ARTICLE IN PRESS - UAc et... · ARTICLE IN PRESS Review ... e Department of Ecology and...

ARTICLE IN PRESS

Perspectives in Plant Ecology Evolution and Systematics 12 (2010) 107ndash129

Contents lists available at ScienceDirect

Perspectives in Plant Ecology Evolution and Systematics

1433-83

doi101

n Corr

E-m

journal homepage wwwelsevierdeppees

Review

Conservation of oceanic island floras Present and future global challenges

Juli Caujape-Castells an Alan Tye b Daniel J Crawford c Arnoldo Santos-Guerra d Ann Sakai eKaty Beaver f Wolfram Lobin g FB Vincent Florens hi Monica Moura j Roberto Jardim kIsildo Gomes l Christoph Kueffer m

a Departamento de Biodiversidad Molecular y Banco de ADN Jardın Botanico Canario lsquolsquoViera y Clavijorsquorsquo Ap de correos 14 de Tafira Alta 35017 Las Palmas de Gran Canaria Spainb Secretariat of the Pacific Regional Environment Programme PO Box 240 Apia Samoac Department of Ecology and Evolutionary Biology and the Natural History Museum and Biodiversity Research Center University of Kansas Lawrence KS 66045 USAd Instituto Canario de Investigaciones Agrarias C Retama 2 38400 Puerto de la Cruz Tenerife Canary Islands Spaine Department of Ecology and Evolutionary Biology 321 Steinhaus Hall University of California at Irvine Irvine CA 92697 USAf Plant Conservation Action Group P O Box 392 Victoria Mahe Seychellesg Botanische Garten der Universitat Bonn Meckenheimer Allee 171 D-53115 Bonn Germanyh UMR PVBMT Faculte des Sciences et Technologies Universite de la Reunion 15 avenue Rene Cassin BP 7151 97715 St Denis La Reunion Francei Department of Biosciences University of Mauritius Reduit Mauritiusj CIBIO Centro de Investigac- ~ao em Biodiversidade e Recursos Geneticos CIBIO-Azores Departamento de Biologia Universidade dos Azores Rua M~ae de Deus 58 Apartado 1422

9501-801 Ponta Delgada Portugalk Jardim Botanico da Madeira Caminho do Meio 9064-251 Funchal Portugall Instituto Nacional de Investigac-ao e Desenvolvimento Agrario S ~ao Jorge dos Orga ~os Santiago Cape Verdem Institute of Integrative Biology ndash Plant Ecology Universitatsstrasse 16 ETH Zentrum CHN CH-8092 Zurich Switzerland

a r t i c l e i n f o

Article history

Received 2 March 2009

Received in revised form

6 October 2009

Accepted 8 October 2009

Keywords

World islands

Threat factors

Endangerment

Conservation research

Conservation policies

Global network

19$ - see front matter amp 2009 Rubel Founda

016jppees200910001

esponding author

ail address julicaujapegmailcom (J Caujap

a b s t r a c t

Current threats to the planetrsquos biodiversity are unprecedented and they particularly imperil insular

floras In this investigation we use the threat factors identified by the Millennium Ecosystem

Assessment as the main drivers of biodiversity loss on islands to define and rank 13 current continuing

threats to the plant diversity of nine focal archipelagos where volcanic origin (or in the Seychelles a

prolonged isolation after a continental origin) has produced a high degree of endemicity and fragility in

the face of habitat alteration We also conduct a global endangerment assessment based on the

numbers of insular endemic plants in the endangered (EN) and critically endangered (CR) IUCN

categories for 53 island groups with an estimated 9951 endemic plant species providing a

representative sample of the worldrsquos insular systems and their floristic richness Our analyses indicate

that isolation does not significantly influence endangerment but plant endemics from very small

islands are more often critically endangered We estimate that between 3500 and 6800 of the estimated

70000 insular endemic plant species worldwide might be highly threatened (CR+EN) and between ca

2000 and 2800 of them in critical danger of extinction (CR) Based on these analyses and on a

worldwide literature review of the biological threat factors considered we identify challenging

questions for conservation research asking (i) what are the most urgent priorities for the conservation

of insular species and floras and (ii) with the knowledge and assets available how can we improve the

impact of conservation science and practice on the preservation of island biodiversity Our analysis

indicates that the synergistic action of many threat factors can induce major ecological disturbances

leading to multiple extinctions We review weaknesses and strengths in conservation research and

management in the nine focal archipelagos and highlight the urgent need for conservation scientists to

share knowledge and expertise identify and discuss common challenges and formulate multi-

disciplinary conservation objectives for insular plant endemics worldwide To our knowledge this is the

most up-to-date and comprehensive survey yet to review the threat factors to native plants on oceanic

islands and define priority research questions

amp 2009 Rubel Foundation ETH Zurich Published by Elsevier GmbH All rights reserved

tion ETH Zurich Published by Elsevier GmbH All rights reserved

e-Castells)

ARTICLE IN PRESS

J Caujape-Castells et al Perspectives in Plant Ecology Evolution and Systematics 12 (2010) 107ndash129108

Contents

Introduction 108

Methods 109

Focal study system 109

Expert survey 110

Endangerment analysis of the worldrsquos insular plant endemics 111

Results and discussion 113

Documentation of plant diversity on oceanic islands status and future prospects 113

The magnitude of plant diversity endangerment on the worldrsquos islands 114

A network of threat factors affecting island plants 114

Small population sizes and fragmentation 114

Lost mutualisms 115

Habitat alteration and destruction 116

Invasive alien plant species 116

Invasive alien invertebrates and pathogens 117

Invasive alien vertebrates 118

Climate change and pollution 118

Need for a better knowledge base 119

First aid measures 119

Conclusions 120

Acknowledgements 121

Appendix 1 121

Appendix 2 121

References 124

Islands are an enormously important source of informationand an unparalleled testing ground for various scientifictheories But this very importance imposes an obligation onus Their biota is vulnerable and precious We must protect itWe have an obligation to hand over these unique faunas andfloras with a minimum of loss from generation to generation

Ernst Mayr (1967)

Introduction

Islands are of particular importance for the conservationof global plant diversity Although they make up only some 5of the Earthrsquos land surface about one quarter of all knownextant vascular plant species are endemic to islands (Kreft et al2008) Indices of vascular plant diversity are markedly higherfor islands than for continental areas (Kier et al 2009) and 20 ofthe 34 biodiversity hotspots defined by Conservation Interna-tional (Myers et al 2000 and updates in httpwwwbiodiversityhotspotsorg) are islands or have an important insular compo-nent In addition to this legacy of a unique evolutionary historyinsular ecosystems are also key to the livelihood economy well-being and cultural identity of 600 million islanders roughly one-tenth of todayrsquos world population (Lutchman et al 2005)

Humans have heavily affected island ecosystems (MillenniumEcosystem Assessment [MEA] 2005 Whittaker and Fernandez-Palacios 2007 Kingsford et al 2009) For instance of some 80documented plant extinctions in the last 400 years about 50 wereisland species (Sax and Gaines 2008) In the near future humanpressure on ecosystems will likely increase more markedly onislands than on continents (Brooks et al 2002 MillenniumEcosystem Assessment [MEA] 2005 Kier et al 2009) The smallpopulation sizes and ranges of island organisms and their uniquecharacteristics resulting from prolonged evolutionary isolationmake them particularly sensitive to anthropogenic change(Frankham 1998) All of these features make islands invaluablebut fragile and vulnerable arks of biodiversity

Island plant conservationists are thus faced with a huge globalconservation challenge Nevertheless contact among researchersand managers of threatened plants on different archipelagos hasbeen limited (for example in Macaronesia Caujape-Castells et al

2006 2007a b) although some local or regional surveys ofendangerment patterns on different oceanic archipelagos have beencarried out (Medail and Quezel 1997 Jaffre et al 1998 Broughtonand McAdam 2002 Sakai et al 2002 Kingston and Waldren 2005Strasberg et al 2005 Danton and Perrier 2006 Maunder et al2008 Reyes-Betancort et al 2008) Wider discussions of conserva-tion issues common to different island systems have happenedsporadically but researchers in different regions have rarelysynthesized their conclusions and global comparisons are lacking

We present here what we believe is the first comprehensivesynthesis of plant conservation issues across major oceanicarchipelagos For nine focal island groups (Azores MadeiraCanary Islands Cape Verde Hawaii Galapagos Juan FernandezMascarenes Seychelles) we review the threats to native floras andthe present state of conservation research and practice The ninearchipelagos belong to eight countries and three oceans AtlanticPacific and Indian We complement this with a literature reviewand an analysis of patterns of endangerment covering 62 additionalisland groups providing quantitative estimates of threats to islandfloras worldwide We compare across archipelagos the patterns ofendangerment threat factors and research results that have beensuccessfully incorporated into management practices We reviewthe present resources and weaknesses in particular archipelagosidentify individual and shared knowledge gaps and use these tosuggest priorities for future research We suggest how existingscientific knowledge may be used to identify additional knowledgegaps launch priority studies to generate the necessary informationand develop strategies to preserve island floras We also discussresults and experiences that extend beyond the local context Webelieve that beyond its local relevance plant conservation researchon oceanic islands can also serve as a model for continents and shedlight on general topics in ecology (Kueffer et al 2010 Kaiser-Bunbury et al 2010)

In particular we address the following questions

What is the level of endangerment of oceanic island vascularplants and how does it differ between island groups

What are the main threats to endemic vascular plants on

oceanic islands and how does the relevance of these factorsvary among island groups

ARTICLE IN PRESS

J Caujape-Castells et al Perspectives in Plant Ecology Evolution and Systematics 12 (2010) 107ndash129 109

FiglsquolsquoMe

end

TabPhy

A

A

C

C

G

H

Ju

M

M

Se

An

(ie

Di

tha

isla

arch

em

som

What is known about the underlying mechanisms throughwhich the different threat factors affect island plant diversity

What future research and management are needed to conserve

island plant diversity

Methods

Focal study system

A sample of nine tropical and sub-tropical oceanic archipelagoswas selected for analysis Galapagos Hawaii and Juan Fernandez

Hawaii

Azores

Cape Verde

Madeira amp SelvagensCanaries

Galaacutepagos

Juan Fernaacutendez0

5

10Madeira amp Selvagens

Canaries

Cape VerdeGalaacutepagos

Hawaii

-10

-5

-15 0 5 10 15

F2 (2

162

)

Azores

MascarenesSeychelles

Juan Fernaacutendez

-10 -5F1 (3593 )

1 Geographical location of the nine focal archipelagos and their position in the

thodsrsquorsquo) This representation explains 5755 of the detected inter-archipelago vari

emic plant diversity

le 1sico-geographical features of the island systems considered and indicators of prote

rchipelago NI dm di Di A nat pro

zores 9 1343 6 615 2332 ca 22a ca 20

anaries 7 95 5 485 7545 ca 15 ca 40

ape Verde 9 576 ca 15 279 4033 2 02c

alapagos 14 1000 4 141 7900 2e 96

awaii 8f 3900 ca 11 128 16636 ca 27 ca 5g

an Fernandez 3 590 1 150 100 1 ca 90

adeira amp Selvagens 2 630 57 57 794 5 ca 67

ascarenes 3 665i 164 740 4528 21 25j

ychelles 4 9930i 45 50 235 7 16

island is defined as any land mass isolated by sea from other land masses at all stages

other than mangroves etc) NI number of main islands dm minimum distance t

maximum distance between two main islands (in Km) A total land area (in km2) n

t is protected by law Ma maximum estimated island age of the present main island

nds (upper estimate in my) h maximum height (in m) C number of censed inha

ipelago in 2007 $ main economic activities of the islands in order of importanc

ployment I industry (including construction and wine industry) M manufactures

a Apart of the 13 existing Nature preserves each of the 9 islands will have a singleb This figure corresponds to stays in traditional hotelsc World resources institute httpearthtrendswriorgtextbiodiversity-protectedcd Data communicated by J Richard Wilson (jrwgeoaudk) the island of Maio hase These occupy 96 of the land area (Galapagos National Park) and the entire sea af 10 smaller land masses included in the total of 124 small islands reefs and shoag This is the estimate of the area really protected and managed for biodiversity (oth

e are protected because they are on military land so access is limited some are mh There are islands no longer above sea level that date back to about 647 Ma (seei To Madagascarj The exact area of the national park in Reunion may change as an area may be exk This is the probable date of separation of Seychelles and India ie complete isolationl Age of youngest main island Silhouette

in the Pacific Ocean the Mascarenes and the granitic islands of theinner Seychelles in the Western Indian Ocean and Madeira (plusthe Selvagens) Azores Canary Islands and Cape Verde inMacaronesia (Atlantic Ocean) This represents a broad geographicsample (Fig 1) encompassing 59 main islands and ca 268 minorislands and ranging widely in physical geographic floristic andsocioeconomic characteristics (Tables 1 and 2) The main islandsof the Seychelles are formed of granitic bedrock of continentalorigin while all other focal islands are true oceanic islands ofvolcanic origin Total endemic vascular plant species richness canbe divided into three size classes fewer than 100 species (AzoresCape Verde Seychelles) 100ndash200 species (Galapagos Juan

Mascarenes

Seychelles

multivariate space defined by the two first axes of the PCA analysis (see section

ability in the impact of the 13 threat factors considered on the decline of insular

ction of the endemic flora

Ma ma h C d T $

8 025 2531 243018 (2006) ca 105 858832b TIA

206 112 3718 2025951 (2008) ca 269 9326116 TIA

76d 32d 2829 426998 (2008) ca 102 312880 TA

4 007 1700 ca 25000 (2006) ca 3 ca150000 TCA

51h 043 4205 1283388 (2007) ca 77 7627819 TAMi

58 10 1320 600 (2005) ca 6 ca 3000 ATG

18 5 1861 246689 (2007) ca 307 988692 TIA

10 3 3070 2061546 (2008) ca 455 1342839 TSM

65k 60l 915 81000 (2002) ca 345 ca 161000 TA

of the annual tidal cycle and able to support at least one species of terrestrial plant

o the mainland (in Km) di minimum distance between two main islands (in km)

at number of nature reserves and national parks pro percentage of the land area

s (upper estimate in my) ma minimum estimated island age of the present main

bitants (census year) d population density T number of tourists that visited the

e [codes are A agriculture livestock and fishery C conservation G government

Mi military S sugar production T tourism and services]

natural park encompassing the areas of relevancy for conservation

ountry-profile-34html

geological formations from the Jurassic (ca 160 mya)

rea around the archipelago (Galapagos Marine Reserve)

ls (see httpwwwsoesthawaiieduGGHCVhaw_volchtml)

er areas are protected in the state natural reserve area but with no enforcement

anaged by The Nature Conservancy)

httpwwwsoesthawaiieduGGHCVhaw_formationhtml)

cised from it to enable geothermal prospecting

of the Seychelles islands The bedrock granite of the islands is up to 750 Ma old

ARTICLE IN PRESS


Fernandez and Madeira-Selvagens) and more than 600 species(Hawaii the Mascarenes and the Canaries) If we consider numberof endemic species per km2 Juan Fernandez (133) is by far themost diverse system followed by Seychelles (029) Madeira-Selvagens (017) the Mascarenes (015) Canaries (008) Hawaii(006) Azores (003) Galapagos (0022) and Cape Verde (0020)

From the vast area of Oceania we included only Hawaii amongour focal archipelagos as the information available for otherarchipelagos from this region is scarce at best ConservationInternational identified plants from Polynesia-Micronesia as a keyinformation gap for red listing (httpwwwconservationorgexplorepriority_areashotspotsasia-pacificPolynesia-MicronesiaPagesconservationaspx) and a recent red-list review for Oceania(Pippard 2009) shows that only ca 5 of Dicots and 1 ofMonocots have been assessed for the IUCN red list However

Table 2Indicators of botanical richness of the nine focal archipelagos and main geographical

Archipelago G sp Isp end red

Azores 1 (1) 72a 5 ca 7 b

Canaries 23 (49) ca 607c ca399 ca 45 ca 30

Cape Verde 1 (7) 82d 30 34 ca 43

Galapagos 7 (35) 180e 40 ca 35 ca 60

Hawaii 32 (288) 929f 624 90gh ca 53gh

Juan Fernandez 12 (33) 133i 101 ca 94 ca 75

Madeira amp Selvagens 5 (11) 136j 94 ca 13 49

Mascarenes 35(81) ca 688gk 547 72 ca 50

Seychelles 12 (13)g ca 70l ca 14g ca 40 71

Total 129 (518) ca 2897 ca 1854 ca 48 ca 54

G number of endemic genera (species) sp total endemic species Isp number of end

red proportion of endemic taxa that are in the red list under some kind of threa

abbreviations are A Africa Am America MED Mediterranean NW new world

a Silva et al (2005a) and Monica Moura (unpubl data)b Although the Azores currently lack a Red List information on anthropogenic threa

needs for the 90 most endangered plants in this archipelago will be available with

Conservationrsquorsquoc Aceves-Ginoves et al (2004) and Arnoldo Santos-Guerra (unpubl data)d Sanchez-Pinto et al (2005) and Wolfram Lobin (unpubl data)e Wiggins and Porter (1971) and Alan Tye (unpubl data)f Wagner et al (2005) and Ann Sakai (unpubl data)g Data for flowering plants onlyh See Sakai et al (2002)i Marticorena et al (1998) and Daniel J Crawford (unpubl data)j Jardim and Sequeira (2008)k httpwwwplantmascorgl Kueffer et al (2007a) and Katy Beaver (unpubl data)

Table 3Ranks (numbered in decreasing order of importance ties allowed) and relevances (A

threat C priority threat factor a potential major driver of plant diversity loss D data

focal archipelagos

Threats AZO CAN CA

Biological

Small population sizes and fragmentation 1C 6B 4CLost mutualisms 11A 12D 5A

Habitat alteration and destruction 1C 4C 1CInvasive alien plant species 1C 2C 8CInvasive alien invertebrates and pathogens 12A 7B 12

Invasive alien vertebrates 10B 1C 13

Climate change and pollution 13A 13D 7A

Socio-economic

Demographic and economic growth 5C 3C 2B

Tourism 9B 5C 6B

Lack of laws or enforcement 6C 10A 11

Poor education and awareness 8B 9B 9B

Overexploitation 4C 8C 3CLack of natural resource management capacity 6C 11C 10

Shadowed cells signal priority threat factors irrespective of their ranked importance

a A grazers D rodents

many of the Oceania archipelagos are included among ourbroader sample of 62 island groups Our focal archipelagos alsodo not include the Caribbean but plant conservation in thatregion is well covered in a recent review by Maunder et al (2008)and many Caribbean islands are included in our broader sample

Expert survey

A questionnaire on floristic geographic and socioeconomicaspects of the nine focal archipelagos was completed by theparticipating authors with help from others knowledgeable abouttheir archipelagos (see Acknowledgements) and the responseswere used to construct Tables 1 and 2 The contributing expertsthen developed a conceptual framework of 13 threat factors on

links of their floras (in order of importance)

Ext Major geographical links of the present endemic flora

1 Western Europe Canaries-Madeira N-A

2 MED Saharan W-A E-A Arabian peninsula S-A NW

3 MED Canaries-Madeira NW-A

3 S-Am coast Andes Caribbean SW-North America

ca 100g Pacific widespread neotropical N-temperate Austro-Malaysia

o6 S-Am Pantropical New Zealand

1 MED

ca 40 AfricaMadagascar Indo-Malaysia

2oExto5 AfricaMadagascar Indo-Malaysia

ca 160

emic species exclusive from a single island end proportion of plant endemicity

t (CRENVU) Ext reported historical extinctions of plants Geographical linksrsquo

ts natural risks biological limitations conservation actions proposed and research

the impending publication of the book lsquolsquoAc-orean Vascular Flora Priorities in

not present or of low relevance B relevant factor but not considered a priority

deficient) of the 13 threats to the endemic plant biodiversity assessed in the nine

P GAL HAW JUA MAD MAS SEY

9B 5C 4C 9B 6C 1C12D 10B 6B 11D 11A 10D

3B 4C 3C 4C 3C 4B

2C 1C 2C 3C 1C 2B

B 7C 1C 10D 6B 10A 3B

A 1C 1C 1C 5C 2C 9ADa

12D 6C 12A 8D 12A 11D

3C 11C 13A 1C 7C 5B

11B 11B 7B 2C 4C 7B

B 3B 7C 5B 13A 8B 6B

10C 9C 9B 12A 9B 9B

3B 8C 11B 7B 13A 8B

C 8C 11C 8B 10B 5C 6B

Abbreviations are the first three letters of each focal archipelagorsquos name

ARTICLE IN PRESS


the nine archipelagos (Table 3) based on the five threatsidentified by the Millennium Ecosystem Assessment [MEA](2005) as the main drivers of biodiversity decline on islandsnamely habitat change climate change invasive alien speciesover-exploitation and pollution For more detailed analysis of thethreats considered important on many of these archipelagoslsquohabitat changersquo and lsquoover-exploitationrsquo were represented in thisassessment by Tourism impact Habitat alteration and destruction(including land use change) Demographic and economic growthOverexploitation and Small population sizes and fragmentationSimilarly lsquoinvasive alien speciesrsquo was split into Invasive alienplants Invasive alien vertebrates and Invasive alien invertebratesand pathogens (see Table 3) Climate change and pollution wereassessed jointly as their predicted effects on plant biodiversity donot lend themselves to easy quantification We also consideredthreats related to socio-economic factors not highlighted by theMEA The ranks of the 13 resulting threat factors (Table 3) werestandardized and used to carry out a principal componentanalysis (PCA Fig 1) based on the matrix of pairwise variancesndashcovariances among them

Though the MEA assesses both past and future threats ourrankings were based only on the current continuing impacts ofthe 13 factors considered not on their influence in the past whentheir relative importance often differed from what it is today Wealso considered only their direct impacts on plant biodiversityrather than indirect impacts via other threat factors So for

POOR EDUCATIONAND AWARENESS

TOURISM

LACK OFMAN

DEMOGRAPHIC amp ECONOMIC GROWTH

INVASIVE ALIEN

VERTEBRATES

SMALL

AND

HABITATALTERATION

INVASIVE ALIEN

DECLINE IN ENDEMIC

INVERTEBRATESAND PATHOGENS

Fig 2 Conceptual representation of some of the major interactions among the 13 th

endemic diversity For clarity not all possible relationships feedbacks or multiple lev

direction(s) of relationships that exacerbate related factors for instance poor educatio

enforcement thereof and could also lead to an increase in demographic growth Two-h

lsquoClimate change and pollutionrsquo because the interactions between this factor and the othe

text) Box fillings indicate factors that were identified as major drivers of plant diversity

of them (light gray filling) and in one or none of them (no filling)

example tourism is ranked in terms of its direct impacts on plantcommunities or populations of rare plants such as visitorpressure or the construction of tourist installations and demo-graphic growth is ranked on the direct impacts of increasingnumbers of residents and their required infrastructure Theexperts then assessed the current or short-term relevance of eachthreat factor based on four categories (see Table 3) and aconceptual representation of the most important interactionsamong the threat factors (Fig 2) was drawn to complement thePCA The main weaknesses and strengths of conservation researchon each focal archipelago were summarized (Appendix 1) to helpidentify future priorities for local conservation action and globalcollaboration

Endangerment analysis of the worldrsquos insular plant endemics

To provide quantitative information on patterns of endanger-ment in insular systems of the world the IUCN red list database(wwwredlistorg) was searched for endangered (EN) and criticallyendangered (CR) island plant species and subspecies After siftingthe records for 62 island groups assigning each taxon to itsislands of distribution and eliminating nine outliers we remainedwith 53 island groups (see Appendix 2) Of the nine outliersAntigua and Barbuda Barbados Montserrat Saint Kitts and Nevisand the Virgin Islands were excluded because of their very lownumber of endemic species andor of inconsistencies between the

LACK OF LAWS ORENFORCEMENT

NATURAL RESOURCESAGEMENT CAPACITY OVEREXPLOITATION

INVASIVEALIEN

PLANTSCLIMATE CHANGEAND POLLUTION

POPULATION SIZES

FRAGMENTATION

LOSTMUTUALISMS

PLANT DIVERSITY

reat factors considered in this paper and of their impact on the decline of plant

el interactions among factors are shown Arrows indicate only the predominant

n could result in a decrease of the effectiveness of laws to protect biodiversity or

eaded arrows are connected by dashed lines We refrained from linking arrows to

rs are probably manifold but not yet well known and thus difficult to quantify (see

loss in 5 or more of the focal archipelagos (dark gray filling) in between 2 and four

ARTICLE IN PRESS


data in the red list and the estimated number of endemic species(ie more CR or EN species than endemics described in thecorresponding literature source) Madagascar Borneo Papua NewGuinea and New Caledonia were excluded because the extremelyhigh numbers of endemics on these islands of continental originfall clearly outside the overall pattern of distribution Correlationanalyses were run between the proportions of EN and CR taxa inthe total endemic flora in each of the remaining 53 archipelagos

35302520151050

EN

25

20

15

10

5

0

C

R

1 2 3LOG1

1 2 3LOG1

Fig 3 Proportions of EN and CR taxa with respect to total endemic species in the 5

archipelago land area (in km2)

Table 4Averages and KruskalndashWallis test significances in five factors tested for the number of

species (CR) and proportion of endangered species (EN) in the 53 islandsisland group

Categories tested Endemicsarea

Island size

Very small [16] 0181

Small [23] 0056

Large [14] 0011

(o00001)nnn

Isolation Index

5 o II o 80 [40] 0052

II 4 80 [13] 0173

(o00001)nnn

Geographic region

Pacific Ocean [17] 0123

Atlantic Ocean [21] 0081

Indian Ocean [10] 0049

Mediterranean Sea [5] 0013

(0278)

Atlantic Ocean

East Atlantic [6] 0150

Caribbean [15] 0053

(0062)

Pacific Ocean

Asia amp Oceania [13] 0058

Eastern Pacific Ocean islands [4] 0334

(0089)

Island size designations are lsquolsquovery smallrsquorsquo areao1000 km2 lsquolsquosmallrsquorsquo 1000 km2oareao(II) correspond to more remote islands (see Appendix 2 for explanation) Numbers in b

each test Numbers in parentheses under the averages indicate the respective P values

nnnPo0001 nn Po 001 n Po005

and the log10 of the archipelagorsquos land area in km2 (Appendix 2Fig 3) We calculated the proportions of lsquolsquohighly threatenedrsquorsquo(CR+EN) and critically endangered (CR) taxa in our sample by(i) dividing the total number of species and subspecies in the(CR+EN) and CR IUCN categories by the total 9951 endemicspecies and (ii) calculating the proportion of (CR+EN) and CR taxain each archipelago relative to the estimated number of endemicspecies in that archipelago and then averaging these proportions

4 5 60AREA

4 5 60AREA

3 archipelagos included in the global statistical analysis plotted against log10 of

endemic vascular plant species per unit area proportion of critically endangered

s analyzed

CR EN CR+EN

4741 11212 15953

4778 4452 9230

1660 1593 3254

(0181) (0044)n (0002)nn

2828 5326 8155

7374 7003 14377

(0063) (0263) (0102)

4135 4392 8527

2772 9240 12012

5507 3486 8993

5083 0106 5189

(0044)n (0002)nn (0831)

3382 3826 7244

2528 11392 13919

(0866) (0062) (0161)

1732 3773 5505

11943 6405 18348

(0006)nn (0212) (0009)nn

15000 km2 and lsquolsquolargersquorsquo area415000 km2 Higher values of the isolation index

rackets after the sub-category names are the numbers of island groups included in

for each variable and category tested

ARTICLE IN PRESS


for the 53 archipelagos Subsequently these proportions wereused to estimate how many of the 70000 plant species thought tobe endemic to the worldrsquos islands (Kreft et al 2008) could behighly threatened (CR+EN) or critically endangered (CR) Finallywe tested the influence of five factors related to island sizeisolation and geographic location on the numbers of endemic taxaper unit area and of the numbers of taxa in the EN and CR IUCNcategories using KruskalndashWallis tests (Table 4) All statisticalanalyses were carried out with XLSTAT 2009101 (Addinsoft2009)

Results and discussion

We first discuss the limitations of plant diversity data onoceanic islands then present the global endangerment analysisfollowed by the threat analysis of the nine focal archipelagos Ourthreat analysis is based on taxa assessed as threatened by theIUCN red-list process so species not assessed by red-list criteriaare not included However focal archipelagos were chosen on thebasis that they have a high proportion of their endemic vascularplant taxa assessed so the main gap is consideration of widelydistributed threatened plant taxa Although some of these may beglobally threatened and should be part of conservation strategiesthey cannot be readily included in this type of analysis Howeverwe consider that an archipelago comparison based on endemicsreveals adequately patterns of threat endangerment and con-servation requirements

Because the threat factors assessed may act differently inarchipelagos not included in the focal group a literature review ofthe impacts of the biological threat factors across archipelagosworldwide complements the above-detailed analysis allowing usto generalize findings to a global scale Finally we highlight somemajor implications of our assessments Important socioeconomicand cultural factors such as differences among archipelagos inthe per capita incomes or in education and awareness are notcomprehensively treated as we focus our analyses on thebiological factors However underlying socioeconomic factorsare included among the threat factors considered and some ofthem are highlighted as priorities to be addressed in many of thefocal archipelagos (Appendix 1) as well as for others included inour broader sample

Documentation of plant diversity on oceanic islands status and

future prospects

Because successful conservation requires reliable identificationand population estimates understanding the state of our knowl-edge of these two factors is critical

Taxonomic knowledge varies sharply between archipelagosPublished floras or checklists exist for most oceanic islandsincluding most of our nine focal archipelagos (eg Wiggins andPorter 1971 Lawesson et al 1987 Acebes-Ginoves et al 2004Silva et al 2005a Sanchez-Pinto et al 2005 Jardim and Sequeira2008) Some are accessible on the Web (eg Hawaii [httpbotanysiedupacificislandbiodiversityhawaiianflora] the Mar-quesas [httpbotanysiedupacificislandbiodiversitymarquesasfloraindexhtm] the Cook Islands [httpcookislandsbishopmuseumorgsearchasp] French Polynesia [httpwwwherbier-tahitipf] La Reunion [httpflorecbnmorg] the West Indies [httppersoonsieduantilleswestindiesindexhtm] the Greater Antil-les [httpwwwnybgorgbscifga]) However for very fewislands have all taxonomic groups been treated thoroughly

DNA sequence data and other non-morphological traits may becritical for producing more objective classifications This isespecially important for endangered populations whose taxo-

nomic rank may affect their legal conservation status Molecularpopulation genetic or phylogenetic investigations are contributingto more refined insights into the population biology andsystematics of endemics in several archipelagos (eg Crawfordet al 1987 Baldwin et al 1991 Francisco-Ortega et al 1996Carine et al 2004 Nuez et al 2004 Ruiz et al 2004 Mooreet al 2006 Archibald et al 2006 Oliva-Tejera et al 2006 Dıaz-Perez et al 2008 Maunder et al 2008 Andrus et al 2009Gonzalez-Perez et al 2009) Currently a sample of the endemicflora of Garajonay National Park (La Gomera Canaries) is beingused to test the potential usefulness of cpDNA sequences incomplementing morphological variables for the identification ofthe whole Canarian Flora (Caujape-Castells et al 2007a b)

Resolution of taxonomic and floristic uncertainties could leadto reductions in the number of endemics on some archipelagoswhereas increased exploration could reveal more Many archipe-lagos have highly dissected or unstable terrain that generates thedistinctive evolutionary patterns seen in their plant endemics butwhich is also a powerful impediment to botanical explorationConsequently large gaps in knowledge of the status anddistribution of endemic plants remain in most archipelagos(Maunder et al 2008 this study) Species thought to be extinctor new populations of rare species continue to be discovered onoceanic islands For example in Galapagos populations of Linum

cratericola (Linaceae) and of both varieties of Scalesia atractyloides

(Asteraceae) were rediscovered after they had been presumedextinct (Mauchamp 1996 Mauchamp et al 1998 Tye 19972002) In Cape Verde three botanical expeditions organised in2006 and 2007 by the Jardın Botanico Canario lsquolsquoViera y Clavijorsquorsquo(JBCVC) to the islands of Santo Anta ~o Sa ~o Nicolau Fogo andSantiago found many previously undocumented populationsof the dragon tree Dracaena draco (Dracaenaceae) which hadbeen considered scarce [Marrero-Rodrıguez and Almeida unpubldata] Since 2000 the botanic garden in La Reunion hasimplemented a specific method to facilitate the rediscovery ofspecies including the production of information leaflets on theprioritized species (Boullet et al 2006 Baret et al 2007) So farseven plant species have been rediscovered in the wild thanks tothis approach including Nesogenes orerensis (Orobanchaceae)rediscovered after 150 years (Baret et al 2006) and the onlyknown wild population of an endemic palm discovered next tohabitation in a coastal site (Lavergne et al 2004)

Other than endemics the status (alien or native) of manyplants on oceanic islands is often unclear (eg Tye 2006a)although palaeobotanical research may be able to clarify statusAn illustrative example is the recent use of pollen records toconfirm as native in the Galapagos six plant species thatwere formerly believed or suspected to be alien species there(Van Leeuwen et al 2008)

Among the nine focal archipelagos there are large differencesbetween the numbers of known recent extinctions documentedIn the Galapagos and Juan Fernandez there is reasonable certaintythat not many anthropogenic extinctions have occurred whileHawaii and the Mascarenes have experienced far more than theothers (Table 2) Further in Hawaii the Canaries Madeira-Selvagens and the Seychelles there may have been manyanthropogenic extinctions before scientific study began as onmost oceanic islands that were settled by humans in earlyprehistory (eg Whittaker and Fernandez-Palacios 2007) Forinstance pollen records show that the legume shrub Kanaloa

kahoolawensis was a dominant species of lowland habitats inHawaii before ca 1550 AD but it was unknown to science untilits discovery in 1992 (Lorence and Wood 1994)

The inception of DNA sequencing has enabled a wealth ofstudies on the origins and post-colonisation history of island plantgroups (eg Baldwin 1992 Moore et al 2006 Dillon et al 2007

ARTICLE IN PRESS


Andrus et al 2009 Jaen-Molina et al 2009) DNA sequences areuseful in detecting phylogenetic patterns important in biodiver-sity conservation planning eg speciose clades such as radiatingcrown groups or species-depauperate sister groups that may beisolated and in danger of extinction DNA sequences also may beused to estimate phylogenetic diversity a quantitative measure ofbiodiversity based on phylogeny (Faith 1992 Faith et al 2004Forest et al 2007)

The magnitude of plant diversity endangerment on the worldrsquos

islands

Data were retrieved from the IUCN red list on 657 endangered(EN) and 739 critically endangered (CR) species and subspeciesrespectively representing 94 and 107 vascular plant families inthe 62 islands and island groups listed in Appendix 2 After theremoval of nine outliers as described above a sample of 53 islandgroups contained an estimated 9951 endemic species andsubspecies There are several reasons to exercise caution in theinterpretation of these data First the IUCN database mayunderestimate the numbers of taxa in the two categoriesconsidered roughly 82 of the EN and 74 of the CR taxaincluded in the global analysis correspond to assessments made in1998 and need updating Further the red-list criteria may notreadily detect changes from naturally small sizes and ranges tocritical states (Martın 2009) For instance the proportion ofCR+EN endemics is higher on the focal archipelagos that have redlists (30ndash75 Table 2) than on the 53 island groups included inthe global analysis (average 28) Second red-list peer review isnot yet implemented for many archipelagos (see httpwwwiucnredlistorgstaticinfo_sources_quality) so the comparisonsare between assessments that may not be as standardized as theyshould be Finally there might be a bias towards higherproportions of threatened taxa for insular regions where con-servation research has been more intense

With these considerations in mind our analysis showedthat the proportions of EN and CR taxa were negatively correlatedwith island area (Fig 3) with stronger correlations for EN(r=0426 P=0001) than for CR (r=0289 P=0036) Bothlsquolsquovery smallrsquorsquo and lsquolsquosmallrsquorsquo islands have significantly highernumbers of endemics per unit area and higher proportions ofEN and (EN+CR) endemics than lsquolsquolargersquorsquo islands (Table 4) althoughthe differences among the three island size groups for CR taxaalone were not significant In our sample ca 25 of the endemictaxa from very small and small islands are highly threatened(either CR or EN) whereas this proportion is much lower in largeislands (ca 3) This indicates higher threats to endemic plantdiversity on smaller islands

Although isolated island groups (isolation index 480 seeTable 4) have more than three times as many endemics per unitarea than islands closer to continents or to other island groups(0173 vs 0052 Po00001) isolation did not influence endan-germent significantly (Table 4)

The islands in the Indian Ocean and the Mediterranean Sea hadsignificantly higher average proportions of CR taxa than those inthe Pacific and the Atlantic (Table 4) however the averageproportion of EN taxa is significantly higher in the Atlantic Oceanthan in the remaining regions with the lowest value in theMediterranean Whereas there are no significant differences atany level examined between islands in the Caribbean and the EastAtlantic both endemics per unit area and the proportion of CRtaxa are significantly lower in the islands of AsiaOceania than inthe islands of the eastern Pacific

In our survey species and subspecies in the (CR+EN)and CR categories account for 507 and 287 respectivelyof the total endemic species and the average proportions

of (CR+EN) and (CR) taxa among the total endemic speciesper archipelago are respectively 968 and 394 Thus of theestimated 70000 plant species endemic to the worldrsquos islands(Kreft et al 2008) between ca 3500 and 6800 insular endemicplant species worldwide might be highly threatened (EN+CR)of which between ca 2000 and 2800 could be in critical danger ofextinction (CR)

A network of threat factors affecting island plants

The PCA representation in Fig 1 describes the relationshipsamong the focal archipelagos in the multivariate space definedby the 13 threat factor rankings (Table 3) and shows aconspicuous lack of correspondence with the geographicalposition of these archipelagos The threat factors that were mostcommonly regarded as major drivers of plant diversity loss(category lsquolsquoCrsquorsquo in Table 3) were lsquoHabitat alteration and destructionrsquoand lsquoInvasive alien plant speciesrsquo in eight archipelagos lsquoSmallpopulation sizes and fragmentationrsquo lsquoDemographic and economicgrowthrsquo lsquoLack of natural resource management capacityrsquo andlsquoInvasive alien vertebratesrsquo were regarded as major drivers in sixarchipelagos lsquoLost mutualismsrsquo and lsquoClimate change and pollu-tionrsquo were mostly assessed as either data deficient or of lowrelevance (categories lsquoDrsquo and lsquoArsquo respectively in Table 3) Theliterature review however indicated that these factors mightalready be or could soon become major threat factors high-lighting either important knowledge gaps or the emergence ofnew threat factors

Some of the threat factors have both direct and indirecteffects while some act primarily as ultimate causes that workthrough other more proximate factors The most importantinteractions between threat factors reveal a semi-hierarchicalarrangement (Fig 2) As an example from Galapagos threatfactor 3 (tourism) is the main driver of the economy and therebythe main cause of threat 4 (human demographic growth)which in turn is the underlying cause behind the increasingnumber of invasive plants (threat 1) and invertebrate pests (7)while threat 9 (poor education about the gravity of threats tobiodiversity) may be seen as a root cause contributing to thepoorly planned development of the economy including tourismSince tourism works primarily through other factors and its directimpacts are relatively limited it is ranked fairly low in importanceon most of the nine focal archipelagos However this does notreflect its underlying importance in contributing to overall lossof biodiversity by working through the other factors consideredhere

Small population sizes and fragmentation

Many plants on oceanic islands have naturally small popula-tions restricted to small geographic areas while others may havebeen brought to this state through prehistoric (eg palms on manyoceanic islands Prebble and Dowe 2008) or more recentanthropogenic impact As highlighted for Hawaii (Sakai et al2002) extinction risk is strongly associated with limited geo-graphic distribution at several scales Taxa endemic to only onearchipelago are at greater risk than taxa with extra-archipelagoranges and single-island endemics are more at risk than multi-island endemics with the smallest islands having the highestproportions of endemic taxa at risk Our analyses (Table 4) showthat these patterns are valid globally

In the three focal archipelagos with the highest endemicity(Table 2) more than 60 of the endemics are single-islandendemics (672 in Hawaii 795 in the Mascarenes and 657 inthe Canaries) and the proportions are similar in Juan Fernandez

ARTICLE IN PRESS


and Madeira-Selvagens (759 and 691 respectively) TheSeychelles (200) the Galapagos (222) Cape Verde (366)and Azores (69) have lower proportions of single islandendemics Overall these data show that most plant endemics inthe focal archipelagos with the richest plant biodiversity havelimited distributions and are likely at higher risk than thosewhere most endemics are more widespread (eg the AzoresCarine and Schaefer 2009)

Rarity may make species vulnerable to extinction for threemain reasons environmental stochasticity demographic stochas-ticity and low genetic variability (Soule and Wilcox 1980Frankel and Soule 1981) Plants restricted to very small areasmay be at higher risk of extinction due to natural catastrophessuch as fires hurricanes or diseases (eg Kingston and Waldren2005) Little is known about the demography of oceanic islandplants relative to continental species but many are slow growing(eg Pattison et al 1998 Schumacher et al 2008 2009) and longlived (eg Carlquist 1974 Hart 2008) For instance many largeindividuals of the dominant tree Metrosideros polymorpha in aHawaiian wet forest were estimated to be 400 years or more ofage with the oldest ones dated at approximately 600 years (Hart2008) Similarly it takes 6ndash7 years for a fruit of the SeychellesCoco-de-Mer (Lodoicea maldivica) to develop (cf Edwards et al2003) Such species may be less vulnerable to short-termdemographic stochasticity and their longevity and naturally lowregeneration over long periods may result in underestimates ofpopulation viability where only adult tree counts are used forpopulation estimates (eg Fleischmann et al 2005) However lifehistory and demography vary considerably among oceanic islandplants not all are slow-growing and long-lived (eg Hamann2001 Schumacher et al 2008 2009)

Detecting populations with reduced genetic variation andincreased inbreeding is particularly important for determiningextinction risk especially in the small isolated populationscharacteristic of many island endemic plants (Barrett and Kohn1991 Frankham and Ralls 1998 Frankham 1998 Cole 2003)Because deleterious genetic consequences related to smallpopulation size can impede population responses to suddenenvironmental shifts including competition from alien invasives(Courchamp et al 2003) the influence of population size on thegenetics and reproduction of insular endemics is of major concernto conservation

Numerous conservation plans for rare and fragmented popula-tions exist but minimum viable population sizes (MVP) aregenerally not known The estimation of MVP for plants is stillunreliable in part because it is heavily influenced by modelsadapted from animal biology (see eg Menges 1991) which areinconsistent with many of the population dynamic parameters ofplants Thus even though the MVP concept may assist conserva-tion planning the only real consensus that thousands to tens ofthousands of individuals are required for an MVP is debatablyrealistic even without entering into the difficult issue of thedefinition and timescale of lsquolsquoviablersquorsquo MVPs and extinctionprobabilities are likely to vary over different populations indifferent environments and with different demographic andgenetic parameters and many of these factors cannot bemeasured precisely (Menges 1991) Much research is still neededto develop a realistic MVP model for plants

Although traits associated with rarity may make some speciesmore prone to extinction in other cases rarity can be related toevolutionary persistence (Fiedler and Ahouse 1992 Kunin andGaston 1997) While species that have been rare for millenniamay have developed adaptations to counter the genetic dis-advantages associated with small population sizes and fragmen-tation historically widespread species that have recently becomerare may be more susceptible to such genetic stresses (eg Millar

and Libby 1991) Because it is often difficult to determine the pasthistory of populations the interpretation of the effects of rarity onthe risk of endangerment may be complicated

Molecular techniques may detect trait states that imperilpopulation survival (eg inbreeding) help predict the relationshipbetween population size and probability of extinction (Shaffer1981) or estimate the population sizes necessary to maintain thegenetic diversity of a species However these predictions can beespecially difficult in plants where they are complicated by theplethora of breeding systems effective population sizes andmethods of pollination and dispersal all of which influence thetransmission of genetic variation between generations (Cole2003)

Francisco-Ortega et al (2000) and Crawford et al (2001)reviewed allozyme diversity in endemic plant species from theCanaries and the Juan Fernandez respectively and concluded thatthe level of genetic cohesion of these insular floras (as measuredby the parameter GST Nei 1973) is much lower than in Hawaii(Helenurm and Ganders 1985) or other insular or mainland areasConsequently both studies called for more intensive conservationmeasures on the grounds that alterations of genetic connectivitycould cause genetic homogenization severe reductions in geneticdiversity among populations or hybridization (Francisco-Ortegaet al 2000) Nonetheless estimates of high differentiation amongnatural populations in many published studies could be aspurious consequence of biased intra-population sampling ratherthan a true representation of the biological characteristics of theseinsular floras (Caujape-Castells 2009a)

Although the number of papers describing genetic variation inisland populations has mushroomed in the last 10 years very fewof them include mainland congeners in comparisons Hence welack robust measurements of the genetic depauperation of islandtaxa In one exception Frankham (1997) showed reducedvariation in island populations for a few plant species studied Itis also unclear to what extent the effects of genetic bottlenecksdiffer between island species that are naturally plastic andwidespread (eg Metrosideros polymorpha in Hawaii) and specieswith a narrow natural niche (eg some species of Lobelia in Hawaiiand Linum species in Galapagos)

Lost mutualisms

Plants depend on pollination and seed dispersal mutualismsfor successful reproduction There is increasing evidence thatisland plants suffer from the extinction or decline of seeddispersers (eg Hansen and Galetti 2009 Hansen et al 2008Whittaker and Fernandez-Palacios 2007 McConkey and Drake2006 Cox et al 1991) and pollinators (eg Mortensen et al2008 Cox and Elmqvist 2000) Further invasive plants andanimals may interfere with plantndashanimal mutualisms in bothpositive and negative ways (see below and Kaiser-Bunbury et al2010)

Plants also depend on other mutualisms especially mycor-rhizae Soils on oceanic islands are often nutrient-poor especiallyin phosphorus and many plants depend on mycorrhizal mutual-isms (Tedersoo et al 2007 Koske et al 2002) It is not known towhat extent mycorrhizal associations have been or will bedisrupted through anthropogenic change but such an effect hasbeen suggested for some endemic trees in the Seychelles(Tedersoo et al 2007)

Finally it is important to conserve lsquolsquokeystonersquorsquo or lsquolsquoumbrellarsquorsquospecies many of which are not red-listed because they arenot endemic or endangered These species are however essentialto the structure and functioning of island ecosystems andprotection of their habitats can also benefit species that are morevulnerable

ARTICLE IN PRESS


Habitat alteration and destruction

Habitat destruction on oceanic islands has been substantialand overall has perhaps been the most important factor in causingdeclines of island plants in the past (eg Rolett and Diamond2004 Kirch and Hunt 1997) Today only a small fraction ofprimary vegetation remains on many oceanic islands (eg Brookset al 2002 Kueffer et al 2004 Kingston and Waldren 2005Whittaker and Fernandez-Palacios 2007 Kingsford et al 2009)however the higher elevations on some high islands have higherproportions of less-disturbed vegetation compared to mostinsular areas (eg Hawaii La Reunion Mueller-Dombois andFosberg 1998 Strasberg et al 2005) In general past habitatdestruction has most heavily impacted species in lowland dry tomesic habitats (eg Strasberg et al 2005 Sakai et al 2002 Kirchand Hunt 1997) although Galapagos is an exception withimpacts having been greater in the mesic highlands than thedry lowlands (Snell et al 2002)

Low rainfall and soil fertility appear to be especially importantpredictors of prehistoric deforestation rates at least on PacificIslands (Rolett and Diamond 2004) In present times deforesta-tion has stopped or reversed on many islands with a strongtourism sector and a service-based economy or alternatively thatare now mostly uninhabited (eg Kueffer et al 2004 Lugo 2004Wilkinson 2004) while on others habitat transformation con-tinues or accelerates in coastal areas (Whittaker and Fernandez-Palacios 2007) Wildfires and soil erosion may lead to furtherhabitat loss (eg Kingston and Waldren 2005 DrsquoAntonio andVitousek 1992)

On some islands that have been heavily deforested secondaryforest has regrown (eg in Ascension Puerto Rico or SeychellesKueffer and Vos 2004 Lugo 2004 Wilkinson 2004) while onothers deforestation rates remain among the highest in theworld (eg some islands in the Caribbean and the PhilippinesBrooks et al 2002) Oceanic islands thus vary widely in their relativecomposition of anthropogenic land degraded secondary vegetationsecondary vegetation of conservation value and relatively undis-turbed habitat (compare Kueffer and Daehler 2009)

Models based on speciesndasharea relationships predict that theextinction risk of species on islands is very sensitive to habitatloss (Brooks et al 2002) Consequently rare and endemic plantspecies are more common in undisturbed than in secondaryvegetation (eg Mueller-Dombois and Fosberg 1998 Perry andMorton 1999 Endress 2002 Strasberg et al 2005) and are oftenconcentrated in small isolated habitat patches (eg Cronk 1980Mueller-Dombois and Fosberg 1998 Wiser et al 2002 Kuefferet al 2004 Kingston and Waldren 2005 Seamon et al 2006) Itis not clear however which species can survive in such restrictedhabitats in the long term (eg Wiser et al 2002) Small habitatfragments may be more vulnerable to disturbances such as fireshurricanes or invasions and edge effects may reduce the effectivesize of fragments of value for plant conservation (eg Mueller-Dombois and Fosberg 1998 Wiser et al 2002 Seamon et al2006) For instance reproduction was negatively correlated withsize of habitat fragments for a species of Dombeya in La Reunion(Gigord et al 1999) Besides the complete transformation ofhabitats more subtle alterations such as disturbance of the forestcanopy or changes in soil stability or fertility may directly orindirectly (eg through the facilitation of invasions) reduce thequality of a habitat for particular native plant species

Invasive alien plant species

Plant invasions on oceanic islands have long been considereddramatic examples of the success of alien plants in colonizing

natural areas (Elton 1958 Cronk and Fuller 1995 Tye et al2002 Tye 2003 Denslow 2003 Kueffer et al 2010) Nowadaysalien plants are an important component of many habitats onmost oceanic islands However the extent to which alien plantinvasions have brought about the decline and extinction of nativeplants on islands is still debated (Simberloff 1995 Sax et al2002 Denslow 2003 Gurevitch and Padilla 2004 Sax andGaines 2008) The assumption that invasive plants causeextinctions has been countered by observations that native andalien plant species richness on different oceanic islands arepositively rather than negatively correlated (Sax et al 2002 Saxand Gaines 2008) and negative impacts of alien plants on nativeplants had rarely (until recently) been rigorously demonstrated(Simberloff 1995 Gurevitch and Padilla 2004) Suggestions thatinvasive plant species drive declines in native plant richness areconfounded by plant invasions and native species declines bothcoinciding with some other environmental change such as habitatdestruction (eg Gurevitch and Padilla 2004) Alien speciesrichness is indeed strongly correlated with indices of economicdevelopment across oceanic islands (Denslow et al 2009 Kuefferet al 2010) possibly in part because anthropogenic habitatalteration is a strong driver of plant invasions (cf Kueffer et al2010) or because economic development favours the deliberateintroduction of a wider range of alien species (Tye 2006a)

The past role of plant invasions versus habitat destruction inthe decline of island plant diversity may thus be difficult todisentangle However it should be of high priority to understandhow invasive plants currently affect endemic plants especiallywhere habitats are heavily disturbed and invaded and populationsizes of many native species are small and studies are nowbeginning to reveal cases in which invasive plants are clearly thedirect cause of declines in native plants on islands

The mechanisms by which invasive plants may affect nativeplants include direct competition interference with plantndashanimalinteractions alteration of habitats and hybridization

CompetitionmdashIt has been argued that oceanic island plants areless competitive than plants of continental origin (cf Denslow2003) In Hawaii many invasive species have higher growth ratesand physiological traits adapted to more extreme conditions thannative species (studies reviewed in Denslow 2003) However inSeychelles growth rates vary considerably within the groups ofnative and invasive woody plants (Fleischmann 1999 Schuma-cher et al 2008 2009) Nevertheless even under low light someinvasive plants show higher growth rates and lower mortalitythan many native species eg Psidium cattleianum (Schumacheret al 2008 2009)

Examples clearly demonstrating that an invasive plant iscompetitively replacing a native plant without other majoranthropogenic disturbances are rare Meyer et al (2003) showedthat successful reproduction of two rare species of Psychotria isnegatively correlated with the density of the invasive Miconia

calvescens Baider and Florens (2006) showed that fruit productionof Sideroxylon grandiflorum in Mauritius was markedly lower inweed-infested areas Denslow (2003) reported that the intro-duced Rubus ellipticus is competitively superior to the Hawaiiannative R hawaiiensis Invasion by the understorey herb Hedychium

gardnerianum can affect overstorey native trees probably throughcompetition for nutrients (Asner and Vitousek 2005) Similarlyalien grasses negatively affect water uptake by native trees in dryhabitats (Cabin et al 2002 Cordell and Sandquist 2008) Treesinvading naturally treeless environments can have particularlysevere competitive effects on native vegetation through shadingas well as via other environmental changes such as fog-dripgeneration (Jager et al 2007 2009)

Further some alien plants can invade relatively undisturbedvegetation rich in rare native species and form dense understorey

ARTICLE IN PRESS


or canopy stands including for instance Clidemia hirta Hedychium

gardnerianum Psidium cattleianum or Miconia calvescens (Meyerand Florence 1996 Meyer 2004 Tassin et al 2006 Asner et al2008 Kueffer et al 2010) In such cases experimental removalof the invasives generally increases native regrowth whichstrengthens the evidence that the invasive species displace nativespecies For instance removal of Hedychium gardnerianum fromforests in La Reunion led to an increase in the regrowth of nativeplants (Lavergne 2005) and the release of a biological controlagent against Miconia calvescens in Tahiti has increased theregeneration of the critically endangered tree Myrsine longifolia

(Meyer et al 2007) However there are also cases whereregeneration of at least some native species is good in alien-dominated forests (eg in Palau [Endress 2002] Puerto Rico[Lugo 2004] or Seychelles [Kueffer et al 2007b]) In some casesestablished alien plants may facilitate native regeneration byhindering invasion by other alien species (cf Kueffer et al2007b) In fact invasion by new alien species is often one result ofremoval of invasive plants (eg Lavergne 2005) However wherethe remaining populations of a highly threatened native speciesare being invaded by introduced plants management of theinvasive species is clearly important as in the cases of thecritically endangered Impatiens gordonii in the Seychelles invadedby Clidemia hirta (Huber and Ismail 2006) and Linum cratericola

invaded by Lantana camara in Galapagos (Simbana and Tye 2009)Invasive species may displace native species in habitats with

relatively minor anthropogenic disturbance through naturaldisturbance processes such as hurricanes Although the nativeflora on many oceanic islands especially atolls and dry islandssuch as Galapagos is of a pioneer fast-growing character (egHamann 1979 1993) the native floras of some others lack light-demanding fast-growing species (Mueller-Dombois 2008) andgaps create an opportunity for fast-growing invasive species toestablish and gain high abundance Examples of invasive speciesprofiting from natural gap formation are Pittosporum undulatum inthe Caribbean (Bellingham et al 2005) or various vines on manyoceanic islands (eg Kueffer et al 2004 Meyer 2004) Invasivespecies may sometimes even accelerate gap formation eg inmontane cloud forests in Seychelles (Kueffer 2006) Even minoranthropogenic impacts such as trails (Baret and Strasberg 2005)may enable light-demanding invasives to establish in naturalareas

In summary invasive species tend to be competitively superiorto native species especially in disturbed and open environmentsalthough alien invasives may occasionally have positive impactson the viability of some native plant species A few invasive plantspecies can successfully invade undisturbed vegetation patcheswhere they may become dominant unfortunately these speciesare difficult to identify and predict (Tye 2006a Kueffer et al2010)

Interference with plantndashanimal interactionsmdashInvasive alienplants may produce flowers or fruits of different quality thannative species (eg Kueffer et al 2009) thus competing withnative species for pollinators or seed dispersers In some casesalien plants may support higher densities of pollinators or seeddispersers in a habitat Alien plants may also facilitate the spreadof pests in natural areas as may be the case with the spread of thepolyphagous insect Aleurodicus dispersus into natural areas inSeychelles (Hazell et al 2008) Kaiser et al (2007) report howcoffee plantations increased pest load on endemic plants innearby natural areas Kaiser-Bunbury et al (2010) review the roleof invasive plants in plantndashanimal mutualisms

Alteration of habitatmdashInvasive alien plants can strongly alterhabitat conditions and thus inhibit native species recovery andhabitat rehabilitation In particular alien grasses in dry habitatsoften accelerate fire cycles thereby hindering the establishment

of native plants or increasing their mortality (eg DrsquoAntonio andVitousek 1992 DrsquoAntonio et al 2000) Invasive nitrogen-fixingtrees may increase soil N availability and thereby favour invasivespecies over native ones (eg Hughes and Denslow 2005) buteven non-N-fixers may change litter and nutrient cycles withgenerally unknown impacts on the balance between native andintroduced species (Jager et al 2009) The impacts and roles ofinvasive species in a habitat and ecosystem context have beenwidely reviewed drawing extensively on oceanic island examples(eg Zavaleta et al 2001 Levine et al 2003 Ewel and Putz 2004Mooney et al 2005) or specifically focusing on islands (egDrsquoAntonio and Dudley 1995 Tassin et al 2006 Reaser et al2007 Kueffer and Daehler 2009) A general conclusion importantfor conservation is that in highly degraded island ecosystemsalien species can play both positive and negative roles so impactshave to be assessed specifically for every habitat (Kueffer andDaehler 2009)

HybridizationmdashThe risk of hybridization between a nativespecies and a closely related alien species may be particularlyproblematic on islands (Francisco-Ortega and Santos-Guerra2001 Levin et al 1996) In the Juan Fernandez cultivation ofseveral species of Dendroseris (Asteraceae) in the CorporacionNacional Forestal (CONAF) garden resulted in the production ofinter-specific hybrids which could endanger the integrity of thespecies Daehler and Carino (2001) have recently reviewedhybridization between native and alien plants with a specialfocus on Hawaii

Invasive alien invertebrates and pathogens

New alien pests have been and still are introduced at a veryrapid rate in oceanic islands but there is relatively littleinformation on their effects on native plants Krushelnycky et al(2005) report that inspection of cargo in one Hawaiian airportduring only 20 weeks in 2000ndash2001 detected 125 alien insectspecies and 16 plant pathogens not known to be established inHawaii Today at least some 2700 alien insect species are presentin Hawaii (Juvik and Juvik 1998) Similarly in Galapagos 463alien insect species had been recorded by 2006 an increase of 186introductions since an inventory in 1998 (Causton et al 2006)almost all of them brought accidentally to the islands

Some alien invertebrates constitute direct threats for theendemic flora The Cottony Cushion Scale Insect Icerya purchasi forinstance caused severe damage to mangroves and a range ofendemic plant species when introduced accidentally to Galapa-gos to the point that some populations of threatened specieswere almost completely lost This damage was successfullycontrolled by the introduction of the ladybird Rodolia cardinalis

as a biological control agent (Causton et al 2006) Invertebratescan also act as disease vectors

In the Seychelles the common coastal Takamaka tree (Calo-

phyllum inophyllum) was strongly affected by an introduced wiltdisease transmitted by an endemic species of bark beetle (Wain-house et al 1998) This outbreak was partly controlled byincreasing attention to hygienic forestry practices and the treespecies is still not threatened (Hill et al 2003) however additionalmeasures are required such as the removal of dead and dying treescombined with under-planting of other native trees

Due to the small area of many islands pests may very rapidlyspread and affect endemic species Alien diseases affect endemicplants on many islands (eg Brooks 2002) and several dramaticexamples have been reported where most individuals of a specieshave been affected by a new pest within less than a year (eg theErythrina Gall Wasp in Hawaii Heu et al 2006 Yalemar et al2009) The coffee berry moth Prophantis smaragdina severely and

ARTICLE IN PRESS


rapidly reduced the reproductive success of threatened endemicplant species on Mauritius (Kaiser et al 2007) On the other handthe separation of islands by ocean can be an opportunity forcontaining the spread of a disease or pest across an archipelago(eg Wainhouse et al 1998)

Beyond a few dramatic examples there is relatively littleinformation on pest impacts on oceanic island plants Howeverbased on extensive survey Messing et al (2007) found alienaphids feeding and reproducing on 64 native Hawaiian plants(16 indigenous species and 48 endemic species) in 32 familiesMessing and Wright (2006) report that a single recent invader(a leafhopper from China) attacks some 70 endemic plants inHawaii including 14 rare and endangered species Indirect effectson native plants through interference with pollinators arereviewed by Kaiser-Bunbury et al (2010)

Invasive alien vertebrates

Perhaps the best-documented ecological disturbance resultingfrom biological invasions is the invasion of island ecosystems byvertebrate herbivores (Holdgate 1967 Atkinson 2001 Courch-amp et al 2003) Besides habitat loss invasive mammals areviewed as one of the most important drivers of past (Drake andHunt 2009) and present (this study) biodiversity loss on islands

Aside from the often dramatically obvious impacts of largeherbivorous ungulates there is also compelling evidence frommany oceanic islands that predation on fruits or seeds of nativeplants by rodents and other invasive animals can strongly limitrecruitment (Meyer and Butaud 2009 Traveset et al 2009) Forinstance Meyer and Butaud (2009) estimated that 499 of theseeds of a species of sandalwood were eaten by rats beforeripening and they suggested that some plant families inPolynesia eg Arecaceae Elaeocarpaceae Rubiaceae Santalaceaeand Sapotaceae are particularly vulnerable to seed predationEspecially during the dry season rodents may also prey onseedlings eg of palms (Kueffer 2006) and on the stem and barkof adult trees (Meyer and Butaud 2009) Baider and Florens(2006) showed that predation by invasive monkeys on unripefruits had a major impact on the regeneration of the very rare treeSideroxylon grandiflorum in Mauritius Monkeys may also virtuallydefoliate whole adult trees (Cheke and Hume 2008) Invasive pigsuproot native plants (eg Culliney 2006 in Hawaii) and feed onfruits and seedlings (eg in Guam Wiles et al 1996) Invasiveanimals may also interfere with pollination andor seed dispersalmutualisms (Campbell and Atkinson 2002 Traveset and Richard-son 2006 Meyer and Butaud 2009 Traveset et al 2009)

Endemic island plants can have high levels of compounds suchas polyphenols or anthocyanins (Hansen et al 2004 Kueffer et al2008) which provide defence against generalist herbivoresNonetheless many insular species have not evolved effectivedefences against vertebrate grazing as a consequence of theirprolonged evolution in the absence of vertebrate herbivores(Carlquist 1974 Atkinson 1989 Bowen and Van Buren 1997Courchamp et al 2003 but see Hansen et al 2004 Hansen andGaletti 2009) Consequently grazing and trampling by introducedcattle goats sheep donkeys horses or deer has greatly affectedvegetation on many oceanic islands (eg Loope et al 1988Courchamp et al 2003 Reaser et al 2007)

Eradication of invasive vertebrate herbivores has been suc-cessful on many small islands (Campbell and Donlan 2005 Smithet al 2006 Howald et al 2007 Cheke and Hume 2008) and hasled to rapid recovery of native vegetation on many oceanicislands In the Galapagos feral goats donkeys pigs and cattlehave been eradicated or controlled on many islands includingsome of the largest eradication projects for these animals in the

world (Campbell et al 2004 Campbell and Donlan 2005 Cruzet al 2005 Carrion et al 2007) Recovery of Galapagosvegetation after eradication has often been rapid (Hamann1979 1993 Tye 2006b) and post-eradication research hasfocused on the lack of recuperation of certain species Forexample the failure of recovery of Opuntia megasperma (Cacta-ceae) after the eradication of goats from Espanola Island led to anexperimental recovery programme and a reassessment of gianttortoise re-introduction to the island (Tye 2006b)

Cessation of free grazing by goats and sheep on the mountainsof Porto Santo and Madeira in 1995 and 2003 allowed theregeneration of natural vegetation and the expansion of endemicspecies that had been confined to the higher peaks until then(Silva et al 2005b) Similarly the eradication of rabbits onDeserta Grande and Selvagem Grande made possible the remark-able recovery of the vegetation of these islands In Mauritiusgoats and rabbits have been eradicated from Round Islandfollowing which native vegetation has recovered greatly rabbitsand rats were also eradicated from Gunnerrsquos Quoin with similarsuccess (Baider and Florens 2005 Cheke and Hume 2008) In theCanaries cessation of traditional grazing after more than twomillennia on the summits of La Palma and Tenerife triggered rapidrecovery of the flora in these areas including several extremelyendangered species (A Santos-Guerra personal observation) Inthe Hawaiian Islands large fencing projects in the national parksled to remarkable recovery of several native plant species (Loopeand Medeiros 1994)

Invasive animals seem to be the key threat factor for manyplants in the Canaries Hawaii Galapagos and Juan Fernandezwhile for others they may be one among several factors involvedin the decline (Meyer and Butaud 2009 Perry and Morton 1999)In rare cases some native plants may profit from the presence ofalien animals (eg Traveset et al 2009) and introduced animalscan also reduce the abundance of some plant invaders (egCampbell and Donlan 2005 Kueffer 2006)

Climate change and pollution

The impacts of climate change on island plant biodiversity arelikely to be substantial (Millennium Ecosystem Assessment[MEA] 2005 Mimura et al 2007) Changes in precipitationpatterns are difficult to predict because most oceanic islandclimates are influenced by large-scale weather systems such asthe position of the inter-tropical convergence zone the Asianmonsoon or the El Nino Southern Oscillation and it is not clearhow these systems will shift in a changing climate (Mimura et al2007 Sachs et al 2009) However an overall trend towardsless rainfall has been suggested for tropical and subtropicaloceanic islands (Giambelluca et al 2008) Climate change mayalso lead to shifts in the trade wind inversion layer therebyaffecting rainfall generated by local orographic effects (Mimuraet al 2007 Loope and Giambelluca 1998 Sachs et al 2009)Climate change could also increase inter-annual variability inprecipitation which might disrupt the highly specific conditionsnecessary to sustain cloud forests and dryland vegetation insubtropical and tropical oceanic islands (Mimura et al 2007Kueffer et al 2007a Loope and Giambelluca 1998) An increasedincidence of extreme events such as hurricanes floods ordroughts can be expected on many islands (Mimura et al2007) promoting enhanced habitat disturbances which couldlead to increased mortality of native species or facilitate invasionby non-native species Sea level rise will reduce the area forcoastal vegetation in some places for instance a 50 loss ofmangrove area is predicted for American Samoa (Mimura et al

ARTICLE IN PRESS


2007) Increased dry periods may also increase the risk of fires(Martın 2009 James 2008)

With changing climate oceanic island plants have feweroptions than mainland plants to migrate to suitable habitat Onhigh islands some altitudinal movement may be possible(Gillespie et al 2008) but on small and low-lying islands it maynot Habitat fragmentation or competition from non-nativespecies may impede migration of many species (James 2008)and oceanic island plant endemics often demonstrate lowdispersability (eg Carlquist 1974)

Some island plants may have sufficient phenotypic plasticity toadapt to novel climatic conditions Some oceanic island plantstolerate environmental stress (eg drought) through a resource-conserving slow growth strategy (Castillo et al 2007 Schuma-cher et al 2008) that probably makes them resilient to somedegree of climate change On high islands there are typicallysome species that are common across broad climatic gradientseg Metrosideros polymorpha or Acacia koa in Hawaii which maybe able to adapt to climate changes through genetic adaptation(Ares et al 2000) or phenotypic plasticity However most in situ

speciation on islands occurred within particular habitats andclimate zones (eg Gillespie et al 2008) as revealed by the manyendemic plants that have narrow distributions in limited habitatsand are particularly vulnerable to extinction (eg Sakai et al 2002for Hawaii) Thus vulnerability to climate change may beparticularly high for rare island species (James 2008)

Some of the characteristics that constitute the lsquolsquoislandsyndromersquorsquo (Carlquist 1974) may also make insular endemicsmore vulnerable to shifts in environmental conditions Inparticular woodiness which allows longer generation timeswhen climatic conditions are stable could represent a serioushindrance to survival in a changing environment (David Bramwellpers commun) Increasing temperatures and aridity are contraryto the conditions supposed by Carlquist (1974) to favour thedevelopment of woodiness (ie a moderate humid climate withlong growing seasons) The percentage of woody endemics withrespect to the total endemic flora in our nine studied archipelagosis high 76 for the Seychelles (Huber and Ismail 2006) ca 73for Hawaii (Sakai et al 2002) 72 for the Canaries (Aldridge1979) 63 for Cape Verde (W Lobin unpubl data) or 52 inMadeira (R Jardim unpubl data) In Juan Fernandez thepercentage of trees sub-trees and shrubs in the flora is smaller(47) but 85 of the native and endemic taxa are perennials(Bernardello et al 2001) The Azores show the lowest percentageof woody endemics (19) but similarly to Juan Fernandez ca 86of the plant endemics are long-lived perennial hemicryptophytes(M Moura unpubl data)

While oceanic islands are often remote and may be lessaffected by pollution air pollution is globally transported anddeposited Increase of CO2 may favour some non-native life formsover native ones (Weltzin et al 2004) Given the very low soilnutrient availabilities and correspondingly finely tuned limitationof biotic processes by several nutrients typical for many oceanicislands (Vitousek 2004 Kueffer 2009) oceanic island vegetationmay be especially sensitive to even relatively small nutrientchanges as illustrated by the well-known fertilization-drivenvegetation changes on islands (Ostertag and Verville 2002Hughes and Denslow 2005)

Need for a better knowledge base

Knowledge of the taxonomy distribution and threat status ofplants on oceanic islands is insufficient Our estimation thatbetween 3500 and 6800 insular plant endemics are endangeredworldwide might be a severe underestimate Urgent efforts are

needed in all archipelagos to train the next generation of planttaxonomists to address these limitations New technologies suchas DNA lsquolsquobarcodingrsquorsquo projects (Faith and Williams 2005) and otherDNA databanks such as those initiated in some insular regions(eg Macaronesia Hawaii) may contribute many new insightsincluding the design of nature reserves through estimates ofphylogenetic diversity per unit area The lsquolsquoMoorea BiocodeProjectrsquorsquo is an initiative to barcode all living organisms includingvascular and non-vascular plants (Check 2006 see also httpmooreaberkeleyedubiocode and httpwwwmooreabiocodeorg) Greater DNA banking of insular endemics is needed tocomplement research on systematics and permit conservationinitiatives based on the new taxonomic information generated

Knowledge of the basic reproductive biology of endemic plantsvaries greatly between the focal archipelagos assessed in thisstudy In Hawaii little is known for all but a very few species (egCarr et al 1986 Sakai et al 1989 1995a b Drake and Morden2008) In Juan Fernandez Anderson et al (2001) and Bernardelloet al (2001) surveyed floral traits breeding systems floral visitorsand pollination systems and presented an extensive discussion ofthe conservation implications of their results In the Mascarenesthere have been numerous studies on breeding systems andpollinators of the flowering plants (eg Humeau and Thompson2001 Litrico et al 2005a b Micheneau et al 2006 2008) InGalapagos studies have centred on the population dynamics ofsome large species (eg Hamann 2001) pollination biology (egMcMullen 1994 2007) and more recently on breeding systems(eg Nielsen et al 2007) Although data are accumulating onaspects of the reproductive biology of various Azorean (Pereira2008) and Canarian endemics (eg Calero and Santos-Guerra1993 Anderson et al 2006 Dupont and Olesen 2006 Crawfordet al 2008 Rodrıguez-Rodrıguez and Valido 2008) a broadknowledge of this aspect of the flora in all Macaronesianarchipelagos is lacking which impedes understanding of thebiological implications of most population genetic studies (egCaujape-Castells et al 2008a b)

Greater knowledge of breeding systems may help to identifywhere reductions in gene flow may lead to increased inbreedingdepression and lowered fitness that may undermine the abilityof endemics to compete with introduced species or limitadaptations to habitatclimatic changes (Frankham 1998Francisco-Ortega et al 2000) Flow charts incorporating biologicalinformation (eg breeding system) plant life history traits rarityand distribution may help to prioritise research on the reproduc-tive biology of endangered oceanic island plants (Kueffer et al2007a Drake and Morden 2008) Established protocols could beadapted for archipelagos lacking plant reproductive studies inorder to promote efficient data collection and its application toconservation In self-incompatible plants molecular techniquescan estimate where stochastic loss of S-alleles has removed cross-compatible genotypes (see Raspe and Kohn 2002) therebyproviding another tool for prioritizing in situ action

First aid measures

Establishment of protected areas and community-basedmanagement areas for in situ conservation is essential to mitigatethreats to plant biodiversity In the nine focal archipelagos theproportion of land area devoted to in situ conservation averages40 (Table 1) but ranges from only about 02 in Cape Verde and5 in Hawaii to almost total protection in Juan Fernandez andGalapagos (ca 90 and 96 respectively) Clearly the land areaprotected should be increased in most archipelagos

However many of the biological threats can undermine theintrinsic survival capabilities of many endangered insular plants

ARTICLE IN PRESS


even if their habitats are protected and managed and thegenerally higher sensitivity of insular plant endemics to environ-mental shifts and natural catastrophes makes it advisable toback up in situ conservation measures by preserving them out oftheir natural areas of occurrence Major investment in ex situ

conservation (ie the propagation and conservation of plants andseeds in Botanic Gardens and other similar facilities) is thus anurgent complementary measure One example of how in situ andex situ strategies can be successfully combined is provided byLotus eremiticus and L pyranthus two Canarian endemic Fabaceaefrom La Palma island that consist of fewer than 10 and 20individuals respectively (Mesa-Coello 2007a b) The lsquoCabildo deLa Palmarsquo decreed a conservation plan (Boletın Oficial de Canarias2006) that comprises the protection and management of theirhabitats the maintenance of clones of most individuals in theCabildorsquos plant nursery to complement the in situ protectionmeasures (Felix Manuel Medina pers commun) and the geneticcharacterisation of all the individuals (Jaen-Molina et al unpubldata)

Within the focal group Hawaii the five Macaronesian archipe-lagos and the Mascarenes all have seed banks with specialized staffand facilities devoted to collecting and properly storing germplasmStarting in 1973 project Artemis of the Universidad Politecnica deMadrid (Gomez-Campo 1986) has also conserved seeds fromMacaronesia distributing duplicates from Canaries and Madeira toother seed banks In Juan Fernandez the seed bank consists ofCONAF personnel simply placing seeds in labelled plastic bagscontaining silica gel in a refrigerator Galapagos and Cape Verde donot have seed banks for endemic plant populations although one isplanned for Galapagos The ca 154 seed accessions collected in CapeVerde by staff of the JBCVC are kept as a trust fund in the seed bankof this institution with a small number of accessions kept at theBerlin Botanical Garden

Despite its importance implementing seed banking in or forsome of the archipelagos presently lacking seed banks could bedifficult for reasons ranging from poor finance or inadequate staffcapacity to legal restrictions on export of live material (seeAppendix 1) However given the political will it seems feasiblethat seeds could be collected by trained local staff and sent toestablished seed banks in other archipelagos or mainland areas(see Table 2) either temporarily until the corresponding archipe-lago can build and maintain its own facility or under a permanentagreement Although seed collections of all endemics may not bepossible it should be a priority to collect critically endangeredspecies and those suffering rapid declines in range or abundanceThe use of molecular genetic information to select populationtargets for seed collection and to maximize the amount of geneticdiversity captured in these species is not yet a common practicethough it is being initiated in the JBCVC (Vilches et al 2004Caujape-Castells 2009b) The seed bank in Mauritius addressesindirectly such intra-species genetic variation by collecting seedsof a given species from several distinct populations whenpossible

Although molecular studies can be carried out on live-collected plant material and herbarium specimens seed bankswould increase the possibilities for identifying priority popula-tions where inbreeding is more severe or for selecting the mostsuitable seed sources for reinforcement of ailing populations(through estimates of evolutionary closeness) Seed bank samplesare frequently used in population reintroduction or enhancementand estimates of the genetic representativeness of collectionswould improve the conservation value and success of theseinitiatives while facilitating monitoring In Madeira a LIFEproject carried out by the Jardim Botanico da Madeira between1999 and 2003 implemented various conservation measures andstudies of eight priority rare endemic species These involved

reintroduction and reinforcement of natural populations seedbanking and restoration of natural habitats in Pico Branco(Porto Santo Island)

In the Azores the Biology Department of the Universidade dos Ac-ores has carried out ex situ conservation of endemics since theearly1990rsquos (Maciel 1995 1996 2004 Moura 1995 1998 2006Toste 1995) whereas in Juan Fernandez understanding seedgermination requirements is a priority for many endemics InGalapagos the stated conservation goal of the Galapagos NationalPark is complete in situ conservation and restoration It has thereforebeen a lesser priority to establish seed banks and other ex situ

collections However extinct populations of the single-islandendemic Scalesia atractyloides may be re-established in the futureusing material from living collections from these populations inCopenhagen Botanic Garden while experiments towards usingcultivated seedlings to reinforce and re-establish populations of theCritically Endangered Linum cratericola on Floreana Island andCalandrinia galapagosa on San Cristobal and of the locally endan-gered Scalesia affinis on Santa Cruz (Atkinson et al 2009) have beenundertaken In the Canaries the government has increased fundingto prioritise in situ recovery plans of threatened species thatoccasionally promotes seed banking or use the existing seedcollections in the reinforcement of natural populations

Conclusions

This review offers the most comprehensive analysis of thethreats to insular endemic plants to date Remote islands havemore endemics per unit area but isolation does not influenceextinction risk significantly whereas island size is a moreimportant predictor of endangerment Critically many threatfactors act synergistically producing impacts that reverberateacross entire ecosystems (Fig 2) Therefore species decline canalso lead to cascading extinctions (Koh et al 2004 Fordham andBrook 2009) and our calculation of between 2000 and 2800endemic insular plant species on the verge of extinction world-wide might be a severe underestimate

The role of specific threat factors depends on the particularendemic and its ecological context Nevertheless we see potentialfor pattern generalisation beyond single cases More data areneeded for assessing the relationships among threats testing theirimpacts and developing ways to prioritise action and strategicallyguide plant conservation Appendix 1 summarises the presentassets and weaknesses for plant conservation in each focalarchipelago and identifies research and political guidelines thatcould help improve local action in the future In addition urgentcoordinated action seems necessary to tackle all the relevantfactors of threat through a holistic conservation approach

The impacts of major threat factors on the endemic plantbiodiversity of the nine focal archipelagos appear to be unrelatedto their geographical proximity (Fig 1) emphasizing the need forenhanced contact among island conservation scientists through-out the world Better communication and coordination of researchefforts among archipelagos with different levels of expertise ineach priority problem would provide opportunities for mutualassistance synergistic action and perhaps common politicalapproaches to plant conservation An island plant conservationnetwork with appropriate communication tools such as awebsite could be instrumental in coordinating knowledgesharing and comparative research among archipelagos Whilethere are examples of regional plant conservation networks(eg for the Mascarenes at httpwwwplantmascorg or forthe Hawaii and the Pacific at httpwwwhearorg) there is noglobal network to date although one existing network whichmight be developed with these objectives is the network of plant

ARTICLE IN PRESS


specialist groups of the IUCN Species Survival Commission Manyecological socioeconomic institutional and methodological chal-lenges specific to islands can only be addressed by sharingexperiences across island groups and oceanic regions The proposednetwork would also facilitate exchange of information on newmethods threats and management techniques and stimulatecommon initiatives between research groups in different countriesIts web site should contain historical and current informationresearch materials such as otherwise inaccessible literature hyper-links to the web pages of research groups lists of plant materialsavailable and other information relevant for research such as permitconditions for collecting and fieldwork These resources would helpresearchers to overcome the diverse hurdles encountered indifferent archipelagos while promoting enhanced collaborationamong scientists working on conservation of island floras

We hope that the present paper will promote the developmentof such a coordinated global research effort to help shareknowledge and expertise to discuss common challenges and toformulate multi-disciplinary conservation objectives for insularplant endemics on a worldwide basis

Acknowledgements

We thank the following researchers who provided insight andhelp with several of the questions discussed in the paper Jo~ao

Table A1Indicators of present and future scientific-political conservation needs in the nine featur

Archipelago Conservation research assets at present

Weaknesses Strengths Suggested improve

Azores Lack of man power

Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level

Canaries Poor funding

Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and

knowledge of biologists

in staff

Good facilities for

research

Good connections among

local plant conservation

centres and researchers

and with other

Macaronesian

archipelagos

The network of nature

reserves and national

parks provides excellent

support for research

Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit

Melo and Luis Silva (Azores) Alfredo J Reyes-Betancort LazaroSanchez-Pinto David Bramwell Aguedo Marrero-Rodrıguez JuliaPerez de Paz Bernardo Navarro Alicia Roca-Salinas and JoseMarıa Fernandez-Palacios (Canaries) Curt Daehler Don DrakeWarren Wagner Loyal Mehrhoff David Lorence and StephenWeller (Hawaii) Jaime Cuevas Alicia Marticorena Marcia Ricciand Tod Stuessy (Juan Fernandez) Miguel Menezes de Sequeira(Madeira) and Claudia Baider and Dominique Strasberg (Mascar-enes) Three anonymous reviewers provided constructive com-ments on an earlier version of the manuscript JCC thanks thesupport of the UNESCO-Unitwin chair for lsquolsquoBiodiversity conserva-tion in Macaronesia and the West of Africarsquorsquo at the Jardın BotanicoCanario lsquolsquoViera y Clavijorsquorsquo CK was partly supported by USDA NRICooperative Research Education and Extension Service Grant no2006-35320-17360 to Curtis C Daehler

Appendix 1

See Table A1

Appendix 2

See Table A2

ed archipelagos according to the priorities identified by the participating scientists

Foresight of urgent future needs in conservation

ments Scientific Political

ation with

n investigation

enefit from

d

sets and

cooperation

of research at

Medium-term monitoring

data to determine population

trends (probability of

extinction) and to clarify the

impact of different threats

Exhaustive molecular work

for taxonomic re-evaluations

and population genetic

analysis development of

plant propagation

techniques

Revise and enforce

legislation

Develop and approve species

specific recovery plans based

on scientific data with very

precise goals and with a legal

nature

Consistent financial support

with the development of a

line of grants exclusively

dedicated to support Azorean

flora and vegetation studies

klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature

Increase the offer of jobs as

field botaniststaxonomists

Stabilise staff who can

support research through

their skills

Carry out and publish basic

reproductive biology studies

for most endemic plants

Train more young local

botanists in modern scientific

research methods

Increase institutional

collaboration at the regional

and international levels

Glean information on

historical collections out of

the archipelago including

typification of the endemic

Flora

Improve funding schemes

Enforce the application of the

existing laws

Improve administrative

control of grazing and

invasive alien species

Relax the bureaucracy

needed at present for local

botanists to sample wild

specimens maybe through a

general lsquolsquoresearcher IDrsquorsquo

Recognize the population as

the minimum biological

conservation unit deserving

legal protection

Implement institutional

support for the facilities

devoted to the

ex-situ preservation of the

flora (herbaria seed and DNA

banks) and to store

information relevant to

biodiversity research

Increase interaction between

the administration and

researchers

ARTICLE IN PRESS

Table A1 (continued )

Archipelago Conservation research assets at present Foresight of urgent future needs in conservation

Weaknesses Strengths Suggested improvements Scientific Political

Cape Verde Lack of funding by

government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with

foreign institutions

Training programs addressed to

local biologists

Basic research activities on

populations of threatened

species

Increase capacity of

staffthrough higher level

training


Galapagos Not highly valued

by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists

Improvements in policy

planning and capacity at local

research and management

institutions

Survey monitoring

population dynamics studies

population management

Enforcement of existing laws

and protocols

More funding

Hawaii Lack of funds

Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding

More interactions of local

groups national and

international conservation

biologists

Alien species control (both

for invasive diseases insect

pests that are affecting

endangered plants and for

invasive plants)

Protection of habitat

Studies of endemic species

(reproductive biology

anticipated responses to

global change y )

Will to control feral

mammals like pigs goats

and deer

Will to set up an effective

biosecurity programme that

keeps new disease pests and

weeds out of Hawaii and has

a rapid response to those that

get through the barriers

Will for funding to effectively

protect conservation zones

Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the

uniqueness and fragility

of the endemic flora

Concerted effort to

educate the local human

population on the

importance of the flora

Programme to assist

locals in identifying new

invasives and in

reporting them to CONAF

personnel

Success in the cultivation

of most endangered

species

Need to document occurrence of

new populations in the

herbarium

Deposit duplicates in herbarium

on mainland Chile

Require that applications for

permission to work in the

islands include agreement to

deposit with CONAF in Chile

publications resulting from

work

Use CONAF personnel to

attempt to limit the

introduction of invasives

Reduce further the goats and

rabbits

Prevent the continued

encroachment of several

aggressive invasives into

native forest

Restoration of the extensive

eroded areas

Studies of the basic biology of

endemics (demography

reproductive biology and

population genetics)

Enforce the rule of no animal

husbandry in the park

Requirement to give

vouchers to CONAF before

anyone working on the flora

can leave the islands

Concerted efforts to prevent

introduction of exotic plant

species

Continued development of

ecotourism that enhances

economy but has minimal

environmental impact

Plan to minimize the conflict

between the local human

population and CONAF on

issues such as cattle grazing

and land use in the national

park lands

Support for professional

conservation biologist in

residence in the islands to

interact with CONAF

personnel

Obtain support from private

foundations foreign

governments etc for basic

research on the flora

Madeira amp

Selvagens

Lack of funds

Lack of human

resources

Large protected areas

Easily accessible plant

populations

Coordination of efforts with the

different Madeiran institutions

working on plant studies and

conservation

Increase funding for continuing

conservation projects

Increase studies on endemic

species (molecular

reproductive biology y )

Increase monitoring

Measures to control invasive

alien species

A effective control

programme for the invasive

alien species

Regional legislation to

conserve Madeiran species

Mascarenes Low government

commitment

(except Reunion)

Most staff poorly

Well curated Herbaria

Floral diversity and

endangerment status

well known

Increase collaboration (local -

island scale- regional -

Mascarene scale- and beyond)

Increase capacity of staff

Improve understanding of

alien species impacts on

native plants and

communities

Proper enforcement of

existing legislation

(Mauritius and Rodrigues

mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with

external institutions

Biological invasions

strategies published for

Mauritius (2008) and

Reunion (2009)

through higher level training

(mostly Mauritius Rodrigues)

Mainstream conservation

Improve the effectiveness

and lower the cost of invasive

alien species control

Develop more cost effective

methods of ecosystem

restoration

Improvement of legislation

particularly regarding non

introduction of potentially


Commitment to a meaningful

ecosystem approach to

conservation as defined for

eg in the National

Biodiversity Strategy and

Action Plan of Mauritius and

Rodrigues

Proper use of the existing line

of funds dedicated to

conservation (Mauritius and

Rodrigues)

Seychelles Lack of local

trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007

Established links with

several overseas research

institutions

Increasing awareness of

the need for conservation

research

Success in propagating

many endemic species

New UNDP-GEF project

on management of


and biosecurity

measures

Widen the network of research

institutions

Actively access local regional amp

international finance

Increase human and financial

capacity for research

Improve curation of the national

herbarium and create seed

collection

Improve the plant diversity

information base including

genetic studies

Improve ex-situ conservation

techniques and protocols

Prioritise species and

habitats for conservation

action

Research in plant-

environment inter-

relationships and biotic

interactions


methodologies for in situ

conservation work

Identify potential climate

change impacts on habitats

and species

Better enforcement of

existing laws

Collaboration with local and

overseas organisations to

find new ways of funding

training research and

conservation efforts

Ensure research projects

include capacity building

element inputs to local data

bases etc

Improve media and

educational awareness of

plant conservation issues and

successes

Encourage the development

of participatory methods of

resolving stakeholder

conflicts over land use

resource use etc

Table A2Summary of data from 62 archipelagos of which 53 were used for the endangerment level analyses Islandisland group areas were taken from the data available at the

UNEP islands website (httpislandsunepch)

Islands Region II Area (km2) Plant endemics

Total EN CR

Andaman and Nicobar Indian 386 71932 1227 10 14

Antigua and BarbudapoundO Caribbean 2770 10 2 0

Azores East Atlantic 757 24345 272 3 0

Bahamas Caribbean 362 120922 1117 2 0

Baleares Mediterranean 317 50210 194 0 8

BarbadosO Caribbean 460 4624 12 1 0

Bermuda Caribbean 910 393 115 1 1

Borneoy O Indian 7330990 16750 101 213

Canaries East Atlantic 304 73499 3607 5 5

Cayman Is Caribbean 447 2623 219 2 0

Comoros Indian 485 20213 1136 2 1

Corse Mediterranean 230 87414 1126 0 4

Crete Mediterranean 280 83497 1159 0 4

Cuba Caribbean 285 1058055 13193 59 22

Dominica Caribbean 410 7873 112 3 0

Fiji AsiaOceania 865 180706 1812 14 22

Galapagos East Pacific 638 76102 4180 8 6

Grenada Caribbean 340 3227 14 1 0

Guadeloupe Caribbean 330 17262 119 5 1

Guam AsiaOceania 860 5410 169 1 1

Hawaii East Pacific 1200 104340 5929 64 109


ARTICLE IN PRESS



Total EN CR

Hispaniola Caribbean 430 739290 11400 15 8

Jamaica Caribbean 480 111899 1852 59 46

Japan AsiaOceania 380 3639315 1222 2 0

Java Indian 393 138794 1250 14 13

Juan Fernandez East Pacific 585 1485 6133 0 15

MadagascarO Indian 5877133 17569 110 66

Madeira amp Selvagens East Atlantic 660 7933 136 0 2

Mariana AsiaOceania 800 4773 181 1 2

Marquesas AsiaOceania 1159 10812 1132 5 3

Martinique Caribbean 420 11666 124 3 0

Mauritius Indian 910 19846 1311 14 68

Moluccas Indian 497 663471 1300 1 4

MontserratO Caribbean 390 1241 12 1 0

New CaledoniaO AsiaOceania 166484 12551 66 28

New Zealand AsiaOceania 786 2610799 11942 7 2

Palau AsiaOceania 815 5161 13 1 0

Papua New GuineaO AsiaOceania 7857530 113250 6 1

Philippines AsiaOceania 496 2959955 13471 32 55

Pitcairn East Pacific 1063 429 114 2 3

Puerto Rico Caribbean 590 92499 1215 16 22

Reunion Indian 730 25352 1234 3 5

Saint Helena East Atlantic 1130 1255 150 7 9

St Kitts and NevisO Caribbean 2631 11 1 0

St Lucia Caribbean 410 6398 111 2 1

St Vincent and Grenadines Caribbean 370 3810 120 1 0

Samoa AsiaOceania 918 30489 1149 1 1

Sao Tome and Principe Caribbean 390 100333 1119 2 0

Sardegna Mediterranean 227 239490 1115 0 8

Seychelles Indian 728 15642 770 4 9

Sicily Mediterranean 50 256624 1188 1 8

Society Is AsiaOceania 1025 16277 1273 0 18

Socotra Indian 36067 1260 26 3

Solomon Islands AsiaOceania 699 265565 130 1 0

Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64

Tonga AsiaOceania 785 6725 125 0 1

Trinidad and Tobago Caribbean 210 53175 1215 1 0

Tristan Da Cunha East Atlantic 1065 1928 140 1 0

Turks and Caicos Caribbean 42 7972 19 2 0

Vanuatu AsiaOceania 564 121058 1150 2 1

Virgin IslandsnO Caribbean 4525 15 5 7

The Isolation Index (II) given in the UNEP islands website for a particular island is the sum of the square roots of the distances to the nearest equivalent or larger island the

nearest island group or archipelago and the nearest continent For archipelagos and island groups II was estimated by averaging the IIs of the islands where such data were

available at the cited UNEP website The total data for the EN and CR endemics retrieved from the IUCN Red-List Database (httpwwwiucnredlistorg) are available

through the permalinks httpwwwiucnredlistorgsearchlink4a4b66e1-ae0a8ae7 (EN) and httpwwwiucnredlistorgsearchlink4a4b672b-0d3a83e0 (CR) The total

numbers of endemic plants per island were obtained from the references indicated by numerical superscriptspound Data unavailable for Barbuda y Only Kalimantan Sabah and Sarawak nData unavailable for St John St Thomas and St Croix O Outlier islands excluded from the

statistical analyses (see section lsquolsquoMethodsrsquorsquo) 1Davis et al (1997) 2Silva et al (2005a) and Monica Moura (unpubl data) 3Acebes-Ginoves et al (2004) and Arnoldo Santos-

Guerra (unpubl data) 4Wiggins and Porter (1971) Flora of the Galapagos Islands and Alan Tye (unpubl data) 5Wagner et al (2005) and Ann Sakai (unpubl data)6Marticorena et al (1998) and Daniel J Crawford (unpubl data) 7Kueffer et al (2007a) and Katy Beaver (unpubl data)


References

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Addinsoft 2009 XLSTAT 2009101 URL httpwwwxlstatcomSAldridge A 1979 Evolution within a single genus Sonchus in Macaronesia In

Bramwell D (Ed) Plants and Islands Academic Press London pp 279ndash291Anderson GJ Bernardello G Stuessy TF Crawford DJ 2001 Breeding system

and pollination of selected plants endemic to Juan Fernandez Islands Am JBot 88 220ndash233

Anderson GJ Bernardello G Opel MR Santos-Guerra A Anderson M 2006Reproductive biology of the dioecious Canary Islands endemic Withaniaaristata (Solanaceae) Am J Bot 93 1295ndash1305

Andrus NG Tye A Nesom G Bogler D Lewis C Noyes R Jaramillo PFrancisco-Ortega J 2009 Phylogenetics of Darwiniothamnus (AsteraceaeAstereae) ndash molecular evidence for multiple origins in the endemic flora of theGalapagos Islands J Biogeogr 36 1055ndash1069

Archibald JK Crawford DJ Santos-Guerra A Mort ME 2006 The utility ofautomated analysis of inter-simple sequence repeat (ISSR) loci for resolvingrelationships in the Canary Islands species of Tolpis (Asteraceae) Am J Bot 931154ndash1162

Ares A Fownes JH Sun W 2000 Genetic differentiation of intrinsic water useefficiency in the Hawaiian native Acacia koa Int J Plant Sci 161 909ndash915

Asner GP Vitousek PM 2005 Remote analysis of biological invasion andbiogeochemical change Proc Natl Acad Sci USA 102 4383ndash4386

Asner GP Hughes RF Vitousek PM Knapp DE Kennedy-Bowdoin TBoardman J Martin RE Eastwood M Green RO 2008 Invasive plantstransform the three-dimensional structure of rain forests Proc Natl Acad SciUSA 105 4519ndash4523

Atkinson IAE 1989 Introduced animals and extinctions In Western D PearlMC (Eds) Conservation for the Twenty First Century Oxford UniversityPress New York pp 54ndash69

Atkinson IAE 2001 Introduced mammals and models for restoration BiolConserv 99 81ndash96

Atkinson R Jaramillo P Tapia W 2009 Establishing a new population ofScalesia affinis a threatened endemic shrub on Santa Cruz Island GalapagosEcuador Conserv Evidence 6 42ndash47

Baider C Florens FBV 2005 Changes in population density of twoendemic plant species on Gunnerrsquos Quoin an offshore islet of Mauritius

ARTICLE IN PRESS


after alien rat and hare eradication In Proceedings of the 19th AnnualMeeting of the Society for Conservation Biology 15ndash19 July 2005 BrasiliaBrazil pp 12

Baider C Florens FBV 2006 Current decline of the lsquolsquoDodo Treersquorsquo a case ofbroken-down interactions with extinct species or the result of new interac-tions with alien invaders In Laurance WF Peres CA (Eds) EmergingThreats to Tropical Forests University of Chicago Press Chicago pp 199ndash214

Baldwin BG 1992 Mol Phyl Evol Phylogenetic utility of the internal transcribedspacers of nuclear ribosomal DNA in plants an example from the Compositae1 3ndash16

Baldwin BG Kyhos DW Dvorak J Carr GD 1991 Chloroplast DNA evidencefor a North American origin of the Hawaiian silversword alliance (Asteraceae)Proc Natl Acad Sci USA 88 1840ndash1843

Baret S Strasberg D 2005 The effects of opening trails on exotic plant invasionin protected areas La Reunion island (Mascarene archipelago Indian Ocean)Rev Ecol (Terre Vie) 60 325ndash332

Baret S Fontaine C Boullet V 2006 Nesogenes orerensis (Cordem) Marais - Plandirecteur de conservation outils drsquoaide a la conservation des especes vegetalesmenacees drsquoextinction Version 2006 Conservatoire Botanique National deMascarin Saint-Leu (Reunion)

Baret S Lavergne C Fontaine C Fossy H Ellama B Boullet V 2007 A globalconservation strategy at the Reunion island scale the national botanicalgarden approach Plant Conservation Research Workshop Seychelles 26ndash28June 2007

Barrett SCH Kohn JR 1991 Genetic and evolutionary consequences of smallpopulation size in plants implications for conservation In Falk DAHolsinger KE (Eds) Genetics and Conservation of Rare Plants OxfordUniversity Press New York pp 3ndash30

Bellingham PJ Tanner EVJ Healey JR 2005 Hurricane disturbance accelerateinvasion by the alien tree Pittosporum undulatum in Jamaican montane rainforests J Veg Sci 16 675ndash684

Bernardello G Anderson GJ Stuessy TF Crawford DJ 2001 A survey of floraltraits breeding systems floral visitors and pollination systems of theangiosperms of the Juan Fernandez Islands (Chile) Bot Rev 67 255ndash308

Boletın Oficial de Canarias 2006 Decreto 1702006 de 21 de noviembre porel que se aprueba el Plan de Recuperacion del Pico de Fuego (lsquolsquoLotus Pyranthusrsquorsquo)y del Picocernıcalo (lsquolsquoLotus Eremiticusrsquorsquo) Boletın Oficial Canarias 23728030ndash28051

Boullet V Picot F Lavergne C Baret S 2006 Plans directeurs de conservationet plans drsquourgence une strategie planifiee de conservation des plantesmenacees appliquee aux regions tropicales insulaires cas de lrsquoıle de laReunion Journees Francophones des Sciences de la Conservation de laBiodiversite (lsquole reveil du Dodo IIrsquo) Museum National drsquoHistoire NaturelleParis 7ndash9 March 2006

Bowen L Van Buren D 1997 Insular endemic plants lack defences againstherbivores Conserv Biol 11 1249ndash1254

Brooks FE 2002 Brown root rot disease in American Samoarsquos tropical rainforests Pacific Sci 56 377ndash387

Brooks TM Mittermeier RA Mittermeier CG Da Fonseca GAB Rylands ABKonstant WR Flick P Pilgrim J Oldfield S Magin G Hilton-Taylor C2002 Habitat loss and extinction in the hotspots of biodiversity Conserv Biol16 909ndash923

Broughton DA McAdam JH 2002 A Red Data List for the Falkland Islandsvascular flora Oryx 36 279ndash287

Cabin RJ Weller SG Lorence DH Cordell S Hadway LJ Montgomery RAGoo D Urakam A 2002 Effects of light alien grass and native speciesadditions on Hawaiian dry forest restoration Ecol Appl 12 1595ndash1610

Calero A Santos-Guerra A 1993 Reproductive biology of the high altitudeCanarian flora In Proceedings of the Fifth Meeting of OPTIMA (Organization forthe Phyto-Taxonomic Investigation of the Mediterranean Area) pp 497ndash502

Campbell DJ Atkinson IAE 2002 Depression of tree recruitment by the Pacificrat (Rattus exulans Peale) on New Zealandrsquos offshore islands Biol Conserv 10719ndash35

Campbell K Donlan CJ Cruz F Carrion V 2004 Eradication of feral goatsCapra hircus from Pinta Island Galapagos Ecuador Oryx 38 328ndash333

Campbell K Donlan CJ 2005 Feral goat eradication on islands Conserv Biol 191362ndash1374

Carine MA Russell SJ Santos-Guerra A Francisco-Ortega J 2004Relationships of the Macaronesian and Mediterranean floras mole-cular evidence for multiple colonizations into Macaronesia and back-colonization of the continent in Convolvulus (Convolvulaceae) Am J Bot 911070ndash1085

Carine MA Schaefer H 2009 The Azores diversity enigma why are there so fewAzorean endemic flowering plants and why are they so widespread JBiogeogr in press

Carlquist S 1974 Island Biology Columbia University Press New YorkCarr GD Powell EA Kyhos DW 1986 Self-incompatibility in the Hawaiian

Madiinae (Compositae) an exception to Bakerrsquos rule Evolution 40430ndash434

Carrion V Donlan CJ Campbell K Lavoie C Cruz F 2007 Feral donkey (Equusasinus) eradications in the Galapagos Biodiv Conserv 16 437ndash445

Castillo JM Leira-Doce P Carrion-Tacuri J Munoz-Guacho E Arroyo-Solıs ACurado G Doblas D Rubio-Casal AE Avarez-Lopez AAA Redondo-GomezS Berjano R Guerrero G De Cires A Figueroa E Tye A 2007 Contrastingstrategies to cope with drought by invasive and endemic species of Lantana inGalapagos Biodiv Conserv 16 2123ndash2136

Caujape-Castells J 2009a General GST and y inflation due to biased intra-population sampling and its consequences for the conservation of theCanarian Flora Conserv Genet in press (101007s10592-009-9842-z)

Caujape-Castells J 2009b Criterios geneticos para conservacion ex situ InBacchetta G Bueno-Sanchez A Fenu G Jimenez-Alfaro B Mattana EPiotto B and Virevaire M (Eds) Conservacion ex situ de plantassilvestres Principado de AsturiasLa Caixarsquorsquo Gijon pp 99ndash115 URLhttpwwwgijonesdocumentosjbaenVerdeDivulgacionCientificaConserv-exsitupdfS

Caujape-Castells J Roca-Salinas A Gomes I Marrero-Rodrıguez A 2006Cavegen Banco de Semillas Banco de Genes y Herbarios colaborando conCabo Verde Rincones Atlantico 3 152ndash153

Caujape-Castells J Gonzalez-Perez M Moura M Perez de Paz J Jaen-MolinaR Polifrone M Silva L Febles-Hernandez R Sosa-Henrıquez P PereiraMJ Fernandez-Palacios O Olangua M Cabrera-Garcıa N Rivero E Leon-Sanchez E Reyes E Rodrigues P Duarte AJ Lourenc-o P 2007BIOMABANC ndash Red de bancos de biodiversidad de la flora MacaronesicaRincones Atlantico 4 184ndash189

Caujape-Castells J Jaen Molina R Marrero-Rodrıguez A Naranjo-Suarez JSantos-Guerra A Reyes-Betancort A Gonzalez-Mancebo M Patino J RosRM Werner O 2007 La flora endemica del parque nacional de Garajonaybajo la perspectiva molecular el codigo de barras molecular como herra-mienta taxonomica URL httpwwwmmaesseccionesel_ministerioorga-nismosoapnoapn_inv_proy05htm5S

Caujape-Castells J Baccarani-Rosas M Cabrera-Garcıa N Naranjo Suarez J SantanaI Marrero M Carque E Mesa-Coello R 2008a Population genetic suggestionsto offset the extinction ratchet in the endangered Canarian endemic Atractylispreauxiana (Asteraceae) Plant Syst Evol 273 191ndash199

Caujape-Castells J Marrero-Rodrıguez A Cabrera-Garcıa N Baccarani-RosasM Vilches-Navarrete B 2008b Population genetics of the endangeredCanarian endemic Atractylis arbuscula (Asteraceae) implications for taxonomyand conservation Plant Syst Evol 274 99ndash109

Causton CE Peck SB Sinclair BJ Roque-Albelo L Hodgson CJ Landry B2006 Alien insects threats and implications for conservation of GalapagosIslands Ann Entomol Soc Am 99 121ndash143

Check E 2006 Treasure island pinning down a model ecosystem Nature 439378ndash379

Cheke AS Hume J 2008 Lost Land of the Dodo T amp AD Poyser LondonCole CT 2003 Genetic variation in rare and common plants Annu Rev Ecol

Evol Syst 34 213ndash237Cordell S Sandquist DR 2008 The impact of an invasive African bunchgrass

(Pennisetum setaceum) on water availability and productivity of canopy treeswithin a tropical dry forest in Hawaii Funct Ecol 22 1008ndash1017

Courchamp F Chapuis JL Pascal M 2003 Mammal invaders on islandsimpact control and control impact Biol Rev 78 347ndash383

Cox PA Elmqvist T Pierson ED Rainey WE 1991 Flying foxes as stronginteractors in South Pacific Island ecosystems a conservation hypothesisConserv Biol 5 448ndash454

Cox PA Elmqvist T 2000 Pollinator extinction in the Pacific islands ConservBiol 14 1237ndash1239

Crawford DJ Stuessy TFM Silva O 1987 Allozyme divergence and theevolution of Dendroseris (Compositae Lactuceae) on the Juan FernandezIslands Syst Bot 12 435ndash443

Crawford DJ Ruiz E Stuessy TF Tepe E Aqeveque P Gonzalez F Jensen RJAnderson GJ Bernardello G Baeza CM Swenson U Silva OM 2001Allozyme diversity in endemic flowering plant species of the Juan FernandezArchipelago Chile ecological and historical factors with implications forconservation Am J Bot 88 2195ndash2203

Crawford DJ Archibald JK Stoermer D Mort ME Kelly JK Santos-Guerra A2008 A test of Bakerrsquos law breeding systems and the radiation of Tolpis(Asteraceae) in the Canary Islands Int J Plant Sci 169 782ndash791

Cronk QCB 1980 Extinction and survival in the endemic vascular flora ofAscension Island Biol Conserv 17 207ndash219

Cronk QCB Fuller JL 1995 Plant Invaders Chapman amp Hall London GlasgowNew York Tokyo

Cruz F Donlan CJ Campbell K Carrion V 2005 Conservation action in theGalapagos feral pig (Sus scrofa) eradication from Santiago Island Biol Conserv121 473ndash478

Culliney JL 2006 Islands in a Far Sea The Fate of Nature in Hawaii RevisedEdition University of Hawaii Press

Daehler CC Carino D 2001 Hybridization between native and alien plants andits consequences In Lockwood JL McKinney M (Eds) Biotic Homogeniza-tion Kluwer New York pp 81ndash102

Danton P Perrier C 2006 Nouveau catalogue de la flore vasculaire de lrsquoarchipelJuan Fernandez (Chili) Acta Bot Gallica 153 399ndash587

Davis SD Heywood VH Herrera-MacBryde O Villa-Lobos J Hamilton AC(Eds) 1997 World Wildlife Fund

DrsquoAntonio CM Vitousek PM 1992 Biological invasions by exotic grasses thegrass fire cycle and global change Annu Rev Ecol Syst 23 63ndash87

DrsquoAntonio CM Dudley TL 1995 Biological invasions as agents of change onislands versus mainland In Vitousek PM Loope LL Adsersen H (Eds)Islands Biological Diversity and Ecosystem Function Springer BerlinHeidelberg New York pp 103ndash121

DrsquoAntonio CM Tunison JT Loh RK 2000 Variation in the impact of exoticgrasses on native plant composition in relation to fire across an elevationgradient in Hawaii Austral Ecol 25 507ndash522

ARTICLE IN PRESS


Denslow JS 2003 Weeds in paradise thoughts on the invasibility of tropicalislands Ann Mol Bot Gard 90 119ndash127

Denslow JS Space JC Thomas PA 2009 Invasive exotic plants in the tropicalPacific Islands patterns of diversity Biotropica 41 162ndash170

Dıaz-Perez A Sequeira M Santos-Guerra A Catalan P 2008 Multiplecolonizations in situ speciation and volcanism-associated stepping-stonedispersals shaped the phylogeography of the Macaronesian red fescues(Festuca L Gramineae) Syst Biol 57 732ndash749

Dillon SL Lawrence PK Henry RJ Price HJ 2007 Sorghum resolved as adistinct genus based on combined ITS1 ndhF and Adh1 analyses Plant SystEvol 268 29ndash43

Drake DR Hunt TL 2009 Invasive rodents on islands integrating historical andcontemporary ecology Biol Invas 11 1483ndash1487

Drake DR Morden CW 2008 Reproductive biology of Hawaiirsquos endangeredflora prioritizing research needs for conservation In Proceedings of the2008 ASPT Meeting 26ndash30 July 2008 University of British ColumbiaVancouver BC

Dupont YL Olesen JM 2006 Andromonoecy and buzz pollination in Solanumspecies (Solanaceae) endemic to the Canary Islands An Jard Bot Madrid 6363ndash66

Edwards PJ Kollmann J Fleischmann K 2003 Life history evolution in Lodoiceamaldivica (Arecaceae) Nordic J Bot 22 227ndash237

Elton CS 1958 The Ecology of Invasions by Animals and Plants MethuenLondon

Endress BA 2002 The importance of endemic species to forest succession inPalau Micronesica 34 141ndash153

Ewel JJ Putz FE 2004 A place for alien species in ecosystem restoration FrontEcol Environ 2 354ndash360

Faith DP 1992 Conservation evaluation and phylogenetic diversity BiolConserv 61 1ndash10

Faith DP Reid CAM Hunter J 2004 Integrating phylogenetic diversitycomplementarity and endemism Conserv Biol 18 255ndash261

Faith DP Williams KJ 2005 How large-scale DNA barcoding programs canboost biodiversity conservation planning linking phylogenetic diversity (PD)analyses to the Barcode of Life Database (BoLD) In Australian EntomologicalSocietyrsquos 36th AGM and Scientific ConferenceSeventh Invertebrate Biodiver-sity and Conservation Conference Australian Systematics Society CanberraAustralia 4ndash9 December 2005 pp 83ndash84

Fiedler PL Ahouse JJ 1992 Hierarchies of cause toward an understanding ofrarity in vascular plant species In Fiedler PL Jain SK (Eds) ConservationBiology Chapman amp Hall New York pp 23ndash47

Fleischmann K 1999 Relations between the invasive Cinnamomum verum andthe endemic Phoenicophorium borsigianum on Mahe island Seychelles ApplVeg Sci 2 37ndash46

Fleischmann K Edwards PJ Ramseier D Kollmann J 2005 Stand structurespecies diversity and regeneration of an endemic palm forest on theSeychelles Afr J Ecol 43 291ndash301

Fordham DA Brook P 2009 Why tropical island endemics are acutelysusceptible to global change Biodiv Conserv in press (101007s10531-008-9529-7)

Forest F Grenyer R Rouget M Davies J Cowling RM Faith DP Balmford AManning JC Proches S van der Bank M Reeves G Hedderson TAJSavolainen V 2007 Preserving the evolutionary potential of floras inbiodiversity hotspots Nature 445 757ndash760

Francisco-Ortega J Crawford DJ Santos-Guerra A Carvalho JA 1996Isozyme differentiation in the endemic genus Argyranthemum (AsteraceaeAnthemideae) in the Macaronesian Islands Plant Syst Evol 202137ndash152

Francisco-Ortega JA Santos-Guerra A Kim SC Crawford DJ 2000 Plantgenetic diversity in the Canary Islands a conservation perspective Am J Bot87 909ndash919

Francisco-Ortega JA Santos-Guerra A 2001 Genes y conservacion de plantasvasculares In Fernandez-Palacios JM Martın-Esquivel JL (Eds) Naturalezade las islas Canarias Ecologıa y Conservacion Ed Turquesa Sta Cruz deTenerife pp 357ndash365

Frankel OH Soule ME 1981 Conservation and Evolution Cambridge UniversityPress Cambridge

Frankham R 1997 Do island populations have less genetic variation thanmainland populations Heredity 78 311ndash327

Frankham R 1998 Inbreeding and extinction island populations Conserv Biol12 665ndash675

Frankham R Ralls K 1998 Inbreeding leads to extinction Nature 392 441ndash442Giambelluca TW Diaz HF Luke MSA 2008 Secular temperature changes in

Hawaii Geophys Res Lett 35 L12702 doi1010292008GL034377Gigord L Picot F Shykoff JA 1999 Effects of habitat fragmentation on Dombeya

acutangula (Sterculiaceae) a native tree on La Reunion (Indian Ocean) BiolConserv 88 43ndash51

Gillespie RG Claridge EM Roderick GK 2008 Biodiversity dynamics inisolated island communities interaction between natural and human-mediated processes Mol Ecol 17 45ndash57

Gomez-Campo C 1986 Proyecto lsquolsquoArtemisrsquorsquo XIII Relacion de endemismosrecolectados con anterioridad a 30061986 Esc T S Ing Agronomos UnivPolitecnica de Madrid Madrid

Gonzalez-Perez MA Sosa PA Rivero E Gonzalez-Gonzalez EA Naranjo A2009 Molecular markers reveal no genetic differentiation between Myricarivas-martinezii and M faya (Myricaceae) Ann Bot 103 79ndash86

Gurevitch J Padilla DK 2004 Are invasive species a major cause of extinctionsTrend Ecol Evol 19 470ndash474

Hamann O 1979 Regeneration of vegetation on Santa Fe and Pinta IslandsGalapagos after the eradication of goats Biol Conserv 15 215ndash236

Hamann O 1993 On vegetation recovery goats and giant tortoises on PintaIsland Galapagos Ecuador Biodiv Conserv 2 138ndash151

Hamann O 2001 Demographic studies of three indigenous stand-forming planttaxa (Scalesia Opuntia and Bursera) in the Galapagos Islands Ecuador BiodivConserv 10 223ndash250

Hansen I Brimer L Molgaard P 2004 Herbivore-deterring secondarycompounds in heterophyllous woody species of the Mascarene IslandsPerspect Plant Ecol Evol Syst 6 187ndash203

Hansen DM Kaiser C Muller CB 2008 Seed dispersal and establishment ofendangered plants on oceanic islands the Janzen-Connell model and the useof ecological analogues PLoS One 3 e2111

Hansen DM Galetti M 2009 The forgotten megafauna Science 324 42ndash43Hart P 2008 Tree Age Growth and Death in an Ancient Hawaiian Wet Forest In

2008 Hawaii Conservation Conference Island Ecosystems The Year of the Reef29-31 July 2008 Abstracts Book Hawaii Conservation Alliance Honolulupp 33

Hazell SP Vel T Fellowes MDE 2008 The role of exotic plants in the invasionof Seychelles by the polyphagous insect Aleurodicus dispersus a phylogeneti-cally controlled analysis Biol Invas 10 169ndash175

Helenurm K Ganders FR 1985 Adaptive radiation and genetic differentiation inHawaiian Bidens Evolution 39 753ndash765

Heu RA Tsuda DM Nagamine WT Yalemar JA Suh TH 2006 Erythrina gallwasp Quadrastichus erythrinae Kim (Hymenoptera Eulophidae) New PestAdvisory no 05-03 Department of Agriculture Hawaii

Hill M Currie D Shah NJ 2003 The impacts of vascular wilt disease of thetakamaka tree Calophyllum inophyllum on conservation value of islands in thegranite Seychelles Biodiv Conserv 12 555ndash566

Holdgate MW 1967 The influence of introduced species on the ecosystems oftemperate oceanic islands Pubis IUCN 9 151ndash176

Howald G Donlan CJ Galvan JP Russell JC Parkes J Samaniego A Wang YVeitch D Genovesi P Pascal M Saunders A Tershy B 2007 Invasiverodent eradication on islands Conserv Biol 21 1258ndash1268

Huber M Ismail S 2006 Suggested IUCN Red List Status of the Endemic WoodyPlants of the Inner Seychelles Unpublished MSc Thesis Institute ofIntegrative Biology ETH Zurich

Hughes RF Denslow JS 2005 Invasion by a N2-fixing tree alters function andstructure in wet lowland forests of Hawaii Ecol Appl 15 1615ndash1628

Humeau L Thompson JD 2001 The allometry of flower size dimorphism indioecious Dombeya species on La Reunion Ecol Lett 4 221ndash228

Jaen-Molina R Caujape-Castells J Reyes-Betancort JA Akhani H Fernandez-Palacios O Perez de Paz J Febles-Hernandez R Marrero-Rodrıguez A 2009The molecular phylogeny of Matthiola R Br (Brassicaceae) inferred from ITSsequences with special emphasis on the Macaronesian endemics Mol PhylEvol in press (101016jympev200908031)

Jaffre T Bouchet P Veillon JM 1998 Threatened plants of New Caledonia isthe system of protected areas adequate Biodiv Conserv 7 109ndash135

Jager H Tye A Kowarik I 2007 Tree invasion in naturally treeless environ-ments impacts of quinine (Cinchona pubescens) trees on native vegetation inGalapagos Biol Conserv 140 297ndash307

Jager H Tye A Kowarik I 2009 Destruction without extinction long-termimpacts of an invasive tree species on Galapagos highland vegetation J Ecolin press (101111j1365-2745200901578x)

James SA 2008 Climate change impacts on native plant communities inMelanesia In Leisz SJ Burke-Burnett J (eds) Climate Change andBiodiversity in Melanesia Bishop Museum Technical Report no 42(8)

Jardim R Sequeira M 2008 The vascular plants (Pteridophyta and Spermato-phyta) of the Madeira and Selvagens archipelagos In Borges PAV Abreu CAguiar AMF Carvalho P Jardim R Melo I Oliveira P Sergio C SerranoARM Vieira P (Eds) A list of terrestrial fungi flora and fauna of Madeira andSelvagens archipelagos Direcc- ~ao Regional do Ambiente da Madeira andUniversidade dos Azores Funchal and Angra do Heroısmo pp 157ndash207

Juvik SP Juvik JO 1998 Atlas of Hawaii University of Hawaii Press third edKaiser CN Hansen DM Mueller CB 2007 Exotic pest insects another

perspective on coffee and conservation Oryx 42 1ndash4Kaiser-Bunbury CN Traveset A Hansen DM 2010 Conservation and restora-

tion of plant-animal mutualisms on oceanic islands Perspectives in PlantEcology Evolution and Systematics 12 131ndash143

Kier G Kreft H Lee TM Jetz W Ibisch PL Nowicki C Mutke J Barthlott W2009 A global assessment of endemism and species richness across island andmainland regions Proc Natl Acad Sci USA 106 9322ndash9327

Kingsford RT Watson JEM Lundquist CJ Venter O Hughes L Johnston ELAtherton J Gawel M Keith DA Mackey BG Morley C Possingham HPRaynor B Recher HF Wilson KA 2009 Major conservation policy issues forbiodiversity in oceania Conserv Biol 23 834ndash840

Kingston N Waldren S 2005 A conservation appraisal of the rare and endemicvascular plants of Pitcairn Island Biodiv Conserv 14 781ndash800

Kirch PV Hunt TL (Eds) 1997 Prehistoric Environmental and LandscapeChange Yale University Press New Haven and London

Koh LP Dunn RR Sodhi NS et al 2004 Species coextinctions and thebiodiversity crisis Science 305 1632ndash1634

Koske RE Gemma JN Flynn T 2002 Mycorrhizae in Hawaiian angiosperms asurvey with implications for the origin of the native flora Am J Bot 79 853ndash862

ARTICLE IN PRESS


Kreft H Jetz W Mutke J Kier G Barthlott W 2008 Global diversity of islandfloras from a macroecological perspective Ecol Lett 11 116ndash127

Krushelnycky PD Loope LL Reimer NJ 2005 The ecology policy andmanagement of ants in Hawaii Proc Haw Entomol Soc 37 1ndash25

Kueffer C Vos P Lavergne C Mauremootoo J 2004 Case Studies on the Statusof Invasive Woody Plant Species in the Western Indian Ocean 1 SynthesisForestry Department Food and Agriculture Organization of the United NationsRome Italy

Kueffer C Vos P 2004 Case Studies on the Status of Invasive Woody PlantSpecies in the Western Indian Ocean 5 Seychelles Forestry Department Foodand Agriculture Organization of the United Nations Rome Italy

Kueffer C 2006 Impacts of woody invasive species on tropical forests of theSeychelles Department of Environmental Sciences Diss ETH no 16602 ETHZurich

Kueffer C Beaver K Kaiser CN (Eds) 2007a Seychelles Plant ConservationResearch Agenda 2008ndash2015 Ministry of Environment Natural Resources andTransport (MENRT) and Plant Conservation Action group (PCA) Seychelles

Kueffer C Schumacher E Fleischmann K Edwards PJ Dietz H 2007b Strongbelowground competition shapes tree regeneration in invasive Cinnamomumverum forests J Ecol 95 273ndash282

Kueffer C Klingler G Zirfass K Schumacher E Edwards P Gusewell S 2008Invasive trees show only weak potential to impact nutrient dynamics inphosphorus-poor tropical forests in the Seychelles Funct Ecol 22 359ndash366

Kueffer C 2009 Reduced risk for positive soil-feedback on seedling regenerationby invasive trees on a very nutrient-poor soil in Seychelles Biol Invas in press(101007s10530-009-9433-4)

Kueffer C Daehler C 2009 A habitat-classification framework andtypology for understanding valuing and managing invasive speciesimpacts In Inderjit (Ed) Management of Invasive Weeds Springer Berlinpp 77ndash101

Kueffer C Kronauer L Edwards PJ 2009 Wider spectrum of fruit traits ininvasive than native floras may increase the vulnerability of oceanic islands toplant invasions Oikos 118 1327ndash1334

Kueffer C Daehler CC Torres-Santana CW Lavergne C Meyer JY Otto RSilva L 2010 A global comparison of plant invasions on oceanic islandsPerspect Plant Ecol Evol Syst in press (101016jppees200906002)

Kunin WE Gaston KJ (Eds) 1997 The Biology of Rarity Causes andConsequences of Rare-common Differences Chapman amp Hall London

Lavergne C Duret C Gigord L 2004 The last wild Red Latan population in theMascarene Archipelago Plant Talk 36 32ndash33

Lavergne C 2005 Invasion par les plantes exotiques envahissantes dans une ıleoceanique Impact ecologique a la Reunion et valeur patrimoniale desecosystemes indigenes envahis Rapport final Programme de Recherche surles Invasions Biologiques INVABIO Conservatoire Botanique National deMascarin et Universite de la Reunion La Reunion France

Lawesson JE Adsersen H Bentley P 1987 An updated and annotated check listof the vascular plants of the Galapagos Islands No 16 Botanical InstituteUniversity of Aarhus Aarhus Denmark

Levin DA Francisco-Ortega J Jansen RK 1996 Hybridization and theextinction of rare species Conserv Biol 10 10ndash16

Levine JM Vila M DrsquoAntonio CM Dukes JS Grigulis K Lavorel S 2003Mechanisms underlying the impacts of exotic plant invasions Proc R SocLond B 270 775ndash781

Litrico I Pailler T Thompson JD 2005a Gender variation and primarysuccession in a tropical woody plant Antirhea borbonica (Rubiaceae) J Ecol93 705ndash715

Litrico I Ronfort J Verlaques R Thompson JD 2005b Spatial structure ofgenetic variation and primary succession in the pioneer tree species Antirheaborbonica on La Reunion Mol Ecol 14 1575ndash1584

Loope LL Hamann O Stone CP 1988 Comparative conservation biology ofoceanic archipelagoes Hawaii and the Galapagos Bioscience 38 272ndash282

Loope LL Giambelluca TW 1998 Vulnerability of island tropical montane cloudforests to climate change with special reference to east Maui Hawaii ClimChange 39 503ndash517

Loope LL Medeiros AC 1994 Impacts of biological invasions on the manage-ment and recovery of rare plants in Haleakala National Park Maui HawaiianIslands In Bowles ML Whelan CJ (Eds) Restoration of Endangered SpeciesCambridge University Press Cambridge UK pp 143ndash158

Lorence DH Wood KR 1994 Kanaloa a new genus of Fabaceae (Mimosoideae)from Hawaii Novon 4 137ndash145

Lugo AE 2004 The outcome of alien tree invasions in Puerto Rico Front EcolEnviron 2 265ndash273

Lutchman I Aalbersberg B Hinchley D Miles G Tiraa A Wells S 2005Marine Protected Areas Benefits and Costs for Islands WWF the NetherlandsURL httpwwwpandaorgcoralS

Maciel GB 1995 Influencia do tempo de conservac- ~ao das sementes na germinac-~ao de especies vasculares endemicas dos Ac-ores Bol Soc Brot 67 171ndash186

Maciel GB 1996 Influencia da temperatura e da luz na germinac- ~ao de especiesvasculares endemicas dos Ac-ores Ac-oreana 8 209ndash218

Maciel GB 2004 Conservac- ~ao de especies vasculares endemicas dos AzoresEcofisiologia da germinac- ~ao de sementes de alguns taxa e identificac- ~ao ecaracterizac- ~ao de microssatelites de Rubus hochstetterorum Seub Unpub-lished PhD Thesis Departamento de Biologia Universidade dos Ac-ores

Marticorena C Stuessy TF Baeza C 1998 Catalogue of the vascular floraof the Robinson Crusoe or Juan Fernandez Islands Chile Gayana Bot 55187ndash211

Martın JL 2009 Are the IUCN standard home-range thresholds for species a goodindicator to prioritise conservation urgency in small islands A case study inthe Canary Islands (Spain) J Nat Conserv 17 87ndash98

Mauchamp A 1996 Scalesia atractyloides one bite from extinction NoticiasGalapagos 57 24ndash25

Mauchamp A Aldaz I Ortiz E Valdebenito H 1998 Threatened species areevaluation of the status of eight endemic plants of the Galapagos BiodivConserv 7 97ndash107

Maunder M Leiva A Santiago-Valentın E Stevenson DW Acevedo-RodrıguezP Meerow AW Mejıa M Clubbe C Francisco-Ortega J 2008 Plantconservation in the Caribbean Island biodiversity hotspot Bot Rev 74197ndash207

Mayr E 1967 The challenge of island faunas Aust Nat Hist 15 369ndash374McConkey KR Drake DR 2006 Flying foxes cease to function as seed dispersers

long before they become rare Ecology 87 271ndash276McMullen CK 1994 Comparative studies on the pollination biology of

Darwiniothamnus tenuifolius (Asteraceae) and Plumbago scandens (Plumbagina-ceae) on Pinta Island and Santa Cruz Island Galapagos Phytologia 78 30ndash38

McMullen CK 2007 Pollination biology of the Galapagos endemic Tournefortiarufo-sericea (Boraginaceae) Bot J Linn Soc 153 21ndash31

Medail F Quezel P 1997 Hot-spots analysis for conservation of plantbiodiversity in the Mediterranean Basin Ann Mo Bot Gard 84 112ndash127

Menges ES 1991 The application of minimum viable population theory toplants In Falk DA Holsinger KE (Eds) Genetics and Conservation of RarePlants Oxford University Press New York pp 45ndash61

Mesa-Coello R 2007a Seguimiento de poblaciones de species amenazadas(2007) Lotus eremiticus A Santos Gobierno de Canarias Sta Cruz de Tenerife

Mesa-Coello R 2007b Seguimiento de poblaciones de species amenazadas(2007) Lotus pyranthus P Perez Gobierno de Canarias Sta Cruz de Tenerife

Messing RH Wright MG 2006 Biological control of invasive species solutionor pollution Front Ecol Environ 4 132ndash140

Messing RH Tremblay MN Mondor EB Foottit RG Pike KS 2007 Invasiveaphids attack native Hawaiian plants Biol Invas 9 601ndash607

Meyer JY 2004 Threat of invasive alien plants to native flora and forestvegetation of Eastern Polynesia Pacific Sci 58 357ndash375

Meyer JY Florence J 1996 Tahitirsquos native flora endangered by the invasion ofMiconia calvescens DC (Melastomataceae) J Biogeogr 23 775ndash781

Meyer JY Florence J Tchung V 2003 The endemic Psychotria (Rubiaceae) ofTahiti (French Polynesia) threatened by the invasive Miconia calvescens(Melastomataceae) status distribution ecology phenology and conservationRev Ecol (Terre Vie) 58 161ndash185

Meyer JY Duplouy A Taputuarai R 2007 Dynamique des population de lrsquoarbreendemique Myrsine longifola (Myrsinacees) dans les forets de Tahiti (PolynesieFranc-aise) envahie par Miconia calvescens (Melastomatacees) apres introduc-tion drsquoun champignon pathogene de lutte biologique premieres investiga-tions Rev Ecol (Terre Vie) 62 17ndash33

Meyer JY Butaud JF 2009 The impacts of rats on the endangered native flora ofFrench Polynesia (Pacific Islands) drivers of plant extinction or coup de gracespecies Biol Invas 11 1569ndash1585

Micheneau C Fournel J Pailler T 2006 Bird pollination in an angraecoid orchid ofReunion Island (Mascarene archipelago Indian Ocean) Ann Bot 97 965ndash974

Micheneau C Fournel J Gauvin-Bialecki A Pailler T 2008 Auto-pollination ina long-spurred endemic orchid (Jumellea stenophylla) on Reunion Island(Mascarene Archipelago Indian Ocean) Pl Syst Evol 272 11ndash22

Millar CI Libby WJ 1991 Strategies for conserving clinal ecotypic and disjunctpopulation diversity in widespread species In Falk DA Holsinger KE (Eds)Genetics and Conservation of Rare Plants Oxford University Press New Yorkpp 149ndash170

Millennium Ecosystem Assessment 2005 Ecosystems and Human Well-beingBiodiversity Synthesis World Resources Institute Washington DC

Mimura N Nurse L McLean RF Agard J Briguglio L Lefale P Payet R SemG 2007 Small islands In Parry ML Canziani OF Palutikof JP van derLinden PJ Hanson CE (Eds) Climate Change 2007 Impacts Adaptation andVulnerability Contribution of Working Group II to the Fourth AssessmentReport of the Intergovernmental Panel on Climate Change CambridgeUniversity Press Cambridge UK pp 687ndash716

Mooney HA Mack RN McNeely JA Neville LE Schei PJ Waage JK (Eds)2005 A New Synthesis Island Press Washington London

Moore MJ Tye A Jansen RK 2006 Patterns of long-distance dispersal inTiquilia subg Tiquilia (Boraginaceae) implications for the origins of amphi-tropical disjuncts and Galapagos Islands endemics Am J Bot 93 1163ndash1177

Mortensen HS Dupont YL Olesen JM 2008 A snake in paradise disturbanceof plant reproduction following extirpation of bird flower-visitors on GuamBiol Conserv 141 2146ndash2154

Moura M 1995 Cultura in vitro de seis taxa vasculares endemicos dos AzoresHypericum foliosum Aiton Lotus azoricus PW Ball Euphorbia azorica SeubertDaboecia azorica Tutin amp Warburg Tolpis azorica (Nutt) P Silva e Leotondonfilii (Hochst) Paiva amp Ormonde Dissertac- ~ao apresentada para provas deAptid~ao Pedagogica e Capacidade Cientıfica Departamento de BiologiaUniversidade dos Azores

Moura M 1998 Conservation of Hypericum foliosum Aiton and endemic Azoreanspecies by micropropagation In Vitro Cellular Development Biology ndash Plant34 244ndash248

Moura M 2006 Desenvolvimento de estrategias para a conservac- ~ao de Viburnumtinus ssp subcordatum (Trel) P Silva multiplicac- ~ao por via seminal e

ARTICLE IN PRESS


vegetativa e estudo da variabilidade genetica das suas populac- ~oes Universi-dade dos Azores Ponta Delgada

Mueller-Dombois D Fosberg FR 1998 Vegetation of the Tropical Pacific IslandsSpringer-Verlag New York

Mueller-Dombois D 2008 Pacific Island forests successionally impoverished andnow threatened to be overgrown by aliens Pacific Sci 62 303ndash308

Myers N Mittermeier RA Mittermeier CG da Fonseca GAB Kent J 2000Biodiversity hotspots for conservation priorities Nature 403 853ndash858

Nei M 1973 Analysis of gene diversity in subdivided populations Proc NatlAcad Sci USA 70 3321ndash3323

Nielsen LR Siegismund HR Hansen T 2007 Inbreeding depression in thepartially self-incompatible endemic plant species Scalesia affinis (Asteraceae)from Galapagos Islands Evol Ecol 21 1ndash12

Nuez F Prohens J Blanca JM 2004 Relationships origin and diversity ofGalapagos tomatoes implications for the conservation of natural populationsAm J Bot 91 86ndash99

Oliva-Tejera F Caujape-Castells J Navarro-Deniz J Reyes-Betancort AScholz S Baccarani-Rosas M Cabrera-Garcıa N 2006 Patterns ofgenetic divergence of three Canarian endemic Lotus (Fabaceae) implica-tions for the conservation of the endangered L kunkelii Am J Bot 931116ndash1124

Ostertag R Verville JH 2002 Fertilization with nitrogen and phosphorusincreases abundance of non-native species in Hawaiian montane forests PlantEcol 162 77ndash90

Pattison RR Goldstein G Ares A 1998 Growth biomass allocation andphotosynthesis of invasive and native Hawaiian rainforest species Oecologia117 449ndash459

Pereira MJ 2008 Reproductive biology of Vaccinium cylindraceum (Ericaceae) anendemic species of the Azores archipelago Botany 86 359ndash366

Perry G Morton JM 1999 Regeneration rates of the woody vegetation ofGuamrsquos Northwest Field following major disturbance land use patternsferal ungulates and cascading effects of the brown treesnake Micronesica 31125ndash142

Pippard H 2009 The Pacific Islands an analysis of the status of species as listedon the 2008 IUCN Red List of Threatened Species IUCN Gland URL httpwwwiucnorgaboutunionsecretariatofficesoceaniaoro_programmesoro_speciesoro_redlistS

Prebble M Dowe JL 2008 The Late Quaternary decline and extinction of palmson oceanic Pacific islands Quat Sci Rev 27ndash28 2546ndash2567

Raspe O Kohn JR 2002 S-allele diversity in Sorbus aucuparia and Crataegusmonogyna (Rosaceae Maloideae) Heredity 88 458ndash465

Reaser JK Meyerson LA Cronk Q De Poorter M Eldrege LG Green E KairoM Latasi P Mack RN Mauremootoo J OrsquoDowd D Orapa W SastroutomoS Saunders A Shine C Thrainsson S Vaiutu L 2007 Ecological andsocioeconomic impacts of invasive alien species in island ecosystems EnvironConserv 34 98ndash111

Reyes-Betancort JA Santos-Guerra A Guma IR Humphries CJ Carine MA2008 Diversity rarity and the evolution and conservation of the CanaryIslands endemic flora An Jard Bot Madrid 65 25ndash45

Rodrıguez-Rodrıguez MC Valido A 2008 Opportunistic nectar-feedingbirds are effective pollinators of bird-flowers from Canary Islands experi-mental evidence from Isoplexis canariensis (Scrophulariaceae) Am J Bot 951408ndash1415

Rolett B Diamond J 2004 Environmental predictors pre-European deforestationon Pacific islands Nature 431 443ndash446

Ruiz E Crawford DJ Stuessy TF Gonzalez F Samuel R Becerra J Silva M2004 Phylogenetic relationships and genetic divergence among endemicspecies of Berberis Gunnera Myrceugenia and Sophora of the Juan FernandezIslands (Chile) and their continental progenitors based on isozymes and nrITSsequences Taxon 53 321ndash332

Sachs JP Sachse D Smittenberg RH Zhang Z Battisti DS Golubic S 2009Southward movement of the Pacific intertropical convergence zone AD 1400ndash1850 Nat Geosci 2 519ndash525

Sakai A Karoly KK Weller SG 1989 Inbreeding depression in Schiedea globosaand S salicaria (Caryophyllaceae) subdioecious and gynodioecious Hawaiianspecies Am J Bot 76 437ndash444

Sakai AK Wagner WL Ferguson DM Herbst DR 1995a Biogeographicaland ecological correlates of dioecy in the Hawaiian Flora Ecology 762530ndash2543

Sakai AK Wagner WL Ferguson DM Herbst DR 1995b Origins of dioecy inthe Hawaiian flora Ecology 76 2517ndash2529

Sakai AK Wagner WL Mehrhoff LA 2002 Patterns of endangerment in theHawaiian flora Syst Biol 51 276ndash302

Sanchez-Pinto L Rodrıguez ML Rodrıguez S Martın K Cabrera A MarreroMC 2005 Spermatophyta In Arechavaleta M Zurita N Marrero MCMartın JL (Eds) Lista preliminar de especies silvestres de Cabo Verde(hongos plantas y animales terrestres) Consejerıa de Medio Ambiente yOrdenacion Territorial Gobierno de Canarias pp 40ndash57

Sax DF Gaines SD 2008 Species invasions and extinction the future of nativebiodiversity on islands Proc Natl Acad Sci USA 105 11490ndash11497

Sax DF Gaines SD Brown JH 2002 Species invasions exceed extinctions onislands worldwide a comparative study of plants and birds Am Nat 160766ndash783

Schumacher E Kueffer C Tobler M Gmur V Edwards PJ Dietz H 2008Influence of drought and shade on seedling growth of native and invasive treesin the Seychelles Biotropica 40 543ndash549

Schumacher E Kueffer C Edwards PJ Dietz H 2009 Influence of light andnutrient conditions on seedling growth of native and invasive trees in theSeychelles Biol Invas 11 1941ndash1954

Seamon JO Mann SS Steele OC Utzurrum RCB 2006 Conservation value ofremnant forest patches tree composition spatial patterns and recruitment inthe Ottoville lowland forest American Samoa Pacific Sci 60 319ndash332

Shaffer ML 1981 Minimum population sizes for species conservation Bioscience31 131ndash134

Silva L Pinto N Press B Rumsay F Carine M Henderson S Sjogren E 2005aList of vascular plants (Pterodophyta and Spermatophyta) In Borges PAVCunha R Gabriel R Frias-Martıns A Silva L Vieira V (Eds) A List of theTerrestrial Fauna (Mollusca e Arthropoda) and Flora (Bryophyta Pteridophytae Spermatophyta) from the Azores Direcc-ao Regional do Ambiente andUniversidade dos Azores Horta Angra do Heroısmo and Ponta Delgada pp131ndash156

Silva M Menezes D Menezes de Sequeira E Menezes de Sequeira M 2005bVascular Plant communities on freira Pterodroma madeira (Mathews 1934)breeding area (oriental Mountains of Madeira) ndash Results on their recovery after 4years without grazing In Proc 48th Ann Symp IAVS Lisbon 24ndash29 July pp 149

Simbana W Tye A 2009 A total population study of a critically endangeredGalapagos plant Linum cratericola reproductive biology and impacts of threatsand protection measures Bot J Linn Soc in press

Simberloff D 1995 Why do introduced species appear to devastate islands morethan mainland areas Pacific Sci 49 87ndash97

Smith DG Shiinoki EK Van der Werf EA 2006 Recovery of nativespecies following rat eradication on Mokolirsquoi Island Orsquoahu Hawaii PacificSci 60 299ndash303

Snell HL Tye A Causton CE Bensted-Smith R 2002 The status of and threatsto terrestrial biodiversity In Bensted-Smith R (Ed) A Biodiversity Vision forthe Galapagos Islands Charles Darwin Foundation and WWF Puerto AyoraEcuador pp 30ndash47

Soule ME Wilcox BA (Eds) 1980 Conservation Biology An EvolutionaryndashEcological Perspective Sinauer Sunderland MA

Strasberg D Rouget M Richardson DM Baret S Dupont J Cowling RM2005 An assessment of habitat diversity and transformation on La ReunionIsland (Mascarene Islands Indian Ocean) as a basis for identifying broad-scaleconservation priorities Biodiv Conserv 14 3015ndash3032

Tassin J Lavergne C Muller S Blanfort V Baret S Le Bourgeois T Triolo JRiviere JN 2006 Bilan des connaissances sur les consequences ecologiquesdes invasions de plants a lrsquoıle de La Reunion (Archipel des Mascareignes OceanIndien) Rev Ecol (Terre Vie) 61 35ndash52

Tedersoo L Suvi T Beaver K K ~oljalg U 2007 Ectomycorrhizal fungi of theSeychelles diversity patterns and host shifts from the native Vateriopsisseychellarum (Dipterocarpaceae) and Intsia bijuga (Caesalpiniaceae) to theintroduced Eucalyptus robusta (Myrtaceae) but not Pinus caribea (Pinaceae)New Phytol 175 321ndash333

Toste M 1995 Micropropagac- ~ao de Rumex azoricus Rech f (Polygonaceae)Trabalho de estagio Departamento de Biologia Universidade dos Azores

Traveset A Richardson DM 2006 Biological invasions as disruptors of plantreproductive mutualisms Trend Ecol Evol 21 208ndash216

Traveset A Nogales M Alcover JA Delgado JD Lopez-Darias M Godoy DIgual JM Bover P 2009 A review on the effects of alien rodents in theBalearic (Western Mediterranean Sea) and Canary Islands (Eastern AtlanticOcean) Biol Invas 11 1653ndash1670

Tye A 1997 Rediscovery of an lsquolsquoextinctrsquorsquo endemic plant ndash the Floreana Flax Linumcratericola Noticias Galapagos 58 2ndash3

Tye A 2002 Threatened species management in an oceanic archipelago theGalapagos Islands In Maunder M Clubbe C Hankamer C Groves M (Eds)Plant Conservation in the Tropics Perspectives and Practice Royal BotanicGardens Kew London pp 323ndash347

Tye A 2003 Plant Research for Conservation in Galapagos Report for the Years1998ndash2003 and Challenges for the Future Charles Darwin Foundation for theGalapagos Islands Puerto Ayora 52pp

Tye A 2006a Can we infer island introduction and naturalization rates frominventory data Evidence from introduced plants in Galapagos Biol Invas 8201ndash215

Tye A 2006b Restoration of the vegetation of the Dry Zone in Galapagos Lyonia9 29ndash50

Tye A Soria M Gardener MR 2002 A strategy for Galapagos weeds In VeitchCR Clout MN (Eds) Turning the Tide The Eradication of Invasive SpeciesIUCN Gland pp 336ndash341

Van Leeuwen JFN Froyd CA van der Knaap WO Coffey EE Tye A WillisKJ 2008 Fossil pollen as a guide to conservation in the Galapagos Science322 1206

Vilches B Roca A Naranjo J Navarro B Bramwell D Caujape-Castells J 2004Estructura espacial de la variacion genetica de Erysimum albescens (Cruciferae)en Gran Canaria implicaciones para la conservacion ex situ en Bancos degermoplasma Bot Macaronesica 25 15ndash30

Vitousek PM 2004 Nutrient Cycling and Limitation Hawaii as a Model SystemPrinceton University Press Princeton

Wagner WL Herbst DR Lorence DH 2005 Flora of the Hawaiian Islandswebsite URL httpbotanysiedupacificislandbiodiversityhawaiianfloraS

Wainhouse D Murphy S Greig B Webber J Vielle M 1998 The role of thebark beetle Cryphalus trypanus in the transmission of the vascular wiltpathogen of takamaka (Calophyllum inophyllum) in the Seychelles For EcolManage 108 193ndash199

ARTICLE IN PRESS


Weltzin JF Belote RT Sanders NJ 2004 Biological invaders in a greenhouseworld will elevated CO2 fuel plant invasions Front Ecol Environ 1 146ndash153

Whittaker RJ Fernandez-Palacios JM 2007 Island Biogeography EcologyEvolution and Conservation second ed Oxford University Press Oxford UK

Wiggins I Porter D 1971 Flora of the Galapagos Islands Stanford UniversityPress

Wiles GJ Schreiner IH Nafus D Jurgensen LK Manglona JC 1996 Thestatus biology and conservation of Serianthes nelsonii (Fabaceae) andendangered Micronesian tree Biol Conserv 76 229ndash239

Wilkinson DM 2004 The parable of Green Mountain Ascension Islandecosystem construction and ecological fitting J Biogeogr 31 1ndash4

Wiser SK Drake DR Burrows LE Sykes WR 2002 The potential for long-term persistence of forest fragments on Tongatapu a large island in westernPolynesia J Biogeogr 29 767ndash787

Yalemar J King C Kaufman L Nagamine W Oishi D 2009 An update on thecurrent status of biological control programs for the Erythrina Gall Wasp(Quadrastichus erythrinae) In 2009 Hawaii Conservation ConferenceIsland Ecosystems Hawaii in a changing climate ecological culturaleconomic and policy challenges and solutions Hawaii Conservation AllianceHonolulu

Zavaleta E Hobbs RJ Mooney HA 2001 Viewing invasive species removal in awhole-ecosystem context Trend Ecol Evol 16 454ndash459


Introduction

Methods

Focal study system

Expert survey



Documentation of plant diversity on oceanic islands status and future prospects

The magnitude of plant diversity endangerment on the worldrsquos islands



Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS


Contents

Introduction 108

Methods 109

Focal study system 109

Expert survey 110

Endangerment analysis of the worldrsquos insular plant endemics 111

Results and discussion 113

Documentation of plant diversity on oceanic islands status and future prospects 113

The magnitude of plant diversity endangerment on the worldrsquos islands 114

A network of threat factors affecting island plants 114

Small population sizes and fragmentation 114

Lost mutualisms 115

Habitat alteration and destruction 116

Invasive alien plant species 116

Invasive alien invertebrates and pathogens 117

Invasive alien vertebrates 118

Climate change and pollution 118

Need for a better knowledge base 119

First aid measures 119

Conclusions 120

Acknowledgements 121

Appendix 1 121

Appendix 2 121

References 124

Islands are an enormously important source of informationand an unparalleled testing ground for various scientifictheories But this very importance imposes an obligation onus Their biota is vulnerable and precious We must protect itWe have an obligation to hand over these unique faunas andfloras with a minimum of loss from generation to generation

Ernst Mayr (1967)

Introduction

Islands are of particular importance for the conservationof global plant diversity Although they make up only some 5of the Earthrsquos land surface about one quarter of all knownextant vascular plant species are endemic to islands (Kreft et al2008) Indices of vascular plant diversity are markedly higherfor islands than for continental areas (Kier et al 2009) and 20 ofthe 34 biodiversity hotspots defined by Conservation Interna-tional (Myers et al 2000 and updates in httpwwwbiodiversityhotspotsorg) are islands or have an important insular compo-nent In addition to this legacy of a unique evolutionary historyinsular ecosystems are also key to the livelihood economy well-being and cultural identity of 600 million islanders roughly one-tenth of todayrsquos world population (Lutchman et al 2005)

Humans have heavily affected island ecosystems (MillenniumEcosystem Assessment [MEA] 2005 Whittaker and Fernandez-Palacios 2007 Kingsford et al 2009) For instance of some 80documented plant extinctions in the last 400 years about 50 wereisland species (Sax and Gaines 2008) In the near future humanpressure on ecosystems will likely increase more markedly onislands than on continents (Brooks et al 2002 MillenniumEcosystem Assessment [MEA] 2005 Kier et al 2009) The smallpopulation sizes and ranges of island organisms and their uniquecharacteristics resulting from prolonged evolutionary isolationmake them particularly sensitive to anthropogenic change(Frankham 1998) All of these features make islands invaluablebut fragile and vulnerable arks of biodiversity

Island plant conservationists are thus faced with a huge globalconservation challenge Nevertheless contact among researchersand managers of threatened plants on different archipelagos hasbeen limited (for example in Macaronesia Caujape-Castells et al

2006 2007a b) although some local or regional surveys ofendangerment patterns on different oceanic archipelagos have beencarried out (Medail and Quezel 1997 Jaffre et al 1998 Broughtonand McAdam 2002 Sakai et al 2002 Kingston and Waldren 2005Strasberg et al 2005 Danton and Perrier 2006 Maunder et al2008 Reyes-Betancort et al 2008) Wider discussions of conserva-tion issues common to different island systems have happenedsporadically but researchers in different regions have rarelysynthesized their conclusions and global comparisons are lacking

We present here what we believe is the first comprehensivesynthesis of plant conservation issues across major oceanicarchipelagos For nine focal island groups (Azores MadeiraCanary Islands Cape Verde Hawaii Galapagos Juan FernandezMascarenes Seychelles) we review the threats to native floras andthe present state of conservation research and practice The ninearchipelagos belong to eight countries and three oceans AtlanticPacific and Indian We complement this with a literature reviewand an analysis of patterns of endangerment covering 62 additionalisland groups providing quantitative estimates of threats to islandfloras worldwide We compare across archipelagos the patterns ofendangerment threat factors and research results that have beensuccessfully incorporated into management practices We reviewthe present resources and weaknesses in particular archipelagosidentify individual and shared knowledge gaps and use these tosuggest priorities for future research We suggest how existingscientific knowledge may be used to identify additional knowledgegaps launch priority studies to generate the necessary informationand develop strategies to preserve island floras We also discussresults and experiences that extend beyond the local context Webelieve that beyond its local relevance plant conservation researchon oceanic islands can also serve as a model for continents and shedlight on general topics in ecology (Kueffer et al 2010 Kaiser-Bunbury et al 2010)

In particular we address the following questions

What is the level of endangerment of oceanic island vascularplants and how does it differ between island groups

What are the main threats to endemic vascular plants on

oceanic islands and how does the relevance of these factorsvary among island groups

ARTICLE IN PRESS


FiglsquolsquoMe

end

TabPhy

A

A

C

C

G

H

Ju

M

M

Se

An

(ie

Di

tha

isla

arch

em

som




Methods

Focal study system


Hawaii

Azores

Cape Verde


Galaacutepagos


5


Canaries


Hawaii

-10

-5

-15 0 5 10 15

F2 (2

162

)

Azores


Juan Fernaacutendez

-10 -5F1 (3593 )






zores 9 1343 6 615 2332 ca 22a ca 20

anaries 7 95 5 485 7545 ca 15 ca 40

ape Verde 9 576 ca 15 279 4033 2 02c

alapagos 14 1000 4 141 7900 2e 96

awaii 8f 3900 ca 11 128 16636 ca 27 ca 5g

an Fernandez 3 590 1 150 100 1 ca 90


ascarenes 3 665i 164 740 4528 21 25j

ychelles 4 9930i 45 50 235 7 16











Mascarenes

Seychelles




Ma ma h C d T $

8 025 2531 243018 (2006) ca 105 858832b TIA

206 112 3718 2025951 (2008) ca 269 9326116 TIA

76d 32d 2829 426998 (2008) ca 102 312880 TA

4 007 1700 ca 25000 (2006) ca 3 ca150000 TCA

51h 043 4205 1283388 (2007) ca 77 7627819 TAMi

58 10 1320 600 (2005) ca 6 ca 3000 ATG

18 5 1861 246689 (2007) ca 307 988692 TIA

10 3 3070 2061546 (2008) ca 455 1342839 TSM

65k 60l 915 81000 (2002) ca 345 ca 161000 TA


















ARTICLE IN PRESS








Cape Verde 1 (7) 82d 30 34 ca 43

Galapagos 7 (35) 180e 40 ca 35 ca 60

Hawaii 32 (288) 929f 624 90gh ca 53gh





Total 129 (518) ca 2897 ca 1854 ca 48 ca 54









focal archipelagos

Threats AZO CAN CA

Biological





Socio-economic


Tourism 9B 5C 6B







Expert survey










1 MED



ca 160








9B 5C 4C 9B 6C 1C12D 10B 6B 11D 11A 10D

3B 4C 3C 4C 3C 4B

2C 1C 2C 3C 1C 2B

B 7C 1C 10D 6B 10A 3B


12D 6C 12A 8D 12A 11D

3C 11C 13A 1C 7C 5B

11B 11B 7B 2C 4C 7B

B 3B 7C 5B 13A 8B 6B

10C 9C 9B 12A 9B 9B

3B 8C 11B 7B 13A 8B

C 8C 11C 8B 10B 5C 6B


ARTICLE IN PRESS





TOURISM

LACK OFMAN


INVASIVE ALIEN

VERTEBRATES

SMALL

AND

HABITATALTERATION

INVASIVE ALIEN

DECLINE IN ENDEMIC














INVASIVEALIEN


POPULATION SIZES

FRAGMENTATION

LOSTMUTUALISMS

PLANT DIVERSITY







ARTICLE IN PRESS



35302520151050

EN

25

20

15

10

5

0

C

R

1 2 3LOG1

1 2 3LOG1






Island size


Small [23] 0056

Large [14] 0011

(o00001)nnn

Isolation Index

5 o II o 80 [40] 0052

II 4 80 [13] 0173

(o00001)nnn

Geographic region





(0278)

Atlantic Ocean


Caribbean [15] 0053

(0062)

Pacific Ocean



(0089)





4 5 60AREA

4 5 60AREA



s analyzed

CR EN CR+EN

4741 11212 15953

4778 4452 9230

1660 1593 3254

(0181) (0044)n (0002)nn

2828 5326 8155

7374 7003 14377

(0063) (0263) (0102)

4135 4392 8527

2772 9240 12012

5507 3486 8993

5083 0106 5189

(0044)n (0002)nn (0831)

3382 3826 7244

2528 11392 13919

(0866) (0062) (0161)

1732 3773 5505

11943 6405 18348

(0006)nn (0212) (0009)nn




ARTICLE IN PRESS







future prospects












ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS


FiglsquolsquoMe

end

TabPhy

A

A

C

C

G

H

Ju

M

M

Se

An

(ie

Di

tha

isla

arch

em

som




Methods

Focal study system


Hawaii

Azores

Cape Verde


Galaacutepagos


5


Canaries


Hawaii

-10

-5

-15 0 5 10 15

F2 (2

162

)

Azores


Juan Fernaacutendez

-10 -5F1 (3593 )






zores 9 1343 6 615 2332 ca 22a ca 20

anaries 7 95 5 485 7545 ca 15 ca 40

ape Verde 9 576 ca 15 279 4033 2 02c

alapagos 14 1000 4 141 7900 2e 96

awaii 8f 3900 ca 11 128 16636 ca 27 ca 5g

an Fernandez 3 590 1 150 100 1 ca 90


ascarenes 3 665i 164 740 4528 21 25j

ychelles 4 9930i 45 50 235 7 16











Mascarenes

Seychelles




Ma ma h C d T $

8 025 2531 243018 (2006) ca 105 858832b TIA

206 112 3718 2025951 (2008) ca 269 9326116 TIA

76d 32d 2829 426998 (2008) ca 102 312880 TA

4 007 1700 ca 25000 (2006) ca 3 ca150000 TCA

51h 043 4205 1283388 (2007) ca 77 7627819 TAMi

58 10 1320 600 (2005) ca 6 ca 3000 ATG

18 5 1861 246689 (2007) ca 307 988692 TIA

10 3 3070 2061546 (2008) ca 455 1342839 TSM

65k 60l 915 81000 (2002) ca 345 ca 161000 TA


















ARTICLE IN PRESS








Cape Verde 1 (7) 82d 30 34 ca 43

Galapagos 7 (35) 180e 40 ca 35 ca 60

Hawaii 32 (288) 929f 624 90gh ca 53gh





Total 129 (518) ca 2897 ca 1854 ca 48 ca 54









focal archipelagos

Threats AZO CAN CA

Biological





Socio-economic


Tourism 9B 5C 6B







Expert survey










1 MED



ca 160








9B 5C 4C 9B 6C 1C12D 10B 6B 11D 11A 10D

3B 4C 3C 4C 3C 4B

2C 1C 2C 3C 1C 2B

B 7C 1C 10D 6B 10A 3B


12D 6C 12A 8D 12A 11D

3C 11C 13A 1C 7C 5B

11B 11B 7B 2C 4C 7B

B 3B 7C 5B 13A 8B 6B

10C 9C 9B 12A 9B 9B

3B 8C 11B 7B 13A 8B

C 8C 11C 8B 10B 5C 6B


ARTICLE IN PRESS





TOURISM

LACK OFMAN


INVASIVE ALIEN

VERTEBRATES

SMALL

AND

HABITATALTERATION

INVASIVE ALIEN

DECLINE IN ENDEMIC














INVASIVEALIEN


POPULATION SIZES

FRAGMENTATION

LOSTMUTUALISMS

PLANT DIVERSITY







ARTICLE IN PRESS



35302520151050

EN

25

20

15

10

5

0

C

R

1 2 3LOG1

1 2 3LOG1






Island size


Small [23] 0056

Large [14] 0011

(o00001)nnn

Isolation Index

5 o II o 80 [40] 0052

II 4 80 [13] 0173

(o00001)nnn

Geographic region





(0278)

Atlantic Ocean


Caribbean [15] 0053

(0062)

Pacific Ocean



(0089)





4 5 60AREA

4 5 60AREA



s analyzed

CR EN CR+EN

4741 11212 15953

4778 4452 9230

1660 1593 3254

(0181) (0044)n (0002)nn

2828 5326 8155

7374 7003 14377

(0063) (0263) (0102)

4135 4392 8527

2772 9240 12012

5507 3486 8993

5083 0106 5189

(0044)n (0002)nn (0831)

3382 3826 7244

2528 11392 13919

(0866) (0062) (0161)

1732 3773 5505

11943 6405 18348

(0006)nn (0212) (0009)nn




ARTICLE IN PRESS







future prospects












ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS








Cape Verde 1 (7) 82d 30 34 ca 43

Galapagos 7 (35) 180e 40 ca 35 ca 60

Hawaii 32 (288) 929f 624 90gh ca 53gh





Total 129 (518) ca 2897 ca 1854 ca 48 ca 54









focal archipelagos

Threats AZO CAN CA

Biological





Socio-economic


Tourism 9B 5C 6B







Expert survey










1 MED



ca 160








9B 5C 4C 9B 6C 1C12D 10B 6B 11D 11A 10D

3B 4C 3C 4C 3C 4B

2C 1C 2C 3C 1C 2B

B 7C 1C 10D 6B 10A 3B


12D 6C 12A 8D 12A 11D

3C 11C 13A 1C 7C 5B

11B 11B 7B 2C 4C 7B

B 3B 7C 5B 13A 8B 6B

10C 9C 9B 12A 9B 9B

3B 8C 11B 7B 13A 8B

C 8C 11C 8B 10B 5C 6B


ARTICLE IN PRESS





TOURISM

LACK OFMAN


INVASIVE ALIEN

VERTEBRATES

SMALL

AND

HABITATALTERATION

INVASIVE ALIEN

DECLINE IN ENDEMIC














INVASIVEALIEN


POPULATION SIZES

FRAGMENTATION

LOSTMUTUALISMS

PLANT DIVERSITY







ARTICLE IN PRESS



35302520151050

EN

25

20

15

10

5

0

C

R

1 2 3LOG1

1 2 3LOG1






Island size


Small [23] 0056

Large [14] 0011

(o00001)nnn

Isolation Index

5 o II o 80 [40] 0052

II 4 80 [13] 0173

(o00001)nnn

Geographic region





(0278)

Atlantic Ocean


Caribbean [15] 0053

(0062)

Pacific Ocean



(0089)





4 5 60AREA

4 5 60AREA



s analyzed

CR EN CR+EN

4741 11212 15953

4778 4452 9230

1660 1593 3254

(0181) (0044)n (0002)nn

2828 5326 8155

7374 7003 14377

(0063) (0263) (0102)

4135 4392 8527

2772 9240 12012

5507 3486 8993

5083 0106 5189

(0044)n (0002)nn (0831)

3382 3826 7244

2528 11392 13919

(0866) (0062) (0161)

1732 3773 5505

11943 6405 18348

(0006)nn (0212) (0009)nn




ARTICLE IN PRESS







future prospects












ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS





TOURISM

LACK OFMAN


INVASIVE ALIEN

VERTEBRATES

SMALL

AND

HABITATALTERATION

INVASIVE ALIEN

DECLINE IN ENDEMIC














INVASIVEALIEN


POPULATION SIZES

FRAGMENTATION

LOSTMUTUALISMS

PLANT DIVERSITY







ARTICLE IN PRESS



35302520151050

EN

25

20

15

10

5

0

C

R

1 2 3LOG1

1 2 3LOG1






Island size


Small [23] 0056

Large [14] 0011

(o00001)nnn

Isolation Index

5 o II o 80 [40] 0052

II 4 80 [13] 0173

(o00001)nnn

Geographic region





(0278)

Atlantic Ocean


Caribbean [15] 0053

(0062)

Pacific Ocean



(0089)





4 5 60AREA

4 5 60AREA



s analyzed

CR EN CR+EN

4741 11212 15953

4778 4452 9230

1660 1593 3254

(0181) (0044)n (0002)nn

2828 5326 8155

7374 7003 14377

(0063) (0263) (0102)

4135 4392 8527

2772 9240 12012

5507 3486 8993

5083 0106 5189

(0044)n (0002)nn (0831)

3382 3826 7244

2528 11392 13919

(0866) (0062) (0161)

1732 3773 5505

11943 6405 18348

(0006)nn (0212) (0009)nn




ARTICLE IN PRESS







future prospects












ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS



35302520151050

EN

25

20

15

10

5

0

C

R

1 2 3LOG1

1 2 3LOG1






Island size


Small [23] 0056

Large [14] 0011

(o00001)nnn

Isolation Index

5 o II o 80 [40] 0052

II 4 80 [13] 0173

(o00001)nnn

Geographic region





(0278)

Atlantic Ocean


Caribbean [15] 0053

(0062)

Pacific Ocean



(0089)





4 5 60AREA

4 5 60AREA



s analyzed

CR EN CR+EN

4741 11212 15953

4778 4452 9230

1660 1593 3254

(0181) (0044)n (0002)nn

2828 5326 8155

7374 7003 14377

(0063) (0263) (0102)

4135 4392 8527

2772 9240 12012

5507 3486 8993

5083 0106 5189

(0044)n (0002)nn (0831)

3382 3826 7244

2528 11392 13919

(0866) (0062) (0161)

1732 3773 5505

11943 6405 18348

(0006)nn (0212) (0009)nn




ARTICLE IN PRESS







future prospects












ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS







future prospects












ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS




islands













ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS











Lost mutualisms




ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS

















ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS



















ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS














ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS













First aid measures



ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS










Conclusions




ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS




Acknowledgements






Lack of consistent

financial support

Dispersal of

resources among

competing

institutionsteams

Material easily

reachable

Increase collabor

renowned foreig

institutions to b

already develope

infrastructure as

knowledge base

Greatly increase

and coordination

regional level


Some staff needs

updated training in

modern research

methods

Poor English

communication

skills of most

specialised staff

Dedication and


in staff

Good facilities for

research




and with other

Macaronesian

archipelagos





Review the chec

plants to publish

Open access of re

existing database

Create a virtual h

Better coordinati

among centres o

Set up a virtual p

all specialised lit


Appendix 1

See Table A1

Appendix 2

See Table A2




ation with

n investigation

enefit from

d

sets and

cooperation

of research at










plant propagation

techniques

Revise and enforce

legislation





nature






klist of wild

a Flora

searchers to all

s

erbarium

on of efforts

f research

df-library with

erature





their skills






research methods








Flora



existing laws












legal protection



devoted to the



banks) and to store





researchers

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS





government and

thus of

institutional

stability to carry

out basic research

Lack of training in

modern research

methods

Openness toward

collaboration with



local biologists



species



training



by responsible

institutions

Under-funded

Programme not

strategically

designed

Dedication of local

biologists




institutions

Survey monitoring




and protocols

More funding


Conflicts of

interests with

other stakeholders

Dedication of local

conservation staffs

Increase funding


groups national and


biologists





invasive plants)





global change y )



and deer









Juan

Fernan-

dez

No well curated

seed bank and no

DNA bank

Lack of one central

herbarium

Lack of trained

conservation

advisors for CONAF

staff

Poor success in

limiting areas

where cattle can

graze

Awareness of the



Concerted effort to


population on the


Programme to assist


invasives and in


personnel


of most endangered

species



herbarium


on mainland Chile






work





rabbits




native forest


eroded areas







Requirement to give






species










park lands




interact with CONAF

personnel


foundations foreign



Madeira amp

Selvagens

Lack of funds

Lack of human

resources



populations




conservation




species (molecular


Increase monitoring


alien species

A effective control


alien species




commitment

(except Reunion)

Most staff poorly



endangerment status

well known







native plants and

communities




mainly)


ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS




trained in ecology

and conservation

(except Reunion)

Lack of good

research facilities

(except Reunion)

Dedicated staff

Openness to

collaboration with





Reunion (2009)









restoration








eg in the National



Rodrigues




Rodrigues)


trained botanists

Although

government sees

the need for plant

conservation

activities and

research an NGO

(PCA) with little

human resources

and time capacity

struggles to be

effective

Plant Conservation

Research Agenda

published in 2007



institutions



research




on management of


and biosecurity

measures


institutions







collection



genetic studies





action

Research in plant-

environment inter-


interactions



conservation work



and species


existing laws









bases etc

Improve media and



successes





resource use etc




Total EN CR








Borneoy O Indian 7330990 16750 101 213



Comoros Indian 485 20213 1136 2 1



Cuba Caribbean 285 1058055 13193 59 22









ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS



Total EN CR




Java Indian 393 138794 1250 14 13

















Reunion Indian 730 25352 1234 3 5













Sulawesi Indian 388 1936405 1500 0 4

Sumatera Indian 170 4430658 11200 32 64















References















ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS

























































ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS

































































ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS






























































ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS


























































ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS











Introduction

Methods

Focal study system

Expert survey







Lost mutualisms







First aid measures

Conclusions

Acknowledgements

Appendix 1

Appendix 2

References

ARTICLE IN PRESS - UAc et... · ARTICLE IN PRESS Review ... e Department of Ecology and...

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