BIOGEOGRAPHY, ECOLOGY AND SYSTEMATICS

JOHN LESLIE DOWE

Australian Cen tre for Tropical Freshwater Research, James Cook Universi ty, Townville, Qld, Australia,

and the Montgomery Botanical Cen ter, Coral Gables, FL, USA

CIIII~~ C S I FtO

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© John Le!:'lit' Dowe 2010

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Natioilil! Lil~rary of Allstnliil. Call11ugl1ing-in-P\lblication entry

D,lwt', John L

Allslr<'lllan pillms: hiog<'o;;raphy, ecology ;:nd system.3th::s I Jolm Leslie Do\V~.

97806430'-)6158 (pbk.)

Inchldes index. Bibliography.

Palms - Australia. Palms - Australia - Identification. P<:tlms - Norfolk Isbnd - Identification.

p.,lms - Loru Howe Island - Identification, Palms - Christmas Island (Indian Ocean) - Identification.

584.50994

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Fwnt cover (cJo;:kwise from top left): Licllain n'11IL';ayi, LivistClln lorophyilfl, GI/lI!liliS llI(lti, Lh:.,isfonl I1Ilul/el'i, Cllr~'mt'lrio aC/lIlJi!ln!n. li.rdwlllapliofl1lX nlc,tillldme, Nypr.fruliIY!11S and Namnlll.'ya i1onu!lllbyi. Photos by the 1l1thor.

Back coyer (c1ock';!.ise frcl11 top left): Li;;';stOIlIl ,'idorin~, Bun~le Bungle Ranges, \·Vestern Austroli,,; Liel/IIM mmSllyi; Cnfflllws II/flli; Ullll.~padi)' J'IJl'fir!nills; and C(i/,yoin nl/;ocrtii. Photos by the author.

Set in Palatilio 9.5/13

Edited by ALirieml€ de Kretser, gig;hting Writing Cover ;md ted design by James Kelly

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Foreword

Despite the long ,Uld venemble tradition of rivalry b eMeen AU$lrcli .. ,md the UK it is obvious thatUl:l Ponunies areon

a losing wicket when it C.omes to plimt diversity. The thrilLs of my own encounters with \.,;Id Australia still r('SoMle today - the first tastes of sclerophyll vegetation on the Sydney sandstones: a formative toad trip through the Atherton Tablehmd!, and the lush. subtropical forests of the Lamington National Park, for example. r have scarcely

dipped my toe in the water, but such experiences can only leave one awt!Strudc and di!:>1inctly envious of the Mlual wonders thai r'\ ustralians can enjoy lm their doorsteps.

The pa lm fa mily makcs only a modest contribution to the Australian flora. but is exceptional nonclhclcss. No other regiQrull palm flora includes representatives o f all five palm subfamilies. It encapsulates flavours of both the

~<ll~iun zmd Paci fic palm Horns while including sevual specialities of its own. Australia contains memonb ie: pl'.hn-dominatffi lar.d~apes, such as the forests o f LiclUlia

ra lllSlllJi in famous !.ocalities such a~ Mission Beach.. Li"Jis­unUl 1Th111·ac On thf! Fhke River system in the Northern TI!r-1"i tory, or the v~st stil:"\d.'l of Kenna palins, Howf!P. forsl.eritna, on the off-shore island of Lord Howe. Significan t among

the endemics is Omr.iopsis, an andent-looking pahn whJSC

,

affinities CIS il. represen t<l IiVE of a lineage othelwise found

only in South Ameri-ca, MadagaS(Ca1" and Juan Fernandez

were detennined onI}' 25 years ago. Similarly dram~tk, the fox taU palm Wodyefia wa~ only described in B83, h avi.ng jusL· been discovered ,.mong spectacular grMite boulders in the Melville Range in Queensland. TI.la t such

finds can be made so recertly hints at the possibility of more great botan.ic:al discov~rie~ to cum!:' in Australia.

ft i:; perhaps surprising filal a comprehensive guide to the pZllms of Austril liil h ilS been Jacking until now, but

readers will not regret the wlit. In typical style, John Dowe

has tackled the subject in grmt dep th and detail, produ-:ing an accoun t SO thorough that it ill unlikely to be supen!8ied for many years to come. 1\ has nrrived at a time Whell the need fnt palm information has never been g loc3 lcr, (0 sen-·ice both horticultural appetires and much-needed con­ser~'ation agendas. This rich source of knowledge and beau tifuJ images is a milesb:me in the botanical histOl)' of Austrlllillllnd 11 fitting h·ibule to the wondef50f its paluso

1,",1illil"ml J. Baker HfJui of Palm ResfmrcJl

R.oyal R()umic GMdens, J<ew

Contents

Preface ix Climate change since t~e Late Cretaceous 21

Acknowledgments xi Co-occurrence of palm species in Australia and New Guinea 24

INTRODUCTION Summary 27

Economic botany 5 Systematic arrangement of fossil palm taxa

Endangerment and conservation from Australia and New Zealand 30

of Australian palms 5 4 DISTRIBUTION AND ECOLOGY 31

2 EARLY DOCUMENTATION OF Australian palms and their environments 31

AUSTRALIAN PALMS 7 Palms in the landscape 31 European plant:ollectors and botanists Phytogeographical regions in Australia 33

in Australia before settlement 7 Distribution patterns 33

Explorers, naturalists and botanists Remote species of Livistona 34 after settlement 3

Allan Cunningham 10 Remote global distribution: Oraniopsis 36

Carl Friedrich Phillip von Martius: Lord Howe Island 37

Historia N<ltur.1Iis Pa/marum 11 Ecological amplitude 37

Ferdinand von Mueller 11 Climate 39

Hermann WencDand and Oscar Dmde: Edaphk preferences 39 Pa/mae Ausiralasicae 11

Fire 3. George Bentham: Flora Australiensis 11

Fire and Uvistona 39 Frederick J\-\anson Bailey 11

Cyclones 42 Karel Domin 12

Adaptation of palm, to wind stres5 42 Odoardo Beccad 13

Altitude and elevatiomI ranges of palm species 45 Max Burret 13

Demography and popLlation dynamics 46 Post-1900 13

Population genetics 4" Recent contribctions to AustraIian palm botany 13

Other ecological aspects 49 Summary 13

Summa.I'Y 49

3 HISTORICAL BIOGEOGRAPHY 15 5 SYSTEMATIC ARRANGEMENT

The fossil record 15 OF THE AUSTRALIAN PALM flORA 51

Aust1'alian and New Zealand palm fossils 16 Phylogenetic relationships of Australian palms 51

vii

ContF.l1l.

F~ mily descriptio" 51 Archontop hoenicinae 110

Key to subfamilie; in Australia 54 Archontophoeni'X 1]1

B~ sselinjjnae 167 6 SUBFAMILY " CALAMOIDEAE 55

LepirJorrbachis rw ClilSsiric.Jtion md relationship!> 5; linospadidnae "" Cafaro"s 56

Lino5,OiIdix 1~1

7 SU8FAMllY 2: NyrOIDEAE 83 l<:!cc:uspadix 2C6

Classification md relationships 83 Howea 200

Nypa 83 Ptychospermatinae 214

Prychosperma 214 S SUBFAMILY 3, CORYPHOIDEAE 87 Carpentaria 213

Classification and relationships 8 ' Wodyetia 225

Llvlstoninae 83 Normanbya 218

Uvisfonil 8' 1{hopalostylldinae 233

Ucuala iJr, Rhopalostylis 233

Caryoteae 141 Hedyscepe 2J7

Caryom 141 Unplac~d Ar~ccac 2,0

Arengt! 14' I-Iydriasreie L.J.U

Coryphcilc 153 11 DOUBTFUL AND EXUUDED NAMES 251

Coryph.l 151

12 FIELD IDENTIFICATION OF AUSTRALIAN 9 SUBFAMILY 4, CEROXYLOIDEAE 159 PALM SPECIES 255

CI<Hsificlltion and relation5hips 159

Oralliopsis 159 Checklist of genera and species 259

Glossa ry 261 10 SUBFAMILY 5, ARECOIOEAE 16j

Abbreviations 2<7 Classification and rela tionships 165

Herbarium acronyms 2&9 Cocoseae 165

Cocos 166 References 271

Areceae 170 Index 2U3

'!iii

Preface

This book provides an updated and thorough systematic Jnd taxonomic treatment of thcAllstralian palm flora. ~hc

only other similarly broad treatment of Australian palms

was prepared over 130 years ago, by German botan:st:; Hermann Wendlan::l and Oscar Drude in 1875 in their Pa/mae AlIstmlasica~, published in the journal Linmea.

Ba~ed on the knowledge of the time, that early work described 26 specie:; and examined the palm flora from aspects of biogeogmphy, relationships and taxonomy. It provided the basil; br much llf OUT underst<Jnding of Aus­

tralian palms well hto the 20th century. Treatments such as Bentham's Flora AlLStralil!nsis of 1878 and F.M. Bailey's

Queensland Flom of 1902 provided timely synopses, but

did not address broader aspects of biogeography, relatxm­ships and dassiilcatlon. Early 20th-century palm botanisls such as Odoardo Becc81i and Max BUlTl"t proVided Se1111"

broader ex~mination of Australian palms, but most often in a context of revision~ and new species description:;. In

the later decades of the 20th century, interest in Australian palms was renewed throllgh the work of A.N. Rood, A.K.

Irvine, J.L. Dowe, D.L. Jones and F.B. Essig; revisions of

Atcl1olttopilOellix, C~1"yota, Hydl'Ulstele, Licunia, LiIlOSprriix, Livistona and Ptycilcsperma have been completed in recent

decades. Sixty species of Australian palms are recognised and.

discussed in lhis book. Thl" book lakes noll" of recen t

advances in biogeographic and phylogenetic resea:·ch,

which allow the Australian palm flora to be incish'ely

placed within a regional and global context. These as~cts aH examined in detail. AustraliaJ1 Palms includes expanded

and thorough descriplLons of genera and species, wiJh additional informalion on diSlributi.OJl, ecology and ~,.pi­

fiCfltion. The desctiptions of both generilll.nd sp~cies wer!'!

obtained £ollowingextensive field-work and examina:ion

of over 1600 specimens in herbaria inAustralia, 50uth~a5t Asia, Europe and the US. The author has !;tudied and col­

lected all mainland palm SpE'Cies in their natural habitats, ~nd has studied th)5e fr0111 off--sllore tenitories as (lJ1ti­

vated specimens in Australian botanic gardens such as

"

The Palmetuffi in Townsville, Flcckcr Botanic Gardens in

Calms and Mt Coot-lha Botanic Gardens in Blisbane, and

in Florida at the Fairchild Tropical Botanical Garden:; and

the Montgomery Botanical Center. This worUs the culmi­nation of over 20 years of research i11to Australian palms.

• STRUCTURE OF THE BOOK

This book is divided into two sections. The first section provides a broad introduction, with dctJiled SU111m~ries

ofbolanical history, historical biogeography, distribution and ecology. The second section deals with sy!;tl:!matic!;,

classification and taxonomy, providing assessment and description of taxa from tilt lellel of subfamily to sub;pe­

cies, ilrranged according to the m05t rLXent classification

of palms in Gmcra Palnumm: Tht! Evo/tltion mId Classlica­nOll afPalms (Dransfield et d. 2(08). Each species is clEarly

illustrated with images of bbit, leavl:!o, flowers, fruit and relevant djag"o~tic features. The fmal chapter offers El key for the field identificaHon cf Australian palm species. TIle

book concludes willl a glossary.

• METHODS

The taxonomic history of each species was invcstiglted and nomcnclarnrc imd typification were revil""i,.ied_ Proto­

logues for all taxa were sourced and SClutiniseci for

validity of publication, ty?fication and adherence to the

appropriate articles in the Inter;ullioll!1[ Code of Bota 1lica/ .I\,'omellclahne (McNeill et a:. 2006). Type spedml2lls were

located at numerous herbal"ia, ilnd imilges of at least one sheet (of those with multiple sheets) are included. Where

types had been lost, misapplied or inappropriately desig­

nated, new typification is presented. In somr= cases, typi­fication is dl:!hlS'llatro for t:I:v< first time. Species descriptlons are based on the examinaion of abou t 1600 herbaTiu111

spcx:imens and living palms in their natural habitats and

cultivation. Descriptions ale based on mature plants. For

Preface

many mC:!"dSuremenls, ruch as srem height and leaf length. only the upper range is provided. O therwise, the range 0: siz@srecordcdIorsome orga.ns,wheresizeismore ccitico\' is provid ed. Phyloger.etic 3s.<;umptioru ami recOllstruC· Hons are based on the most recent anaJyS€s as referenced in the text, mostly accepting those presented in Generr PalmaTurn (Drans field f!t al. 2008). Au thor abbreviati.ons foUow Brummitt and PuwelJ (1992), hCIDariU1J1 acronyms

,

an> accnrding to HOlmgren and [·Iolmgren (1998), joum~1 abbrevi;ttion6 follow lIri dsnn md Smith (1991) and publi4

cation abbreviations follow Stafleu and Cowan (1976-98).

Spe<:imcn.s examined in<:ompHing species descripHons and associated with typificulion may be viewed as ~n Exed spreadslwet ... t htlp: / 1v.~'I'.'l.montgomerybotanicll .org/ Pages/Researeh.htm.