Canoe plants

One of the major human migration events was the maritime settlement of the islands of the Indo-Pacific by the Austronesian peoples, believed to have started from at least 5,500 to 4,000 BP (3500 to 2000 BCE). These migrations were accompanied by a set of domesticated, semi-domesticated, and commensal plants and animals transported via outrigger ships and catamarans that enabled early Austronesians to thrive in the islands of Maritime Southeast Asia (also known as 'Island Southeast Asia'. e.g.: Philippines, Indonesia), Near Oceania (Melanesia), Remote Oceania (Micronesia and Polynesia), Madagascar, and the Comoros Islands.

They include crops and animals believed to have originated from the Hemudu and Majiabang cultures in the hypothetical pre-Austronesian homelands in mainland China, as well as other plants and animals believed to have been first domesticated from within Taiwan, Maritime Southeast Asia, and New Guinea. Some of these plants are sometimes also known as "canoe plants", especially in the context of the Polynesian migrations. Domesticated animals and plants introduced during historic times are not included.

Plants

Domesticated, semi-domesticated, and commensal plants carried by Austronesian voyagers include the following:

''Aleurites moluccanus'' (candlenut)

The candlenut ('' Aleurites moluccanus'') was first domesticated in Island Southeast Asia. Remains of harvested candlenuts have been recovered from archaeological sites in Timor and Morotai in eastern Indonesia, dated to around 13,000 BP and 11,000 BP respectively. Archaeological evidence of candlenut cultivation is also found in Neolithic sites of the Toalean culture in southern Sulawesi dated to around 3,700 to 2,300 BP. Candlenut were widely introduced into the Pacific Islands by early Austronesian voyagers and became naturalized to high volcanic islands.

Candlenut has a very wide range of uses and every part of the tree can be harvested. They were primarily cultivated for the high oil content in their nut kernels. They were used widely for illumination, prior to the introduction of other light sources, hence the name "candlenut". The kernels were skewered on coconut midribs that were then set alight. Each kernel takes about three minutes to burn and thus the series could act as a torch. This tradition of making candlenut torches exists in both Southeast Asia and Oceania. Candlenut oil extracted from the nuts can also be used directly in lamps. They can also be utilized in the production of soaps, ointments, and as preservatives for fishing gear. Other traditional uses include using the timber for making small canoes and carvings; the sap for varnish and resins; the nut shells for ornamentation (most notably as leis), fish-hooks, toys, and the production of black dyes; the bark for medicine and fiber; and so on. Some non-toxic varieties are also used as condiments or ingredients in the cuisines of Southeast Asia and the Pacific.

The Proto-Malayo-Polynesian word for candlenut is reconstructed as ''*kamiri'', with modern cognates including Hanunó'o, Iban, and Sundanese ''kamiri''; Javanese and Malay ''kemiri''; and Tetun ''kamii''. However the Oceanian words for candlenut are believed to be derived instead from Proto-Austronesian ''*CuSuR'' which became Proto-Malayo-Polynesian ''*tuhuR'', originally meaning "string together, as beads", referring to the construction of the candlenut torches. It became Proto-Eastern-Malayo-Polynesian and Proto-Oceanic ''*tuRi'' which is then reduplicated. Modern cognates including Fijian, Tongan, Rarotongan, and Niue ''tui-tui''; and Hawaiian ''kui-kui'' or ''kukui''.

''Alocasia macrorrhizos'' (giant taro)

The giant taro (''Alocasia macrorrhizos'') was originally domesticated in the Philippines, but are known from wild specimens to early Austronesians in Taiwan. From the Philippines, they spread outwards to the rest of Island Southeast Asia and eastward to Oceania where it became one of the staple crops of Pacific Islanders. They are one of the four main species of aroids (taros) cultivated by Austronesians primarily as a source of starch, the others being '' Amorphophallus paeoniifolius'', ''Colocasia esculenta'', and '' Cyrtosperma merkusii'', each with multiple cultivated varieties. Their leaves and stems are also edible if cooked thoroughly, though this is rarely done for giant taro as it contains higher amounts of raphides which cause itching.

The reconstructed word for giant taro in Proto-Austronesian is ''*biRaq'', which became Proto-Oceanic ''*piRaq''. Modern cognates for it in Island Southeast Asia and Micronesia include Rukai ''vi'a'' or ''bi'a''; Ifugao ''bila''; Ilocano, Cebuano, and Bikol ''biga''; Tiruray ''bira''; Ngaju ''biha''; Malagasy ''via''; Malay and Acehnese ''birah''; Mongondow ''biga''; Palauan ''bísə''; Chamorro ''piga''; Bima ''wia''; Roti and Tetun ''fia''; Asilulu ''hila''; and Kowiai ''fira''. In Oceania, cognates for it include Wuvulu and Aua ''pia''; Motu and 'Are'are ''hira''; Kilivila and Fijian ''via''; and Hawaiian ''pia''. Note that in some cases, the cognates have shifted to mean other types of taro.

''Amorphophallus paeoniifolius'' (elephant foot yam)

The elephant foot yam ('' Amorphophallus paeoniifolius'') is used as food in Island Southeast Asia, Mainland Southeast Asia, and South Asia. Its origin and center of domestication was formerly considered to be India, where it is most widely utilized as a food resource in recent times. But a genetic study in 2017 has shown that Indian populations of elephant foot yams have lower genetic diversity than those in Island Southeast Asia, therefore it is now believed that elephant foot yams originated from Island Southeast Asia and spread westwards into Thailand and India, resulting in three independent domestication events. From Island Southeast Asia, they were also spread even further west into Madagascar, and eastwards to coastal New Guinea and Oceania by Austronesians. Though they may have spread south into Australia without human intervention.

The elephant foot yam is one of the four main species of aroids (taros) cultivated by Austronesians primarily as a source of starch, the others being ''Alocasia macrorrhizos'', ''Colocasia esculenta'', and '' Cyrtosperma merkusii'', each with multiple cultivated varieties. Elephant foot yam, however, is the least important among the four and was likely only eaten as a famine crop, since it contains more raphides that cause irritation if not cooked thoroughly.

''Artocarpus''

Numerous species of '' Artocarpus'' are traditionally cultivated or harvested from semi-domesticated or wild populations in Island Southeast Asia and Micronesia for food, timber, traditional medicine, and other uses. They include '' Artocarpus anisophyllus'' (entawak), ''Artocarpus heterophyllus'' (jackfruit or nangka), '' Artocarpus integer'' (cempedak), '' Artocarpus lacucha'' (lakuch), '' Artocarpus mariannensis'' (Marianas breadfruit), '' Artocarpus odoratissimus'' (tarap or marang), and ''Artocarpus treculianus'' (tipuho), among many others. The most important species pertaining to the Austronesian expansion however, are '' Artocarpus camansi'' (breadnut or seeded breadfruit) and ''Artocarpus altilis'' (breadfruit).

''Artocarpus altilis'' (breadfruit)

According to DNA fingerprinting studies, the wild seeded ancestor of ''Artocarpus altilis'' is the ''Artocarpus camansi'', which is native to New Guinea, the Maluku Islands, and the Philippines. ''A. camansi'' was domesticated and selectively bred in Polynesia, giving rise to the mostly seedless ''Artocarpus altilis''. Micronesian breadfruit also show evidence of hybridization with the native ''Artocarpus mariannensis'', while most Polynesian and Melanesian cultivars do not. This indicates that Micronesia was initially colonized separately from Polynesia and Melanesia through two different migration events which later came into contact with each other in eastern Micronesia.

The reconstructed Proto-Malayo-Polynesian word for breadfruit is '' *kuluʀ'', which became Proto-Oceanic ''*kulur'' and Proto-Polynesian ''*kulu''. Modern cognates include Sundanese and Malay ''kulur'' or ''kelur''; Acehnese ''kulu''; Iban ''kurur''; Cebuano ''kulo'' or ''kolo''; Muna ''kula''; Mussau ''ulu''; Kapingamarangi ''gulu''; Wayan Fijian ''kulu''; Emae ''kuro''; Tuamotuan, Takuu, and Rarotongan ''kuru''; Tahitian ''uru''; Samoan and Hawaiian ''ulu''; and Māori ''kuru''. Note that in Māori, ''kuru'' is only mentioned in tradition, but does not refer to the plant because breadfruit did not survive into Aotearoa. Also note that it is believed that breadfruit only reached western Island Southeast Asia (Java, Sumatra, Malay Peninsula) during the recent centuries, as a result of trade with the Maluku Islands.

Another notable reconstructed word for breadfruit is Proto-Oceanic ''*maRi'' or ''*mai''. It is a common root for words for breadfruit in Micronesia, northern and western New Guinea, the Solomon Islands, the Admiralty Islands, St Matthias Islands, New Caledonia, and parts of the Central Pacific. The term itself may have originally been for ''Artocarpus mariannensis'' instead of ''Artocarpus altilis''. Cognates include Pohnpeian, Mokil, and Ngatik ''māi''; Palauan, Satawal, and Tuvaluan ''mai''; Puluwat ''mais''; Yapese ''maiyah''; and Tongan, Niuean, and Marquesan ''mei''.

''Artocarpus heterophyllus'' (jackfruit)

The jackfruit (''Artocarpus heterophyllus'') was domesticated independently in South Asia and Southeast Asia, as evidenced by the fact that the Southeast Asian names for the fruit are not derived from the Sanskrit roots. It was probably first domesticated by Austronesians in Java or the Malay Peninsula. The word for jackfruit in Proto-Western-Malayo-Polynesian is reconstructed as ''*laŋkaq''. Modern cognates include Sundanese, Javanese, Malay, Balinese, and Cebuano ''nangka''; Tagalog, Pangasinan, Bikol and Ilocano ''langka''; Chamorro ''lanka'' or ''nanka''; Kelabit ''nakan''; Wolio ''nangke''; Ibaloi ''dangka''; and Lun Dayeh ''laka''. Note, however, that the fruit was only recently introduced to Guam via Filipino settlers when both were part of the Spanish Empire.

Bambusoideae (bamboos)

Various species of bamboo ( subfamily Bambusoideae) are found throughout Island Southeast Asia, Mainland Southeast Asia, East Asia, and South Asia. In Austronesian regions, different types of bamboos have different names, as well as the products made from them. They are used variously as building materials, fishing gear, musical instruments, knives, water and food vessels, and so on. Bamboo shoots are also a food source in Southeast Asia. A few species of bamboo were carried by Austronesian settlers as they colonized the Pacific islands. They include the ohe (''Schizostachyum glaucifolium''), the common bamboo ('' Bambusa vulgaris''), and the thorny bamboo ('' Bambusa bambos'').

Reconstructed Proto-Austronesian words that referred to bamboo include ''*qauR'', ''*kawayan'', ''*buluq'', and ''*betung''. The latter entered Proto-Malayo-Polynesian and Proto-Oceanic as ''*bitung'', with cognates including Sundanese ''awi bitung''; Fijian ''bitu''; and Tongan ''pitu''. Most terms for bamboo in Polynesia, however, originated from Proto-South-Central-Pacific ''*kofe'' (originally from Proto-Polynesian ''*kofe'', "root"). Modern cognates include Tongan and Niue ''kofe''; Tokelau, Marquesan, Tuamotuan, and Māori ''kohe''; Rarotongan ''koe''; Samoan and Tahitian ''ofe''; and Hawaiian ''ohe''. Some names have also shifted refer to bamboo-like plants; especially in islands where they were not introduced into or did not survive, like in Aotearoa.

''Benincasa hispida'' (wax gourd)

''Broussonetia papyrifera'' (paper mulberry)

Paper mulberry ('' Broussonetia papyrifera''), better known as " tapa cloth tree" in the Pacific, originates from subtropical regions in mainland Asia and is one of the best evidence for the mainstream "Out of Taiwan" hypothesis of the Austronesian expansion. Various genetic studies have traced the origins of paper mulberry populations in the Remote Pacific all the way to Taiwan via New Guinea and Sulawesi. In the Philippines, which was along the expansion path, paper mulberry are mostly descendants of modern introductions in 1935. It is presumed that ancient introductions of paper mulberry went extinct in prehistory due to its replacement with hand-woven fabrics, given that paper mulberry generally only survives under human cultivation. However, its absence in the Philippines further underlines its origins in Taiwan, and not within Island Southeast Asia. Additionally paper mulberry populations in New Guinea also show genetic inflow from another expansion out of Indochina and South China.

It is believed to be the most widely transported fiber crop in prehistory, having been transported along with the full range of the Austronesian expansion, as opposed to most of the other commensal crops in Oceania. Paper mullbery is present in almost every island or island group in Polynesia, including Rapa Nui and Aotearoa. Some populations have gone recently extinct after they stopped being cultivated, like in the Cook Islands and Mangareva, although accounts and prepared barkcloth and herbarium specimens of them exist in museum collections gathered by Europeans during the Colonial Period. They were spread by Polynesians primarily through vegetative propagation with cuttings and root shoots. They were rarely cultivated from seeds as most plants were harvested prior to flowering, when the stems reach around in diameter, as described by 18th century European accounts. It is also unknown if the feral plants reproduced sexually as the plants are dioecious and require both male and female specimens to be present in one island.

Paper mulberry is primarily used in the Pacific Islands to make barkcloth ('' tapa'' in most Polynesian languages). Barkcloth, can also be made from other members of the mulberry family (Moraceae), including ''Ficus'' (figs) and '' Artocarpus''. Barkcloth was also occasionally made from '' Pipturus'' nettles, especially in Hawaii. However the highest quality of barkcloth was from paper mulberry.

Barkcloth was mainly used for clothing among ancient Austronesians and is traditionally made using characteristic stone or wooden beaters which are among the most common artifacts found in Austronesian archaeological sites. Numerous archaeological remains of barkcloth beaters in southern China has been regarded as evidence that the pre-Taiwan Austronesian homelands were located in the region prior to the southward expansion of the Han Dynasty, particularly around the Pearl River Delta. The oldest such remains is from the Dingmo Site in Guangxi, dated to around 7,900 BP. Barkcloth remained an important source of clothing fabrics in pre-colonial Melanesia, Polynesia, and parts of Indonesia. However, it has been mostly replaced by woven fiber clothing in most of Island Southeast Asia and Micronesia.

There are numerous names for paper mulberry throughout Austronesia, the most general can be reconstructed to Proto-Central Eastern Malayo-Polynesian ''*malaw'', which also refers to the loincloth and other items of clothing made from paper mulberry bark. Its cognates including Selaru ''mal''; Asilulu ''mala ai''; Buli ''māl''; Numfor ''mār''; Tanga, Tolai, and Gedaged ''mal''; Rennellese ''mago''; Kairiru ''myal''; Lusi, Kove, Manam, Gitua, Mota, Niue, Futunan, Samoan, Tuvaluan, Nukuoro, Anuta, and Hawaiian ''malo''; and Arosi, Rarotongan, and Maōri ''maro''.

In Eastern Polynesia, terms for paper mulberry can also be reconstructed to Proto-Central Eastern-Polynesian ''*aute'', with cognates including Tahitian and Rarotongan ''aute''; Marquesan ''ute''; Hawaiian ''wauke''; Rapa and Maōri ''aute''.

In most of Polynesia, the term for barkcloth can also be reconstructed from Proto-Nuclear-Polynesian ''*taba'', meaning "bark", with cognates including Wayan ''taba''; Tongan, Samoan, Mangareva, and Rarotongan ''tapa''; and Hawaiian ''kapa''. Other terms widely used for barkcloth and paper mulberry are derived from the Proto-Polynesian reconstructed word ''*siapo'', with cognates including Niue, Tongan, and Marquesan ''hiapo''; and Samoan and East Futunan ''siapo''. The term for barkcloth beater, however, can be reconstructed more extensively back to Proto-Malayo-Polynesian ''*ike'', with cognates including Uma ''ike''; Sa'a ''iki''; Bauan, Tongan, and East Futunan ''ike''; and Samoan and Hawaiian ''ie''.

''Calophyllum inophyllum'' (mastwood)

Mastwood (''Calophyllum inophyllum'') is a widespread timber tree native to tropical Asia. It is notable for its ability to grow to massive sizes in sandy or rocky beaches of island and coastal habitats, as well as its habit of sending out arching large trunks over the water where its seeds are dispersed via the currents. Due to these characters, mastwood are of particular importance to traditional shipbuilding of the larger Austronesian outrigger ships and were carried with them as they migrated to Oceania and Madagascar.

Other species of the genus ''Calophyllum'' were also used similarly, like ''Calophyllum soulattri'', ''Calophyllum peekelii'', and ''Calophyllum goniocarpum''. The wood grain of the members of the genus are characteristically interlocked, which make them harder to work with but also makes them stronger as well as being more suitable for carving intricate shapes. They were comparable in importance to how oaks were in European shipbuilding and timber industries. In many parts of Polynesia, mastwood groves planted in ''marae'' were considered sacred and abodes of spirits. Mastwood were also carved into religious objects like '' tiki''. They are also commonly mentioned in the chants and folklore of Polynesia.

Various parts of the mastwood were integral to the manufacture of outrigger canoes. The large curving limbs were commonly carved into the dugout canoes that formed the keel of the Austronesian outriggers ships. The strakes, which are attached to the keel by the uniquely Austronesian technique of "sewing" them with a combination of dowels and lashed lugs instead of nails, can also be made from mastwood, but it is more commonly made from other "softer" timber species like '' Artocarpus''. Other pieces became masts, outrigger floats, and outrigger spars. Smaller curving limbs can also be carved into the ribs of the boat.

Aside from shipbuilding, tamanu oil extracted from the fruit kernels were important in Polynesian culture. The oils, as well as poultices made from leaves and flowers, are also commonly used for traditional medicine. The leaves contain compounds that are poisonous to fish and can be used as fish poison.

The reconstructed Proto-Austronesian word for mastwood is ''*bitaquR'', with modern cognates including Ilocano ''bittáug''; Ifugao ''bitául''; Bikol, Cebuano, Maranao, Mansaka and Manobo ''bitáog'' or ''bitaug''; Nias ''bito''; Palauan ''btáəs''; Wetan ''witora''; and Asilulu ''hataul''. The Western Malayo-Polynesian words for mastwood is derived from the doublet Proto-Austronesian ''*bintaŋuR'', with cognates including Iban, Malay and Toba Batak ''bintangur'' or ''bentangur''; Tontemboan ''wintangor''; and Malagasy ''vintáno''. In Proto-Oceanic, the reconstructed word is ''pitaquR'', with cognates including Nauna ''pitɨ''; Loniu ''pitow''; Nali ''pirow''; Seimat ''hita''; Aua ''piaw''; Pohnpeian ''isou''; Rotuman ''hefau''; Fijian ''vetau'', Tongan ''fetau''; Niue, Samoan, and Tuvaluan ''fetau''; Nukuoro ''hedau''; and Rennellese ''hetau''. In most of these languages, the name specifically refers to ''C. inophyllum'', although in Ifugao, Maranao, Nias, Wetan, and Fijian, the name has become more generalized to large timber trees.

Another set of cognates for ''C. inophyllum'' in Proto-Oceanic can be reconstructed as ''*tamanu''. Its difference from ''*pitaquR'' is unclear, but given the distinction between the terms in the Mussau reflex, ''*tamanu'' probably originally referred to specimens of the tree that grow in island interiors and not on the coastlines. Modern cognates include Mussau, Tongan, Niue, Samoan, and Rarotongan ''tamanu''; Fijian ''damanu''; and Hawaiian ''kamani''.

''Cananga odorata'' (ylang-ylang)

'' Cananga odorata'', with its large, aromatic flowers, is used for ornamentation. It is not known whether it is native to Polynesia and Melanesia or introduced.

''Citrus''

Numerous species of Citrus are native to Island Southeast Asia, Mainland Southeast Asia, East Asia, South Asia and Near Oceania. The Austronesians cultivated and gathered a variety of citrus for food, medicine and washing with the thorns being used as piercing implements for tattooing. '' Citrus hystrix'', '' Citrus macroptera'', and '' Citrus maxima'' were also among the canoe plants carried by Austronesian voyagers eastwards into Micronesia and Polynesia.

''Cocos nucifera'' (coconut)

The region between Southwest Asia and Melanesia is the center of origin for coconuts ('' Cocos nucifera''), where it shows greatest genetic diversity. A study in 2011 identified two highly genetically differentiated subpopulations of coconuts, one originating from Island Southeast Asia (the Pacific group) and the other from the southern margins of the Indian subcontinent (the Indo-Atlantic group). The Pacific group is the only one to display clear genetic and phenotypic indications that they were domesticated; including dwarf habit, self-pollination, and the round "''niu vai''" fruit morphology with larger endosperm-to-husk ratios. The distribution of the Pacific coconuts correspond to the regions settled by Austronesian voyagers indicating that its spread was largely the result of human introductions.

It is most strikingly displayed in Madagascar, an island settled by Austronesian sailors at around 2,000 to 1,500 BP. The coconut populations in the island show genetic admixture between the two subpopulations indicating that Pacific coconuts were brought by the Austronesian settlers that later interbred with the local Indo-Atlantic coconuts.

Most words for "coconut" in Austronesian languages are derived from proto-Malayo-Polynesian ''* niuʀ''. Modern cognates include Tagalog ''niyog''; Chamorro ''niyok''; Malay ''nyiur'' or ''nyior''; Tetum ''nuu''; Drehu ''nu''; Gilbertese ''nii''; Hawaiian, Samoan, Tongan, Fijian, and Rapa Nui ''niu''; and Malagasy '' nio''.

Genetic studies of coconuts have also confirmed pre-Columbian populations of coconuts in Panama in South America. However, it is not native and display a genetic bottleneck resulting from a founder effect. A study in 2008 showed that the coconuts in the Americas are genetically closest related to coconuts in the Philippines, and not to any other nearby coconut populations (including Polynesia). Such an origin indicates that the coconuts were not introduced naturally, such as by sea currents. The researchers concluded that it was brought by early Austronesian sailors to the Americas from at least 2,250 BP, and may be proof of pre-Columbian contact between Austronesian cultures and South American cultures, albeit in the opposite direction than what early hypotheses like Heyerdahl's had proposed. It is further strengthened by other similar botanical evidence of contact, like the pre-colonial presence of sweet potato in Oceanian cultures. During the colonial era, Pacific coconuts were further introduced to Mexico from the Spanish East Indies via the Manila galleons.

In contrast to the Pacific coconuts, Indo-Atlantic coconuts were largely spread by Arab and Persian traders into the East African coast. Indo-Atlantic coconuts were also introduced into the Atlantic Ocean by Portuguese ships from their colonies in coastal India and Sri Lanka; first being introduced to coastal West Africa, then onwards into the Caribbean and the east coast of Brazil. All of these introductions are within the last few centuries, relatively recent in comparison to the spread of Pacific coconuts.

''Coix lacryma-jobi'' (Job's tears)

''Colocasia esculenta'' (taro)

The taro (''Colocasia esculenta''), sometimes referred to as the "true taro", is one of the most ancient cultivated crops and pre-dated the Austronesian expansion. Taro is found widely in tropical and subtropical regions of South Asia, East Asia, Southeast Asia, Papua New Guinea, and northern Australia and is highly polymorphic, making taxonomy and distinction between wild and cultivated types difficult. It is believed that they were domesticated independently multiple times, with authors giving possible locations as New Guinea, Mainland Southeast Asia, and northeastern India, based largely on the assumed native range of the wild plants. However, more recent studies have pointed out that wild taro may have a much larger native distribution than previously believed, and wild breeding types may also likely be indigenous to other parts of Island Southeast Asia.


Archaeological traces of taro exploitation have been recovered from numerous sites pre-dating the Austronesian expansion, though whether these were cultivated or wild types can not be ascertained. They include the Niah Caves of Borneo, dating to <40,000 BP; Ille Cave of Palawan, dated to at least c. 11,000 BP; Kuk Swamp of New Guinea, dated to 10,200 to 9,910 cal BP; and Kilu Cave in the Solomon Islands dated to around c. 28,000 to 20,000 BP. In the case of Kuk Swamp, there is evidence of formalized agriculture emerging by about c. 10,000 BP, with evidence of cultivated plots, though which plant was cultivated remains unknown.

Regardless, taro were definitely among the cultivated plants of Austronesians as well as preceding populations in Island Southeast Asia. However, their importance in Island Southeast Asia had largely been replaced by rice, although they are still planted at the margins of rice paddies in some communities. They remained a staple in the islands of Melanesia and Polynesia