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California State University, Northridge Selection on Larval Traits in Early Post-Settlement Temperate and Tropical Reef Fishes

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California State University, Northridge Selection on Larval Traits in Early Post-Settlement Temperate and Tropical Reef Fishes CALIFORNIA STATE UNIVERSITY, NORTHRIDGE SELECTION ON LARVAL TRAITS IN EARLY POST-SETTLEMENT TEMPERATE AND TROPICAL REEF FISHES A thesis submitted in partial fulfillment of the requirements For the degree of Master of Science in Biology By Heidi Elise Block December 2011 The thesis of Heidi Elise Block is approved: Dr. Larry G. Allen Date Dr. Mia S. Adreani Date Dr. Mark A. Steele, Chair Date California State University, Northridge ii ACKNOWLEDGEMENTS First of all, I would like to thank Dr. Mark Steele for helping me with all aspects of this project: from completing my scientific dive certification and teaching me how to catch gobies, to patiently editing many drafts of my thesis, I would not have been able to complete this project without his continuing support. In addition, I would like to thank my committee members Drs. Larry Allen and Mia Adreani for their help in editing my thesis, and general support during my project. I also would like to thank Dr. Steven Dudgeon for providing me with a good statistical foundation, without which I would not have been able to complete the analyses required for my study. I would like to acknowledge all of the people who helped me in the field and lab: Katie Field, David Sinkiewicz, Edwin Leung, Mike Schram, Jenna Krug, Steve Doo, Natalie Low, Jason Selwyn, Barbara Sanchez, Jessica Baker and Audrey Guzowski. Without all of their help this project would not have been possible. I would like to give a special thanks to Katie Field and David Sinkiewicz, for spending many hours underwater helping me to collect fish and complete surveys, and to Edwin Leung and Mike Schram, for volunteering to read close to a thousand of my otoliths. The Wrigley Institute for Environmental Studies provided me with financial and logistical support for the Catalina portion of this project, for which I am very grateful. I would also like to thank Northeastern University’s Three Seas Program for providing me with the opportunity and financial support to complete my study in Moorea; and the Gump South Pacific Research Station for their logistical support. Lastly, I would like to acknowledge California State University, Northridge for providing additional funding for this project. iii TABLE OF CONTENTS Signature Page ii Acknowledgements iii Abstract v Chapter 1: General introduction 1 Chapter 2: Spatially and temporally inconsistent selection on 6 larval traits in two common kelp forest fishes Introduction Methods Results Discussion Tables and Figures Chapter 3: Selective mortality on larval traits in two coral reef fishes 37 in Moorea, French Polynesia Introduction Methods Results Discussion Tables and Figures Chapter 4: Conclusions 60 Literature Cited 63 iv ABSTRACT SELECTION ON LARVAL TRAITS IN EARLY POST-SETTLEMENT TEMPERATE AND TROPICAL REEF FISHES By Heidi Elise Block Master of Science in Biology In marine organisms with complex life histories it is possible for events occurring during earlier life stages to affect an individual’s subsequent survival. In fishes, it has been proposed that settlement, the transition from a pelagic habitat to a benthic one, is a critical period in determining survivorship. Selection (i.e., non-random mortality) may occur on specific larval traits, which could influence an individual’s probability of survival after settlement. Herein I examine the effects of larval traits on the survival of juveniles shortly after settlement in two temperate and two tropical reef fishes. This was achieved by comparing traits of individuals sampled just after settlement with those of individuals from the same cohort sampled one month later, which evaluated selective mortality occurring over a cohort’s first month of life on the reef. The two temperate species studied were señorita, Oxyjulis californica, and blackeye goby, Rhinogobiops nicholsii. Collections were completed at four sites at Santa Catalina Island, California, and two times between June and August of 2009. For señorita, I examined four larval traits: planktonic larval duration (PLD), size at settlement, metamorphosis band width, and pre-settlement growth rates. For the blackeye goby I examined two traits: size at settlement and pre-settlement growth rates. This study v revealed that selection was inconsistent in both species. There were differences in both the direction and intensity of selection between the different sites as well as times. Differences in selection intensity were not related to variation in habitat characteristics or conspecific densities. Unlike selection, I found some consistent differences in larval traits between cohorts in señorita. For the señorita I found that individuals settling earlier in the summer had longer PLDs and slower growth rates than those settling later in the summer. In Moorea, French Polynesia I studied the blue green chromis, Chromis viridis, and the Gnatholepis species complex, Gnatholepis scapulostigma/anjerensis. This study was conducted at two sites on the North side of the island. Unlike the temperate study, only one cohort was examined due to logistical constraints. I examined three larval traits for both “species”: PLD, size at settlement, and pre-settlement growth rates. For the Gnatholepis species complex, I did not find any evidence of selection or any differences between the two study sites for any of the larval traits measured. For Chromis viridis, I found that there was selection for larger sizes at settlement at both sites. However, selection on the other two larval traits was inconsistent between the two sites, with one site showing selection for faster pre-settlement growth rates and the other showing selection for longer PLDs. For this study I also explored whether differences in habitat characteristics or predator densities between the two sites might be influencing these patterns. These two sites were not significantly different in terms of habitat or predator densities and therefore these factors could not explain differences in selection between sites. vi These two studies reveal that selection on larval traits can vary greatly on relatively small scales: with sites less than 3 km apart exhibiting differences in the direction or strength of selection on a larval trait. This small-scale variation in selection may be one reason why variability in larval traits is maintained in these species. Understanding the causes of variation in larval traits in these populations and how variation is being maintained, through processes such as selective mortality, will help to understand and predict recruitment dynamics and ultimately community structure. vii Chapter 1 General Introduction Many organisms exhibit complex life history strategies, e.g. amphibians, plants, insects, fishes, and marine invertebrates (Wilbur 1980). Complex life histories involve multiple life stages, which often provide individuals with the chance of higher dispersal during a particular phase. In many amphibians for instance, there is a larval (tadpole) life stage that is fully aquatic and has limited dispersal potential; whereas after metamorphosis into the terrestrial adult stage, individuals have much higher potential for dispersal (Wilbur 1980). The opposite of this pattern is usually seen in plants, in which the highest dispersal is for seeds as compared to sessile adults. Dispersal is important for populations because it can decrease crowding and competition, as well as increase genetic diversity and decrease inbreeding (Hedgecock 1986, Pechenik 1999). Many benthic marine organisms have complex life histories with a benthic- associated adult stage and a planktonic larval stage. These different stages provide benefits, and present challenges for marine organisms. One of the major benefits of having multiple life stages is the possibility for dispersal to new environments. Dispersal to new locations can, as mentioned earlier, decrease crowding and potential competition at one location as well as increase genetic diversity within a population by providing a wider range of suitable mates. Although some mobile marine organisms disperse as juveniles and adults, many benthic reef based organisms rely on larval stages for dispersal. Dispersal of marine larvae is important in connecting populations, which can be important for sustaining them. For example, if one population experiences a decline, a nearby population can supplement that population through larval dispersal. Due to the 1 importance of larval dispersal in population sustainability, larval dispersal and connectivity have been widely studied for the purposes of fisheries management and conservation (Levin 2006, Cowen and Sponaugle 2009). One of the challenges associated with larval dispersal is the possibility that the larvae will never find suitable settlement habitat, and will die in the plankton. When larvae leave their natal reef they are leaving behind known suitable habitat in order to find another potentially better location, which may or may not exist. Additionally, there is extremely high mortality of larvae in the plankton, as well as at the time of settlement (Steele and Forrester 2002, Almany 2003, Doherty et al. 2004, Almany and Webster 2006). The settlement transition has been considered a critical period due to this high mortality (Searcy and Sponaugle 2001, Doherty et al 2004, Almany and Webster 2006). For organisms with complex life histories there is the possibility that events occurring during one life stage might influence their success later in life. This is one reason that the settlement transition, the transition from pelagic to
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