, Volume 152, Issue 1, pp 37–47 | Cite as

Maternal investment in reproduction and its consequences in leatherback turtles

  • Bryan P. Wallace
  • Paul R. Sotherland
  • Pilar Santidrian Tomillo
  • Richard D. Reina
  • James R. Spotila
  • Frank V. Paladino


Maternal investment in reproduction by oviparous non-avian reptiles is usually limited to pre-ovipositional allocations to the number and size of eggs and clutches, thus making these species good subjects for testing hypotheses of reproductive optimality models. Because leatherback turtles (Dermochelys coriacea) stand out among oviparous amniotes by having the highest clutch frequency and producing the largest mass of eggs per reproductive season, we quantified maternal investment of 146 female leatherbacks over four nesting seasons (2001–2004) and found high inter- and intra-female variation in several reproductive characteristics. Estimated clutch frequency [coefficient of variation (CV) = 31%] and clutch size (CV = 26%) varied more among females than did egg mass (CV = 9%) and hatchling mass (CV = 7%). Moreover, clutch size had an approximately threefold higher effect on clutch mass than did egg mass. These results generally support predictions of reproductive optimality models in which species that lay several, large clutches per reproductive season should exhibit low variation in egg size and instead maximize egg number (clutch frequency and/or size). The number of hatchlings emerging per nest was positively correlated with clutch size, but fraction of eggs in a clutch yielding hatchlings (emergence success) was not correlated with clutch size and varied highly among females. In addition, seasonal fecundity and seasonal hatchling production increased with the frequency and the size of clutches (in order of effect size). Our results demonstrate that female leatherbacks exhibit high phenotypic variation in reproductive traits, possibly in response to environmental variability and/or resulting from genotypic variability within the population. Furthermore, high seasonal and lifetime fecundity of leatherbacks probably reflect compensation for high and unpredictable mortality during early life history stages in this species.


Reproductive output Optimal egg size theory Clutch size Egg size Neonate size 



We thank the field biologists, especially V. Saba, E. Price, V. Izzo, and N. Sill, and Earthwatch volunteers, and the Park Rangers and Administration for their collective conservation effort at PNMB and specifically for assistance with data collection for this project. We are grateful to J. Moore and M. Sims for statistical assistance. Financial support was provided by Earthwatch Institute, the Betz Chair of Environmental Science, Drexel University, the Schrey Chair of Biology, Indiana University–Purdue University Fort Wayne, a Faculty Development Grant from Kalamazoo College, and the Leatherback Trust. All procedures conformed to conditions of Costa Rican Ministerio del Ambiente y Energía permits and were conducted with appropriate Institutional Animal Care and Use Committee approval.


  1. Bell BA, Spotila JR, Paladino FV, Reina RD (2003) Low reproductive success of leatherback turtles, Dermochelys coriacea, is due to high embryonic mortality. Biol Conserv 115:131–138CrossRefGoogle Scholar
  2. Bennett AF (2003) Experimental evolution and the Krogh principle: generating biological novelty for functional and genetic analyses. Physiol Biochem Zool 76:1–11PubMedCrossRefGoogle Scholar
  3. Bjorndal KA, Carr A (1989) Variation in clutch size and egg size in the green turtle nesting population at Tortuguero, Costa Rica. Herpetologica 45:181–189Google Scholar
  4. Bowden RM, Harms HK, Paitz RT, Janzen FJ (2004) Does optimal egg size vary with demographic stage because of a physiological constraint? Funct Ecol 18:522–529CrossRefGoogle Scholar
  5. Caut S, Guirlet E, Jouquet P, Girondot M (2006) Influence of nest location and yolkless eggs on the hatching success of leatherback turtle clutches in French Guiana. Can J Zool 84:908–915CrossRefGoogle Scholar
  6. Congdon JD (1989) Proximate and evolutionary constraints on energy relations of reptiles. Physiol Zool 62:356–373Google Scholar
  7. Congdon JD, Gibbons JW (1985) Egg components and reproductive characteristics of turtles: relationships to body size. Herpetologica 41:194–205Google Scholar
  8. Congdon JD, Gibbons JW (1987) Morphological constraint on egg size: a challenge to optimal egg size theory? Proc Natl Acad Sci 84:4145–4157PubMedCrossRefGoogle Scholar
  9. Congdon JD, Nagle RD, Dunham AE, Beck CW, Kinney OM, Yeomans SR (1993) The relationship of body size to survivorship of hatchling snapping turtles (Chelydra serpentina): an evaluation of the “bigger is better” hypothesis. Oecologia 121:224–235CrossRefGoogle Scholar
  10. Dunham AE, Grant BW, Overall KL (1989) Interfaces between biophysical and physiological ecology and the population ecology of terrestrial vertebrate ectotherms. Physiol Zool 62:335–355Google Scholar
  11. Finkler MS, Claussen DL (1997) Within and among clutch variation in the composition of Chelydra serpentina eggs with initial egg mass. J Herpetol 31:620–624CrossRefGoogle Scholar
  12. Frazier J, Salas S (1984) The status of marine turtles in the Egyptian Red Sea. Biol Conserv 30:41–67CrossRefGoogle Scholar
  13. Hays GC, Speakman JR (1991) Reproductive investment and optimum clutch size of loggerhead sea turtles (Caretta caretta). J Anim Ecol 60:455–462CrossRefGoogle Scholar
  14. Jordan MA, Snell HL (2002) Life history trade-offs and phenotypic plasticity in the reproducion of Galápagos lava lizards (Microlophus delanonis). Oecologia 130:44–52Google Scholar
  15. Lack D (1947) The significance of clutch size. Ibis 89:302–352Google Scholar
  16. Lack D (1948) The significance of clutch size: some inter-specific comparisons. Ibis 90:25–45Google Scholar
  17. Miller JD (1997) Reproduction in sea turtles. In: Lutz PL, Musick JA (eds) The biology of sea turtles. CRC, Boca Raton, Fla., pp 51–82Google Scholar
  18. Mrosovsky N (1983) Ecology of nest site selection of leatherback turtles, Dermochelys coriacea. Biol Conserv 26:47–56CrossRefGoogle Scholar
  19. Nussey DH, Clutton-Brock TH, Elston DA, Albon SD, Kruuk LEB (2005) Phenotypic plasticity in a maternal trait in red deer. J Anim Ecol 74:387–396CrossRefGoogle Scholar
  20. Olsson M, Shine R (1997) The limits to reproductive output: offspring size versus number in the sand lizard (Lacerta agilis). Am Nat 149:179–188CrossRefGoogle Scholar
  21. Price ER, Wallace BP, Reina RD, Spotila JR, Paladino FV, Piedra R, Velez E (2004) Size, growth, and reproductive output of adult female leatherback turtles Dermochelys coriacea. Endang Spec Res 5:1–8Google Scholar
  22. R Development Core Team (2005) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. ISBN: 3-900051-07-0. URL:
  23. Reina RD, Mayor PA, Spotila JR, Piedra R, Paladino FV (2002) Nesting ecology of the leatherback turtle, Dermochelys coriacea, at Parque Nacional Marino Las Baulas, Costa Rica: 1988–1989 to 1999–2000. Copeia 2002:653–664CrossRefGoogle Scholar
  24. Reznick D, Nunney L, Tessier A (2000) Big houses, big cars, superfleas and the costs of reproduction. Trends Ecol Evol 15:421–425PubMedCrossRefGoogle Scholar
  25. Roff DA (1992) The evolution of life histories: theory and analysis. Chapman and Hall, New YorkGoogle Scholar
  26. Roff DA (2002) Life history evolution. Sinauer, Sunderland, Md.Google Scholar
  27. Rowe J (1994) Reproductive variation and the egg size-clutch trade-off within and among populations of painted turtles (Chrysemys picta bellii). Oecologia 99:35–44CrossRefGoogle Scholar
  28. Saba VS, Santidrian Tomillo P, Reina RD, Spotila JR, Musick JA, Evans DA, Paladino FV (in press) The effect of the El Niño Southern Oscillation on the reproductive frequency of eastern Pacific leatherback turtles. J Appl EcolGoogle Scholar
  29. Shine R (1992) Relative clutch mass and body shape in lizards and snakes: is reproductive investment constrained or optimized? Evolution 46:828–833CrossRefGoogle Scholar
  30. Sinervo B (1999) Mechanistic analysis of natural selection and a refinement of Lack’s and William’s principles. Am Nat 154:S26–S42CrossRefGoogle Scholar
  31. Smith CC, Fretwell SD (1974) The optimal balance between size and number of offspring. Am Nat 108:499–506CrossRefGoogle Scholar
  32. Spitze K (1991) Chaoborus predation and life-history evolution in Daphnia pulex: temporal pattern of population diversity, fitness, and mean life history. Evolution 45:82–92CrossRefGoogle Scholar
  33. Spitze K, Burnson J, Lynch M (1991) The covariance structure of life-history characters in Daphnia pulex. Evolution 45:1081–1090CrossRefGoogle Scholar
  34. Spotila JR, Dunham AE, Leslie AJ, Steyermark AC, Plotkin PT, Paladino FV (1996) Worldwide population decline of Dermochelys coriacea: are leatherback turtles going extinct? Chel Conserv Biol 2:209–222Google Scholar
  35. Steyermark AC, Spotila JR (2001) Effects of maternal identity and incubation temperature on hatching and hatchling morphology in snapping turtles, Chelydra serpentina. Copeia 2001:129–135CrossRefGoogle Scholar
  36. Steyermark AC, Williams K, Spotila JR, Paladino FV, Rostal DC, Morreale SJ, Koberg MT, Arauz R (1996) Nesting leatherback turtles at Las Baulas National Park, Costa Rica. Chel Conserv Biol 2:173–183Google Scholar
  37. Tinkle DW, Ballinger RE (1972) Sceloporus undulatus: a study of the intraspecific comparative demography of a lizard. Ecology 53:570–584CrossRefGoogle Scholar
  38. Tiwari M, Bjorndal KA (2000) Variation in morphology and reproduction in loggerheads, Caretta caretta, nesting in the United States, Brazil, and Greece. Herpetologica 56:343–356Google Scholar
  39. Trivers RL (1972) Parental investment and sexual selection. In: Campbell BG (ed) Sexual selection and the descent of man. Aldine, Chicago, Ill., pp 136–179Google Scholar
  40. Van Buskirk J, Crowder L (1994) Life-history variation in marine turtles. Copeia 1994:66–81CrossRefGoogle Scholar
  41. Wallace BP, Sotherland PR, Spotila JR, Reina RD, Franks BR, Paladino FV (2004) Biotic and abiotic factors affect the nest environment of embryonic leatherback turtles, Dermochelys coriacea. Physiol Biochem Zool 77:423–432PubMedCrossRefGoogle Scholar
  42. Wallace BP, Sotherland PS, Bouchard SS, Santidrian Tomillo P, Reina RD, Spotila JR, Paladino FV (2006a) Egg components, egg size, and hatchling size in leatherback turtles. Comp Biochem Physiol Part A 145:524–532CrossRefGoogle Scholar
  43. Wallace BP, Kilham SS, Paladino FV, Spotila JR (2006b) Energy budget analyses reveal resource limitation for eastern Pacific leatherback turtles (Dermochelys coriacea). Mar Ecol Prog Ser 318:263–270Google Scholar
  44. Williams GC (1966) Natural selection, the costs of reproduction, and a refinement of Lack’s principles. Am Nat 100:687–690CrossRefGoogle Scholar
  45. Zar JH (1999) Biostatistical analysis, 4th edn. Prentice Hall, Upper Saddle River, N.J.Google Scholar

Copyright information

© Springer-Verlag 2007

Authors and Affiliations

  • Bryan P. Wallace
    • 1
  • Paul R. Sotherland
    • 2
  • Pilar Santidrian Tomillo
    • 3
  • Richard D. Reina
    • 4
  • James R. Spotila
    • 3
  • Frank V. Paladino
    • 5
  1. 1.Nicholas School of the Environment and Earth SciencesDuke University Center for Marine ConservationBeaufortUSA
  2. 2.Department of BiologyKalamazoo CollegeKalamazooUSA
  3. 3.Department of Bioscience and BiotechnologyDrexel UniversityPhiladelphiaUSA
  4. 4.School of Biological SciencesMonash UniversityMelbourneAustralia
  5. 5.Department of BiologyIndiana–Purdue UniversityFt. WayneUSA

Personalised recommendations