Agrobiodiversity, Social Institutions, and Indigenous Farming Practices: A Case Study of the Rukai in Wutai, Taiwan

Abstract

Agrobiodiversity plays a crucial role in maintaining sustainable agricultural systems and supporting local livelihoods, but its integration into indigenous social institutions and cultures remains under-researched. We used a qualitative approach entailing in-depth interviews and participant observation to conduct research among the Rukai indigenous farmers in Taiwan focusing on: (1) how Rukai farmers practice agrobiodiversity, and (2) how these agricultural practices integrated into social institutions and cultural norms. Our findings reveal that food security and risk minimization are the prime considerations to preserve agrobiodiversity. Rukai farmers practice swidden cultivation utilizing five mixed cropping systems in seven different ecosystems that vary by elevation, length of exposure to sunlight, and temperature, intercropping, and regular fallows to maintain soil fertility. Importantly, Rukai agricultural practices are integrated into four social institutions that regulate the distribution and management of land and resources, the timing and nature of agricultural activities, and social recognition and status to ensure maximum production throughout the year and reducing risks posed by climate change.

Full text

Human Ecology (2023) 51:1127–1140
https://doi.org/10.1007/s10745-023-00463-4
on TEK play a crucial role in enhancing community
adaptive capacity by providing social, economic, and
ecological security and promoting resilience and sustain-
able development (Ingty, 2017) through the integration of
traditional agricultural practices, indigenous knowledge,
and local governance systems (FAO, 2004; Williams &
Kramer, 2019).
There is a great deal of research on agrobiodiversity
and TEK in indigenous communities (Berkes, 2018;
Cajete, 2012; Corvalán et al., 2020; Das et al., 2022;
Lu et al., 2006; Mylliemngap, 2021; Sarria et al., 2022;
Zimmereret al., 2020). However, prior investigations
have predominantly focused on examining the inuence
of religion, social values, and taboos on environmental
and biodiversity protection (Berkes, 2018; Gadgil et al.,
2021; Rath & Ormsby, 2020). The social dimensions of
biodiversity have received comparatively less attention
(Galicia-Gallardo et al., 2023; Ingty, 2017; Minato et
al., 2010; Skarbø, 2014). Though the social-ecological
network approach has proved useful to assess the impor-
tance of social network structures, composition, and
connectivity patterns in facilitating farmers’ access to
agrobiodiversity (Isaac & Matous, 2017; Labeyrie et al.,
Introduction
Agro-ecosystem management strategies rooted in tradi-
tional ecological knowledge (TEK) provide heightened
adaptability and resilience in the face of climate change
(Hosen et al., 2020; Ingty, 2017; McMillen et al., 2014;
Wyllie de Echeverria and Thornton, 2019). TEK encom-
passes an extensive reservoir of knowledge, practices,
and beliefs pertaining to environmental stewardship and
sustainability generated over generations of engagement
between indigenous and local communities and their sur-
rounding ecosystems (Berkes, 2018). Agrobiodiversity
refers to the range of plants, animals, and microorgan-
isms cultivated by a community, and is shaped by human
choices that are also inuenced by social values (Altman
& Mesoudi, 2019). Local indigenous institutions based
Hong-Zen Wang
hongzen@g-mail.nsysu.edu.tw
1 Department of Sociology, Center for Austronesian Studies,
National Sun Yat-sen University, 70 Lianhai Road,
Kaohsiung 804, Taiwan
Abstract
Agrobiodiversity plays a crucial role in maintaining sustainable agricultural systems and supporting local livelihoods, but
its integration into indigenous social institutions and cultures remains under-researched. We used a qualitative approach
entailing in-depth interviews and participant observation to conduct research among the Rukai indigenous farmers in Tai-
wan focusing on: (1) how Rukai farmers practice agrobiodiversity, and (2) how these agricultural practices integrated into
social institutions and cultural norms. Our ndings reveal that food security and risk minimization are the prime consid-
erations to preserve agrobiodiversity. Rukai farmers practice swidden cultivation utilizing ve mixed cropping systems in
seven dierent ecosystems that vary by elevation, length of exposure to sunlight, and temperature, intercropping, and regu-
lar fallows to maintain soil fertility. Importantly, Rukai agricultural practices are integrated into four social institutions that
regulate the distribution and management of land and resources, the timing and nature of agricultural activities, and social
recognition and status to ensure maximum production throughout the year and reducing risks posed by climate change.
Keywords Traditional ecological knowledge · Agrobiodiversity · Climate change · Swidden agriculture · Indigenous
farming practices · Social institutions · Rukai indigenous farmers · Wutai township · Pingtung county · Taiwan
Accepted: 29 November 2023 / Published online: 7 December 2023
© The Author(s) 2023
Agrobiodiversity, Social Institutions, and Indigenous Farming
Practices: A Case Study of the Rukai in Wutai, Taiwan
Qing-XiongBa1· Hong-ZenWang1· Mei-HsiangWang1
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Human Ecology (2023) 51:1127–1140
2021), these studies do not address the inuence of social
structures and institutional design.
To address this gap, we examine the case of Rukai
indigenous farmers in Taiwan to demonstrate how tradi-
tional agrobiodiversity farming practices based on TEK
are embedded in social institutions. Specically, we
focus on two research questions: rst, how Rukai farmers
practice agrobiodiversity, and second, how these swidden
farming practices are integrated into social institutions
and cultural norms.
We rst provide an overview of our research site and
methodology, and the importance of food security in sus-
taining tribal livelihoods. We then examine agrobiodi-
versity in our study site and analyze the diverse mixed
cropping practices implemented across dierent land
zones characterized by varying elevation and temperature
gradients. We next explore four vital social institutions to
elucidate the complex interrelationships among farming
systems, agrobiodiversity, and social institutions. Our
conclusion highlights the implications of our ndings.
Research Site and Methods
We conducted ethnographic eldwork in Wutai Township,
Pingtung County, Taiwan, which is predominantly inhab-
ited by indigenous Rukai (Fig. 1). The Rukai are distributed
across three dierent regions in Taiwan, in areas ranging
from 430 to 2,736 m.a.s.l. Wutai Township comprises seven
villages: Wutai, Adiri, Kinulane, Kabalalradhane, Karume-
medesane, Labuwane, and Kudrengere (in Chinese: Wutai,
Ali, Jilu, Shenshan, Jiamu, Dawu, and Guchuan respec-
tively). As of February 2023, there were 1,104 households
and a total population of 3,324 in the township, with the
largest village being Wutai village, with a population of
1,277 (Neipu Household Aairs Oce, 2023).
Historically, the Rukai in the Wutai region practiced swid-
den farming subsistence agriculture and only exchanged
surplus produce with other neighboring indigenous com-
munities, including Paiwan and Bunun tribes. Following
a short war, the Japanese government ocially colonized
Rukai territory in 1914, and the Wutai Rukai were forced to
cultivate paddy rice elds in Kabalalradhane in the 1930s to
conform with the ocial ideology that sedentary agriculture
Fig. 1 Map of Wutai Region
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Human Ecology (2023) 51:1127–1140
was superior to shifting cultivation. However, rudimentary
cultivation techniques, diculties in maintaining the water
supply system, and the lack of connection between rice
and traditional culture led to the abandonment of the paddy
elds immediately after the Japanese left and the Rukai re-
established their traditional farming practices (Matsuoka,
2012/2018).
In the aftermath of World War II, Taiwan came under
the control of the Chinese colonial government, which
implemented various agricultural initiatives in indigenous
areas in the name of development. One such initiative
involved the introduction of cash crops, including the cul-
tivation of pears (Pyrus serotina Rehd. cv. Hengshan), to
the Wutai region during the 1960s. However, inadequate
technical cultivation skills, such as grafting techniques,
coupled with insucient management practices related to
fruit thinning, pest control, and the establishment of eec-
tive distribution channels, contributed to the abandonment
of the pear cultivation project within a few years (Ba et
al., 2021). However, from 2010 other cash crops, such as
coee and ice jelly (Ficus pumila var. awkeotsang) have
gradually attracted the interest of indigenous farmers in the
Wutai, Kuchapungane, and Makazayazaya regions due to
rising market demand (Taiban, 2014, pp. 157-8). Never-
theless, challenges persist around marketing, management
techniques, and environmental and physical constraints. As
a result, in 2017, traditional crop cultivation continues to
occupy in excess of 60% of the total cultivated area in Wutai
(Wutai Village Oce, 2022, p. 314, Tables 130 and 131).
The Rukai are a hierarchical society, with two classes:
chiefs, who nominally own the land, and commoners (vil-
lagers). Consequently, there are two categories of land
ownership: land leased from the chiefs by the villagers for
cultivation, and land permanently acquired by the villagers
through gifting the chiefs during public ceremonies recog-
nized by the whole village (Qiao, 2001).
We selected the Wutai area for our research for two rea-
sons, in addition to the rst author’s indigenous background
and his knowledge of this region. Firstly, unlike many other
indigenous communities in Taiwan, the Rukai were not
forced to relocate from the hills to the foot of the moun-
tains during Japanese and Chinese colonization, and their
social structures and norms have remained relatively intact.
Secondly, most of the agricultural work is done by elderly
Rukai who adhere to traditional socio-cultural practices that
preserve agrobiodiversity.
The primary data was collected by the rst author, a
Rukai indigenous scholar who grew up here. From 2015
to 2017, he returned to his hometown and worked as a
farmer, meticulously documenting various aspects of farm-
ing practices, agrobiodiversity systems, social interactions,
and socio-cultural information. Due to the close-knit nature
of the Wutai community, he had easy access to interviews
and conducted formal interviews as well as informal con-
versations during daily life, primarily in the Rukai lan-
guage and occasionally in Mandarin. Formal interviews
were audio recorded and later transcribed. In cases where
clarication was needed, the interviewees were contacted
by phone. Most of the interviewees were indigenous Rukai
people over 70 years old, whose life histories and farming
experiences provided valuable insights for the research. To
Table 1 Demographic background of interviewees
Name Sex Social
Class
Age Occupation
Uselrepe ka
Tarudralumu
F Commoner
(Widow)
82 Farmer,
Housewife
Kalrimadraw ka
Pacekele
M Commoner 81 Farmer
Kuringici ka Pacekele M Commoner 79 Farmer, Hunter
Caeve ka Alegeane M Commoner 77 Farmer, Hunter
Kwalrimi ka Kapulru F Commoner 77 Farmer,
Housewife
Kainwane ka
palavelave
M Commoner 77 Farmer
Kedevese ka Pacekele F Commoner 76 Farmer,
Housewife
Lavane ka Manigaii F Commoner 75 Farmer,
Housewife
Tivulrangane ka
Tavelengane
M Commoner 75 Farmer, Hunter
Dremedremane ka
Salrebelrebe
F Commoner 75 Farmer
Asyane ka pakidavai F Commoner 74 Farmer,
Housewife
Capadisi ka Mabalriyu F Commoner
(Widow)
73 Farmer,
Housewife
Lebetai ka Aruladenge F Commoner 72 Farmer,
Housewife
Upelrenge M Commoner 72 Farmer
Aras ka Pacengelraw M Noble 72 Farmer
Palri M Commoner 70 Farmer
Pulravale ka
Kaelreane
M Commoner 67 Farmer
Cugange ka Abalyusu M Noble 62 Farmer, Hunter
Legeaii ka
Palrangelrange
M Commoner 62 Farmer
Rethamekane
Kadarulane
M Commoner 56 Coee Farmer,
Retired Civil
Servant
Dralrawsu Pacekele M Commoner 55 Coee Farmer,
Teacher
Acudusu Ragyusane M Commoner 43 Coee Farmer,
Carpenter
Notes:
1. The age of the inter viewees corresponds to their act ual age at the
time of the eld survey
2. The names used here are real names, as the inter viewees expressed
their desire to have their names recorded to highlight their contribu-
tions to this research
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Human Ecology (2023) 51:1127–1140
encounters the need to participate in tribal activities, partic-
ularly funeral ceremonies, which often require adjustments
to previously established team schedules. Over time, the
two Han Taiwanese scholars gained an increasing aware-
ness that tribal norms, which prioritize familial and social
relationships, take precedence over considerations of e-
ciency and predictability. Consequently, throughout the col-
laborative process, both parties were compelled to navigate
a delicate balance between these diverging social norms.
Food Security Among Taiwan’s Indigenous Tribes
Prior to the imposition of the Japanese colonial government
over the indigenous tribes of Taiwan in 1914, each operated
as an independent entity that maintained varying degrees of
friendly or hostile relations with one another. For example,
a survey conducted in the 1930s revealed that the Wutai
had friendly relations with the Tarumake, Adiri, Kucapun-
gane, Kinulane, Kabalalradhane, and Karumemedesane,
but hostile relations with the Oponoho, Sagarane, Timuru,
Talamakao, and Valjulu (Yap, 2017, p. 133). Each tribe
has its own territory, and encroaching on another’s land
could lead to disputes, head-hunting, or war. This custom
remains strong among the Wutai. For instance, the Labu-
wane monopolize eco-tourism on the River Thadarane in
their traditional territory, utilizing jeeps to transport tourists
between scenic spots and accommodations. Wutai villagers,
despite belonging to the same Rukai ethnic group and being
only 20 min away from the Labuwane by car, are prohibited
from operating this business due to their longstanding lack
of direct friendly relations. Consequently, it is imperative
for each group to maintain the highest level of food security
to avert disasters, including conicts with hostile neighbors.
One strategy is to forge alliances to facilitate mutual
assistance during times of food scarcity. Adiri village is
situated in a cold and wet zone, and thus has a distinct cul-
tivation cycle from Wutai so that at times of potential food
shortages in one village, the other may have surpluses to
share. Another strategy to mitigate the risk of food scar-
city is the adoption of agrobiodiversity methods to mitigate
potential crop failure. Cultivating various crops at dierent
times throughout the year guarantees a consistent food sup-
ply. These agricultural practices necessitate collective social
organization (Torres-Vitolas et al., 2019; Handoyo et al.,
2022).
Crop Varieties and Local Classication
Sasaki and Fukano (1976/2013: 48) conducted an agri-
cultural survey of the Rukai village of Kinulane, located
in Wutai Township. They collected samples of two types
of millet: glutinous (11) and japonica (6) millet, which
further update and supplement these data, his fellow authors
accompanied him on multiple visits to the region between
2021 and 2023 and conducted additional interviews with
Rukai farmers below the age of sixty to gather information
about the younger generation’s current cultivation practices
of cash crops. Prior to the interviews, the participants were
informed about the research objectives and required to sign
an interview consent form, ensuring the condentiality of
the interview data. As a gesture of appreciation, partici-
pants were oered a small gift and provided with a US$30
honorarium. In addition to ethnographic eld data, we also
collected land ownership information from the Wutai Town-
ship Oce and utilized GIS to identify lands owned by the
same family across dierent climate zones.
It is important to address our positionality as researchers.
Within Rukai culture, individuals are required to contribute
to the society to attain social recognition. In the past, hunters
would share their captured prey with fellow tribe members
and provide information about changes in the agricultural
environment, for which they were publicly recognized by
wearing a lily ower. Despite being the rst indigenous
scholar from the Wutai village to obtain a doctoral degree,
the rst author is still expected to actively contribute to the
village community to earn social respect. Accordingly, he
strives to enhance the understanding and appreciation of
Rukai culture among a wider audience in mainstream soci-
ety through various means such as educational outreach,
participation in television programs, and writing popular
science articles. In addition, the three authors maintain close
social relationships with the Wutai villagers, actively sup-
porting the establishment of cultural tourism initiatives and
collaborating with local township oces to promote Rukai
culture. As part of their commitment to preserving TEK, the
authors have plans to edit and publish an academic book on
Rukai culture. These endeavors align with the ethical guide-
lines followed throughout the research process.
In addition, in contrast to the universalistic approach of
traditional Western science, indigenous knowledge systems
emphasize the contextual specicity and localized nature of
knowledge, which incorporates belief and social systems
(Watts, 2013). Integrating these two distinct epistemolo-
gies poses a signicant challenge for both indigenous and
non-Indigenous scholars in Taiwan and elsewhere. There-
fore, the participation of indigenous scholars in scientic
research can foster more meaningful dialogues with con-
temporary scientic knowledge production systems. It
is noteworthy that the two Han Taiwanese scholars bring
their own cultural values and epistemological perspectives
into the research collaboration, guided by their indigenous
partner’s insights. This prompts them to remain vigilant
and acknowledge any biases or preconceptions that may
emerge. For instance, the indigenous scholar frequently
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Human Ecology (2023) 51:1127–1140
horticultural fruit crops, and fruit and vegetable crops.
However, according to the local classication system,
crops are organized into only four categories: food
crops (taleke), potato crops (lawrasi), vegetable crops
(lacenge), and hobby crops (sapwane). This divergence
in crop classication reects the distinct conceptual
frameworks and values that the local community assigns
to dierent crops. For the Wutai, the cultivation of crops
primarily secures food, and consequently, all crop types
except for hobby crops are considered essential sources
of subsistence.
Sweet potato (Ipomoea batatas) is a highly adaptable
food crop that can be grown in various areas and is avail-
able throughout the year, serving as the main staple food.
As it is widely available, it is not considered culturally
sacred. Although not the main staple food, millet holds
signicant cultural value as the sole crop used to pro-
duce wine, which is a requirement for all rituals. Millet is
extensively used in events such as weddings, birthdays,
house building, childbirth, rituals, and marriage, and is a
sacred symbol in Rukai society.
Fruits classied as horticultural produce are typi-
cally considered as food. Mango (Mangifera indica L.)
is used as a shade tree for resting in elds, and farm-
ers usually consume it as a snack during breaks. Papaya
(Carica papaya L.) is considered a supplementary food
and is grown around houses. Due to their secondary
importance, these crops receive less attention in terms
of management. Indigenous farmers lack a tradition or
knowledge of fruit plantations, making it challenging for
them to transition to commercial fruit plantations.
Traditional “vegetables” (lacenge) are plants that are
easily gathered during farm work. For instance, wild let-
tuce (Pterocypsela indica (L.) C. Shih) and black night-
shade (Solanum nigrum Linn.) grow in the wild and
are generally cooked with millet, and fruits of chayote
(Sicyos edulis) (Rukai: haitulri) are cooked with sweet
potatoes or pumpkins.
were classied based on their seed starch properties.
They report: “… the number of millet varieties grown in
the village may be more than several times the number
collected by the author[s]” (ibid.). In our 2018 survey
of Wutai village, 13 strains of glutinous millet (Strain
Durai) and two strains of japonica millet (Strain Cipaer-
ane) were identied (Table 2; Fig. 2).
The crops cultivated currently in Wutai village com-
prise 26 distinct species that can be further subdivided
into 78 local strains (Table 3), and grouped into nine
modern categories: cereal crops, legume crops, tuber
crops, root crops, oil crops, hobby crops, sugar crops,
Table 2 Millet landraces and uses
Name of Millet
Landrace
Uses in Food and Social Functions
1* Darangidangi ka
paedeane
millet wine, millet cake, boiled millet
2 Lamarwaane millet wine, millet cake, boiled millet
3 Makasasagarane millet wine, millet cake, boiled millet
4 Palralramu millet wine, millet cake, boiled millet
5 Saidhipane millet wine, millet cake, boiled millet
6 Salalai millet wine, millet cake, boiled millet
7 Talralrelreba millet wine, millet cake, boiled millet
8 Ababaake millet wine, millet cake, boiled mil-
let, symbol of bumper harvest
9 Paedeane millet wine, millet cake, boiled mil-
let, bridewealth
10 Tivukale ka paedeane
/ kaduradhane
best ingredients for millet wine, mil-
let cake, bridewealth
11 Salalai ka
Darangidangi
millet wine, millet cake,
headpieces, bridewealth, capacity for
cultivation
12 Darangidangi ka
ciparepare
millet wine, millet cake, decoration
13 Lrulrubungu millet wine, millet cake, ceremonies
for men
14 Sinabucyane** porridge for the sick or athletes
15 Latwatwaane** porridge for the sick or athletes,
headpieces, bridewealth, decoration
Notes:
* The numbers for each landrace are the same as in Fig. 2
** 1–13 Waxy type (Durai); 14–15 Non-waxy foxtail millet (Cipaer-
ane)
Fig. 2 Millet varieties in Wutai
region. Notes: 1 ~ 13 Waxy millet
(Durai), 14 ~ 15 Non-waxy millet
(Cipaerane)
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Human Ecology (2023) 51:1127–1140
Table 3 Crop categorizations
Modern Crop
Categorization
English Name (species) Rukai Name Rukai Names of the Landraces
Cereals or
grain crops
Foxtail millet
(Setaria italica (L.) P. Beauv.)
Becenge Strain Durai: Darangidangi, Paedeane, Makasasag-
aran, Darangidangi ka ciparepare, Saidhipane, Lrul-
rubungu, Palralramu, Tivukale, Salalai, Talralreba,
Ababaake, Lamarwaane, Darangidangi ka paedeane
Strain Cipaerane: Latwatwaane, Sinamucyane
Sorghum
(Sorghum bicolor (L.) Moench)
Rumagai Ka Rumagadhane, pairangane ka rumagai
Maize (Zea mays L.) Lapanai Durai ka Lapanai (kalapanadhane) (yellow), lapanai
ki pairange (white)
Formosa frost grass
(Spodiopogon formasanus Rendle)
Lalrumai Lalrumai
Formosa lambsquarters
(Chenopodium formosanum Koidz.)
Baae Duduli ka baae, bwaburavane ka baae, Ethenge,
darangidangi (rabungu)
Adlay (Coix lacryma-jobi L.) Mudai Mudai
Legumes for seed Pigeon pea
(Cajanus cajan (L.) Huth)
Karidrange Karidrange (ve nuts), Kapitu (seven nuts), Maka-
teteve (red nuts)
Cowpea
(Vigna unguiculata (L.) Walp.)
Lepelepe Kalepelepane, Kadaida (wild, edible), Kavuthyange
(small), Kalepelepane, Alrumu
Lima bean
(Phaseolus limensis Macfad.)
Thadare Thadare
Tuber crops Taro
(Colocasia esculenta (L.) Schott)
Taai Upland taro: Rithiliyu, Drangadrangale, Baliyu,
Kulrailraili, Kataiyane, Lreleve, Taburicaca, Dreke,
Kawdawdaare, Taruimu, Makapakalrukalru,
Paddy taro: Drulrungu
Root crops Cassava
(Manihot esculenta Crantz)
Udunku Duduli ka udunku, Lrawlrapungu ka udunku, Bwa-
buravane ka udunku
Common yam
(Dioscorea batatas Decne.)
Tuba Katubane (red), Bakwai (particularly small)
Nigaimo
(Dioscorea opposita Thunb.)
Lrawlraapungu (white)
Sweet potato
(Ipomoea batatas (L.) Lam.)
Urasi Peleti, Amurwane, Ulralrebenge, Makatanakavusu,
Pasavavaau, Usavane, Lriteteete, There, Kasapiti,
Senedhane
Oil crops Sesame (Sesamum indicum L.) Kulungu Kulungu
Peanut (Arachis hypogaea L.) Makapairange Bukulruui, Kamakapairangane
Stimulants Areca nut (Areca catechu L.) Sabiki Sabiki
Betel pepper (Piper betle L.) Drangaw Lawviyu, kadravangane
Coee
(Coea canephora pierre ex A. Froehner)
Kakuri Kakuri
Fruits Banana (Musa × paradisiaca L.) Belebele Kabelebelaane (banana)
Plantain(Musa basjoo) Lrasa (plantain)
Mango (Mangifera indica L.) Kamadha Kamadha
Vegetables Pumpkin
(Cucurbita moschata Duchesne)
Gulrudulrane Thulululrungu (oval), Capyapyake (round at),
Tububu (oval with tummy)
Chinese chive
(Allium tuberosum Rottler)
Kucai Daidapiri ka kucai
Shallot
(Allium cepa var. aggregatu G. Don)
Kucai Tungangadra ka kucai/ragiyu
Sugar plants Sugarcane
(Saccharum ocinarum L.)
Cubusu Tavarikwaai for sugar manufacturing, kacubusane
(longer internode), parangane ka cubusu (plain
sugar cane)
Source: Qing-xiong Ba
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Human Ecology (2023) 51:1127–1140
(Carrera-Garcia et al., 2012). Specically, the land use prac-
tices of the Wutai Rukai people are shaped by the geographic
variations (Fig. 3; Table 4) and various cropping systems,
including fallow, rotational plantation, mixed cropping, and
intercropping. This has led to the establishment of a highly
Land Classication and Mixed Cropping
The practice of agrobiodiversity in Wutai is inuenced by
the intricate interplay between genetic resources, cultur-
ally managed ecosystems, and the physical environment
Table 4 Five dierent mixed cropping systems in seven dierent zones
Cropping Systems Agricultural Drekai
(wet and cold)
Kabiceacelrake
(buer zone)
Labelabe
(warm and hot)
Altitude (m) 900>1200 600 ~ 900 < 600
Temperature 13.0 ~ 20.8 20.8 ~ 22.6 22.6 ~ 24.4
Sunlight Time (Short) (A) Dry Taro Mixed Cropping (B) Dry Taro, Peanut, Millet
or Sweet Patato Mixed Cropping
(C) Dry Taro, Peanut, Millet or
Sweet Patato Mixed Cropping
Sunlight Time (Long) (D) Dry Taro or Millet Mixed Cropping (E) Dry Taro, Peanut, Millet or
Sweet Patato Mixed Cropping
(F) Peanut, Millet or
Sweet Patato Mixed Cropping
Water Swamp (G) Paddy Taro Mixed Cropping (G) Paddy Taro Mixed Cropping
Villages Adiri Kinulane, Wutai, Kabalalradhane Karumemedesane,
Kudrengere, Labuwane
Notes:
1. The letters in parentheses, A BCDEFG, represent the eight dierent agricultural zones in Fig. 3
2. G represents rivers with water swamp, which ows through Kabiceacelrake and Labelabe zones
Fig. 3 Farmland classication by elevation and sunlight length in Wutai. Note: Please refer Table 4 for an explanation of the various physical
conditions in each zone and the corresponding farming practices
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Human Ecology (2023) 51:1127–1140
potatoes or taro in rotation. The choice of crops in rotation
is determined by the farmer’s preference (see Cavechia et
al., 2014).
The Labelabe zone (C and F zones) located below
600 m.a.s.l. is characterized by a warm and hot climate,
with temperatures ranging between 22.6 and 24.4 °C. This
zone is suitable for cultivating millet mixed crops, peanut
mixed crops, and sweet potato mixed crops. Karumemede-
sane, Kudrengere, and Labuwane villages have farming
areas in this zone. While most farmers in Labelabe culti-
vate warm climate crops, such as millet, peanut, and sweet
potato, some also engage in dry taro mixed cultivation due
to insucient sunlight in some areas resulting in low soil
temperature and high humidity, making upland taro mixed
cultivation a viable alternative.
The Rukai refer to riverbank (G zone) soil as “valrwalru
ka Acyacilai” (water swamp). The relatively at river land
is primarily cultivated in paddy taro elds. Owning such a
eld is considered a symbol of wealth due to their scarcity.
Paddy taro holds signicant importance in social and cul-
tural events, such as marriages, ceremonial rites, and wel-
coming visitors.
Mixed Cropping Systems and Agrobiodiversity
In Wutai mixed cropping is widely practiced. There are
ve main types of mixed cropping systems prevalent in the
region: millet (babecengane), upland taro (tataitaai), peanut
(maka pairairaange), sweet potato, and paddy taro (dradrul-
rungane) mixed cropping systems (Table 4). The choice of a
mixed cropping system depends on various factors such as
physical space, correlation with climatic conditions, topo-
graphical variations, soil properties, and short- or long-term
crops. These factors contribute to the high level of complex-
ity in the plantation system and result in maximum agro-
biodiversity in the region. Clearing new land for planting
crops requires extensive labor investment and small-scale
swidden farming relies on ecosystem services for favorable
agricultural conditions (Marquardt et al., 2013).
Long-Term Timeline: Sedentary or Shifting Farming
Agricultural cultivation practices in Wutai can be catego-
rized as short- and long-term. For the long term, it is sed-
entary cultivation in the Kabiceacelrake buer zone, while
shifting cultivation in the other two zones. For yearly plan-
tation, farmers use mixed plantation and intercropping.
In Kabiceacelrake buer zone, agricultural land is often
located near the tribal community, allowing easy access so
that farming practices in these areas tend to be sedentary.
For example, millet mixed cropping is often utilized for
two years before the soil fertility is depleted, when farmers
intricate plantation system that serves to maximize agrobio-
diversity and minimize risk of food shortages in the region.
The Rukai people of Wutai classify their lands into eight
distinct categories, among which the sacred type is deemed
o-limits for agricultural activities. The Sacred Drekai zone,
situated above 1,200 m in elevation, is considered uninhab-
itable due to the presence of malevolent spirits. Although the
low temperature renders the Sacred Drekai zone unsuitable
for crop cultivation, it plays a crucial role in maintaining
biological reproduction, ecological integrity, and providing
natural resources and water for the population. By conserv-
ing this area, the Rukai facilitate the biological reproduction
of wild ora and fauna and contribute to water supply ben-
ets for downstream agricultural zones (Byers et al., 2001;
Ormsby & Bhagwat, 2010; Rath & Ormsby, 2020; Suchiang
et al., 2020; Wadley & Colfer, 2004;).
Below 1,200 m, the land is classied into three catego-
ries based on altitude: the agricultural Drekai zone (cold and
damp), the Kabiceacelrake zone (buer zone between cold
and hot), and the Labelabe zone (warm and hot). The zones
are further classied based on the length of sunshine, result-
ing in six types of lands for farming (Fig. 3, A to F). The last
type of cultivated land is paddy taro land, mainly planted in
riverbank elds (G zone, Fig. 3).
The agricultural Drekai zone (A and D zones), located
between 900 and 1,200 m.a.s.l. is characterized by a rela-
tively wet and cold environment with an average tem-
perature of 13-20.8 °C. The choices of crops planted are
determined by variations in the topography and duration of
sunlight on cultivated land. In areas with a shorter duration
of sunlight (A zone), farmers typically practice mixed crop-
ping of dry taro with other crops, whereas in areas with a
longer duration of sunlight (B zone), mixed cropping of mil-
let is more common with other crops such as Taiwan quinoa,
corn, pumpkin, and dry taro. Millet cultivation in this zone
typically starts in November or December, earlier than in
the Labelabe (warm and hot) zone, where it begins in Feb-
ruary. The Adiri village locates in the agricultural Drekai
zone.
Kabiceacelrake zone (B and E zones) is situated between
600 and 900 m.a.s.l., with average temperature ranging
from 20.8 to 22.6 degrees, which is suitable for mixed
cropping of dry taro, millet, peanuts, and sweet potatoes.
The villages in this zone, Kinulane, Wutai, and Kabalalrad-
hane, grow mixed crops to fulll their individual household
needs. Any surplus crops are shared or redistributed among
the community members. Due to the inuence of Japanese
colonization, xed cultivation practices remain widespread
in this zone. To maintain soil fertility and promote sustain-
able land use, crop rotation with mixed cropping is neces-
sary. People cultivate millet mixed with other crops for two
years, followed by the cultivation of peanuts, and then sweet
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Human Ecology (2023) 51:1127–1140
Intercropping is practiced exclusively in Kabalalrad-
hane village due to the zone’s gentle slope of less than 30
degrees that provides sucient space for growing dier-
ent crops in a single plot on a terraced eld. In areas with
steeper slopes, intercropping is not practical. For peanut
mixed cropping, farmers plant the peanuts on the ridge,
with corn grown in the ditch to increase food production.
Social Institutional Support for Agrobiodiversity
Wutai is one of the few indigenous societies in Taiwan
that has maintained traditional farming practices and did
not experienced forced migration during the Japanese
and Chinese colonial periods. They retain four social
institutions that exemplify the close relationship between
social and agricultural activities that maintains agrobio-
diversity: land use norms, social reciprocity, rites and
ceremonies, and social competition for honor.
Land Use Norms
Due to the shifting cultivation practices, each household
in the Wutai region owns several small plots of land in
dierent climate zones that they cultivate with various
crops, while letting one or two lie fallow. For instance, on
interviewee reported he had seven plots scattered across
three climate zones - two in agricultural Rekai zone,
three in Kabiceacelrake buer zone, and one in warm and
hot Labelabe zone. It is customary for each household to
secure lands in dierent climate zones to diversify their
risks (Fig. 4).
Although household plots have similar natural envi-
ronments, farmers usually plant dierent crops from
their neighbors, allowing them to exchange and share
their produce after harvest. This practice not only pre-
serves more plant varieties but also connects dierent
farmlands and promotes opportunities for seed selection,
cross-fertilization, and higher regional agrobiodiversity.
Furthermore, viewed from a community perspective, this
land distribution reduces the risk of pest infestation and
signicantly reduces the risks of monocropping. The col-
lective land norm system observed in Wutai village is
an example of the multi-actor landscape approach rec-
ommended by some scholars for achieving functional
agrobiodiversity (Gerits et al., 2021). In fact, the risks
of monocropping are so signicant in an area where two
Rukai tribes resettled after the Morakot typhoon disas-
ter of 2009 (Huang et al., 2021) that some households
returned to traditional mixed cropping farming. A com-
mon saying among the villagers of Wutai recognizes the
benets of mixed cropping: “Taiwan quinoa and millet
are good companions,“ similar to the “Three Sisters” in
move to peanut mixed cropping for a year since the nitro-
gen xing capabilities of peanut rhizobia generate organic
fertilizer. Following the peanut harvest, farmers frequently
burn the remaining plant material and mix the ashes into the
soil as fertilizers. In the fourth year, farmers practice either
dry taro or sweet potatoes mixed cropping, culminating the
ve-year cycle, when a fresh cycle of millet mixed cropping
begins.
In the remaining two zones, shifting cultivation predomi-
nates due to its eectiveness in maintaining soil fertility. A
survey conducted in 1972 indicated that the average fallow
time for the Wutai area was between four and ve years
(Sasaki and Fukano, 1976/2013). However, as the popula-
tion has increased fallows have shortened.
In the cold and wet agricultural Rekai zone, upland
taro mixed cropping is favored due to its longer grow-
ing period and limited options for suitable crops. Taro
is planted in the rst year, followed by baby taro in the
second year without plowing. In the third year, the soil
is plowed and turned to prepare for baby taro planting
without plowing in the fourth year. After the harvest in
the fth year of crops like sweet potatoes and pumpkins,
the land is left semi-fallow for approximately ve years
on average to restore soil fertility. Whether or not to con-
tinue the fallow after ve years is determined by farmers’
evaluation of the land’s fertility.
In the warm and hot Labelabe zone, the growth period
of crops such as millet, peanuts, and sweet potatoes is
shorter. On sun-facing slopes, millet mixed cropping
predominates during the initial two years, followed by
peanut mixed cropping in the third and fourth years. In
the fth year, sweet potatoes unharvested for the previ-
ous four years are allowed to propagate and grow. This
ve-year crop rotation cycle continues until soil fertility
becomes depleted, after which the land is left fallow.
Short-Term Timeline: Mixed Cropping and
Intercropping in a Yearly Cycle
To maximize yearly cultivation production and minimize
the risk of insect pests, Wutai farmers commonly practice
mixed cropping and intercropping using a variety of dier-
ent seeds carefully selected to prevent spatial and nutritional
competition in a plot. For instance, in the millet mixed crop-
ping system in the Kabiceacelrake buer zone 14 dier-
ent crops are cultivated throughout the year, with harvests
occurring sequentially: corn in April, Taiwan quinoa in
April and May, millet in June and July, sorghum in Sep-
tember, cowpea and Formosan Frost Grass in October, and
pigeon pea (Cajanus) in November and December. Pump-
kin and yam are harvested at varying intervals throughout
the year, ensuring year-round food security.
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Human Ecology (2023) 51:1127–1140
Social Reciprocity: Seed Preservation and
Agricultural Product Sharing
The economic life of humans is organized by markets,
redistribution, or reciprocity (Polanyi, 1957). Although the
market economy has encroached upon Wutai for over half
a century, traditional reciprocity remains an essential guid-
ing principle for the people’s economic actions. For exam-
ple, they collectively reclaim farmland and harvest crops
through a labor exchange system. Surplus crops are given
to neighbors, friends, or relatives as gifts. In Wutai, each
household has its preferred varieties of millet, so if a house-
hold wants to grow a dierent variety, they simply ask their
neighbors. Such reciprocity spreads the risk of losing millet
varieties while collectively maintaining a diversity of seeds.
Certain root crops, including pumpkin, yam, and ground-
nut are left on fallow land to allow the stems to cover the
ground, protecting soil moisture from evaporation and pre-
serving topsoil and organisms beneath. The root crops in
these fallow elds typically comprise varieties that the vil-
lagers do not favor and consume only in cases of severe
food shortage.
Crops for Rites and Ceremonies
Since millet is used in ceremonies, the strains selected are
not necessarily those that generate the highest yields. For
instance, in Kinulane, with a population of only 167 people,
more than 20 millet varieties are planted in a small area
reecting their social and religious roles as well as agrobio-
diversity. Millet holds signicant importance as both a stable
essential and sacred crop in Taiwan’s indigenous commu-
nity and both waxy and non-waxy varieties are utilized for
dierent ritual occasions. Only specic glutinous varieties,
such as palralramu, kawapane, and makasasagarane (local
many Native American nations (Kapayou et al., 2023).
When discussing the use of traditional methods for pest
and disease control in farming, one interviewee noted:
“Millets, just like coee, can also be aected by
epidemics. I once visited [the] Guanshan area and
observed that the rice elds were infested with leaf-
hoppers, and ultimately, they had to resort to burn-
ing as a solution. Therefore, I frequently advise
young people that in cases where their millet elds
are plagued by ladybugs and leafhoppers that can-
not be eectively managed, they may ultimately
need to resort to employing traditional pest control
methods, i.e., mixed cropping methods.“
In the past, crops were grown primarily for food security.
However, due to the inuence of the market economy,
some arable land has been used to grow cash crops to
fulll expanding market demands, such as the popular
ice jelly, coee, and the traditional crop of Taiwan quinoa
(Huang et al., 2021). Although some arable land is used
for cash crops, two norms must be followed: land leased
from chiefs cannot be used for cash crops1 and cash crops
must be planted on fallow or marginal land.
When considering cash crop cultivation and the mar-
ket risks involved, Wutai farmers typically utilize fallow
land. As the population ages, more land is left unused, and
older farmers abandon distant plots that are inaccessible
by motor vehicles. Furthermore, with the importation of
many foods, there is no mass demand for traditional sub-
sistence crops.
1 Chiefs do not take cash crops as tributes, as they are unfamiliar with
the practice and not well-versed in selling such products in the market.
Fig. 4 Land ownership and
distribution by individual family
in Wutai. Notes: 1. The data
were collected from the Wutai
Township Oce and analyzed
using a GIS system to identify
lands owned by the same family
scattered across dierent zones.
To illustrate the land norms,
we selected three families as
examples. 2. Each household is
represented by a unique graphic
symbol. Round shaped lands are
owned by Kuri family, scattered
in three dierent climate zones
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Human Ecology (2023) 51:1127–1140
or nd wild foods such as mushrooms, yams, or bamboo
shoots, or fallow elds with sweet potatoes. In the spring
of 2020, the rst author accompanied a senior hunter to the
mountains to locate water sources due to insucient rain-
fall. The hunter remarked that this was the good time to nd
water sources since the land was dry, and if water was pres-
ent, it would run year-round. Hunters’ knowledge of their
micro-climate enables the community to identify and better
cope with long-term climate changes. The social prestige
accrued motivates hunters to preserve a sustainable and bio-
diverse cropping environment for the entire community.
In addition to hunting prestige, farmers who produce the
best millets in terms of weight and quality are also accorded
an honor. A competitor must exhibit over 120 bundles of
millet in public, approximately 480 kg, with full millet ears.
The winner is entitled to wear a millet-hat. Through these
social competitions and honors, the Rukai people uphold
social order and sustain their agricultural traditions.
Concluding Remarks
Set in the framework of traditional ecological knowl-
edge and the pursuit of sustainable biodiversity, our study
emphasizes the social dimensions of ecology through an
examination of the traditional farming practices and land
classication systems of the Wutai Rukai tribe in Taiwan,
elucidating the intricate interplay between agrobiodiver-
sity, social institutions, and ecological sustainability. Our
ndings reveal a multifaceted system that safeguards food
security and cultivates a thriving agricultural environment
characterized by diversity and resilience.
We identied seven distinct spaces utilized for farming,
including three zones categorized by altitude, two by the
length of sunlight, and one riverside zone. These zones are
cold and wet agricultural Rekai, cold and hot Kabiceacel-
rake buer zone, hot Labelabe zone, and the riverbank zone.
Five mixed cropping systems are practiced: millet, peanut,
sweet potato, upland taro, and paddy taro, with the last
being unique to the riverbank zone.
The Rukai people have developed sophisticated planting
methods, such as ve-year cycles involving swidden farm-
ing, rotational planting, and fallow to preserve soil fertility.
On a yearly basis, dierent crops are planted and harvested
at dierent times to ensure food security, and intercrop-
ping is practiced on atter land with slopes of less than 30
degrees.
Four local social institutions sustain agrobiodiversity
within the Wutai Rukai tribe: land norms, social reciprocity,
cultural rituals, and social competition. Land norms prohibit
planting cash crops in food cropping lands and encourage
spatially scattered land ownership to diversify risk. Social
reciprocity involves preserving dierent seed varieties,
landraces without a formal scientic classication), are used
for the harvest festival, while other varieties are reserved for
marriage or men’s ceremonies. According to a 1972 survey,
these were the most common millet varieties in the Kinu-
lane zone (Sasaki and Fukano, 1976/2013: 48).
Kyatudariti (man’s rite) is conducted by men to ensure
that their farming or hunting tools function well in the
coming year for a bountiful harvest. During the ceremony,
women are responsible for chasing away any animal or
stranger that may interrupt the ritual. The specic variety of
millet used is durai, a glutinous millet that readily adheres
to the tools and shaped like an animal’s hoof. Marriage cer-
emonies constitute one of the most signicant occasions for
social exchange among households in the same or dierent
tribes. The exchange of gifts between both parties during the
ceremony can serve to underscore the economic and social
status of each household. In contemporary times, customary
bridal gifts include a diverse range of commodities, includ-
ing millet, paddy taro, millet wine, millet cake, betel nut,
piper betel, sugar cane, and millet seeds, thus contributing
to the circulation resources and contributing to the preserva-
tion of biodiversity. Paddy taro holds such signicance that
even people who had to relocate to the plains following the
Morakot typhoon disaster in 2009 return to the mountains
to procure it whenever a wedding takes place. Paddy taro
elds are typically small and fragmented, situated close to
the river, which hampers large-scale production and conse-
quently renders it even more valuable and widely utilized in
various ceremonies to showcase the generosity and sincer-
ity of the owners who are willing to share their resources.
Planting ample paddy taro is a crucial responsibility to
support the cultural practices and to earn the respect in the
community.
Social Competition and Acquired Status
Social competition and honor are integral to Rukai society,
which is characterized by a class-based hierarchy where
chiefs and noble families enjoy inherent prestige, while
commoners can acquire social recognition through their
exceptional performance. A hunter who captures more than
ve male wild hogs and shares the meat with their com-
munity in public is entitled to wear a single wild lily as a
badge of honor. Winning a second lily requires the hunter
to capture another 20 wild hogs and bring them back to the
village by himself without assistance. Hunters play a crucial
role in supplying animal protein and micro-climate farming
information to the community, thereby gaining high status.
Embarking on a hunting expedition, they carefully observe
the micro-environmental changes, such as water ow or
plant growth, which are vital for farmers. During times of
drought or food scarcity, hunters can identify water sources,
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Human Ecology (2023) 51:1127–1140
the conservation and restoration of ecosystems. By harness-
ing traditional indigenous knowledge and cultural prac-
tices, society can preserve its agrobiodiversity system and
enhance its capacity to cope with and adapt to extreme cli-
mate change.
Although the transition to commercial cropping is a
common global phenomenon, it does not inevitably lead to
signicant declines or losses in biodiversity (Rerkasem et
al., 2009), as is the case in Wutai. However, Wutai soci-
ety is also experiencing signicant social transformation.
Approximately 40% of their land is now dedicated to
cash crop cultivation, which is primarily managed by the
younger generation, aged below 50 years old. Traditional
crops are no longer nancially viable for sustaining their
households. These younger farmers employ distinct cultiva-
tion methods that dier from those practiced by the older
generation. For example, older farmers adhere strictly to
organic coee cultivation, avoiding the use of fertilizers and
embracing organic practices to maintain soil fertility. How-
ever, the younger generation argues that organic cultivation
may compromise the avor of coee. A young farmer we
interviewed insisted that coee requires substantial nutrient
inputs that may not be adequately supplied through organic
cultivation methods. Future research should monitor the
impact of market-oriented agricultural production on both
the local ecology and social relations and its implications
for sustainability.
We should note two limitations inherent in our research.
Firstly, we rely primarily on ethnographic data collected
during a specic time period, which may restrict the ability
to capture long-term dynamics and changes in agrobiodi-
versity farming practices. Conducting longitudinal stud-
ies and gathering extended-term data could oer a more
comprehensive understanding of these practices over time.
Secondly, while our research focuses on the integration of
agrobiodiversity farming practices into social institutions
and cultural norms within the Rukai community, it does not
extensively explore other inuential factors such as market-
oriented economic activities, national policy frameworks, or
the impact of an aging population on the sustainability and
eectiveness of these practices.
Acknowledgements We would like to express our sincere apprecia-
tion to Professor Ludomir Lozny, the Managing Editor of the journal,
for his dedicated eorts in polishing our article. We also extend our
gratitude to the two reviewers for their insightful comments, which
have signicantly enhanced the overall quality of the paper. Any errors
in the paper remain the sole responsibility of the authors.
Author Contribution The rst author conducted the fundamental eld
survey and documentation, particularly regarding crop investigation
and agricultural cultivation methods. Based on the eld data collected
by the rst author, the second author collaborated with the other two
authors to discuss the structure and analytical concepts of the entire
paper. The rst and second author drafted and revised this article. The
planting dierent crops from neighbors, and exchanging
products after harvest. Cultural rituals require dierent
crops for various ceremonies, such as the Kyatudariti cere-
mony and weddings, which circulate dierent plants among
communities and maintain agrobiodiversity. Social compe-
titions and honors for e.g., hunting success and producing
the best millets, motivate community members to preserve a
biodiverse agricultural environment and uphold their social
order.
By integrating traditional ecological knowledge with
intricate social cultivation management systems, the agri-
cultural system of the Rukai in the Wutai region achieves
two objectives: (1) ensuring the maximum production of
food crops throughout the year, and (2) reducing the risk of
decreased food production due to climate change. The for-
mer is accomplished through agricultural practices such as
swidden farming, fallow periods, mixed cropping, rotational
cropping, and intercropping, which contribute to mainte-
nance of soil fertility and pest control. The latter is achieved
by institutionally distributing the same household’s lands
across dierent climatic zones and through social reciproc-
ity to mitigate the impacts of extreme weather conditions.
In response to the risks imposed by extreme climatic con-
ditions, the Rukai indigenous community employs two dis-
tinct strategies to address food security concerns. The rst
strategy involves agricultural practices that minimize poten-
tial impacts on crop yields, such as land distribution across
dierent climate zones and deliberately leaving certain crop
varieties (e.g., tuba, kalri, kelrenge) that are less favored
for consumption unharvested until they are needed during
periods of food insecurity, underscoring the community’s
commitment to preserving and utilizing diverse crop variet-
ies irrespective of their economic protability. The second
strategy to address food security risks centers on a coopera-
tive framework guided by principles of moral economics.
Through reciprocal exchanges and food redistribution, the
community establishes a collaborative system to eectively
tackle food security challenges (Ba et al., 2021).
Our Wutai case study holds signicant implications for
climate change adaptation, ecological sustainability, con-
servation, and the recognition of traditional knowledge
as a valuable resource in addressing climate change chal-
lenges and promoting resilient and sustainable agricultural
systems. The observed agrobiodiversity farming practices,
such as mixed cropping methods, small land plots across
dierent climate zones enhances habitat heterogeneity and
provides ecological niches for a variety of plant species. The
sustainable land management practices, including swidden
farming, rotational plantation, and fallow periods, contrib-
ute to soil fertility and ecological sustainability. The inte-
gration of agrobiodiversity practices into social institutions
and cultural norms, like the sacred Rekai zone, facilitates
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Human Ecology (2023) 51:1127–1140
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third author participated in at least three eld trip interviews in Wutai
and engaged in thorough discussions with the other two authors re-
garding the structure and analytical concepts of the paper, after reading
the initial draft.
Funding This research was supported by the National Science and
Technology Council, Taiwan, under the collaborative three-year re-
search project “Socially embedded indigenous agriculture: produc-
tion, distribution and consumption of agricultural products in Wutai,
Taiwan.“ (National Science and Technology Council, 112-2420-H-110
-002; 112-2420-H-110 -003; 112-2420-H-110 -004).
Data Availability Photos in the paper, as well as a copy of the research
application approval document for the land ownership data provided
by the Wutai Township Oce, are available upon request.
Declarations
Ethical Approval The research projects of the three authors have
been approved by the National Cheng Kung University Governance
Framework for Human Research Ethics (No. 110–494, 110–499, and
110–501, respectively). The authors adhere strictly to the codes and
guidelines established by the Committee. For further information,
please refer to the following website: https://rec.chass.ncku.edu.tw/en/
node/19140.
Competing interests The authors declare no competing interests.
Open Access This article is licensed under a Creative Commons
Attribution 4.0 International License, which permits use, sharing,
adaptation, distribution and reproduction in any medium or format,
as long as you give appropriate credit to the original author(s) and the
source, provide a link to the Creative Commons licence, and indicate
if changes were made. The images or other third party material in this
article are included in the article’s Creative Commons licence, unless
indicated otherwise in a credit line to the material. If material is not
included in the article’s Creative Commons licence and your intended
use is not permitted by statutory regulation or exceeds the permitted
use, you will need to obtain permission directly from the copyright
holder. To view a copy of this licence, visit http://creativecommons.
org/licenses/by/4.0/.
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