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Rare Marine Macroalgae of Southern Australia

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Rare Marine Macroalgae of Southern Australia Rare Marine Macroalgae of Southern Australia by Fiona Jean Scott B.Sc. M.Sc. Institute for Marine and Antarctic Studies and School of Geography and Environmental Studies Submitted in fulfilment of the requirements for the Doctor of Philosophy University of Tasmania December 2012 Declaration of Originality This thesis contains no material which has been accepted for a degree or diploma by the University or any other institution, except by way of background information and duly acknowledged in the thesis, and to the best of my knowledge and belief no material previously published or written by another person except where due acknowledgement is made in the text of the thesis, nor does the thesis contain any material that infringes copyright. Signed: ______________________ ii Authority of Access This thesis may be made available for loan. Copying and communication of any part of this thesis is prohibited for two years from the date this statement was signed; after that time limited copying and communication is permitted in accordance with the Copyright Act 1968. iii Acknowledgements I sincerely thank Graham Edgar, Jamie Kirkpatrick and Neville Barrett for guidance and support throughout my candidature. Abbreviations and Sources of Data AVH: Australia’s Virtual Herbarium, Council of Heads of Australasian Herbaria, viewed March 2009, http://avh.ala.org.au/ AD: State Herbarium of South Australia, http://www.flora.sa.gov.au/index.html PERTH: Western Australian Herbarium, http://www.dec.wa.gov.au/content/category/41/831/1821/ MEL: National Herbarium of Victoria,, Royal Botanic Gardens Board, Melbourne, MELISR database, dated 03/03/2009. http://www.rbg.vic.gov.au/science/information-and-resources/national- herbarium-of-victoria HO: Tasmanian Herbarium, Hobart http://www.tmag.tas.gov.au/collections_and_research/tasmanian_herbarium2, GA: Geosciences Australia, http://www.ga.gov.au/ CSIRO: Commonwealth Scientific Industry and Research Organisation, CSIRO Atlas of Regional Seas data. http://www.marine.csiro.au/~dunn/cars2009/ FS: Fiona Scott personal herbarium Diving and logistic assistance Neville Barrett, Matt Cameron, Toni Cooper, Marlene Davey, Graham Edgar, Amelia Fowles, Dane Jones, Gerald Kraft, Tania Mendo, Iona Mitchell, Elisabeth Oh , Prue Scott, Rick Stuart-Smith, Jemina Stuart-Smith , Robert Wills, Carolina Zagal. Technical assistance Justin Hulls and Mark Morphett (ArcGIS), Austin Pepper (database formatting), Russell Thomson (Random Forests), Brett Summerell (AVH data), Robert Baldock, Rex Hunt and Fred Gurgel (AD herbarium data), Timmothy Newlands and Lynnette Cave (HO herbarium data), Ann Preece (proof-reading), Parks Tasmania (collecting permits). Financial assistance National Environmental Research Program (NERP) / Commonwealth Environmental Research Facilities (CERF) Marine Biodiversity Hub, (PhD scholarship awarded to Fiona Scott). The Winifred Violet Scott Charitable Trust (grant to Fiona Scott). Australian Research Council (ARC) linkage project ‘Incorporation of vulnerability and irreplaceability into marine protected area planning’ (funding to Graham Edgar, the candidate’s principal PhD supervisor). iv Abstract The aims of this thesis were to document the pattern of distribution of the apparently rare marine algal species endemic to southern Australia, test the validity of the apparent rarity and seek causes of the patterns. A review of the concepts and causes of rarity with special reference to the marine macroalgae of temperate Australia is preceded by a brief background to macroalgal taxonomy, growth patterns, reproduction and ecology. One hundred and forty-two species met the selected rarity criterion of occurring at five or fewer localities, as data-based in the Australian Virtual Herbarium. Seventy-one rare species were of broad range (>500 km), forty-five of narrow range (50–500 km), and twenty-six of restricted range (<50 km). The characteristics common to > 60% of rare macroalgae were that of small size and of filamentous or coarsely-branched habit. Of the three major taxonomic groups (Divisions Chlorophyta, Heterokontophyta and Rhodophyta) rare species within the Heterokontophyta (brown algae) were proportionally under-represented. Recognised rare species are discussed in terms of taxonomic affiliation, range, form and function, and co-occurrence of species in localised areas. Six ‘centers of rarity’, areas with high proportions of rare species, were identified. These were Rottnest I WA, King George Sound WA, Eucla WA, Fowlers Bay SA, Port Phillip Bay VIC and D’Entrecasteaux Channel TAS. Using the random forest procedure, analyses indicated that there are particular environments and distinct environmental extremes that favour high proportions of rare species. Associations were found between high proportions of rare species and extremes of sand substratum, nitrate, phosphate, and chlorophyll-a. Overall, the analyses suggested that rare species are predominantly associated with low-nutrient environments and sandy substrata, whereas highly productive waters appear to be bereft of rare species. The potential association of rare species with rare environments was tested. Fifteen environmental domains were identified, and Port Phillip Bay VIC and south-east Tasmania emerged as areas particularly favourable for rare species. However, the v spatial extent of domains was not related to the mean proportion of threatened species within them. Other possible explanations for concentrations of rare species or high proportions of rare species, are associations with habitat complexity, or associations with escape routes along glacial drainage pathways during sea level change. Such conditions pertain to the areas with the highest concentrations of rare species. A field program of target-searching for rare species was undertaken to elucidate rare species distributions. Results from underwater visual census surveys (UVC) supported the patterns of rare species found in historical herbarium records. A total of 489 macroalgal species were recorded collectively from 111 UVC surveys. Sixteen rare species were observed, representing ~11% of the 142 rare species. Notable range- extensions are reported for 7 rare species, and specific life history phases are newly recorded for 2 rare species. A novel find that comprises a new family and probable new order, Entwisleia bella gen. et sp. nov. (F. Entwisleiaceae fam. nov.), was described upon a suite of morphological features that distinguish it from other known taxa. Planned genetic analyses will elucidate the ordinal classification of the new taxon and contribute to the formal proposal of the species. Fifty rare marine macroalgae have been recorded from sites within sanctuary (no-take) zones of existing marine protected areas. A general paucity of adequate abundance and seasonality data thwarts confirmation of occurrence and/or persistence of these species within sanctuary zones, and therefore they cannot be firmly considered as adequately safeguarded. vi Table of Contents Acknowledgements …………………………………………………………….. iv Abstract ………………………………………………………………………… v Chapter 1: Rare marine macroalgae– Introduction ………………………….... 1 1.1 Marine macroalgae: An overview……………………………………….. 1 1.1.1 Algal taxonomy ……………………………………………………….. 1 1.1.2 Growth patterns and forms …………………………………...……….. 4 1.1.3 Reproduction and life histories ………………………………..….…… 5 1.1.4 Aspects of macroalgal ecology ……………………………………...... 6 1.2 Why southern Australia is so interesting ……………………………….. 8 1.2.1 Biogeography of Australian taxa ………………………………….….. 8 1.2.2 Biogeography of southern Australian taxa …………………………… 10 1.2.3 Endemism of flora and fauna …………………………………………. 14 1.3 Rarity ………………………………………………………………….… 19 1.3.1 Concepts of rarity …………………………………………..……….… 19 1.3.2 Types of rarity ………………………………………………….….….. 21 1.3.3 Causes of rarity ………………………………………………….……. 23 1.3.4 Assessment of threat …………………………………………….……. 27 1.3.5 Macroalgal rarity ……………………………………………….….…. 31 1.4. Means of investigating distributions of rare species …………………… 31 1.4.1 Field surveys …………………………………………………………. 32 1.4.2 Historical records ………………………………………………….…. 33 1.5 Structure of thesis ………………………………………………..…….. 35 1.6. Bibliography ……………………………………………………...……. 36 Chapter 2: Rare marine macroalgae– Taxonomy and distribution ……....…… 48 2.1 Introduction and background …………………………………………... 48 2.2 Materials and methods ……………………………………………….…. 49 2.2.1 Areas of interest ………………………………………………………. 49 2.2.2 Sources of data ………………………………………………..…….... 49 2.2.3 Definitions and criteria …………………………………………….…. 50 2.2.4 Geographic rarity estimated from literature ……………………….….. 53 2.2.5 Functional groups of rare species ………………………………….….. 53 vii 2.2.6 Statistical analyses …………………………………………..…….….. 54 2.3 Results ………………………………………………………………… 54 2.3.1 Rare species …………………………………………………………. 54 2.3.2 Patterns of rare species distributions ………………………………… 60 2.3.3 Co-occurrence of rare species ……………………………………….. 64 2.3.4 Functional groups of rare species …………………………………… 66 2.3.5 Range-restricted rare species ……………………………………….. 66 2.3.6 Species richness …………………………………………………….. 69 2.4 Discussion ……………………………………………………………... 72 2.4.1 General observations …..……………………………………………. 72 2.4.2 Estimates of rare species numbers ……………………………….…. 75 2.4.3 Patterns of macroalgal rarity and richness ………………………….. 77 2.5 Bibliography …………………………………………………………… 79 Chapter 3: Concentrations of rare macroalgae ….………………………….…. 82 3.1 Introduction …………………………………………………………… 82 3.2 Materials and methods ….……………………………………………… 83 3.2.1 Study
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