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Mechanisms of Coexistence Between Two Octopus Species in a South Florida Lagoon

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Mechanisms of Coexistence Between Two Octopus Species in a South Florida Lagoon MECHANISMS OF COEXISTENCE BETWEEN TWO OCTOPUS SPECIES IN A SOUTH FLORIDA LAGOON by Chelsea Bennice A Dissertation Submitted to the Faculty of The Charles E. Schmidt College of Science In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Florida Atlantic University Boca Raton, FL May 2019 Copyright 2019 by Chelsea Bennice ii MECHANISMS OF COEXISTENCE BETWEEN TWO OCTOPUS SPECIES IN A SOUTH FLORIDA LAGOON by Chelsea Bennice This dissertation was prepared under the direction of the candidate's dissertation advisor, Dr. W. Randy Brooks, Department of Biological Sciences, and has been approved by all members of the supervisory committee. It was submitted to the facu1ty of the Charles E. Schmidt College of Science and was accepted in partial fulfillment of the requirements for the degree of Doctor ofPhilosophy. SUPER~ORY ~E: f;V.r~ W. Randy Brooks, Ph.D. Disserta · Advisor ._,l£11111r~. 4~~ Roge:7 anion, Ph.D. ~Q Colin Hughes, Ph.D. ta.J!mtlrR"' ~Cimi, Ph.D. ean, The Charles E. Schmidt College of Science /}WeJ, t1.,. l 1d!'J Khaled Sobhan, Ph.D. Date Interim Dean, Graduate College iii ACKNOWLEDGEMENTS I express sincere gratitude to my committee members for all of their guidance and support, and special thanks to my major advisors for their persistence, patience, and encouragement during the completion of my PhD project. I thank Florida Atlantic University’s scientific divers for accompanying me on research dives throughout this study; specifically, Liana Houston, Jessica Pate, Rachel Shanker, Danielle Bartz, Jeanette Wyneken, and Marianne Porter. Thank you to FAU’s diving and boating safety program for providing SCUBA tanks and compressed air. Laz Ruda discovered Macrotritopus defillippi at BHB and invited Roger Hanlon to dive this site, which led to this research project. I am grateful for funding from the Broward Shell Club, Palm Beach Fishing Club, and the Animal Behavior Society. Underwater photographs and octopus observations were also acquired from SCUBA divers: Sandra Edwards, David Sanchez, Anne DuPont and Linda Ianniello. We are grateful to Stephanie Farrington for her guidance on CPCe software, Andrew Rayburn for his guidance on spatial statistics, and Conor Sullivan for the development of the octopus 24-h camera. Thank you to my family and friends for their love and support. iv ABSTRACT Author: Chelsea Bennice Title: Mechanisms of coexistence between two octopus species in a South Florida Lagoon Institution: Florida Atlantic University Dissertation Advisor: Dr. W. Randy Brooks Degree: Doctor of Philosophy Year: 2019 Theoretically, sympatric species must partition resources or space to allow for coexistence. Determining empirically the specific resources each species exploits and species’ interactions (e.g., intra- and interspecific competition) can sometimes be challenging, thus the data are relatively sparse for certain taxa. This paucity of data exists for octopuses. Therefore, I chose to study niches of two sympatric octopuses (Octopus vulgaris and Macrotritopus defilippi) in an intracoastal habitat. Specifically, I assessed (1) spatial distribution of octopus home or “den” space, (2) habitat association, (3) octopus abundance, (4) foraging activity periods, (5) diet, and (6) associated substrates and behaviors used during foraging events. Octopus den locations were marked by GPS to quantify spatial patterns of both species and their v spatial relationship to each other. Habitat associations were measured by quantifying photoquadrats of den and surrounding habitats. For foraging activity periods, a video camera was placed near an octopus den for 24-h observation to determine when each octopus species leaves/returns from foraging. Underwater video recording was used to determine associated foraging substrates and behaviors for both species. Prey remains from octopus’ dens and video recordings indicating prey consumption were used to determine diets of the two octopus species. Video recordings from the 24-h camera and foraging behavior events also provided observations of intra- and interspecific interactions. Results revealed that the two species are interspersed throughout the shallow Florida lagoon and are both abundant during the spring months (March, April, May). Although both species are interspersed throughout the lagoon, their den and surrounding habitat association differed. O. vulgaris was associated with hard bottom and M. defilippi was associated with soft bottom, thus they may not compete strongly for habitats. Each species used different foraging strategies and different primary prey, which may also lessen competition and facilitate coexistence. O. vulgaris had peak foraging activity during night hours, foraged mostly on hard bottom and mainly consumed bivalves while M. defilippi had peak foraging activity during day hours, foraged mostly on soft bottom and mainly consumed crustaceans. Octopuses also had species-specific foraging behaviors, with O. vulgaris using parachute attack and M. defilippi using flounder swimming and tripod stance. Additional intra- and interspecific interactions were video recorded and included: fishes following octopuses, predation attempts, agonistic encounters, cannibalism, and tactile communication. This study identified ecological and behavioral components that may facilitate coexistence of these vi sympatric species, provided insight into cephalopod niches and ecology, and provided baseline conservation requirements for sand-dwelling cephalopods, both of which may be using this site as a mating and nursery habitat. vii MECHANISMS OF COEXISTENCE BETWEEN TWO OCTOPUS SPECIES IN A SOUTH FLORIDA LAGOON LIST OF TABLES ............................................................................................................. xi LIST OF FIGURES .......................................................................................................... xii INTRODUCTION .............................................................................................................. 1 General overview of species coexistence and niche partitioning ................................... 1 Cephalopod niche partitioning ........................................................................................ 3 CHAPTER I: FINE-SCALE HABITAT PARTITIONING FACILITATES SYMPATRY BETWEEN TWO OCTOPUS SPECIES IN A SHALLOW FLORIDA LAGOON .......... 5 ABSTRACT .................................................................................................................... 5 INTRODUCTION .......................................................................................................... 6 MATERIALS AND METHODS .................................................................................... 8 Description of species ................................................................................................. 8 Study site ..................................................................................................................... 8 Spatial distribution ...................................................................................................... 9 Habitat association .................................................................................................... 10 Octopus abundance ................................................................................................... 13 RESULTS ..................................................................................................................... 13 Spatial distribution .................................................................................................... 14 Habitat association .................................................................................................... 15 viii Octopus abundance ................................................................................................... 16 DISCUSSION ............................................................................................................... 17 CHAPTER II: OCTOPUS COEXISTENCE FACILITATED BY FORAGING AND DIET STRATEGIES IN A SOUTH FLORIDA LAGOON ............................................. 30 ABSTRACT .................................................................................................................. 30 INTRODUCTION ........................................................................................................ 31 MATERIALS AND METHODS .................................................................................. 32 Description of species ................................................................................................... 32 Study Site ...................................................................................................................... 33 Foraging activity periods .............................................................................................. 33 Diet ................................................................................................................................ 35 Foraging strategies (foraging substrates & behaviors) ................................................. 36 RESULTS ......................................................................................................................... 39 Foraging activity periods .............................................................................................. 39 Diet ................................................................................................................................ 40 Foraging strategies (substrates & behaviors
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