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Paper mulberry (Broussonetia papyrifera) as a commensal model for human mobility in Oceania: anthropological, botanical and genetic considerations | Papers by Andrea Seelenfreund et al.

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Paper mulberry (Broussonetia papyrifera (L.) Vent.) was one of the most widely distributed crop species in prehistoric Oceania, occurring from continental East Asia to the Polynesian islands. Its broad distribution is largely due to human-mediated
   PLEASE SCROLL DOWN FOR ARTICLE This article was downloaded by: [University of Otago]  On: 19 February 2011 Access details: Access Details: [subscription number 927826668]  Publisher Taylor & Francis  Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK New Zealand Journal of Botany Publication details, including instructions for authors and subscription information: Paper mulberry ( Broussonetia papyrifera ) as a commensal model forhuman mobility in Oceania: anthropological, botanical and geneticconsiderations D. Seelenfreund a ; AC Clarke b ; N. Oyanedel a ; R. Piña ac ; S. Lobos a ; EA Matisoo-Smith b ; A. Seelenfreund da Department of Biochemistry and Molecular Biology, Universidad de Chile, Santiago, Chile b Department of Anatomy and Structural Biology and Allan Wilson Centre for Molecular Ecology andEvolution, University of Otago, Dunedin, New Zealand c Biology Department, UniversidadMetropolitana de Ciencias de la Educación, Santiago, Chile d Escuela de Antropología, UniversidadAcademia de Humanismo Cristiano, Santiago, ChileFirst published on: 22 December 2010 To cite this Article Seelenfreund, D. , Clarke, AC , Oyanedel, N. , Piña, R. , Lobos, S. , Matisoo-Smith, EA andSeelenfreund, A.(2010) 'Paper mulberry ( Broussonetia papyrifera  ) as a commensal model for human mobility in Oceania:anthropological, botanical and genetic considerations', New Zealand Journal of Botany, 48: 3,, First published on: 22December 2010 (iFirst) To link to this Article: DOI: 10.1080/0028825X.2010.520323 URL: Full terms and conditions of use: article may be used for research, teaching and private study purposes. Any substantial orsystematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply ordistribution in any form to anyone is expressly forbidden.The publisher does not give any warranty express or implied or make any representation that the contentswill be complete or accurate or up to date. The accuracy of any instructions, formulae and drug dosesshould be independently verified with primary sources. The publisher shall not be liable for any loss,actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directlyor indirectly in connection with or arising out of the use of this material.  Paper mulberry ( Broussonetia papyrifera ) as a commensal model for humanmobility in Oceania: anthropological, botanical and genetic considerations D Seelenfreund a , AC Clarke b , N Oyanedel a , R Pin ˜a a,c , S Lobos a , EA Matisoo-Smith b and A Seelenfreund d * a Department of Biochemistry and Molecular Biology, Universidad de Chile, Santiago, Chile; b Department of Anatomy and Structural Biology and Allan Wilson Centre for Molecular Ecology and Evolution, University of Otago, Dunedin, New Zealand; c Biology Department, Universidad Metropolitana de Ciencias de la Educacio´n,Santiago, Chile; d  Escuela de Antropologı´a, Universidad Academia de Humanismo Cristiano, Santiago, Chile ( Received 30 May 2010; final version received 30 August 2010 )Paper mulberry ( Broussonetia papyrifera (L.) Vent.) was one of the most widely distributed cropspecies in prehistoric Oceania, occurring from continental East Asia to the Polynesian islands. Itsbroad distribution is largely due to human-mediated dispersal during colonization of the islandsof Near and Remote Oceania. We explore the potential for analyses of genetic variation in papermulberry and the value of such data for the development of a new commensal model species forreconstructing patterns of human mobility in Oceania. We introduce and discuss paper mulberryas another commensal species and outline key features for its contribution to the understandingof human migration and post-colonization interaction. Here, we describe some of the extant B. papyrifera populations in Remote Oceania and Taiwan that were sampled for initial studies.We argue that the unique characteristics of this species and its importance in ancient Pacificisland societies may provide the opportunity to collect valuable genetic data with which we canaddress several key questions in Pacific prehistory. Keywords: Broussonetia papyrifera ; paper mulberry; Polynesia; migration; genetic markers;Pacific prehistory; Lapita dispersal; Rapa Nui srcins Introduction Paper mulberry ( Broussonetia papyrifera (L.)Vent.; Moraceae) is a dioecious, perennialspecies occurring naturally in Taiwan andSoutheast Asia. It is an economically importantplant, and in its native range is used for makingpaper, rope and feeding domestic animalssuch as cattle. It is also used as a medicinaland ornamental plant (Matthews 1996). Thedispersal of paper mulberry into the Pacific andultimately the Polynesian Triangle is thought tobe associated with the Austronesian expansionand later the spread of the Lapita culturalcomplex (Matthews 1996). The dispersal of paper mulberry and other plant and animalspecies is believed to be part of the Lapitapeoples’ colonizing strategy of using ‘trans-ported landscapes’. The use of this strategy, itis believed, was to maximize the likelihood of survival as they moved out into the Pacificisland environments which had increasinglylimited terrestrial resources (Kirch 2000).Although we can find no record of archaeo-logical remains of paper mulberry, such aspreserved wood or pollen, being identified inany Lapita site, the association of this plantwith Lapita peoples is assumed based on itshistoric distribution and significance in Pacificsocieties. Paper mulberry was distributed asfar as Hawai‘i, Rapa Nui (Easter Island) and *Corresponding author. Email: New Zealand Journal of Botany 2010, 1  Á  17, iFirst article ISSN 0028-825X print/ISSN 1175-8643 online # 2010 The Royal Society of New ZealandDOI: 10.1080/0028825X.2010.520323  D o w nl o ad ed  B y : [ U ni v e r si t y  of  O t a g o]  A t : 19 :00 19  F eb r u a r y 2011  New Zealand in Polynesian prehistory. (Wehave chosen to use the name Rapa Nui, whichis the name used and preferred by the islanderstoday.)Paper mulberry is called lu-a-shu in Taiwan, ai masi  in Fiji, lau‘a in Samoa, hiapo in Tongaand lafi  in Futuna. In Eastern Polynesia itscommon name is indicative of its shared recenthistory: wauke in Hawai‘i, aute in Tahiti andNew Zealand, ‘ ute in the Marquesas and mahute on Rapa Nui (Matthews 1996; Neich& Pendergrast 1997). However, the relationshipbetween these Eastern Polynesian cognates andterms further west remains unclear. Through-out Polynesia, the bark of paper mulberry isused to make textiles, known generally as tapa.However, the name for barkcloth in Samoais siapo , and in Tonga barkcloth is known as ngatu, whereas in Fiji it is generally knownas masi  (Ewin 2009). Green (1979) reconstructsthe Proto Polynesian word for the papermulberry plant as * siapo , which is clearlyrelated to the name still used in Tonga for theplant and in Samoa, for the barkcloth.In the past, paper mulberry was used for themanufacture of cloaks, skirts, loin cloths andritual gifts (R Kennedy 1934; Kooijman 1972;Bell 1988). The making of barkcloth and itsuse and role in Polynesian culture were bothwidespread and significant, and remain so inlocationssuchasTonga,andsomeislandsinFiji.In those islands where barkcloth productionhas ceased (due, in part, to the availability of cheaper industrial made cloth such as cotton),paper mulberry has generally gone extinct and,furthermore, is often deliberately eliminated(A Seelenfreund, personal observation). Despitethis, the association of paper mulberry withmany economic, political and ritual uses makesit one of the most important cultivated plants of Oceania (Matthews 1996).Richer understandings of human mobility inOceaniaareincreasinglybeingobtainedthroughmolecular studies of associated commensalplants, animals and pathogens. In an Oceanicsense, commensals are those species that weremovedbyhumans(usuallydeliberately)betweenislands, often as a part of expansion and settle-ment. The use of commensal models for humanmobility include analyses of molecular datausing population genetics, phylogenetic andphylogeographic methods. These models arecomplemented by (and necessarily include) re-levant biological, linguistic and archeologicalinformation, or Kirsh and Green’s ‘triangula-tion’ approach (2001). The genetic relationshipsare used to infer patterns of plant and animaldispersal and therefore indicate patterns of hu-man mobility. Whereas human genetic researchreveals patterns of migration and settlement,studies of commensal species potentially gofurther by revealing additional patterns of hu-man activity, such as trade, that may notnecessarily be associated with settlement. Com-mensaldatacanalsoenhanceourunderstandingof agricultural practices.A number of plant and animal species inthe Pacific have been used to model humanmobility. The integration of these studies, alongwith analysis of human genetic variation, hassignificantly enriched our understanding of prehistoric human-mediated dispersal andcolonization histories in the Pacific. Commensalanimal species studied include the Pacific rat( Rattus exulans ; Matisoo-Smith & Robins2004), pig ( Sus scrofa ; Larson et al. 2007),chicken ( Gallus gallus ; Storey et al. 2007), thelizard ( Lipunia noctua ; Austin 1999) and theland snail ( Partula hyalina ; Lee et al. 2007).Commensal plant species studied include bread-fruit ( Artocarpus spp.; Zerega et al. 2004), bottlegourd ( Lagenaria siceraria ; Clarke et al.2006), ti ( Cordyline fruticosa ; Hinkle 2007),yams ( Dioscorea spp.; Lebot et al. 1998), taro( Colocasia esculenta ; Caillon et al. 2006) andbananas ( Musa spp.; J Kennedy 2008). All of these have provided important new informationregarding population srcins and interactions inthe Pacific.Despite improved understandings of thedirection and timing of human settlementof Remote Oceania, there remain many2 D Seelenfreund  et al.  D o w nl o ad ed  B y : [ U ni v e r si t y  of  O t a g o]  A t : 19 :00 19  F eb r u a r y 2011  unanswered questions and topics of debate. Forexample, it has recently been suggested thatpost-Lapita movements from Island SoutheastAsia through the low islands of the Carolines,Kiribati and Tuvalu into West Polynesia mayexplain some of the cultural, biological andgenetic diversity observed within RemoteOceania in terms of people and their commensalplants and animals (Addison & Matisoo-Smith2010). There also remain questions about thesettlement and interaction histories of specificislands, particularly Rapa Nui. While muchrecent debate has focused on dating the initialhuman colonization of Rapa Nui (Hunt & Lipo2006) and the subsequent impact of humanarrival (Diamond 2006), the specific origin(s)of the founding population or populations andthe process of colonization are issues that havenot yet been resolved (Green 2000a, 2005;Martinsson-Wallin & Crockford 2002).There is the potential to explore these andother key questions in Pacific prehistory byanalysing commensal species within an inte-gratedframeworkofbiological,anthropologicaland genetic data. Paper mulberry may be parti-cularlyusefulasacommensalmodelspecies.Itisvery widely distributed in Polynesia and, of the  70 plant species prehistorically introducedinto Polynesia by humans (Whistler 1991; Yen1991), paper mulberry was one of only  20plant species that was successfully introduced toRapa Nui (Me ´traux 1971; Orliac & Orliac 1998)and one of only six plant species introduced toNew Zealand (Sykes 1969; Matthews 1996).Here we present our preliminary explorationof the use of paper mulberry as a commensalmodel species. We identify key questions inPacificprehistoryforwhichgeneticdataofpapermulberry could be most valuable and identifywhat data need to be obtained in the futureto fully utilize paper mulberry as such amodel species. We hope this approach will beusefulforothersundertakingcommensalspeciesresearch and will provide a framework forunderstanding the dispersal of paper mulberryin Oceania. The distribution and biology of paper mulberry The genus Broussonetia belongs to the largefamily Moraceae, which is widely distributedin tropical, sub-tropical and, to a lesser extent,temperate regions (Mauseth 1991; Sharma1993). Broussonetia (along with the genus Morus ) belong to the large and morphologicallydiverse tribe Moreae, which has a wide geo-graphic distribution in Asia and Island South-east Asia (Zerega et al. 2005). Broussonetia papyrifera is a deciduous dioecious plant, withserrated leaves ranging from 8 to 20cm long.Leaf shape is highly variable, even on the sameplant,rangingfromovatetodeeplythree-tofive-lobed, with the lobes being strongly lacerate inthe most dissected leaves. Leaves are denselytomentose,especiallyontheabaxialsurface.Thepetiole is nearly as long as the lamina. Plantvarieties are distinguished by leaf shape, and onsome islands these varieties are designatedby different names (e.g. Hawai‘i) (B Mulloy,personal communication). Paper mulberry canbe easily distinguished from other mulberryspecies by the milky sap and the fuzzy surfaceof the leaves. Plants can be multiplied by bothseeds and cuttings. Its habit of producing rootsuckers, by which it can become a nuisance, isanotherdistinguishingfeatureofpapermulberry(Whistler & Elevitch 2006). Although it is nativeto temperate climates, it is highly adaptable todifferentlatitudes(Andrews1940)andsoiltypes,especially volcanic (Whistler & Elevitch 2006).Paper mulberry is native to southern China,Japan and Taiwan, and was introduced byAustronesian settlers to the islands of the Pacificas far east as Rapa Nui, north to Hawai‘i andsouth to New Zealand, and even to such remoteislandsasPitcairn(Whistler &Elevitch2006).Inthe Pacific islands, paper mulberry is normallyonly seen as a small tree or shrub and is usuallyprunedtokeepitunder3mtall,whichmayaffectthe species’ ability to flower in cultivation.Indeed, Florence (2004) and Matthews (1996),state that flowering is unknown or infrequentovermostofthetree’sdistributioninthePacific.The lack of flowers in Pacific individuals may Paper mulberry, a new commensal model for human mobility in Oceania 3  D o w nl o ad ed  B y : [ U ni v e r si t y  of  O t a g o]  A t : 19 :00 19  F eb r u a r y 2011  also limit the ability to diagnose sex. Whistler &Elevitch (2006) and Whistler (2009) state thatonlymaletreesoccurinPolynesia,butRapaNuipopulations of  B. papyrifera have been seen toproduce fruits and therefore are female (A.Seelenfreund unpublished observations; Fig. 1).It is not known, however, if these are fertile(sterile fruits have been observed in Hawai‘i(Meilleur et al. 1997)). Under cultivation in thePacific, propagation is mainly through rootshoots, which are produced freely. Althoughthe red fruits are consistent with dispersal bybirds in the plant’s native range, there is noevidence for birds being an effective vector forthe interisland dispersal of paper mulberry inthe Pacific. Human-mediated propagationfrom seeds has not been recorded in the Pacific(plants not reaching flowering age beforebeing harvested for their bark is probably alsoa factor; Florence 1997). It is not clear whetherabandoned (feral) plants may sometimes repro-duce sexually and whether there is gene flowwith cultivated plants that have been allowedto flower. In terms of dispersal, there may havebeen some natural dispersal within an island(either through vegetative or sexual reproduc-tion), but dispersal between islands (at leastin Remote Oceania) was certainly humanmediated. It is possible that both sexes of papermulberry occur in Remote Oceania, althoughwhether this situation has varied through time(prehistory and the historical period) and space(someislandsandnotothers)remainsunknown.If both sexes are present in the same place andtime, this would have obvious implications forwhether breeding populations can establish, andthereforethetypeofbiologicalpatternswemightexpect to see. Fig. 1 (A,B) Paper mulberry ( Broussonetia papyrifera ) plant from Rapa Nui (Easter Island) with femaleinflorescence. (C) Syncarp on a plant from Rapa Nui. (D) Feral plants, in the foreground, growing betweenboulders inside Rano Kao volcano, Rapa Nui. 4 D Seelenfreund  et al.  D o w nl o ad ed  B y : [ U ni v e r si t y  of  O t a g o]  A t : 19 :00 19  F eb r u a r y 2011
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