Integrative and Comparative Biology Advance Access originally published online on March 31, 2006
Integrative and Comparative Biology 2006 46(3):217-223; doi:10.1093/icb/icj029
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Legacies in life histories
* Department of Ecology and Evolution, Stony Brook University Stony Brook, NY 11794-5245, USA
National Center for Ecological Analysis and Synthesis Santa Barbara, CA 93101, USA
Bodega Marine Laboratory, University of California Davis, CA 94923, USA
Correspondence: 1E-mail: padilla{at}life.bio.sunysb.edu
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 Synopsis IntroductionLegacies in marine invertebrate...The legacies of Larry...Larry McEdward's studentsLarry McEdward's publicationsReferences |
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Complex life-histories are common in nature, have many important biological consequences, and are an important focal area for integrative biology. For organisms with complex life-histories, a legacy is something handed down from an ancestor or previous stage, and can be genetic, nutritional/provisional, experiential, as well as the result of random chance and natural variation in the environment. As we learn more about complex life-histories, it becomes clear that legacies are inexorably linked in the short- and long-term through ecology and evolution. Understanding the consequences and drivers of life-history patterns can therefore only be understood by considering all types of legacies and integrating legacies across the entire life cycle. Larry McEdward was a leader in the field of ecological physiology, and evolutionary ecology of marine invertebrate larvae with complex life-histories. Through his scientific work and publications, devotion to students, colleagues, family, and friends, Larry has left a lasting legacy that will impact the future development of the field of larval ecology and complex life-histories.
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SynopsisIntroductionLegacies in marine invertebrate...The legacies of Larry...Larry McEdward's studentsLarry McEdward's publicationsReferences |
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Complex life-histories are an important focal area of study that integrates across disciplines including ecology, evolution, development, physiology, and behavior. They are common in nature, found in almost all major taxa of both plants and animals, and have many important biological consequences. Among animals, the greatest diversity of life histories is found in marine invertebrates. Virtually all animal phyla originated in marine systems and have a larval form (Levin and Bridges 1995
; Young and others 2002). Evidence suggests that the ancestral condition for many animal phyla is a complex life-history (Jägersten 1972; Nielsen 1995). These complex life-histories result in early life stages that are physiologically, morphologically, and ecologically different from later stages. Furthermore, there are intriguing phylogenetic patterns of complex life-histories among species of marine invertebrates—including convergence of adult forms with different larval types, and convergence of larval forms with different adults. It is therefore not surprising that complex life-histories of marine invertebrates have fascinated scientists since the mid-1800s when Thompson discovered the larvae of crabs and barnacles (see Young 1990 for an excellent review of the history of larval ecology), and have proven to be vital for studies of physiology, life-history theory, ecology, and most recently evolution of development and biogeography.
Larry McEdward was a leader in the field of ecological physiology, and evolutionary ecology of marine invertebrate larvae, until 2001 when he unexpectedly passed away—a significant loss to the field. As a tribute to Dr. McEdward, we assembled leaders in the studies of complex life-histories of marine invertebrates for a symposium held at the annual meeting of the Society for Integrative and Comparative Biology in San Diego, California in January of 2005. This symposium combined forward thinking syntheses of ideas for guiding future studies of complex life-histories of marine animals. Since the last major review of this area (McEdward 1995) there have been several major advances in our understanding of complex life-histories in marine invertebrates, including the ecological forces that drive evolutionary shifts in life histories, the developmental mechanisms that allow for these shifts, the ecological and evolutionary roles of dispersal via larvae, and the linking of larval condition with later life-history stages. The papers derived from our symposium and contained in this volume synthesize recent major research advances, and focus on the future directions, especially those that result from Dr. McEdward's work. We hope that this collection facilitates new questions and lays the foundation for work on complex life-histories for the next several decades.
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Legacies in marine invertebrate complex life-histories |
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SynopsisIntroductionLegacies in marine invertebrate...The legacies of Larry...Larry McEdward's studentsLarry McEdward's publicationsReferences |
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A general concept that emerged from the symposium was the importance of different types of legacies for the evolution and ecology of complex life-histories. According to The American Heritage Dictionary (2000)
a legacy is something handed down from an ancestor, a predecessor or from the past. The origin of legacy comes from the Medieval Latin legatio and from Latin legare, which mean to bequeath. For larvae resulting from complex life-histories there are many types of legacies (Fig. 1), 3 of which seem particularly important: genetic, provisioning, and experiential legacies.
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Genetic legacies
The genetic composition of an individual is a legacy bequeathed directly from parents, and ultimately through phylogenetic contributions of ancient ancestors (Fig. 1). Understanding the causes for and consequences of genetic legacies has been a primary goal of studies of life history, such as the evolution of life-history traits within lineages (Byrne 2006
; Zeng and others 2006) and the patterns of development and mechanisms that alter gene expression and function (Wray 2006). In addition, genetic legacies can be used as a tool to reconstruct long-term evolutionary histories, which can then be used to examine broad scale patterns of dispersal and range expansion (Paulay and Meyer 2006). For example, Byrne (2006) has been able to reveal a stunning diversity of life histories among closely related Asteroids in southern Australia. She has also shown that this life-history diversity is seen in very closely related species with similar morphologies, some so similar that they form cryptic species pairs. This work clearly shows that life-history stages can be extremely labile and decoupled through evolutionary time, and thus do not necessarily change in a unidirectional pattern as had previously been assumed. Life history and growth form also show repeated evolutionary origins in ascidians (Zeng and others 2006), and it is genetic and phylogenetic legacies that allow us to unravel these historic patterns.
Thus, although there is some element of phylogenetic inertia in many aspects of the development, morphology, and behavior driven by historic genetic legacies, variation in genetic mechanisms that impact development and function in the present shows that these features can be labile, allowing similar mechanisms to operate in different ways, as well as converge of form with a divergence of function (Wray 2006). Certainly much of the work on the evolution of development (EvoDevo) has shown that genetic legacies provide the opportunity for flexibility as well as constraints that can drive alternative strategies and solutions.
Provisioning legacies
Provisions (materials including energy, nutrients, etc.) acquired during a life stage and passed on or provided to (for example, nurse eggs) a subsequent stage are also an important legacy in complex life-histories (Fig. 1). An obvious provisional legacy is the parental material and energy contributed to eggs, which can affect subsequent development, such as whether an egg is fertilized (Levitan 2006), or when larvae must feed to avoid starving (McEdward and Miner 2006). Because much of the theory developed to explain the evolution of different developmental modes in marine animals has focused on the importance of egg size (Vance 1973), and therefore maternal provisioning, the effects of provisional legacies demand further attention (see McEdward 1997; Levitan 2000).
Energy and nutritional provisioning to eggs impacts egg size, which can impact fertilization success (Levitan 2006), as well as development time and the need for larval feeding (McEdward and Miner 2006), which impact dispersal potential (Paulay and Meyer 2006; Levin 2006) and has been the focus of life-history models (Havenhand 1995; McEdward and Miner 2006). There has also been recent recognition that some larvae have facultative feeding, and that lecithotrophic and planktotrophic life histories are not distinct categories. Therefore, larval feeding mode is not necessarily dictated by genetics, but can be flexible and dependent on environmental conditions as well as the nutritional legacies of parental investment. These new findings dictate the need for a revision and re-evaluation of how we approach life-history models (McEdward and Miner 2006).
Although provisional legacies almost certainly have important consequences throughout ontogeny for organisms with complex life-histories, relatively few of these legacies have been investigated. One type of provisional legacy that is starting to receive more attention is how nutritional stores accumulated as larvae influence post-metamorphic juvenile size, growth and performance (Emlet 2006; Pechenik 2006). This work clearly demonstrates that we need to include ontogeny after the larval stage when considering complex life-histories. Having enough energy or nutritional stores just sufficient for metamorphosis is not the end of the story—the amount, and perhaps quality, of resources that larvae acquire can influence the success of later life stages (Emlet 2006; Pechenik 2006). Equally important are the potential tradeoffs that larvae face between gathering nutrition and other demands such as locomotion (Strathmann and Grünbaum 2006), which will impact the amount of provisional legacy that a larva can accumulate for later life stages. The importance of provisional legacies for later life stages is potentially not only important for species with feeding larvae, but also for those with non-feeding larvae, especially depending on the length of the larval phase, and environmental conditions. However, this question has yet to be explored for those species with non-feeding larvae.
Environmental/experiential legacies
Environmental or experiential legacies result from experiences during ontogeny that influence later stages (Fig. 1). For example, the large distance dispersal of many larvae is determined by water currents that carry them into waters with different types of characteristics, thus the larval experience can be quite variable among individuals within the same species, and sometimes among individuals within the same population (Levin 2006). Many larvae have been shown to be capable of phenotypically plastic responses to short-term variability in the environment, and these responses can have not only short term but also lasting impacts (Hadfield and Strathmann 1996). Miner and Vonesh (2004) demonstrated that larvae can not only alter their morphology in response to the mean difference in food concentration, but also to the amount of variation in food concentration. Larvae reared on a constant diet developed larger feeding structures than those fed the same mean density of food, but that experienced higher variance in food concentrations, as might be expected in the real world given the patchiness of natural phytoplankton populations. Exposure to predators during the larval phase can also impact behavior and morphology, including shell growth and thickness in gastropod larvae (D. Vaughn, unpublished data). Such short-term changes due to phenotypically plastic responses will have a lasting legacy on individuals, especially as they may impact factors important for feeding efficiency, locomotion, growth, and survivorship during the larval phase.
Experiences and local environmental fluctuations and conditions can affect all life stages and can impact such important life-history features such as phenology, age, and size of first reproduction, number of reproductive bouts, as well as persistence and longevity (Hadfield and Strathmann 1996). Environmental and experiential legacies have received the least amount of attention to date, but are undoubtedly important for the evolution of complex life-histories.
Integration among types of legacies
Each type of legacy does not operate in isolation, but rather in concert with all other legacies throughout the life history of an individual (Fig. 1). Each provides and responds to feedbacks, opportunities, and constraints of the others. Legacies link all different life-history stages, therefore focusing solely on eggs or larvae is not sufficient. In addition, different types of legacies interact and therefore an integrative approach that considers how the combined effects of different legacies influence the evolution and ecology of species with complex life-histories is needed.
Parental experience and physiological state can alter the allocation of provisional legacies to offspring, thus changing important life-history parameters such as development time and whether or not larvae need to or can feed in the plankton (Gibson and Gibson 2004). The length of the larval phase can have a number of impacts, including the potential for dispersal, range expansion, and larval transport (Levin 2006; Paulay and Meyer 2006). Feedbacks between provisional and experiential legacies can impact large scale patterns of dispersal and biogeographic patterns, both of which are currently important for conservation, patterns of biodiversity, as well as local enhancement and recovery of species in protected areas and for fisheries enhancement and restoration (Levin 2006).
Morphological legacies also have impacts on larval performance and post-larval life-history. At a basic level, morphology is genetically controlled, and the result of long evolutionary histories of body plans and larval forms (Raff 1996). Environmentally induced feedbacks can modulate larval morphologies (through phenotypic plasticity, for example, Miner and Vonesh 2004) and behaviors that then further impact performance and energy or nutritional acquisition as well as survivorship. Indeed, the contrasting tradeoffs between alternative necessities of planktonic larvae such as locomotion and feeding (Strathmann and Grünbaum 2006) can impact developmental morphology and larval nutritional acquisition, both known to drive and be driven by feedbacks between the environment and larval development, morphology, and behavior. Environmental experiences and other factors that impact survival and fitness ultimately shape long-term evolutionary patterns and legacies. Even phenotypically plastic traits and environmental responses are being recognized as important drivers of genetic legacies through genetic assimilation (Strathmann and others 1992; West-Eberhard 2003).
As we learn more about complex life-histories, it becomes clear that legacies are inexorably linked in the short term as well as the long term through ecology and evolution. Understanding the consequences and drivers of life-history patterns can therefore only be understood by considering all types of legacies and integrating legacies across the entire life cycle.
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The legacies of Larry MCEdward |
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SynopsisIntroductionLegacies in marine invertebrate...The legacies of Larry...Larry McEdward's studentsLarry McEdward's publicationsReferences |
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As scientists, our ideas and research are a lasting legacy through our published work. Much like the phylogenetic legacies of organisms, our work is built on and influenced by the writings of and interactions with those who preceded us—specially teachers and mentors who helped us develop our scientific philosophies, skills and approaches to answering questions. Similarly, we impart a legacy on our colleagues, students, and those we train. We not only influence their research and scientific products through direct interactions and input, but we also create environments that can facilitate thought and the development of new ideas and approaches.
Larry has left many different types of legacies, not only through his research and publications, thoughts and ideas, but also through the time and devotion to his students, colleagues, friends, and family. Larry also inherited many important legacies from his colleagues and mentors, John Lawrence, Richard Strathmann, and Fu-Shiang Chia. These legacies not only affected his mentoring, teaching, and research, but were also passed on to his students and fellow colleagues. Larry trained 6 Masters and 7 Ph.D. students and influenced countless numbers of graduate and undergraduate students he taught at the University of Florida and at the Friday Harbor Laboratories. Larry's devotion to his family and friends, passions, and personal work-hard play-hard ethic, sense of humor and generous nature will provide lasting legacies for those who knew him, just as the development of the field of larval ecology and complex life-histories through his published science will have a legacy on the future scientific research.
We miss you Larry.
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Larry McEdward's students |
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*Students in 2001.
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Larry McEdward's publications |
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SynopsisIntroductionLegacies in marine invertebrate...The legacies of Larry...Larry McEdward's studentsLarry McEdward's publicationsReferences |
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McEdward LR, Miner. BG 2006. Calculation, estimation, and interpretation of egg provisioning in marine invertebrate life cycles. Integr Comp Biol. doi: 10.1093/icb/icj026.
Miner BG, McEdward LA, McEdward LR. 2005. The relationship between egg size and the duration of the facultative feeding period in marine invertebrate larvae. J Exp Mar Biol Ecol 321:135–44.
Reitzel AM, Miles CM, Heyland A, Cowart JD, McEdward LR. 2005. The contribution of the facultative feeding period to echinoid larval development and size at metamorphosis: a comparative approach. J Exp Mar Biol Ecol 317:189–201.
Reitzel AM, Miner BG, McEdward LR. 2004. Relationships between spawning date and larval development time for benthic marine invertebrates: a modeling approach. Mar Ecol Prog Ser 280:13–23.
McEdward LR, BG Miner. 2003. Fecundity-time models of reproductive strategies in marine benthic invertebrates: fitness differences under fluctuating environmental conditions. Mar Ecol Prog Ser 256:111–21.
McEdward LR, Qian PY. 2001. Effects of the duration and timing of starvation during larval life on the metamorphosis and juvenile size of the polychaete Hydroides elegans. J Exp Mar Biol Ecol 261:185–97.
McEdward LR, Jaeckle WB, Komatsu M. 2002. Phylum Echinodermata: Asteroidea. In: Young CM, Rice ME, Sewell M, editors. An atlas of invertebrate larvae. San Diego: Academic Press. p 499–512.
Miner BG, Cowart JC, McEdward LR. 2002. Egg energetics for the facultative planktotroph Clypeaster rosaceus (Echinodermata: Echinoidea), revisited. Biol Bull 202:97–9.
McEdward LR, Miner BG. 2001. Larval and life cycle patterns in echinoderms. Can J Zool 79:1125–70.
McEdward LR, Miner BG. 2001. Larval ecology of echinoids. In: Lawrence JM, editor. Biology and ecology of edible sea urchins. Amsterdam: Elsevier. p 59–78.
McEdward LR, Morgan KH. 2001. Interspecific relationship between egg size and the level of parental investment per offspring in echinoderms. Biol Bull 200:33–50.
McEdward LR. 2000. Introduction; larval evolution. Sem Cell Dev Biol 11:383–4.
McEdward LR. 2000. Adaptive evolution of larvae and life cycles. Sem Cell Dev Biol 11:403–9.
McEdward LR. 2000. The origin and evolution of marine invertebrate larvae. Q Rev Biol 75:191–2. Book Review.
McEdward LR, Herrera JC. 1999. Body form and skeletal morphometrics during larval development of the sea urchin Lytechinus variegatus. J Exp Mar Biol Ecol 232:151–76.
McEdward LR. 1997. Reproductive strategies of marine benthic invertebrates revisited: facultative feeding by planktotrophic larvae. Am Nat 150:48–72.
McEdward LR, Janies DA. 1997. Relationships among development, ecology, and morphology in the evolution of echinoderm larvae and life cycles. Biol J Linn Soc 60:381–400.
McEdward LR. 1996. Experimental manipulation of parental investment in echinoid echinoderms. Am Zool 36:169–79.
Herrera JC, McWeeney S, McEdward LR. 1996. Diversity of energetic strategies among echinoid larvae and the transition from feeding to nonfeeding development. Oceanol Acta 19:313–21.
McEdward LR, Hadfield MG. 1996. Flexibility in life cycles. Oceanol Acta 19:468.
Nichols D, McEdward LR, Smith AB. 1996. Evolution of life history traits. Oceanol Acta 19:469–70.
McEdward LR. 1995. Ecology of marine invertebrate larvae. Boca Raton: CRC Press.
McEdward LR. 1995. Evolution of pelagic direct development in the starfish Pteraster tesselatus (Asteroidea: Velatida). Biol J Linn Soc 154:299–327.
Janies DA, McEdward LR. 1994. A hypothesis for the evolution of the concentricycloid water-vascular system. In: Wilson WH Jr, Stricker SA, Shinn GL, editors. Reproduction and development of marine invertebrates. Baltimore: Johns Hopkins Press. p 246–57.
Janies DA, McEdward LR. 1994. Heterotopy, pelagic direct development, and new body plans in velatid asteroids. In: David B, Guille A, Féral J-P, Roux M, editors. Echinoderms through time. Proceedings of the 8th International Echinoderms Conference, Dijon, France. Rotterdam, Netherlands: Balkema Press. p 319–24.
McEdward LR and Janies DA. 1993. Life cycle evolution in asteroids: what is a larva? Biol Bull 184:255–68.
Janies DA, McEdward LR. 1993. Highly derived coelomic and water-vascular morphogenesis in a starfish with pelagic direct development. Biol Bull 185:56–76.
McEdward LR. 1992. Morphology and development of a unique type of pelagic larva in the starfish Pteraster tesselatus. Biol Bull 182:177–87.
McEdward LR, Chia FS. 1991. Size and energy content of eggs from echinoderms with pelagic lecithotrophic development. J Exp Mar Biol Ecol 147:95–102.
Paulay G, McEdward LR. 1990. A simulation model of island reef morphology: The effects of sea level fluctuations, growth, subsidence, and erosion. Coral Reefs 9:51–62. Received best paper award from the journal Coral Reefs for 1990.
Qian PY, McEdward LR, Chia FS. 1990. Effects of delayed settlement on survival, growth, and reproduction in the spionid polychaete, Polydora ligni. Int J Invertebr Repr Dev 18:147–52.
McEdward LR, Carson SF, Chia FS. 1988. Energetic content of eggs, larvae, and juveniles of Florometra serratissima and the implications for the evolution of crinoid life histories. Int J Invertebr Repr Dev 13:9–21.
McEdward LR. 1988. Experimental embryology as a tool for studying the evolution of echinoderm life histories. In: Smith A, Paul CRC, editors. Echinoderm phylogeny and evolutionary biology. Oxford: Clarendon. p 189–96.
Sinervo B, McEdward LR. 1988. Developmental consequences of an evolutionary change in egg size: an experimental test. Evolution 42:885–900.
McEdward LR, Strathmann RR. 1987. The body plan of the cyphonautes larva of bryozoans prevents high clearance rates: comparison with the pluteus and a growth model. Biol Bull 172:30–45.
Emlet RB, McEdward LR, Strathmann RR. 1987. Echinoderm larval ecology viewed from the egg. In: Jangoux M, Lawrence JM, editors. Echinoderm studies. Volume 2. Rotterdam: A.A. Balkema. p 55–136.
McEdward LR, Carson SF. 1987. Variation in organic content and its relationship with egg size in the starfish Solaster stimpsoni. Mar Ecol Prog Ser 37:159–69.
McEdward LR, Coulter LK. 1987. Egg volume and energetic content are not correlated among sibling offspring of starfish: Implications for life history theory. Evolution 41:914–7.
McEdward LR. 1986. Comparative morphometrics of echinoderm larvae. I. Some relationships between egg size and initial larval form in echinoids. J Exp Mar Biol Ecol 96:251–65.
McEdward LR. 1986. Comparative morphometrics of echinoderm larvae. II. Larval size, shape, growth, and the scaling of feeding and metabolism in echinoplutei. J Exp Mar Biol Ecol 96:267–86.
Strathmann RR, McEdward LR. 1986. Cyphonautes' ciliary sieve breaks a rule of biological inference. Biol Bull 171:694–700.
McEdward LR. 1985. An apparatus for measuring and recording the depth dimension of microscopic organisms. Trans Am Microsc Soc 104:194–200.
McEdward LR. 1985. Effects of temperature on the body form, growth, ETS activity, and development rate of an echinopluteus. J Exp Mar Biol Ecol 93:169–81.
Day RL, McEdward LR. 1984. Aspects of the physiology and ecology of pelagic larvae of marine benthic invertebrates. In: Steidinger KA, Walker LM, editors. Marine plankton life cycle strategies. Boca Raton, FL: CRC Press. p 93–120.
McEdward LR. 1984. Morphometric and metabolic analysis of the growth and form of an echinopluteus. J Exp Mar Biol Ecol 82:259–87.
McEdward LR, Lawrence JM. 1981. Respiratory electron transport system activity during larval development of Lytechinus variegatus Lamarck (Echinodermata: Echinoidea). Comp Biochem Physiol B 70:653–5.
Diehl WJ, McEdward LR, Proffitt E, Rosenberg V, Lawrence JM. 1979. The response of Luidia clathrata (Echinodermata: Asteroidea) to hypoxia. Comp Biochem Physiol A 62:669–71.
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Acknowledgements |
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This paper was part of a symposium entitled: "Complex Life Histories of Marine Invertebrates—A Tribute to Larry McEdward," supported by the Division of Invertebrate Zoology and the Division of Ecology and Evolution of SICB, and NSF (IBN-0450894). This work was conducted while D.K.P. was a Sabbatical Fellow at the National Center for Ecological Analysis and Synthesis, a Center funded by NSF (Grant #DEB-0072909), the University of California, and the Santa Barbara campus. Contribution number 2294, Bodega Marine Laboratory, University of California, Davis.
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2Present address: Department of Biology, Western Washington University Bellingham, WA 98225-9160, USA.
From the symposium "Complex Life-Histories in Marine Benthic Invertebrates: A Symposium in Memory of Larry McEdward" presented at the annual meeting of the Society for Integrative and Comparative Biology, January 4–8 2004, at San Diego, California.
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Byrne, M. 2006. Life history diversity and evolution in the Asterinidae. Integr Comp Biol doi: 10.1093/icb/icj033.
Emlet, R. 2006. Linking stages of life history: how larval quality translates into juvenile performance for an intertidal barnacle (Balanus gladula). Integr Comp Biol doi: 10.1093/icb/icj023.
Gibson, GD and AJF Gibson. 2004. Heterochrony and the evolution of poecilogony: generating larval diversity. Evolution 58:2704–17.[Medline]
Hadfield, MG and MF Strathmann. 1996. Variability, flexibility and plasticity in life histories of marine invertebrates. Oceanol Acta 19:323–34.
Havenhand, N. 1995. Evolutionary ecology of larval types. In McEdward, L (Ed.). Ecology of marine invertebrate larvae Boca Raton, FL CRC Press pp. 79–121.
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Levin, LA. 2006. Recent progress in understanding larval dispersal: new directions and digressions. Integr Comp Biol doi: 10.1093/icb/icj024.
Levin, LA and TS Bridges. 1995. Pattern and diversity in reproduction and development. In McEdward, LR (Ed.). The ecology of marine invertebrate larvae Boca Raton, FL CRC Press pp. 1–48.
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