details). Note that the outliers (pygmy blue and sperm whales) were obtained from Lockyer (1976; see Table A2 and text for discussion).  Asymptotic weights using equation (2) were obtained, based on the following criteria: i) species of the same sex, with length-weight and VBGF parameters from the same locality; ii) species from the same body of water; and iii) species with different sex/locality. Details of the methods used in solving for asymptotic weights are indicated in Table A2. Values of the parameter b of the length-weight relationship ranged from 2.31 to 3.97, with 120 estimates computed through condition factors (and the assumption of allometric growth; thus b=3), while 53 were obtained from regression analyses of several length-weight data pairs. Figure 1 shows the distribution of b  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  5  Frequency  values for these 53 populations 80 Seals, sea lions, walruses, (mode at 2.74 and median at purpoises, dolphins 2.86). The outliers at b=3.75 and killer whales 70 and 4.00 were obtained from 60 Lockyer (1976, Table 1), which were based on weight of parts Fur, elephant, crabeater, 50 leopard and Weddell seals and not on whole individuals. Minke and sperm whales Lockyer (1976) notes that fluid 40 losses may account for the high b values and weights calculated 30 Humpback from these L/W relationships. and pygmy blue whales Discounting these outliers, we 20 Humpback get a spread of b values whale 10 between 2.50 and 3.50 with a Ringed seal mean at 2.86. This appears to 0 justify our use of b=3 values to 3.0 3.5 4.0 4.5 5.0 5.5 6.0 6.5 estimate the coefficient a from Ф' = log K + 2* log L∞ (year-1; cm) condition factors for other species for which several L/W data pairs are not available. Figure 2. Frequency distribution of the growth performance index Ф' for 179 Thus, we were able to obtain populations of marine mammals. asymptotic weight values for all of the populations for which asymptotic length values were available (see Table A2). Asymptotic lengths ranged from 110 cm for a female Enhydra lutris (Linnaeus, 1758) (sea otter) from the Aleutian Islands (Alaska) to 2,190 cm for a female Balaenoptera musculus brevicauda (pygmy blue whale) from an unspecified location. The distribution of growth performance indices calculated for these 179 populations (Figure 2) indicated that, in general, seals, sea lions, walruses and dolphins (i.e., smaller 2.0 1.5  -1  K (year ; log10)  1.0  Whales Seals and sea lions  0.5 Dolphins  Fur seal Humpback whale  Sea otter Polar bear  0.0  Elephant seals Gray whale  -0.5 -1.0  Minke whale  -1.5  Weddell seal Walrus  Killer whale  Bowhead whale  Pygmy blue whale  Sperm whale  -2.0 3.0  4.0  5.0  6.0  7.0  8.0  9.0  W∞ (g; log10) Figure 3. Auximetric plot of 179 populations of marine mammals (see Table A2 for details). Note that, in this plot, the growth of killer whales (which are basically large dolphins), and that of fur and elephant seals are similar to that of minke whales. Also note that the growth of polar bears is similar to that of seals and sea lions.  6  Growth of marine mammals, Palomares, M.L.D., et al.  marine mammals), have indices between 3.5 and 4.5, while larger marine mammals tend to have indices higher than 4.5. These indices might be useful as a quick and easy test for the reliability of growth parameter estimates, notably in cases where the age-at-length or frequency distribution data might be biased or based on a small number of samples, not representing the population. Similarly, the auximetric plot of W∞ (log10; g) and K (log10; year-1;) in Figure 3, indicates that: a) sea otters, small species of dolphins, seals, sea lions and polar bears have similar growth patterns, typical of small marine mammals with W∞ ranging from 104 to 105 g; b) there is a medium sized group, i.e., walruses, Weddell seals, fur and elephant seals and killer whales, with W∞ ranging from 105 to 107 g; and c) the group of marine mammals, with W∞ ranging from 107 to 108 g, which include male fur and elephant seals and the great whales. Note that female fur and elephant seals grow in a fashion similar to sea otters, seals and sea lions. Figure 3 also indicates a downward trend in the growth performance of marine mammals, from smaller marine mammals with fast metabolic rates (K values around 3.2 year-1). Overall, we find, as we did previously for fishes (Pauly et al., 2000; Pauly, 1979), and, as we document in this report, for seabirds (Karpouzi & Pauly, 2008) and aquatic reptiles (Dar et al., 2008), that auximetric plots (i.e., plots of logK vs logW∞) can be used to show and interpret patterns in the growth of marine mammals.  ACKNOWLEDGEMENTS This study was made possible by the generous support by the Oak Foundation (Geneva, Switzerland), Dr. Andrew Wright (Vancouver, Canada) and the Pew Charitable Trusts (Philadelphia, USA).  REFERENCES Allen, R.C., Keay, I., 2006. Bowhead whales in the Eastern Arctic, 1611-1911: population reconstruction with historical whaling records. Environment and History 12, 89-113. Amano, M., Miyazaki, N., 2004. Composition of a school of Risso's dolphins, Grampus griseus. Mar. Mamm. Sci. 20(1), 152-160. Arnould, J.P.Y., Warneke, R.M., 2002. Growth and condition in Australian fur seals (Arctocephalus pusillus doriferus) (Carnivora: Pinnipedia). 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A new method for estimating growth and mortality parameters from length-frequency data. Fishbyte 4(1), 12-14. Winship, A.J., Trites, A.W., Calkins, D.G., 2001. Growth in body size of the Steller Sea Lion (Eumetopias jubatus). J. Mammal. 82(2), 500-519. Yoshida, H., Shirakihara, M., Takemura, A., Shirakihara, K., 1994. Development, sexual dimorphism, and individual variation in the skeleton of the finless porpoise, Neophocaena phocaenoides, in the coastal waters of western Kyushu, Japan. Mar. Mamm. Sci. 10(3), 266-282.  10  Growth of marine mammals, Palomares, M.L.D., et al.  APPENDIX Table A1. Summary of 173 populations of 61 species of marine mammals for which length-weight relationships were found (t=tonnes; kg=kilograms; m=meters). Spec. No. 1  Species  Stock  Locality  Method  Sex  b  a  Arctocephalus australis  a  Rio Grande, Brazil  a from cf  F  3.00  0.0488  Fossi et al. (1997; Tab. 1)  Arctocephalus gazelle  b c a  Rio Grande, Brazil San Clemente, Argentina Not specified  a from mean cf a from cf a from cf  M F F  3.00 3.00 3.00  0.0544 0.0385 0.0081  idem idem Trites & Pauly (1998; Tab. 2)  Arctocephalus forsteri  b a  Not specified New Zealand  a from cf a from cf  M F  3.00 3.00  0.00396 0.0191  Idem Dickie & Dawson (2003; p. 177)  Arctocephalus pusillus doriferus  b a  a from cf Recomputed kg  M F  3.00 3.13  0.0216 0.00993  idem Arnould & Warneke (2002; p. 56)  Recomputed from juv./adults, kg a from cf  M  3.30  0.004726  idem  F  3.00  0.00841  Trites & Pauly (1998; Tab. 2)  (South American fur seal) 2  Source  (Antarctic fur seal) 3  (New Zealand fur seal) 4  Arctocephalus tropicalis  a  New Zealand Seal Rocks, Bass Strait, Australia Seal Rocks, Bass Strait, Australia Not specified  Balaena mysticetus  b a  Not specified Not specified  a from cf a from cf  M F  3.00 3.00  0.00508 0.00384  idem Trites & Pauly (1998; Tab 4)  Balaenoptera acutorostrata  b a  Not specified Washington  a from cf a from cf  male F  3.00 3.00  0.00393 0.00927  idem Lockyer (1976; p. 272)  b c d  Unspecified, Antarctic Unspecified, Antarctic Not specified  F M mixed  3.00 3.00 2.31  0.0112 0.0133 1.189  idem idem Lockyer (1976; Tab. 1)  e f  Unspecified, Antarctic Unspecified, Antarctic  unsexed unsexed  3.00 3.23  0.00687 0.00264  Lockyer (1976; p. 272) Lockyer (1976; Tab. 2)  Balaenoptera bonaerensis  a  Southern Ocean  a from mean cf a from mean cf Recomputed from t and m a from mean cf Recomputed from t and m a from cf (pregnant)  F  3.00  0.0115  Tamura & Konishi (2006; Tab. 5)  Balaenoptera musculus brevicauda  b a  Southern Ocean Unspecified, Antarctic  a from cf a from cf  M F  3.00 3.00  0.0115 0.00666  idem Lockyer (1976; p. 269)  b c  Unspecified, Antarctic Unspecified, Antarctic  a from mean cf Recomputed from t and m  M mixed  3.00 3.97  0.00644 0.0000046  idem Lockyer (1976; Tab. 2)  (Australian fur seal)  b 5  (Subantarctic fur seal) 6  (bowhead whale) 7  (minke whale)  8  (Antarctic minke whale) 9  (pygmy blue whale)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  11  Table A1. Continued. Spec. No. 10  Callorhinus ursinus  Species  Stock a  b  a  Locality Sanriku, Japan  Method a from mean cf  Sex F  3.00  0.019  Source Ikemoto et al. (2004; Tab. 1)  Sanriku, Japan Sanriku, Japan Not specified Not specified (pregnant) Not specified Not specified  a from mean cf a from mean cf  Cystophora cristata  b c d e f a  M mixed F F M F  3.00 3.00 2.74 2.67 2.83 3.00  0.0194 0.019 0.0608 0.0979 0.0432 0.0115  Idem Idem Hunter (2005; Tab. A.8) idem idem Trites & Pauly (1998; Tab. 4)  Delphinus delphis  b a  a from cf a from mean cf  M F  3.00 3.00  0.00471 0.0124  idem Kastelein et al. (2000; Tab. 1)  Enhydra lutris  b a  Not specified Hawke Bay, North Island, New Zealand Northeast, USA western Alaska  a from mean cf a from cf  unsexed F  3.00 3.00  0.0119 0.0119  Kastelein et al. (2000; Tab. 3) Estes (1980, p. 2)  Erignathus barbatus  b a  western Alaska Not specified  a from cf a from cf  M F  3.00 3.00  0.0147 0.0107  Idem Trites & Pauly (1998; Tab. 2)  Eschrichtius robustus  b a  Not specified California, USA  a from cf a from mean cf  M F  3.00 3.00  0.0128 0.0107  Idem Lockyer (1976; p. 268)  b c d e  California, USA California, USA Bering Sea Northern Pacific  a from mean cf a from cf a from cf Recomputed from t and m  M unsexed F mixed  3.00 3.00 3.00 3.28  0.00933 0.0108 0.0131 0.0014  Idem Idem Idem Lockyer (1976; Tab. 2)  a  Not specified  F  2.92  0.0332  Hunter (2005; Tab. A.8)  b  Alaska  from kg  F  2.89  0.0363  Idem  c  Alaska (pregnant)  from kg  F  2.79  0.0692  Idem  a  Mediterranean Sea, Italy  from  F  3.00  0.0153  b  Mediterranean Sea, Italy  from  F  3.00  0.0152  Storelli & Marcotrigiano (2000; Tab. 1) Idem  c  Mediterranean Sea, Italy  from  F  3.00  0.0146  Idem  a  Not specified  mixed  2.86  0.0522  Hunter (2005; Tab. A.8)  (northern fur seal)  11  a from cf  (hooded seal) 12  (common dolphin) 13  (sea otter) 14  (bearded seal) 15  (gray whale)  16  Eumetopias jubatus (steller sea lion)  17  Grampus griseus (Risso's dolphin)  18  Halichoerus grypus (grey seal)  Recomputed and m Recomputed and m Recomputed kilograms Recomputed kilograms Recomputed kilograms  12  Growth of marine mammals, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 19  Species  Histriophoca fasciata  Stock a  Locality Not specified  Method a from cf  b  a  Sex F  3.00  0.0104  Source Trites & Pauly (1998; Tab. 4)  (ribbon seal)  Hydrurga leptonyx  b a  Not specified Not specified  a from cf a from cf  M F  3.00 3.00  0.0104 0.0141  Idem Idem  Lagenodelphis hosei  b a  Not specified Not specified  a from cf a from cf  M mixed  3.00 3.00  0.0117 0.00519  Idem Idem  Lagenorhynchus obliquidens  a  Not specified  mixed  2.82  0.035  Hunter (2005; Tab. A.8)  Leptonychotes weddellii  a  Unspecified, Antarctic  mixed  2.53  0.202  Hunter (2005; Tab. A.8)  24  Lobodon carcinophaga  a  Not specified  a from cf  F  3.00  0.0123  Trites & Pauly (1998; Tab. 4)  25  Megaptera noveangliae  b a  Not specified California, USA  a from cf a from mean cf  M F  3 3  0.0112 0.0171  Idem Lockyer (1976; p. 272)  b c  Unspecified, Antarctic Unspecified, Antarctic  F F  3 2.95  0.0103 0.0158  Idem Lockyer (1976; Tab. 2)  d  Puget Sound, Washington, USA Bering Sea Bering Sea Not specified  a from cf Recomputed from t and m a from cf  F  3  0.0104  Lockyer (1976; p. 272)  F M mixed  3 3 2.95  0.0121 0.0129 0.062  Idem Lockyer (1976; p. 272) Lockyer (1976; Tab. 1)  M  3.02  0.0281  Haley et al. (1991; Tab. 1)  F  3  0.0116  Trites & Pauly (1998; Tab. 2)  20  (leopard seal) 21  (Fraser's dolphin) 22  (Pacific white-sided dolphin) 23  (Weddell seal)  (crabeater seal)  (humpback whale)  e f g  a  Año Nuevo State Reserve, California, USA Not specified  a from cf a from cf Recomputed from t and m Recomputed from kg and m a from cf  Monachus schauinslandi  b a  Not specified Not specified  a from cf a from cf  M F  3 3  0.00462 0.0118  Idem Trites & Pauly (1998; Tab. 4)  Monodon monoceros  b a  Not specified Western Greenland  a from cf a from mean cf  M F  3 3  0.0106 0.0161  Idem Garde et al. (2007, p. 57-58)  Neophocaena phocaenoides  b a  Western Greenland Kyushu around Nagasaki and Kanmon Pass, Japan Kyushu around Nagasaki and Kanmon Pass, Japan Not specified  a from mean cf a from mean cf  M F  3 3  0.0168 0.0157  Idem Shirakihara et al. (1993; Tab. 2)  a from mean cf  M  3  0.0144  Shirakihara et al. (1993; Tab. 3)  mixed  3  0.00576  Trites & Pauly (1998; Tab. 4)  Mirounga angustirostris  a  27  Mirounga leonine  28  26  (northern elephant seal) (southern elephant seal) (Hawaiian monk seal) 29  (narwhal) 30  (finless porpoise)  b c  a from cf  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  13  Table A1. Continued. Spec. No. 31  Species  Odobenus rosmarus  Stock a  Locality Not specified  Method a from cf a from cf  Sex F  b  a  3  0.0175  Source Trites & Pauly (1998; Tab. 2)  M mixed  3 3.2  0.0143 0.006  Idem Hunter (2005; Tab. A.8)  mixed F  2.58 3  0.208 0.0113  Idem Trites & Pauly (1998; Tab. 4)  3 2.81  0.00469 0.0645  Idem Hunter (2005; Tab. A.8)  3  0.0095  Trites & Pauly (1998; Tab. 4)  (walrus) 32  Orcinus orca  b a  Not specified Not specified  Otaria flavescens  b a  Not specified Not specified  Pagophilus groenlandicus  b a  Not specified Not specified  a from cf  M mixed  Phoca largha  a  Not specified  a from cf  F  Phoca vitulina  b a  Not specified Not specified  a from cf  M mixed  3 2.89  0.0102 0.0404  Idem Hunter (2005; Tab. A.8)  Phocoena phocoena  a  Not specified  F  2.43  0.216  Idem  Phocoenoides dalli  b c a  Not specified Not specified Not specified  a from cf  M mixed mixed  2.74 2.63 3  0.051 0.083 0.00576  Idem Hunter (2005; Tab. A.8) Trites & Pauly (1998; Tab. 4)  Physeter macrocephalus  a  Japan  a from mean cf  3  0.00893  Lockyer (1976; p. 273)  b c  Japan Japan  M mixed  3 3.18  0.00964 0.0029  Lockyer (1976; p. 272-273) Lockyer (1976; Tab. 1)  d e  Natal, South Africa Natal, South Africa  F F  3 3.55  0.0131 0.00023  Lockyer (1976; p. 273) Lockyer (1976; Tab. 2)  f g h i j k  Natal, South Africa Bering Sea Bering Sea Iceland Canada Antarctic and Pacific  M M unsexed M M mixed  3 3 3 3 3 2.74  0.0131 0.00918 0.00797 0.00997 0.0139 0.0649  Lockyer (1976; p. 273) Idem Idem Idem Idem Lockyer (1976; Tab. 2)  l m n a  Unspecified, Antarctic Not specified Not specified Not specified  a from mean cf Recomputed from t and m a from mean cf Recomputed from t and m a from cf a from mean cf a from mean cf a from cf a from cf Recomputed from t and m a from mean cf a from cf a from cf a from cf  unsexed F M F  3 3 3 3  0.0109 0.00584 0.00462 0.00626  Lockyer (1976; p. 273) Trites & Pauly (1998; Tab. 2) Idem Idem  b  Not specified  a from cf  M  3  0.00635  Idem  (killer whale) 33  a from cf  (South American sea lion) 34  (harp seal) 35  (larga seal) 36  (Harbour seal) 37  (harbour porpoise) 38  (Dall's porpoise) 39  F  (sperm whale)  40  Pontoporia blainvillei  (franciscana dolphin)  14  Growth of marine mammals, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 41  Species  Locality Caspian Sea  Method a from cf  Stock a b c d e  Caspian Sea Caspian Sea northern Caspian Sea northern Caspian Sea  f  northern Caspian Sea  g  northern Caspian Sea  a from cf a from cf a from mean a from mean (pregnant) a from mean pregnant) a from mean  a  Svalbard  b  Svalbard  c  Kongsfjorden, Svalbard  d  Kongsfjorden, Svalbard  Pusa sibirica  a  Lake Baikal  Recomputed from kg and m Recomputed from kg and m Recomputed from kilograms Recomputed from kilograms a from mean cf  Stenella frontalis  b c a  Lake Baikal Lake Baikal Not specified  a from mean cf a from mean cf a from cf  Steno bredanensis  b a  Not specified Not specified  Tursiops truncates  b a  Ursus maritimus  b  a  Sex F  3  0.0341  Source Ikemoto et al. (2004; Tab. 1)  M mixed F F  3 3 3 3  0.0285 0.033 0.031 0.0362  Idem Idem Watanabe et al. (2002;Tab. 1) Idem  cf (non-  F  3  0.027  Idem  cf  M  3  0.0327  Idem  F  3.15  0.0145  Hunter (2005; Tab. A.8)  M  3.26  0.00832  Idem  F  3  0.0257  Krafft et al. (2007; Tab. 2)  male  3  0.0350  Idem  F  3  0.0248  Ikemoto et al. (2004; Tab. 1)  M mixed F  3 3 3  0.021 0.023 0.00562  Idem Idem Trites & Pauly (1998; Tab. 4)  a from cf a from cf  M F  3 3  0.00567 0.00529  Idem Idem  Not specified Not specified  a from cf a from cf  M F  3 3  0.00518 0.00348  Idem Trites & Pauly (1998; Tab. 2)  b a  Not specified Svalbard  a from cf a from cf  M F  3 3  0.00367 0.0253  Idem Derocher & Wiig (2002; Tab. 1)  Arctocephalus pusillus  b a  Svalbard Not specified  a from cf a from cf  M F  3 3  0.0342 0.0101  Idem Trites & Pauly (1998; Tab. 4)  Arctocephalus townsendi  b a  Not specified Guadalupe, Mexico  a from cf a from mean cf  M F  3 3  0.00444 0.0151  Idem Gallo-Reynoso et al. (1996; Table 1)  Pusa caspica  (Caspian seal)  42  Pusa hispida  (ringed seal)  43  cf cf  (Baikal seal) 44  (Atlantic spotted dolphin) 45  (rough-toothed dolphin) 46  (bottlenose dolphin) 47  (polar bear) 48  (South African fur seal) 49  (Guadalupe fur seal)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  15  Table A1. Continued. Spec. No. 50  Species  Balaenoptera borealis  Stock a  Japan  Locality  b c  Japan Japan  d  Japan  e f a  Natal, South Africa Unspecified, Antarctic Japan  b c  Japan Japan  d  Japan  a  Unspecified, Antarctic  b c  Unspecified, Antarctic Unspecified, Antarctic  d e  Unspecified, Antarctic Not specified  f a  Newfoundland, Canada Unspecified, Antarctic  b c  Unspecified, Antarctic Unspecified, Antarctic  d e f g h i j k  California, USA Korf Bay, Kamchatka, Russia Natal'ya Bay, Russia Far East Far East Iceland Commander Island, Russia Not specified  a  Japan  Method a from mean cf  Sex F  b  a  3  0.00559  Source Lockyer (1976; p. 271)  M mixed  3 2.43  0.00617 0.356  Lockyer (1976; p. 270) Lockyer (1976; Tab. 1)  unsexed  2.43  0.334  Lockyer (1976; Tab. 2)  3 3 3  0.00856 0.00639 0.00622  Lockyer (1976; p. 271) Idem Idem  M mixed  3 2.74  0.00623 0.0429  Idem Lockyer (1976; Tab. 1)  unsexed  2.74  0.0404  Lockyer (1976; Tab. 2)  3  0.00612  Lockyer (1976; p. 269)  M mixed  3 3.09  0.00636 0.00304  Lockyer (1976; p. 268-269) Lockyer (1976; Tab. 2)  unsexed mixed  3 3.25  0.00593 0.000917  Lockyer (1976; p. 269) Lockyer (1976; Tab. 1)  unsexed F  3 3  0.00473 0.00554  Lockyer (1976; p. 269) Lockyer (1976; p. 270)  a from mean cf Recomputed from t and m a from cf a from cf  M unsexed  3 2.53  0.0056 0.207  Lockyer (1976; p. 270) Lockyer (1976; Tab. 2)  3 3  0.00581 0.00598  Lockyer (1976; p. 270) Idem  a from cf a from cf a from cf a from mean cf a from cf Recomputed from t and m  F F M M M mixed  3 3 3 3 3 2.9  0.00617 0.00619 0.00583 0.00573 0.00504 0.0127  Idem Idem Lockyer (1976; p. 269) Idem Idem Lockyer (1976; Tab. 1)  mixed  3.08  0.00634  Hunter (2005; Tab. A.8)  (sei whale)  51  Balaenoptera brydei  a from mean cf Recomputed from t and m Recomputed from t and m a from cf a from cf a from mean cf  F M F  (Bryde's whale)  52  Balaenoptera musculus  a from mean cf Recomputed from t and m Recomputed from t and m a from mean cf  F  (blue whale)  53  Balaenoptera physalus  a from mean cf Recomputed from t and m a from mean cf Recomputed from t and m a from cf a from mean cf  (fin whale)  54  Berardius bairdii  (Baird's beaked whale)  F F  16  Growth of marine mammals, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 55  Species  Cephalorhynchus hectori  Stock a  Locality Not specified  Method  b  a  Sex mixed  2.53  0.1689  Idem  Source  (Hector's dolphin) 56  Delphinapterus leucas  a  St. Lawrence, Canada  mixed  2.61  0.156  Idem  Globicephala melas  b c a  Hudson Bay, Canada Hudson Bay, Canada Faeroe Island (postnatal)  mixed mixed mixed  2.56 2.54 2.5  0.182 0.452 0.23  Idem Idem Idem  Pseudorca crassidens  a  Not specified  mixed  2.44  0.216  Idem  Stenella attenuate  a  Not specified  F  2.61  0.0696  Idem  Stenella coeruleoalba  b c a  Not specified Not specified Not specified (postnatal)  M mixed F  2.87 2.93 2.91  0.0193 0.0126 0.0183  Idem Idem Idem  Stenella longirostris  b c a  Not specified (postnatal) Not specified Not specified  M mixed F  2.98 2.93 2.61  0.0139 0.0171 0.0696  Idem Idem Idem  b  Not specified  M  2.87  0.0193  Idem  (white whale) 57  (long-finned pilot whale) 58  (false killer whale) 59  (Pantropical spotted dolphin) 60  (striped dolphin) 61  (long-snouted spinner dolphin)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  17  Table A2. Summary of 179 populations of 47 marine mammal species for which von Bertalanffy growth parameters were found. Spec. No. 1 2 3  4  5 6 7 8  9  10  Species  Arctocephalus australis  (South American fur seal)  Arctocephalus gazelle (Antarctic fur seal)  Arctocephalus forsteri  (New Zealand fur seal)  Arctocephalus pusillus doriferus (Australian fur seal)  Arctocephalus tropicalis (Subantarctic fur seal) Balaena mysticetus (bowhead whale)  Balaenoptera acutorostrata (minke whale)  Balaenoptera bonaerensis (Antarctic minke whale)  Balaenoptera musculus brevicauda (pygmy blue whale)  Callorhinus ursinus (northern fur seal)  N  W∞  K  to  Locality  a  Isla de Lobos, Uruguay  253  F  118  72  0.30  -0.67  a  Not specified  -  F  220  87  0.44  -0.68  b  Idem  -  M  331  143  0.13  -0.66  a  New Zealand  F  119  32  0.41  -  b  Kangaroo Island, South Australia  -  F  137  50  0.33  -1.55  c  Idem  -  M  184  135  0.17  -8.18  a  Seal Rocks, Bass Strait, Australia  163  F  163  84  0.36  -1.91  b  Idem  69  M  600  7072  0.30  -0.88  a  Amsterdam Island, southern Indian Ocean  108  F  139  23  0.62  -  a  Alaska  -  unsexed  1602  16000  0.032  -22.2  a  Not specified  -  M  833  7688  0.17  -4.30  a  Idem  -  F  907  8581  0.14  -4.30  b  Idem  -  ♂  833  6647  0.17  -4.30  a  Idem  170  F  2190  70000  0.08  -16.2  b  Idem  218  M  2110  60500  0.09  -15.5  a  Eastern Bering Sea, California  6493  F  128  36  0.31  -2.06  b  Idem  9630  F  130  42  0.19  -7.32  57  Sex  L∞  Stock  (cm)  (year-1)  (kg)  (year)  Comments/Source Length-at-age; 0-28.5 years. Average W∞ from Tab. 1 (1a, 1c). Lima & Paez (1995; Fig. 1). From generalized VBGF. W∞ from Tab. 1(2b). McLaren (1993; Tab. 1). Idem W∞ from Tab. 1 (3a). Dickie & Dawson (2003; Tab. 1). W∞ from Tab. 1(3a). McKenzie et al. (2007; Tab. 2). W∞ from Tab. 1 (3b). McKenzie et al. (2007; Tab. 2). W∞ from Tab. 1 (4a). Arnould & Warneke (2002; Tab. 1) From logistic curve. W∞ from Tab. 1(4b). Arnould & Warneke (2002, Abstract); Hunter (2005; Tab. A.8). From Gompertz equation. W∞ from Tab. 1 (5a). Dabin et al. (2004; p. 1045). Average W∞ from Tab. 1 (6a, 6b). George et al. (1999; p. 575) W∞ from Tab. 1 (7c). Hunter (2005, Tab. A.8). W∞ from Tab. 1 (8a). Hunter (2005, Tab. A.8). W∞ from Tab. 1(8b). Hunter (2005, Tab. A.8). From m to cm. W∞ from Tab. 1 (9a). Branch (2008, Tab. 3). From m to cm. W∞ from Tab. 1 (9b). Branch (2008, Tab. 3). Length at age; non-pregnant females; 0-15 years.Average W∞ from Tab. 1 (10a, 10d-e). Trites & Bigg (1996; Tab. 1). Length at age; pregnant females; 4-23 years. Average W∞ from Tab. 1 (10a, 10d-e). Trites & Bigg (1996; Tab. 1).  18  Growth of marine mammals, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No. 10  11  12 13  Species  Callorhinus ursinus (northern fur seal)  Cystophora cristata (hooded seal)  Delphinus delphis  (common dolphin)  Enhydra lutris (sea otter)  Stock  Locality  L∞  W∞  K  to  N  Sex  2008  M  266  303  0.08  -3.69  (cm)  (kg)  (year-1)  (year)  c  Idem  d  Pribilof Island, Alaska  137  F  127  39  0.38  -1.83  e  Idem  306  M  308  818  0.08  -3.13  f  Not specified  -  F  198  124  0.26  -0.67  g  Idem  -  M  396  942  0.03  -0.42  a  Idem  -  F  280  252  0.20  -0.62  b  Idem  -  M  311  141  0.16  -0.61  a  Hawke Bay, North Island, New Zealand  F  196  93  0.20  -6.99  a  Not specified  -  F  148  39  0.20  -  b  Idem  -  M  148  48  0.22  -  c  Western Aleutian Islands, Alaska  102  F  118  20  2.49  -0.22  d  Idem  90  M  117  24  2.63  -0.21  4  -  F  110  16  0.53  -2.35  f  Aleutian Islands, Alaska Idem  -  F  123  22  0.82  -1.55  g  Idem  -  M  119  25  0.38  -2.51  h  Idem  -  M  132  33  0.61  -2.05  i  California, USA  -  F  128  25  -  -  j  Idem  -  F  127  24  -  -  e  Comments/Source Length at age; 0-16 years. W∞ from Tab. (10b, 10f). Trites & Bigg (1996; Tab. 1). Length at age; 0-10 years. Average W∞ from Tab. 1 (10a, 10d-e). Scheffer & Wilke (1953; Tabs. 1-2). Length at age; 0-10 years. Average W∞ from Tab. 1 (10b, 10f). Scheffer & Wilke (1953; Tabs. 1-2). From generalized VBGF. Average W∞ from Tab. 1 (10d-e). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (10f). McLaren (1993; Tab. 1) From generalized VBGF. W∞ from Tab. 1 (11a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (11b). McLaren (1993; Tab. 1). Length at age; 2-27 years. W∞ from Tab. 1 (12a). Kastelein et al. (2000; Fig. 3). L∞ from Lmax; K from theta of female pups (13c). W∞ from Tab. 1 (13a). Jefferson et al. (1993). L∞ from maximum length; K from theta of female pups (13c). W∞ from Tab. 1 (13b). Jefferson et al. (1993). Length at age; female pups; 0-3 years. W∞ from Tab. 1 (13a). Schneider (1973; Tab. 3). Length at age; male pups; 0-3 years. W∞ from Tab. 1 (13b). Schneider (1973; Tab. 3). W∞ from Tab. 1 (13a). Laidre et al. (2006; Tab. 2). Idem W∞ from Tab. 1 (13b). Laidre et al. (2006; Tab. 2). Idem W∞ from Tab. 1 (13a). Laidre et al. (2006; p. 985). Idem  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  19  Table A2. Continued. Spec. No. 13  14  15 16  17 18  19  Species  Enhydra lutris (sea otter)  Erignathus barbatus (bearded seal)  Eschrichtius robustus (gray whale)  Eumetopias jubatus (steller sea lion)  Grampus griseus (Risso's dolphin)  Halichoerus grypus (grey seal)  Histriophoca fasciata (ribbon seal)  Stock  Locality  N  Sex  L∞  (cm)  W∞  (kg)  K  to  (year-1)  (year)  k  Idem  -  M  119  25  -  -  l  Idem  -  M  118  24  -  -  a  Barents Sea  -  mixed  306  338  0.21  -0.70  b c d  Sea of Okhotsk Bering-Chukchi Sea Eastern Canada California and Washington, USA  -  mixed mixed mixed  271 300 326  233 319 516  0.29 0.25 0.18  -0.74 -0.71 -0.73  -  F  1297  23346  0.25  -2.84  a a  Gulf of Alaska  -  F  360  913  0.34  -0.65  b c d  Idem Shelikof Alaska Idem  -  M F M  486 304 454  2137 567 1766  0.17 0.20 0.17  -0.65 -0.66 -0.64  e  Alaska  201  F  230  255  0.54  -1.05  f  Idem  235  M  307  579  0.26  -1.50  a  Taiji, Japan  -  F  271  298  0.49  -2.09  b  Idem  -  M  273  305  0.57  -1.62  a  Eastern Canada  -  F  271  475  0.18  -0.60  b c d  Idem Farne Islands, England Idem  -  M F M  328 241 290  821 338 573  0.14 0.18 0.16  -0.58 -0.53 -0.54  a  Sea of Okhotsk  -  F  245  153  0.47  -0.62  b  Idem  -  M  261  185  0.57  -0.62  c  Idem  -  mixed  254  17  0.52  -0.64  d  Bering Sea  -  F  242  148  0.37  -0.63  e  Idem  -  M  262  187  0.46  -0.64  Comments/Source W∞ from Tab. 1 (13b). Laidre et al. (2006; p. 985). Idem From generalized VBGF. Average W∞ from Tab. 1 (14a-b). McLaren (1993; Tab. 1). Idem Idem Idem W∞ from Tab. 1 (15a). Kastelle et al. (2003; p. 26). From generalized VBGF. Average W∞ from Tab. 1 (16b-c). McLaren (1993; Tab. 1). Idem Idem Idem Length at age; 0-24 years. Average W∞ from Tab. 1 (16b-c). Winship et al. (2001; Tab. 3). Length at age; 0-18 years. Average W∞ from Tab. 1 (16b-c). Winship et al. (2001; Tab. 3). Average W∞ from Tab. 1 (17a-c). Amano & Miyazaki (2004; Fig. 2). Idem From generalized VBGF. W∞ from Tab. 1 (18a). McLaren (1993; Tab. 1). Idem Idem Idem From generalized VBGF. W∞ from Tab. 1 (19a). McLaren (1993; Tab. 1). Idem From generalized VBGF. Average W∞ from Tab. 1 (19a-b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (19a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (19b). McLaren (1993; Tab. 1).  20  Growth of marine mammals, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No. 19 20  21 22  23  Species  Histriophoca fasciata (ribbon seal)  Hydrurga leptonyx (leopard seal)  Lagenodelphis hosei (Fraser's dolphin)  Lagenorhynchus obliquidens (Pacific white-sided dolphin)  Leptonychotes weddellii (Weddell seal)  Stock  25  26  Lobodon carcinophaga (crabeater seal)  Megaptera novaeangliae (humpback whale)  Mirounga angustirostris  (northern elephant seal)  N  Sex  L∞  (cm)  W∞  (kg)  K  (year-1)  to  Idem  -  mixed  253  168  0.42  -0.63  a  Antarctic  -  F  539  221  0.36  -0.69  b  Idem  -  M  497  1434  0.47  -0.69  a  Southeast Brazil  mixed  236  69  0.48  -1.05  a  North Pacific  F  186  88  0.71  -1.29  11 -  Comments/Source  (year)  f  b  Idem  -  M  195  100  0.38  -2.06  c  Idem  -  mixed  191  95  0.46  -1.75  a  South Orkney Island  -  F  558  1795  0.62  -0.73  From generalized VBGF. Average W∞ from Tab. 1 (19a-b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (20a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (20b). McLaren (1993; Tab. 1). Length at age; 0-19 years. W∞ from Tab. 1 (21a). Siciliano et al. (2007; Tab. 6). W∞ from Tab. 1 (22a). Heise (1997; Tab. 2). Idem W∞ from Tab. 1 (22a). Hunter (2005; Tab. A8). From generalized VBGF. W∞ from Tab. 1 (23a). McLaren (1993; Tab. 1).  -  F  399  770  0.37  -0.73  Idem  c d e f g  McMurdo Sound, Antarctica Idem Idem Idem Idem Idem  -  F M M mixed mixed  394 410 382 396 383  743 824 687 756 692  0.21 0.46 0.30 0.38 0.27  -0.74 -0.73 -0.73 -0.72 -0.74  a  Not specified  -  F  393  747  0.66  -0.73  b  Idem  -  M  389  659  0.61  -0.74  c  Idem  -  mixed  391  702  0.64  -0.72  a  Northwest Atlantic  -  mixed  1050  51000  1.96  -0.26  b  Idem  -  mixed  1145  65410  0.98  -0.46  c  Northern Atlantic  11  F  1394  33000  0.25  -3.18  d  Idem  12  M  1124  18327  0.84  -1.00  a  Not specified  -  F  492  3851  0.15  -0.67  b  Idem  -  M  911  250000  0.16  -0.68  Idem Idem Idem Idem Idem From generalized VBGF. W∞ from Tab. 1 (24a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (24b). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (24a-b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (25g). Stevick (1999; Fig. 4). Idem Length at age; not a good fit. Average W∞ from Tab. 1 (25a, 25f). Stevick (1999; Tab. 1). Length at age; not a good fit. W∞ from Tab. 1 (25f). Stevick (1999; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (26a). McLaren (1993; Tab. 1). Idem  b  24  Locality  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  21  Table A2. Continued. Spec. No. 27  28 29  30  31  31  32  Species  Mirounga leonine  (southern elephant seal)  Monachus schauinslandi (Hawaiian monk seal)  Monodon monoceros (narwhal)  Neophocaena phocaenoides (finless porpoise)  Odobenus rosmarus (walrus)  Odobenus rosmarus (walrus)  Orcinus orca  (killer whale)  Stock  Locality  N  Sex  a  Macquarie Island  -  F  b  South Georgia  -  c  Idem  a  Not specified  a  West Greenland  b  L∞  (cm)  W∞  (kg)  K  (year-1)  to  (year)  410  802  0.18  -0.68  F  471  12147  0.27  -0.67  -  M  1444  14000  0.17  -0.68  -  mixed  354  497  0.15  -0.73  24  F  396  1000  -  -  Idem  38  M  457  1603  -  -  a  Kyushu, Japan  46  F  148  51  0.74  -1.00  b  Idem  51  M  150  48  0.71  -1.00  a  Foxe Basin, Northwest Territories, Canada  90  F  275  364  0.31  -1.86  b  Idem  103  M  312  433  0.20  -2.71  c  Foxe Basin, Nunavut, Canada  -  M  576  2735  0.25  -0.86  d  Hudson Bay, Canada  -  F  402  1137  0.26  -0.87  e  Idem  -  M  432  1153  0.12  -0.87  f  Unspecified, Alaska  -  F  422  1311  0.22  -0.87  g  Idem  -  M  470  1481  0.10  -0.87  h  Unspecified, Russia  -  F  475  1879  0.16  -0.88  i  Idem  -  M  552  2411  0.10  -0.87  j  Northwest Greenland  34  F  269  341  -  -  k  Idem  54  M  314  443  -  -  a  Norway, coastal waters  173  F  564  3196  0.17  -4.17  b  Idem  143  M  650  4854  0.10  -5.81  Comments/Source From generalized VBGF. W∞ from Tab. 1 (27a). McLaren (1993; Tab. 1). Idem From generalized VBGF. W∞ from Tab. 1 (27b). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (28a-b). McLaren (1993; Tab. 1). W∞ from Tab. 1 (29a). Garde et al. (2007; p. 52). W∞ from Tab. 1 (29b). Garde et al. (2007; p. 52). Length at age. W∞ from Tab. 1 (30a). Shirakihara et al. (1993; Tab. 1). Length at age. W∞ from Tab. 1 (30b). Shirakihara et al. (1993; Tab. 1). W∞ from Tab. 1 (31a). Garlich-Miller & Stewart (1998; Tab. 1). W∞ from Tab. 1 (31b). Garlich-Miller & Stewart (1998; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (31b). McLaren (1993; Tab. 1). W∞ from Tab. 1 (31a). Knutsen & Born (1994). W∞ from Tab. 1 (31b). Knutsen & Born (1994). Length at age. Average W∞ from Tab. 1 (32a-b). Christensen (1984; Fig. 4). Idem  22  Growth of marine mammals, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No. 32  33  34 35 35  36  Species  Orcinus orca  (killer whale)  Otaria flavescens  (South American sea lion)  Pagophilus groenlandicus (harp seal)  Phoca largha (larga seal)  Phoca largha (larga seal)  Phoca vitulina  (Harbour seal)  Stock  Locality  N  Sex  L∞  (cm)  W∞  (kg)  K  (year-1)  to  (year)  c  British Columbia and Washington  27  F  618  4180  0.15  -  d  Idem  29  M  704  6151  0.12  -  e  Holland, Netherlands  1  F  618  4180  0.15  -  a  Southern Brazil  32  F  194  83  0.31  -2.00  b  Idem  94  M  254  77  0.30  -1.60  a  Not specified  -  mixed  240  315  0.31  -0.57  a  Bering-Okhotsk Sea  -  F  225  109  0.36  -0.56  b  Idem  -  M  246  152  0.44  -0.53  c  Hokkaido, Japan  -  F  209  87  0.19  -0.57  d  Idem  -  M  216  103  0.16  -0.55  b c d  Commander, Aleutian and Pribilof Islands Idem Norway Idem  e  Gulf of Alaska  f g h i j k l m n  Idem Idem Idem Aleutian, Alaska Idem Denmark/Sweden Idem Nova Scotia, Canada Idem  a  -  F  167  107  0.20  -4.49  -  M F M  175 210 226  123 207 256  0.23 0.24 0.22  -3.80 -0.63 -0.65  -  F  203  189  0.22  -0.64  -  F M M F M F M F M  150 162 226 218 245 207 228 223 249  78 98 257 231 323 200 263 247 340  0.31 0.30 0.22 0.09 0.17 0.26 0.26 0.36 0.40  -3.03 -2.76 -0.62 -0.66 -0.65 -0.62 -0.63 -0.63 -0.63  Comments/Source L∞ from Powell-Wetherall Plot; K from theta (32e). Z/K=0.628. Average W∞ from Tab. 1 (32a-b). Bigg & Wolman (1975). L∞ from Powell-Wetherall Plot; K from theta (32e). Z/K=1.05. Average W∞ from Tab. 1 (32a-b). Bigg & Wolman (1975). Growth increments; Gulland and Holt Plot; 1-12 years. Average W∞ from Tab. 1 (32a-b). Kastelein & Vaughan (1989; Tab. 1). W∞ from Tab. 1 (33a). Rosas et al. (1993; p. 141, 143). W∞ from Tab. 1 (33b). Rosas et al. (1993; p. 141, 143). From generalized VBGF. W∞ from Tab. 1 (34a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (35a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (35b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (35a). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (35b). McLaren (1993; Tab. 1). From generalized VBGF. W∞ from Tab. 1 (36a). McLaren (1993; Fig. 40). Idem Idem Idem From generalized VBGF. W∞ from Tab. 1 (36a). McLaren (1993; Fig. 39). Idem Idem Idem Idem Idem Idem Idem Idem Idem  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  23  Table A2. Continued. Spec. No. 36  37  38  39  40  41  Species  Phoca vitulina  (Harbour seal)  Phocoena phocoena (Harbour porpoise)  Phocoenoides dalli (Dall’s porpoise)  Physeter macrocephalus (Sperm whale)  Pontoporia blainvillei  (Franciscana dolphin)  Pusa caspica (Caspian seal)  Stock  L∞  W∞  K  to  Locality  N  Sex  o  British Columbia  -  F  217  227  0.23  -0.64  p q r s t  Idem Idem Idem Hokkaido, Japan Idem  -  F F M F M  236 151 167 224 250  292 79 108 249 345  0.12 0.37 0.24 0.22 0.16  -0.65 -2.52 -3.69 -0.62 -0.64  a  Sea of Azov  45  F  145  39  0.76  -  b  Idem  53  M  132  32  0.91  -  c  Black Sea  41  F  132  32  0.71  -  d  Idem  48  M  123  26  1.21  -  e  Western Greenland  -  F  155  46  0.48  -  f  Idem  -  M  143  40  0.46  -  -  F  186  37  0.58  -1.39  -  F M  188 192  38 41  0.40 0.50  -2.78 -1.60  (cm)  (year-1)  (kg)  (year)  Comments/Source From generalized VBGF. W∞ from Tab. 1 (36a). McLaren (1993; Fig. 38). Idem Idem Idem Idem Idem W∞ from Tab. 1(37a). Gol’din (2004; Tab. 1). W∞ from Tab. 1(37b). Gol’din (2004; Tab. 1). W∞ from Tab. 1(37a). Gol’din (2004; Tab. 1). W∞ from Tab. 1(37b). Gol’din (2004; Tab. 1). W∞ from Tab. 1(37a). Lockyer et al. (2001; Tab. 3). W∞ from Tab. 1(37b). Lockyer et al. (2001; Tab. 3). Length-at-age. W∞ from Tab. 1 (38a). Ferrero & Walker (1999; Figs. 8-9). Idem Idem Average W∞ from Tab. 1 (39d-e). Evans et al. (2004; p. 248). Average W∞ from Tab. 1 (39d-e). Bannister (1969). Length-at-age. Average W∞ from Tab. 1 (39a-n). Lockyer (1981; Abstract).  b c  Western Aleutian Islands Idem Idem  a  Tasmania, Australia  -  F  1082  15100  0.16  -2.58  b  Western Australia  -  mixed  1052  14300  0.12  -4.12  c  Not specified  -  M  1858  65100  0.05  -5.37  a  Paraná and Sao Paulo (25°00' - 25°58'S), Brazil  18  F  129  13  0.33  -3.07  W∞ from Tab. 1 (40a). Barreto & Rosas (2006; Tab. 3).  b  Idem  23  M  113  9  1.00  -0.90  W∞ from Tab. 1 (40b). Barreto & Rosas (2006; Tab. 3).  c  Rio Grande do Sul (29°20' - 33°45'S), Brazil  48  F  146  20  0.57  -1.71  W∞ from Tab. 1 (40a). Barreto & Rosas (2006; Tab. 3).  d  Idem  59  M  130  14  1.14  -0.71  a  Not specified  F  185  202  0.25  -0.61  a  W∞ from Tab. 1 (40b). Barreto & Rosas (2006; Tab. 3). From generalized VBGF. Average W∞ from Tab. 1 (41a, 41d-f). McLaren (1993; Tab. 1).  24  Growth of marine mammals, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No. 42  Species  Pusa hispida (ringed seal)  Stock  Locality  a  Sea of Okhotsk  aa  High Canada, Arctic  b  Sea of Okhotsk  c  Idem  d  N  Sex  L∞  (cm)  W∞  (kg)  K  (year-1)  to  (year)  F  161  118  0.11  -0.62  mixed  181  184  0.1  -0.61  M  164  146  0.15  -0.61  mixed  162  130  0.12  -0.63  Chukchi Sea  F  172  144  0.27  -0.58  e  Idem  M  167  154  0.21  -0.6  f  Idem  mixed  169  149  0.24  -0.61  g  Baltic Sea  F  198  222  0.23  -0.63  h  Baltic Sea  M  205  294  0.23  -0.62  i  Idem  mixed  204  265  0.25  -0.61  j  Barents Sea  F  178  160  0.22  -0.62  k  Idem  M  186  215  0.29  -0.61  l  Idem  mixed  181  185  0.26  -0.63  F  180  167  0.17  -0.6  m  Bering Sea  Comments/Source From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1) From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1) From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1) From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1) From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1) From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  25  Table A2. Continued. Spec. No. 42  43 44 45  Species  Pusa hispida (ringed seal)  Pusa sibirica (Baikal seal)  Stenella frontalis  (Atlantic spotted dolphin)  Steno bredanensis  (rough-toothed dolphin)  Stock  Locality  N  Sex  L∞  (cm)  W∞  (kg)  K  (year-1)  to  Comments/Source  (year)  n  Idem  M  184  210  0.08  -0.61  o  Idem  mixed  180  183  0.11  -0.56  p  Svalbard  F  166  129  0.15  -0.58  q  Idem  144  F  130  61  0.17  r  Idem  102  F  128  58  0.18  s  Idem  131  M  130  70  0.34  t  Idem  170  M  128  67  0.43  u  Idem  M  186  216  0.31  -0.62  v  Idem  mixed  172  157  0.22  -0.6  w  Western Canada, Arctic  F  160  116  0.14  -0.65  x  Western Canada, Arctic  M  169  161  0.13  -0.67  y  Idem  mixed  164  136  0.15  -0.68  z  Southeast Canada, Arctic  mixed  154  112  0.1  -0.59  a  Not specified  F  178  140  0.42  -0.62  a  Southeast Brazil  27  mixed  225  64  0.14  -5.56  a  Idem  13  mixed  259  91  0.32  -2.97  From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1) Average W∞ from Tab. 1 (42a, 42c). Krafft et al. (2006; Tab 1). Idem Average W∞ from Tab. 1 (42b, 42d). Krafft et al. (2006; Tab 1). Idem From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a, 42c). McLaren (1993; Tab. 1) From generalized VBGF. Average W∞ from Tab. 1 (42b, 42d). McLaren (1993; Tab. 1). From generalized VBGF. Average W∞ from Tab. 1 (42a-d). McLaren (1993; Tab. 1). Idem From generalized VBGF. Average W∞ from Tab. 1 (43a). McLaren (1993; Tab. 1). Length at age; 0-23 years. Average W∞ from Tab. 1 (44a-b). Siciliano et al. (2007; Tab. 1). Length at age; 0.5-24 years. Average W∞ from Tab. 1 (45a-b). Siciliano et al. (2007; Tab. 5).  26  Growth of marine mammals, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No. 46  47  Species  Tursiops truncates  (bottlenose dolphin)  Ursus maritimus (polar bear)  Stock  Locality  a  Idem  b  N 21  Sex  L∞  (cm)  W∞  K  (year-1)  (kg)  to  (year)  mixed  305  101  0.14  -6.24  North-Central Gulf of Mexico  F  242  49  0.48  -1.19  c  Idem  M  253  59  0.36  -1.77  d  Indian River Lagoon, Florida, USA  72  F  114  5  0.45  e  Idem  118  M  124  7  0.36  -0.01  a  Svalbard  F  194  185  0.75  -0.27  b  Idem  M  225  390  0.537  -0.4  Comments/Source Length at age; 0-26 years. Average W∞ from Tab. 1 (46a-b). Siciliano et al. (2007; Tab. 3). From Gompertz curve; <1-30 years. W∞ from Tab. 1 (46a). Mattson et al. (2006; Fig. 6). From Gompertz curve; <1-30 years. W∞ from Tab. 1 (46b). Mattson et al. (2006; Fig. 6). From Gompertz equation. W∞ from Tab. 1 (46a). Stolen et al. (2002; Tab. 1). From Gompertz equation. W∞ from Tab. 1 (46b). Stolen et al. (2002; Tab. 1). W∞ from Tab. 1 (47a). Hunter (2005; Tab. A.8). W∞ from Tab. 1 (47b). Hunter (2005; Tab. A.8)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  27  LIFE-HISTORY PATTERNS IN MARINE BIRDS 1 Vasiliki S. Karpouzi  Former address: The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email: v.karpouzi@fisheries.ubc.ca Current address: BC Ministry of Environment, Environmental Stewardship Division, Ecosystems Branch, 2nd floor, 10470 - 152nd St, Surrey BC, V3R 0Y3, Canada  Daniel Pauly  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email: d.pauly@fisheries.ubc.ca  ABSTRACT The parameters of the von Bertalanffy growth equation for seabirds were estimated from previously published growth curves to allow within and between group comparisons of their life-history patterns. Overall, growth data were available for 447 seabird populations breeding around the globe, corresponding to 137 species, 13 families and four orders. A negative relationship between the logarithmic values of W∞ and K was identified for the orders of Charadriiformes, Pelecaniformes, Procellariiformes, and Sphenisciformes, as well as all seabird species combined. The values of the slope b ranged from -0.32 for the Sphenisciformes, to -0.12 for the Pelecaniformes, with a mean slope of -0.21, when all seabirds were considered.  INTRODUCTION Seabirds can be broadly characterized as long-lived species, with delayed sexual maturation and breeding, as well as low annual reproductive rates. Many species have a life span well over 30 years (e.g., most species of albatrosses; Schreiber & Burger, 2002). In addition, most species start breeding when they are three years or older (e.g., over ten years in some albatross species; Schreiber and Burger, 2002). Most species lay not more than three eggs, and in some instances chick rearing lasts for a long time (e.g., 380 days in the Wandering albatross; Schreiber & Burger, 2002). These life-history characteristics have been shaped as an evolutionary response to conditions of living in the marine environment, i.e, reflecting the patchy and unpredictable distribution of marine resources (Ricklefs, 1990; Hamer et al., 2002; Weimerskirch, 2002), which poses challenges to seabirds in finding food and provisioning chicks. Nonetheless, Weimerskirch (2007) has recently suggested that prey dispersal may not be as unpredictable as we once thought. As a result, specialization of seabirds for use of a particular marine habitat may be the driving force for the evolution of a particular life history strategy (Weimerskirch, 2007). Life-history strategies have been studied for a number of marine organisms (fish: e.g., Adams, 1980; Froese & Pauly, 1998; Pauly, 1998; Stergiou, 2000; marine mammals: e.g., Herzing, 1997; Trites & Pauly, 1998; sea turtles: e.g., Fraser & Ehrhart, 1985; van Buskirk & Crowder, 1994; marine birds: e.g., Ricklefs, 1990; Visser, 2002; Weimerskirch, 2007). In addition, examining relationships between life history traits and developing empirical equations has been proven useful for comparisons among different taxonomic groups (e.g., Peters, 1983; Froese & Pauly, 1998; Visser, 2002). Growth, in particular, has been described in seabirds with the use of several mathematical equations. Three of the most popular ones are listed in Table 1 (see also Table 8.1 in Peters, 1983). For all three equations, parameters are estimated by fitting the selected model to size-at-age data for each individual or population under study. These equations allow Cite as: Karpouzi, V.S., Pauly, D. 2008. Life history patterns in marine birds. In: Palomares, M.L.D., Pauly, D. (Eds.), Von Bertalanffy Growth Parameters of Non-Fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 27-53.  1  28  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  the estimation of the key parameters of chick growth, such as the growth constant, K, and the asymptotic weight, W∞ and a measure of how rapidly this approached (K1, KG, K). In the present study, we compiled information on growth parameters of seabird chicks, in an attempt to explore their life-history patterns. In addition, we established empirical relationships between life-history traits, in order to investigate potential differences in growth rates for a few seabird orders.  Table 1: Three equations most frequently used to describe chick growth in marine birds. Growth curve  Equation  Description  1) Logistic  Wt = W∞ / (1 + e -KL·(t-tL) )  -KG·(t-tG)  2) Gompertz  Wt = W∞ · e -e  3) Von Bertalanffy (VB)  Wt = W∞ · (1 - e -K·(t-t0) )b  W∞: asymptotic weight; KL: logistic growth rate constant; tL: the time of inflection point, which corresponds  to the age of 50% of asymptotic weight of chicks. W∞: asymptotic weight; KG: Gompertz growth rate constant; tG: the time of inflection point. W∞: asymptotic weight; K: VB growth rate constant; to: the theoretical ‘age’ the chick would have at weight zero; b: exponent indicating isometric growth pattern, when its value is 3.  METHODOLOGY In the present study, we gathered all available information pertinent to growth patterns in seabird chicks, from studies conducted since 1937. Overall, growth data were available for 447 seabird populations, corresponding to 137 species, 13 families and four orders (Tables 2 to 4). For the purpose of this paper, we defined as seabird population a number of seabirds belonging to the same species and breeding at a certain location at a certain year (Tables 2 to 4). We gathered information using the following databases: (a) Aquatic Sciences and Fisheries Abstracts; (b) Web of Science - Thomson Scientific; (c) BioSciences Information Service of Biological Abstracts; and (d) the Searchable Ornithological Research Archive, which cover peer-reviewed journals and other literature sources. We also used some unpublished theses and technical reports that were available to us, and extracted information from the online database of Birds of North America, Cornell University (http://bna.birds.cornell.edu/BNA/). The form of the VBGF used here is:  Wt = W∞ · (1-e-K·(t-t0))3  …1)  where Wt is the weight at age t, W∞ the asymptotic size (here the size of a chick if it were to continue growing forever in the manner described by the equation), K is a parameter of dimension time-1 (here: year-1), and to adjusts the function such that Wt=0 at t=to. In case where the original graph of chick body weight-at-age was not available, we obtained data for the following life-history parameters, to fit the VBGF: (a) the asymptotic weight, W∞ (g), the growth constant, KL (in days-1), and the inflection point, tL (in days), of the logistic growth curve (Table 2); and (b) the asymptotic weight, W∞ (in g), the growth constant, KG (in days-1), and the inflection point, tG (in days), of the Gompertz growth curve (Table 3). When information on KL and tL was not available, we used the following equations respectively to estimate the missing values:  KL = 0.962· W∞-0.31 (Visser, 2002); and  ... 2  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  29  tL = ln (W∞/W0 - 1) / KL  … 3)  where Wo is the initial weight of the chicks at hatching (Navarro and Bucher, 1990).  VBGF  200  Logistic  Weight (g)  150 100 50 0 -25  0  25  50  -50  (b)  5000 Gompertz VBGF 2500  0 -25  When the chick body weight-at-age graph was provided in the original studies, data was traced and re-analyzed using the least square optimization technique to get growth parameter estimates in terms of the VBGF. An example is given in Figure 2, for the growth of Little shearwater (Puffinus assimilis) chicks of New Zealand.  250  (a)  Weight (g)  For each seabird population, we calculated seven data points of time when chick body weight is equal to 1%, 5%, 10%, 25%, 50%, 75%, 90%, 95%, and 99% of W∞, using the growth curve published by the corresponding study, and then used these data points to re-express growth parameters in terms of the VBGF (see Figure 1 for examples). As the VBGF and the other growth functions share one parameter (W∞), we kept this constant, and used the least square optimization technique to estimate the other VBGF parameters (K, to), given the data points. An example is given in Figure 1, for the growth of the Crested auklet (Aethia cristatella) chicks of the Okhotsk Sea (Figure 1a), and of the Least auklet (Aethia pussila) chicks of the Pribilof Islands, Alaska (Figure 1b).  0  25  50  75  100  125  150  Time (days)  Figure 1. Comparison of the von Bertalanffy growth function (VBGF; black dot, solid line) with the growth curve originally used (open dot, dotted line) to describe weight-at-age data (not shown). (a) Crested auklet (Aethia cristatella) from the Okhotsk Sea, described with the logistic growth function (Kitaysky, 1999). (b) Light-mantled albatross (Phoebetria palpebrata) from Macquarie Island, Southern Ocean, described with the Gompertz growth function (Terauds and Gales, 2006).  The standard deviation (SD) was also estimated as a measure of effectiveness of the least-squares optimization. For all seabird populations, SD was then re-expressed as a % deviation (%D), i.e., relative to Wt=0.5* W∞. Pauly et al. (1996) proposed the auximetric plot as another tool for the comparison of within- and between- species growth patterns. The auximetric plot is a double logarithmic plot of the parameters K and the asymptotic size (W∞ or L∞) (Pauly et al., 1996; Froese and Pauly, 2000). In such a plot, each set of growth parameters represents a point, with the different points for a species or higher taxon forming an ellipsoid cluster of points, whose surface area is related to the ‘growth space’ occupied by a given species or higher taxon (Pauly et al., 1996; Froese and Pauly, 2000).  30  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  Table 2. Growth parameters of seabird chicks from the logistic growth curve (in normal font), and also estimated for this paper (in bold), using the von Bertalanffy (VB) growth model. W∞ (in g): asymptotic weight of chicks for both the logistic and VB growth models; tl (in days): inflection point of logistic curve; to (in years): hypothetical ‘age’ chicks would have had at zero weight; Kl (in days-1) and K (in years-1): growth coefficients for the logistic and VB models respectively. AK: Alaska; CA: California; WA: Washington State; and NY: New York State. Species Alcidae  Area (Year)  tl  Kl  W∞  K  to  Source  Aethia cristatella Cepphus columba Cepphus columba Cepphus columba Cepphus columba Cepphus columba Cepphus columba Cepphus columba Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Cerorhinca monocerata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula cirrhata Fratercula corniculata Fratercula corniculata Fratercula corniculata Fratercula corniculata Fratercula corniculata Fratercula corniculata  Okhotsk Sea (1994) Farallon Is, AK (1989) Farallon Is, AK (1990) Farallon Is, AK (1991) Farallon Is, AK (1992) Farallon Is, AK (1993) Farallon Is, AK (1994) Farallon Is, AK (1995) Destruction Is, WA (1974) Destruction Is, WA (1975) Destruction Is, WA (1979) Destruction Is, WA (1980) Destruction Is, WA (1981) Protection Is, WA (1975) Protection Is, WA (1976) Protection Is, WA (1979) Protection Is, WA (1980) Protection Is, WA (1981) Okhotsk Sea (1994) Buldir Is, AK (1975) Ugaiushak Is, AK (1976) Barren Is, AK (1976) Chowiet Is, AK (1976) Shumagin Is, AK (1976) Wooded Is, AK (1976) Ugaiushak Is, AK (1977) Barren Is, AK (1977) Sitkalidak, AK (1977) Cathedral Is, AK (1977) Buldir Is, AK (1975) Barren Is, AK (1976) Chowiet Is, AK (1976) Shumagin Is, AK (1976) Ugaiushak Is, AK (1977) Barren Is, AK (1977)  14.17 *14.01 *12.75 *15.86 *15.90 *15.91 *13.95 *15.26 *23.20 *27.74 *32.78 *38.44 *39.49 *24.61 *27.71 *25.88 *25.15 *32.59 20.41 *19.35 *16.27 *18.53 *14.54 *12.90 *16.13 *12.72 *18.62 *15.88 *15.71 *20.13 *16.07 *12.61 *12.95 *12.89 *17.31  0.129 ¤0.150 ¤0.155 ¤0.143 ¤0.143 ¤0.143 ¤0.150 ¤0.145 0.074 0.068 0.058 0.049 0.049 0.076 0.071 0.076 0.078 0.061 0.118 0.074 0.125 0.111 0.091 0.145 0.120 0.153 0.110 0.126 0.127 0.075 0.122 0.113 0.144 0.139 0.114  233 401 359 466 468 469 398 443 335 395 400 394 412 412 432 432 430 440 621 360 600 600 330 520 550 555 595 590 580 300 440 280 405 380 445  20.96 24.37 25.19 23.24 23.24 23.24 24.37 23.56 12.02 11.05 9.43 7.96 7.96 12.35 11.54 12.35 12.67 9.91 19.17 12.02 20.31 18.04 14.79 23.56 19.50 24.86 17.87 20.47 20.64 12.18 19.82 18.36 23.40 22.59 18.52  -0.043 -0.032 -0.033 -0.031 -0.031 -0.031 -0.032 -0.031 -0.080 -0.080 -0.093 -0.111 -0.108 -0.072 -0.073 -0.069 -0.067 -0.085 -0.034 -0.090 -0.040 -0.045 -0.077 -0.038 -0.044 -0.034 -0.045 -0.041 -0.040 -0.086 -0.043 -0.059 -0.038 -0.041 -0.046  Kitaysky (1999) Shultz and Sydeman (1997) Shultz and Sydeman (1997) Shultz and Sydeman (1997) Shultz and Sydeman (1997) Shultz and Sydeman (1997) Shultz and Sydeman (1997) Shultz and Sydeman (1997) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Wilson and Manuwal (1986) Kitaysky (1999) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983) Wehle (1983)  Oceanodroma homochroa  Farallon Is, CA (1985)  *13.56  0.108  49  17.55  -0.061  Ainley et al. (1990)  Larus atricilla Larus ridibundus Sterna hirundo Sterna paradisaea  Florida (1972) The Netherlands (2000) Great Gull Is, NY (1968) Shetland Is (1975)  *13.45 9.90 8.22 7.50  ¤0.162 0.200 0.246 0.288  310 237 113 111  26.32 32.50 39.97 46.80  -0.029 -0.026 -0.021 -0.016  Dinsmore and Schreiber (1974) Eising and Groothuis (2003) LeCroy and Collins (1972) Furness (1978)  Pelecanoides georgicus Pelecanoides urinatrix  S Georgia (1982) S Georgia (1982)  14.90 15.60  0.145 0.146  148 139  23.56 23.72  -0.032 -0.030  Roby (1991) Roby (1991)  Phaethon lepturus  Seychelles (2002)  35.00  ¤0.155  362  25.19  0.027  Hydrobatidae  Laridae  Pelecanoididae  Phaethontidae  Ramos and Pacheco (2003)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  31  Table 2 continued. tl  Kl  W∞  K  20.00 21.40 17.10 16.10 21.40 18.10 19.60  0.104 0.168 0.152 0.158 0.123 0.122 0.125  951 793 836 642 1110 1148 1154  16.90 27.30 24.70 25.67 19.99 19.82 20.31  -0.047 -0.004 -0.023 -0.023 -0.028 -0.037 -0.031  Montague (1982) Montague (1982) Hodgson (1975) Jones (1978) Kinsky (1960) Richdale (1940) Gales (1987)  Shetland Is. (1975) 16.96 0.176 * Estimated using equation 2 described in the methodology. ¤ Estimated using equation 3 described in the methodology.  1167  28.60  -0.014  Furness (1978)  Species Spheniscidae  Eudyptula minor  Area (Year) Victoria, Australia (1980) Victoria, Australia (1981) Tasmania (1970) New Zealand (1975) New Zealand (1958) New Zealand (1938) New Zealand (1983)  to  Source  Stercorariidae  Catharacta skua  RESULTS 300  200 Weight (g)  Table 2 summarizes the logistic growth parameters as extracted from the corresponding studies, as well as the growth parameters we re-expressed using the VBGF. Similarly, Table 3 summarizes growth parameters derived from the Gompertz growth model, which we then re-expressed using the VBGF. Lastly, Table 4 summarizes VBGF parameters re-estimated after tracing and re-analyzing originally published weight-at-age data. W∞ values ranged from 36 g for the Wilson’s storm petrel (Oceanites oceanicus) chicks (Table 4), to 15,243 g for the Wandering albatross (Diomedea exulans) chicks both from the Crozet Islands, Southern Ocean (Table 4).  100  0 -20  0  20  40  60  80  Time (days)  Figure 2. Von Bertalanffy growth function (solid line) for Little shearwater (Puffinus assimilis) chicks from Lady Alice Island, New Zealand, re-estimated using original weight-at-age data (black dot) published by Booth et al. (2000).  The most intensively studied species were the Pigeon guillemot (Cepphus columba), the Rhinoceros auklet (Cerorhinca monocerata), the Atlantic puffin (Fratercula arctica), the Tufted puffin (Fratercula cirrhata), the Horned puffin (Fratercula corniculata), the Common tern (Sterna hirundo), the Black-legged kittiwake (Rissa tridactyla), the Thick-billed murre (Uria lomvia), and the Blue penguin (Eudyptula minor). They were represented by more than 10 seabird populations each (Tables 2 to 4), and comprised 31% of all seabird populations compiled (138 out of 447; Tables 2 to 4). All the above-mentioned seabird species, with the exception of the Blue penguin, belong to the order Charadriiformes (Tables 2 to 4). KL values ranged from 0.049 days-1 for the Rhinoceros auklet chicks of Destruction Island, off the coast of Washington State (Table 2), to 0.288 days-1 for the Arctic tern (Sterna paradisaea) chicks of Shetland Islands, UK (Table 2). KL values were not available for nine seabird populations (Table 2). These were estimated using equation 2 described above. The values of tL ranged from 7.5 days for the Arctic tern chicks of Shetland Islands, UK (Table 2), to 39.5 days for the Rhinoceros auklet chicks of Destruction Island, off the coast of Washington State (Table 2). Values of tL were lacking for 35 seabird populations (Table 2). These were estimated using equation 3 described in the methodology section.  32  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  Table 3. Growth parameters of seabird chicks from the Gompertz growth curve (in normal font), and also estimated for this paper (in bold), using the von Bertalanffy growth function (VBGF). W∞ (in g): the asymptotic weight of chicks for both the Gompertz curve and VBGF; tG (in days) and KG (in days-1): the inflection point and the growth constant for the Gompertz curve respectively; to (in years) and K (in years-1): the hypothetical ‘age’ chicks would have at zero weight and the growth constant for VBGF respectively. AK: Alaska. Species Diomedeidae  Phoebastria immutabilis Phoebastria nigripes Phoebetria palpebrata  tG  KG  W∞  K  to  19.00 17.90 32.40  0.050 0.056 0.047  2836 2714 4760  14.84 16.62 13.95  -0.037 -0.010 -0.006  Area (Year) Hawaii (1987) Hawaii (1987) Macquarie Is (2000)  Source Sievert and Sileo (1993) Sievert and Sileo (1993) Terauds and Gales (2006)  KG values ranged from 0.047 days-1 for the Light-mantled albatross (Phoebetria palpebrata) chicks of Macquarie Island, Southern Ocean (Table 3), to 0.056 days-1 for the Black-footed albatross (Phoebastria nigripes) chicks of Hawaii (Table 3). In addition, tG ranged from 17.9 days for the Black-footed albatross chicks of Hawaii (Table 3), to 32.4 days for the Light-mantled albatross chicks of Macquarie Island, Southern Ocean (Table 3). When re-expressed through VBGF, the logistic growth curve deviated from VBGF by 13%, while the Gompertz curve deviated by only 3%. This suggests that the VBGF and the Gompertz curves are equivalent.  A negative relationship between the logarithmic values of W∞ and K was identified for the orders of Charadriiformes, Pelecaniformes, Procellariiformes, and Sphenisciformes as well as for all seabird species combined (Table 4, Figure 3). Each order was represented by 239, 50, 111 and 47 seabird populations respectively (Table 4, Figure 3). The values of the slope ranged  Table 4. Regression equations between the von Bertalanffy growth parameters K and W∞, for four orders and all seabird species combined. SE(b): Standard error of the slope. r: The correlation coefficient. N: The number of seabird populations representing each order. All regressions were statistically significant (P<0.05). Order Charadriiformes Pelecaniformes Procellariiformes Sphenisciformes All seabirds  Regression LogK=2.18-0.31LogW∞ LogK=1.63-0.12LogW∞ LogK=1.79-0.18LogW∞ LogK=2.35-0.32LogW∞ LogK=1.93-0.21LogW∞  SE(b) 0.03 0.05 0.02 0.05 0.01  r -0.53 -0.35 -0.61 -0.70 -0.62  N 239 50 111 47 445  P P<0.05 P<0.05 P<0.05 P<0.05 P<0.05  2.0  1.5 -1  K (log; year )  Computed K values ranged from 3.22 years-1 for the Wandering albatross chicks of the Crozet Islands, Southern Ocean (Table A1), to 61.46 years-1 for the Cory’s shearwater (Calonectris diomedea) chicks of Selvagem Grande, of the Madeira archipelago (Table A1). Moreover, to values ranged from 0.111 years for the Rhinoceros auklet chicks of Destruction Island, off the coast of Washington State (Table 2), to 0.001 years for the Whiskered auklet (Aethia pygmaea) chicks of Buldir Island, Alaska (Table A1), the Rhinoceros auklet chicks of Teuri Island, Japan (Table A1), and the Masked booby (Sula dactylatra) chicks, of Kure Atoll, Hawaii (Table A1).  1.0  Charadriiformes, n=239  0.5  Pelecaniformes, n=50 Procellariiformes, n=111 Sphenisciformes, n=47 0.0 0.0  1.0  2.0  3.0  4.0  5.0  W∞ (log; g)  Figure 3. Auximetric plot for the four orders of Charadriiformes, Sphenisciformes, Procellariiformes and Pelecaniformes (see text).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  33  from -0.32 for Sphenisciformes, to -0.12 for Pelecaniformes (Table 4). All regressions were statistically significant (P<0.05; Table 4). This justifies the use of auximetric plots in seabirds.  DISCUSSION In the present study, relationships were established between the life-history parameters K and W∞, for a number of seabird populations around the globe. However, these relationships were based on information available for about 39% of the world’s seabirds (137 out of 351 in Karpouzi et al., 2007). Thus, these relationships are provisional and subject to change when additional information becomes available. Nonetheless, they can be particularly useful in estimating approximate values of K from W∞, and hence to obtain a preliminary growth curve for species without growth data. In addition, they allow us to compare growth patterns of seabirds to those from other groups of organisms, whose growth has also been described using the VBGF. Similar studies that investigate the relationship between the growth parameters K and asymptotic size have also been conducted mainly for fish species of marine and freshwater ecosystems. In particular, Winemiller and Rose (1992) analyzed life-history patterns of 216 North American marine and freshwater fish species, belonging to 57 families. Pauly (1998) analyzed growth parameters for 4826 fish populations listed in FishBase (www.fishbase.org; Froese and Pauly, 2000). Stergiou (2000) explored life-history patterns for 40 fish species from Greek waters, belonging to 20 families, and compared them with those from Pauly (1998). Starck and Ricklefs (1998) compiled information on the growth parameters KL, KG and W∞ for 1117 populations, belonging to 557 bird species, from both terrestrial and aquatic ecosystems. Out of these, 366 belonged to marine birds and represented 114 seabird species, and 13 families (Starck and Ricklefs, 1998). Ricklefs et al. (1998) and later Visser (2002) used these data to examine the relationship between KL and W∞. Their analyses revealed that growth rates are particularly low for many pelagic seabird species, and tend to be higher in species that feed close to shore, such as the larid species. In addition, highest growth rates are observed among penguin species (Ricklefs et al., 1998; Visser, 2002). The VBGF parameters can be linked by the relation W∞=a*K-b (e.g., Beverton and Holt, 1959; Adams, 1980; Pauly, 1980; Charnov, 1993; Pauly, 1998; Froese and Pauly, 2000). For fish, the exponent b takes values that generally range from -0.27 to -0.80 (Charnov, 1993; Stergiou, 2000). The value of b for all Greek fish stocks is equal to -0.32 (Stergiou, 2000). In contrast, the value of b equals -0.57 for the 4826 populations analyzed by Pauly (1998). When all seabird populations were taken into account, the slope on the auximetric plot was equal to -0.21 (Table 5). This value was heavily influenced by 53% of the K and W∞ values of Charadriiformes (239 out of 447 seabird populations; Table 5). Hence, it may be subject to change when growth parameter estimates from other seabird species belonging to the other three orders becomes available. The auximetric plot revealed differences in the growth potential of the seabird species included in this study (Figure 3). Indeed, the growth spaces occupied by the four orders of seabirds seem to reflect differences in the seabirds’ breeding biology (e.g., adult foraging behaviour during chick-rearing; e.g., Fernández et al., 2001; parental feeding strategies; e.g., Ydenberg, 1989). In particular, tern and gull species with generally smaller body size exhibited faster growth rates (Figure 3). These are species that produce large clutches, which tend to transport food from areas close to shore to feed their young (e.g., Hulsman and Smith, 1988). On the other hand, alcid species, also characterized by small body size, grow more slowly (Figure 3). Alcid species produce single-egg clutches, and exhibit a more pelagic, nocturnal foraging behaviour (e.g., Sealy, 1973; Ricklefs, 1982, 1990). As a result, provisioning rates, and consequently growth rates of chicks, are reduced (e.g., Sealy, 1973; Ricklefs, 1982, 1990). Some procellariiform species (e.g., storm petrels of the family Hydrobatidae) also displayed a growth pattern similar to that of the alcid species (Table 4; Figure 3). Storm petrels are small in size. However, the growth rates of their chicks are relatively low (Table 4; Figure 3). Storm petrels feed far from nesting colonies on prey that is sparse and unpredictably distributed. The single-clutch size may suggest that their ability to deliver energy to the brood is severely limited (e.g., Place et al., 1989). Slow growth may be an adaptation to reduce the rate at which chicks require energy for development, thus making it easier for parents to utilize more distant and sparse food resources for breeding (e.g., Ricklefs et al., 1980; Place et al., 1989).  34  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  The large-bodied Sphenisciformes (i.e., penguins) exhibited high growth rates (Tables 2 and 4; Figure 3). A similar growth pattern was also observed by Visser (2002). This has been interpreted as an adaptation to the severe Antarctic conditions that shorten the breeding season. Indeed, faster growth rates enable chicks to leave the colony before the beginning of the winter (Volkman and Trivelpiece, 1980; Visser, 2002). Lastly, albatross species with large body size exhibited slow growth rates (Tables 3 and 4; Figure 3). This growth pattern is typical of the albatross family, which is dominated by long-distance foragers (e.g., Fernández et al., 2001). 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Growth parameters of seabird chicks re-estimated for this paper using the von Bertalanffy growth function (VBGF), from body weight-at-age data published in the original studies. W∞ (g): the asymptotic weight of chicks; K (years-1) and to (in years): the growth constant and the hypothetical ‘age’ chicks would have at zero weight respectively. Species Alcidae  Aethia cristatella Aethia pusilla Aethia pygmaea Alca torda Alle alle  Brachyramphus marmoratus Cepphus carbo Cepphus columba  Cepphus grylle  Cerorhinca monocerata  Cyclorrhynchus psittacula  Area (Year) Buldir Is, Alaska (1996) Buldir Is, Alaska (1997) St Lawrence Is, Alaska (1987) Kiska Is, Alaska (2003) Pribilof Is, Alaska (1982) St Lawrence Is, Alaska (1987) Buldir Is Alaska (1998) Machias Seal Is (1995) Machias Seal Is (2003) Franz Josef Land (1993) Svalbard (1978) Svalbard (1984) Svalbard (1987) Svalbard (1992) Barren Is, Alaska (1978) Barren Is, Alaska (1979) Teuri Is, Japan (1989) Farallon Is, California (1985) Mandarte Is, British Columbia (1960) Mitlenatch Is, British Columbia (1985) Prince William Sound, Alaska (1978) Queen Charlotte Is, British Columbia (1991) Piqiuliit, Nunavut (1983) Pitsiulak, Nunavut (1981) Pitsiulak, Nunavut (1982) Pitsiulak, Nunavut (1983) Québec (1977) Cleland Is, British Columbia (1969) Protection Is, Washington (1989) Protection Is, Washington (1990) Protection Is, Washington (1991) Teuri Is, Japan (1994) Teuri Is, Japan (1995) Teuri Is, Japan (1996) Teuri Is, Japan (1997) Teuri Is, Japan (1998) Triangle Is, British Columbia (1978) Buldir Is, Alaska (1991)  W∞ 376 358 299 80 114 95 113 189 208 152 138 136 178 138 152 167 806 447 476 421 607 412 404 386 408 447 448 455 355 392 455 593 615 550 329 439 406 266  K  to  20.47 20.62 32.75 33.60 34.41 35.40 32.92 44.81 54.25 44.37 41.05 37.53 30.53 37.80 42.84 37.16 19.45 28.59 27.19 29.61 26.04 29.82 28.10 28.53 26.76 26.09 25.60 10.00 7.15 6.01 5.69 10.78 10.20 9.35 7.10 12.07 22.04 26.47  -0.011 -0.011 -0.018 -0.018 -0.018 -0.019 -0.018 -0.022 -0.022 -0.021 -0.015 -0.019 -0.019 -0.019 -0.021 -0.020 -0.017 -0.016 -0.017 -0.015 -0.015 -0.016 -0.016 -0.017 -0.018 -0.017 -0.018 -0.054 -0.076 -0.091 -0.104 -0.005 -0.001 -0.006 -0.072 -0.004 -0.026 -0.017  Source Fraser et al. (1999) Fraser et al. (1999) Piatt et al. (1990) Major et al. (2006) Roby and Brink (1986) Piatt et al. (1990) Hunter et al. (2002) Bond et al. (2006) Bond et al. (2006) Stempniewicz et al. (1996) Clark and Ydenberg (1990) Clark and Ydenberg (1990) Konarzewski and Taylor (1989) Stempniewicz et al. (1996) Simons (1980) Hirsch et al. (1981) Minami et al. (1995) Ainley and Boekelheide (1990) Drent (1965) Emms and Verbeek (1991) Oakley (1981) Vermeer et al. (1993) Cairns (1987) Cairns (1987) Cairns (1987) Cairns (1987) Cairns (1981) Summers and Drent (1979) Wilson (1993) Wilson (1993) Wilson (1993) Takahashi et al. (2001) Takahashi et al. (2001) Takahashi et al. (2001) Takahashi et al. (2001) Takahashi et al. (2001) Vermeer and Cullen (1982) Hipfner and Byrd (1993)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  45  Appendix 1. Continued Species  Fratercula arctica  Fratercula cirrhata Fratercula corniculata  Area (Year) Bleiksøy, Norway (1982) Bleiksøy, Norway (1986) Bleiksøy, Norway (1987) Farne Is, UK (1963) Gannet Is, Newfoundland (1996) Gannet Is, Newfoundland (1997) Gannet Is, Newfoundland (1998) Gull Is, Newfoundland (1998) Hornøy, Norway (1980) Hornøy, Norway (1981) Is May, UK (1975) Is May, UK (1992) Is May, UK (1995) Machias Seal Is (1997) Machias Seal Is (1999) Machias Seal Is (2003) Røst, Norway (1983) Røst, Norway (1984) Røst, Norway (1985) Røst, Norway (1988) Røst, Norway (1989) Røst, Norway (1990) Røst, Norway (1991) Røst, Norway (1992) Røst, Norway (1993) Røst, Norway (1996) Røst, Norway (1999) Røst, Norway (2000) W Scotland, UK (1975) Wales, UK (1977) Wales, UK (1978) Destruction Is, Washington (1975) Prince William Sound, Alaska (1995) Triangle Is, British Columbia (2000) Duck Is, Alaska (1995) Duck Is, Alaska (1996) Duck Is, Alaska (1997) Duck Is, Alaska (1998) Duck Is, Alaska (1999)  W∞ 280 118 221 195 317 442 438 236 387 372 334 265 310 367 221 379 377 222 292 182 326 304 306 368 228 219 260 188 339 353 337 528 604 517 511 371 472 303 402  K  to  19.00 36.54 14.86 27.51 22.11 18.01 20.00 26.61 29.49 32.72 28.87 31.23 31.28 26.58 28.49 19.74 27.15 35.49 16.10 26.37 33.76 31.81 29.50 32.27 43.54 37.92 43.68 53.82 21.81 27.07 23.81 25.50 25.82 29.38 20.31 26.23 20.43 33.01 31.80  -0.039 -0.030 -0.053 -0.028 -0.028 -0.032 -0.029 -0.028 -0.018 -0.020 -0.018 -0.024 -0.019 -0.019 -0.029 -0.029 -0.017 -0.020 -0.040 -0.030 -0.019 -0.018 -0.018 -0.018 -0.019 -0.021 -0.020 -0.022 -0.025 -0.016 -0.017 -0.017 -0.017 -0.027 -0.033 -0.025 -0.039 -0.026 -0.021  Source Barrett et al. (1987) Barrett and Rikardsen (1992) Barrett and Rikardsen (1992) Pearson (1968) Baillie and Jones (2003) Baillie and Jones (2003) Baillie and Jones (2003) Baillie and Jones (2003) Barrett et al. (1987) Barrett and Rikardsen (1992) Harris (1978) Wernham and Bryant (1998) Cook and Hamer (1997) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Barrett et al. (1987) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Anker-Nilssen and Aarvak (2002) Harris (1978) Ashcroft (1979) Hudson (1979) Burrell (1980) Piatt et al. (1997) Gjerdrum (2004) Harding et al. (2003) Harding et al. (2003) Harding et al. (2003) Harding et al. (2003) Harding et al. (2003)  46  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  Appendix 1. Continued Species  Ptychoramphus aleuticus  Uria aalge  Uria lomvia  Area (Year) California Channel Is (2001) California (1959) Farallon Is, California (1971) Triangle Is, British Columbia (1996) Triangle Is, British Columbia (1997) Triangle Is, British Columbia (1998) Triangle Is, British Columbia (1999) Farne Is, UK (1963) Is May, UK (1992) St Lawrence Is, Alaska (1972) Sweden (1974) Sweden (1975) Sweden (1976) Sweden (1977) Wales, UK (1987) Cape Hay, Northwest Territories (1979) Coats Is, Nunavut (1991) Coats Is, Nunavut (1994) Coats Is, Nunavut (1995) Coburg Is, Northwest Territories (1979) Digges Is, Nunavut (1999) Prince Leopold Is, Nunavut (2000) Prince Leopold Is, Nunavut (2001) Prince Leopold Is, Nunavut (2002) St Lawrence Is, Alaska (1972)  W∞  K  to  150 155 192 118 149 136 187 169 267 229 320 278 291 292 234 215 268 268 231 247 137 305 200 117 211  26.45 30.42 21.59 32.70 31.15 27.99 25.48 55.21 54.34 38.73 36.74 43.02 37.97 39.64 45.19 44.67 41.66 37.92 41.69 41.48 56.41 33.09 36.92 41.05 48.51  -0.027 -0.016 -0.014 -0.018 -0.018 -0.021 -0.019 -0.023 -0.022 -0.026 -0.021 -0.021 -0.022 -0.021 -0.022 -0.021 -0.021 -0.024 -0.023 -0.021 -0.023 -0.019 -0.021 -0.024 -0.023  Source Ackerman et al. (2004) Thoresen (1964) Manuwal (1974) Hedd et al. (2002a) Hedd et al. (2002a) Hedd et al. (2002a) Hedd et al. (2002a) Pearson (1968) Harris and Wanless (1995) Johnson and West (1975) Hedgren and Linnman (1979) Hedgren and Linnman (1979) Hedgren and Linnman (1979) Hedgren and Linnman (1979) Hatchwell (1991) Birkhead and Nettleship (1981) de Forest and Gaston (1996) Hipfner et al. (2006) Hipfner et al. (2006) Birkhead and Nettleship (1981) Hipfner et al. (2006) Gaston et al. (2005) Gaston et al. (2005) Gaston et al. (2005) Johnson and West (1975)  Amsterdam Is (1984) Crozet Is (1986) Crozet Is (1994) Crozet Is (2000) Midway Atoll, Hawaii (1965) Macquarie Is (2001) S Georgia (1977) Albatross Is, Australia (1998) Amsterdam Is (1996) Amsterdam Is (1997) Amsterdam Is (2001) S Georgia (1976) S Georgia (1996) S Georgia (1976) S Georgia (1996)  8818 12249 11557 15243 2478 3741 3247 5986 2921 2492 2732 5090 3755 5540 4002  7.58 7.06 8.35 3.22 15.72 16.58 16.46 10.84 22.29 14.42 31.01 12.02 17.18 12.92 17.93  -0.028 -0.033 -0.038 -0.006 -0.037 -0.011 -0.017 -0.060 -0.041 -0.086 -0.029 -0.025 -0.003 -0.023 -0.006  Jouventin et al. (1989) Lequette and Weimerskirch (1990) Weimerskirch and Lys (2000) Mabille et al. (2004) Fisher (1967) Terauds and Gales (2006) Thomas et al. (1983) Hedd et al. (2002b) Weimerskirch et al. (2001) Weimerskirch et al. (2001) Pinaud et al. (2005) Ricketts and Prince (1981) Huin and Prince (2000) Ricketts and Prince (1981) Huin and Prince (2000)  Baja California, Mexico (1988) Barbuda (1971)  1424 1369  10.58 9.15  -0.021 -0.042  Carmona et al. (1995) Diamond (1973)  Diomedeidae  Diomedea amsterdamensis Diomedea exulans Phoebastria immutabilis Phoebetria palpebrata Thalassarche cauta Thalassarche chlororhynchos Thalassarche chrysostoma Thalassarche melanophris  Fregatidae  Fregata magnificens  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  47  Appendix 1. Continued Species Hydrobatidae  Fregetta tropica Garrodia nereis Hydrobates pelagicus Oceanites oceanicus  Oceanodroma furcata  Oceanodroma leucorhoa  Oceanodroma tristrami Pelagodroma marina Laridae  Anous minutus Anous stolidus  Anous tenuirostris  Chlidonias niger Creagrus furcatus Gygis alba Larus argentatus Larus atricilla  Area (Year) Crozet Is (1982) S Shetland Is (1996) Chatham Is, New Zealand (1987) Shetland Is, UK (1992) Crozet Is (1982) S Shetland Is (1996) W Antarctic Peninsula (1986) Barren Is, Alaska (1976) Barren Is, Alaska (1977) Barren Is, Alaska (1978) Queen Charlotte Is, British Columbia (1983) Kent Is, New Brunswick (1962) Kent Is, New Brunswick (1972) Kent Is, New Brunswick (1983) Kent Is, New Brunswick (1988) Queen Charlotte Is, British Columbia (1983) Laysan Is, Hawaii (1991) Selvagem Grande (1996) Victoria, Australia (2003) Victoria, Australia (2003) Hawaii (1981) Manana Is, Hawaii (1972) Puerto Rico (1989) Seychelles (1995) Seychelles (1996) Seychelles (2001) Tern Is, Hawaii (1989) Houtman Abrolhos, Australia (1991) Seychelles (1995) Seychelles (1996) Seychelles (1997) Seychelles (2001) Seychelles (2002) The Netherlands (1995) Galápagos (1966) Galápagos (1967) Hawaii (1981) Appledore Is, New Hampshire (1973) Germany (1996) Florida (1976)  W∞  K  to  50 118 74 40 36 59 58  32.70 20.74 22.53 23.69 32.36 33.99 26.92  -0.023 -0.022 -0.019 -0.024 -0.023 -0.017 -0.025  Jouventin et al. (1985) Hahn (1998) Plant (1989) Bolton (1995) Jouventin et al. (1985) Quillfeldt and Peter (2000) Obst and Nagy (1993)  87 80 86 76 73 58 72 76 65 90 58 74 63  26.24 21.67 28.28 30.29 27.30 24.26 24.29 17.50 27.30 26.89 42.52 34.22 16.12  -0.021 -0.028 -0.018 -0.020 -0.018 -0.023 -0.022 -0.041 -0.020 -0.009 -0.019 -0.017 -0.042  Boersma et al. (1980) Boersma et al. (1980) Simons (1981) Vermeer et al. (1988) Ricklefs et al. (1985) Ricklefs et al. (1980) Ricklefs et al. (1985) Ricklefs and Schew (1994) Vermeer et al. (1988) Marks and Leasure (1992) Campos and Granadeiro (1999) Underwood and Bunce (2004) Underwood and Bunce (2004)  117 171 180 214 187 226 222 110 100 106 104 100 83 78 701 752 117 1084 746 353  33.44 31.73 33.19 21.94 27.39 19.06 25.36 34.26 34.61 37.33 28.84 38.66 44.31 41.13 20.37 16.55 18.58 18.87 30.91 25.89  -0.019 -0.018 -0.018 -0.027 -0.017 -0.020 -0.024 -0.020 -0.019 -0.019 -0.026 -0.019 -0.019 -0.019 -0.015 -0.027 -0.021 -0.017 -0.012 -0.017  Pettit et al. (1984a) Brown (1976a) Morris and Chardine (1992) Ramos et al. (2006) Ramos et al. (2006) Ramos et al. (2006) Megyesi and Griffin (1996) Surman and Wooller (1995) Ramos et al. (2006) Ramos et al. (2006) Ramos et al. (2006) Ramos et al. (2006) Ramos et al. (2006) Beintema (1997) Harris (1970a) Harris (1970a) Pettit et al. (1984a) Dunn and Brisbin (1980) Wilkens and Exo (1998) Schreiber and Schreiber (1980)  Source  48  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  Appendix 1. Continued Species  Larus audouini Larus californicus Larus fuscus Larus glaucescens Larus modestus Larus occidentalis Larus ridibundus Larus schistisagus Procelsterna cerulea Rissa brevirostris Rissa tridactyla  Sterna albifrons Sterna anaethetus Sterna caspia Sterna dougallii Sterna elegans Sterna fuscata  Area (Year) Columbretes Is, Spain (2000) Turkey (1974) California (1986) Farne Is, UK (1963) Mandarte Is, British Columbia (1978) Squab Is, Alaska (1979) Squab Is, Alaska (1980) Chile (1986) Farallon Is, California (1970) San Nicolas Is, California (1968) Germany (1986) The Netherlands (2002) Teuri Is, Japan (1984) Teuri Is, Japan (1985) Nihoa Is, Hawaii (1981) St George Is, Alaska (1993) Bleiksøy, Norway (1986) Farne Is, UK (1963) Middleton Is, Alaska (1996) Middleton Is, Alaska (1997) Newfoundland (1970) Norway (1973) Norway (1974) Norway (1976) Prince William Sound, Alaska (1996) Prince William Sound, Alaska (1997) Prince William Sound, Alaska (1998) Prince William Sound, Alaska (1999) St George Is, Alaska (1993) Portugal (2003) Great Barrier Reef (1980) Penguin Is, Australia (1990) California (1978) New Zealand (1993) Great Barrier Reef (1986) Rhode Is (1967) Rhode Is Sound (1990) California (1999) Hawaii (1972)  W∞ 620 743 897 717 1308 1326 2189 302 902 904 325 395 1612 1668 63 422 503 218 402 430 415 518 474 476 497 534 451 459 544 61 128 119 624 622 92 124 107 221 193  K  to  30.40 21.24 19.40 15.56 19.38 22.25 12.43 19.11 23.70 24.65 26.58 20.28 16.67 16.38 28.54 29.68 24.97 35.00 31.26 25.53 35.85 25.50 30.58 30.90 29.46 25.96 28.70 30.08 26.44 30.60 23.08 27.28 30.25 27.58 24.12 34.79 42.46 26.67 24.01  -0.007 -0.018 -0.010 -0.030 -0.013 -0.017 -0.026 -0.005 -0.016 -0.016 -0.018 -0.023 -0.015 -0.013 -0.008 -0.018 -0.017 -0.020 -0.018 -0.023 -0.018 -0.018 -0.017 -0.018 -0.017 -0.016 -0.018 -0.017 -0.017 -0.018 -0.031 -0.024 -0.015 -0.018 -0.029 -0.018 -0.018 -0.020 -0.025  Source Villuendas and Sarzo (2003) Witt (1977) Jehl et al. (1990) Pearson (1968) Verbeek and Morgan (1980) Murphy et al. (1984) Murphy et al. (1984) Guerra et al. (1988) Coulter (1979) Schreiber (1970) Nelsen and Brandl (1987) Müller et al. (2005) Watanuki (1992) Watanuki (1992) Rauzon et al. (1984) Lance and Roby (2000) Barrett (1989) Pearson (1968) Gill et al. (2002) Gill et al. (2002) Maunder and Threlfall (1972) Barrett and Runde (1980) Barrett and Runde (1980) Barrett and Runde (1980) Suryan et al. (2002) Suryan et al. (2002) Suryan et al. (2002) Suryan et al. (2002) Lance and Roby (2000) Paiva et al. (2006) Hulsman and Langham (1985) Garavanta and Wooller (2000) Schew et al. (1994) Barlow and Dowding (2002) Milton et al. (1996) LeCroy and Collins (1972) Nisbet et al. (1995) Dahdul and Horn (2003) Brown (1976b)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  49  Appendix 1. Continued Species  Sterna hirundo  Sterna paradisaea  Sterna sandvicensis Sterna sumatrana Sterna virgata  Area (Year) Bird Is, Massachusetts (1999) Couquet Is, UK (1966) Farne Is, UK (1963) Germany (1999) Machias Seal Is (1995) Machias Seal Is (1996) Machias Seal Is (1997) Machias Seal Is (1999) Machias Seal Is (2000) Machias Seal Is (2001) Machias Seal Is (2002) Machias Seal Is (2003) Québec (1983) Rhode Is (1967) The Netherlands (1989) The Netherlands (1990)  Farne Is, UK (1963) Machias Seal Is (1996) Machias Seal Is (1997) Machias Seal Is (1998) Machias Seal Is (2002) Québec (1983) Svalbard (1986) The Netherlands (1989) The Netherlands (1990) Farne Is, UK (1963) The Netherlands (1998) Great Barrier Reef (1986) Crozet Is (1982) S Shetland Is (1979) S Shetland Is (1981) S Shetland Is (1991)  W∞  K  to  136 181 86 150 149 185 127 142 125 109 134 170 145 102 108 124  37.74 27.70 38.96 33.83 31.18 23.99 33.96 36.34 34.90 41.60 33.91 26.11 37.28 35.24 39.27 37.44  -0.019 -0.017 -0.021 -0.019 -0.018 -0.017 -0.019 -0.018 -0.019 -0.019 -0.017 -0.016 -0.018 -0.020 -0.018 -0.017  Source Apanius and Nisbet (2006) Langham (1972) Pearson (1968) Becker and Wink (2003) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Chapdelaine et al. (1985) LeCroy and Collins (1972) Klaassen et al. (1994) Klaassen (1994)  73 90 119 151 104 125 143 103 140 114 245 133 94 166 216 159  38.42 44.12 40.63 27.39 40.25 38.74 34.68 40.63 37.90 43.36 34.80 26.55 44.17 32.50 34.48 32.91  -0.021 -0.019 -0.019 -0.017 -0.018 -0.018 -0.018 -0.018 -0.017 -0.022 -0.018 -0.018 -0.017 -0.017 -0.016 -0.018  Pearson (1968) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Bond et al. (2006) Chapdelaine et al. (1985) Klaassen et al. (1989) Klaassen et al. (1994) Klaassen (1994) Pearson (1968) Stienen and Brenninkmeijer (2002) Hulsman and Smith (1988) Weimerskirch and Stahl (1988) Jabłoński (1995) Jabłoński (1995) Klaassen (1994)  3812  20.72  -0.006  Schreiber (1976)  126 134  25.09 31.40  -0.028 -0.015  Jouventin et al. (1985) Jouventin et al. (1985)  Pelecanidae  Pelecanus occidentalis  Florida (1972)  Pelecanoides georgicus Pelecanoides urinatrix  Crozet Is (1982) Crozet Is (1982)  Pelecanoididae  50  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  Appendix 1. Continued Species Phaethontidae  Phaethon lepturus  Phaethon rubricauda  K  to  433 387 442 360 985 988 813 624 781 726 801 746  18.40 20.84 25.12 21.53 15.11 11.64 23.85 15.80 22.05 21.57 20.88 19.99  -0.029 -0.027 -0.012 -0.015 -0.031 -0.038 -0.016 -0.030 -0.015 -0.011 -0.014 -0.017  Diamond (1975) Diamond (1975) Schaffner (1990) Ramos and Pacheco (2003) Diamond (1975) Diamond (1975) Schreiber (1994) Schreiber (1994) Fleet (1974) Schreiber (1994) Schreiber (1994) Schreiber (1994)  E Bic Reef, Québec (1978) E Bicquette Is, Québec (1978) Grand Metis Is, Québec (1978) Shoals Is, New Hampshire (1972) SW Razade Reef, Québec (1978) W Bicquette Reef, Québec (1978) Chile (1997) S Africa (1993) Israel (2001)  1997 2446 2131 3188 2288 3462 1565 477 514  21.88 19.09 21.33 19.32 22.07 14.45 12.13 33.14 32.43  -0.013 -0.009 -0.011 -0.009 -0.011 -0.007 -0.007 -0.015 -0.012  DesGranges (1982) DesGranges (1982) DesGranges (1982) Dunn (1975) DesGranges (1982) DesGranges (1982) Kalmbach and Becker (2005) Kopij (1996) Shmueli et al. (2003)  Argentina (1993) Heard and McDonald Is (1993) S Georgia (1989) Greece (1994) Israel (2001) Bleiksøy, Norway (1986) Farne Is, UK (1963) Is May, UK (1998) Norway (1995)  2475 3312 2944 2735 2282 2712 1027 1854 2046  19.58 19.93 17.92 21.18 21.26 15.85 20.60 22.77 22.19  -0.012 -0.012 -0.027 -0.012 -0.004 -0.011 -0.013 -0.011 -0.007  Punta et al. (2003) Green (1997) Wanless and Harris (1993) Goutner et al. (1997) Shmueli et al. (2003) Barrett (1989) Pearson (1968) Daunt et al. (2001) Østnes et al. (2001)  Madeira (1995) Azores (1995) Portugal (1987) Selvagem Grande (1969) Selvagem Grande (1991) S Shetland Is (1992) Shetland Is, UK (1997) Shetland Is, UK (1997) Shetland Is, UK (1998)  142 1040 1042 895 977 582 959 879 993  34.45 22.59 25.59 20.83 61.46 20.61 29.38 26.20 25.12  -0.017 -0.016 -0.015 -0.026 -0.023 -0.026 -0.008 -0.017 -0.017  Nunes and Vicente (1998) Ramos et al. (2003) Granadeiro (1991) Zino (1971) Hamer and Hill (1993) Weidinger (1998) Phillips and Hamer (2000) Gray et al. (2003) Gray et al. (2003)  Area (Year) Aldabra Atoll (1968) Aldabra Atoll (1969) Puerto Rico (1986) Seychelles (2002) Aldabra Atoll (1968) Aldabra Atoll (1969) Christmas Is (1967) Christmas Is (1991) Green Is, Hawaii (1965) Johnston Atoll (1986) Johnston Atoll (1991) Johnston Atoll (1992)  W∞  Source  Phalacrocoracidae  Hypoleucos auritus  Hypoleucos brasiliensis Microcarbo africanus Microcarbo pygmaeus Notocarbo atriceps Phalacrocorax carbo Strictocarbo aristotelis  Procellariidae  Bulweria bulwerii Calonectris diomedea  Daption capense Fulmarus glacialis  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  51  Appendix 1. Continued Species  Fulmarus glacialoides Halobaena caerulea Lugensa brevirostris Macronectes giganteus Macronectes halli Pachyptila belcheri  Pachyptila desolata Pachyptila salvini Pachyptila turtur Pagodroma nivea Procellaria aequinoctialis Procellaria cinerea Pseudobulweria rostrata Pterodroma arminjoniana Pterodroma atrata Pterodroma axillaris Pterodroma hypoleuca Pterodroma incerta Pterodroma lessoni Pterodroma leucoptera Pterodroma macroptera Pterodroma mollis Pterodroma nigripennis Pterodroma phaeopygia Pterodroma pycrofti Puffinus assimilis Puffinus gravis Puffinus griseus Puffinus huttoni Puffinus opisthomelas  Area (Year) Prydz Bay, Antarctica (1989) Crozet Is (1982) Prince Edward Is (1983) Crozet Is (1982) Prince Edward Is (1980) Prince Edward Is (1977) Prince Edward Is (1977) Falkland Is (1978) Falkland Is (2003) Falkland Is (2004) Falkland Is (2005) S Georgia (1992) Crozet Is (1982) Prince Edward Is (1981) S Georgia (1983) Dronning Maud Land, Antarctica (1985) Prince Edward Is (1981) S Georgia (1986) Kerguelen Is (1988) Prince Edward Is (1982) New Caledonia (2004) Mauritius (1978) Pitcairn Is (1990) Chatham Is, New Zealand (1997) Midway Atoll, Hawaii (1981) Gough Is (2001) Kerguelen Is (1987) New South Wales, Australia (2001) Prince Edward Is (1980) Prince Edward Is (1982) Crozet Is (1982) Prince Edward Is (1980) Lord Howe Is, Australia (1990) Galápagos (1986) Galápagos (1966) Hawaii (1981) New Zealand (2001) Lord Howe Is, Australia (1989) New Zealand (1994) Gough Is (2001) New Zealand (1944) New Zealand (1999) Natividad Is, Mexico (1998)  W∞ 1059 222 198 308 356 4505 5194 239 263 181 155 225 164 158 182 322 1429 1496 1394 1247 583 513 379 328 239 762 108 279 444 621 295 341 237 536 423 540 227 222 278 1157 1147 507 395  K  to  28.59 28.72 34.51 19.76 29.66 15.30 14.43 29.44 19.12 29.15 31.60 35.35 19.51 36.08 33.60 17.51 20.36 18.99 13.14 17.31 15.52 22.83 25.86 26.44 24.75 5.89 16.13 17.07 20.41 12.84 18.60 22.44 21.31 18.03 22.04 19.84 50.45 28.83 24.13 21.12 11.50 23.17 25.96  -0.018 -0.019 -0.020 -0.052 -0.022 -0.008 -0.011 -0.016 -0.028 -0.018 -0.016 -0.013 -0.035 -0.020 -0.016 -0.036 -0.016 -0.013 -0.038 -0.015 -0.047 -0.024 -0.015 -0.013 -0.022 -0.118 -0.023 -0.038 -0.022 -0.037 -0.038 -0.024 -0.024 -0.032 -0.034 -0.018 -0.025 -0.014 -0.016 -0.020 -0.035 -0.023 -0.019  Source Norman and Ward (1992) Jouventin et al. (1985) Fugler et al. (1987) Jouventin et al. (1985) Schramm (1983) Cooper et al. (2001) Cooper et al. (2001) Strange (1980) Quillfeldt et al. (2007) Quillfeldt et al. (2007) Quillfeldt et al. (2007) Reid et al. (1999) Jouventin et al. (1985) Berruti and Hunter (1986) Prince and Copestake (1990) Røv (1990) Berruti et al. (1985) Hall (1987) Zotier (1990a) Newton and Fugler (1989) Villard et al. (2006) Gardner et al. (1985) de L. Brooke (1995) Gardner (1999) Pettit et al. (1982) Cuthbert (2004) Zotier (1990b) O’Dwyer et al. (2006) Schramm (1983) Newton and Fugler (1989) Jouventin et al. (1985) Schramm (1983) Hutton and Priddel (2002) Cruz and Cruz (1990) Harris (1970b) Simons (1985) Gangloff and Wilson (2004) Priddel et al. (2003) Booth et al. (2000) Cuthbert (2005) Richdale (1945) Cuthbert and Davis (2002) Keitt et al. (2003)  52  Life-history patterns in marine birds, Karpouzi, V., Pauly, D.  Appendix 1. Continued Species  Puffinus pacificus  Puffinus puffinus  Puffinus tenuirostris Thalassoica antarctica  Area (Year) Kilauea Point, Hawaii (1978) Kilauea Point, Hawaii (1979) Kilauea Point, Hawaii (1980) Manana Is, Hawaii (1978) Manana Is, Hawaii (1979) Manana Is, Hawaii (1984) Tern Is, Hawaii (1979) Faeroe Is (1981) Wales, UK (1995) Wales, UK (1996) Wales, UK (1999) Great Dog Is, Australia (1995) Dronning Maud Land, Antarctica (1984) Dronning Maud Land, Antarctica (1985) Dronning Maud Land, Antarctica (1992) Prydz Bay, Antarctica (1989)  W∞  K  to  489 479 456 427 441 476 503 427 559 525 680 930 640 1058 852 1057  14.00 21.89 18.10 19.19 18.88 14.50 15.86 28.02 23.71 22.96 20.55 12.85 23.89 15.57 22.14 30.53  -0.039 -0.025 -0.031 -0.029 -0.030 -0.044 -0.033 -0.015 -0.016 -0.018 -0.010 -0.032 -0.022 -0.027 -0.023 -0.017  Source Pettit et al. (1984b) Pettit et al. (1984b) Pettit et al. (1984b) Pettit et al. (1984b) Pettit et al. (1984b) Fry et al. (1986) Pettit et al. (1984b) Bech et al. (1982) Hamer and Hill (1997) Hamer et al. (1998) Gray et al. (2005) Hamer et al. (1997) Røv (1990) Haftorn et al. (1991) Lorentsen (1996) Norman and Ward (1992)  Crozet Is (2000) Heard and McDonald Is (1992) Prince Edward Is (1989) Macquarie Is (1956) Macquarie Is (1994) Macquarie Is (1995) Macquarie Is (1996) Prince Edward Is (1985) Prince Edward Is (1985) S Georgia (1986) S Georgia (1998) S Georgia (1999) S Georgia (2000) Penguin Is, Australia (1989) Penguin Is, Australia (1990) Penguin Is, Australia (1991) New Zealand (1937) New Zealand (1938) New Zealand (1940) New Zealand (1984) New Zealand (1985) New Zealand (1986)  5797 11010 10033 2786 2763 2551 2808 1902 2609 4739 4369 3502 4302 1018 1190 1242 6026 7563 5640 6543 6078 4184  12.22 9.38 7.56 17.35 14.87 11.67 14.96 20.26 15.57 14.56 11.89 18.00 14.00 25.46 23.54 21.77 13.37 9.74 13.10 14.11 13.76 17.69  -0.024 -0.007 -0.037 -0.010 -0.004 -0.013 -0.010 -0.015 -0.035 -0.017 -0.030 -0.015 -0.026 -0.016 -0.017 -0.016 -0.004 -0.020 -0.012 -0.003 -0.013 -0.010  de Margerie et al. (2004) Moore et al. (1998) van Heezik et al. (1993) Warham (1963) Hull et al. (2004) Hull et al. (2004) Hull et al. (2004) Brown (1987) Brown (1987) Williams (1990) Barlow and Croxall (2002) Barlow and Croxall (2002) Barlow and Croxall (2002) Wienecke et al. (2000) Wienecke et al. (2000) Wienecke et al. (2000) van Heezik (1991) van Heezik (1991) van Heezik (1991) van Heezik (1990) van Heezik (1990) van Heezik (1990)  Spheniscidae  Aptenodytes patagonicus Eudyptes chrysocome  Eudyptes chrysolophus  Eudyptula minor Megadyptes antipodes  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  53  Appendix 1. Continued Species  Pygoscelis adeliae  Pygoscelis antarctica  Pygoscelis papua Spheniscus demersus Spheniscus magellanicus  Area (Year) Humble Is, Antarctica (1989) Humble Is, Antarctica (1990) Lützow-Holm Bay, Antarctica (1989) Lützow-Holm Bay, Antarctica (1990) Lützow-Holm Bay, Antarctica (1991) Ross Is, Antarctica (1970) Torgersen Is, Antarctica (1989) Torgersen Is, Antarctica (1990) S Shetland Is (1980) S Shetland Is (1990) S Shetland Is (1991) S Shetland Is (1992) S Shetland Is (1993) S Shetland Is (1980) S Africa (1974) Argentina (1991) Argentina (1992) S Chile (1997)  W∞  K  to  4134 5749 5107 3983 2567 3420 3651 5190 3914 4058 4643 4621 3170 6739 1930 3840 5030 3667  17.47 8.67 16.52 20.84 32.00 22.97 16.77 9.57 22.55 22.33 17.05 19.06 21.54 13.55 14.62 12.63 9.53 18.18  -0.016 -0.072 -0.032 -0.021 -0.020 -0.015 -0.042 -0.064 -0.012 -0.020 -0.014 -0.015 -0.013 -0.012 -0.026 -0.018 -0.028 -0.008  Source Salihoglu et al. (2001) Salihoglu et al. (2001) Watanuki et al. (1992) Watanuki et al. (1992) Watanuki et al. (1992) Ainley and Schlatter (1972) Salihoglu et al. (2001) Salihoglu et al. (2001) Taylor (1985) Croll et al. (2006) Croll et al. (2006) Croll et al. (2006) Moreno et al. (1994) Taylor (1985) Cooper (1977) Frere et al. (1998) Frere et al. (1998) Radl and Culik (1999)  S Georgia (2001) S Georgia (2002) S Georgia (2003) Prydz Bay, Antarctica (1990) S Shetland Is (2001) E Greenland (1975)  2199 1808 1938 1726 1347 306  16.89 20.53 20.55 14.86 23.05 31.88  -0.013 -0.012 -0.011 -0.030 -0.012 -0.015  Phillips et al. (2004) Phillips et al. (2004) Phillips et al. (2004) Wang and Norman (1993) Ritz et al. (2005) de Korte (1986)  Baccalieu Is, Newfoundland (1979) Magdalen Is, Québec (1979) Québec (1965) Scotland, UK (1962) Scotland, UK (1976) S Africa (1967) S Africa (1974) S Africa (1988) Victoria, Australia (1995) Victoria, Australia (1999)  4123 4477 4708 4746 4732 3390 3671 3461 3668 3457  17.54 15.48 15.15 15.39 15.33 15.81 14.81 15.41 15.82 16.58  -0.006 -0.011 -0.011 -0.008 -0.008 -0.009 -0.007 -0.009 -0.006 -0.007  Montevecchi et al. (1984) Kirkham and Montevecchi (1982) Poulin (1968) Nelson (1964) Wanless (1984) Jarvis (1974) Cooper (1978) Navarro (1991) Gibbs et al. (2000) Bunce (2001)  Ascension Is (1960) Kure Atoll, Hawaii (1965) Lord Howe Is, Australia (2002) Galápagos (1964) Lobos de Tierra Is, Peru (1979) Galápagos (1963)  1952 2107 2260 1939 1669 956  17.60 18.25 17.74 14.74 19.54 9.63  -0.009 -0.001 -0.011 -0.005 -0.020 -0.014  Dorward (1962) Kepler (1969) Priddel et al. (2005) Duffy and Ricklefs (1981) Duffy and Ricklefs (1981) Nelson (1969)  Stercorariidae  Catharacta antarctica Catharacta maccormicki Stercorarius longicaudus  Sulidae  Morus bassanus  Morus capensis Morus serrator Sula dactylatra Sula nebouxii Sula sula  54  Growth of marine reptiles, Palomares, M.L.D., et al.  GROWTH OF MARINE REPTILES 1 M.L. Deng Palomares  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email:m.palomares@fisheries.ubc.ca  Christine Dar  The SeaLifeBase Project, WorldFish Center, Khush Hall, IRRI, Los Baños, Laguna, Philippines; Email: c.dar@cgiar.org  Gary Fry  CSIRO Marine and Atmospheric Research, PO Box 120, Cleveland Qld Australia 4163; Email: gary.fry@csiro.au  ABSTRACT Growth data were obtained from the scientific literature and re-expressed according to the von Bertalanffy Growth Function (VBGF) using a variety of methods. This resulted in 103 population estimates of VBGF parameters for 27 species of marine reptiles, i.e., marine iguana, saltwater crocodile, 8 species of sea turtles and 16 species of aquatic snakes. A frequency distribution of the growth performance index (Ф’) values indicate two peaks, i.e., index values at 2.75 for sea turtles and marine iguana and at 3.75 for sea snakes, while saltwater crocodiles have index values at the tail of the distribution (4.75-5.25). The auximetric plot of log10K against log10W∞ indicates that like marine mammals, seabirds and invertebrates, marine reptiles exhibit the same growth patterns as those of fish and thus, their growth can be expressed according to the VBGF.  INTRODUCTION Marine reptiles consist of 77 species belonging to 4 major groups, i.e., marine iguana (1 species), saltwater crocodile (1), sea turtles (8) and sea snakes (67; Kharin, 2008, see Bell, 1843). Sea turtles are circumglobal whereas sea snakes are distributed mostly around the eastern Indo-Pacific region. The marine iguana, Crocodylus porosus, is endemic to the Galapagos Islands (Kruuk & Snell, 1981). The crocodile, Amblyrhynchus cristatus, is the only crocodile inhabiting marine and freshwaters in the Indo-Pacific (Mead et al., 2002). There are 9 reptile species included in the IUCN Red List of Threatened Species (2007), which include 7 species of sea turtles, i.e., 3 are listed as critically endangered (Dermochelys coriacea, Eretmochelys imbricata, Lepidochelys kempii), 3 as endangered (Caretta caretta, Chelonia mydas, Lepidochelys olivacea) and Natator depressus as data defficient, the marine iguana, Amblyrhynchus cristatus is listed as vulnerable, and the Atlantic salt marsh snake, Nerodia clarkii as of least concern (see also Ineich & Laboute 2002). These 9 species represent 12% of all species of marine reptiles existing in the world, which is a high percentage; it is due to the fact that these animals grow to large sizes and have slower metabolic and turn over rates. In spite of reptiles’ vulnerability to extinction, studies on reptilian life history are few and usually serological in nature (e.g., Rogers, 1902-1903) and are usually on terrestrial species, e.g., on sexual dimorphism and diet (e.g., Shine et al., 2002; Camilleri & Shine, 1990), reproductive strategies (e.g., Shine, 1988; Lemen & Voris, 1981), body size (e.g., Boback & Guyer, 2003) and patterns of growth (e.g., Shine & Charnov, 1992). Most studies focusing on sea snakes lack discussions on the growth of these  1 Cite as: Palomares, M.L.D., Dar, C., Fry, G., 2008. Growth of marine reptiles. In: Palomares, M.L.D., Pauly, D. (Eds.), Von Bertalanffy Growth Parameters of Non-fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 54-81.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  55  animals and even the review of sea snake biology by Dunson (1975), though comprehensive, did not include growth. Recent interest, notably on the effect of trawl fisheries to sea snake populations, e.g., in Australia (Ward, 2001; Fry et al., 2001; Ward, 2000) indicate a need for growth parameters for sea snakes. In response to this, growth data were compiled from various published references. In addition, these were used to confirm that the von Bertalanffy Growth Function (VBGF) can indeed describe the growth of marine reptiles as suggested for snakes and lizards in Shine & Charnov (1992) and for sea turtles in Jones et al. (2008) even though they have body shapes, i.e., elongate or box-like, different from fish, for which the VBGF has been largely used. The data assembled here and the VBGF parameters estimated will be made available via the online information system, SeaLifeBase (www.sealifebase.org) in the hope that they could be used in the assessment and management of marine reptilian stocks.  MATERIAL AND METHODS Growth parameter estimation Growth data for species in their natural environment (wild) representing a reasonable range of sizes were assembled from various published literature, i.e., (i) growth parameter estimates; (ii) length-at age or growth increment data; and (iii) length-frequency distributions (carapace length (CL) for sea turtles and snout-vent length (SVL) for sea snakes, marine crocodile and iguana). Data in (i), if expressed in functions other than the von Bertalanffy Growth Function (VBGF), e.g., Gompetz and logistic curves, were reexpressed as VBGF parameters. Data in (ii) were fitted directly to the von Bertalanffy growth function (von Bertalanffy, 1957):  Lt = L∞(1-e-K(t-t0))  … 1)  where Lt is the length at age t, L∞ is the asymptotic length, K is a growth coefficient (growth rate towards the maximum), and t0 is the age at which length is zero. Data in (iii) were fitted to the Powell-Wetherall Plot (PW-Plot; see Pauly, 1998; Wetherall, 1986; Powell, 1979) to estimate L∞, based on the assumption that the resulting distribution is representative of the population. The PW-Plot consists of pointer parameters, i.e., mean (Lmean) and cut off (Li) lengths. A series of mean lengths (Lmean), computed from successive cut-off lengths (Li+1), minus the Li (i.e., Lmean-Li) are plotted against Li. The resulting downward trend of points were fitted to a linear regression where L∞ is estimated as a/-b and Z/K as (1+b)/(-b), Z being the total instantaneous mortality of an exploited population, and conversely, is equivalent to M (natural mortality) if the population being studied is not exploited (see below). In cases where the data in (iii) were comprised of successive length-frequency distributions, the data were fitted to the VBGF using a non-parametric, robust approach known as ELEFAN (Pauly, 1987; 1998), implemented in the FiSAT software package (Gayanilo et al., 1996). In cases where only L∞ estimates were available, e.g., results of the PW-Plot, values of K were obtained using the growth performance index (Ф’) defined by Pauly & Munro (1984) as Ф’ = log10 K + 2·log10L∞, and mean values of Ф’, available from L∞ and K pairs for: (a) the same species in different localities; (b) other species in the same genus; (c) other species in the same family. Estimates of K obtained in this fashion are marked as such in SeaLifeBase and thus can be ignored when only independent estimates are sought. Asymptotic weight estimation Asymptotic weight, W∞, was estimated using the length-weight relationship of the form  W = a · Lb  … (2)  where a is a multiplicative term equivalent to the y-intercept of the log-log transformed linear regression, L the asymptotic length, and b the isometric weight growth parameter, equivalent to the slope of the regression. In cases where sufficient length-weight data pairs were not available for linear regression analyses, condition factors (c.f.) using individual length-weight pairs were estimated with  56  Growth of marine reptiles, Palomares, M.L.D., et al.  c.f. = W · 100/L3, where W is the weight in grams, and L the length in centimeters (Pauly, 1984b). The value of the length-weight parameter a was then obtained as a = c.f./100, assuming that b=3. All lengths are expressed in centimeters and weights in grams. Mortality estimation The total instantaneous mortality (Z) of a given population is defined as:  Nt2 = Nt1·e-Z·(t2-t1)  … 3)  where Nt1 and Nt2 is the population size at time t1 and t2, respectively (Koch et al., 2007). The parameter Z is the sum of natural mortality (M) and fishing mortality (F). As marine reptiles are exploited, either by a target fishery or as by-catch, we can assume that the mortalities inferred from equation (3) refer to total mortality. The data in (iii), as discussed above, were plotted with the Powell-Wetherall Plot to infer Z/K assuming that the samples are representative of the population in the juvenile and adult phases (Wetherall, 1986; Wetherall et al., 1987). Where applicable, length-frequency samples were converted to catch curves, using the growth parameters obtained from FiSAT ((Pauly, 1987; 1998; Gayanilo et al., 1996) to obtain estimates of total mortality. All mortalities are expressed in years-1.  RESULTS AND DISCUSSION Growth data were found for 92 populations of 26 marine reptile species. Sea turtles represent half of this available data, perhaps due to the fact that they are endangered and thus the need to study and understand their biology instigates baseline studies. Sea snakes, the other group which is well-represented in this study, are an important by-catch in Australian trawl fisheries protected under Schedule 1 of the National Parks and Wildlife Regulations since 1994 (Milton, 2005; 2001) and are thus the subject of research programs in Australia. Here, survey data, graciously provided by the Australian Fisheries Research and Development Corporation and Commonwealth Scientific and Industrial Research Organisation in collaboration with fishers from the Australian Northern Prawn Fishery were used to obtain length-frequency distributions analyzed with ELEFAN to estimate L∞ for 13 species of sea snakes from the Gulf of Carpentaria. Table 1. Summary of marine reptile growth data obtained from the scientific literature. Details of growth data are included in Table A1. Order  Family  Crocodilia Squamata  Crocodylidae Acrochordidae Colubridae Hydrophiidae Iguanidae Cheloniidae Dermochelyidae ‐   Testudines Total  No. spp 1 1 1 15 1 7 1 27  No. stocks 3 2 2 20 4 58 3 92  L∞ K  L∞ Z/K  L/W  c.f.  Z  Lm  2 2 2 10 1 26 1 43  2 21 2 31 3 69  1 1 36 11 3 52  2 102 5 4 5 103  1 2 4 1 4 12  53 2 55  The results of this study, summarized in Table 1, show that life history data on marine reptiles do exist, though not standardized in a format that could be readily used for management purposes. The standardization performed here included the following: (i) converting length units in centimeters (cm), weight units in grams (g) and age units in years; (ii) expressing lengths in the same length type, i.e., snoutvent length for most marine reptiles and carapace length for sea turtles; (iii) re-expressing growth through the VBGF; and (iv) converting L/W relationships to cm and g. All conversions were straighforward except for item (ii), notably for sea turtles where curved and straight carapace lengths (CCL and SCL, respectively) are used. Empirical equations based on simple linear regression of paired SCL and CCL data were adapted from Teas (1993; p. 3) and used to convert values of CCL∞ to SCL∞ (see details in Table A1) and vice versa depending on the length type used in the length-weight relationships. The frequency histogram of the L/W relationship coefficient b values obtained for 52 populations using linear regressions of length and weight data pairs (Figure 1) shows a clear peak at size class b=3 (median  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  57  value is 2.96 and the mode is at 3.00; s.d.=0.36; sample variance of 0.13). This suggests that marine reptiles grow like marine mammals (see Palomares et al., 2008), seabirds (see Karpouzi & Pauly, 2008) and fish (see Carlander, 1969; 1977).  -1  K (year ; log10)  Frequency  The asymptotic lengths obtained 16 ranged from 28.6 cm (Amblyrhynchus cristatus, 14 Sea turtles and sea snakes Galapagos Islands) to 323 cm (Crocodylus porosus, Northern 12 Saltwater crocodile, Territory, Australia). Sea snakes sea turtles and sea snakes 10 ranged in size from 66.8 cm (Emydocephalus ijimae, 8 Zamamijima, Ryukyu Island) to 257 cm (Hydrophis elegans, Gulf 6 of Carpentaria, Australia) while sea turtles ranged in size from 4 56.2 cm (Lepidochelys kempii, Sambine Pass, Gulf of Meixco, 2 USA) to 168 cm (Chelonia 0 mydas, Great Inagua, Bahamas). 2.00 2.25 2.50 2.75 3.00 3.25 3.50 3.75 The auximetric grid plotting L/W relationship coefficient 'b' log K against log W∞ (Figure 2) indicates that marine iguanas Figure 1. Distribution of length-weight relationship coefficient b of 53 grow similarly to sea snakes, populations of marine reptiles (see TableA1 for details). while saltwater crocodiles, though clearly a group apart, grow more similarly to sea turtles. Note however, three outliers, i.e., Chelonia mydas (Table A2, 21g), Lepidochelys kempii (see Table A2, 24a) and Dermochelys coriacea (Table A2, 27d). Though the sample size range (26-72 cm) of the C. mydas population is wide enough to include juveniles and adults, this range probably represent sub-adult populations given that the largest length in the sample is only 70% of Lmax (Lmax = 105 cm CL; Schneider, 1990) and only 43% of the largest L∞ reported for this species (i.e., 168 cm SCL; see Table A2, 21k). The same could be argued for the L. kempii population (sample length range=20-60 cm) which grow to Lmax=75 cm CL (Carr and Caldwell, 1956). However, note that the growth parameters for this population were obtained from a single length-frequency histogram using the Powell-Wetherall Plot and K from Ф’. The last sea turtle 1.0 Lepidochelys kempii outlier, D. coriacea came Chelonia mydas (24a) Sea snakes (21g) from measurements of 0.5 Marine iguanas turtles sampled from tropical Saltwater areas (see Jones et al., 2008) crocodiles 0.0 and reared in captivity in Dermochelys Vancouver, Canada to more coriacea (27d) -0.5 than 2 years of age, i.e., to only 60% of the recorded Lmax (257 cm CL; Márquez, -1.0 1990). The growth Astrotia stokesii (8a) parameters of the outlier sea -1.5 snake population of Astrotia Sea turtles stokesii were obtained from -2.0 survey samples of the -2 -1 -1 0 1 1 2 2 3 3 AFRDC, CSIRO and NPF W∞ (kg; log10) (Australia) and the PowellWetherall Plot. Though the Figure 2. Auximetric plot of von Bertalanffy growth parameters for 92 L∞ estimate may be viable, populations of 26 species of marine reptiles (see Table A2 for details). Note the K estimate, obtained similarity of growth performance of sea snakes with marine iguanas and saltwater from the average Ф’ for crocodiles with sea turtles. The 3 outlier populations of sea turtles are based on species in the family length-frequency samples with narrow length ranges, i.e., juveniles, while the outlier snake population’s K was estimated from the average Φ' of species in the family Hydrophiidae.  58  Growth of marine reptiles, Palomares, M.L.D., et al.  Hydrophiidae and is not an independent estimate.  25  20 Frequency  Average values of Z/K for sea snakes and sea turtles are 2.07 (s.e. = 0.231) and 1.4 (s.e. = 0.115), respectively (see Figure 3). These values are comparable with those reported for fishes, i.e., 1.00-2.00 (Beverton and Holt 1956, Pauly 1998). The Z/K values available for marine iguanas are 1.24 and 1.76, also within the range given for fishes. Values of natural mortality for 12 species of marine reptiles ranged from 0.16 (Chelonia mydas, Bahia Magdalena, Mexico) to 4.83 (Amblyrhynchus cristatus, Genovesa, Galapagos Island).  30  15  10  5  0 1  2  3  4  5  Z/K  Figure 3. Frequency distribution of Z/K values obtained for sea snakes and sea turtles with average values of 2.07 (s.e. = 0.231) and 1.4 (s.e. = 0.115), respectively (see Table A2 for details).  Length at maturity assembled for sea snakes ranged from 42.5 cm (Thalassophis anomalus, Sourabaya, Java, Indonesia) to 145 cm (Disteira kingii, northern Australia). This data set was used with the available growth parameter estimates for the same population to obtain a frequency histogram of the reproductive load for sea snakes, i.e., Lm/L∞ values (Figure 4), which ranged from 0.400 (Astrotia stokesii, northern Australia) to 0.832 (Disteira kingii, northern Australia), with a mean value of 0.582 (s.e. = 0.0221). Only two populations of sea turtles, i.e., Lepidochelys kempii, have available Lm data and for which reproductive load were calculated (see Figure 4). Figure 4 emulates the negative trend found for fish (Pauly 1984a) and reiterates the result that marine reptiles grow like fish.  ACKNOWLEDGEMENTS  120  100 Reproductive load (Lm*100/L∞)  This study is the first to compile data on growth of marine reptiles. Only 16 percent of the total species stated in the introduction were covered in this study. However, more work is necessary in order to further understand the biology of marine reptiles and help prevent marine reptile species, e.g., sea turtles, from being farther endangered.  Lepidochelys kempii  80  60  40  20  We wish to thank the 0 Australian Fisheries Research 0.0 0.5 1.0 1.5 2.0 2.5 3.0 and Development Corporation, Asymptotic length (log; cm) the Commonwealth Scientific and Industrial Research Figure 4. Reproductive load plotted against asymptotic length of 35 Organisation and the Northern populations of sea snakes (12 species) and 2 populations of sea turtles, Lepidochelys kempii. Note negative trend emulating what has been found for Prawn Fishery fishers for fish (see details in Table A3). providing the sea snake survey data used here. 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Carbone, F. Trillmich., 1996. Lekking in marine iguanas, female grouping and male reproductive strategies. Anim. Behav. 52, 581-596. Wikelski, M., B. Gall, F. Trillmich., 1993. Ontogenetic changes in food intake and digestion rate of the herbivorous marine iguana (Amblyrhynchus cristatus, Bell). Oecologia 94, 373-379. Wikelski, M., F. Trillmich., 1997. Body size and sexual dimorphism in marine iguanas fluctuate as a result of opposing natural and sexual selection, an island comparison. Evolution 51(3), 922-936.  62  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A1. Length-weight data for marine reptiles used to obtain Figure 1. TL=total length; SVL=snout-vent length; SCL=straight carapace length; CCL=curved carapace length; CL=carapace length. Spec. No. 1  Stock No. a  Species  Locality  n  Sex  Type  b  a  c.f.  r2  Method/Source  Crocodylus porosus  Cape York Peninsula, Australia  11  M  TL  3.60  0.0001  -  0.998  Grigg et al. (1998; Tab. 1 p. 1793).  Acrochordus granulatus  Phangnga Bay, Thailand  45  F  SVL  3.00  0.0005  0.0521  -  Phangnga Bay, Thailand  19  M  SVL  3.00  0.0004  0.0384  -  Cerberus rynchops  Muar River, Malaysia  14  unsexed  SVL  3.01  0.0006  -  0.992  Acalyptophis peronii  East coast, northern Australia  -  unsexed  SVL  3.00  0.0011  0.1095  -  b  Groote, northern Australia  1  M  SVL  3.00  0.0008  0.0797  -  c  Gulf of Carpentaria, Australia  22  F  SVL  3.29  0.0002  -  0.974  d  Gulf of Carpentaria, Australia  24  M  SVL  2.70  0.0028  -  0.937  e  Gulf of Carpentaria, Australia  50  unsexed  SVL  3.00  0.0007  -  0.851  f  Mornington, northern Australia  -  mixed  SVL  3.00  0.0007  0.0670  -  g  Weipa, northern Australia  9  mixed  SVL  3.00  0.0008  0.0765  -  a from c.f. of data from Wangkulangkul et al. (2005; Fig. 2, p. 259). a from c.f. of data from Wangkulangkul et al. (2005; Fig. 2, p. 259). Jayne et al. (1988; Tab. 5, p. 10). Results maybe biased because N is small. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from mean c.f. of survey data from AFRDC, CSIRO, NPF. a from mean c.f. of survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59).  (Crocodilla, Crocrodylidae) 2  a  (Squamata, Acrochordidae) b 3  a  (Squamata, Colubridae) 4  a  (Squamata, Hydrophiidae)  5  a  Aipysurus apraefrontalis  northwestern Shelf, Australia  1  M  SVL  3.00  0.0007  0.0700  -  6  a  Aipysurus duboisii  East coast, northern Australia  -  M  SVL  3.00  0.0008  0.0814  -  b  Groote, northern Australia  3  F  SVL  3.00  0.0006  0.0612  -  c  Gulf of Carpentaria, Australia  8  F  SVL  3.00  0.0006  -  -  d  Gulf of Carpentaria, Australia  11  M  SVL  3.00  0.0005  -  -  e  Gulf of Carpentaria, Australia  20  unsexed  SVL  2.90  0.0009  -  0.720  f  Mornington, northern Australia  2  M  SVL  3.00  0.0006  0.0583  -  g  Weipa, northern Australia  3  mixed  SVL  3.00  0.0005  0.0486  -  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  63  Table A1. Continued. Spec. No. 7  8  8  9  Stock No. a  Species  Aipysurus eydouxii  Locality East coast, northern Australia  n  Sex  Type  b  a  c.f.  r2  -  F  SVL  3.00  0.0012  0.1156  -  b  Groote, northern Australia  12  mixed  SVL  3.00  0.0012  0.1236  -  c  Gulf of Carpentaria, Australia  75  F  SVL  2.48  0.0099  -  0.843  Gulf of Carpentaria, Australia  28  F  SVL  3.00  0.0008  0.0845  -  e  Gulf of Carpentaria, Australia  24  M  SVL  2.50  0.0096  -  0.786  f  Gulf of Carpentaria, Australia  30  M  SVL  3.00  0.0010  0.0973  -  g  Gulf of Carpentaria, Australia  104  unsexed  SVL  2.60  0.0061  -  0.866  h  Mornington, northern Australia  -  mixed  SVL  3.00  0.0014  0.1364  -  i  Weipa, northern Australia  18  F  SVL  3.00  0.0012  0.1233  -  East coast, northern Australia  -  unsexed  SVL  3.00  0.0010  0.1008  -  b  Groote, northern Australia  7  mixed  SVL  3.00  0.0015  0.1485  -  c  Gulf of Carpentaria, Australia  36  F  SVL  3.62  0.0001  -  0.954  d  Gulf of Carpentaria, Australia  19  F  SVL  3.00  0.0013  0.1281  -  e  Gulf of Carpentaria, Australia  36  M  SVL  3.00  0.0011  -  0.881  f  Gulf of Carpentaria, Australia  12  M  SVL  3.00  0.0012  0.1233  -  Gulf of Carpentaria, Australia  74  unsexed  SVL  3.52  0.0001  -  0.900  h  Mornington, northern Australia  2  F  SVL  3.00  0.0013  0.1316  -  i  Torres Strait, Australia  1  M  SVL  3.00  0.0011  0.1136  -  j  Weipa, northern Australia  14  mixed  SVL  3.00  0.0012  0.1188  -  Darwin, northern Australia  1  M  SVL  3.00  0.0012  0.1231  -  East coast, northern Australia  -  unsexed  SVL  3.00  0.0011  0.1081  -  a  g  a b  Aipysurus laevis  Aipysurus laevis  Astrotia stokesii  Method/Source a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59).  64  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 9  10  11  Stock No. c  n  Sex  Type  b  a  c.f.  r2  Gulf of Carpentaria, Australia  71  F  SVL  3.58  0.0001  -  0.895  d  Gulf of Carpentaria, Australia  16  F  SVL  3.00  0.0015  0.1547  -  e  Gulf of Carpentaria, Australia  57  M  SVL  3.07  0.0008  -  0.856  f  Gulf of Carpentaria, Australia  10  M  SVL  3.00  0.0010  0.1016  -  g  Gulf of Carpentaria, Australia  128  unsexed  SVL  3.58  0.0001  -  0.881  h  Mornington, northern Australia  21  mixed  SVL  3.00  0.0013  0.1327  -  i  Weipa, northern Australia  33  mixed  SVL  3.00  0.0011  0.1149  -  East coast, northern Australia  2  mixed  SVL  3.00  0.0002  0.0150  -  b  Gulf of Carpentaria, Australia  27  F  SVL  3.09  0.0001  -  0.946  c  Gulf of Carpentaria, Australia  23  F  SVL  3.00  0.0002  0.0173  -  d  Gulf of Carpentaria, Australia  14  M  SVL  2.38  0.0046  -  0.810  e  Gulf of Carpentaria, Australia  12  M  SVL  3.00  0.0002  0.0169  -  f  Gulf of Carpentaria, Australia  47  unsexed  SVL  3.00  0.0002  -  0.899  g  Mornington, northern Australia  -  mixed  SVL  3.00  0.0002  0.0246  -  h  Torres Strait, Australia  -  F  SVL  3.00  0.0002  0.0233  -  i  Weipa, northern Australia  -  mixed  SVL  3.00  0.0003  0.0260  -  East coast, northern Australia  1  M  SVL  3.00  0.0050  0.5011  -  b  East coast, northern Australia  -  mixed  SVL  3.00  0.0006  0.0618  -  c  Groote, northern Australia  -  mixed  SVL  3.00  0.0006  0.0618  -  d  Gulf of Carpentaria, Australia  153  F  SVL  2.40  0.0101  -  0.710  e  Gulf of Carpentaria, Australia  94  F  SVL  3.00  0.0006  0.0553  -  f  Gulf of Carpentaria, Australia  84  M  SVL  2.64  0.0031  -  0.815  a  a  Species  Astrotia stokesii  Disteira kingii  Disteira major  Locality  Method/Source Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  65  Table A1. Continued. Spec. No. 11  Stock No. g  n  Sex  Type  b  a  c.f.  r2  Method/Source  Gulf of Carpentaria, Australia  55  M  SVL  3.00  0.0005  0.0508  -  h  Gulf of Carpentaria, Australia  240  unsexed  SVL  2.54  0.0052  -  0.765  i  northwest Australia  3  Unsexed  SVL  3.00  0.0006  -  -  j  northwest Australia  1  Female  SVL  3.00  0.0007  0.0690  -  k  northwest Australia  2  Male  SVL  3.00  0.0006  -  -  l  Torres Strait, Australia  1  F  SVL  3.00  0.0009  0.0880  -  Weipa, northern Australia  17  mixed  SVL  3.00  0.0006  0.0579  -  a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of survey data from AFRDC, CSIRO, NPF. Masunaga et al. (2003; Fig. 2 & 4). a, p. 464 & 467). Masunaga et al. (2003; Fig. 2 & 4). b, p. 464 & 466). Survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from mean c.f. of survey data from AFRDC, CSIRO, NPF. a from mean c.f. of survey data from AFRDC, CSIRO, NPF. a from mean c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59).  Species  Disteira major  m  Locality  12  a  Emydocephalus annulatus  Gulf of Carpentaria, Australia  1  M  SVL  3.00  0.0010  0.1032  -  13  a  Emydocephalus ijimae  Zamamijima, Ryukyu Island  58  F  SVL  2.94  0.0012  -  0.976  Zamamijima, Ryukyu Island  52  M  SVL  2.53  0.0050  -  0.970  Gulf of Carpentaria, Australia  33  F  SVL  3.26  0.0002  -  0.922  b  Gulf of Carpentaria, Australia  24  M  SVL  3.15  0.0003  -  0.884  c  Gulf of Carpentaria, Australia  69  unsexed  SVL  3.33  0.0001  -  0.9400  d  Mornington, northern Australia  -  mixed  SVL  3.00  0.0006  0.0556  -  e  Weipa, northern Australia  39  mixed  SVL  3.00  0.0006  0.0621  -  Gulf of Carpentaria, Australia  2  F  SVL  3.00  0.0005  -  -  b  Gulf of Carpentaria, Australia  5  M  SVL  3.00  0.0057  -  -  c  Gulf of Carpentaria, Australia  7  unsexed  SVL  3.00  0.0006  -  -  d  Mornington, northern Australia  2  M  SVL  3.00  0.0005  0.0545  -  e  Weipa, northern Australia  5  mixed  SVL  3.00  0.0006  0.0621  -  b 14  15  a  a  Enhydrina schistosa  Hydrophis caerulescens  16  a  Hydrophis czeblukovi  northwestern Shelf, Australia  1  F  SVL  3.00  0.0008  0.0817  -  17  a  Hydrophis elegans  East coast, northern Australia  -  mixed  SVL  3.00  0.0004  0.0353  -  66  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 17  Stock No. b  Species  Hydrophis elegans  Locality Groote, northern Australia  n  Sex  Type  b  a  c.f.  r2  -  mixed  SVL  3.00  0.0004  0.0444  -  c  Gulf of Carpentaria, Australia  230  F  SVL  3.26  0.0001  -  0.898  d  Gulf of Carpentaria, Australia  231  F  SVL  3.00  0.0003  0.0314  -  e  Gulf of Carpentaria, Australia  207  M  SVL  3.01  0.0003  -  0.915  f  Gulf of Carpentaria, Australia  283  M  SVL  3.00  0.0003  0.0293  -  g  Gulf of Carpentaria, Australia  490  unsexed  SVL  3.17  0.0001  -  0.929  h  Mornington, northern Australia  -  mixed  SVL  3.00  0.0003  0.0308  -  i  northwest Australia  6  Unsexed  SVL  3.00  0.0003  -  -  j  northwest Australia  3  Female  SVL  3.00  0.0003  -  -  k  northwest Australia  3  Male  SVL  3.00  0.0003  -  -  l  Weipa, northern Australia  -  mixed  SVL  3.00  0.0003  0.0291  -  18  a  Hydrophis inornatus  Gulf of Carpentaria, Australia  1  F  SVL  3.00  0.0005  0.0525  -  19  a  Hydrophis macdowelli  Gulf of Carpentaria, Australia  11  F  SVL  3.22  0.0002  -  0.904  b  Gulf of Carpentaria, Australia  3  M  SVL  3.00  0.0005  -  -  c  Gulf of Carpentaria, Australia  14  unsexed  SVL  2.97  0.0007  -  0.846  d  Mornington, northern Australia  7  mixed  SVL  3.00  0.0006  0.0550  -  e  northwestern Shelf, Australia  1  M  SVL  3.00  0.0005  0.0478  -  f  Weipa, northern Australia  1  F  SVL  3.00  0.0006  0.0637  -  East coast, northern Australia  -  unsexed  SVL  3.00  0.0010  0.1018  -  b  Groote, northern Australia  -  mixed  SVL  3.00  0.0010  0.0976  -  c  Gulf of Carpentaria, Australia  73  F  SVL  2.88  0.0016  -  0.640  20  a  Hydrophis ornatus  Method/Source a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. a from mean c.f. of survey data from AFRDC, CSIRO, NPF. Results maybe biased because N is small. Survey data from AFRDC, CSIRO, NPF. Results maybe biased because N is small. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  67  Table A1. Continued. Spec. No. 20  21  22  Stock No. d  n  Sex  Type  b  a  c.f.  r2  Gulf of Carpentaria, Australia  42  F  SVL  3.00  0.0008  0.0765  -  e  Gulf of Carpentaria, Australia  82  M  SVL  2.23  0.0316  -  0.5610  f  Gulf of Carpentaria, Australia  45  M  SVL  3.00  0.0007  0.0699  -  g  Gulf of Carpentaria, Australia  166  unsexed  SVL  2.51  0.0085  -  0.568  h  Mornington, northern Australia  -  mixed  SVL  3.00  0.0007  0.0701  -  i  northwestern Shelf, Australia  2  M  SVL  3.00  0.0008  0.0756  -  j  Torres Strait, Australia  -  F  SVL  3.00  0.0006  0.0625  -  k  Weipa, northern Australia  -  mixed  SVL  3.00  0.0007  0.0656  -  Gulf of Carpentaria, Australia  24  F  SVL  3.00  0.0003  -  -  b  Gulf of Carpentaria, Australia  8  M  SVL  3.00  0.0004  -  -  c  Gulf of Carpentaria, Australia  32  unsexed  SVL  2.45  0.0053  -  0.758  d  Mornington, northern Australia  4  mixed  SVL  3.00  0.0004  0.0393  -  Darwin, northern Australia  1  mixed  SVL  3.00  0.0016  0.1592  -  a  a  Species  Hydrophis ornatus  Hydrophis pacificus  Lapemis hardwickii  Locality  b  East coast, northern Australia  70  mixed  SVL  3.00  0.0016  0.1637  -  c  Groote, northern Australia  7  mixed  SVL  3.00  0.0013  0.1292  -  d  Gulf of Carpentaria, Australia  309  F  SVL  2.99  0.0011  -  0.903  e  Gulf of Carpentaria, Australia  220  F  SVL  3.00  0.0013  0.1297  -  f  Gulf of Carpentaria, Australia  175  M  SVL  2.82  0.0024  -  0.927  g  Gulf of Carpentaria, Australia  177  M  SVL  3.00  0.0012  0.1156  -  h  Gulf of Carpentaria, Australia  535  unsexed  SVL  2.97  0.0012  -  0.933  i  Mornington, northern Australia  -  mixed  SVL  3.00  0.0012  0.1190  -  j  northwest Australia  1  Male  SVL  3.00  0.0009  -  -  Method/Source a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from mean c.f. of survey data from AFRDC, CSIRO, NPF. a from mean c.f. of survey data from AFRDC, CSIRO, NPF. Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Ward (2000; Tab. 2, p. 158). Survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of survey data from AFRDC, CSIRO, NPF.  68  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 22  Stock No. k  n  Sex  Type  b  a  c.f.  r2  Method/Source  Sabah, Malaysia  391  F  SVL  3.00  0.0016  0.1560  -  Sabah, Malaysia  363  M  SVL  3.00  0.0015  0.1451  -  Weipa, northern Australia  -  mixed  SVL  3.00  0.0011  0.1109  -  Fiji  -  F  SVL  2.64  0.0031  0.8150  -  b  Fiji  -  M  SVL  2.40  0.0101  0.7100  -  c  Vanuatu  -  F  SVL  2.54  0.0052  0.7650  -  d  Vanuatu  -  M  SVL  2.38  0.0046  0.8100  -  unknown  -  F  SVL  3.07  0.0008  0.8560  -  unknown  49  M  SVL  3.58  0.0001  0.8950  -  unknown  -  F  SVL  3.09  0.0001  0.9460  -  unknown  -  M  SVL  3.00  0.0002  0.8990  -  near Orchid Island, Taiwan  70  F  SVL  3.00  0.0008  0.0841  -  near Orchid Island, Taiwan  141  M  SVL  3.00  0.0008  0.0775  -  Gulf of Carpentaria, Australia  2  F  SVL  3.00  0.0008  -  -  Weipa, northern Australia  2  F  SVL  3.00  0.0008  0.0807  -  Genovesa, Galapagos Island  41  M  SVL  3.00  0.0458  4.579  -  Genovesa, Galapagos Island  15  M  SVL  3.00  0.0424  4.244  -  a from c.f. of data from Hin et al. (1991; Tab. 2, p. 466). a from c.f. of data from Hin et al. (1991; Tab. 2, p. 466). a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Cogger and Heatwole (2006; Tab. 1, p. 248). a from c.f. of data from Tu et al. (1990; Tab. 1, p. 120). a from c.f. of data from Tu et al. (1990; Tab. 1, p. 120). a from mean c.f. of survey data from AFRDC, CSIRO, NPF. a from c.f. of data from Fry et al. (2001; Tab. 2, p. 59). a from c.f. of territorial males from Wikelski et al. (1996; Tab. 1, p. 587). a from c.f. of marginal males from Wikelski et al. (1996; Tab. 1, p. 587).  Species  Lapemis hardwickii  l m 23  24  a  a  Laticauda colubrina  Laticauda frontalis  b 25  a  Laticauda saintgironsi  b 26  a  Laticauda semifasciata  b 27  a  Pelamis platurus  b 28  a  Amblyrhynchus cristatus  Locality  (Squamata, Iguanidae) b  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  69  Table A1. Continued. Spec. No. 28  29  Stock No. c  n  Sex  Type  b  a  c.f.  r2  Method/Source  Genovesa, Galapagos Island  16  M  SVL  3.00  0.0458  4.578  -  d  Genovesa, Galapagos Island  11  M  SVL  3.00  0.0448  4.476  -  e  Genovesa, Galapagos Island  30  M  SVL  3.00  0.0463  4.628  -  Caretta caretta  Curacao  23  unsexed  SCL  2.95  0.1655  -  0.986  431  unsexed  SCL  2.82  0.000282  Chelonia mydas  USA (Cheasapeake, Florida), UK, France, Japan Gulf coast of Florida, USA  208  unsexed  CL  2.91  0.1674  -  0.993  b  Saurashtra Coast, Gujarat, India  69  unsexed  CCL  3.00  0.1145  11.45  -  c  USA (Florida), Mexico (Baja California), Tortuguero, Ascension, Suriname, Baja California, Mexico Milman, Great Barrier Reef, Australia Persian Gulf (Shidvar, Ommolkaran, Nakhillo and Queshm Islands), Iran Honduras, Cayman, Barbados, Suriname  426  unsexed  SCL  2.90  0.000206  200 25  unsexed F unsexed  SCL CCL CCL  3.00 3.00 2.96  0.1519 0.0922 0.1275  112  unsexed  SCL  2.74  0.000278  Florida, USA  78  unsexed  CL  2.49  0.8919  USA (Cheasapeake, Florida), UK, France  145  unsexed  SCL  2.84  0.000247  Northern Territory, Australia  85  F  CCL  3.00  0.0001  0.0111  -  b  Primeira Islands, Mocambique  1  unsexed  CL  3.00  0.1166  11.66  -  c  Hawaii, Brazil, Suriname, Mozambique  40  unsexed  SCL  2.68  0.000479  205  unsexed  CCL  3.00  0.0001  0.0105  -  2  unsexed  SCL  3.00  0.0897  -  -  16 102  unsexed F  CCL CCL  3.67 2.41  0.0044 1.8253  -  0.853 0.746  a from c.f. of small males from Wikelski et al. (1996; Tab. 1, p. 587). a from c.f. of single territories from Wikelski et al. (1996; Tab. 3, p. 589). a from c.f. of leks from Wikelski et al. (1996; Tab. 3, p. 589). Nagelkerken et al. (2003; Tab. 1, p. 186). Not a good representative of the population. Wabnitz (2008; Tab. 1 p. xx); weight in kg. Carr & Cadwell (1956; Tab. 2, p. 15). a from c.f. of data from Kannan et al. (2005; Tab. 2, p. 5). Wabnitz (2008; Tab. 1 p. xx); weight in kg. Seminoff et al. (2003; p. 1355). Loop et al. (1995; Tab. 2, p. 247). Morabaki & Elmi (2005; Tab. 1, p. 7). Wabnitz (2008; Tab. 1 p. xx); weight in kg. Carr & Cadwell (1956; Tab. 2, p. 15). Wabnitz (2008; Tab. 1 p. xx); weight in kg. a from c.f. of data from Whiting et al. (2007; Tab. 3, p. 205); weight in kg. a from c.f. of data from Hughes (1972; Tab. 1, p. 129). Wabnitz (2008; Tab. 1 p. xx); weight in kg. a from c.f. of data from Schäuble et al. (2006; Tab. 2, p. 192); weight in kg. a from mean c.f. of data from Jones et al. (2008; Tab. 3). James et al. (2007; Fig. 3, p. 248). James et al. (2005; Fig. 3, p. 199).  a  Species  Amblyrhynchus cristatus  Locality  (Testudines, Cheloniidae) b 30  31  a  a b c  Eretmochelys imbricata  d 32  a  Lepidochelys kempi  b 33  a  Lepidochelys olivacea  34  a  Natator depressus  Field Island, Australia  35  a  Dermochelys coriacea  Florida, USA  0.970  0.990 14.99 9.224 -  0.844 0.990  -  0.951 0.960  0.840  (Testudines, Dermochelydae) b c  Nova Scotia, Canada St. Croix, US Virgin islands  70  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A1. Continued. Spec. No. 35  Stock No. d  Species  Dermochelys coriacea  Locality  n  Sex  Type  b  a  c.f.  r2  101  unsexed  SCL  2.81  0.2640  -  -  1  unsexed  SCL  3.00  0.1180  11.80  -  e  University of British Colombia, Vancouver, Canada Unknown, American Samoa  f  Unknown, Hawaii  3  unsexed  SCL  3.00  0.1149  -  -  g  Unknown  1  unsexed  SCL  3.00  0.1118  11.18  -  h  western Australia, Australia  2  unsexed  SCL  3.00  0.0656  -  -  Method/Source Jones et al. (2008; Tab. 3). a from c.f. of data from Jones et  al. (2008; Tab. 3).  a from mean c.f. of data from Jones et al. (2008; Tab. 3). a from c.f. of data from Jones et al. (2008; Tab. 3). a from mean c.f. of data from Jones et al. (2008; Tab. 3).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  71  Table A2. Growth parameter estimates assembled for marine reptiles and used to obtain Figure 2. Sex: U=unsexed or mixed; F=females; M=males. Type: TL=total length; SVL=snout-vent length; SCL=straight carapace length; CCL=curved carapace length; CL=carapace length. Spec. No.  Stock No.  1  a  Species  Crocodylus porosus  L∞  Type  W∞ kg  (or Z)  Z/K  years-1  K  Φ'  K  Φ'  U  323.1  TL  144  -  0.710  4.87  -  b  Queensland, Australia  907  U  265.2  TL  70.5  -  2.500  5.25  -  c  W. Australia  736  U  262.1  TL  67.6  -  0.750  4.71  -  Phangnga Bay, Thailand  77  F  93.4  SVL  0.425  4.04  -  -  0.829  Phangnga Bay, Thailand  42  M  72.2  SVL  0.145  1.61  -  -  1.386  Muar River, Malaysia  181  U  85.0  SVL  0.360  (0.99)  0.270  3.29  -  Muar River, Malaysia  597  U  76.8  SVL  0.265  (1.53)  0.410  3.38  -  Acalyptophis peronii  G. Carpentaria, Australia  50  U  137.7  SVL  1.79  3.29  -  -  0.382  a  Acrochordus granulatus  a  Cerberus rynchops  (Squamata, Colubridae) b  a  (Squamata, Hydrophiidae)  Method/Comments/Source  from  7665  b  4  Sex  N. Territory, Australia  (Squamata, Acrochordidae)  3  N  cm  (Crocodilia, Crocodylidae)  2  Locality  5  a  Aipysurus duboisii  G. Carpentaria, Australia  20  U  114.6  SVL  0.796  0.990  -  -  0.551  6  a  Aipysurus eydouxii  G. Carpentaria, Australia  106  U  112.8  SVL  1.32  3.05  -  -  0.569  L∞ & K from length frequency analysis of data from Messel and Vorlicek (1986; Tab. 1a, p. 7684). Unexploited; 61-305 cm. W∞ from Tab 1 (1a). L∞ & K from length frequency analysis of data from Messel and Vorlicek (1986; Tab. 1a, p. 7684). Unexploited; 61-244 cm. W∞ from Tab 1 (1a). L∞ & K from length frequency analysis of data from Messel and Vorlicek (1986; Tab. 1a, p. 7684). Unexploited; 61-244 cm. W∞ from Tab 1 (1a). L∞ from single length frequency histogram from Wangkulangkul et al. (2005; Fig. 2, p. 260). Exploited; 35-81 cm. W∞ from Tab 1 (2a). K=ave. Φ'; all sea snakes. L∞ from single length frequency histogram from Wangkulangkul et al. (2005; Fig. 2, p. 259). Exploited 35-65 cm. W∞ from Tab 1 (2b). K=ave. Φ'; all sea snakes. L∞ & K from length frequency analysis of data from Jayne et al. (1988; Tab. 1, p. 5 rows 1-2). Unexploited; 24-64 cm. W∞ from Tab 1 (3a). L∞ & K from length frequency analysis of data from Jayne et al. (1988; Tab. 1, p. 5 rows 3-6). Unexploited; 1984-1987; 24-64 cm. W∞ from Tab 1 (3a). L∞ from single length frequency histogram from FRDC/CSIRO/NPF survey data. Exploited; 70126 cm. W∞ from Tab 1 (4e). K=ave. Φ' Hydrophiidae. L∞ from single length frequency histogram from FRDC/CSIRO/NPF survey data. Exploited; 80110 cm. W∞ from Tab 1 (6e). K=ave. Φ'; Hydrophiidae. L∞ from single length frequency histogram from FRDC/CSIRO/NPF survey data. Exploited; 35-96 cm. W∞ from Tab 1 (7g). K=ave. Φ'; Hydrophiidae.  72  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No.  Stock No.  7  a  Aipysurus laevis  8  a  Astrotia stokesii  9  a  Disteira kingii  10  a  Disteira major  11  a  Emydocephalus ijimae  Species  13  W∞  Z/K  K  from  Method/Comments/Source  -  -  0.375  0.020  -  -  0.223  1.07  2.44  -  -  0.239  SVL  2.63  1.23  -  -  0.233  79.8  SVL  0.449  -  3.820  4.39  -  M  66.8  SVL  0.210  -  2.540  4.05  -  69  U  102.1  SVL  0.628  1.38  -  -  0.752  Muar, Malaysia  295  F  113.9  SVL  0.890  (1.13)  0.600  3.89  -  Muar, Malaysia  359  M  103.4  SVL  0.583  (1.52)  0.740  3.90  -  Australian continental shelf Australian continental shelf G. Carpentaria, Australia  306  M  170.1  SVL  1.33  -  0.310  3.95  -  276  F  221.3  SVL  3.09  -  0.170  3.92  -  525  U  257.0  SVL  5.01  (0.590)  0.170  4.05  -  L∞ from single L/F histogram; FRDC/CSIRO/NPF survey data. Exploited; 60-130cm. W∞ from Tab 1 (8g). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram; FRDC/CSIRO/NPF survey data. Exploited; 55-176 cm. W∞ from Tab 1 (9g). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram; FRDC/CSIRO/NPF survey data. Exploited; 60-170 cm. W∞ from Tab 1 (10f). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram; FRDC/CSIRO/NPF survey data. Exploited; 60-160 cm. W∞ from Tab 1 (11h). K=ave. Φ'; Hydrophiidae. Direct fitting of VBGF to age at length from Masunaga and Ota (2003; Fig. (2,4)a, p. 464,467); t0= 0; unexploited; 42-83 cm. W∞ from Tab 1 (13a). Direct fitting of VBGF to age at length from Masunaga and Ota (2003; Fig. (2,4)b, p. 464 & 466). t0=-0.42; unexploited; 30-68 cm. W∞ from Tab 1 (13b). L∞ from single L/F histogram from FRDC/CSIRO/NPF survey data. Exploited; 35-96 cm. W∞ from Tab 1(14c). K=ave. Φ'; same species. L∞ & K from L/F analysis of data from Voris and Jayne (1979; Fig. 1(b,d,f,h), p. 311). Exploited; 21-102 cm. W∞ from Tab 1 (14a). L∞ & K from L/F analysis of data from Voris and Jayne (1979; Fig. 1(a,c,e,g), p. 310). Exploited; 21-100 cm. W∞ from Tab 1 (14b). VBGF parameters from Ward (2001; Tab. 3, p. 196); t0=-0.93; T=28-29°C; exploited; 75-193 cm. W∞ from Tab 1 (17e). VBGF parameters from Ward (2001; Tab. 3, p. 196); t0=-2.1; T=28-29°C; exploited; 82-218 cm. W∞ from Tab 1 (17c). L∞ & K from L/F analysis; FRDC/CSIRO/NPF survey data. Exploited; 40-221 cm. W∞ from Tab 1 (17g).  Sex  G. Carpentaria, Australia G. Carpentaria, Australia G. Carpentaria, Australia G. Carpentaria, Australia Zamamijima, Ryukyu Island  74  U  138.8  131  U  48  Type  (or Z)  years-1  SVL  4.00  2.16  180.1  SVL  10.0  U  174.2  SVL  248  U  176.4  -  F  -  G. Carpentaria, Australia  b c  a  a b c  Zamamijima, Ryukyu Island  Enhydrina schistosa  Hydrophis elegans  K  Φ'  N  kg  b  12  L∞  Locality  cm  Φ'  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  73  Table A2. Continued. Spec. No.  Stock No.  14  a  Hydrophis ornatus  15  a  16  a  W∞  Z/K  K  K  from  Method/Comments/Source  -  -  0.432  1.42  -  -  0.203  1.51  -  0.410  3.71  -  SVL  1.48  -  0.440  3.75  -  132.0  SVL  2.45  (2.98)  0.750  4.12  -  F  94.4  SVL  1.31  3.06  -  -  0.812  363  M  86.2  SVL  0.929  2.38  -  -  0.974  Indo-Pacific  -  F  170.4  SVL  2.39  2.55  -  -  0.249  b  Indo-Pacific  -  M  136.6  SVL  1.34  1.89  -  -  0.388  c  Indo-Pacific  1294  U  125.8  SVL  0.454  1.47  -  -  0.457  d  Mabualau and Toberua, Fiji  352  F  150.8  SVL  1.73  2.97  -  -  0.318  e  Mabualau and Toberua, Fiji  648  M  90.8  SVL  0.504  0.763  -  -  0.878  L∞ from single L/F histogram from FRDC/CSIRO/NPF survey data. Exploited; 60120 cm. W∞ from Tab 1 (20g). K=ave. Φ';Hydrophis. L∞ from single L/F histogram from FRDC/CSIRO/NPF survey data. Exploited; 120180 cm. W∞ from Tab 1 (21c). K=ave. Φ'; Hydrophis. VBGF parameters from Ward (2001; Tab. 3, p. 196); t0=-0.86; exploited; 42-120 cm. W∞ from Tab 1 (22d). VBGF parameters from Ward (2001; Tab. 3, p. 196); t0=-0.57; exploited; 50-126 cm. W∞ from Tab 1 (22f). L∞ & K from L/F analysis of data from FRDC/CSIRO/NPF survey data. Exploited; 30121 cm. W∞ from Tab 1 (22h). L∞ from single L/F histogram from Hin et al. (1991; Fig. 1, p. 467). Exploited; 30-80 cm. W∞ from Tab 1 (22k). K=ave. Φ'; same species. L∞ from single L/F histogram from Hin et al. (1991; Fig. 1, p. 468). Exploited; 34-84 cm. W∞ from Tab 1 (22l). K=ave. Φ'; same species. L∞ from single L/F histogram from Heatwole et al. (2005; Fig. 24, p. 44). Exploited; 25-156 cm. W∞ from Tab 1 (23a). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Heatwole et al. (2005; Fig. 24, p. 44). Exploited; 30-125 cm. W∞ from Tab 1 (23b). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Heatwole et al. (2005; Fig. 24, p. 44). Exploited; 15-165 cm. W∞ from Tab 1 (23d). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Shetty and Shine (2002; Fig. 1b, p. 48). Unexploited; 30140 cm. W∞ from Tab 1 (23a). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Shetty and Shine (2002; Fig. 1a, p. 48). Unexploited; 30-91 cm. W∞ averaged from Tab 1 (23b). K=ave. Φ; Hydrophiidae.  N  Sex  G. Carpentaria, Australia  178  U  129.4  Hydrophis pacificus  G. Carpentaria, Australia  32  U  Lapemis hardwickii  Australian continental shelf Australian continental shelf G. Carpentaria, Australia Sabah, Malaysia  227  Sabah, Malaysia  b c d e 17  L∞  Φ'  Species  a  Laticauda colubrina  Locality  Type  kg  (or Z)  years-1  SVL  1.70  1.40  188.8  SVL  2.01  F  112.2  SVL  184  M  113.0  549  U  391  cm  Φ'  74  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No.  Stock No.  18  a  19  W∞  Z/K  K  K  from  Method/Comments/Source  -  -  0.480  L∞ from single L/F histogram from Heatwole et al. (2005; Fig. 25, p. 45). Exploited; 30-111 cm.  4.44  -  -  0.776  0.443  1.38  -  -  0.424  SVL  1.08  1.24  -  -  1.516  29.8  SVL  1.12  (4.83)  1.400  3.09  -  U  41.1  SVL  3.17  1.76  -  -  0.737  1600  U  73.4  SCL  55.5  2.09  -  -  0.102  Florida, Georgia, S. Carolina, USA Cayman Islands  118  U  110.0  SCL  175  -  0.031  2.58  -  250  U  110.3  SCL  174  1.34  -  -  0.048  d  Florida, USA  1234  U  110.9  SCL  178  -  0.044  2.79  -  e  Florida, USA  41  U  94.7  SCL  114  -  0.115  3.01  -  f  Florida, USA  51  U  96.1  SCL  119  -  0.059  2.73  -  Locality  N  Sex  Indo-Pacific  -  F  122.7  b  Indo-Pacific  -  M  c  Indo-Pacific  192  Genovesa, Galapagos Island  b  c  a  Laticauda saintgironsi  Amblyrhynchus cristatus (Squamata, Iguanidae)  20  L∞  Φ'  Species  a  Caretta caretta  Type  kg  (or Z)  years-1  SVL  0.359  1.68  96.5  SVL  1.82  U  130.6  SVL  41  M  28.6  Genovesa, Galapagos Island  318  U  Sta.Fe, Galapagos Island  8000  Azores  cm  Φ'  (Testudines, Cheloniidae) b c  W∞ from Tab 1 (25a). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Heatwole et al. (2005; Fig. 25, p. 45). Exploited; 35-86 cm. W∞ from Tab 1 (25b). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Heatwole et al. (2005; Fig. 25, p. 45). Exploited; 20-120 cm. W∞ averaged from Tab 1 (25a and b). K=ave. Φ'; Hydrophiidae. L∞ from single L/F histogram from Wikelski et al. (1996; Fig. 6, p. 590). Unexploited; territorial males; 21-29 cm W∞ from Tab 1 (28a). K=Φ'; same species. L∞ & K from L/F analysis of data from Wikelski and Trillmich (1997; Fig. 8, p. 928). Unexploited; 11-27 cm; narrow size range may not be a good representative of the population. W∞ from Tab 1 (28b). L∞ from single L/F histogram from Wikelski and Trillmich (1997; Fig. 2, p. 926). Unexploited; 1137 cm. W∞ from Tab 1 (28c). K=Φ'; same species. L∞ from single L/F histogram from Bjorndal et al. (2003; Tab. 1, p. 735). Unexploited; 10-70 cm. W∞ from Tab 1 (29b). K=ave. Φ'; same species. VBGF parameters from Epperly et al. (2001; Tab. 8 p. 46). Unexploited; 45-110 cm; markrecapture. W∞ from Tab 1 (29b). L∞ from single L/F histogram from Epperly and Teas (2002; Tab. 1, p. 468). Unexploited; 1-101 cm. W∞ from Tab 1 (29b). K=ave. Φ'; same species. VBGF parameters from Bjorndal et al. (2001; Fig. 1, p. 242). Unexploited; stranded sea turtles; 46-87 cm. W∞ from Tab 1 (29b). VBGF parameters from Epperly et al. (2001; Tab. 8 p. 46). Unexploited; 53-77 cm; markrecapture. W∞ from Tab 1 (29b). VBGF parameters from Epperly et al. (2001; Tab. 8 p. 46). Unexploited; 38-110 cm; markrecapture. W∞ from Tab 1 (29b).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  75  Table A2. Continued. Spec. No.  Stock No.  20  g  21  L∞  W∞  Z/K  K  K  Φ'  from  Method/Comments/Source  0.057  2.72  -  -  0.051  2.81  -  250  3.32  -  -  0.037  SCL  160  -  0.052  2.77  -  96.7  SCL  121  -  0.064  2.78  -  U  120.0  SCL  222  2.57  -  -  0.041  92  F  101.1  SCL  127  1.78  -  -  0.083  Bahia Magdalena, Mexico  718  U  102.5  SCL  132  2.60  -  -  0.094  c  Bahia Magdalena, Mexico  212  U  101.0  SCL  127  (0.160)  0.040  2.61  -  d  Baja California, Mexico  200  U  106.1  SCL  146  2.87  -  -  0.088  e  Cayman Islands  176  U  88.2  SCL  85.8  0.704  -  -  0.127  VBGF parameters from Epperly et al. (2001; Tab. 8 p. 46). Unexploited population; 28-110 cm; mark-recapture. W∞ from Tab 1 (29b). VBGF parameters from Bjorndal et al. (2001; Fig. 2, p. 243). Unexploited; stranded sea turtles; 46-87 cm. W∞ from Tab 1 (29b). L∞ from single L/F histogram from Teas (1993; Tab. 6-7, p. 16-18). Unexploited; 0-121 cm. W∞ from Tab 1 (29b). K=ave. Φ'; same species. VBGF parameters from Epperly et al. (2001; Tab. 8 p. 46). Unexploited; with 45-76 cm; mark-recapture. W∞ from Tab 1 (29b). VBGF parameters from Epperly et al. (2001; Tab. 8 p. 46). Unexploited; 62-104 cm; markrecapture. W∞ from Tab 1 (29b). L∞ from single L/F histogram from Teas (1993; Tab. 5-7, p. 14-18). Unexploited; 0-121 cm. W∞ from Tab 1 (29b). K=ave. Φ'; same species. L∞ from single L/F histogram from Broderick et al. (2003; Fig. 2a, p. 98). Growth rate: 11 cm CCL year-1; 0.27 cm CCW year-1; unexploited; 76-106 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species. L∞ from single L/F histogram from Koch et al. (2006; Fig. 3, p. 331). Unexploited; juveniles; 35-91 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species. Seasonalized VBGF parameters C= 0, ts= 0.75, t0= 0 from Koch et al. (2007; p. 35). Unexploited; 43-73 cm. Growth 3x higher in summer (0.28 cm month-1) than winter 0.09 cm month-1; ave. growth rate 1.62 cm year-1. W∞ from Tab 1 (30c). L∞ from single L/F histogram from Seminoff et al. (2003; Fig. 4, p. 1359). Unexploited; 46-100 cm. W∞ from Tab 1 (30c). K= ave. Φ' for same species. L∞ from single L/F histogram from Epperly and Teas (2002; Tab. 1, p. 468). Unexploited; 1-81 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species.  Species  Locality  N  Sex  Florida, USA  19  U  96.1  h  G. Mexico, USA  570  U  i  G. Mexico, USA  1639  j  N. Carolina, USA  k  S.E. USA  l  W. Atlantic  Type  kg  (or Z)  years-1  SCL  119  -  105.7  SCL  155  U  125.1  SCL  57  U  106.9  54  U  6727  Alagadi beach, Cyprus  b  a  Caretta caretta  Chelonia mydas  cm  Φ'  76  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No.  Stock No.  21  f  L∞  W∞  Z/K  K  K  Φ'  from  0.320  3.39  -  -  0.740  3.85  -  117  -  0.074  2.85  -  SCL  121  -  0.072  2.85  -  92.6  SCL  98.7  -  0.082  2.85  -  U  168.0  SCL  554  -  0.025  2.85  -  571  U  82.2  SCL  69.9  -  0.122  2.92  -  Great Inagua, Bahamas  509  U  158.6  SCL  469  -  0.035  2.94  -  n  Great Inagua, Bahamas  363  U  162.8  SCL  506  -  0.033  2.94  -  o  Great Inagua, Bahamas  211  U  84.4  SCL  75.5  -  0.114  2.91  -  p  Gulf of Mexico, USA  357  U  96.5  SCL  111  0.611  -  -  0.106  q  Queensland, Australia  94  U  98.8  SCL  119  0.628  -  -  0.087  r  Watamu, Kenya  1666  U  110.3  SCL  164  -  0.070  2.98  -  Species  Locality  N  Sex  Pamlico, N. Carolina, USA  226  U  87.7  g  Great Inagua, Bahamas  964  U  h  Great Inagua, Bahamas  884  i  Great Inagua, Bahamas  j  Type  kg  (or Z)  years-1  SCL  84.4  (0.760)  98.3  SCL  117  U  98.3  SCL  839  U  99.4  Great Inagua, Bahamas  772  U  k  Great Inagua, Bahamas  691  l  Great Inagua, Bahamas  m  Chelonia mydas  cm  Φ'  Method/Comments/Source L∞ & K from L/F analysis of data from Epperly et  al. (2007; Fig. 6, p. 590). Unexploited; 20-80  cm. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992; 26-72 cm. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). VBGF parameters from Bjorndal et al. (1995; Tab. 1, p. 74). Unexploited; 1983-1992. W∞ from Tab 1 (30c). L∞ from single L/F histogram from Teas (1993; tab. 9-10, p. 22-24). Unexploited; 0-91 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species. L∞ from single L/F histogram from Robins (2007; Fig. 2, p. 163). Exploited; 20-101 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species. Direct fitting of VBGF to age at length from Watson (2006; Fig. 5.7, p. 45). Unexploited; 33115 cm; t0=-0.75; growth rates 0.468-10.67 cm (CCL) year-1, average growth 5.18 cm year-1. W∞ from Tab 1 (30c).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  77  Table A2. Continued. Spec. No.  Stock No.  21  s  Species  Chelonia mydas  t 22  a  Chelonia mydas agassizii  23  a  Eretmochelys imbricata  W. Atlantic Yaeyama, Okinawa, Japan Bahia Magdalena, Mexico Great Barrier Reef, Australia  L∞  N  Sex  1393  U  118.8  50  U  52  W∞  Z/K  K  K  Φ'  from  Method/Comments/Source  -  -  0.070  1.84  -  -  0.126  123  -  -  2.70  0.050  SCL  58.3  1.82  -  -  0.119  L∞ from single L/F histogram from Teas (1993; Tab. 8-10, p. 20-24). Unexploited; 0-111 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species. L∞ from single L/F histogram from Sakai et al. (2000; Fig. 2a, p. 379). Exploited; 32-81 cm. W∞ from Tab 1 (30c). K=ave. Φ'; same species. VBGF parameters from Lyons et al. (2002; Fig. 2). C=0.65, ts=0.72; unexploited; 43-72 cm; mark-recapture. W∞ from Tab 1 (30c). L∞ from single L/F histogram from Limpus (1992; Fig. 4, p. 498). Unexploited; 35-88 cm. W∞ from Tab 1 (31d). K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Moncada et al. (1999; Tab. 1, p. 258). Unexploited; 31-92 cm. W∞ from Tab 1 (31b). K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Teas (1993; Tab. 15-16, p. 34-36). Unexploited; juveniles; 050 cm. W∞ from Tab 1 (31a). K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Beggs et al. (2007; Fig. 3, p. 162). Unexploited; 77-103 cm. W∞ from Tab 1 (31b). K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Teas (1993; Tab. 14-16, p. 32-36). Unexploited; 0-81 cm. W∞ from Tab 1 (31d). K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Schmid (2000; Fig. 1-3, p. 10). Unexploited; 20-60 cm. W∞ from Tab 1 (32a). L∞ from single L/F histogram from Epperly and Teas (2002; Tab. 1, p. 468). Unexploited; 1-51 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Schmid (2000; Fig. 1-3, p. 10). Unexploited; 25-60 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Schmid (2000; Fig. 1-3, p. 10). Unexploited; 20-56 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ & K from L/F analysis of data from Epperly et al. (2007; Fig. 7, p. 288). Unexploited; 25-59 cm. W∞ from Tab 1 (32b).  Type  kg  (or Z)  years-1  SCL  203  1.73  88.8  SCL  87.4  U  100.0  SCL  106  U  85.7  cm  Φ'  b  Cuban Archipelago  6789  F  99.0  SCL  89.6  2.01  -  -  0.101  c  G. Mexico, USA  117  U  61.7  SCL  35.7  2.71  -  -  0.260  d  NeedHam's Point, Barbados W. Atlantic  1310  F  99.4  SCL  90.7  1.39  -  -  0.089  169  U  86.0  SCL  58.9  1.61  -  -  0.134  Cape Canaveral, Florida, USA Cayman Islands  147  U  66.2  SCL  31.1  (3.09)  1.30  3.76  -  631  U  61.5  SCL  23.5  1.22  -  -  0.408  c  Cedar Keys, USA  253  U  61.1  SCL  23.1  1.22  -  -  0.413  d  Chesapeake Bay, USA  38  U  61.6  SCL  23.6  1.86  -  -  0.406  e  E. Pamlico, N. Carolina, USA  67  U  61.6  SCL  23.5  (0.720)  0.770  3.46  -  e 24  Locality  a b  Lepidochelys kempii  78  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A2. Continued. Spec. No.  Stock No.  24  f  W∞  Z/K  K  from  Method/Comments/Source  0.129  2.79  -  -  0.317  3.09  -  40.1  1.42  -  -  0.280  SCL  31.9  1.96  -  -  0.329  70.7  SCL  34.9  -  0.200  3.00  -  U  71.1  SCL  35.4  -  0.210  3.03  -  189  U  56.2  SCL  18.2  2.12  -  -  0.488  102  U  62.7  SCL  24.8  2.07  -  -  0.392  1028  U  77.6  SCL  45.4  1.93  -  -  0.256  Withlacoochee and Crystal Rivers, USA G. Mannar, India  76  U  56.7  SCL  18.7  2.09  -  -  0.479  99  U  73.8  CCL  49.2  0.758  -  -  0.177  N. Territory, Australia  85  F  77.0  CCL  50.6  2.87  -  -  0.163  VBGF parameters from Coyne (2000; Tab. 4, p. 59). Unexploited; recaptured wild and head-start turtles; W∞ from Tab 1 (32a). VBGF parameters from Coyne (2000; Tab. 4, p. 59). Unexploited; stranded head-start turtles; W∞ from Tab 1 (32b). L∞ from single L/F histogram from Teas (1993; Tab. 12-13, p. 28-30). Unexploited; 0-71 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Coyne (2000; Fig. 21, p. 45). Unexploited; stranded sea turtles; 15-61 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. Fitted VBGF from Schmid and Woodhead (1998; Eq. 2, p. 96). To= -0.32; Unexploited; 22-67 cm; mark-recapture. W∞ from Tab 1 (32b). Fitted VBGF from Schmid and Woodhead (1998; Eq. 1, p. 96); t0=-0.31; unexploited; 22-68 cm; tag-recapture. W∞ from Tab 1 (32b). L∞ from single L/F histogram from Coyne (2000; Fig. 21, p. 45). Unexploited; 20-51 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Schmid (2000; Fig. 1-3, p. 10). Unexploited; 20-56 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Teas (1993; Tab. 11-13, p. 26-30). Unexploited; 0-71 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Schmid (2000; Fig. 1-3, p. 10). Unexploited; 25-50 cm. W∞ from Tab 1 (32b). K=ave. Φ'; same species. L∞ from single L/F histogram from Bhupathy and Saravanan (2006; Fig. 2, p. 140). Exploited; 4672 cm; narrow range, may not be a good representative of the population. W∞ from Tab 1 (33c). L∞ assummed as SCL; K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Whiting et al. (2007; Fig. 4, p. 205). Unexploited; 64-76 cm. W∞ from Tab 1 (33a). L∞ assummed as SCL; K=ave. Φ'; Cheloniidae.  N  Sex  Lepidochelys kempii  Florida, USA  36  U  69.4  g  G. Mexico, USA  114  U  h  G. Mexico, USA  722  i  NMFS Statistical Zone, USA  j  Type  kg  (or Z)  years-1  SCL  33.0  -  62.3  SCL  24.3  U  74.3  SCL  256  U  68.5  N G. Mexico, Atlantic coast, USA N. G. Mexico, USA  96  U  58  Sambine Pass, G. Meixco, USA Apalachee Bay, USA  n  W Atlantic  o  l m  a  b  Lepidochelys olivacea  K  Φ'  Locality  k  25  L∞  Species  cm  Φ'  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  79  Table A2. Continued. Spec. No.  Stock No.  25  c  26  a  Species  W∞  Z/K  K  from  Method/Comments/Source  -  -  0.136  2.68  -  -  0.095  156  2.12  -  -  0.074  SCL  521  1.17  -  -  0.057  162.2  SCL  420  1.94  -  -  0.033  U  150.1  SCL  338  1.57  -  -  0.039  101  U  155.0  SCL  370  -  0.270  3.81  -  243  U  105.0  SCL  124  ‐  -  -  0.140  L∞ from single L/F histogram from Robins (2007; Fig. 2, p. 163). Exploited; 20-71 cm. W∞ from Tab 1 (33b). L∞ assummed as SCL; K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Limpus et al. (2006 ; Fig. 2, p. 10). Unexploited; nesting females; 89-99 c; Woongarra coast not included. W∞ from Tab 1 (34a). L∞ assummed as SCL; K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Robins (2007; Fig. 2, p. 163). Exploited; 20 to 101 cm. W∞ from Tab 1 (34a). L∞ assummed as SCL; K=ave. Φ'; Cheloniidae. L∞ from single L/F histogram from Teas (1993; Tab. 18-19, p. 40-42). Unexploited; 100-160 cm. W∞ from Tab 1 (35d). K=Φ'; same species. L∞ from single L/F histogram from James et al. (2007; Fig. 1, p. 248). Unexploited; 125-166 cm. W∞ from Tab 1 (35d). K=Φ'; same species. L∞ from single L/F histogram from James et al. (2007; Fig. 1, p. 248). Unexploited; 100-156 cm. W∞ from Tab 1 (35d). K=Φ'; same species. VBGF parameters from Jones et al. (2008; Tab. 1). To= -0.12; unexploited; maintained in captivity from hatchlings to > 2-years; W∞ from Tab 1 (35d). L∞ from single L/F histogram from Teas (1993; Tab. 17-19, p. 38-42). Unexploited; 10-101 cm. W∞ from Tab 1 (35d). K=Φ'; same species.  Sex  Lepidochelys olivacea  Queensland, Australia  31  U  84.3  Natator depressus  Curtis Island, Australia  48  F  Queensland, Australia  76  G. Mexico, USA  b  Type  kg  (or Z)  years-1  CCL  69.8  1.99  100.5  CCL  107  U  114.2  CCL  41  U  164.1  Nova Scotia, Canada  120  U  c  off coast, France  82  d  Vancouver, Canada  e  W. Atlantic  a  Dermochelys coriacea  (Testudines, Dermochelydae)  K  Φ'  N  b  27  L∞  Locality  cm  Φ'  80  Growth of marine reptiles, Palomares, M.L.D., et al.  Table A3. Maturity data assembled for sea snakes and sea turtles used to obtain Figure 4. Spec. No. 1  Stock No. a  Species  Acalyptophis peronii  Locality  Sex  Lm  northern Australia  unsexed  (cm) 71.6  northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia Sourabaya, Java, Indonesia Sourabaya, Java, Indonesia northern Australia northern Australia northern Australia Sourabaya, Java, Indonesia Sourabaya, Java, Indonesia northern Australia northern Australia Sourabaya, Java, Indonesia Sourabaya, Java, Indonesia  M F M unsexed F M unsexed M F unsexed M F M F unsexed unsexed M F unsexed M F M M F unsexed F M M F F M M F  89 71.6 91 91 47.2 64 47.2 103 102 103 72 81.7 81.7 82.3 145 82.3 71 84 71 70 70 79 79 65 65 84 70 70  (Squamata, Hydrophiidae) 2 3 4 5 6 7 8 9 10  11 12 13  b c a a b c a b c a b c a b c a b c a b c a a b c d e a b a b a b  Aipysurus apraefrontalis Aipysurus duboisii Aipysurus eydouxii Aipysurus laevis Astrotia stokesii Disteira kingii Disteira major Emydocephalus annulatus Enhydrina schistosa  Hydrophis brookii Hydrophis caerulescens Hydrophis cyanocinctus  Comments/Remarks Lm max 114 cm from Milton (2001). Lm range 70.3 - 113.9 cm from Fry et al. (2001). Lm range 70.2 - 110.8 cm from Fry et al. (2001). Lm min 92 cm from Fry et al. (2001). Lm max 117 cm from Milton (2001). Lm range 91 - 116.2 cm from Fry et al. (2001). Lm range 57 - 116.5 cm from Fry et al. (2001). Lm max 85 cm from Milton (2001). Lm range 54.7 - 78 cm from Fry et al. (2001). Lm range 39.2 - 85 cm from Fry et al. (2001). Lm max 130 cm from Milton (2001). Lm range 64 - 106 cm from Fry et al. (2001). Lm range 71.2 - 130 cm from Fry et al. (2001). Lm range 59.5 - 122 cm from Fry et al. (2001). Lm range 71.4 - 138 cm from Fry et al. (2001). Lm max 138 cm from Milton (2001). Lm max 165 cm from Milton (2001). Lm range 66.1 - 162 cm from Fry et al. (2001). Lm range 78.9 - 157.2 cm from Fry et al. (2001). Lm max 165 cm from Milton (2001). Lm range 53 - 163.5 cm from Fry et al. (2001). Lm range 61.5 - 143.1 cm from Fry et al. (2001). Lm min 880 cm from Fry et al. (2001) Length at the beginning of maturity from Bergman Length at the beginning of maturity from Bergman Lm max 102.4 cm from Milton (2001). Lm range 47.1 - 101.5 cm from Fry et al. (2001). Lm range 56 - 88.1 cm from Fry et al. (2001). Length at the beginning of maturity from Bergman Length at the beginning of maturity from Bergman Lm range 71 - 84 cm from Fry et al. (2001). Lm range 76 - 94.7 cm from Fry et al. (2001). Length at the beginning of maturity from Bergman Length at the beginning of maturity from Bergman  (1943). (1943).  (1943). (1943). (1943). (1943).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  81  Table A3. Continued. Spec. No. 14 15 16  Stock No. a a b c a  Species  Hydrophis czeblukovi Hydrophis elegans Hydrophis fasciatus  b 17 18 19 20 21  a b c d a b c a b c a  Hydrophis inornatus  Hydrophis ornatus Hydrophis pacificus Lapemis hardwickii  b  22  c d e a  Laticauda semifasciata  b 23  a  Thalassophis anomalus  b 24  a  Caretta caretta  (Testudines, Cheloniidae) 25  a  Chelonia mydas  26  a b  Lepidochelys kempii  Locality northern Australia northern Australia northern Australia northern Australia Sourabaya, Java, Indonesia Sourabaya, Java, Indonesia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia northern Australia Sourabaya, Java, Indonesia Sourabaya, Java, Indonesia northern Australia northern Australia northern Australia near Orchid Island, Taiwan near Orchid Island, Taiwan Sourabaya, Java, Indonesia Sourabaya, Java, Indonesia Merritt Island, Florida, USA Merritt Island, Florida, USA Gulf of Mexico, USA Florida, USA  Sex  Lm  Comments/Remarks  F unsexed M F F  (cm) 98 118 89 118 65  Lm min 98 cm from Fry et al. (2001). Lm max 227 cm from Milton (2001). Lm range 51.2 - 172 cm from Fry et al. (2001). Lm range 90.4 - 22.7 cm from Fry et al. (2001). Length at the beginning of maturity from Bergman (1943).  M  60  Length at the beginning of maturity from Bergman (1943).  F unsexed M F unsexed M F unsexed M F M  92 63.5 78 63.5 80 85 80 135 141 44  Lm min 92 cm from Fry et al. (2001). Lm max 91.2 cm from Milton (2001). Lm range 76 - 91.2 cm from Fry et al. (2001). Lm range 35.1 - 82 cm from Fry et al. (2001). Lm max 163 cm from Milton (2001). Lm range 81.2 - 126 cm from Fry et al. (2001). Lm range 70 - 157.4 cm from Fry et al. (2001). Lm max 165 cm from Milton (2001). Lm min 141 cm from Fry et al. (2001). Lm range 135 - 165 cm from Fry et al. (2001). Length at the beginning of maturity from Bergman (1943).  F  44  Length at the beginning of maturity from Bergman (1943).  unsexed M F M  67.7 54 67.7 70  F  80  M  42.5  Length at the beginning of maturity from Bergman (1943).  F  42.5  Length at the beginning of maturity from Bergman (1943).  unsexed  -  unsexed  -  unsexed unsexed  60 62.5  Lm max 125 cm from Milton (2001). Lm range 44.2 - 118 cm from Fry et al. (2001). Lm range 33 - 113 cm from Fry et al. (2001). Minimun SVL at sexual maturity; Ave. water temperature of Kuroshio current 26°C (21-29°C). Lm range 70-80 cm from Tu et al. (1990). Tu et al. (1990)  Estimated VBGF range of age at maturity; tm= 12-30. Lm range 74-92 cm from Frazer and Ehrhart (1985). Estimated VBGF range of age at maturity; tm= 18-27. Lm range 88-99 cm from Frazer and Ehrhart (1985). Stranded head-starts; tm= 10 from Coyne (2000). Recaptured wild and head-started turtles; tm= 9 from Coyne (2000).  82  Growth of leatherback sea turtles, Jones, T.T. et al.  GROWTH OF LEATHERBACK SEA TURTLES (DERMOCHELYS CORIACEA) IN CAPTIVITY, WITH INFERENCES ON GROWTH IN THE WILD 1  T. Todd Jones  Department of Zoology, University of British Columbia, 6270 University Boulevard., Vancouver, BC, V6T 1Z4, Canada; E-mail: tjones@zoology.ubc.ca  Mervin Hastings  Department of Zoology, University of British Columbia, 6270 University Boulevard., Vancouver, BC, V6T 1Z4, Canada and Conservation and Fisheries Department, Ministry of Natural Resources and Labour, Government of the British Virgin Islands, Road Town, Tortola, BVI  Brian Bostrom  Department of Zoology, University of British Columbia, 6270 University Boulevard., Vancouver, BC, V6T 1Z4, Canada  Daniel Pauly  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada  David R. Jones  Department of Zoology, University of British Columbia, 6270 University Boulevard., Vancouver, BC, V6T 1Z4, Canada  ABSTRACT Leatherback turtles (Dermochelys coriacea) are critically endangered with current population trends in the Pacific indicating that they are nearing extinction. Their recovery will depend on coupling strong conservation measures with knowledge of their life history, particularly growth. Until now, however, there was considerable uncertainty on the growth on both juvenile and adults in the wild. The research reported here marks the first time that several leatherback juveniles have been maintained for over two years in captivity, and we discuss our experiences raising these leatherbacks from hatchlings (50 g) to juveniles (> 40 kg) for studies on their early growth. We derived a length-weight relationship of the form W (kg) = 0.000264 · SCL (cm)^2.806, which fitted both ours, and 10 turtles sampled from the wild. Also, a von Bertalanffy growth curve was derived whose parameters (SCL∞ = 155 cm; K = 0.266 year-1 and t0 = 0.12 year) predicts, for a length at first maturity of 135 cm, an age of 7 years, in agreement with earlier studies of the hard parts of leatherbacks. These results are in agreement with the known biology of leatherbacks; some of their implications for the study of leatherback biology are discussed.  1 Cite as: Jones, T.T., Hastings, M., Bostrom, B., Pauly, D., Jones, D.R., 2008. Growth of leatherback sea turtles (Dermochelys coriacea) in captivity, with inferences on growth in the wild. In: Palomares, M.L.D., Pauly, D. (Eds.), Von Bertalanffy Growth Paramters of Non-fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 82-91.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  83  INTRODUCTION All seven species of marine turtle are threatened, with several species listed as endangered or critically endangered (IUCN, 2007). Detailed knowledge of their life-history, notably the time they spend in various feeding grounds and their age at first maturity is essential for conservation (Seminoff et al., 2002; Chaloupka & Musick 1997). This requires a knowledge of growth rate at all stages (or of size-at-age), which is best summarized by the parameters of a growth equation, e.g., the von Bertalanffy Growth Function (VBGF) for length and weight (von Bertalanffy, 1938). Once a ‘standard growth curve’ has been established, it is then straightforward to evaluate growth in different populations, which should aid in our understanding of the variability in geographically separated foraging grounds and allow quantitative and qualitative comparisons of the foraging areas, based on the ability of a habitat to meet the ecological requirements of marine turtles (Bjorndal & Jackson, 2003; Bjorndal et al., 2000; Bjorndal & Bolten, 1988). Most studies of marine turtle growth have focused on the cheloniid species (see Chaloupka & Musick, 1997, Palomares et al., 2008) while relatively few have focused on leatherbacks. This is not surprising when considering the near-exclusive oceanic lifestyle of leatherbacks, of which the female go on land only for nesting, and the near impossibility of maintaining them in captivity (Jones et al., 2000). Yet, leatherbacks are listed as critically endangered (IUCN, 2007) and may be nearing extinction in the Pacific (Spotila et al., 2000). Within two decades the number of adult females in the Pacific declined from ~ 91,000 to under 3,000 (Spotila et al., 2000; 1996). We need to have information on the basic biology of leatherbacks, including demographics and life-history patterns, if we are to stop, and hopefully reverse, their decline. Leatherbacks are the largest (Buskirk & Crowder, 1994) of the marine turtles, but there are few reports on adult growth rates (Price et al., 2004; Zug & Parham, 1996). The growth of juvenile leatherbacks in the wild, moreover, is completely unknown, due to their distribution being largely unknown, thus precluding marking-recapture studies of their growth. Marking-recapture studies with marine turtles other than leatherbacks suggest they reach sexual maturity at an age of 20-30 years (Chaloupka & Musick, 1997), but recent evidence based on the study of hard parts in wild leatherbacks suggests an early attainment of minimum nesting sizes, i.e., as early as 3-6 years (Rhodin, 1985), or 6 years (Zug & Parham, 1996). Herein, we describe how we derived the parameters of the VBGF for length and weight growth in leatherback, by combining and harmonizing the results of several studies, notably our own growth experiment on captive leatherbacks, i.e., 20 hatchlings raised from emergence to > 2 years of age in the laboratory. We then suggest, in the light of the coherence of the results obtained, that the growth curves presented below can serve as standard growth curves for leatherback turtles.  MATERIALS AND METHODS Captive rearing experiments Leatherback turtles were obtained on Canada CITES import permit CA05CWIM0039 and British Virgin Islands CITES Export certificate CFD062005. These animals are housed and maintained for research purposes and we meet all the ethical animal care standards as put forth by the Canadian Council for Animal Care (CCAC) and the UBC Animal Care Committee (UBC Animal Care Protocol: A04-0323). Twenty hatchlings (emergence July 2nd, 2005) were transported from Tortola, BVI to the Animal Care Center, Department of Zoology, University of British Columbia. Animals were reared at the South Campus Animal Care facility using protocols developed by Jones et al. (2000). The three main obstacles to overcome in rearing leatherbacks are (i) their oceanic-pelagic nature (no recognition of barriers), (ii) designing a food matching their gelatinous food in the wild, and (iii) water quality. As leatherbacks are oceanic-pelagic animals, which do not recognize vertical (tank walls) and horizontal barriers (tank bottom), the animals were tethered to PVCTM pipes secured across the tops of the tanks. Animals < 10 kg were attached to the tether using VelcroTM and cyanoacrylate cement attaching the tether  84  Growth of leatherback sea turtles, Jones, T.T. et al.  to the posterior portion of their carapace, thus confining them to a section of the tank. Each hatchling could swim or dive in any direction, but was unable to contact other turtles or the tank’s bottom and walls. Upon reaching ≥ 10 kg the juveniles were secured to the tether with a harness made of TygonTM tubing. The harness circled each shoulder like a backpack and then looped around the caudal peduncle of the animal. Harnessing the leatherbacks is necessary as they swim continuously and, failing to recognize physical barriers, would abrade their skin against such barriers, which would lead to infections and usually death (Jones et al., 2000). The turtles were fed 3 to 5 times daily to satiation during the first 2-months of age and 3 times daily to satiation when > 2 months of age on a squid gelatin diet. The diet consists of squid (Pacific Ocean squid; mantle and tentacles only), vitamins (ReptaviteTM) and calcium (Rep-CalTM), blended with flavorless gelatin and hot water. As the wild diet of leatherbacks consists solely of gelatinous zooplankton (i.e., jellyfish; see Pauly et al. 2008), it is necessary for the food to have the proper texture and consistency. The food was weighed (Ek-1200 A; Stites Scale Inc., 3424 Beekman Street, Cincinnati, OH 45223) prior to feeding and notes were made as to individual food mass intake per day. The food had a water content of 90 % water, and an energy content of 20.16 ± 0.39 kJg-1 (dry weight). Random food samples were dried in a desiccating oven at 60°C for 24 to 72 hours to determine the dry to wet weight ratio. The dried homogenized samples were then sent to the Southwest Fisheries Science Center of NOAA (La Jolla, California, USA) for analysis with a bomb calorimeter (Parr Instrument Co.). The turtles were maintained in large oval tanks (5 m long x 1.5 m wide x 0.3 m deep) containing ~ 2,500 l of re-circulated/filtered salt water. Water temperature was maintained at 24 ± 1 oC. Four fluorescent fixtures (40 W UVA/B; Repti-Glow 8) suspended 0.5 m above each pool provided full spectrum radiation on a 12/12 hour cycle; also, each tank received ambient light. Water quality was maintained to the following levels pH = 8.0 to 8.3; salinity = 28-33, and ammonia < 0.1 mg-1. Water quality for each pool was maintained by four systems: a biological filter, a sand filter (Triton II™), an ultraviolet filter (Aqua Ultraviolet™ 114 W UV water sterilizer) and a protein skimmer. The turtles were weighed and measured on emergence, at 3 and 7 days of age, then weekly. Straight carapace length (SCL), the distance from the center of the nuchal notch to the caudal peduncle (posterior of the carapace), was used for all length measurements, and performed with a digital caliper to the nearest 0.1 mm. The turtles were weighted using an Ek-1200 A scale (Stites Scale Inc., 3424 Beekman Street, Cincinnati, OH 45223) from hatching to weights of 1.2 kg (± 0.1g), and an ADAM CPW-60 scale (Dynamic Scales, 1466 South 8th Street, Terre Haute, IN 47802) for weights ≥ 1.2 kg (± 0.02 kg). Length-weight relationships and growth curves We fitted the available length-weight data pairs (Table 1 and 2) with a length weight relationship of the form:  W = a·Lb  … 1)  where W is the weight in kg, L the SCL in cm, a is a multiplicative parameter of dimension L·W-1, and b is an exponent usually taking values near 3 (which then indicates isometric growth, and allows interpretation of ‘a’ as a condition factor; Pauly, 1984). Equation (1) was fitted by first transforming the data of Table 1 into log10Wi - log10Li pairs, and fitting these by a linear regression of the form:  log10Wi = α+b·log10Li  … 2)  where antilog α = a, and all other parameters are as defined previously. The VBGF for length has the form:  Lt = L∞(1 - e-K(t-t0))  … 3)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  85  where Lt is the predicted length at age t, L∞ is the mean the adults of the population in question would reach if they were to grow for a very long time (indefinitely, in fact), K is a growth parameter (not a growth rate) of dimension time-1, and t0 is the age of the turtles at length=0. It is a property of the VBGF that its first derivative (dl/dt) declines linearly with length, reaching zero at L∞. Hence, its parameter K can be estimated by plotting observed growth increments (Δl/Δt) against the mid-lengths of the increments (Pauly, 1984; Gulland & Holt, 1959), or  Yi = a - K X i  … 4)  where Yi = Li2-Li1/t2-ti1, X i = Li1+Li2/2, and Li1 and Li2 are length measurements taken at the start and end of an arbitrary time interval ti1 to ti2. Also, we have L∞ = a/K. This method leads to robust estimate of K, provided that the intervals ti1 to ti2 are relatively short, as in this case (Gulland & Holt, 1959). Its main advantage is that it provides for visualization of the data, and thus to identify outliers or incompatible data sets (Pauly, 1984). The method can also be modified to allow for estimation of K even when growth Y i/(L∞ - X ¯ increments are available only for juveniles. In such cases, a forcing value of L∞ is used, and K = ¯ ) (Pauly, 1984). We used 155 cm SCL (mean length of nesting females) as forcing value of L , based on i ∞ studies in both the Atlantic (Boulon et al., 1996) and the Pacific (Price et al., 2004). Another approach to fitting the VBGF is iterative, non-linear fitting (e.g., Fabens, 1965). Here, this was performed using the Sigma Plot software, with L∞=155 cm as constraint, given that the narrow range of the length-at age data fitted (Table 1) would not have otherwise lead to convergence. The VBGF for weight has the form:  Wt = W∞(1 - e-K(t-t0))b  … 5)  where W∞ is the weight corresponding to L∞, e.g., as estimated by Equation (1), b the exponent of that same length-weight relationship, and all other parameters are defined as for the VBFG for length (Equation 3).  RESULTS AND DISCUSSION 80 70 60 Body w eight (kg)  The hatchlings averaged 0.046 ± 0.001 kg body mass and 6.32 ± 0.13 cm SCL (straight carapace length) upon emergence. All hatchlings began feeding on the formulated squid gelatin by 3-5 days post emergence. Four turtles survived 18 months post emergence, with only 2 surviving more than 2 years. The largest animal was 42.65 kg and 72.0 cm SCL at 26 months old (age at death). Due to space constraints, we give here only a subset of the length and weight measurements taken during the life span of all 20 hatchlings (Table 2). Despite the deaths, the feeding regime seemed adequate, as assessed by the fact that our captive animals matched the condition of wild leatherbacks (Figure 1). The  Hawaii longline  50 40 30  this study Florida 2006  20  Western Australia unknown Florida 2005  10  American Samoa Western Australia  0 0  20  40 60 Straight carapace length (cm)  80  100  Figure 1. Plot of weight vs. length in 20 leatherbacks turtles maintained in captivity, from hatchlings to > 2-years (this study, Table 1) compared with weight vs. length from strandings and by-catch (Table 2). The overlap between the two data sets suggests that conditions for the captive turtles corresponded to those in the wild (c.f. with Figure 2).  86  Growth of leatherback sea turtles, Jones, T.T. et al.  relationship we obtained from the N = 101 log-transformed length and weight data pairs in Tables 1 and 3 (r2 = 0.998) is:  W = 0.000264·L2.806  … 6)  where W is the weight in kg and L the SCL in cm. 60 50  Body weight (kg)  The length and weight data pairs from our study match those of leatherback taken from the wild (Figure 1), and hence equation (4) may be proposed as standard L-W relationship for leatherback turtles. On the other hand, the data in Figure 1, and Equation (4) suggest that the turtles raised by Deraniyagala (1939) and Bels et al. (1988) suffered from sub-optimal condition, notably inadequate nutrition (i.e., algae, beef heart, and French bread; see Table 2), resulting in elevated mortality (Table 2), emaciation (Figure 2), and reduced growth (see below).  40 30 20 10 0 0  20  40  60  80  Straight carapace length (cm)  Figure 2. Length-weight relationships of leatherback turtles. Solid black line: relationship based on length and weight (open dots) of the turtles we maintained in captivity from hatchlings to > 2-years (this study, Table 1). Thin black line: relationship based on the turtles (black squares) reared by Bels et al. (1988). Dotted line: relationship based on the turtles (black triangles) reared by Deraniyagala (1939). The low weight at length of the turtles reared by Bels et al. and Deraniyagala suggest that they suffered from less than optimal conditions (c.f. with Figure 1).  Figure 3 contrasts the growth rates obtained in this study (Table 1) with those reported by Deraniyagala (1939) and Bels et al. (1988). Despite much variability, our turtles exhibited higher growth rates than theirs. Moreover, the juvenile growth rates we obtained appear compatible with the adult growth rates reported by Price et al. (2004). Figure 3 also demonstrates the compatibility of our results with those Zug and Parham (1996), who found that juvenile leatherback growth rates were 31.6 cm year-1 for juveniles 8-37 cm SCL and 23.1 cm year-1 for juveniles 37-65 cm SCL [data converted from curved-carapace lengths using the equation of Tucker & Frazer (1991)]. Our growth rate data, combined with a value of L∞ set at 155 cm allows estimation of a preliminary value of K = 0.232 from the slope of the plot in Figure 3. Fitted non-linearly, the same inputs yielded the VBGF for length:  Lt = 155(1-e-0.266(t+0.12))  … 7)  The resulting curve is shown in Figure 4, and contrasted with a curve based on the length-at-age data of Deraniyagala (1939) and Bels et al. (1988). As might be seen, our juvenile growth data suggest faster growth than theirs, as also shown in Figure 3. Using 135 cm SCL as the minimum size at nesting, based on Boulon et al. (1996) for the Atlantic and Price et al. (2004) for the Pacific, Equation (8) suggests that it would take leatherbacks 7 years to reach sexual maturity, in agreement with the 6 years proposed by Zug & Parham (1996). Combining Equation (6) with (7) leads, finally, to a VBGF for the growth in weight in leatherbacks, i.e.:  Wt= 370(1-e-0.266 (t+0.12))2.806  ... 8)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  which can be used to predict mean weight at any age.  75 65 Growth increment (cm year-1)  Major assumptions have been made in the experimental design of this study and for the results to have any validity they must be addressed. Firstly, the VBGF requires that growth be monotonic throughout postnatal development, as it displays no inflection points (Choulpka & Musick, 1997). Therefore, polyphasic growth data, or displaying an initial lag phase, would require another growth function, e.g., the Gompertz, logistic or others. However, the leatherback turtles we raised, and our longitudinal sampling (repeated sampling on the same individuals; Choulpka & Musick, 1997) resulted in growth data exhibiting neither polyphasic growth, nor a lag phase. Therefore, the use of the VBGF is justified in our case, and by extension, in leatherbacks as a whole. We also suspect this to be the case in other species of marine turtles, as well.  87  55 45 35 25 15 5 0 -5 0  50  100  150  Straight carapace length (mid-length, cm)  Figure 3. Plot of growth rates (Δl/Δt) against the corresponding midlengths of the growth increments in leatherback turtles, computed from Table 1 (open dots, our study), the studies of Deraniyagala (1939) and Bels et al. (1988) (black triangles), Zug & Parham (1996) (2 black dot) and adult growth rates from Price et al. (2004) (open squares). The solid line links the means of the values from our study (open dots) and L∞ = 155 cm (SCL); its slope allows a preliminary estimation of K = 0.232 year-1. The data points from Deraniyagala (1939) and Bels et al. (1988) were omitted, as their turtles probably experienced suboptimal condition (c.f. Fig. 2, and see text).  160  Straight carapace length (cm)  Captive growth does not 140 necessarily reflect wild growth. 120 However, our captive specimens exhibited the same length-weight 100 relationships as wild juvenile leatherbacks (stranded or by80 catch; Fig 1.), suggesting 60 appropriate rearing conditions at least compared with earlier 40 captive growth studies. On the 20 other hand, the problem of accelerated growth in captivity, 0 seem to be limited to cheloniids 0 5 10 15 20 (Swingle et al., 1993;Wood & Age (years) Wood, 1980), and may not occur in leatherbacks, whose chondroFigure 4. Von Bertalanffy Growth Functions for leatherback turtles: Solid osseous development line: VBGF with a fixed value of L∞ = 155 cm, K = 0.266 year-1and t0 = -0.12 characteristic suggests rapid year, based on length-at-age data in Table 1 (this study, open dots) fitted growth (Rhodin et al., 1996; with SigmaPlot™ version 10. Dotted line: same L∞ and fitting method, with Rhodin, 1985). Also, Zug & K = 0.185 year-1 and t0 = -0.03 year, derived from the length-at-age data in Parham (1996), whose growth Table 3 (i.e., from studies of Deraniyagala, 1939 and Bels et al., 1988, black data match ours almost perfectly triangles). The sub-optimal conditions suggested to have occurred in these (Figure 3), found rapid growth studies affected the growth of the turtles. rates in wild leatherbacks (15 adults and 2 juveniles) and stated that the early captive growth pattern of leatherbacks closely matches the growth curves of wild individuals.  88  Growth of leatherback sea turtles, Jones, T.T. et al.  Our findings confirm that leatherbacks mature a younger age (6-7 years, see above), but at a larger size than cheloniid turtles. For example, loggerheads take > 15 years to reach a sexually mature size of about 90 cm carapace length (Frazer & Ehrhart, 1985; Mendoca, 1981), whereas green turtles take > 20-30 years to reach sexual maturity at a carapace length of about 100 cm (Frazer & Ladner, 1986; Frazer & Ehrhart, 1985; Mendoca, 1981; Limpus & Walter, 1980). Similarly, green turtles with size of 30 cm spend nearly 20 years in juvenile habitats, before they acquire adult features (Seminoff et al., 2002; Bjorndal & Bolten, 1988). Table 1. Length and weight of 20 turtles raised in captivity from hatchling to ages of over 2 years, using the protocol and feed described in the text. N = 20 ≤ 12 months; 4 from 12 to 18 months; 2 from 18 months to > 24 months. Turtle ID Dc 7 Dc 7 Dc 7 Dc 8 Dc 8 Dc 8 Dc 9 Dc 9 Dc 9 Dc 9 Dc 10 Dc 10 Dc 10 Dc 10 Dc 10 Dc 10 Dc 10 Dc 10 Dc 10 Dc 11 Dc 11 Dc 11 Dc 11 Dc 11 Dc 11 Dc 12 Dc 12 Dc 12 Dc 12 Dc 12 Dc 12 Dc 12 Dc 12 Dc 13  Age (days) 1 31 73 1 31 73 1 31 73 157 1 31 73 157 206 304 402 500 628 1 31 73 150 206 255 1 31 73 150 206 304 402 479 1  Weight (kg) 0.048 0.139 0.355 0.046 0.129 0.342 0.047 0.123 0.326 1.280 0.046 0.124 0.335 1.220 2.180 5.420 10.900 12.060 21.240 0.046 0.105 0.264 0.943 2.000 2.960 0.046 0.111 0.303 1.146 2.460 5.620 10.420 13.040 0.046  SCL (cm) 6.37 9.25 13.17 6.10 8.78 13.29 6.41 8.82 12.85 20.69 6.42 9.03 12.99 20.20 25.10 34.46 44.57 47.50 55.80 6.04 8.23 11.90 18.38 23.65 26.74 6.44 8.55 12.59 19.71 25.73 34.47 43.87 48.40 6.19  Turtles experience strong ontogenic habitat shifts. Thus, green turtles enter the oceanicpelagic habitat as posthatchling, and then turn into coastal-benthic feeders as juveniles (Bjorndal & Bolten, 1988), which probably induce a shift from an omnivorous to a herbivorous diet. These ontogenic habitat, diet and hence niche shifts may be the reason why the somatic growth of marine turtles often appears to be polyphasic (Hendrickson, 1980; Chaloupka & Musick, 1997). Leatherbacks, however,  Turtle ID Dc 13 Dc 13 Dc 13 Dc 13 Dc 14 Dc 14 Dc 14 Dc 14 Dc 14 Dc 14 Dc 14 Dc 14 Dc 14 Dc 15 Dc 15 Dc 15 Dc 16 Dc 16 Dc 16 Dc 16 Dc 16 Dc 17 Dc 17 Dc 17 Dc 18 Dc 18 Dc 18 Dc 19 Dc 19 Dc 19 Dc 19 Dc 19 Dc 19 Dc 19  Age (days) 31 73 157 206 1 31 101 157 206 304 402 507 611 1 31 122 1 31 73 157 248 1 31 73 1 31 66 1 31 73 157 206 304 402  Weight (kg) 0.115 0.305 1.260 2.140 0.048 0.115 0.489 1.180 2.160 5.460 11.000 17.280 25.600 0.046 0.133 0.580 0.045 0.119 0.360 1.320 3.420 0.046 0.144 0.367 0.047 0.131 0.263 0.046 0.135 0.346 1.280 2.400 6.360 13.780  SCL (cm) 8.61 12.59 20.04 23.67 6.32 8.75 14.99 20.29 24.93 34.27 44.14 52.60 61.50 6.43 9.05 15.01 6.13 8.52 13.16 20.67 28.38 6.41 9.32 13.79 6.38 9.19 11.57 6.34 9.20 13.06 20.39 25.73 35.03 47.31  Turtle ID Dc 19 Dc 20 Dc 20 Dc 20 Dc 20 Dc 20 Dc 20 Dc 21 Dc 21 Dc 21 Dc 22 Dc 22 Dc 22 Dc 23 Dc 23 Dc 23 Dc 24 Dc 24 Dc 24 Dc 24 Dc 24 Dc 24 Dc 25 Dc 25 Dc 25 Dc 26 Dc 26 Dc 26 Dc 27 Dc 27 Dc 27 Dc 27 Dc 27 -  Age (days) 500 1 31 73 150 206 297 1 31 87 1 31 129 1 31 122 1 31 73 150 206 332 1 31 108 1 31 108 1 31 101 150 213 -  Weight (kg) 20.360 0.047 0.131 0.349 1.180 2.480 5.440 0.045 0.119 0.300 0.047 0.127 0.701 0.047 0.140 0.754 0.048 0.117 0.301 1.020 2.360 5.580 0.046 0.117 0.375 0.046 0.132 0.496 0.045 0.125 0.558 0.900 1.520 -  SCL (cm) 55.40 6.55 9.26 13.49 20.00 26.33 34.74 6.29 8.81 12.05 6.37 9.11 16.00 6.24 9.29 17.15 6.43 8.72 12.24 19.21 25.78 35.13 6.16 8.83 13.61 6.33 9.24 15.03 6.35 8.91 14.85 17.98 21.50 -  Table 2. Length and weight of 10 loggerhead turtles taken from the wild (stranded or as by-catch). Date, location and source are given for each turtle, except one, for which only the length and weight are known. Weight SCL Source Date Location (kg) (cm) Aug-93 Sep-05 Mar-06 Apr-98 Apr-99 Apr-06 Jul-06 Jul-02 1983 Unknown  American Samoa Florida (2005) Florida (2006) Hawaii Hawaii Hawaii Hawaii W. Australia W. Australia Unknown  7.00 0.19 3.10 44.50 74.10 35.45 33.60 1.85 3.30 0.17  39.0 10.4 25.0 70.4 85.3 70.0 67.5 20.0 31.0 11.5  MTN (1994; no 66, p. 3-5) J. Wyneken (pers. comm.) J. Wyneken (pers. comm.) NOAA (NMFS/PIFSC) NOAA (NMFS/PIFSC) NOAA (NMFS/PIFSC) NOAA (NMFS/PIFSC) MTN (2004; no. 104, p. 3-5) MTN (2004; no.104, p. 3-5) M. Conti (pers. comm.)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  Table 3. Length and weight at age of leatherback turtles raised from the hatchling stage to ages of over 1 year Deraniyagala (1939; initial N = 10; food: algae, beef hearts and French bread) and Bels et al. (1988; initial N = 14; food: mussels). Deraniyagala lost 90% his turtles in the first month, with 2 lasting 169 days, and 1 from day 169 to 662. Bels et al. lost 70% of their turtle within 2-months, with 1 lasting from day 183 to 1351.  89  are oceanic-pelagic animals throughout their life-history (Bolten, 2003) and do not exhibit an ontogenetic diet shift; the diet consists solely of gelatinous zooplankton, throughout all life-history stages (Salmon et al., 2004; Bjorndal, 1997). This, then, would justify the use of the VBGF.  Eckert (2002) used reports of visual sightings and incidental captures in north Atlantic to show that leatherbacks do not move above ~30̊ N and into water < 26 ̊C until they are over 100 cm in carapace length, corresponding given Equation (6) and (7), to an age of 3.8 years, and a weight of 108 kg, respectively. The latter value, used as an input for the leatherback thermoregulatory model of Bostrom & Jones (2007), suggest that these leatherbacks could maintain body temperatures 1.63 to 8.15 ̊C above ambient temperatures. This would allow them to move into colder waters where they can exploit different assemblages and perhaps greater abundance of gelatinous zooplankton, without their metabolism and growth being much reduced by the lower ambient temperatures. A review of reptilian growth by Avery (1994) showed that growth was not affected by cooler temperatures when the organisms were allowed to behaviorally thermoregulate. Although leatherback thermoregulation is endogenously driven, it is also a consequence of a large mass and locomotion (Bostrom & Jones, 2007). Thus, the benefit of higher body temperatures with regards to growth rates would not be lost to increased thermoregulatory costs. Deraniyagala (1939) Age Weight SCL (days) (kg) (cm) 1 0.033 5.9 21 0.096 8.5 22 -7.3 32 -8.5 32 -8.9 46 -10.2 91 -13.7 169 -16.0 183 -25.4 195 -25.5 203 2.438 -218 3.005 30.2 308 -35.0 344 -35.6 466 4.536 36.8 562 6.804 43.3 586 7.258 43.3 624 7.265 43.5 662 -42.0  Bels et al. (1988) Age Weight SCL (days) (kg) (cm) 1 0.046 6.1 41 0.047 6.2 85 0.075 9.6 239 0.312 14.7 478 0.950 21.2 506 1.125 22.8 726 3.720 -847 4.500 -928 8.020 47.0 1140 20.000 61.7 1200 28.500 82.0 1351 49.500 85.0 ----------------------  The decline in the Pacific leatherback population is daunting. The presumed cause is decades of intense egg harvest at most nesting beaches, exacerbated by widespread incidental by-catch from fisheries practices (Eckert & Sarti, 1997). Although the numbers of adults are higher in the Atlantic (~30,000), fishing practices continue to take their toll and the numbers from artisanal fisheries is unknown but probably severe (Peckham et al., 2007). The good news is that with 7 years time to first nesting, leatherbacks still have a chance, as there is potential for a rapid rebound (at least compared with the slowgrowing cheloniids) if fisheries by-catch can be reduced through moratoria and regulation.  ACKNOWLEDGEMENTS This work would not have been possible without the help and cooperation of Gaverson ‘Gary’ Frett and Arlington ‘Zeke’ Pickering of the Conservation and Fisheries Department (CFD), British Virgin Islands. As well, we thank the CFD, BVI for granting us permission to study and rear leatherbacks. Our gratitude also goes to Ms Colette Wabnitz (Fisheries Centre, UBC) for invaluable assistance, and Dieta Lund (Zoology, UBC) for keeping track of our rearing data. We thank Jeanette Wyneken, Bob Prince, M. Conti and NOAA, National Marine Fisheries Service, Pacific Islands Fisheries Science Center for data on stranded, and bycaught juvenile leatherbacks. We also thank Ashley Houlihan, Katerina Kwon, Amir Shamlou, Dieta Lund, Erika Kume, AndrewYamada, Thea Sellman, Oliver Claque, Brian Woo, Angela Stevenson and Dana Miller, all UBC undergraduate students for their care of the leatherbacks housed at the Animal Care Center, Department of Zoology, UBC as well as Art Vanderhorst and Sam Gopaul (Turtle emergency care), Bruce Gillespie and Vincent Grant (for everything mechanical) and Chris Harvey-Clark, Bob George and Tamara Godbey for clinical assistance. 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LENGTH–WEIGHT RELATIONSHIPS AND ADDITIONAL GROWTH PARAMETERS FOR SEA TURTLES 1  Colette Wabnitz  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email:c.wabnitz@fisheries.ubc.ca  Daniel Pauly  The Sea Around Us Project, Fisheries Centre, UBC, 2202 Main Mall, Vancouver, B.C V6T 1Z4, Canada; Email:m.pauly@fisheries.ubc.ca  ABSTRACT To facilitate field and other work on sea turtles, composite length-weight relationships, based on a wide range of sizes sampled by various authors, are presented for five species, viz. Kemp’s ridleys (Lepidochelys kempi), olive ridleys (Lepidochelys olivacea), loggerheads (Caretta caretta), greens (Chelonia mydas), and hawkbills (Eretmochelys imbricata). Also, 38 pairs of growth parameters of the von Bertalanffy growth function (VBGF; K; L∞ and W∞) are presented for four species, leaving only the growth of the olive ridley undocumented.  INTRODUCTION There are seven living species of sea turtles: flatback (Natator depressus), green sea turtle (Chelonia mydas), hawksbill (Eretmochelys imbricata), Kemp's Ridley (Lepidochelys kempi), leatherback (Dermochelys coriacea), loggerhead (Caretta caretta), and olive ridley (Lepidochelys olivacea). Populations of all these species are threatened throughout the world because of overexploitation, disease, incidental capture by fishers, and destruction of critical nesting habitat (Lutcavage et al., 1997; Mortimer et al., 2000; Lewison et al., 2004; Peckham et al., 2008). Intensive, and sometimes sophisticated research has been conducted to quantify these impacts and inform management practices (e.g., Chaloupka & Balazs, 2007; Bailey et al., 2008; e.g., Sims et al., 2008). In the process, however, basic biological data are frequently overlooked. This applies particularly to morphometric relationships, whose validity is often taken for granted, although they tend to be based on too small a range of sizes to be of any use in building more elaborate models, e.g., turtle growth studies. This contribution presents key morphometric data for 5 species of sea turtles, namely Kemp’s ridleys (L. kempi), olive ridleys (L. olivacea), loggerheads (C. caretta), greens (C. mydas), and hawkbills (E. imbricata), and complements two other works in this volume, Jones et al. (2008) for leatherbacks and Palomares et al. (2008) for reptiles (including sea turtles).  MATERIAL AND METHODS The relationship between total length (L) and weight (W) for most animals is expressed by the equation:  W = a•Lb  …1)  whose parameters (a, b) are estimated by the antilog of the intercept, and the slope, respectively, of a regression of the log10 W against log10 L. The value of b is generally close to 3, implying ‘isometry’, i.e., the shape of the animal in question remaining the same as they get older and gain in size. 1 Cite as: Wabnitz, C., Pauly, D., 2008. Length–weight relationships and additional growth parameters for sea turtles. In: Palomares, M.L.D., Pauly, D. (Eds.), Von Bertalanffy Growth Parameters of Non-fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 92-101.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  93  Table 1. Empirical equations used to convert curved carapace length (CCL; cm) into straight carapace length (SCL; cm) measurements for individual species. Species  Lepidochelys kempi Lepidochelys olivacea Caretta caretta Chelonia mydas Eretmochelys imbricata Eretmochelys imbricata  R2 0.99 0.91 0.97 0.93 n.a. 0.99  Equation SCL = 0.957 * CCL - 0.696 SCL = 0.818 * CCL + 9.244 SCL = 0.948 * CCL – 1.442 SCL = 0.932 * CCL + 0.369 SCL= 0.939 * CCL - 0.154 SCL = 0.935 * CCL + 0.449  Reference Plotkin (2007) Whiting et al. (2007) Teas (1993) Peckham et al. (2008) CITES (2002) Limpus (1992) – for Australia  Sea turtles can be measured in a number of ways, requiring standardisation before datasets can be compared. Straight carapace length (SCL) and curved carapace length (CCL) are the most commonly used measurements taken of sea turtles. As their name implies, CCL measurements are taken over the curve of the carapace whereas straight measurements are taken with a set of callipers. Although variations exist in how these measurements can be taken (e.g., notch to notch [NN] or notch to tip [NT]), authors most often do not detail the specific technique used in measuring individuals beyond curved or straight. For the purposes of this analysis, we assumed discrepancies to be minimal. Where necessary, data were converted to SCL using empirical equations listed in Table 1, based on linear regression of paired CCL and SCL data for the species in question. To ensure that the parameters of length-weight relationships are estimated properly (Safran, 1992), length-weight data pairs from different studies were compiled to cover the widest possible range of sizes, and all developmental stages, i.e., juveniles, subadults, and adults (Table 2). Table 2. Length weight relationships for 5 species of sea turtles; a and b are parameters in the equation of the type W=a L3. Species  Location  a  b  r2  N  Lepidochelys kempi  Chesapeake, Florida, UK & France  0.000247  2.834  0.958  145  Size range (SCL; cm) 19-67  Caretta caretta  Chesapeake, Florida, UK & France, Japan  0.000282  2.823  0.966  431  12-105  Chelonia mydas  Florida, Tortuguero, Ascension, Suriname, Baja, Solomon Islands  0.000206  2.895  0.992  449  5-124  Lepidochelys olivacea  Hawaii, Brazil, Suriname, Mozambique, Thailand, Australia  0.000479  2.673  0.9955  46  4-74  Eretmochelys imbricata  Honduras, Cayman, Barbados, Suriname  0.000278  2.736  0.988  112  22-99  References  Carr & Caldwell (1956); Byles (1988); Campbell & Sulak (1997); Coles (1999); Witt et al. (2007) Byles (1988); Sato et al. (1995); Barichivich et al. (1997); Campbell & Sulak (1997); Coles (1999); Witt et al. (2007) Carr & Caldwell (1956); Pritchard et al. (1969); Barichivich et al. (1997); Campbell & Sulak (1997); (2000); Gilbert (2005); Seminoff et al. (2006); CCC (Unpublished); Krueger (unpublished); Seminoff & Jones (Seminoff & Jones) Pritchard et al. (1969); Hughes (1972); Chantrapornsyl (1992); Work & Balazs (2002); de Castilhos & Tiwari (2007); WWF-Australia (WWF-Australia) Pritchard et al. (1969); Beggs et al. (2007); Blumenthal et al. (2008); Dunbar et al. (2008)  94  Length-weight relationship and growth of sea turtles, Wabnitz, C. & Pauly, D.  Although other growth curves exist to describe the growth of sea turtle (e.g. Bjorndal & Bolten, 1988; Chaloupka, 1998; Bjorndal et al., 2000a; Chaloupka et al., 2004), we have used the von Bertalanffy growth function (VBGF; von Bertalanffy, 1938) to ensure compatibility with the other growth parameters in this report. The VBGF for length has the form:  Lt = L∞(1 – e-K(t-t0))  …2)  where Lt is the predicted length at age t, L∞ (also Linf) is the mean the adults of the population in question would reach if they were to grow for a very long time (indefinitely, in fact), K is a growth parameter (not a growth rate) of dimension time-1, and t0 is the age the turtles at length = 0. Using the parameters K (quantifying the curvature of the VBGF), and L∞ (or W∞, Winf) one can then summarize and compare growth data by means of so called auximetric plots (Pauly, 1998). The parameters K and L∞ used for this analysis were taken from the published literature (see Table 3). Length-weight (L/W) relationships for each species, as described in Table 2, were then used to calculate W∞ (Table 3).  RESULTS AND DISCUSSION Table 1 summarizes available relationships between SCL and CCL, while Table 2 summarizes the L/W relationships and related data. The r2 values for all L/W relationships were greater than 0.95. Estimates of parameter b ranged from 2.673 for olive ridleys to 2.895 for green turtles. When split into individual ‘populations’ for each species b spanned values between 2.495 and 3.134. This increased range in estimates reflected differences in ‘population’ sample sizes and length ranges. The L/W relationships for all 5 species, and the ‘population’ data used to derive them, are presented in Figure 1. One potential application of such length-weight relationships is the computation of biomass estimates from length-frequency distributions. This is of great value when, for example, site and season-specific weights have not been collected due to logistical difficulties and/or lack of time required to record weight in the field. Although weight can be reliably estimated from length using equations such as those presented here, it should be noted that the exact relationship between length and weight may differ depending on the ‘condition’ of individual animals. Condition may reflect differences in food availability and population densities at individual sites (Bjorndal et al., 2000a), and is likely to vary between seasons and years for a given population. In instances where the individuals of a population remain below the average curve, its individuals can be considered comparatively ‘skinny’; conversely, when individuals lie above the curve, they can be considered ‘stout’. Notably, the compiled data presented here highlight the importance of obtaining ‘true’ estimates of population parameters through comprehensive sampling of a species size range. Relationships derived from morphometric data for a location-specific population may be biased by being representative of only a narrow size range. For example, because the majority of sea turtle programs operate on nesting beaches, length-weight data pairs are likely to be primarily, if not solely, collected from mature females. This can lead to erroneous population-level L/W relationships, as the juvenile-subadult phase is missing.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D. 50  180  UK/France Chesapeake Florida  45  140  35  120 Weight (kg)  Weight (kg)  UK/France Chesapeake Florida Japan  160  40  95  30 25 20  100 80 60  15  40  10 5  20  W = 0.000247 SCL2.834 R2 = 0.96; n=145  0 0  10  20  30  40  50  60  70  W = 0.000282 SCL2.823 2 R = 0.97; n=431  0  80  0  20  Straight Carapace Length (cm)  40  60  80  120  100  Straight Carapace Length (cm)  A  B 300  70  Suriname Tortuguero Florida Baja Ascension Solomons  250  60 50 Weight (kg)  200 Weight (kg)  Suriname Hawaii Brazil Mozambique Australia Thailand  150  100  40 30 20  50  10 W = 0.000206 SCL2.896 R2 = 0.99; n=449  0 0  20  40  60  80  100  120  W = 0.000479 SCL2.678 R2 = 0.99; n=46  0 0  140  20  40  60  80  100  Straight Carapace Length (cm)  Straight Carapace Length (cm)  C  D 100  Suriname Barbados Cayman Honduras  90 80  Weight (kg)  70 60 50 40 30 20 10  2.736  W = 0.000278 SCL R2 = 0.99; N=112  0 0  20  40  60  80  Straight Carapace Length (cm)  E  100  120  Figure 1. Correlations between straight carapace length (SCL, cm) and weight (W, kg) for five species of sea turtles: A. Kemp’s ridley (Lepidochelys kempi); B. loggerhead (Caretta caretta); C. green (Chelonia mydas); D. olive ridley (Lepidochelys olivacea); E. hawksbill (Erytmochelys imbricata) discussed here.  ACKNOWLEDGMENTS CW would like to thank E. Harrison, B. Krueger, and TT Jones for the provision of unpublished biometric data for nesting green turtles at Tortuguero, Costa Rica; foraging hawksbills in Barbados and the Solomon Islands; green and loggerheads in Baja respectively. B. Hunt is kindly acknowledged for providing useful comments and constructive suggestions. This is a contribution of the Sea Around Us Project, initiated and funded by the Pew Charitable Trusts, Philadelphia.  0.0 -0.2 -0.4 -0.6 -0.8  -1  Table A1 summarizes the growth parameters (K, L∞ and W∞), while the auximetric plot of Figure 2, which does not include outliers, shows that these growth parameters are mutually consistent.  Length-weight relationship and growth of sea turtles, Wabnitz, C. & Pauly, D.  K (year ; log10)  96  -1.0 -1.2 -1.4 -1.6  y = -0.762x + 0.458 R2 = 0.70  -1.8 -2.0 0  0.5  1  1.5 2 W∞ (kg; log10)  2.5  3  3.5  Figure 2. Auximetric plot of von Bertalanffy growth parameters for 38 data pairs of four species of sea turtles (see Table 3 for details). Dark circles represent data for Lepidochelys kempi, open circles Caretta caretta, dark squares Chelonia mydas, and open squares Erytmochelys imbricata  REFERENCES Bailey, H., Shillinger, G., Palacios, D., Bograd, S., Spotila, J., Paladino, F., Block, B., 2008. 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Marine Biology 151, 873-885. Work, T., Balazs, G.H., 2002. Necropsy findings in sea turtles taken as bycatch in the North Pacific longline fishery. Fishery Bulletin 100, 876-880. WWF-Australia, 2008. Olive ridley turtle tracking: Turtle bios. Accessed 2008. http://www.wwf.org.au/ourwork/oceans/ oliveridleytrackingbios/#milika. Zug, G.R., Kalb, H.J., Luzar, S.J., 1997. Age and growth in wild Kemp's ridley seaturtles Lepidochelys kempii from skeletochronological data. Biological Conservation 80, 261-268.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  99  Table A1. Additional growth parameter estimates for 4 species of sea turtles. Method: MR=Mark recapture; SC=Skeletochronology; LF=Length frequency. All data are from wild sea turtles except for data by Caillouet (1995) for L. kempii. Reported average lengths from http://www.nmfs.noaa.gov/pr/species/turtles /loggerhead.htm. Species (reported average length; cm)  Lepidochelys kempii (56-79)  Caretta caretta (92)  (year-1)  (SCL; cm)  (kg)  W∞  Sample size  Gulf of Mexico  0.317  62.3  24.4  117a  Atlantic: Gulf of Mexico Atlantic: Cape Canaveral Atlantic: Cape Canaveral  0.129  80.0  49.5  36  21.5–60.3  Schmid & Witzell (1997) [MR]  0.577  61.1  23.0  12c  21.5-60.3  0.594  60.8  22.7  10  21.5-60.3  Atlantic Gulf of Mexico Atlantic: Gulf of Mexico Gulf of Mexico: Cedar Keys Atlantic Gulf of Mexico Atlantic Gulf of Mexico  0.215 0.219 0.079  58.9 70.5 87.7  20.8 34.6 64.2  56 15 70  Probably underestimated due to lack of adult sized Kemp’s ridley turtles in the database; Schmid (1995) [MR] 60% 20-40cm; probably underestimated due to lack of adult sized Kemp’s ridley turtles in the database; Schmid (1995) [MR] Zug et al. (1997) [SC] Zug et al. (1997) [SC] Zug et al. (1997) [SC]  0.085  91.4  72.2  24  Schmid (1998) [SC]  0.167 0.210 0.115 0.053  73.2 71.1 74.9 97.0  38.5 35.4 41.0 85.4  38 58 109 660  21.7–50.5 20-61  Turtle Expert Working Group (2000)b [SC, MR] Turtle Expert Working Group (2000) [SC, MR] Snover et al. (2007) [SC] Bjorndal & Bolten (1997) [LF]  Atlantic: Cape Canaveral Atlantic: Cape Canaveral  0.059  96.1  118  51c  38.2-110  0.037  112  185  17  38.2–110  Chesapeake Bay Atlantic (Florida, Georgia & South Carolina) Azores, North Atlantic Florida, Mosquito lagoon Florida  0.076 0.031  112 110  182 174  83 118  13-42 45–110  0.072  98.9  129  574  10-64  0.120  94.6  114  28  53.3-77.3  0.115  94.7  114  41  53.3–77.  North Carolina  0.052  107  160  57  45.1–75.8  Area  K  L∞  Size range (cm)  Comments; reference [method] Caillouet et al. (1995) [MR]  80%<80 cm SCL; 20%>80cm; Schmid (1995) [MR - Adults include males and females] Growth model for captures and recaptures by the contract vessel; size range for study but not specified for N=19; Schmid (1995) [MR] Klinger & Musick (1995) [SC] Size range for study, no specified for N=118; Henwood (1987) [MR] Assuming 105.5 CCL, where CCL=1.388+(1.053)(SCLnt); Bjorndal et al. (2000b) [LF] Frazer & Ehrhart (1985) [MR] Size range based on 8 individuals with specified lengths, 20 adults with lengths not specified, and 13 individuals with no specified lengths but assumed <82 cm; Frazer (1987) [MR] Braun-McNeill et al. 2002 in Epperly et al.(2001) [MR]  100  Length-weight relationship and growth of sea turtles, Wabnitz, C. & Pauly, D.  Table A1. Continued. Species (reported average length; cm)  Caretta caretta  (year-1)  (SCL; cm)  (kg)  W∞  Sample size  Florida  0.064  96.7  121  54  Size range (cm) 62.2–104.2  Georgia, Cumberland island Georgia, Cumberland island Georgia, Cumberland island Georgia, Cumberland island Georgia, Cumberland island Gulf of Mexico Florida, Atlantic coast  0.096  96.8  121  69  >49.76-103  0.098  102  138  25  >49.76–103  0.086  95.4  116  25  >49.76–103  0.106  108  163  26  >36.04–103  Reported in CCL and converted to SCL using SCL=(0.948×CCL)–1.442 ; Teas (1993); Parham & Zug (1997) [SC–1979 ; regression growth protocol] Reported in CCL and converted to SCL using SCL=(0.948×CCL)–1.442 ; Teas (1993); Parham & Zug (1997) [SC–resampled 1979 data–correction factor protocol] Reported in CCL and converted to SCL using SCL=(0.948×CCL)–1.442 ; Teas (1993); Parham & Zug (1997) [SC–resampled 1979 data–regression growth protocol Parham & Zug (1997) [SC – 1980 correction factor protocol]  0.074  109  170  26  >36.04–103  Parham & Zug (1997) [SC – 1980 regression growth protocol]  0.051 0.044  106 111  155 178  570 1234  >36.04–103 42.2-81.03  Texas Great Barrier Reef, Australia  0.030 0.060  144 105  372 151  819 172  46-87 63–90.3  Bjorndal et al .(2001) [LF] Reported in CCL and converted to SCL using SCL=(0.948×CCL)–1.442 ; Teas (1993); Bjorndal et al. (2001) [LF] Bjorndal & Bolten (1997) [LF] Reported in CCL and converted to SCL using SCL=(0.948×CCL)–1.442 ; Teas (1993); Frazer et al. (1994) [MR]  Florida, Mosquito lagoon Florida, Atlantic Inagua, Bahamas US Virgin Islands Watamu, Kenya  0.089  109  157  11  27.7->69.6  Frazer & Ehrhart (1985) [MR]  0.026 0.072  182 99.7  694 122  976 964  25-70 25-70  Bjorndal & Bolten (1997) [LF] Bjorndal & Bolten (1995) [LF]  0.048  148  379  41  25.6–62.3  0.068  117  195  563  31-108  Area  K  L∞  Comments; reference [method] Foster (1994) [MR]  (92)  Chelonia mydas (91)  Size range at first capture; Boulon & Frazer (1990) [MR] Reported in CCL and converted to SCL using SCL=0.932*CCL+0.369 ; Peckham et al. (2008) ; Watson (2006) [MR]  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  101  Table A1. Continued. Species (reported average length; cm)  Eretmochelys imbricata (63-90)  Area St Thomas, Virgin islands Mona Island, Puerto Rico Queensland, Australia  (year-1)  (SCL; cm)  (kg)  W∞  Sample size  0.071  100  88.9  9  Size range (cm) 36-43  0.036  100  88.9  15  -  0.048  100  88.9  41  33-82  K  L∞  Comments; reference [method] Boulon (1994) as in Heppell & Crowder (1996) [MR] Van Dam and Diez (1994) as in Heppell & Crowder (1996) [MR] Reported in CCL and converted to SCL using SCL=SCL=0.935*CCL+0.449; Limpus (1992) as in Heppell & Crowder (1996)  102  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  A PRELIMINARY COMPILATION OF LIFE-HISTORY DATA FOR MEDITERRANEAN MARINE INVERTEBRATES 1 Charalampos A. Apostolidis Konstantinos I. Stergiou  Department of Zoology, School of Biology, Aristotle University of Thessaloniki, UP Box 134, 54124 Thessaloniki, Greece; E-mail: chapost@gmail.com, kstergio@bio.auth.gr  ABSTRACT Quantitative information on the life-history traits of fish is available online through FishBase (www.fishbase.org). This is not the case for marine invertebrates, although these organisms are of primary importance to marine ecosystems and are being heavily exploited. In order to fill this gap for the Mediterranean at least, we surveyed the primary and grey scientific literature and collected the following type of information on Mediterranean marine invertebrates: (i) length-weight relationships; (ii) maximum length (Lmax) and age (Tmax); (iii) length conversion relationships; (iv) von Bertalanffy growth parameters; and (v) length at maturity (Lm). Overall, we collected data for 246 stocks of 48 species belonging to 5 major groups (Decapoda, Bivalvia, Cephalopoda, Holothuroidea and Anthozoa). We established empirical relationships to predict asymptotic length (L∞) from Lmax and Lm from L∞. Finally, we analyzed growth parameters using the auximetric plot at the group (Decapoda and Bivalves) and species level (Aristaemorpha foliacea, Nephrops norvegicus and Plesionika martia).  INTRODUCTION Growth parameters and length-weight relationships are important not only for theoretical aspects, e.g., life-history trade-offs (Binohlan & Pauly, 2000; Charnov, 1993), but for practical reasons as well, e.g., conservation and management. In addition, compilations of historical growth data are of paramount importance for establishing baselines (Pauly, 1995). Compared to fish, invertebrate stocks are expected to be less vulnerable to overfishing, primarily due to their small body size (Jennings et al., 1998). Yet, their high economic value, and thus the high fishing effort they experience, combined with the absence or low mobility of most invertebrate species, can change this (Thorpe et al., 2006). In addition, many benthic invertebrates are keystone components for the Mediterranean ecosystems (Coll et al., 2006; 2007). Growth parameters and length-weight relationships have been assembled for fishes from different aquatic ecosystems of the world and are available online through FishBase (www.fishbase.org, Froese & Pauly, 2008). Though various compilations exist for marine invertebrates (e.g., Relini et al., 1999; Ramirez Llorda, 2002), they were done in a less systematic fashion than presented here, and are not available online (as the data presented here will be through SeaLifeBase, www.sealifebase.org). In this report, we present a preliminary compilation of life-history data (i.e., maximum length and age, length-weight relationships, von Bertalanffy growth parameters and length at first maturity) for Mediterranean marine invertebrates (Decapoda, Cephalopoda, Bivalvia, Holothuroidea and Anthozoa). This complements previous collections of life-history data for Mediterranean fishes (see Stergiou & Karpouzi, 2002; Stergiou et al., 2006), and will (i) allow the study and comparison of the patterns and propensities in the life-history of main organisms embedded in the Mediterranean; and (ii) facilitate the construction of ecosystem models of the Mediterranean Sea.  1 Cite as: Apostolidis, C.A., Stergiou, K.I., 2008. A preliminary compilation of life-history data for Mediterranean marine invertebrates. In: Palomares, M.L.D., Pauly, D. (Eds.), Von Bertalanffy Growth Parameters of Non-fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 102-121.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  103  MATERIALS AND METHODS We gathered peer-reviewed and grey literature (i.e., local journals, national and international conference proceedings, technical reports and theses) reporting growth parameters for Mediterranean marine invertebrates using the Aquatic Sciences and Fisheries Abstracts (ASFA), the Web of Science, and Google Scholar. We collected the following type of information (Table 1): (i) maximum age and length, Tmax (years) and Lmax (cm), respectively; (ii) length type reported (i.e., carapace length (CL); total length (TL); shell length (SHL); shell height (SHH); mantle length (ML); vertical length (VL)); (iii) morphometric relationships between CL and TL for Decapoda and SHL and SHH for Bivalvia; (iv) the parameters a and b of the length-weight relationship (W=aLb) and length at maturity, Lm (cm); and (v) the von Bertalanffy growth parameters, L∞ (cm), K (year-1) and t0 (year), and the method used to estimate them. We also collected auxiliary information on the sampling characteristics (i.e., sampling gear, frequency, date and region and sample size). In all cases, the parameter a of the length-weight relationship was originally estimated by the authors using millimeters as length unit. We converted all estimates to cm using the formula a’ (cm) = a (mm)*10b (Binohlan & Pauly, 1998; Stergiou & Moutopoulos, 2001). We presented Tmax only when it was estimated from growth rings (presumed to be annual), which applied to 11 bivalve stocks. When multiple methods were used by the original authors for the estimation of growth parameters, we selected the results of the method with the best fit. We estimated the Lmax/L∞ and Lm/L∞ ratios and established empirical relationships to predict L∞ from Lmax and Lm from L∞. Growth parameters were plotted with a double logarithmic scale (i.e., through an auximetric plot, Pauly et al., 1996) in order to view the relationships between K and L∞ and compare growth patterns among different Mediterranean groups and species. For 11 records, L∞ was expressed in TL or SHH. For 3 out of these 11 records, we used the known morphometric relationships to convert from one length type to another. For 8 cases, no conversion equation was found; we excluded them from the analysis.  RESULTS Overall, our dataset is based on 102 publications, of which 60 (59%) were published in sources not covered by the Science Citation Index (SCI). Most of the information gathered refers to Spain (86 stocks), Italy (83 stocks) and Greece (45 stocks), followed by Algeria (14 stocks), Croatia and Tunisia (6 stocks each) and France and Portugal (3 stocks each) (Figure 1). In total, 92% of the collected information refers to the northern Mediterranean.  Country  We collected growth parameters for 246 invertebrate stocks belonging to 48 species and 29 families (Table A1), representing 5 major groups: (i) Decapoda: 28 species (57%) and 202 stocks (82%); (ii) Bivalvia: 10 species (24%) and 27 stocks (12%); (iii) Cephalopoda: 5 species (10%) and 11 stocks (4%); (iv) Holothuroidea: 3 species (6%) and 3 stocks (1%); and (v) Anthozoa: 1 species Spain (2%) and 1 stock. The best-studied species in terms of growth were Aristeus Italy antennatus (43 stocks), followed by Greece Nephrops norvegicus (30 stocks), and Aristaeomorpha foliacea (25 stocks) Algeria (Figure 2), all of which are highlycommercial species. Tunisia Sample size ranged from 10 individuals for Holothuria sanctori (Algeria) to 31,082 individuals for Donax trunculus (Southern Adriatic Sea) (Table A1). For 54% of the stocks for which there was available information on sampling frequency such information was derived from monthly (61%), seasonal (17%), yearly (2%) and bimonthly (2%)  Croatia France Portugal 0  20  40  60  80  100  Number of stocks  Figure 1. Distribution of growth information per Mediterranean country.  104  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  sampling. For the remaining cases, either the sampling was irregular (16%), or the analysis was based on a single sample (3%). Information on the sampling gear was not available for 40% of the populations. For the remaining 60%, samples were mainly collected by trawling (79%) followed by other gears (22%) (i.e., scuba diving, hand dredges, lift nets, trammel nets, trawling box, etc.).  A. antennatus N. norvegicus  Species  A. foliacea P. longirostris P. martia M. kerathurus P. nobilis  Information on the ageing method was Others unavailable for 12% of the stocks (Table 0 20 40 60 80 100 120 A1). For the remaining stocks, growth Number of stocks was studied using length-frequency analysis (91%), shell rings reading (7%) Figure 2. The best studied invertebrate species in the and tag-recapture data (2%). LengthMediterranean Sea. frequency analysis was the only method used for the Decapoda, Cephalopoda and Holothuroidea. For the estimation of VBGF parameters, the ELEFAN software (Pauly, 1987) was used in 70% of the stocks followed by non-linear fitting of age-at-length data (24%). The method used for estimating the parameters of the VBGF was not available for 37% of the 246 cases. The K parameter varied between 0.03 year-1 for Nephrops norvegicus (Catalan Sea) and 2.06 year-1 for Palaemon adspersus (Balearic Islands) (Figure 3). The mean K value for Decapoda and Bivalvia was 0.51 year-1 (s.e. = 0.02; n = 31) and 0.52 year-1 (s.e. = 0.02; n = 210) respectively, with the two means being significantly different (t-test; t = -2.957; P = 0.003). Longevity (Tmax) from growth readings reported for 11 Bivalve stocks (Table A1) ranged between 4 and 28 years for Pinna nobilis in Carboneras (Spain) and the Thermaikos Gulf (Greece) respectively. Lmax was reported for 146 invertebrate stocks (Decapoda: 126 stocks; Bivalves: 11 stocks; Cephalopoda: 6 stocks; and Holothuria: 3 stocks). The Lmax/L∞ ratio ranged between 0.39 for N. norvegicus (Catalan sea) and 1.19 for A. antennatus (Ionian Sea), with a mean value of 0.88 (s.e. = 0.01). The relationship between L∞ and Lmax was established for: (a) Decapoda: logCL∞ = 0.0471+1.0266logCLmax (r2 = 0.93; n = 126; s.e.slope = 0.025; P < 0.001), (b) Bivalvia: logSHL∞ = 0.0708+0.9611logSHLmax (r2 = 0.99; n = 11; s.e.slope = 0.038; P < 0.001), and (c) all stocks combined: logL∞ = 0.0653+0.9920logLmax (r2 = 0.96; n = 146; s.e.slope = 0.016; P < 0.001).  40  Values of Lm were also obtained for 29 Decapoda stocks (7 species), 2 Bivalvia stocks (2 species), and 3 Cephalopoda stocks (2 species). The Lm/L∞ ratio ranged between 0.30 for A. antennatus (Ibiza Channel, Spain) and 0.66 for Melicertus kerathurus (Amvrakikos Gulf, Greece), with a mean value 0f 0.48 (s.e. = 0.017). The relationship between Lm and L∞ is presented in Figure 4.  n=246  Number of stocks  35 30 25 20 15 10 5 0  0.1  0.3  0.5  0.7  0.9  1.1  1.3  1.5  1.7  1.9  2.1  -1  Growth coefficient (K; year )  Figure 3. Distribution of K values for Mediterranean invertebrates.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D. 16 14 Length at maturity (Lm; cm)  Length-weight relationships were reported only for Decapoda (82 stocks and 23 species) and Bivalvia (10 stocks and 6 species). Length-weight relationships were reported only for 36% of the stocks (and 59% of the species) for which VBGF parameters were available. For Decapoda, b ranged between 1.4 for female Sergestes arcticus (Catalan Sea) and 3.82 for female Polycheles typhlops (Catalan Sea) (mean = 2.83; s.e. = 0.043). For bivalves, b ranged between 2.78 for Chamelea gallina (Adriatic Sea) and 3.33 for Venus verrucosa (Italy) (mean = 3.02; s.e. = 0.072).  105  12 Lm = 0.487·L∞ - 0.054 R2 = 0.92; s.e.slope = 0.026 n=34; P<0.001  10 8 6 4 2 0 0  5  10  15  20  25  30  35  Asymptotic length (L∞; cm)  Figure 4. The relationship between length at maturity (Lm) and We also gathered 7 morphometric asymptotic length (L∞) for 34 Mediterranean marine invertebrate relationships for 7 stocks and 5 species stocks of Mediterranean invertebrates, useful for the conversion of SHL and TL into SHH and CL, respectively, and vice versa (Table A1). Auximetric analysis was based on 194 and 29 sets of growth parameters for Decapoda and Bivalvia, respectively, but was not performed for groups represented by few cases (i.e., Cephalopoda, Holothuroidea, and Anthozoa) (Figure 5). Table 1. Auximetric relationships for 3 Mediterranean invertebrate species. Species Aristaeomorpha foliacea Nephrops norvegicus Plesionika martia  n 25 30 8  Relationship logk=0.124-0.533logCL∞ logk=0.225-1.336logCL∞ logk=0.416-1.590logCL∞  r2 0.207 0.664 0.758  s.e.slope 0.218 0.180 0.367  P 0.022 <0.001 0.005  The same was true for Mytilus galloprovincialis (L∞ = 12.5, K = 0.048, Ligurian Sea) and P. nobilis (L∞ = 67.13, K = 0.006, Mar Menor Lagoon). The plots revealed a significant negative linear relationship for both Decapoda and Bivalvia, the former with a steeper slope (Figure 5). The two slopes were significantly different at the 0.05 level (ANCOVA, P=0.0175). Auximetric relationships were estimated for A. foliacea, N. norvegicus and Plesionika martia (Table 1). This was not done for the rest where a low number of growth parameters were available (n<4) or the relationship was statistically not significant (P>0.05).  DISCUSSION Most of the species presented in our compilation are either of high commercial value or are discards of Mediterranean fisheries (Machias et al., 2001; Sanchez et al., 2004; Gokce et al., 2007). Concerning the articles collected, 41% were derived from SCI journals and the remaining from grey literature sources. This is very close to what was reported by Stergiou & Tsikliras (2006) for Mediterranean fishes, indicating the importance of the grey literature in the study of Mediterranean marine ecosystems. In addition, biological information on Mediterranean invertebrates is not equally distributed geographically, i.e., southern Mediterranean countries are strongly underrepresented. Marine invertebrates are a very diverse group. This diversity is also reflected in the various length types reported (i.e., CL, ML, SHL), which highlights the importance of conversion equations in order for comparisons to be done. All empirical relationships presented here displayed a strong fit and were in accordance with similar relationships estimated for fish (e.g., Froese & Binohlan, 2000). The only exception was the high slope (1.027) of the L∞-Lmax relationship for Decapoda, probably reflecting the underestimation of L∞ for organisms with small body size (Froese & Binohlan, 2000). The Lmax/L∞ ratio, which had a mean value of 0.88 (s.e. = 0.01), is similar to that reported by Stergiou (2000) for Greek  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  0.4 log10CL∞ = -0.844·log10K+0.173  0.2  -1  marine fishes. The highly significant relationships between L∞ and Lmax and Lm and L∞ can be used to predict L∞ and Lm for less studied species or stocks in data-poor situations such as in the southern Mediterranean.  Growth coefficient (log10 K; year )  106  2  R = 0.409; s.e.slope =0.073 n = 194; P < 0.001  0.0 -0.2 -0.4 -0.6 -0.8 -1.0 -1.2 -1.4  -1.6 With respect to the K-L∞ -1.8 relationships, the intercept 0.0 0.5 1.0 1.5 2.0 2.5 cannot be compared across Asymptotic carapace length (log CL ; cm) groups because of the different length types used. However, slopes are 0.4 log SHL = -0.364·log K-0.194 0.2 comparable and the slope = 0.106 R = 0.304; s.e. 0.0 for Decapoda was found to n = 29; P = 0.002 -0.2 be steeper than for -0.4 Bivalvia (0.36) and for -0.6 Mediterranean fishes -0.8 (0.39, n=1029, Apostolidis -1.0 & Stergiou unpublished -1.2 data). This relationship is -1.4 -1.6 known as the growth -1.8 trade-off and the slope has 0.0 0.5 1.0 1.5 2.0 2.5 been related to other lifeAsymptotic shell length (log SHL ;cm) history parameters and has a metabolic basis Figure 5. Relationship between growth coefficient (K) and asymptotic length (L∞) for decapod crustaceans (upper panel) and and bivalve mollusks (lower panel). (Charnov, 1993; 2007). In addition, the slopes for the 3 invertebrate species presented here are higher than those of the groups in which these species belong (see Charnov, 1993). 10  ∞  -1  Growth coefficient (log10 K; year )  10 2  10  ∞  10  slope  ∞  ACKNOWLEDGEMENTS The authors wish to thank Dr Daniel Pauly for pushing us to assemble the data presented here.  REFERENCES AA.V.V., 1993. La valutazione delle risorse demersali dei mari italiani. Atti del seminario nazionale delle unitá operative italiane svoltosi presso l'Istituto di technologia della pesca e del pescatodi Mazara del Vallo. NTR-ITPP Special Publication 2, 1-246. Abella, A.J., Righini, P., 1998. Biological reference points for the management of Nephrops norvegicus stocks in the northern Tyrrhenian Sea. J. Natur. Hist. 32, 1419-1430. Abelló, P., Marfín, P., 1993. 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N denotes the sample size. S denotes sex, i.e., F: female; M: male; and C: combined. Length-based parameters, i.e., asympotic length (L∞), length at maturity (Lm) and maximum length (Lmax) are in cm. The growth coefficient (K) is in year-1 and to in year. Length-weight relationship coefficients a and b dimensionless. AM denotes the ageing method of original data set (LF: length frequency analysis; T: tag-recapture data; SR: shell rings readings) and M denotes the method used for the estimation of the von Bertalanffy growth parameters (NL: non-linear estimation; El: Elefan software; GH: Gulland-Holt plot; FW: Ford-Walford plot). Species Decapoda  Country  Aristaeomorpha foliacea  Italy  Greece  N  S  L∞  K  t0  a  b  Lm  Tmax  Lmax  AM  M  LT  C. Tyrrhenian Sea  -  F  7.32  0.620  0.19  -  -  3.05  -  -  LF  -  CL  Tyrrhenian Sea  -  F  7.32  0.483  -0.44  0.5241  2.69  -  -  -  LF  -  CL  S. Tyrrhenian Sea  -  F  7.10  0.470  -0.28  -  -  -  -  -  -  -  CL  Sardinian Channel  -  F  5.10  0.620  0.00  -  -  -  -  -  -  -  CL  Sardinian Channel  -  M  5.10  0.635  -  -  -  -  -  -  -  CL  Sardinian Sea  -  F  7.54  0.456  0.58  -  -  3.90  -  -  LF  -  CL  Sicilian Channel  -  F  6.55  0.670  0.00  -  -  -  -  7.00  LF  -  CL  Sicilian Channel  -  M  4.15  0.960  0.28  -  -  -  -  -  LF  -  CL  Sicilian Channel  -  F  6.58  0.520  -0.23  -  -  4.20  -  -  -  -  CL  Ionian Sea  -  F  6.98  0.450  -0.18  -  -  -  -  6.50  LF  -  CL  Ionian Sea  -  F  6.98  0.450  -0.18  -  -  -  -  6.90  LF  -  CL  Ionian Sea  -  M  4.97  0.420  -0.34  -  -  -  -  4.40  LF  -  CL  W. Ionian Sea  295  F  6.60  0.450  -  -  -  -  -  6.50  LF  -  CL  W. Ionian Sea  386  M  5.00  0.420  -  -  -  -  -  4.50  LF  -  CL  Tyrrhenian Sea Sardinian Sea Aegean Sea  20819 14660 1963  C C C  7.20 7.07 6.21  0.396 0.538 0.600  0.00 0.27 -0.34  -  -  -  -  7.20 7.00 6.00  LF LF LF  NL NL NL  CL CL CL  N.E. Ionian Sea  -  F  7.25  0.430  -  -  -  -  -  7.00  LF  -  CL  N.E. Ionian Sea  -  M  6.00  0.400  -  -  -  -  -  5.90  LF  -  CL  Locality  -  Reference I Leonardi & Ardizzone (1994; in Spedicato et al., 1999a) Spedicato et al. (1998; in Spedicato et al., 1999a) Spedicato et al. (1994; in Papaconstantinou & Kapiris 2003) Mura et al. (1997; in Papaconstantinou & Kapiris 2003) Mura et al. (1997; in Papaconstantinou & Kapiris 2003) Cau et al. (1994; in Spedicato et al., 1999a) Ragonese et al. (1994; in Spedicato et al., 1999a) Ragonese et al. (1994; in Papaconstantinou & Kapiris 2003) Ragonese et al. (2004) Matarrese et al. (1997; in Spedicato et al., 1999a) Tursi et al. (1998; in Fiorentino, 2000) Tursi et al. (1998; in Fiorentino, 2000) D'Onghia et al. (1998a; in Politou et al., 2004) D'Onghia et al. (1998a; in Politou et al., 2004) Cau et al. (2002) Cau et al. (2002) Cau et al. (2002) Anonymous (2001; in Politou et al., 2004) Anonymous (2001; in Politou et al., 2004)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  113  Table A1. Continued. Species  Aristaeomorpha foliacea  Country Greece  Locality Aegean Sea N.E. Ionian Sea  Algeria  Aristeus antennatus  Spain  N 1963  S C  L∞  K  t0  a  b  Lm  Tmax  Lmax  6.21  0.600  -0.34  -  -  -  -  6.00  AM LF  M NL  LT CL  -  F  7.25  0.430  -  -  -  -  -  7.00  LF  -  CL  N.E. Ionian Sea  -  M  6.00  0.400  -  -  -  -  -  5.90  LF  -  CL  E. Ionian Sea E. Ionian Sea  392 498  F M  6.66 4.70  0.370 0.450  -  -  -  -  -  6.20 5.10  LF LF  NL NL  CL CL  E. Ionian Sea  -  F  6.40  0.460  -0.19  -  -  -  -  6.20  LF  EL  CL  E. Ionian Sea  -  M  4.70  0.564  -0.13  -  -  -  -  4.00  LF  EL  CL  Algerian Coasts  -  F  6.90  0.505  -  -  -  -  -  6.70  LF  -  CL  Algerian Coasts  -  M  4.45  0.660  -  -  -  -  -  4.50  LF  -  CL  Ibiza Channel  -  F  7.30  0.363  -0.41  0.7323  2.48  2.19  -  5.90  LF  EL  CL  Ibiza Channel  -  M  5.50  0.380  -0.43  0.7928  2.40  1.81  -  3.70  LF  EL  CL  Catalan Sea  -  F  7.60  0.300  -0.07  -  -  -  -  6.10  LF  EL  CL  Catalan Sea  -  M  5.40  0.250  -  -  -  -  -  3.80  LF  EL  CL  6962  C  7.60  0.382  0.20  -  -  -  -  4.40  LF  NL  CL  -  F  7.30  0.390  -0.08  -  -  -  -  -  -  -  CL  5844 1792 2765 1464 2678 1052 1910 961 2291 908 4049 1784  F M F M F M F M F M F M  7.40 4.60 7.30 4.40 7.30 4.50 7.40 4.40 7.40 4.60 7.40 4.40  0.380 0.468 0.521 0.435 0.364 0.410 0.521 0.390 0.510 0.531 0.387 0.400  -  0.7628 0.7730 0.7083 0.7365 0.7657 0.7789 0.5480 0.7610 0.7083 0.7676 0.7378 0.7836  2.42 2.32 2.47 2.42 2.44 2.36 2.46 2.39 2.47 2.38 2.47 2.35  2.93 2.23 2.67 2.15 2.85 2.15 2.78 2.18 2.49 2.21 2.69 2.13  -  6.10 3.80 6.10 3.80 6.00 3.70 6.30 3.60 6.50 3.50 6.40 3.60  LF LF LF LF LF LF LF LF LF LF LF LF  EL EL EL EL EL EL EL EL EL EL EL EL  CL CL CL CL CL CL CL CL CL CL CL CL  Algerian Coasts Murcia Balearic Balearic Balearic Balearic Balearic Balearic Balearic Balearic Balearic Balearic Balearic Balearic  Islands Islands Islands Islands Islands Islands Islands Islands Islands Islands Islands Islands  Reference I Cau et al. (2002) Anonymous (2001; in Politou et al., 2004) Anonymous (2001; in Politou et al., 2004) Politou et al. (2004) Politou et al. (2004) Papaconstantinou & Kapiris (2003) Papaconstantinou & Kapiris (2003) Yahiaoui et al. (1994; in Politou et al., 2004) Yahiaoui et al. (1994; in Politou et al., 2004) García-Rodriguez & Esteban (1999) García-Rodriguez & Esteban (1999) Demestre (1990; in Company & Sardá, 2000) Demestre (1990; in Company & Sardá, 2000) Cau et al. (2002) Martínez-Baños (1996; in Orsi Relini & Relini, 1998) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999) Carbonell et al. (1999)  114  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  Table A1. Continued. Species  Aristeus antennatus  Country Italy  Greece  Locality  N  S  L∞  K  t0  a  b  Lm  Tmax  Lmax  AM  M  LT  Ligurian Sea  -  F  6.30  0.142  5.26  -  -  -  -  -  LF  EL  CL  Ligurian Sea  -  F  7.70  0.213  -0.02  -  -  -  -  7.10  LF  -  CL  Ligurian Sea  -  M  4.60  0.213  -0.02  -  -  -  -  -  LF  -  CL  Tyrrhenian Sea  -  F  6.68  0.558  -0.23  0.7424  2.48  3.50  -  6.31  LF  -  CL  Tyrrhenian Sea  -  F  8.67  0.258  -  -  -  -  -  6.60  LF  -  CL  Tyrrhenian Sea  -  -  6.94  0.337  -  -  -  -  -  -  LF  -  CL  Sardinian Sea  -  F  7.68  0.340  0.37  -  -  -  -  -  LF  -  CL  Sicilian Channel  798  F  6.91  0.532  0.00  -  -  -  -  6.60  LF  EL  CL  Ionian Sea  -  F  7.72  0.350  -0.36  -  -  3.80  -  -  LF  -  CL  Ionian Sea  -  M  5.46  0.990  -0.14  -  -  2.50  -  6.50  LF  -  CL  Tyrrhenian Sea  -  F  6.77  0.490  0.00  -  -  -  -  -  LF  -  CL  Ionian Sea  -  F  6.60  0.930  -  -  -  -  -  -  -  -  CL  Ionian Sea  -  M  5.50  0.990  -  -  -  -  -  -  -  -  CL  Ionian Sea  -  F  7.72  0.350  -0.36  -  -  -  -  6.60  LF  -  CL  Ionian Sea  -  M  5.15  0.400  -0.35  -  -  -  -  3.90  LF  -  CL  Tyrrhenian Sea Sardinian Sea  8834 9452  C C  7.56 7.94  0.197 0.214  -0.29 -0.08  -  -  -  -  6.40 6.30  LF LF  NL NL  CL CL  Sicilian Channel  -  C  6.91  0.532  -  -  -  -  -  6.60  LF  -  CL  E. Ionian Sea  7273  F  6.60  0.390  0.38  1.2835  2.05  -  -  6.20  LF  EL  CL  E. Ionian Sea  1345  M  5.80  0.430  -0.46  1.2216  2.06  -  -  4.50  LF  EL  CL  Reference I Orsi Relini & Relini (1985) Orsi Relini & Relini (1998a) Orsi Relini & Relini (1998b) Spedicato et al. (1995; in Spedicato et al., 1999b) Arculeo et al. (1994; in Spedicato et al., 1999b) Arculeo et al. (1994; in Spedicato et al., 1999b) Cau et al. (1994; in Spedicato et al., 1999b) Ragonese & Bianchini (1996) Matarrese et al. (1997; in Spedicato et al., 1999b) D'Onghia et al. (1994; in Spedicato et al., 1999b) Colloca et al. (1998; in Spedicato et al., 1999b) Matarrese et al. (1992; in Papaconstantinou & Kapiris, 2001) Matarrese et al. (1992; in Papaconstantinou & Kapiris, 2001) Tursi et al. (1998; in Fiorentino, 2000) Tursi et al. (1998; in Fiorentino, 2000) Cau et al. (2002) Cau et al. (2002) Levi et al. (1998; in Cau et al., 2002) Papaconstantinou & Kapiris (2001) Papaconstantinou & Kapiris (2001)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  115  Table A1. Continued. Species  Aristeus antennatus  Country  Locality  N  S  L∞  K  Algeria  t0  b  Lm  Tmax  Lmax  AM  M  LT  Reference I Yahiaoui et al. (1986; in Fiorentino, 2000) Nouar (2001) Nouar (2001) Campillo (1994; in Orsi Relini & Relini, 1998) Dos Santos & Cascalho (1994; in Orsi Relini & Relini, 1998) Vafidis et al. (2004) Vafidis et al. (2004) Vafidis et al. (2004) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Rossetti et al. (2006) Rossetti et al. (2006) Conides et al. (1990; in Stergiou et al., 1997) Conides et al. (1990; in Stergiou et al., 1997)  -  -  F  6.51  0.370  -  -  -  -  -  -  -  CL  Algerian Coasts Algerian Coasts  -  F M  6.75 3.75  0.350 0.425  -  -  -  -  -  -  LF LF  -  CL CL  France  Lion Gulf  -  F  6.36  0.525  -0.26  -  -  -  -  -  -  -  CL  Portugal  Algarve  -  F  7.54  0.360  -0.30  -  -  -  -  -  -  -  CL  Chlorotocus crassicornis  Greece  Geryon longipes  Spain  Medorippe lanata  Italy  201 164 365 203 35 238 725 639  F M C F M C F M  1.90 1.74 1.73 5.30 7.50 7.55 3.35 3.11  0.400 0.480 0.480 0.300 0.500 0.540 1.050 1.575  -0.30 -0.27 -0.31 -  0.4224 0.4121 0.4620 -  3.14 3.20 3.10 -  2.10 -  -  4.91 6.97 6.97 2.90 2.90  LF LF LF LF LF LF LF LF  EL EL EL EL EL EL EL EL  CL CL CL CL CL CL CL CL  Melicertus kerathurus  Greece  N. Aegean Sea N. Aegean Sea N. Aegean Sea Catalan Sea Catalan Sea Catalan Sea E. Ligurian Sea E. Ligurian Sea Amvrakikos Gulf Amvrakikos Gulf Amvrakikos Gulf Amvrakikos Gulf Amvrakikos Gulf Gabes Gulf Gabes Gulf Catalan Sea Catalan Sea Catalan Sea C. Adriatic Sea C. Adriatic Sea Catalan Sea Catalan Sea Catalan Sea  -  F  24.74  0.572  -0.30  -  -  -  -  -  LF  FW  TL  -  M  24.17  0.470  -0.37  -  -  -  -  -  LF  FW  TL  -  F  6.97  1.062  0.24  -  -  4.60  -  6.20  LF  NL  CL  Conides et al. (2006)  -  M  6.27  1.253  -  -  -  -  -  -  LF  NL  CL  Conides et al. (2006)  5505  C  5.97  1.047  -  -  -  -  -  6.20  LF  NL  CL  Conides et al. (2006)  -  F M F M C F M F M C  5.43 3.70 2.9 3.05 3.05 2.16 2.37 2.75 2.92 2.95  0.600 0.780 0.250 0.320 0.320 0.460 0.480 0.400 0.400 0.400  -0.86 -0.96 -0.76 -0.59 -  0.8494 0.9093 0.8712 1.0715 1.0233 0.6805 0.6540 0.6794  3.31 3.06 3.23 2.96 3.23 3.14 3.15 3.15  -  -  2.72 2.87 2.87 2.30 2.50 2.51 2.60 2.60  LF LF LF LF LF LF LF LF LF LF  EL EL EL NL NL EL EL EL  CL CL CL CL CL CL CL CL CL CL  Ben Meriem (2004) Ben Meriem (2004) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Gramitto & Froglia (1998) Gramitto & Froglia (1998) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000)  Tunisia  Munida intermedia  Spain Italy  Munida tenuimana  Spain  55 76 131 61 67 128  0.05  a  116  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  Table A1. Continued. Species  Nephrops norvegicus  Country Italy  Spain  Greece  Algeria  Locality E. Ligurian Sea E. Ligurian Sea Sicilian Channel Sicilian Channel Ligurian Sea Ligurian Sea Tyrrhenian Sea Tyrrhenian Sea Adriatic Sea Adriatic Sea Alboran Sea Alboran Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea Euboikos Gulf Euboikos Gulf W.C. Aegean Sea W.C. Aegean Sea E.C. Aegean Sea E.C. Aegean Sea  N 37 32 46 61 30 88 49 39 38 40 79 79  S F M F M F M F M F M F M F M F M F M  L∞  K  t0  a  b  Lm  Tmax  Lmax  5.77 7.21 5.30 6.20 7.74 8.90 8.78 9.98 8.18 12.08 9.39 9.13 17.11 9.42 7.00 8.20 9.03 9.32  0.214 0.169 0.140 0.130 0.110 0.110 0.080 0.090 0.100 0.060 0.090 0.120 0.030 0.090 0.100 0.100 0.090 0.100  0.00 0.00 -0.50 -0.50 -1.32 -1.08 -1.26 -1.39 -1.36 -1.92 -1.61 -1.08 -1.80 -0.81 -2.07 -0.69 -1.27 -1.10  0.5935 0.3072 -  3.13 2.86 -  3.20 3.10 -  -  5.50 6.30 6.00 7.50 5.40 6.50 4.80 6.00 6.60 7.90 5.20 6.30  AM LF LF LF LF LF LF LF LF LF LF LF LF LF LF LF LF  M NL NL NL NL NL NL NL NL NL NL NL NL EL EL NL NL  LT CL CL CL CL CL CL CL CL CL CL CL CL CL CL CL CL CL CL  -  F  6.90  0.090  -  -  -  -  -  -  LF  EL  CL  -  M  8.60  0.060  -  -  -  -  -  -  LF  EL  CL  -  F  6.70  0.100  -  -  -  -  -  -  LF  EL  CL  -  M  8.70  0.060  -  -  -  -  -  -  LF  EL  CL  Thracian Sea  -  F  6.60  0.140  -  -  -  -  -  5.50  LF  EL  CL  Thracian Sea  -  M  7.30  0.120  -  -  -  -  -  6.60  LF  EL  CL  Toroneos & Siggitikos Gulfs  -  F  6.60  0.130  -  -  -  -  -  5.30  LF  EL  CL  Toroneos & Siggitikos Gulfs  -  M  8.30  0.110  -  -  -  -  -  7.20  LF  EL  CL  Beni-saf Beni-saf Beni-saf Beni-saf  -  F M F M  6.20 7.98 7.10 8.70  0.170 0.140 0.130 0.120  -  -  -  -  -  -  LF LF LF LF  -  CL CL CL CL  Reference I Abella & Righini (1998) Abella & Righini (1998) Ragonese et al. (2004) Ragonese et al. (2004) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1998) Sardá & Lleonart (1993) Sardá & Lleonart (1993) Mytilineou et al. (1998) Mytilineou et al. (1998) Mytilineou et al. (1993; in Stergiou et al., 1997) Mytilineou et al. (1993; in Stergiou et al., 1997) Mytilineou et al. (1993; in Stergiou et al., 1997) Mytilineou et al. (1993; in Stergiou et al., 1997) Papaconstantinou et al. (1994; in Stergiou et al., 1997) Papaconstantinou et al. (1994; in Stergiou et al., 1997) Papaconstantinou et al. (1994; in Stergiou et al., 1997) Papaconstantinou et al. (1994; in Stergiou et al., 1997) Djabali et al. (1990) Djabali et al. (1990) Djabali et al. (1990) Djabali et al. (1990)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  117  Table A1. Continued. Species  Palaemon adspersus  Country Greece  Spain  Palinurus elephas  Parapenaeus longirostris  Italy  Italy  Greece  Portugal  Algeria  Pasiphaea multidentata  Spain  N  S  L∞  K  t0  a  b  Lm  Tmax  Lmax  AM  M  LT  Messolongi Lagoon  -  F  7.90  0.165  -  -  -  -  -  6.30  LF  FW  TL  Messolongi Lagoon  -  M  6.50  0.165  -  -  -  -  -  6.00  LF  FW  TL  Balearic Islands  2506  F  4.78  2.065  -  0.0186  2.96  -  -  -  LF  GH  TL  Balearic Islands  888  M  3.41  1.076  -  0.0160  3.01  -  -  -  LF  GH  TL  Corsica  -  F  16.60  0.151  -0.35  -  -  -  -  -  T  -  CL  Corsica  -  M  13.59  0.185  -0.34  -  -  -  -  -  T  -  CL  -  F  4.44  0.740  -0.13  -  -  -  -  -  LF  -  CL  Locality  C.Tyrrhenian Sea C.Tyrrhenian Sea Sicilian Channel Sicilian Channel Sicilian Channel  -  M  3.31  0.930  -0.05  -  -  -  -  -  LF  -  CL  -  C F M  3.05 4.09 3.43  0.630 0.710 0.730  -0.19 -  1.1340 -  2.27 -  2.40 1.90  -  -  LF -  EL -  CL CL CL  Tyrrhenian Sea  -  C  4.59  0.670  -0.25  -  -  -  -  -  LF  -  CL  Tyrrhenian Sea  -  C  5.17  0.640  -  -  -  -  -  -  LF  -  CL  Tyrrhenian Sea  -  C  4.61  0.720  -  -  -  -  -  -  LF  -  CL  Ionian Sea  -  F  4.77  0.740  -0.19  -  -  -  -  -  LF  -  CL  Ionian Sea  -  M  3.55  0.540  -0.19  -  -  -  -  -  LF  -  CL  Greek Seas  -  F  3.72  0.520  -0.30  -  -  -  -  -  LF  -  CL  Greek Seas  -  M  3.37  0.620  -0.16  -  -  -  -  -  LF  -  CL  Algarve  -  F  4.40  0.700  -0.30  1.1230  2.31  2.40  -  -  LF  -  CL  Algarve  -  M  3.60  0.900  -0.30  1.1616  2.19  2.00  -  -  LF  -  CL  Algerian Coasts Algerian Coasts Catalan Sea Catalan Sea Catalan Sea  161 276 650  F M F M C  4.44 3.55 4.85 4.44 5.00  0.545 0.570 0.850 0.770 0.800  -  0.3157 0.3096 0.2511  2.61 2.65 2.84  -  -  4.67 4.27 4.67  LF LF LF LF LF  EL EL EL  CL CL CL CL CL  Reference I Klaoudatos & Tsevis (1987; in Stergiou et al., 1997) Klaoudatos & Tsevis (1987; in Stergiou et al., 1997) Manent & AbellaGutiérrez (2006) Manent & AbellaGutiérrez (2006) Marin (1985; in Secci & Cau, 1999) Marin (1985; in Secci & Cau, 1999) Ardizzone et al. (1990; in Tursi et al., 1999) Ardizzone et al. (1990; in Tursi et al., 1999) Levi et al. (1995) Ragonese et al. (2004) Ragonese et al. (2004) Carbonara et al. (1998; in Tursi et al., 1999) Carbonara et al. (1998; in Tursi et al., 1999) Carbonara et al. (1998; in Tursi et al., 1999) D'Onghia et al. (1998b; in Tursi et al., 1999) D'Onghia et al. (1998b; in Tursi et al., 1999) Anonymous (1999; in Sombrino et al., 2005) Anonymous (1999; in Sombrino et al., 2005) Ribeiro-Cascalho (1988; in Sombrino et al., 2005) Ribeiro-Cascalho (1988; in Sombrino et al., 2005) Nouar (2001) Nouar (2001) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000)  118  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  Table A1. Continued. Pasiphaea sivado  Species  Country Spain  Plesionika acanthonotus  Spain  Plesionika antigai  Greece  Plesionika edwardsii  Spain  Plesionika gigliolii  Spain  Plesionika heterocarpus  Greece Spain  Plesionika martia  Spain Greece  Polycheles typhlops  Spain  Processa canaliculata  Spain  Locality Catalan Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea N. Aegean Sea N. Aegean Sea N. Aegean Sea Catalan Sea Catalan Sea Catalan Sea W. Mediterranean Sea W. Mediterranean Sea Catalan Sea Catalan Sea Catalan Sea N. Aegean Sea N. Aegean Sea N. Aegean Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea N. Aegean Sea N. Aegean Sea N. Aegean Sea W. Ionian Sea W. Ionian Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea Catalan Sea  N 144 4156 276 64 121 192 560 384 944 209 239 453  S F M C F M C F M C F M C  L∞  K  t0  a  b  Lm  Tmax  Lmax  2.60 2.75 2.95 1.90 1.84 1.90 1.39 1.27 1.27 3.10 3.20 3.10  0.550 0.620 0.550 0.550 0.500 0.550 0.980 0.680 0.730 0.650 0.800 0.700  -0.79 -0.27 -0.11 -  0.1988 0.2106 0.2307 0.9239 0.8203 0.7757 0.6902 0.7727 0.6991  2.92 2.76 2.68 2.55 2.97 3.13 3.09 2.92 3.07  -  -  2.32 2.43 2.43 1.79 1.62 1.79 2.90 2.72 2.90  AM LF LF LF LF LF LF LF LF LF LF LF LF  M EL EL EL EL EL EL EL EL EL EL EL EL  LT CL CL CL CL CL CL CL CL CL CL CL CL  Reference I Company & Sardá (2000) Company et al. (2001) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Vafidis et al. (2004) Vafidis et al. (2004) Vafidis et al. (2004) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000)  -  F  3.10  0.800  0.15  0.8387  2.81  -  -  2.91  LF  EL  CL  García-Rodriguez et al. (2000)  -  M  2.60  0.800  -0.05  0.7900  2.94  -  -  2.88  LF  EL  CL  García-Rodriguez et al. (2000)  140 144 285 9468 11465 20933 129 50 188  F M C F M C F M C  2.05 2.00 2.10 1.78 1.61 1.56 2.30 2.24 2.27  0.750 0.550 0.750 1.450 1.170 1.090 0.900 1.000 0.900  -0.17 -0.28 -0.31 -  0.9781 1.1562 0.9160 0.7662 0.7603 0.7277  2.60 2.92 2.84 2.99 3.09 3.10  -  -  1.86 1.60 1.86 2.02 1.94 2.02  LF LF LF LF LF LF LF LF LF  EL EL EL EL EL EL EL EL EL  CL CL CL CL CL CL CL CL CL  Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Vafidis et al. (2004) Vafidis et al. (2004) Vafidis et al. (2004) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000)  208 149 370 1643 1491 3134 8231 6943 76 134 210 53 90 154  F M C F M C F M F M C F M C  3.04 2.75 3.01 2.37 2.30 2.40 3.05 2.80 4.80 3.20 4.95 2.30 2.10 2.30  0.390 0.540 0.500 0.710 0.580 0.730 0.440 0.500 0.350 0.500 0.450 1.100 0.700 1.100  -0.74 -0.19 -0.79 -  0.6239 0.6059 0.5753 0.7086 0.7230 0.1321 0.2435 0.2422 0.4624 0.4199 0.4266  3.04 3.08 3.20 2.85 2.84 3.82 3.01 3.03 3.14 3.45 3.37  1.55 -  -  2.67 2.39 2.67 2.60 2.50 4.67 3.00 4.67 2.00 1.99 2.00  LF LF LF LF LF LF LF LF LF LF LF LF LF LF  EL EL EL EL EL EL EL EL EL EL EL EL EL EL  CL CL CL CL CL CL CL CL CL CL CL CL CL CL  Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Vafidis et al. (2004) Vafidis et al. (2004) Vafidis et al. (2004) Maiorano et al. (2002) Maiorano et al. (2002) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000)  -  -  -  -  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  119  Table A1. Continued. Processa nouveli  Species  Country Spain  Scyllarides latus  Italy  Sergestes arcticus  Spain  Sergia robusta  Spain  Solenocera membranacea  Spain  Squilla mantis  Italy  N 24 38 77  S F M C  L∞  K  t0  a  b  Lm  Tmax  Lmax  1.35 1.35 1.35  1.110 1.100 1.100  -  0.4885 0.5509 0.5290  2.60 3.24 3.01  -  -  1.20 1.12 1.20  59  C  12.72  0.200  -  0.3905  3.01  -  -  35 160 77 231  F C F C  1.65 1.70 2.48 2.43  0.700 0.800 0.640 0.640  -  0.2588 0.2848 0.3932 0.3976  1.40 2.31 3.03 2.92  -  -  Catalan Sea  1367  F  3.15  0.600  -  -  -  -  -  Catalan Sea  322  M  2.40  0.500  -  -  -  -  Catalan Sea Catalan Sea Catalan Sea  246 661 907  F M C  2.85 2.16 2.90  0.650 0.560 0.650  -  0.5301 0.5558 0.5560  3.38 2.89 2.91  E. Ligurian Sea  -  F  22.00  1.450  -  -  E. Ligurian Sea  -  M  22.50  1.300  -  C. Adriatic Sea  -  F  4.19  0.450  Locality Catalan Sea Catalan Sea Catalan Sea Sicily & Linosa Islands Catalan Sea Catalan Sea Catalan Sea Catalan Sea  C. Adriatic Sea Spain  Ebro Delta Ebro Delta  AM LF LF LF  M EL EL EL  LT CL CL CL  Reference I Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000)  -  T  EL  CL  Bianchini et al. (1997)  1.37 1.37 2.25 2.25  LF LF LF LF  EL EL EL EL  CL CL CL CL  3.00  LF  EL  CL  -  2.10  LF  EL  CL  -  -  2.68 2.03 2.68  LF LF LF  EL EL EL  CL CL CL  -  -  -  -  -  -  TL  -  -  -  -  -  -  -  TL  -  1.5351  3.04  -  -  -  -  -  CL  Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Demestre & Abelló (1993) Demestre & Abelló (1993) Company & Sardá (2000) Company & Sardá (2000) Company & Sardá (2000) Righini & Baino (1996; in Piccinetti-Marfin, 1999) Righini & Baino (1996; in Piccinetti-Marfin, 1999) Froglia (1996; in Maynou et al., 2005) Froglia (1996; in Maynou et al., 2005) Abelló & Martín (1993) Abelló & Martín (1993)  -  M  4.12  0.530  -  1.5351  3.04  -  -  -  -  -  CL  1768 1732  F M  20.00 20.00  1.300 1.600  -  -  -  -  -  18.00 19.00  LF LF  EL EL  TL TL  -  C  3.50  0.160  -0.02  -  -  -  13  -  SR  NL  SHH  Peharda et al. (2002)  -  C  3.15  0.170  -0.04  -  -  -  16  -  SR  NL  SHH  Peharda et al. (2002)  -  C  3.01  0.150  -0.02  -  -  -  13  -  SR  NL  SHH  Peharda et al. (2002)  C  6.27  0.240  -0.32  0.1047  3.08  SR  -  SHL  Bivalvia  Arca noae  Callista chione  Croatia  Greece  Italy  Marina, E. Adriatic Sea Mali Ston Gulf, E. Adriatic Sea Malo Jezero, E. Adriatic Sea Thassos Island  -  -  -  -  Thassos Island  -  C  5.78  0.260  -0.15  0.1047  3.08  -  16  -  SR  -  SHL  -  -  C  9.04  0.208  0.14  0.1350  3.25  -  -  -  SR  -  SHL  Leontarakis & Richardson (2004) Leontarakis & Richardson (2004) AA.VV. (1993; in Marano et al., 1999a)  120  Compilation of life-history data for Mediterranean marine invertebrates, C.A. Apostolodis & K.I. Stergiou  Table A1. Continued. Locality  N  S  L∞  K  t0  a  b  Lm  Tmax  Lmax  AM  M  LT  Adriatic Sea  -  C  4.16  0.480  -0.01  -  -  -  -  -  SR  -  SHL  Adriatic Sea  -  C  4.28  0.790  -0.03  0.3260  2.78  -  -  -  -  -  SHL  Tyrrhenian Sea  -  C  3.91  0.500  -0.30  0.4622  2.70  -  -  -  -  -  SHL  Italy  E. Ligurian Sea  -  C  3.67  0.500  -0.31  0.1733  2.70  -  -  -  -  -  SHL  Spain  S. Adriatic Sea Catalan Sea  31082 -  C C  4.76 4.18  0.300 0.710  0.00 -0.35  -  -  1.84 -  -  3.70 3.60  LF SR  EL NL  SHL SHL  France  -  -  C  3.60  0.956  0.70  -  -  -  -  -  LF  -  SHL  Ensis siliqua  Italy  E. Ligurian Sea  -  C  14.10  0.700  -0.15  0.0096  3.08  -  -  Modiolus barbatus  Croatia  -  C  5.98  0.210  -0.10  -  -  -  13  Mytilus galloprovincialis  Italy  Mali Ston Gulf C. Tyrrhenian Sea  -  C  11.17  0.680  -0.75  -  -  -  -  Species  Chamelea gallina  Donax trunculus  Paphia aurea  Country Italy  Italy  Pecten jacobaeus  Croatia  Pinna nobilis  Greece  Ancona Northern Adriatic Sea Thermaikos Gulf  -  -  -  -  SHL  6.55  SR  NL  SHL  11.15  LF  NL  SHL  Ardizzone et al. (1996)  4.80  -  -  SHL  Froglia et al. (1998; in Marano et al., 1999e)  13  14.20  SR  GH  SHL  Peharda et al. (2003)  -  28  69.00  SR  NL  SHL  C  4.47  0.440  0.37  0.1909  2.97  1.50  70  C  12.79  0.420  -0.22  -  -  -  112  C  73.77  0.063  -0.22  -  -  -  France  Port-Cros  -  C  86.30  0.053  0.22  -  -  -  10  -  SR  -  SHL  Spain  -  C C C  49.41 45.27 68.98  0.210 0.280 0.220  -0.08 -0.07 -0.11  -  -  -  13 8 4  61.00  SR SR SR  NL NL NL  SHL SHL SHL  Galinou-Mitsoudi et al. (2005) Moreteau & Vicente (1988) Richardson et al. (1999) Richardson et al. (1999) Richardson et al. (1999)  C  72.31  0.160  -  -  -  -  -  78.00  T  GH  SHL  Siletić & Peharda (2003) Breber (1985) Arneri et al. (1991; in Marano et al., 1999f) Arneri et al. (1991; in Marano et al., 1999f) Vacchi et al. (1996; in Marano et al., 1999f) Brizzi et al. (1992; in Marano et al., 1999f)  Tapes decussata  Italy  Aguamarga Rodalquilar Carboneras S.E. Adriatic Sea Venice Lagoon  -  C  5.37  0.440  -  -  -  -  -  3.98  -  -  SHL  Venus verrucosa  Italy  Mafredonia  -  C  5.34  0.280  -1.26  0.1197  3.33  -  -  -  -  -  SHL  Bari  -  C  4.28  0.260  -1.34  0.1591  3.21  -  -  -  -  -  SHL  Genova Gulf  -  C  5.78  0.157  1.04  -  -  -  -  -  -  -  SHL  Trieste Gulf  -  C  7.54  0.189  -  -  -  -  -  -  -  -  SHL  E. Ligurian Sea  -  F  10.58  0.200  -  -  -  6.00  -  -  LF  -  ML  E. Ligurian Sea  -  M  7.94  0.270  -  -  -  5.00  -  -  LF  -  ML  Croatia  47  Cephalopoda  Loligo media  Reference I Arneri et al. (1995; in Marano et al., 1999b) Vaccarella et al. (1996; in Marano et al., 1999b) Costa et al. (1987; in Marano et al., 1999b) Costa et al. (1987; in Marano et al., 1999c) Zeichen et al. (2002) Ramón et al. (1995) Bodoy (1982; in Ramón et al., 1995) Costa et al. (1987; in Marano et al., 1999d) Peharda et al. (2006)  Italy  Auteri et al. (1987; in Belcari, 1999) Auteri et al. (1987; in Belcari, 1999)  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  121  Table A1. Continued. Eledone cirrhosa  Species  Country Italy  Illex coindetii  Spain  Octopus vulgaris  Spain  Sepia officinalis  Locality Ligurian Sea Ligurian Sea Catalan Sea Catalan Sea  -  N 217 202 416 371  S F M F M  L∞  K  t0  a  b  Lm  Tmax  Lmax  19.28 15.56 29.27 25.67  0.387 0.422 0.205 0.202  -0.03 -0.07 -  -  -  -  -  15.50 24.00 18.00  AM LF LF LF LF  M EL EL FW FW  LT ML ML ML ML  -  C  30.00  0.720  -  -  -  -  -  -  -  -  ML  Reference I Orsi Relini et al. (2006) Orsi Relini et al. (2006) Sánchez (1984) Sánchez (1984) Guerra (1979; in Belcari & Sartor, 1999) Zguidi (2002; in Ezzeddine & El Abed, 2004) Ezzeddine-Najai & El Abed (2001) Ezzeddine-Najai & El Abed (2001) Ezzeddine-Najai & El Abed (2001)  Tunisia  Gabes Gulf  -  C  29.60  0.560  -0.23  -  -  14.50  -  -  LF  -  ML  Tunisia  Tunisian coasts  -  F  27.06  0.831  -0.05  -  -  -  -  26.00  LF  EL  ML  Tunisian coasts  -  M  29.51  0.723  -0.06  -  -  -  -  27.00  LF  EL  ML  Tunisian coasts  2459  C  28.58  0.739  -0.07  -  -  -  -  27.00  LF  EL  ML  15 10 26  C C C  18.72 16.10 14.34  0.690 0.250 0.660  -  -  -  -  -  16.50 16.50 13.15  LF LF LF  EL EL EL  VL VL VL  Mezali & Semroud (1998) Mezali & Semroud (1998) Mezali & Semroud (1998)  C  35.00  0.060  -  -  -  -  -  -  -  -  -  Garcia (1984; in Campisi & Murenu, 1999)  Holothuroidea  Holothuria polii Holothuria sanctori Holothuria tubulosa  Algeria Algeria Algeria  Sidi-Fredj Sidi-Fredj Sidi-Fredj  Anthozoa  Corallium rubrum  -  -  -  122  Growth estimates of spiny lobster, Garces, L.  GROWTH ESTIMATES OF THE SPINY LOBSTER, PANULIRUS LONGIPES IN CAPTIVITY 1 Len R. Garces  The WorldFish Center - Philippine Office, Khush Hall, IRRI, College, Los Baños, Laguna, Philippines; Email: l.garces@cgiar.org  ABSTRACT Growth determination studies were conducted on spiny lobsters, Panulirus longipes (A. Milne-Edwards, 1868) in captivity to determine their growth rates and to estimate von Bertalanffy growth parameters, i.e., asymptotic carapace length (L∞) and the growth constant (K) using the Gulland and Holt method. Molting frequency in smaller individuals was higher than in larger animals. The mean single molt increments of lobsters (2.0-4.9 cm) ranged from 0.2 to 0.27 cm carapace length and 6.1 to 15.0 g total weight, with mean intermolt days of 32.5 to 60.1 days. Mean intermolt days of group-reared lobsters (4.04.9 cm) were significantly higher (P<5%) than for smaller lobsters (2.0-3.9 cm), both group- and individually-reared. Spiny lobsters which were reared in captivity had an estimated asymptotic carapace length of 6.9 cm and 7.6 cm and K of 0.68 year-1 and 0.51 year-1 for group- and individually-reared lobsters, respectively.  INTRODUCTION Knowledge of growth is essential to the basic understanding of the biology of any organism and may provide useful information both for culture and resource management considerations of commercially important species, such as spiny lobsters. Although extensive studies have been conducted on the Western Australian spiny lobster (Panulirus cygnus George, 1962), little is known about its counterpart in the Philippines, Panulirus longipes (A. Milne-Edwards, 1868). This study, conducted from January to December 1987, investigated the growth of the spiny lobster P. longipes in captivity in order to obtain estimates of the von Bertalanffy growth parameters, asymptotic carapace length (L∞) and growth constant (K), as part of a larger study on their biology and ecology (Garces, 1988).  MATERIALS AND METHODS Acquisition of experimental animals Live P. longipes were bought from fishermen in Bolinao, Pangasinan, Philippines, who collected them in the coral reef areas off the coast of Balingasay (Figure 1). These lobsters inhabit reef flats or areas deeper down the seaward portion of the outer reef. Growth determination The lobsters were held in a compartmentalized wooden tank (2.4 x 1.2 x 0.9 m), i.e., within individual stocking compartments (0.4 x 0.3 x 0.9 m) and group stocking compartments (0.6 x 0.5 x 0.9 m). Stocking density was approximately 10 lobsters·m-2. Hollow blocks were provided as shelters to simulate natural crevices. The experimental setup was provided with flow-through sea water at a rate of 2.3 l·min-1 during daytime when the pumps are running with twenty hour aeration. Every afternoon, lobsters were fed ad libitum with clams (Family Veneridae) and/or gastropods (Strombus sp.). Excess food was removed every morning to prevent fouling. 1 Cite as: Garces, L., 2008. Growth estimates of the spiny lobster, Panulirus longipes in captivity. In: Palomares, M.L.D., Pauly, D. (Eds.) Von Bertalanffy Growth Parameters of Non-fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 122-126.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  123  Carapace length (CL), total weight (TW) and sex of each lobster were determined prior to growth studies. The lobsters were tagged with colored wires tied to the base of an antenna to ensure accurate monitoring of an individual's growth. The date of each molt was recorded and the respective CL and TW were measured after 3 days when the new shell had hardened. Newly-molted lobsters were then retagged. All molting incidences were treated as single molts and grouped into 1-cm size classes. First molts in captivity and succeeding molts (i.e., second and third molts) were treated separately because part of the inter-molt period prior to the first molt in captivity was spent in the wild. This was done to eliminate the probable differences in the growth increments (Chittleborough, 1975). Growth Parameter Estimates  Figure 1. Map of study area in Bolinao, Pangasina, Philippines.  A Gulland and Holt Plot (Pauly, 1984) was used to estimate the asymptotic length (L∞) and the growth constant (K) of P. longipes longipes. A plot of size increments per unit time against mean size (for the increment in question) gives a straight line whose slope is an estimate of the value of K. Statistical Analysis Growth data such as CL increments and intermolt days were tested using unbalanced 2-way nested ANOVA. This was done to determine the differences in growth performance among the size groups and between individually and group-reared lobsters per size class. Multiple mean comparisons were also done.  RESULTS Of the 54 P. longipes reared in captivity at the Bolinao Marine Laboratory, 20 were reared individually, while 34 were reared in groups. The mean carapace length increment (CLInc) increased with size with 3.0-3.9 cm CL size class exhibiting the highest increments (Table 1, Figure 2). In terms of total weight increment (TWInc), larger individuals had greater TWInc than smaller animals (Figure 3). Similarly, mean intermolt days (IntD) increased with increasing size, while the percentages of CLInc and TWInc decreased with increase in size.  Figure 2. Mean carapace length increment per molt per size class of Panulirus longipes reared in experimental tanks at the Bolinao Marine Laboratory, University of the Philippines Marine Science Institute at Bolinao, Pangasinan, Philippines.  Figure 3 shows that the total weight increments of individuals held in isolation were higher than those held in groups. However, those in groups had higher CLInc except those in size class 3.0-3.9 cm CL (Figure 2). Moreover, lobsters held in groups exhibited shorter mean IntD than individually held animals, except those in size class 4.0-4.9 cm CL (Table 1).  124  Growth estimates of spiny lobster, Garces, L.  Table 1. Differences in growth based on single molts for the succeeding molt per size class for individually and group-reared spiny lobsters Panulirus longipes from tank experiments at the Bolinao Marine Laboratory, University of the Philippines - Marine Science Institute at Bolinao, Pangasinan, Philippines. Standard deviations of size increments and intermolt days are in brackets. Treatment Indiv. Grouped Indiv. Grouped Indiv. Grouped   Class size (cm) 2.0-2.9 3.0-3.9 4.0-4.9    Sample size 5 11 4 7 7 11   Mean Length (CL, cm) 2.6 2.5 3.5 3.6 4.4 4.6   Length increments (CL, cm) 0.20 (0.24) 0.21 (0.29) 0.27 (0.23) 0.24 (0.41) 0.22 (0.20) 0.23 (0.24)  Although carapace length increments did not differ significantly between individually and group-reared lobsters and among size classes, mean intermolt days of group reared lobsters with size 4.0-4.9 cm CL were significantly higher (P < 5%) than those of smaller lobsters (2.0-3.9 cm CL). Mean intermolt days of group and individually reared lobsters of the same size class were not significantly different (Table 1).  Mean Weight (TW, g) 22.3 17.9 45.0 53.6 88.5 117.0  Weight increments (TW, g) 6.8 (6.5) 6.1 (7.2) 13.8 (5.0) 10.0 (18.3) 15.0 (11.2) 14.0 (19.5)  Number of molts 11 14 4 7 10 15   Mean intermolt days 38.4 (6.7) 32.6 (7.6) 49.0 (12.4) 45.3 (8.8) 58.8 (10.5) 60.1 (16.3)  Table 2. Growth parameters etimated via the Gulland and Holt Plot for Panulirus longipes reared in experimental tanks at the Bolinao Marine Laboratory, University of the Philippines - Marine Science Institute at Bolinao, Pangasinan, Philippines Stocking  L∞  K  r  Sample size  6.88 7.56  0.68 0.51  -0.8958 ‐0.8631   19 7   (CL, cm) Group Individual  Length range (CL, cm) 2.02-5.72 2.63‐4.72  Preliminary growth estimates for P. longipes from the Gulland and Holt Plot are presented in Table 2. Individually reared lobsters attained higher asymptotic length (L∞) values (7.6 cm) than those held in groups (6.9 cm). In contrast, those in groups had higher K values than individually reared lobsters at 0.68 year-1 and 0.51 year-1, respectively (Figures 4a and 4b). This may suggest that group reared animals grow faster than those held in isolation.  DISCUSSION Growth of P. longipes, as in other decapod crustaceans took place discontinuously in a series of steps when ecdysis occurs. Therefore, growth is determined by the increase in CL and TW as well as molting frequency. Growth rates of P. longipes were highest in smaller individuals and decreased with increasing size. This finding is comparable with results of similar studies on P. argus (Travis, 1954) and Jasus lalandii (Fielder, 1964). Similar conclusions were also derived by Berry (1971) for P. homarus and Gomez & Junio (1985) for P. ornatus, P. versicolor and P. longipes based on carapace length increments and molting frequency. Decreasing growth rates in larger animals may be more influenced by increasing intermolt periods with size rather than smaller carapace length or total weight increments. The mean single molt increment of P. longipes (2.0-4.9 cm CL) ranged from 0.20-0.27 cm CL and 6.115.0 g TW, with mean intermolt days of 32.5-60.1 (Table 1). This is comparable with results obtained by Gomez & Juinio (1985) using the same species wherein the average single molt increment was 0.17 cm for CL ranging from 5.52-6.44 cm, and an average of 80 intermolt days (or 4.55 molts·year-1).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D. & Pauly, D.  125  Figure 3. Mean total weight increment per molt per size class of Panulirus longipes reared in experimental tanks at the Bolinao Marine Laboratory, University of the Philippines - Marine Science Institute at Bolinao, Pangasinan, Philippines.  The slight decrease in carapace length increments of the lobsters at 4.0-4.9 cm CL suggests that the growth rate may be slightly depressed as they reach sexual maturity (Figure 3). As observed by Travis (1954) in his work on P. argus, the growth rate of juveniles was rapid and decreased as they approached sexual maturity. Moreover, Berry (1971) pointed out that a decline in CL increment was observed at 5.0 cm CL in P. homarus, the size at sexual maturity. Similarly, Gomez & Juinio (1985) reported that the smallest egg-bearing female P. longipes is 4.18 cm CL. Therefore, the size at first sexual maturity for P. longipes may be about 4.0 cm CL.  Figure 4. Growth estimates of Panulirus longipes (pooled both sexes) from the Gulland and Holt Plot. A: Group stocking. B: Individual stocking.  Table 3. Growth parameters and growth performance indices (Φ’= log10K +2log10L from Pauly & Munro 1984) of three species of Panulirus from five different localities. Note clear morphological differences between the generally larger males and the smaller females. Species  P. homarus P. longipes P. longipes P. longipes P. penicillatus P. penicillatus  Sex M F both both both M F M F  Location Durban, S. Africa Durban, S. Africa Aquaria/Australia Bolinao, Pangasinan Philippines Enewetok Atoll, Marshall Islands Sta. Ana, Cagayan, Philippines  (cm) 12.0 9.42 11.3 8.8 13.4 14.6 9.65 16.1  L∞  (year-1) 0.177 0.337 0.459 0.379 0.181 0.211 0.580 0.131  K  Φ’  Source; Remarks  3.406 3.476 3.768 3.469 3.511 3.653 3.732 3.530  15.3  0.172  3.604  Smale (1978) Smale (1978) Chittleborough (1976) This study; lobsters reared in groups This study; lobsters reared individually Ebert and Ford (1986) Ebert and Ford (1986) Arellano (1989) ; K is estimated from Φ’, based on two other values for male lobsters Arellano (1989) ; K is estimated from Φ’, based on two other values of female lobsters  Growth increments per molt of P. longipes were apparently not affected by crowding since food supply was in excess. Also those lobsters held in groups had higher growth rates than those held in isolation. This observation is similar with results of earlier studies on P. cygnus in Western Australia, where Chittleborough (1975) reported that individually reared juveniles grew less than when they were held in  126  Growth estimates of spiny lobster, Garces, L.  groups. In addition, laboratory and field studies (Chittleborough 1976) showed that limited food supply is the primary cause of retarded growth. Finally, results obtained from this study also indicate that the growth parameter estimates for P. longipes are comparable with those of other species (Table 3).  REFERENCES Arellano, R.V., 1989. Estimation of growth parameters in Panulirus penicillatus using a wetherall plot and comparisons with other lobsters. Fishbyte 7(2),13-15. Berry. P.F., 1971. The biology of the spiny lobster Panulirus homarus (Linnaeus) off the east coast of Southern Africa. Invest. Rep. Oceanogr. Res. Inst. (Durban) 28, 1-75. Chittleborough, R.G., 1975. Environmental factors affecting growth and survival of juvenile western rock lobsters Panulirus longipes (Milne-Edwards). Aust. J. Mar. Freshwater Res. 26, 177-196. Chittleborough, R.G., 1976. Growth of juvenile Panulirus longipes cygnus George on coastal reefs compared with those reared under optimal environmental conditions. Aust. J. Mar. Freshwater Res. 27, 279-295. Fielder, D.R., 1964. The spiny lobster, Jasus lalandii (A. Milne-Edwards) in South Australia. I. Growth of captive animals. Aust. J. Mar. Freshwater Res. 15, 77-92. Garces, L.R., 1988. Natural Diet, Feeding and Growth in Captivity of the Spiny Lobster, Panulirus longipes longipes (A. MilneEdwards) (Decapoda: Palinuridae). University of the Philippines, College of Science, Diliman, Quezon City. MS thesis. Gomez, E.D., Juinio, A.R., 1985. Biological and Ecological Studies on Spiny Lobsters, Panulirus spp. University of the Philippines, Marine Science Institute, Terminal Report (unpublished). Pauly, D., 1983. Some Simple Methods for Assessment of Tropical Fish Stocks. FAO Fish. Tech. Pap. No. 234. Pauly, D., 1984. Fish Population Dynamics in Tropical Waters: A Manual for Use with Programmable Calculators. ICLARM Stud. Rev. No. 8. Travis, D.F., 1954. The molting cycle of the spiny lobster, Panulirus argus Latreille, I. Molting and growth in laboratory-maintained individuals. Biol. Bull. 107(30), 350-433.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  127  DEVELOPMENT AND GROWTH OF EDIBLE OYSTERS (OSTREIDAE) IN PAPUA NEW GUINEA 1 J. L. Maclean 2  Formerly of the Department of Agriculture Stock and Fisheries, Fisheries Research Station, Kanudi, Papua New Guinea  M.L. Deng Palomares  The Sea Around Us Project, Fisheries Centre, University of British Columbia, 2202 Main Mall, Vancouver, B.C. V6T 1Z4, Canada; Email: m.palomares@fisheries.ubc.ca  ABSTRACT This study is based on hitherto-unpublished field and laboratory work conducted by the first author in the early 1970s, but still considered useful; the second author provided updates and a more recent context. Larval development rates to the trochophore stage of the Papua New Guinea oysters Crassostrea amasa (Iredale) and ecomorphs of C. echinata (Quoy & Gaimard) are compared in different thermohaline regimes. The conspecificity of these ecomorphs is reflected in the similar thermohaline conditions that produce optimum development rates. Embryos of the C. echinata ecomorphs appear to prefer warmer less saline waters than C. amasa, the latter preferring almost oceanic conditions. These differences are reflected in the respective habitats of adult oysters. At least eight oyster species occur around the Papua New Guinea coastline. Three rock oysters (Crassostrea spp.) were studied with respect to their farming potential. The mangrove oyster (C. echinata) appeared suitable by its size and excellent condition attained, but the period of good condition was not predictable and collectors failed to attract spat. The Pacific oyster C. gigas, may be considered for introduction as a mariculture species, as it has been successfully introduced and farmed in other countries, but the high temperature would likely hinder reproduction and settlement, and seedlings would have to be imported for each new generation.  INTRODUCTION In the 1980s, attempts to establish farms on the Papuan coast, in Milne Bay, Galley Reach and Yule Island, of native Papua New Guinea oysters, e.g., Saccostrea cucullata (Born, 1778), were unsuccessful. Water temperatures or salinities were believed to be the cause of these failed experiments. Observations in Port Moresby harbor in 1972 and 1973 showed that oysters were spawning throughout most of the year in both hyper and hyposaline conditions. Peaks in settlement suggested that larval development was more successful in certain combinations of salinity and temperature than others. Previous work indicated ranges of these parameters experienced by oysters in vivo, which is information useful in aquaculture. However, available information on these projects, in Department of Agriculture, Stock and Fisheries files, is inadequate to assess the potential of the native oysters for farming. The work described in this paper includes unpublished experiments of the first author, who examined some aspects of oyster biology relevant to farming, including seasonality of settlement and condition factor of local species in the Port Moresby area. A series of experiments carried out in 1973 to determine the rate of development and success of larval cultures of these oysters at various salinities and 1 Cite as: Maclean, J.L., Palomares, M.L.D. 2008. Development and growth of edible oysters (Ostreidae) in Papua New Guinea. In: Palomares, M.L.D., Pauly, D. (Eds.) Von Bertalanffy Growth Parameters of Non-fish Marine Organisms. Fisheries Centre Research Reports 16(10). Fisheries Centre, University of British Columbia [ISSN 1198-6727], pp. 127-137. 2 Present address: 1901A Skyland Plaza Condominium, Sen. Gil Puyat Avenue, San Antonio Village, Makati City, Philippines; Email: jaymaclean2007@gmail.com.  128  Growth of edible oysters in Papua New Guinea, Maclean, J.L., Palomares, M.L.D.  temperatures is presented and discussed. In addition, the growth of members of the Family Ostreidae is compared with the widely-used mariculture species, the Pacific oyster, Crassostrea gigas (Thunberg, 1793), and the wisdom of a possible introduction of this species for mariculture is discussed. Species The Papua New Guinea coastline provides habitats for a number of species of edible oysters belonging to the Family Ostreidae. In the study done by the first author in the 1970s, he found that sub-littoral isolated individuals of Pycnodonte hyotis (Linnaeus, 1758), Ostrea folium Linnaeus, 1758 and Ostrea trapezina Lamarck, 1819 were common. A small intertidal Ostrea sp. forms clusters but it is too small (1.5 cm diameter) for culture purposes. There are several rock oysters (Crassostrea spp.) which could be considered for farming, occurring around most of the mainland and outer islands. Two are clustering species, forming dense discrete intertidal zones in harbours and bays, the black lip, Crassostrea echinata and C. amasa, the milky oyster. The third occurs as large individuals on mangrove roots or stones and is known locally as the mangrove oyster. It has also been identified as C. echinata. In Lombrum harbour, Manus Island, very large isolated individuals of C. tuberculata occur. The specific and generic classification of Indo-Pacific oysters is controversial, and the three Crassostrea species may in fact be subspecies of Saccostrea cuccullata, i.e., S. c. echinata, S. c. camasa and S. c. tuberculata (P. Dinamani, pers. comm. to J. Maclean).  Figure 1. Species of Ostreidae occurring in Papua New Guinea. Left panel, top to bottom: Crassostrea echinata from Port Moresby harbour wharf piles; Port Moresby harbour mangroves; Bootless Bay and Fairfax harbours mangroves. Right panel, top to bottom: Pycnodonte hyotis (Port Moresby harbour); Ostrea trapezina (Port Moresby harbour); and different forms of C. amasa (Port Moresby harbour).  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  129  Habitat The three oyster varieties considered for farming studies had different habitats. C. amasa preferred stone surfaces on more exposed areas, (west and outer Port Moresby harbour) C. echinata preferred vertical structures in sheltered areas (wharf area of Port Moresby Harbour), while the mangrove oyster appeared to favour exposed areas with a fast current, on stones and mangroves (Bootless Bay reef shallows). Size Only one species, the mangrove oyster, attains a commercially acceptable size in natural conditions for ‘in shell” sale. Table 1 shows the average sizes of 120 individuals of each of the three species. C. echinata was always flat (Figure 1) and thin shelled, indicating rapid growth. The other two species were well cupped and of thicker shell suggesting slower growth rate.  Table 1. Average volume of 120 oysters in cm3. Species Black lip Milky Mangrove  Whole 9.0 9.2 43.0  Shells 6.2 o.5 34.4  Meat 1.8 1.3 6.6  MATERIALS AND METHODS Larval cultures Gonad material was removed from adult oysters by small spatula. Eggs were sieved through a series of collars of different mesh size to remove gross foreign matter, placed in Petri dishes of filtered seawater of selected salinity and washed in similar water by pipette. A small quantity of sperm was added, and where there was foreign matter in the sperm or an excess of it, the eggs were washed again after a few minutes. Dishes and containers of seawater had previously been placed in a water bath held at the desired temperature. Temperature accuracy was to ±0.1°C. Temperatures chosen were 25, 27.5, 30, 32.5 and 35°C. Salinities were dilutions of normal seawater by distilled water from 100% seawater, 80 %, 60 % to 40 % seawater. For C. amasa, 70 % seawater was also used. After fertilization the eggs were inspected every five minutes in the order that sperm was added; times of first occurrence of polar bodies, cell divisions and finally movement of the larvae were recorded. Duplicate dishes were always used. Onset of movement of the trochophore was the last stage measured. Larvae generally failed to reach the next, D shaped veliger stage. The few that did survive would not accept available food, sterile mono-specific cultures of Chlorella pyrenoidosa and Dunaliella tertiolecta. The algae 3 were maintained in media and conditions as described by Loosanoff & Davis (1963). Success of cultures was measured by fertilization rate and numbers of abnormal larvae 4 . Seasonality of settlement  Figure 2. Asbestos cement spat collector used in Port Moresby harbour consisting of 12 plates, each 48” x 3” set 0.5” apart. The plates are supported by a steel frame and the whole collector suspended by ropes below a jetty. A free standing version on shore was abandoned due to vandalism.  To determine settlement periods of the oysters, intertidal spat collectors were placed in representative areas in Fort Moresby Harbor and Bootless Bay. For C. amasa, bunks of fibro collectors (Figure 2) (Thomson, 1950) were set out near Napa Napa. However, after three successive collectors were vandalized, this site was abandoned. For C. echinata, similar fibro collectors were hung below the Navy wharf in the harbor from March 1972 to October 1973. Results are shown in Table 2.  3 Starter cultures from Commonwealth Scientific and Industrial Research Organization (CSIRO) Marine Biochemistry Section, Sydney. Isochrysis and Monochrysis cultures were also supplied but died, presumably through temperature stress. 4 While no laboratory cultures of oysters passed the D stage veliger, one culture did produce a light spatfall on fibro plates in an outdoor concrete tank 2 x 1 x 1 m. Larvae were added to the tank at the trochophore stage into filtered seawater enriched with the Chlorella and Dunaliella cultures, though not in sufficient quantity to tint the water (about 10,000 c.c.-1). Salinity was approximately 29‰ and temperature range 25 to 27.5°C. Since this experiment was preceded by a number of unsuccessful ones, settlement was not monitored daily. Spat were first observed on the 14th day from fertilisation, when they were perhaps two days old. The larval period at these temperatures was then about 12-13 days. Settlement would undoubtedly have occurred in less time in warmer water.  130  Growth of edible oysters in Papua New Guinea, Maclean, J.L., Palomares, M.L.D.  Table 2 shows there was a major spat fall within the period MarchMay 1972, and minor spatfalls from November to April, 1973. Outside these periods there was virtually no settlement. In the case of mangrove oysters, cement bricks were set out in the adult habitat in Bootless Bay from November 1972 to November 1973. No settlement was recorded during this period. Empty shells held in netting bags were set out from May to November, 1973, but these also failed to attract spat. Seasonality of spawning  Table 2. Spat settlement of black-lipped oyster, Crassostrea echinata, on fibro collector plates, Port Moresby harbour. Period submerged Mar 2-28 1972 Mar 28 - May 15 Jun 9 - Jul 6 Jul 6 - Sep 7 Sep 5 - Oct 1 Oct 1 - Nov 10 Nov 10 - Feb 2 1973  Upper surface -  Lower surface -  119.0  86.0  -  0.2  -  0.0 16.3  Feb 23 - Apr 19  5.8  Apr 19 - Jun 4 Jun 4 - Jul 12 Jul 12 - Sep 5 Sep 6 - Oct 16  -  1.5 -  2.0 27.5 32.9  0.4 3.0  0.3  Remarks medium barnacle growth heavy barnacle, and Pinctada settlement; medium sea squirts, mainly on lower plates. light algal covering heavy barnacle settlement heavy sea squirt settlement plates fairly clean, few barnacles or sea squirts fairly clean clean to very light algae  Gonad smears of the three varieties of oysters were examined microscopically each month to determine sex and gonad stage. In practice, it was possible to detect three states of gonad development only: ripe, spending and spent or immature. Ripe oysters had full gonads, in which eggs or sperm were clearly distinguishable; spending oysters were semi-flaccid, not full, but still contained recognizable eggs and spermatozoa. Spent oysters contained very small gonads in most of which were very few eggs or Table 3. Gonad analysis of three species of Crassostrea from Port Moresby, Papua New Guinea. Species  C. amasa  Year  Month  No.  1972  June July September October December January February June September October June July August September October November December February March April May June July September November December January February March April June July August September October November  84 20 30 30 30 30 60 30 30 30 41 20 47 30 49 22 30 30 22 30 30 30 60 30 30 30 30 30 18 30 30 30 30 30 30 30  (Milky oyster)  1973  C. echinata  1972  (Black lip oyster)  1973  C. echinata  1972  (Mangrove oyster) 1973  Sex ratio 0.20 0.40 0.16 0.07 0.20 0.67 0.33 0.27 0.17 0.37 0.29 0.60 0.26 0.27 0.16 0.14 0.47 0.37 0.14 0.27 0.23 0.17 0.35 0.53 0.47 0.37 0.53 0.23 0.56 0.26 0.37 0.43 0.40 0.53 0.57 0.37  Ripe 43 9 5 2 2 1 5 9 3 13 4 6 5 4 2 4 2 1 2 -  Females Spending 10 2 15 22 10 3 24 11 20 13 5 5 15 12 35 15 8 11 15 14 14 18 31 12 16 15 11 20 4 21 12 12 14 4 9 19  Spent  Ripe  13 1 5 4 12 6 16 6 5 6 15  12 5 1 6 6 5 -  8 6 4 3 8 3 8 5 7 6 2 3 1 3 1 7 5 10 4 -  Males Spending 3 5 2 6 11 20 8 5 11 1 6 6 8 8 3 14 11 3 8 7 5 21 16 14 11 16 5 10 8 11 13 12 16 17 11  Spent 5 -  8  5 1 2 -  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  spermatozoa. Females contained mainly small oocytes, while males contained spermatocytes. Interestingly, it was only when gonad material was virtually indistinguishable that no viable eggs were present. In all months of the year, viable sperm could be obtained and eggs fertilized from specimens of all three species. The results of gonad analysis are shown in Table 3. In C. amasa, significant proportions of ripe males and females were present in two months only, June and July, 1972. In all other months sampled, most individuals were spending. C. echinata males were ripe from June to August, 1972, although some females remained so until November. In the majority of sampled months, most individuals were spending. The same was true of the mangrove oyster. In this species, no ripe males and few ripe females were found.  Table 4. Condition factors, i.e., volume of meat (M) as a ratio of the volume of the whole animal (W) and of shell space (S), of edible oysters from the Port Moresby area in 1972-73. Each sample comprises 30 specimens. Species  Year  Month  Crassostrea amasa  1972  February March April May June July August September October November December January February June September October February March April June July August September October November December January February March April April bis May June July July bis September October December January February April June July August September October November  (Milky oyster)  1973  C. echinata  1972  (Black lip oyster)  In all three species, the sex ratio varied considerably from month to month, without obvious pattern. Condition Oysters were easily obtainable as intact specimens from their substrate and initially condition was determined each month in terms of volume of meat as a percentage of whole volume. Later, condition was also measured in terms of volume of meat as a percentage of shell space (whole volume minus volume of valves). Thirty oysters were used for each determination. Meats were drained and volumes found by displacement of water. Condition factors by month are shown in Table 4. Condition as a factor of whole volume varied erratically from month to month indicating wide variation in relative volume of shells between individuals.  131  1973  C. echinata  1972  (Mangrove oyster) 1973  M/W (% vol.) 12.2 8.1 11.6 8.3 8.2 8.4 14.6 16.5 10.5 15.2 20.1 11.7 16.5 11.7 14.0 6.7 19.0 13.0 14.4 23.6 6.3 14.3 19.7 11.8 18.6 5.3 12.8 23.7 25.0 15.7 26.8 24.7 20.4 13.7 14.5 19.6 25.0 21.8 27.0 15.2 23.3 13.0 14.7 20.2 9.7 16.5 15.6  M/S (% vol. 80.6 62.1 50.0 53.2 60.6 100.0 33.0 57.9 26.5 42.2 49.0 55.6 58.0 80.0 53.7 71.8 75.5 55.6 69.2 99.6 100.0 89.1 95.2 96.8 57.8 95.4 74.8 54.4 66.3 69.0  The number of determinations of condition as a factor of shell space was insufficient to show any cyclic patterns. Black lip oysters never attained “good” condition (say over 90% full) in sampled months but there appeared to be a tendency for better condition in the latter part of the year. Even where two samples were taken in the same month (April and July 1973), condition had changed significantly. Milky oysters were full on one sampling occasion only. Mangrove oysters on the other hand exhibited good condition from October 1972 to March 1973 and in July 1973. However in October and November 1973 their condition was relatively poor. Larval sampling  Near surface phytoplankton hauls were made fortnightly using a 48 micron net at a station in Port Moresby harbor, May 1972-October 1973, to estimate relative abundance of bivalve larvae. The number of larvae in the samples was always very low and no seasonal variation was detected. Sub-samples were analyzed by Drs. Dinamani and Booth, New Zealand taxonomists, who found possible ostriids in one haul  132  Growth of edible oysters in Papua New Guinea, Maclean, J.L., Palomares, M.L.D.  only, 9 August, 1973. Evidently, the sampling method was inadequate to monitor oyster spawning in this situation.  Table 5. Average monthly water temperature, salinity and rainfall in the Port Moresby area. Year  Water temperature  1972  Temperatures were taken at least three times per week at a depth of 0.3 meters near Port Moresby, between 0900 and 1000 hours. Average monthly temperatures are shown in Table 5. Note that in 1973 water temperatures began to warm in August, whereas in 1972 the minimum occurred in September. Conditions were warmer in 1973 when the minimum recorded water temperature was 26°C. In 1972 the minimum was 24.4°C.  1973  Salinity  Month May June July August September October November December January February March April May June July August September  Water temp. (°C) 27.8 27.0 25.9 25.3 24.9 26.4 27.9 28.8 30.3 29.7 30.4 30.0 28.6 28.4 26.5 26.9 28.9  Salinity (‰)  Rainfall (mm)  38.0 37.7 37.5 36.8 36.6 37.0 36.1 34.7 32.9 33.7 35.6 36.0 36.1 36.8  133 5 8 0 0 5 2 2 236 103 216 37 150 56 19 0 0  Water samples were taken from Port Moresby harbor at a depth of 0.3 meters concurrently with temperature recording. Salinity was determined by titration with silver nitrate, standardized against “standard” sea water. Average monthly salinities are included in Table 5. Minimum salinities in March 1973 coincided with the middle of the rainy season, as shown in the rainfall data in Table 5. The hypersaline conditions that exist in the dry season are noteworthy.  RESULTS The eggs of the three species of Crassostrea are pear shaped on removal from the gonads and rounded off quickly in seawater. Average diameters are shown in Table 6. In low salinities (20% seawater) they became turgid and were not subsequently viable. There was some enlargement of eggs in 40% seawater but fertilization did take place in C. echinata, though not in C. amasa.  Table 6. Early development rates of Papua New Guinea oysters (in minutes). Parameter Temperature (°C) Salinity Egg diameter Polar bodies First cleavage Second cleavage Third cleavage Trochophore  Crassostrea amasa (Milky oyster) 30 34.5 45 10-20 min 25-30 min 35-60 min 75-85 min 190 min  C. echinata  (Black lip oyster) 30 33 41 10-20 min 30 min 35-50 min 65-70 min 180 min  C. echinata (Mangrove oyster) 30 34.5 47 5-10 min 20 min 25-45 min 45-55 min 165 min  Development of embryos was rapid at most temperatures and salinities. The stages distinguished and an example of times taken to reach them are shown in Table 6. Rate of development of individual embryos was uniform. Most attained the four cell stage within five minutes of the first observed in a given experiment, while the time lag was greater in later stages. Duplicate dishes invariably gave identical results. Development rate was influenced by both salinity and temperature in all three oyster varieties. Higher temperatures gave more rapid development in all salinities up to 32.5°C in the case of the two C. echinata forms and 30°C for C. amasa. Above these temperatures development was slower. In general lower salinities also retarded development. It would be spurious to provide results for all observed stages. The stages reached were proportional in development time, i.e. an experiment exhibiting fastest formation of polar bodies, also showed most rapid cleavages and trochophore formation. Emphasis is placed on the trochophore stage to explain the results. Fertilization rate in cultures was generally close to 100% and marked reduction only occurred at lower extremes of salinity and temperature. Abnormal embryonic development was also observed in extreme  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  133  experimental conditions. Usually this was manifest by the third cleavage in grossly mis-shaped cells. Rarely did abnormal embryos reach trochophore stage. Conditions which produced more than approximately 5% abnormal embryos are shown as shaded areas in Figure 3, i.e., approximate isopleths of times of trochophore formation at different salinities and temperatures for the three oyster varieties, which shows that:     Figure 3. Curves of equal development time in minutes to the trochophore stage at different salinities and temperatures. Shaded area denotes conditions which produced significant proportions of abnormal embryos. Dots represent development time results used to construct curves. T = temperature (°C); S = salinity expressed as % seawater where S100 = 33‰. Top left panel: Crassostrea echinata (Black lip oyster). Top right panel: C. echinata (Mangrove oyster). Bottom left panel: C. amasa (Milky oyster).  •  The effects of salinity and temperature are different for each variety.  •  Different combinations of salinity and temperature result in equivalent development rates, e.g. C. amasa reached trochophore stage at much the same time in 100% seawater at 25°C as it did in 60‰ seawater at 32.5°C.  •  There appears to be a single optimum temperature-salinity regime for each variety at the focus of each series of isopleths (see Table 7).  •  The optimum temperature-salinity values for the two C. echinata ecomorphs are very similar, as is the suggested maximum rate of trochophore development at the focus of isopleths, about 150 minutes.  •  The optimum development rate conditions for C. amasa are cooler and more saline than those of the two C. echinata forms. Minimum development time of C. amasa to trochophore stage is about 175 minutes.  •  The conditions producing significant numbers of abnormal embryos are similar in the C. echinata varieties, which appear to ‘tolerate’ higher temperatures better than C. amasa.  134  Growth of edible oysters in Papua New Guinea, Maclean, J.L., Palomares, M.L.D.  DISCUSSION Most published data on this subject relate to temperate species, as summarised by Dinamani (1974). Some of his data are reproduced here as Table 8 for comparison, with the addition of equivalent data for the tropical species Crassostrea forskali Chemnitz, which lives in waters of constant high salinity (40‰) and water temperatures in the range 16-30°C in the Red Sea (Eisawy, 1974).  Table 7. Optimum temperature-salinity regimes for species of Crassostrea in Papua New Guinea. Species  C. amasa C. echinata  Temperature (°C) 31.5  Salinity (‰) 33.0  34.5  29.7  33.8  30.0  (Black lip oyster)  C. echinata  (Mangrove oyster)  Table 8. Early development rates of other species of Crassostrea. Parameter Temperature (°C) Polar bodies First cleavage Second cleavage Third cleavage Trochophore  C. forskali  C. angulata  C. gigas  C. virginica  C. commercialis  C. glomerata  27 15-35 min 45-50 min 5 hr  20-23 40-60 min 70-80 min 80-90 min 14 hr  25 50-70 min 100 min 180 min 180 min 24-30 hr  23-25 25-65 min 45 min 50-120 min 55-195 min 8-9 hr  25 90 min 120 min 6 hr  17-18 30-45 min 90 min 120 min 180 min 12-18 hr  Salinities and temperatures in Port Moresby harbour vary from lows of S = 31‰, T = 25°C, to highs of S = 39‰, T = 31.5°C. These extremes do not occur concurrently, since cooler temperatures are associated with the hypersaline dry season. At the lower in vitro extremes, C. echinata ecomorphs would reach the trochophore stage in 4 hours, while for C. amasa it would require about 5 hours. In in vivo conditions, these times would be shorter. Consequently embryonic development rates of Papua New Guinea oysters are faster than those of the species in Table 8. Open sea temperatures in the tropics rarely exceed 30°C. The markedly higher temperature ‘preferred’ or tolerated by C. echinata ecomorphs suggests a sheltered habitat with high insolation, while the low ‘optimum’ salinity for development is associated with dilution by freshwater. C. amasa embryos clearly prefer more oceanic conditions. These preferences are confirmed in the habitats of the adult oysters in the Port Moresby area. C. amasa forms bands or aggregations on rocks in those parts of the outer harbour exposed to prevailing trade winds. C. echinata (Black lip oyster) prefers vertical structures such as wharf pylons sheltered parts of the harbour while Mangrove oyster, as the name implies, is found mainly on mangrove roots. No rivers enter the harbour and dilution is by seasonal run off. Preference for a more oceanic habitat by C. amasa would suggest less tolerance to excessive dilution of seawater. This is the case within the Great Barrier Reef where heavy mortality of C. amasa has occurred during the height of the rainy season in areas where nearby growing C. echinata (Mangrove oyster) are unaffected. The optimum thermohaline conditions for development of the two C. echinata forms are very similar indicating a close phyletic relationship, as expected. Simple larval culture experiments as described here would be very useful in assessing taxonomic hypotheses based on morphological features of oysters. The combined results of settlement, gonad and condition data indicate that the oysters in the Port Moresby region do not possess a clear reproductive cycle as such, but probably spawn sporadically throughout most of the year. Spawning peaks occur, as evidenced by patchiness of settlement in the case of black lip, and period of good condition (glycogen storage) in the milky and mangrove oysters. However, there are no seasonal patterns to these peaks. Settlement of black lip oysters was negligible during the dry seasons (winter) but the condition of adults varied erratically suggesting that some spawning continued, but larvae failed to survive in the hypersaline waters there. Changes in sex ratio provide another expression of spawning activity. In general, sex ratios in populations of Crassostrea spp. change with age, older individuals being predominantly females (Dinamani, 1974a). In C. glomerata the New Zealand rock oyster, the percentage of females also increases during the breeding season (Dinamani, 1974a). It has not been possible to age Papua New Guinea oysters, but the age  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  135  distribution would have been unlikely to vary markedly from sample to sample, and the irregular changes in sex ratio may reflect sporadic spawning activities of tropical oyster species. The mangrove oyster, by its size, and attainment of a high condition factor, would appear the most promising species for farming. However, it does not settle in a discrete zone, and attempts to collect spat have proven fruitless. Further the period of good condition in summer 1972 was not repeated in 1973. Since oysters are suitable for sale only when in good condition, mangrove oysters could be unreliable from the marketing point of view. There were experiments in 1956 and 1961-1963 of the potential of several bays in Papua New Guinea for oyster culture (Appendix 1) that showed some positive data in terms of their settlement and perhaps growth, but they were not followed up. The varying and as yet unexplained results of those and the present experiments indicate that the indigenous oysters would be difficult to farm commercially. Consideration has been given to the introduction of Pacific oysters, Crassostrea gigas, to Papua New Guinea, as this species grows rapidly to large sizes (see Figure 4). They were introduced in 1971 to Fiji where the local oysters are too small for farming. Pacific oysters there have shown rapid growth and fattening at some sites. Specimens of 10.5 cm long in excellent condition could be seen 12 months after seeding near Suva. 0.8 Crassostrea spp.  0.6 0.4 0.2 0.0  -1  K (year ; log10)  Pacific oysters were also introduced to Mauritius in 1971. There, experiments over a 3 year period concluded that the high temperatures (averaging 22.6°C in winter and 29.6 in summer) in most areas caused stress conditions in the oysters resulting in stunted growth and heavy losses. Mature oysters also suffered exhaustion by producing gametes over an excessively long reproductive season as a result of the warm temperatures (Brusca & Ardill, 1974).  -0.2 -0.4 C. virginica  -0.6  C. gigas  -0.8 -1.0 0.5  1.0  1.5  2.0  L∞ (cm; log10)  Areas of lowered salinity (estuaries) were found to show Figure 4. Auximetric plot of log10K vs log10L∞ values for 41 populations of 8 best survival and growth of species of oysters (Crassostrea ariakensis, C. cortesiensis, C. gigas, C. Pacific oysters both in Fiji and iridescens, C. madrasensis, C. rhizophorae, C. tulipa, C. virginia). Note Mauritius. Water temperatures steeper slopes of growth efficiency for C. gigas (Pacific oyster) and C. in the Port Moresby area are virginica (Atlantic oyster), cultured oyster species native to temperate areas warmer than in Mauritius. It is and introduced in tropical waters, e.g., the Pacific islands. Von Bertalanffy doubtful whether successful parameter estimates are available from SeaLifeBase (www.sealifebase.org). spawning conditions could be attained for this species in Papua New Guinea. Optimum conditions for development of Pacific oysters are temperatures of 23-250C and salinities of 23-28%. A temperature of 30°C is said to be the upper lethal limit to larval development (Fujiya, 1970). An industry based on this species in Papua New Guinea would therefore probably require imported seed each generation. Even then the ability of the oysters to tolerate the apparently stressful conditions would have to be proven by trial shipments to various estuarine sites before any large scale planting is attempted. There were early concerns of ecological problems resulting from the introduction of C. gigas in Australia and New Zealand (Dinamani, 1974b; Medcof & Wolf, 1975) that do not seem to have been realized. In Australia, C. gigas was introduced to Tasmania in the 1940s; it now grown commercially in Tasmania, South Australia, and one locality in central New South Wales, where it has been introduced. Elsewhere in  136  Growth of edible oysters in Papua New Guinea, Maclean, J.L., Palomares, M.L.D.  southeast Australian waters it has been declared a “noxious fish”. Its introduction in New Zealand sometime before 1970 was said to be accidental but it is now cultured in the north island. 5 Another constraint to the development of an oyster industry in Papua New Guinea is the prevalence of seasonal red tides (Pyrodinium bahamense) in many areas and associated risk of paralytic shellfish poisoning. Deaths from eating oysters have been recorded (Maclean, 1973). Oyster mortality during the red tides is rare. However, it would be necessary to ban sales of the oysters for a specified period during and after the red tide season. The alternative is to locate farms in Pyrodinium red tide free zones. From evidence on hand, this would restrict oyster farm localities to the mainland, west of Port Moresby on the south coast, and west of Lae on the north coast.  ACKNOWLEDGEMENTS Thanks to Dr. P. Dinamani for helpful advice on taxonomy. Thanks are also due to Daniel Pauly who convinced the first author to resurrect this contribution from an old pile of unpublished work.  REFERENCES Brusca, G., Ardill, D., 1974. Growth and survival of the oysters Crassostrea gigas, C. virginica and Ostrea edulis in Mauritius. Rev. Agricole et Sucrière de l’Ille Maurice 53, 111-131. Dinamani, P., 1974a. Reproductive cycle and gonadial changes in the New Zealand rock oyster Crassostrea glomerata. N.Z. J. Mar. Freshwater Res. 8 (1), 39-65. Dinamani, P., 1974b. Pacific oyster may pose threat to rock oyster. N.Z. Catch 74 1(6), 5-9. Dinamani, P., 1974c. Embryonic and larval development in the New Zealand rock oyster, Crassostrea glomerata (Gould, 1850). Veliger 15(4), 295-299. Eisawy, A., 1974. Spawning and larval development of the Red Sea oyster, Crassostrea forskali Chemnitz. Bull. Inst. Ocean Fish. A.R.E. 4, 205-219. Fujiya, M., 1970. Oyster farming in Japan. Helgolander wiss. Meeresunters.20, 464-479 Loosanoff, V.L., Davis, H.C., 1963. Rearing of bivalve mollusks. Advances in Marine Biology 1, 1-136. Maclean, J.L., 1973. Red tide and paralytic shellfish poisoning in Papua New Guinea. Papua New Guinea Agricul. J. 24(4), 131-138. Medeof, J.C., Wolf, P.H., 1975. Spread of Pacific oyster worries N.S.W. culturists. Australian Fisheries 34 (7): 32-38. Stenzel, H., 1971. Oysters. Volume 3 in Bivalvia part N of Treatise on Invertebrate Paleontology. R.C. Moore Editor. Geological Society of America Inc. and the University of Kansas, N.Y. Thomson, J.M.. 1950. The effects of the orientation of cultch material on the setting of the Sydney rock oyster. Aust. J. Mar. Freshwater Res. 1(1), 139-149.  5  From various internet sources.  Von Bertalanffy Growth Parameters of Non-fish Marine Organisms, Palomares, M.L.D., Pauly, D.  137  APPENDIX 1: NOTES ON EARLY OYSTER FARMING EXPERIMENTS IN PAPUA NEW GUINEA These notes are based on entries in a rediscovered file, containing carbon copies of reports to the Chief, Division of Fisheries, Department of Agriculture, Stock and Fisheries, Konedobu. The (hard copy) files have been passed to SeaLifeBase. The earliest recorded experiments were by biologist William Reed, who set out oyster spat collectors at two sites (Manumanu and Galley Reach) in the Central District west of Port Moresby and in Port Moresby harbor itself during 1956.The oyster species were not identified. The collectors were mangrove branches and shells of a large cockle. At Manumanu, spat that settled in March 1956 grew up to 2 ¼” by end October, although most were 1 ¼” and “not fat”. An interesting observation was that this growth suggested that 90% of oysters on natural substrates were less than a year old. ● At Galley Reach, collectors were set out monthly from May to October 1956. Siltation was a problem. However, some spat reached 2” in diameter after 5 months growth. ● In Port Moresby Harbor, the experiments began in March and ended in October 1956. It was noted that while March seemed to be the best time for settlement of spat, the combined observations at all sites, including those in the Central District, suggested that this “best” time corresponded with the time when the collectors were first deployed, i.e., before they began to be covered with silt and other organisms. The best collecting level was found to be 3 feet above low tide datum. Some spat grew to ¾” in diameter from March through June. In August 1960, a new marine biologist, M. Stuart-Fox carried out an examination of experimental oyster racks in a creek at the head of Milne Bay, at the eastern end of the PNG mainland. The origin of these experiments is not recorded but there was little to show for them due to mortality of oysters from gastropod (Morula sp.) infestation and burial under silt. However, in the area were “mangrove oysters”, which were “not very large” at 2-3” in diameter, and “rock oysters” at the mouth of the creek that were 4” in diameter. Stuart-Fox concluded that Milne Bay “provides ideal conditions for the natural growth of oysters attached to both rocks and mangroves and if a market could be assured, a profitable commercial oyster fishery could be established”. He then carried out a one-year project, October 1960-October 1961, to investigate the best conditions in Milne Bay for oyster settlement and growth. ●  The 36 sets of collectors were set out in 10 selected areas, mainly of suspended inverted cockle shells; a few collectors of coconut husks and asbestos (fibro) plates were used for comparison. The oysters were not identified. Shells attracted the most spat, averaging 20 per square inch, as against 5 on the coconut husks. However, the oysters were very difficult to remove without damage from the shells, a comparatively easy task with coconut husks. Asbestos plates were found too fragile and attracted vast numbers of serpulid tube worms on the underside, up to 150 per square inch. At some depths barnacles were a problem on the shells. Other fouling organisms included algae, sponges and polyzoa that covered the oysters at times. Morula caused significant mortality at some sites and depths. Density of settlement was correlated with salinity, being higher in higher salinities. Temperature varied little, in the different locations and over time; the few recordings were between 26.5 and 29 degrees centigrade. The oyster growth rates, up to 1”diameter in three months at one site, were encouraging enough to warrant continuation of the experiment after its first 6 months, for which purpose a “Fisheries Boy Grade II” was sent to train a local “boy” to take care of the experiment. However, the 6th month report was apparently the last from Mr Stuart-Fox. The next and last entry in the file is a report from the next biologist Mr Win Filewood, who visited the project site two years later, on 22-26 July 1963. The trainee used by Stuart-Fox was still employed in the work. Filewood inspected the condition of the oysters growing on coconut husk collectors from three of the sites, adding that those “growing in other areas were said to have been silted out, or affected by worms or otherwise interfered with”. He proposed that future work focus on one area and “as Mr Stuart-Fox’s work as already shown that spat-collection can be carried out fairly readily, growth should be now the principal concern. The rest of his report provides details of a proposed continuation of the project using rather sophisticated growing trays.  

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