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A New Species of Bidens (Asteraceae: Heliantheae) from Starbuck Island Provides Evidence for a Second Colonization of Pacific Islands by the Genus

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A New Species of Bidens (Asteraceae: Heliantheae) from Starbuck Island Provides Evidence for a Second Colonization of Pacific Islands by the Genus A new species of Bidens (Asteraceae: Heliantheae) from Starbuck Island provides evidence for a second colonization of Pacific islands by the genus by Cynthia Sayre BSc, (1994), University of British Columbia A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES Department of Botany We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA July 2001 ©Cynthia Sayre, 2001 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the head of my department or by his or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department The University of British Columbia Vancouver, Canada ABSTRACT An unknown Bidens was recently discovered on Starbuck Island, a small, remote coral atoll in the South Pacific and one of the least visited islands in the world. I determined the identity and evolutionary origin of this Bidens using both morphological analyses and phylogenetic relationships in Bidens (Asteraceae; Heliantheae) and Coreopsis (Asteraceae; Heliantheae) using variation in nuclear rDNA internal transcribed spacer (ITS) sequences. The ITS analyses include all species of Bidens sequenced to date, including five from this study and sequences from Ganders et al. (2000) and a selection of Coreopsis sequences from Kim et al. (1999). Using morphological and molecular data 1 tested the following hypotheses: (1) Starbuck Island Bidens is an endemic species that evolved from a second colonization of Pacific Islands that has not led to further speciation or adaptive radiation, independent of the colonization that radiated in the Hawaiian Islands and French Polynesia; (2) Starbuck Island Bidens is B. aurea or B. serrulata; (3) Starbuck Island Bidens recently evolved from either B. aurea or B. serrulata. Based on ITS sequences, B. aurea and B. serrulata are rejected as being the same as or even closely related to Starbuck Island Bidens. Surprisingly, Bidens cynapiifolia, a weedy species that occurs from Mexico to Brazil and is widely introduced elsewhere, appears as a very close sister species to Starbuck Island Bidens. However, the two taxa are distinct morphologically. Starbuck Island Bidens is described as a new species, endemic to Starbuck Island. Also revealed in the ITS phylogeny are new sectional and generic relationships. Two species of Cosmos (Asteraceae; Heliantheae), originally included as outgroup species, appear nested within Coreopsis and are most closely related to section Pseudoagarista from Mexico. Bidens mitis (section Meduseae) is nested within section Bidens, indicating that there is no support for the recognition of section Meduseae. Bidens section Psilocarpaea remains polyphyletic, now composed of four distinct clades. Both Bidens laevis (section Bidens) and Bidens squarrosa (section Greenmania) cluster with the other species in their respective sections. My data indicate that section Greenmania is monophyletic and section Bidens is monophyletic if section Meduseae is included in it. It is now known ii that one of the South Pacific Bidens species, Starbuck Island Bidens, is not closely related to the Hawaiian and Marquesan species of Bidens. To fully understand the evolutionary relationships among indigenous Pacific island Bidens a complete ITS phylogeny should be constructed that includes the remaining South Pacific Bidens species not yet sequenced. iii TABLE OF CONTENTS ABSTRACT ii LIST OF TABLES : vi LIST OF FIGURES vii ACKNOWLEDGEMENTS ix Chapter 1 GENERAL INTRODUCTION 1 1.1 Starbuck Island Bidens 1 1.1.1 Possible origin of Starbuck Island Bidens 4 1.1.2 Description of Starbuck Island 4 1.1.3 Flora of Starbuck Island 8 1.1.4 Bidens present on the Line Islands 8 1.2 The genus Bidens 9 1.2.1 General biology and classification 9 1.2.2 Pacific Island Bidens 10 1.3 Characteristics of island taxa 10 1.3.1 Long distance dispersal 10 1.3.2 Evolutionary change in dispersal characters 11 1.3.3 Adaptive radiation and island endemism 12 1.4 ITS phylogeny of Bidens : 12 1.5 Hypotheses 13 Chapter 2 METHODS 16 2.1 ITS sequencing • 16 2.1.1 Specimen sources 16 2.1.2 DNA Extraction and amplification 16 2.1.3 Sequencing reaction 18 2.1.4 Data analysis 18 2.2 Morphological studies 23 2.2.1 Seed source '• 23 2.2.2 Growth environment 23 2.2.3 Traits measured 24 2.2.4 Chromosome counts 24 Chapter 3 RESULTS , 26 3.1 ITS Sequence Phylogeny 26 3.1.1 Species for which ITS sequences were obtained 26 3.1.2 ITS sequence phylogeny of Bidens and Coreopsis 26 3.1.3 Continental sister species of Starbuck Island Bidens 27 3.1.4 A new continental sister species of the Hawaiian and Marquesan Bidens 27 3.1.5 Sectional relationships revealed within Bidens 27 3.1.6 Relationship of Cosmos to Bidens and Coreopsis 31 3.2 Morphology of Starbuck Island Bidens 31 3.2.1 Floral and achene characters 32 3.2.2 Vegetative characters 34 Chapter 4 DISCUSSION 36 4.1 Starbuck Island Bidens: a new endemic Pacific island species 36 4.1.1 Evolutionary origin of Starbuck Island Bidens 36 4.1.2 Possible autopolyploid origin 37 4.1.3 Evidence for evolutionary adaptations to island life 38 4.1.4 The future of Starbuck Island Bidens 39 4.2 Sectional relationships within Bidens 40 iv 4.3 Generic relationships between Bidens, Coreopsis and Cosmos 41 4.4 Implications for future research 41 Chapter 5 TAXONOMIC TREATMENT 43 Chapter 6 REFERENCES 46 APPENDIX 50 Appendix A: Alignment of ITS 1 and ITS 2 sequences used in phylogenetic analyses. Alignment is also available in TreeBASE 50 V LIST OF TABLES Table 1. Information on plant species (Garnett, 1983; IUCN, 1991; Kepler and Kepler, 1994), protection status (Dahl, 1986; IUCN, 1991) and human activity (Garnett, 1983; UNEP/IUCN, 1988) in the Line Islands. Islands are listed in order from north to south 9 Table 2. Specimen sources for Bidens species sequenced. Herbaria are identified by the standard international abbreviation as listed in the Index Herbariorum (Holmgren et al, 1981) 17 Table 3. Sequences and sources used in phylogenetic analysis. Sequences are deposited in the National Center for Biotechnology Information (GenBank) or the National Center for Genome Resources (GSBO). Sequences for newly sequenced species will be available in GenBank July 31, 2002 19 Table 4. Mean, median, minimum and maximum values for floral, achene and vegetative traits of Starbuck Island Bidens 31 vi LIST OF FIGURES Figure 1. Starbuck Island Bidens on Starbuck Island. Photograph used with permission, by A. K. Kepler, 1995 2 Figure 2. Starbuck Island Bidens on Starbuck Island, detail of leaves and flower head. Photograph used with permission, by A. Whistler, 1995 3 Figure 3. Map of the Pacific Ocean and location of Starbuck Island 6 Figure 4. Map of the Line Islands and the location of Starbuck Island within the archipelago. The eleven islands, from north to south, are: Kingman Reef, Palmyra, Teraina (formerly Washington), Tabuaeran (formerly Fanning), Kiritimati (formerly Christmas), Jarvis, Maiden, Starbuck, Caroline (now Millennium), Vostok and Flint 7 Figure 5. Aerial view of Starbuck Island. Photograph used with permission, by A. K. Kepler, 1995 7 Figure 6. ITS phylogeny by Ganders et al. (2000). Percentages represent parsimony bootstrap values, while whole numbers represent the minimum number of base-pair changes required for each step 14 Figure 7. Diagram of internal transcribed spacers (ITS) and the 5.8s gene of ribosomal DNA, including primer locations and orientation 18 Figure 8. One of 10 trees with the highest maximum likelihood score from 44 equally most parsimonious trees for 53 taxa. Tree length 768. Numbers shown are bootstrap percentages from the parsimony analysis. Branch lengths are proportional to the number of substitutions. Clades are identified by capital letters. Underlined species were sequenced in this study 28 Figure 9. Strict consensus of 44 equally parsimonious trees of length 768, for 53 taxa. Clades are identified by capital letters 29 Figure 10. Maximum likelihood tree for 45 taxa with 8 duplicate taxa omitted. The analysis used optimized parameters with 7% of sites invariable, a transition/trans version ratio of 1.80 and a gamma shape parameter of 0.93. Branch lengths are proportional to the estimated number of substitutions. Clades are identified by capital letters 30 Figure 11. Photographs showing floral characteristics of Starbuck Island Bidens. Plants grown at the University of British Columbia from seed collected by A. Whistler on Starbuck Island. (A) Heads, side view. (B) Heads, face view. (C) Ray florets from three plants. (D) Head with rays removed. (E) Ray floret with sterile stamens. (F) Disc florets. (G) Inner involucral bracts. (H) Outer involucral bracts. (I) Chaffy bracts 33 Figure 12. Photograph of Starbuck Island Bidens achenes. Achenes represent variation among three plants grown at the University of British Columbia, from seed collected by A. Whistler on Starbuck Island 34 Figure 13. Photograph of leaves from a single plant of Starbuck Island Bidens. Plant was grown at the University of British Columbia, from seed collected by A. Whistler on Starbuck Island 34 Figure 14. Photograph of leaf variation in Starbuck Island Bidens. Leaves are from fourteen plants grown at the University of British Columbia, from seed collected by A. Whistler on Starbuck Island 35 Figure 15. Bidens cynapiifolia. Illustration adapted from Sheriffs monograph on Bidens (Sheriff, 1937). (a) Portion of fruiting specimen (x 0.85).
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