Paper mulberry (Broussonetia papyrifera) as a commensal model for human mobility in Oceania: anthropological, botanical and genetic considerations

New Zealand Journal of Botany Vol. 48, Nos. 3
4, September
December 2010, 231
247

Paper mulberry (Broussonetia papyrifera) as a commensal model for human mobility in Oceania: anthropological, botanical and genetic considerations

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D Seelenfreunda, AC Clarkeb, N Oyanedela, R Pin˜aa,c, S Lobosa, EA Matisoo-Smithb and A Seelenfreundd* a Department of Biochemistry and Molecular Biology, Universidad de Chile, Santiago, Chile; bDepartment of Anatomy and Structural Biology and Allan Wilson Centre for Molecular Ecology and Evolution, University of Otago, Dunedin, New Zealand; cBiology Department, Universidad Metropolitana de Ciencias de la Educacio´n, Santiago, Chile; dEscuela 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 crop species in prehistoric Oceania, occurring from continental East Asia to the Polynesian islands. Its broad distribution is largely due to human-mediated dispersal during colonization of the islands of Near and Remote Oceania. We explore the potential for analyses of genetic variation in paper mulberry and the value of such data for the development of a new commensal model species for reconstructing patterns of human mobility in Oceania. We introduce and discuss paper mulberry as another commensal species and outline key features for its contribution to the understanding of 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 Pacific island societies may provide the opportunity to collect valuable genetic data with which we can address several key questions in Pacific prehistory. Keywords: Broussonetia papyrifera; paper mulberry; Polynesia; migration; genetic markers; Pacific prehistory; Lapita dispersal; Rapa Nui origins

Introduction Paper mulberry (Broussonetia papyrifera (L.) Vent.; Moraceae) is a dioecious, perennial species occurring naturally in Taiwan and Southeast Asia. It is an economically important plant, and in its native range is used for making paper, rope and feeding domestic animals such as cattle. It is also used as a medicinal and ornamental plant (Matthews 1996). The dispersal of paper mulberry into the Pacific and ultimately the Polynesian Triangle is thought to be associated with the Austronesian expansion and later the spread of the Lapita cultural complex (Matthews 1996). The dispersal of paper mulberry and other plant and animal

species is believed to be part of the Lapita peoples’ colonizing strategy of using ‘transported landscapes’. The use of this strategy, it is believed, was to maximize the likelihood of survival as they moved out into the Pacific island environments which had increasingly limited terrestrial resources (Kirch 2000). Although we can find no record of archaeological remains of paper mulberry, such as preserved wood or pollen, being identified in any Lapita site, the association of this plant with Lapita peoples is assumed based on its historic distribution and significance in Pacific societies. Paper mulberry was distributed as far as Hawai‘i, Rapa Nui (Easter Island) and

*Corresponding author. Email: [email protected] ISSN 0028-825X print/ISSN 1175-8643 online # 2010 The Royal Society of New Zealand DOI: 10.1080/0028825X.2010.520323 http://www.informaworld.com

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232 D Seelenfreund et al. New Zealand in Polynesian prehistory. (We have chosen to use the name Rapa Nui, which is the name used and preferred by the islanders today.) Paper mulberry is called lu-a-shu in Taiwan, ai masi in Fiji, lau‘a in Samoa, hiapo in Tonga and lafi in Futuna. In Eastern Polynesia its common name is indicative of its shared recent history: wauke in Hawai‘i, aute in Tahiti and New Zealand, ‘ute in the Marquesas and mahute on Rapa Nui (Matthews 1996; Neich & Pendergrast 1997). However, the relationship between these Eastern Polynesian cognates and terms further west remains unclear. Throughout Polynesia, the bark of paper mulberry is used to make textiles, known generally as tapa. However, the name for barkcloth in Samoa is siapo, and in Tonga barkcloth is known as ngatu, whereas in Fiji it is generally known as masi (Ewin 2009). Green (1979) reconstructs the Proto Polynesian word for the paper mulberry plant as *siapo, which is clearly related to the name still used in Tonga for the plant and in Samoa, for the barkcloth. In the past, paper mulberry was used for the manufacture of cloaks, skirts, loin cloths and ritual gifts (R Kennedy 1934; Kooijman 1972; Bell 1988). The making of barkcloth and its use and role in Polynesian culture were both widespread and significant, and remain so in locations such as Tonga, and some islands in Fiji. In those islands where barkcloth production has 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). Despite this, the association of paper mulberry with many economic, political and ritual uses makes it one of the most important cultivated plants of Oceania (Matthews 1996). Richer understandings of human mobility in Oceania are increasingly being obtained through molecular studies of associated commensal plants, animals and pathogens. In an Oceanic sense, commensals are those species that were

moved by humans (usually deliberately) between islands, often as a part of expansion and settlement. The use of commensal models for human mobility include analyses of molecular data using population genetics, phylogenetic and phylogeographic methods. These models are complemented by (and necessarily include) relevant biological, linguistic and archeological information, or Kirsh and Green’s ‘triangulation’ approach (2001). The genetic relationships are used to infer patterns of plant and animal dispersal and therefore indicate patterns of human mobility. Whereas human genetic research reveals patterns of migration and settlement, studies of commensal species potentially go further by revealing additional patterns of human activity, such as trade, that may not necessarily be associated with settlement. Commensal data can also enhance our understanding of agricultural practices. A number of plant and animal species in the Pacific have been used to model human mobility. The integration of these studies, along with analysis of human genetic variation, has significantly enriched our understanding of prehistoric human-mediated dispersal and colonization histories in the Pacific. Commensal animal species studied include the Pacific rat (Rattus exulans; Matisoo-Smith & Robins 2004), pig (Sus scrofa; Larson et al. 2007), chicken (Gallus gallus; Storey et al. 2007), the lizard (Lipunia noctua; Austin 1999) and the land snail (Partula hyalina; Lee et al. 2007). Commensal plant species studied include breadfruit (Artocarpus spp.; Zerega et al. 2004), bottle gourd (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) and bananas (Musa spp.; J Kennedy 2008). All of these have provided important new information regarding population origins and interactions in the Pacific. Despite improved understandings of the direction and timing of human settlement of Remote Oceania, there remain many

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Paper mulberry, a new commensal model for human mobility in Oceania 233 unanswered questions and topics of debate. For example, it has recently been suggested that post-Lapita movements from Island Southeast Asia through the low islands of the Carolines, Kiribati and Tuvalu into West Polynesia may explain some of the cultural, biological and genetic diversity observed within Remote Oceania in terms of people and their commensal plants and animals (Addison & Matisoo-Smith 2010). There also remain questions about the settlement and interaction histories of specific islands, particularly Rapa Nui. While much recent debate has focused on dating the initial human colonization of Rapa Nui (Hunt & Lipo 2006) and the subsequent impact of human arrival (Diamond 2006), the specific origin(s) of the founding population or populations and the process of colonization are issues that have not yet been resolved (Green 2000a, 2005; Martinsson-Wallin & Crockford 2002). There is the potential to explore these and other key questions in Pacific prehistory by analysing commensal species within an integrated framework of biological, anthropological and genetic data. Paper mulberry may be particularly useful as a commensal model species. It is very widely distributed in Polynesia and, of the 70 plant species prehistorically introduced into Polynesia by humans (Whistler 1991; Yen 1991), paper mulberry was one of only 20 plant species that was successfully introduced to Rapa Nui (Me´traux 1971; Orliac & Orliac 1998) and one of only six plant species introduced to New Zealand (Sykes 1969; Matthews 1996). Here we present our preliminary exploration of the use of paper mulberry as a commensal model species. We identify key questions in Pacific prehistory for which genetic data of paper mulberry could be most valuable and identify what data need to be obtained in the future to fully utilize paper mulberry as such a model species. We hope this approach will be useful for others undertaking commensal species research and will provide a framework for understanding the dispersal of paper mulberry in Oceania.

The distribution and biology of paper mulberry The genus Broussonetia belongs to the large family Moraceae, which is widely distributed in tropical, sub-tropical and, to a lesser extent, temperate regions (Mauseth 1991; Sharma 1993). Broussonetia (along with the genus Morus) belong to the large and morphologically diverse tribe Moreae, which has a wide geographic distribution in Asia and Island Southeast Asia (Zerega et al. 2005). Broussonetia papyrifera is a deciduous dioecious plant, with serrated leaves ranging from 8 to 20 cm long. Leaf shape is highly variable, even on the same plant, ranging from ovate to deeply three- to fivelobed, with the lobes being strongly lacerate in the most dissected leaves. Leaves are densely tomentose, especially on the abaxial surface. The petiole is nearly as long as the lamina. Plant varieties are distinguished by leaf shape, and on some islands these varieties are designated by different names (e.g. Hawai‘i) (B Mulloy, personal communication). Paper mulberry can be easily distinguished from other mulberry species by the milky sap and the fuzzy surface of the leaves. Plants can be multiplied by both seeds and cuttings. Its habit of producing root suckers, by which it can become a nuisance, is another distinguishing feature of paper mulberry (Whistler & Elevitch 2006). Although it is native to temperate climates, it is highly adaptable to different latitudes (Andrews 1940) and soil types, especially volcanic (Whistler & Elevitch 2006). Paper mulberry is native to southern China, Japan and Taiwan, and was introduced by Austronesian settlers to the islands of the Pacific as far east as Rapa Nui, north to Hawai‘i and south to New Zealand, and even to such remote islands as Pitcairn (Whistler & Elevitch 2006). In the Pacific islands, paper mulberry is normally only seen as a small tree or shrub and is usually pruned to keep it under 3 m tall, which may affect the species’ ability to flower in cultivation. Indeed, Florence (2004) and Matthews (1996), state that flowering is unknown or infrequent over most of the tree’s distribution in the Pacific. The lack of flowers in Pacific individuals may

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234 D Seelenfreund et al. also limit the ability to diagnose sex. Whistler & Elevitch (2006) and Whistler (2009) state that only male trees occur in Polynesia, but Rapa Nui populations of B. papyrifera have been seen to produce 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 the Pacific, propagation is mainly through root shoots, which are produced freely. Although the red fruits are consistent with dispersal by birds in the plant’s native range, there is no evidence for birds being an effective vector for the interisland dispersal of paper mulberry in the Pacific. Human-mediated propagation from seeds has not been recorded in the Pacific (plants not reaching flowering age before being harvested for their bark is probably also

a factor; Florence 1997). It is not clear whether abandoned (feral) plants may sometimes reproduce sexually and whether there is gene flow with cultivated plants that have been allowed to flower. In terms of dispersal, there may have been some natural dispersal within an island (either through vegetative or sexual reproduction), but dispersal between islands (at least in Remote Oceania) was certainly human mediated. It is possible that both sexes of paper mulberry occur in Remote Oceania, although whether this situation has varied through time (prehistory and the historical period) and space (some islands and not others) remains unknown. If both sexes are present in the same place and time, this would have obvious implications for whether breeding populations can establish, and therefore the type of biological patterns we might expect to see.

Fig. 1 (A,B) Paper mulberry (Broussonetia papyrifera) plant from Rapa Nui (Easter Island) with female inflorescence. (C) Syncarp on a plant from Rapa Nui. (D) Feral plants, in the foreground, growing between boulders inside Rano Kao volcano, Rapa Nui.

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Paper mulberry, a new commensal model for human mobility in Oceania 235 Extant B. papyrifera in Polynesia Fieldwork during 2008, by one of us (AS), revealed that a large number of paper mulberry plants still grow on some islands of Polynesia, but that it has gone extinct on several others, such as Mangareva, the Cook Islands and apparently New Zealand, where it is no longer cultivated. We performed a combination of basic botanical and anthropological fieldwork at a number of locations in Polynesia involving field observations, interviews with local informants and growers, the collection of fresh leaf material and limited examination of herbarium material. In general, the botanical varieties recognized are based on a small number of morphological traits. On each island visited, the number of plants, their density and availability varied. Leaf morphology was also variable to some degree (Fig. 2), ranging from ovate to multi-lobed and highly dissected. The sampling for our research was not carried out by a botanist, so we did not record a number of botanical features which might be

useful for understanding diversity in the species, and the recognition of different varieties. Today on some islands people recognize several varieties according to leaf morphology. At present, we cannot distinguish between the effect of phenotypic plasticity and genetically distinct lineages in explaining the observed morphological diversity. Because visits to the islands were short and most plants observed were cultivated, fruits and flowers were not observed, except on Rapa Nui. Sampling was not uniform across all islands. The number of individual samples collected per island varied from 1 in Ua Huka in the Marquesas, to 19 in Tonga, 32 in Taiwan and 38 on Rapa Nui (Table 1). This variation was due to several factors, including the plant population size on each island, which is determined by the impact of 19th and 20th century colonization, access to western goods, its contemporary use for the tourist industry or its continuity within traditional wealth and ritual contexts. The number of samples collected also differed depending on whether the sampling was performed

Fig. 2 Diverse paper mulberry (Broussonetia papyrifera) leaf morphologies from: (A) Samoa (B,C) Tonga and (D) Rapa Nui (Easter Island).

236 D Seelenfreund et al. Table 1 Locations from which paper mulberry plants were sampled for this study. Locations (country and/or island group)

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Chile

Island

Locations within island

Rapa Nui

Roiho. Poike, Maunga Toa Toa, Rano Raraku, Rano Kao, Mataveri, Ara Tataki re reo, Hanga Oteo, Pu Toki Toki Santiago, Parque Arrieta Haka Hau

mainland French Polynesia. Marquesas island Ua Pou group Nuku Hiva Fatu Hiva Tahuata Ua Huka French Polynesia, Society Islands Tahiti USA, Hawaii Hawai’i Western Samoa Savaii Upolu Tonga Tongatapu Fiji Taiwan

Viti Levu Taiwan

Taiohae Omoa, Hanavave Vaitahu, Hapatoni Arboretum Pirae, Punauia Kona Palauli, Safu’a, Siutu, Salailua, Faga Maangiangi, Apia, Siumu Foui, Tekiu, Kanokupolu, Liahona, Vaini, Niutoua, Navutoka, Pelehake, Faatai, Hamula Suva Hua Lian river, Da Han river, Lang Yang river, Da Du river, Lao Nong river, Bei Nan river, Jhuo Shuei river, Zen Wun river

by one of us, or if samples were procured through third parties (as in the case of Taiwan, Fiji, Pitcairn, Hawai‘i and some samples from Tahiti). A further complicating factor is that it often was impossible to separate or recognize individual plants because of paper mulberry’s tendency to produce suckers, which can appear as separate individuals. This was not significant when collecting isolated specimens, such as those on Tahiti or on the north coast of Rapa Nui, but was definitely relevant in Tonga, where B. papyrifera is planted in large fields. Fig. 3 shows islands sampled for this study.

The Taiwan populations Samples were collected from eight major river systems in Taiwan. Samples from individual trees were procured from coastal and inland locations from each drainage. Leaves were generally shallow to deeply lobed and somewhat lacerate.

The Tongan samples For this research, only plants on Tongatapu were observed and sampled. In Tonga, we recognized what appeared to be two varieties. The first and most abundant variety has deeply lobed, palmate leaves with lacerate lobes (see Fig. 2). The other variety has shallowly threelobed leaves. One voucher of an accession of this variety held in the herbarium of the Hawai‘i National Tropical Botanical Garden, has a female inflorescence (apparently not forming fruits) (DH Lorence, Hawaiian National Tropical Botanic Gardens Herbarium, Hawai’i, email communication). No flowers were observed on either of the varieties. The two cultivars are generally grown together, however, there is a tendency to see plantations in which one of the varieties dominates. We did observe one plant that had the two types of leaves on the same branch. Plants are extensively cultivated in plantations outside the villages and also in home gardens. Plants attain a height of 2 m before being harvested.

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Paper mulberry, a new commensal model for human mobility in Oceania 237

Fig. 3 Map of the Pacific showing paper mulberry (Broussonetia papyrifera) sampling locations.

The Samoan samples Samples were obtained from both Savai‘i and Upolu. On Upolu, most of the samples were collected on the southern side of the island around Si’umu village on the southern coast. Apparently, plants used to be more plentiful in the past, but today few isolated stands remain. People informed us that the plants were ripped out due to their invasiveness. Plants are generally grown in the back of the gardens, usually behind the pig pens and often on volcanic and rocky ground. One plant was also sampled from the backyard of a private home in Apia, the main town on Upolu. Only one large plantation was observed on Savai‘i in Sala‘ilua village. All other stands were of no more than five or six isolated plants with shallowly threelobed leaves. No flowers or fruit were observed.

Samples from Fiji The samples from Fiji were collected for us on the island of Viti Levu. No further information on the abundance of the plants on this island was available. The samples were of young,

ovate leaves but the mature plants would likely possess leaves with three to five lacerate lobes. Their shape is similar to, although slightly more lobed than, those observed and collected on Samoa, although less lobed than the samples from Tonga.

Samples from Hawaii In Hawaii samples were collected on the Island of Hawai‘i (Big Island) at Waimea, and represent three different varieties as recognized by the growers: mara mara lima (star-shaped and somewhat lacerate lobed leaves), poa aha (ovate unlobed leaves) and maui (somewhat wedgeshaped shallow to unlobed leaves). Specimens held in the Hawaii National Tropical Botanical Garden are of what is considered the ‘feral’ Hawaiian cultivar, with entire (unlobed) or shallowly three-lobed leaves found in places like Poomau Canyon and Waialae Canyon where it apparently propagates vegetatively (DH Lorence, Hawaiian National Tropical Botanic Gardens Herbarium, Hawai’i, email communication). B. papyrifera plants in the

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La¯ wa’i Canoe Gardens on Kaua‘i had three different types of leaves and their inner bark was different as well. The poa aha variety gives the whitest type of bark and is preferred by many of the modern-day barkcloth makers. Some of the plants on Kaua‘i were wiped out a few years ago by a flood (S Kauka, Kaua‘i, email communication; B Mulloy, Molokai, personal communication). French Polynesia sampling Broussonetia papyrifera still grows in the Marquesas, to a limited extent in the Society Islands, and in the Austral Islands. We did not find any plants in the Gambier Islands, or in the Tuamotus. Marquesas In the Marquesas archipelago, paper mulberry is found mainly on the islands of Fatu Hiva and Tahuata, in the southern group of islands, and on Ua Pou in the northern group. Fatu Hiva is the only island where it is cultivated and which therefore has a significant plant population. It is grown in gardens around the houses, often mixed with banana plants. The main plantations are found around the village of Omo‘a, where paper mulberry handicrafts provide the main income for the population. At Hana Vave, the other village of Fatu Hiva, located in a different valley, we only found a single plant growing beside a soccer field. On Ua Pou, most plantations are located up-valley, away from the villages. Only isolated plants are found in old gardens inside the villages. The Gambier Islands Although J Florence (2004) mentions that B. papyrifera is present in the Gambier Islands, today no paper mulberry plants appear to survive. In the past, apparently there were two varieties. Laval (1968) describes one called eute, and another which was called puri; eute being the wild variety, whereas puri were the

cultivated and very valued plants. The puri plantations were under the protection of a priest and dedicated to the god Rongo. The old name for the island of Akamaru was ‘Rari te hiapo’ (hiapo being the name of barkcloth in Niue and for the plant in Tonga). In an old song, hiapo is associated with Akamaru. Buck (1938: 249) states that there was also a plant called iapo but it, ‘. . . is said to have disappeared from Mangareva, but it is probable that iapo was an alternate name for the paper mulberry, and the connection between the old name and the plant has been forgotten’. In 1966, the French botanic expedition to Mangareva mentions both eute and puri, however, it is not clear from the publication if plant specimens were actually seen or samples collected (Huguenin 1974). Austral Islands Although we did not sample from this island group, there are records that B. papyrifera is still grown on Rurutu and Rapa (Florence 2004). The herbarium at the Muse´e des Iles, in Tahiti, holds only two samples, one from Rurutu, and the other from Rapa (V MuLiepmann, email communication). These were not examined for this study. Society Islands Extensive paper mulberry plantations were described during the early contact period for Tahiti, where the bark was used for making scented and dyed barkcloth, one of the most prized presentation and exchange objects in the past. Leaf samples were collected by J Banks and D Solander in 1772 and are kept in the US National Herbarium. The specimen shown in Fig. 4 is labelled as being collected in the ‘Friendly Islands’. We cannot find any record of any islands Cook visited being called the Friendly Islands (‘Friendly Islands’ is often used for Tonga, but Cook did not visit this island group on his first voyage). It is most likely that that specimen was actually collected

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Paper mulberry, a new commensal model for human mobility in Oceania 239

Fig. 4 This specimen of paper mulberry (Broussonetia papyrifera (L.) Vent.) was collected in 1769 by Joseph Banks and Daniel Solander during Captain James Cook’s first voyage (1768
1771). It is likely, despite the ‘Friendly Islands’ label, that this plant is from the Society Islands. The label was certainly added later because the taxonomic revision resulting in the movement of paper mulberry from Morus to Broussonetia did not occur until 1799 (Tabl. regn. veg. 1799. 3:547). Specimen US 1276390; image courtesy of the United States National Herbarium, National Museum of Natural History, Smiths onian Institution, USA.

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240 D Seelenfreund et al. in the Society Islands, where Banks observed the manufacture of tapa from ‘Morus papyrifera’ and made extensive notes in his journal (Beaglehole 1962). George and Johan Reinhold Forster, in 1774, made some observations on the cultivation of paper mulberry, noting that the plants were cultivated ‘in plots that were regularly weeded and manured’ (JR Forster 1778: 21, 445): and planted under other trees, (GA Forster 1777: 279), both in the valleys and on the hillsides (GA Forster 1777: 119, 270). JR Forster also tells us that . . . they plant the young shoots of the aouta, in regular rows, at the distance of about 18 inches, or two feet; they lop off the leaves and branches that are sprouting out, which operation increases the main shoot, and in-vigorates its strait growth . . .

and . . . as soon as the saplings have attained the size of an inch in diameter, and the height of about six or eight feet, they are drawn up, the roots and tops are then cut off, and such parts of the root as have young shoots, are carefully preserved and planted again . . . (JR Forster 1778: 445; also GA Forster 1777: 277)

Propagation seems to have been primarily by cuttings and root shoots, as attested by several early descriptions. However, it is possible that the plants seeded, as noted by Anderson (Oliver 1974: 256): ‘the cloth-plant . . . is raised by seeds brought from the mountains’. Lepofsky (2003) based on Whistler (1991) asserts that the plants could not seed. However, our observations on Rapa Nui and those made by Meilleur et al. (1997) in Hawai‘i demonstrate the ability of Polynesian plants to produce female flowers. The ability for seed to be produced in the Polynesian range has not yet been documented. Morrison in 1787 describes two varieties of the plants of the ‘youte (morus papyrifera), or Chinese paper mulberry tree, which are here called myeree and poorow. They use the bark

of these to make their finest white cloth, and they cultivate large plantations of it for that purpose’ (Morrison 2010: 141). He recorded that the name for tapa ‘. . . was Huboo or parrawye’ (paraui) and that it ‘. . . is made of the youte or cloth plant’ (Morrison 2010: 152). In the more recent ethnographic literature, the name for the paper mulberry is ‘aute (Lepofsky 2003). Today few plants grow on Tahiti or on any of the other islands of the Society group. Leaf samples were collected from a small tree, about 3 m tall, which grows in the gardens of the Muse´e des Iles at Punaauia. This tree is supposed to have been planted 40 years ago from a plant collected at B Danielsson’s garden at Paea (V Mu-Liepmann, email communication). Other samples were collected from private gardens; however, no specific information on the history of each plant was recorded. The leaves of the plants are somewhat lobed and the young leaves generally show little or no lobing. A single stand of plants was found and sampled on the island of Raiatea. These specimens had reportedly been imported from Tahiti. However, even the plants on Tahiti, that provided the cuttings for the Raiatean plants, had originally been brought from Fatu Hiva, in the Marquesas. The woman who was growing them used the plants for medicinal purposes. No plants could be found on the island of Tahaa. We did not sample on the other islands of the Society group, and inquiries made through local people did not prove fruitful. Samples from Pitcairn Leaves of Pitcairn samples are similar in shape to those from Rapa Nui and the Marquesas. No record was obtained on the history of the Pitcairn plants. It is not known if these plants are actual descendants of the plants brought by the women of the Bounty, as related by Morrison (2010), if they were already on the island or were reintroduced later. Barrow et al. (1833) observed Morus chinensis growing in the gardens and the use of bedding and clothing made of paper mulberry: ‘The women are

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clothed in the paper mulberry white cloth extending in folds from the shoulders to the feet, and so loose as entirely to conceal their figure’ (Barrow et al. 1833: 159, 161). The samples collected in Adamstown were fully grown leaves with three shallow lobes, or no lobes with only serrated edges when young. The Rapa Nui populations On Rapa Nui, paper mulberry is found growing in small and scattered populations, the only exceptions being the plants which grow inside Rano Kao crater and in the collapsed lava tubes in the area of Roiho. It is likely that these are remnants of old plantations, such as those observed by Cook and Forster in 1774. Cook saw ‘. . . in several places Otaheitian cloth plant; but was poor and weak . . . .’ (Cook 1777: 285). Johann Reinhold Forster describes the plants as being ‘. . . from two to four feet high, and planted in rows, among very large rocks, where the rain had washed a little soil together’ (Forster 1778
1780: 427). Later, in 1786, La Pe´rouse mentions the plantations inside Rano Kao crater where ‘. . . the marsh was surrounded by finest plantations of bananas and mulberry trees’ (La Pe´rouse 1797: 81). He also noted that paper mulberry plants needed to be protected from the winds and that ‘. . . the height of these trees never exceeded that of the walls by which they are sheltered’ (La Pe´rouse 1797: 73). To help ensure samples were derived from in situ prehistoric lineages, leaves were collected preferably from plants growing in isolated localities around the island and in abandoned or semi-abandoned garden enclosures. On the north-western side of the island, plants grow in places that were occupied before missionary times in the early 19th century. Here the plants grow often between rocks or boulders or in old rock gardens. Usually each locality holds no more than one or two adult plants and a number of smaller plants which clearly have grown from the root system of the older ones. Plants do not attain a height over 1.2 m (Fig. 1).

Paper mulberry collections from Rapa Nui were made by F Fuentes in 1911 for the Santiago National Museum of Natural History. Samples collected (accession number SGO 058300) show druplets, indicative of female plants. No information is provided on the exact provenance or point of collection of the samples. In his report, Fuentes (1913) states that paper mulberry ‘. . . is common inside Rano Kao crater and in the centre of the island’, presumably around Roiho, two localities where plants still grow. Skottsberg (1956) in 1919 observed neither flowers nor fruits among Rapa Nui populations. Collections made in 1953 by Volosky (accession number SGO 075595) on Rapa Nui also bear druplets, but again, no precise collection location is given for these samples. In 1971, F Sudzuki also collected samples with druplets, but once again, no precise collecting location is given. G Zizka (1991) states that he never saw the plants with fruits. An old Rapa Nui name for paper is para para (Martı´ nez 1913; Fuentes 1960). The old Rapa Nui word pararaha was used for something flat or as a verb for flattening something, parehe kahu referred to a piece or scrap, or a rag, and paruai is translated as linen (Fuentes 1960). Rapa Nui barkcloth is generally much darker than the tapa produced today in Samoa, Tonga or in the Marquesas. This might be due to the plants generally having little water and their growth is stunted by wind, therefore plants are very woody and the barkcloth is usually tough and ligneous. As a final comment, we need to point out the importance of recording the provenance of the plants collected, especially on islands such as Tahiti with a high percentage of migrant Polynesian populations, where plants may have been transported in recent or historic times. For example, within the Marquesas some of the plants we found growing on the island of Hiva Oa had recently been re-introduced from Fatu Hiva. Whenever possible, special care should be taken to register the particular history of the plants or the families who own them, or alternatively search in situ for prehistoric

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lineages of paper mulberry growing in or near abandoned archaeological sites where there is no record of historic-period cultivation or habitation, as, for example, inside Rano Kao crater or the north coast of Rapa Nui (Fig. 1D). Given the multiplicity of possible transport events and alternative explanations for the distribution of paper mulberry, we cannot over emphasize the need to work with the local community, who often know the particular history of a plant or plant population, which could help to sort out some of these issues. Potential for molecular studies of paper mulberry Because paper mulberry is of little commercial importance in western society, except for making special quality papers, there is little genetic information available for this species and also for other species of this genus. There is only one report describing the genetic diversity of Taiwanese accessions of B. papyrifera (Ho & Chang 2006) and our own analysis of ribosomal DNA internal transcribed spacer (rDNA ITS) marker sequences from Polynesian and Taiwanese samples (Pin˜a et al. 2010). There are, however, genetic resources (e.g. EST libraries, microsatellite and sequencing markers) for some commercially important members of the Moraceae, including the white mulberry (Morus alba) and common fig (Ficus carica). It may be possible to use some of these markers from other moraceous species in paper mulberry. Even if relatively few markers can be successfully transferred, it is likely that this is still a faster and less-expensive method than developing markers de novo for paper mulberry. There are a number of considerations in choosing a new marker system for paper mulberry. The markers need to be informative at the intraspecific level (although some sequence data from outgroup species may be useful). For several reasons, the amplified fragment length polymorphism (AFLP) technique (Vos et al. 1995; Meudt & Clarke 2007) is an ideal marker system: it is useful when variability is low (such as might be expected in paper mulberry
a crop

species that has undergone a recent radiation), the broad genome coverage allows for reticulating patterns of evolution to be resolved, and, because no a priori sequence knowledge is required, data can be generated very quickly. A limitation of AFLP is the need for high-quality (preferably fresh) tissue for DNA extraction. Another weakness is that AFLP is a dominant marker system (Meudt & Clarke 2007). A second strategy is to use high copy number chloroplast DNA (cpDNA) and rDNA ITS (nuclear) markers. Although these markers often have limited resolution in resolving recent radiations, their high copy number means they are often useful for amplifying low-quality and -quantity DNA as might be expected from herbarium material and tapa cloth. Finally, a third strategy is to take advantage of the small (686 Mbp; Ohri & Kumar 1986), diploid (2n26; Oginuma & Tobe 1995) genome for the development of co-dominant markers, such as microsatellite or sequencing markers. These markers have the advantage of being potentially better for starting material of variable quality (ranging from fresh leaf tissue to herbarium material of up to 250 years old), and for a long-term project in which laboratory work may be carried out by a number of different researchers. Significant disadvantages, however, are the high labour and financial investments before data are generated, and the potentially low resolution unless a large number of markers are developed. Value of B. papyrifera for addressing key questions in Pacific prehistory Lapita origins As a result of recent archaeological and genetic evidence (Kirch 2000; Larson et al. 2007; Matisoo-Smith 2007, 2009; Summerhayes 2007), a general consensus is emerging that various components of the Lapita cultural complex had different origins and all came together as Lapita in the Bismarck Archipelago
supporting Green’s Triple I model (1991, 2000b). The dispersal history for B. papyrifera in the Pacific is different to that of many other

Paper mulberry, a new commensal model for human mobility in Oceania 243

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presumably Lapita-associated plant species in that it seems to have solely an Asian origin. By contrast, other cultivated plants, such as breadfruit, sugarcane, taro and kava, were likely obtained and incorporated within the Lapita complex in Near Oceania (Lebot 1991; Yen 1991). This fact makes B. papyrifera an especially interesting plant species to test the different archaeological models of human expansion out of Asia and into Oceania. Polynesian origins As suggested earlier, it has recently been suggested that the origins of Polynesian populations may not be as simple as has often been portrayed. In general, the consensus for the last 30 years or so has been that Polynesian origins lie solely with Lapita peoples who arrived in West Polynesia around 2900 BP. Addison & Matisoo-Smith (2010), based on analyses of archaeological and human and commensal animal genetics, have proposed an alternative hypothesis in which a later population movement, occurring sometime around 2000
1500 BP and coming more directly or recently out of Island Southeast Asia may have moved through Micronesia and into Samoa. It is suggested that this group of new migrants then mixed with the Lapita colonists, who were only living at low population densities in this most eastern Lapita outpost. From the Samoa/Tonga/Fiji region of Western Polynesia, this now mixed population expanded east into central Eastern Polynesia and also dispersed to the west, back into ‘Melanesia’ and the small offshore islands in Vanuatu, New Caledonia, the Solomon Islands and Papua New Guinea (islands now generally referred to as the Polynesian outliers). Genetic analyses of paper mulberry accessions from both Near and Remote Oceania could be used to test some of these hypotheses. Rapa Nui origins and interactions Analyses of paper mulberry could also provide important and unique new insights into the

origin(s) of the people who settled Rapa Nui. Other than the analyses of ancient and modern human DNA (Hagelberg & Clegg 1993; Gonza´lez-Pe´rez 2006; Lie et al. 2007; Thorsby et al. 2009), only a small number of the genetic studies of Pacific species have included Rapa Nui samples. Chickens and rats were the only commensal animals successfully introduced by humans to Rapa Nui in prehistory and genetic analyses of these species have been inconclusive in terms of addressing settlement history. One of the reasons for this is the lack of mitochondrial DNA (mtDNA) variation within Polynesia, particularly for the Pacific rat, Rattus exulans. Barnes et al. (2006) studied mtDNA variation from archaeological remains of the Pacific rat recovered from several sites on Rapa Nui, but found that all belonged to a single haplotype, identified previously as the R9 haplotype (Matisoo-Smith & Robins 2004). This haplotype is the most common found in Polynesia, and has been identified in all possible source locations. As the authors point out, while this result might suggest only a single introduction, it does not rule out the possibility of arrivals to Rapa Nui by colonists who did not bring Pacific rats with them, or the possibility of multiple migrations to Rapa Nui by Polynesians bringing with them rats of a single genetic population, but which is found on numerous islands (Barnes et al. 2006). Storey et al. (2007), in their study of mtDNA variation in Pacific chicken (Gallus gallus) remains, report two mtDNA lineages on Rapa Nui, but unfortunately they did not analyse bones from any possible central Eastern Polynesian source populations. Without data on mtDNA of central Polynesian chickens, specifically those from the Marquesas, Australs and Gambiers (Storey et al. 2008), we cannot know if the variation seen in Rapa Nui is due to multiple introductions or merely the presence of both lineages in a single founding population. Genetic analyses of introduced plants may provide the necessary additional evidence allowing us to choose between the various hypotheses

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244 D Seelenfreund et al. regarding origins and interaction on Rapa Nui, particularly if the levels of genetic variation within Polynesia are relatively high. To our knowledge, no genetic analyses on the Polynesian-introduced plants on Rapa Nui have been undertaken. Paper mulberry is one of the few plants that was introduced to almost every island or island group in Polynesia, including Rapa Nui, and thus provides a unique opportunity to address the question of the immediate origins of Rapa Nui plants and by association, the origins of the Polynesian colonists who settled the island.

Using herbarium material and tapa cloth When studying extant plant populations it is important to consider the extent to which we can make inferences about prehistoric diversity. One way to reduce the effects of recent reintroductions of plants is to examine herbarium specimens that were collected before modernday reintroductions or extinctions (Fig. 4), such as those that happened recently on Mangareva or the Cook Islands. Unfortunately, unlike studies of modern plants, analyses such as AFLP can often not be used for this kind of material because high molecular mass DNA is needed yet is rarely obtained from anything but fresh samples. This material, however, might be suitable for using microsatellite markers, and we are currently investigating this possibility. A second approach we intend to pursue is to look at the finished products made out of paper mulberry bark. Collections of tapa cloth, made since the first European contact (which at the most are 300 years old), are held in various museums around the world, and could eventually be tested using molecular analyses. Museum collections could also provide a partial solution to the issue of chronological control, because most are of known provenance and their approximate date of manufacture are known.

Conclusions In conclusion, there are several attributes which make paper mulberry an excellent candidate for its study as a commensal species and for resolving questions around human mobility and the use and development of crops in Remote Oceania. Most importantly, paper mulberry is very widely distributed, meaning that it can be used to determine mobility patterns across a broad geographic area. Therefore, it may be possible to use paper mulberry to test models of human expansion from Asia into Near Oceania, the Lapita expansion into Remote Oceania, patterns of mobility (trade and back-voyaging) in Eastern Polynesia, and the settlement of, and interaction with, Rapa Nui in prehistory. More detailed questions around the dispersal of cultivars, and the movement (and loss) of the plant in the historic period have the potential to inform our understanding of cultural practices in the Pacific. We aim to fully develop a paper mulberry commensal model, and assess its ability to be another proxy for tracking human movement in the Pacific. This is achieved using a multidisciplinary approach, integrating botany, anthropology and genetics. Acknowledgements The present research is funded by Fondecyt, Grant 1080061 from the Government of Chile. We wish to thank the following people and institutions. For research permits, we thank M. Rauch, R. Criso´stomo from National Parks in Chile, (CONAF) and Dr P. Frogier, from the Department of Research of French Polynesia, De´le´gation a` la Re´che´rche, Tahiti. Thanks are extended also to S. Tihoni from Aranui cruises for free passage to the Marquesas, Dr K-Y. Ho, for samples from Taiwan; A. Bernacchi, J. Edmunds, for field assistance on Rapa Nui; Dr J-F. Butaud, F. Timau, T. Tereino, G.M. Teiki Teetini, M. Autuche, P. Ebb, H. Roedel, V. Vaucherot, in Tahiti, the Marquesas and Raiatea, French Polynesia; M. Atioo, F. Tunupopo, W. Chaplin, O. O’ola, in Samoa; C. Walter, for the Pitcairn samples; V. Campbell, Dr H. Sykes, V. Bonito, for

Paper mulberry, a new commensal model for human mobility in Oceania 245 samples from Fiji; B. Mulloy and M. Go´mez, for samples from Hawai‘i; and finally D. Ragone for establishing contacts. We thank all of them for their invaluable assistance.

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