Life History Characteristics of Glassfish, Ambassis Jacksoniensis, Adjacent
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Life history characteristics of glassfish, Ambassis jacksoniensis, adjacent to saltmarsh within a large and permanently-open estuary Jack J. McPhee Doctor of Philosophy (Environmental and Life Sciences) Supervisors: Dr Maria Schreider (Environmental and Life Sciences) Dr Margaret Platell (Environmental and Life Sciences) “Ambassis jacksoniensis” - Illustrated by Corrine Edwards a ACKNOWLEDGEMENTS I would like to express my sincerest gratitude to a series of people, without whom this PhD would not have been possible. First and foremost, I would like to give a warm thank you to my two supervisors, Dr Maria Schreider and Dr Margaret Platell. Maria, your encouragement to push on, not for self-benefit, but for the greater scientific good is a trait that I have valued since your teaching during my undergraduate years. Your motive to work hard in order to seek the truth (Без муки нет науки) is a characteristic that often reminds me why I was inspired to pursue a scientific career to begin with – спасибо! Margaret, your genuinely friendly and inquisitive attitude towards the project, and science in general, is a characteristic that has also shaped me over the years. Your genuine care for the organisms and environments that we study is a continual reminder that such scientific pursuits are not only for the benefit of the scientific community, but are of equal importance to the organisms that we are studying. While Maria’s traditional, clinical, to the point (i.e. “eschew obfuscation”) scientific perspective helped me “cut the fat” during my studies, Margaret has brought the “seasoning,” the fun, the flavour. Margaret, to me you are truly perspicacious in the field of estuarine ecology and I thank you for eliciting me into this world. The two of you are genuinely altruistic scientists in my eyes and I am hugely grateful for all your inspiration. Thank you for being wonderful teachers, colleagues and friends. I would also like to thank the numerous enthusiastic colleagues that assisted me in the often wild and uncomfortable sampling occasions. A special thank you to Jeff Law, Ryan Hefford, Wayne Davis, Stuart Bowers, James Brooker and Daniel Camilleri. Thank you Gordon Thomson, from Murdoch University, for preparing the gonadal slides for microscopic validation. Thank you Macquarie Geotech for conducting the calorimetric analyses. Thank you Natalie Moltschaniwskyj, for your guidance with the life history and reproduction chapter. I would also like to thank the University of Newcastle for their provision of funding and resources throughout the duration of the project. Thank you Corrine Edwards for illustrating the cover of my thesis. Lastly, I would like to thank my family and friends (particularly my Mum and Dad, Micaela and to my dogs, Willow, Jessie and Pippa), who have all supported me and kept me sane and motivated throughout the entire process of my PhD. The collection of fish and zooplankton was authorised by the Department of Primary Industries (Permit No. P11/0085-1.0) and UoN Animal Ethics Committee (Permit No. A-2012-219). In memory of Dr Kenneth Zimmerman - the man who inspired me to study marine science. i STATEMENT OF ORIGINALITY This thesis contains no material that has been accepted for the award of any other degree or diploma in any university or other tertiary institution and, to the best of my knowledge, contains no material previously published or written by another person, except where due reference has been made in the text. _________________________________________________________ Jack James McPhee ii PUBLICATIONS RESULTING FROM THIS THESIS The following article has been published by Estuarine, Coastal and Shelf Science and a copy of the publication is attached at the end of this thesis (see Appendix A). Other chapters/components of the research are currently being prepared for submission to various ecological journals for publication. McPhee, J. J., Platell, M. E., and Schreider, M. J. (2015). Trophic relay and prey switching – A stomach contents and calorimetric investigation of an ambassid fish and their saltmarsh prey. Estuarine, Coastal and Shelf Science, 167, 67-74. iii ABSTRACT Saltmarsh vegetation, which typically occurs in intertidal areas within estuaries globally, provides an important habitat and feeding ground for estuarine organisms such as crustaceans, gastropods, birds and fish (some of which are of economic importance). Within south-east Australian estuaries, saltmarsh vegetation is both typically bordered by mangroves and tidally inundated three or four times per month during the high tide of the spring tidal cycle (during the day high tide in summer and during the night high tide in winter). In recent decades, saltmarsh vegetation has declined globally due to anthropogenic influence, and in Australia, ‘Coastal Saltmarsh’ is now listed as an Endangered Ecological Community under the Threatened Species Conservation Act 1995. This study was conducted within a representative and relatively “unmodified” saltmarsh habitat (Empire Bay Wetland) in a large and permanently open estuary, Brisbane Water Estuary, located in south-eastern Australia. This study, which was conducted at two markedly different times of the year during 2012, examined the general “response” of the estuarine fish (using seine nets) and zooplankton (using plankton nets) assemblages to tidal inundation, with further emphasis being placed on selected biological and ecological characteristics of the abundant estuarine ambassid, Ambassis jacksoniensis. Abundances of A. jacksoniensis (mean standard length=37.3 mm, ±0.021 (SE)) and overall fish diversity were greater in nightly winter catches than daily summer catches, which is consistent with previous evidence of important feeding times for estuarine fish (including A. jacksoniensis) upon saltmarsh- derived zooplankton (e.g. crab zoeae released by saltmarsh-dwelling grapsid crabs), during ebb tides that drain saltmarsh following its inundation. Indeed, zooplankton assemblages were dominated by crab zoeae during ebb tides following saltmarsh inundation, while calanoid copepods dominated these assemblages at other times. Moreover, stomach content analyses of A. jacksoniensis showed that crab zoeae were heavily preyed upon during such times, with dietary “switching” to caridean decapods being evident when crab zoeae were not abundantly present within the water column (i.e. during flood tides and during ebb tides that did not follow saltmarsh inundation; as shown within zooplankton assemblages). Despite their high abundance within zooplankton assemblages, calanoid copepods were not preyed upon by A. jacksoniensis, which is likely to reflect the relatively fast escape responses of iv calanoids to predators. Further, stomach fullnesses of A. jacksoniensis were generally highest during ebb tides on days of saltmarsh inundation, implying that feeding was most marked at these times. Trophic relay is an ecological model that involves the movement of biomass and energy from vegetation, such as saltmarshes, within estuaries to the open sea via a series of predator-prey relationships. Therefore, the trophic relationship between saltmarsh-dwelling grapsid crabs (which feed on saltmarsh-derived detritus and microphytobenthos), A. jacksoniensis and their predators (which include economically important fish, such as Acanthopagrus australis, Platycephalus fuscus and Argyrosomus japonicus, provides evidence of partial trophic relay within this system, and thus highlights the ecological and economic importance of saltmarsh within this system. The trophic relationship between A. jacksoniensis and its zooplanktonic prey (e.g. crab zoeae, which is of a red/orange colour) was further investigated, for the first time, by comparisons of the calorimetric contribution of its potential prey (i.e. crab zoeae, and the far paler caridean decapods and calanoid copepods), which found no difference in the energetic densities among such potential prey, suggesting that prey (i.e. zooplankton) abundance and/or prey visibility (due to colour) has a stronger relationship than prey energetic density to the diets of A. jacksoniensis. The feeding ecology of A. jacksonsiensis was also explored, for the first time, in light of its various life history characteristics (e.g. the seasonality of sex ratios, sexual maturity and somatic/reproductive growth), with links being made between saltmarsh-derived tropic relay and energetic requirements for reproductive purposes. Thus, the gonads of A. jacksoniensis were found to be generally maturing and ripe during summer, while juvenile/inactive and spent gonads were prevalent during winter, consistent with previous evidence that A. jacksoniensis spawn during summer with a lull during winter. The sex ratios of A. jacksoniensis were also heavily female-biased during summer before equalising (to approximately 1:1) during winter, suggesting that male A. jacksoniensis may avoid the shallow sampling locations (seagrass adjacent to the saltmarsh/mangroves) in a strategy to counteract visual predation from fish and birds during daytime (summer) before returning to these waters during the night winter, during a lull in spawning, for important feeding opportunities. Female A. jacksoniensis, alternatively, may remain in such vulnerable locations due to increased energetic requirements for reproductive purposes (as demonstrated in male vs female somatic/gonadal growth v analyses). These findings therefore suggest that the seasonal timing of spawning for A. jacksoniensis may be linked to their feeding