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OVERVIEW of the GRASSLANDS of OCEANIA 3R3 Gillison, A.N. (1994 *Oo"*i"* of the Gtasslands of Oceania. In Ihtwai Taprate Grasslatf,s: Evasia, Africa, tuuth Arerica and Oreania, etl. R.T. Corplantl In firaslestens of the l/orJd ed-in-chief D.ll. cmdalL, 88. pp. 303-313. Elserrier Scientific hrblishing Co., Amsterdam. Chapterl2 OVERVIEWOF THE GRASSLANDSOF OCEANIA A.N. CILLISON INT'RODUCTION foundedby variationin climatc,soils, tcrrain, is- land size and patternsof land use. Some of the The grasslandsof Oceania arc distributedin land areastreated here are outside Oceaniascnsrl fragmcntcd units over a region extending some stricto and includc "outlicrs" such as Hawaii, the 105degrees longitudinally [from Australia(120'E) Philippinesand the sub-Antarcticislands. These to Pitcairn Island (13,5'W)]and 78 degreeslat- havebeen includcd to providc a morc appropriatc itudinally Ifrom thc Northern Mariana Islands regional biogeographicframework. A broad indi- (23"N)to the sub-Antarcticislands (55'S)1. Within cation of the regional distribution of major grass this vast territory, the land arca of approximately tribcsis givcnin Thble12.1. 8 509400 km2 supports a human population of Comparativestudies of grasslandswithin Ocea- about 2-5060000,the area per head ranging from nia are made complex by extreme variation in nearly 0.3 ha in Tirvalu zrnd Nauru to 14 ha in physical environmcnts: although semiarid Aus- Papua Ncw Guinea and 44 ha in Australia.Al- tralia containsextensive areas of "pure" grassland, though thc proportion of land which has been the south-wcstPacific island grasslands are mostly convertedto grasslandtcnds to vary directly with fragmentedmosaics distributed along locally vari- populationpressure, clirect relationships are con- able - often steep - physical environmental TABLE I2.I Compositionof the grassflora (?, contributcd by cach major tribc) in sclectedlocalitics in and near Oceania.compared with the gkrbal avcragcl Locality Agrostcac Andropogoncae Avcncae Eragrostoac Fcstuccire Paniceae Others Ilawaii (belowr'. 1300n) 5.'7 | 1.4 6.8 8.{) 17.0 35.2 t-5.9 Luzon, Philippincs 2.3 33.7 l.l 7.4 2.9 -33.7 18.9 PapuaNew Guinea 5.5 29.6 1.6 8.3 11 22.2 22..2 Kimbcrlcv Dist., W Austrahir 28.2 10.9 9.I 3.6 29.1 19.I De Grel'. Wcstern Australia l.l 1-5.7 r0.l 22.5 27.(l 23.6 South-wcstcrnQueensland 1.1 10.6 8.0 2l .3 1.8 18.6 35.3 SouthernS. Alrstralia 10.,1 12.3 2.0 26.0 9.7 3-s.I Thsmania 21.2 3.0 14.1 2.0 :1.0 28.4 South-westernNcw Zcaland 25.3 29.3 30.7 | 1.7 Ave rage u.,+ 15.2 10.8 9.0 13.0 19.9 23.7 Normal distribution spectrum(global) 8.2 I 1.9 6.3 IJ.I r6.5 2.4.3 rThc valucs usedarc from Hartley (1950),cxccpt for New Guinea,which arc basedon listsof grassgencra extractcdfrom Henty (1969).Hawaii and the Philippincsarc not parts ol Oceania.tensu stri(to. but are of intcrcst for biogcographiccomprrisr)n. 303 304 A.N. GII-LISON gradientswhich may be variously associatedwith Antarctic islands.ln thc tropical Pacific, on thc woody species.Similarly, in thc sub-Antarcfic is- othcr hand, thc closcsterlilnitics of New Zealand lands,grasslands are componentsof ecosystems are with the upper montane and sLrbalpincfbrma- rather than ccosystcmsin their own right. tionsof New Guinea. The fbllowing filur chirptersare concerncdwith Partly becauscof a sparsefossil record, thc ori- the grasslandsof Oceania:Australia (Chapter l3), gin of the gcne pool from which Occanic grass- Ncw Zealand (Chaptcr 14), the strr-Antarctic is- lands havc been dcrivcd is unclcar. Thkhtajan lands (Chapter 15) and the south-west Pacific (19{19)and othcrs have argucd a case for likely (Chapter16.) centrcs of origin for thc Attgiospcrms,one focal area being north-casternAustralia - a hypothesis which tends to be supported by thc more rcccnt ORIGINSAND AFFINITIBS findingso1' Audlcy-Charles (1987). A numericerl analysisof world grassgencra by Simon and Ja- Uncertainty surrounds the origins of grassland cobs(1990) postulates that, with the exccptionof in Oceania.Whatever the pre-humanevtllutionary the pooids,the currently recognize.lgroups of thc determinants.the close interaction between man Poaceacarc Condwanan in origin. Thc analysis and grasslandin both Australia (up to 40 000ycars by Simon and Jacobs(1990) has dcmonstratcd or r.rlorewith fire) and Papua New Guinea (at closeaffinitics between thc tropical Paciflcgrasses least 9000 ycars with slash-and-burnagriculture) ernd those of thc cold tempcratc (mesotherm/ suggcstsit is logical to include humans as a "nat- microthcrm) oceauic islands,whercas grasslands ural", albeit reccnt, evolutionary component for of microthcrm/mesothermAustralia and New thcseareas (Cillison, 1972). Zcaland have closer ties with Hawaii and the As with large arcas of south-casternAsia and Antarctic zonc. The affinitiesof thc sub-Antarctic the Philippines,in recent times many grasslands islandgrasslands with SouthAnterica (Gre mmen, have becn initiated by slash-and-burnagriculture 19t12;scc also Chapter 15) are also reflectcdirr and by fire-bascdhunting practiccs.ln Australia, thc findingsof Simon and Jacttbs,whcrc, at thc before thc arrival of European man in thc 1780s, genericlevel, therc are evidcnt links betweentrop- aboriginal huntcrs manipulatedthe spatial extcnt ical Asia, Australia and parts of the western Pa- and structural and floristic composition of many cillc. woodland savannasand grasslandsby "patch"- Although providing a useful lbcus firr regional burning in ordcr Io c()nccntralcgr()ups of grass- geographic studies. phytogcographiczrnalyses ol eating animals such as kangarclosand wallabies. grass gcncra alonc are inappropriatc for cctl- Although the use of hrc tbr hunting is also logical intcrpretation.Therc is a nced ttl take widespread in Melanesia and Polynesia,thcre is into accountvariability in cnvironmentas well as no parallcl in Australia fbr aboriginal slash-and- in plant functions and floristics in orclcr to pro- burn subsistenccagriculture. vide clearcr insights into evolution zrnddevelop- This background, together with the uncertain mcnt and into responscsto envirttnmentalchange. origin of many so-calledexotics, also crcates prob- Throughout Oceaniathere are "grasslands"which lems in dcfining"naturalness". There is continu- arc closely associatedrvith, or sttmetimcs dom- ing debatc about thc origins of the New Zealand inirtcd by, cypcraceousor other graminoids.Thc alpine and grasslandbiotas that have developed pervasivedistributicln patterns of thesegraminoids only recently and continuc to evolve rapidly (scc and their close ecologicaltics with true grasses Chapter l4). The New Zealandgrassland vcgeta- suggestthat they should be included in studics tion dcveloped following a major orogerryin thc involvinggrasslands. Pliocene and Late Tertiary and was latcr subject Instcad of cclncentratingsolely on grass taxa to limiting climatcsunder Pleistoccneglaciation. (c.g. genera),somc of thc questionsconcerning The generallack of drought in New Zealand, and grasslandorigins may be bctter focuscdusing mul- the relatively cool oceanic climate, distinguishit tivariatc analytical techniqucs involving identifi- from the continentalenvironment of Australiaand able, rcsponsc-basedattributes of graminoidsas givc it a closeraffinity with Thsmaniaand the sub- a whole. Functional groups in granrinoiclscan bc 305 OVERVIEWOF THE GRASSLANDS OF OCEANIA the islandsin the sub-Antarcticcon- characterized in part by specific forms of acid Miocene.For likely that the climate has remained decarboxylationand associatedenzymic pathways vergenceit is past 10000 years (see (Prendergast,1989) and by other functional char- relativelyconstant over the acteristics(Gillison, 1988).By examiningthe spa- Chapter15). tial and temporal distribution of these functional groups along defined physicalenvironmental gra- VEGETATION dients,potentially useful models cou,d be derivcd which woulcl assistin interpreting graminoid ori- Bioclimate classification gins. For Australia and the larger islands (New Zealancl, New Guinea) and parts of the sub- Within Oceania, present-day climates rangc Antarctic, various authors have argued that his- from extremes of seasonal tropical lowland on torical climatic fluctuations,and in particular os- the one hand and alpine on the other to cold cillatory glaciation, have profoundly influenced non-seasonalregimes in the sub-Antarctic' Be- the establishmentand maintenanceof grasslands' there are broad climatic similaritiesbetween Periglacialactivity in parts of the sub-Antarctic cause and low-altitude/high- (e.g. Macquarie Island) also affectsthe establish- high-altitude/low-latitude latitude domains,the use of terms such as "trop- ment of grasses(l-offler et al., 1983; see also ical" and "temperate" can misleadwhen compar- Chapter 15). These processesfuel speculation ing grasslandclimates between regions (Oliver, about grasslandorigins. Apart from geomorphic processeslgrassland domains have a broad his- 1919). The bioclimatic framework used in Chapter 16 iory of expansionand maintenanceby man-made (see also Gillison, 1983) to compare the grass- fire, except perhaps in the sub-Antarcticislands' lands of the south-westPacific has been extended In New Zealand it is argued that the origin of following discussionto apply to Oceania grasslandsfollowing the arrivalof early Polynesian in the categoriesof havebeen accom- as a whole. This is based on broad N settlersover 1000years ago may the whole plant to and not be purely py- optimal growth responsesof I panied by climate change, of defined thermal domains.The use of a response- z rogenic.In PapuaNew Guinea, the connection
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