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Meghan Dueck Miniaturization in Alestidae (Ostariophysi: Characiformes) by Meghan Dueck A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Systematics and Evolution Department of Biological Sciences University of Alberta © Meghan Dueck, 2020 Abstract Alestidae (African tetras) is one of the most species-rich families of the order Characiformes and exhibits a pronounced diversity of body size. Two independent miniaturization events (a drastic phylogenetic decrease in body size) have been identified in Alestidae. Miniaturization is frequently accompanied by morphological novelties, structural reductions and losses, hyperossifications, and increased intraspecific variation, which can create difficulties when establishing natural classifications. Traditionally, all “small” species of Alestidae with reduced multicuspid teeth were grouped in the tribe Petersiini. This tribe has since been recognized as polyphyletic, but a limited understanding of the intrarelationships of Alestidae persists. Few osteological descriptions for small-bodied alestids are available but are needed to assess the effects of miniaturization on osteological features of these fish. I conducted morphological studies of continuous measurements and meristic characters within a sample of a small-bodied alestid, Hemigrammopetersius barnardi (Herre) (n = 161), and a petersiin with a comparatively larger body size, Petersius conserialis Hilgendorf (n = 17). Specimens of H. barnardi, P. conserialis, and juveniles of an alestid that attains a larger adult size, Alestes dentex (Linnaeus), were cleared and counterstained for bone and cartilage for comparative osteological analyses and to identify potential convergent features resulting from a decrease in body size. Measurements in the examined sample of H. barnardi was normally distributed; all specimens had a relatively small body size (19.2 – 34.9 mm standard length) compared with the reported maximum standard length of 70 mm for H. barnardi. Potential sexual dimorphism was identified in the anal fin of cleared and stained specimens of H. ii barnardi, supporting previous observations in the literature. I compared osteological descriptions for H. barnardi and P. conserialis, which represent some of the few osteological descriptions for small-bodied characiforms and will serve as a foundation for future comparisons. I describe a size series for P. conserialis that addresses previous discrepancies in the literature regarding the presence or absence of a parietal fontanelle and reports the presence of a parietal fontanelle in all examined specimens. I document negative allometric growth between standard length and the length and width of the parietal fontanelle. Finally, I compared specimens of H. barnardi and P. conserialis to juvenile specimens of A. dentex. This allowed the identification of features that are convergent due to a decrease in body size. I found that certain bones, including the supraorbital and postcleithrum 3, are absent in H. barnardi but are present in A. dentex and P. conserialis. In addition, I found reductions to the lateral line system, such as the loss of the anterodorsal branch of the circumorbital sensory canal on the dermosphenotic and an incomplete lateral line, in H. barnardi in comparison with other examined species. These features arise towards the end of development and have been reported as lost or reduced in other small and miniature characiforms. Caution should be used when including features that are susceptible to developmental truncation for phylogenetic analyses as they are likely to independently occur with body size decreases and may not be indicative of close phylogenetic relationships. iii Acknowledgements I am forever indebted to the many individuals who have substantially contributed to my thesis work and continual development as a scientist. I would not be where I am today without their commitment and passion. 1. I wish to express my deepest gratitude to my supervisor, Dr. Alison Murray, for her expertise, encouragement, and countless revisions. It was an honor to work with Dr. Murray, her commitment to her students and their learning has tremendously enriched my experience as a graduate student and I will be forever grateful. 2. A huge thank you to Dr. Heather Proctor for her invaluable feedback throughout the project and for being on my supervisory committee. 3. I would like to pay my special regards to John Acorn, for taking the time to read my thesis as the arm’s length examiner. 4. Thank you to additional faculty and staff from the Department of Biological Sciences at the University of Alberta. They made the entire process of my masters exceptionally easier by assisting with mind-boggling administrative requirements to providing thought-provoking conversations and advise on select skills. I would like to extend my gratitude to Dr. Aaron LeBlanc, Dr. Audrey Reid, Dr. Braden Barr, Dr. Corwin Sullivan, Mark Wolansky, Dr. Michael Caldwell, Dr. Robert Holmes, and Shelley Scott. 5. I am grateful to the Department of Biological Sciences and the Government of Alberta for providing funding that enabled this project. 6. A huge thank you to Emily Drystek, who set aside her valuable time to provide advice on resume-writing, interview tips, and general skills to help succeed outside academics. 7. My deepest gratitude to past and present fellow graduate students: Aaron Dyer, Annie McIntosh, Curtis Dinn, Emily Upham-Mills, Dr. Gregory Funston, Dr. Ilaria Paparella, Jasdeep Kaur, Katherine Bramble, Luthfur Rahman, Mark Powers, Matthew Rhodes, Oksana Vernygora, Rebekah Vice, Samantha Hamilton, Sinjini Sinha, Sydney Mohr, and Yan-yin Wong. iv 8. I am thoroughly grateful to those who have supported my previous research endeavors, encouraged my research interests, and continually check in on me and offer help in any possible way. I would specifically like to thank Dr. Bernadette Quémerais, Dr. Binal Gandhi, Eduardo Reyes-Serratos, Dr. Mrinal Das, Dr. Paige Lacy, Samineh Kamravaei, Dr. Sindhu Nair, and Dr. Vivek Gandhi. Finally, words will never express how exceptionally grateful I am for the extensive patience, understanding, and support of my family and friends. Specifically, my parents (Irv and Denise Dueck), sibling and roomie (Jessica Dueck), best friends (Nicole Roshko and Jenna LaBerge), and boyfriend and icecream-provider (Gavin Bradley). If it wasn’t for them, I would have shed a lot more tears. v Table of Contents Chapter 1: Introduction ......................................................................................................................................... 1 1.1 General Introduction .................................................................................................................................... 1 1.2 Alestidae ...................................................................................................................................................... 2 1.3 Miniaturization in Fish .................................................................................................................................. 4 1.4 Thesis Objectives.......................................................................................................................................... 6 1.5 References ................................................................................................................................................... 7 Chapter 2: Variation in the small African fish Hemigrammopetersius barnardi (Characiformes: Alestidae) ......... 12 2.1 Introduction ............................................................................................................................................... 12 2.2 Materials and Methods .............................................................................................................................. 14 2.2.1 Materials ............................................................................................................................................ 14 2.2.2 Measurements .................................................................................................................................... 14 2.3 Results ....................................................................................................................................................... 16 2.3.1 General Body Form and External Characteristics .................................................................................. 16 2.3.2 Intraspecific Variation in External Measurements ................................................................................ 16 2.3.3 Skull.................................................................................................................................................... 17 2.3.4 Jaws and Teeth ................................................................................................................................... 18 2.3.5 Opercular Bones ................................................................................................................................. 19 2.3.6 Suspensorium ..................................................................................................................................... 19 2.3.7 Infraorbital Bones ............................................................................................................................... 20 2.3.8 Branchial Arches and Pharyngeal Region ............................................................................................
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