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BIOL 4900 Honours Thesis Towards an Understanding of Shallow Water Marine Hydroids on Cape Breton Coastlines

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BIOL 4900 Honours Thesis Towards an Understanding of Shallow Water Marine Hydroids on Cape Breton Coastlines BIOL 4900 Honours Thesis Towards an understanding of shallow water marine hydroids on Cape Breton coastlines Matthew S. A. Penney Cape Breton University Sydney, Nova Scotia April, 2017 Supervisor: Dr. Timothy A. Rawlings Towards an understanding of shallow water marine hydroids on Cape Breton coastlines Matthew Penney April 2017 A thesis submitted in partial fulfillment of the requirements for BIOL4900 in the Department of Biology, Cape Breton University, Sydney, Nova Scotia, Canada. The undersigned grants permission for the Cape Breton University Library to store and provide access to this thesis and for the Chair of the Department of Biology, upon request, to allow extensive copying of the material contained in this thesis, in whole or in part, for the purposes of teaching and research by faculty and students of Cape Breton University. Requests for permission to copy or make other use of the material contained in this thesis, in whole or in part, should be addressed to: Chair, Department of Biology Cape Breton University P.O. Box 5300 Sydney, NS B1P 6L2 Canada Student Signature: ________________________________ Date: _______________________ Table Of Contents Abstract ......................................................................................................................................................... 1 Introduction .................................................................................................................................................. 3 Materials & Methods .................................................................................................................................... 8 Sample Collection ..................................................................................................................................... 8 Isolation, Processing, & Fixing................................................................................................................... 9 Morphological Analysis ............................................................................................................................. 9 Molecular Analysis .................................................................................................................................. 10 Phylogenetic Analysis .............................................................................................................................. 14 Results ......................................................................................................................................................... 16 Sample Collection ................................................................................................................................... 16 16S Amplification & Sequencing ............................................................................................................. 17 Identifications ......................................................................................................................................... 18 Species Profiles ....................................................................................................................................... 18 Discussion.................................................................................................................................................... 26 Comparison To Regional Surveys ............................................................................................................ 26 Genetic Connectivity ............................................................................................................................... 26 Clarification of Species-Level Identities .................................................................................................. 28 Monophyly of Bougainvillia .................................................................................................................... 30 Study Limitations .................................................................................................................................... 30 Future Studies ............................................................................................................................................. 32 Conclusion ................................................................................................................................................... 33 Figures ......................................................................................................................................................... 34 Tables .......................................................................................................................................................... 50 Acknowledgments ....................................................................................................................................... 55 References .................................................................................................................................................. 56 Abstract Distributions of North Atlantic marine animals can be expected to shift in response to climate change. Tracking distributions temporally requires an understanding of what species are currently present. Hydroids (Cnidaria, Hydrozoa, Hydroidolina) are conspicuous taxa in the fouling community of shallow water marine environments in Cape Breton, yet surveys of them specific to Cape Breton are sparse and outdated. Consequently, the identity and diversity of local species needs to be clarified. For this study, hydroids were sampled from shallow water environments around Cape Breton and preserved for morphological and molecular analysis. Key morphological features including tentacle anatomy and arrangement, gonophore anatomy, colony structure, and the presence of hydrothecae were examined for family and genus level designations. Sequences were generated for the 16S rRNA barcoding gene and used in BLAST searches to find strong genetic matches on GenBank. Molecular phylogenetic analyses were then performed to refine identifications and determine genetic relationships to other populations. The use of both molecular and morphological analyses has enabled the identification of thirteen taxa to at least genus level and eleven to species level. In total, 14 out of 18 sequences strongly matched GenBank sequences, with similarities ranging from 98%-100%. Two taxa, Podocorynoides minima and Pachycordyle michaeli, do not appear to have been recorded previously in the region. Phylogenetic analysis of genetic connectivity indicated strong genetic relationships with conspecifics in the United States and more distant relationships to conspecifics in Europe. Additionally, it revealed potential cryptic species in Obelia dichotoma. Two genera, Hydractinia and Sarsia, have yet to be identified to species level. Having reliable morphological and molecular databases to identify species, especially for Hydrozoa, is important for monitoring changing distributions over time. This study represents the first stage in the development of a useful dataset for identifying local hydroids using 16S sequence data and accompanying 1 | P a g e morphological descriptions. 2 | P a g e Introduction Hydrozoa is a class within the phylum Cnidaria (Collins, 2002; Daly et al., 2007). Hydrozoans are characterized by septae-lacking polyps which have a chitinous perisarc. In species with a medusoid life stage, reproduction occurs via budding and medusae have a velum and dual nerve rings (Bouillon et al., 2006; Daly et al., 2007). Symmetry of hydrozoans is tetramerous, polymerous, or biradial (Bouillon and Boero, 2000). Members are typically carnivorous, though some feed on phytoplankton and bacteria, and can be heavy consumers of larvae and other marine organisms (Bouillon et al., 2006). The Subclass Hydroidolina is one of two major clades within Hydrozoa, along with Trachylina (Collins, 2002). It includes three major orders: Anthoathecata, Leptothecata, and Siphonophora (Collins, 2000). According to Collins (2000; 2002), Hydroidolina traditionally includes five main groups: Capitata, e.g. Sarsia tubulosa (Schuchert 2010); Filifera, e.g. Bougainvillia muscus (Schuchert, 2007); Hydridae, e.g. the genus Hydra (Schuchert, 2010); Leptomedusa, e.g. the genus Obelia (Cornelius, 1995a); and Siphonophora, e.g. Physalia physalis (Ulster Museum, 1997). However, the monophyly of Filifera has been disputed (Cartwright et al., 2008). There are solitary and colonial species of hydroids. Colonial varieties can exhibit specialized polyps, e.g. gastrozooids for feeding, gonozooids for reproduction, and dactylozooids for defense, all connected via stolons with a continuous gastrovascular canal; the stolon is usually covered by a perisarc. Many hydroids are typically benthic, a notable exception being the clade Siphonophora (Bouillon et al., 2006). Reproduction is indirect, with an asexual polyp stage budding off sexual adult medusae. Hydroids are largely polymorphic, the only exception being 3 | P a g e the clade Hydridae (Collins, 2002), and also have a diverse cnidome, with 21 forms of cnidocysts in total (Bouillon and Boero, 2000). Hydroid species can inhabit a wide range of salinities, as evidenced by the euryhaline Cordylophora caspia (Folino-Rorem and Indelicato, 2005) and marine species such as Plumularia setacea (Schuchert, 2014). They can also be found ranging from shallow to deep waters, with one record of hydroids fouling deep-sea instruments located at a 3690m depth (Blanco et al., 2013). Benthic hydroids
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