Effects of Landscape Structure on Medicinal Plant Richness in … · 2018-09-26 · Effects of Landscape Structure on Medicinal Plant Richness in Home Gardens: Evidence for the Environmental

Effects of Landscape Structure on Medicinal Plant Richnessin Home Gardens: Evidence for the Environmental ScarcityCompensation Hypothesis

MONIKA KUJAWSKA*,1, FERNANDO ZAMUDIO2, LÍA MONTTI

3,4, AND

VERONICA PIRIZ CARRILLO5

1Institute of Ethnology and Cultural Anthropology, University of Lodz, ul. Lindleya 3/5, 90-131, Lodz,Poland2Interacciones ecológicas y conservación, Instituto Multidisciplinario de Biología Vegetal (CONICET-UNC), Av. Vélez Sarsfield 299, 5000, Córdoba, Argentina3Instituto de Investigaciones Marinas y Costeras, UNMdP-CONICET & Instituto de Geología de Costasy del Cuaternario, UNMdP-CIC, Deán Funes 3350-CP 722, Buenos Aires, Argentina4Instituto de Biología Subtropical, UNaM-CONICET, Bertoni 85, CP 3370, Puerto Iguazú, Misiones,Argentina5Tucuman, Argentina*Corresponding author; e-mail: [email protected]

Introduction

Over the last few decades, there has been a surgeof home garden studies utilizing diverse methodo-logical approaches and conceptual frameworks.Scholars have formulated questions related to plantuse, richness, diversity, composition, and structurewithin these social-productive units (Padoch and de

1 Received 26 July 2017; accepted 2 May 2018;published online 30 May 2018

Electronic supplementary material The online versionof this article (https://doi.org/10.1007/s12231-018-9417-3) contains supplementary material, which is avail-able to authorized users.

Economic Botany, 72(2), 2018, pp. 150–165© The Author(s) 2018

Our research involves of how Paraguayan migrants who are living in Misiones, Argentina,managemedicinal plants in home gardens, and how this practice can be related to the landscape.We examine the relationship between the richness of home garden medicinal plants andlandscape variables (e.g., distance to the forest) by applying PLS analysis, which combinesprincipal component analysis with linear regression. We surveyed 60 home gardens localized ina rural area, and we characterized the surrounding landscape with geospatial tools. Paraguayans’home gardens are extremely diverse sites (total of 136medicinal species), where both native (82)and introduced species (50) are managed. People who live close to the native forest or mixed useareas (e.g., farms, secondary vegetation) tend to possess less native plants in their gardensbecause they are available nearby. While gardeners, who live in proximity to tree crops (e.g.,pine plantations), have reduced access to wild medicinal resources; therefore, their effort isconcentrated on maintaining native plants. These results reflect a relationship between accessi-bility to medicinal plants in the landscape and the management practices in the home gardens, aneglected driver in explaining the richness and composition of the medicinal plants in homegardens so far. Thus, we contributed evidence in support of the environmental scarcity compen-sation hypothesis. Finally, our study supports the idea that home gardens appear to function as aspringboard for plant domestication.

Key Words: Agrobiodiversity, Domestication, Ethnobotany, Geospatial tools, Land use, Mi-grants, Paraguayan farmers, Plant management, Atlantic Forest.

Jong 1991; Parihaar et al. 2014; Smith et al. 2006;Sujarwo and Caneva 2015). It has been acknowl-edged that both intrinsic and extrinsic factors affectthe richness of species in home gardens. Commonintrinsic factors are, for example, the size of thegarden, its age, and its proximity to the dwelling,among others (Blanckaert et al. 2004; Coomes andBan 2004; Sander and Vandebroek 2016). On theother hand, research indicates that factors extrinsicto the garden, such as distance to urban centers,kinship, size of family groups, forms of manage-ment, and networks of exchange, may explain thespecies richness and diversity of home gardens(Díaz-Reviriego et al. 2016; Ellen and Platten2011; Lamont et al. 1999; Peroni et al. 2016;Sander and Vandebroek 2016). For example, stud-ies have shown that peri-urban and urban homegardens may host a greater diversity of species thanthose situated in remote places. This is explained bywider opportunities for knowledge and plant ex-change, especially in intercultural contexts(Bernholt et al. 2009; Heckler 2007; WinklerPrins2002). More recent studies, however, have demon-strated that biodiversity in home gardens differenti-ates along the rural-peri-urban gradient, withoutnecessarily increasing or declining (Peroni et al.2016; Poot-Pool et al. 2015). While the richnessof useful tree and shrub species, mainly native ones,tends to decrease, the number of herbaceous species,especially introduced ornamental ones, increases inurban space (Poot-Pool et al. 2015).

Home gardens are one of the oldest forms of landuse (Kumar and Nair 2004). They represent reser-voirs of currently cultivated and potentially usefulresources (Alvarez-Buylla et al. 1989), and provide awide range of ecosystem services (Calvet-Mir et al.2012). Importantly, there is a considerable consen-sus in favor of acknowledging rural and urban gar-deners for their in situ preservation of biological andgenetic diversity (Galuzzi et al. 2010; Heraty andEllstrand 2016; Kumar and Nair 2006; Saleko et al.2014). Moreover, home gardens are suitable placesfor the study of gardeners’ choices concerning theselection of plant species for protection or evencultivation, and are very interesting sites for researchon plant domestication (Blanckaert et al. 2004;Casas et al. 1996; Smith 1996).

As some studies suggest, a large proportion ofnative plants are moved to and cared for in domesticgardens (Blanckaert et al. 2004; Larios et al. 2013;Peroni et al. 2016). In addition, a remarkable pro-portion of spontaneously growing plants, especiallymedicinally useful ones, are spared by gardeners

(Blanckaert et al. 2004). This is explained by thestrong interaction between home garden plants andthe surrounding vegetation (Blanckaert et al. 2004).While there are studies analyzing the relationshipbetween landscape and human accessibility, such asdistance from human settlements and plant diversi-ty in the local landscape (see Thomas et al. 2009),little attention has been paid so far to the relation-ship between the structure and composition of thesurrounding landscape and the diversity and man-agement of home gardens. Haines-Young (2009)affirms that understanding of the relationships be-tween land use and biodiversity is fundamental tounderstanding the links between people and theirenvironment, while the way in which land is man-aged are key drivers of changes in biodiversity.Hence, changes in land use influence the structure,function, and dynamics of socio-ecological systems(Orozco et al. 2015).

Access to medicinal resources may be influencedby the structure, composition and distribution ofnative vegetation, productive activities related toland use (agriculture, forestry, etc.), the presenceof roads, the intensity of use of these resources,and the requirements and habits of medicinal spe-cies. Deforestation itself affects access to these spe-cies by decreasing availability (see Shanley and Luz2003). While the debate about the relative contri-bution of primary and secondary vegetation to theacquisition of medicinal plants persists, evidencebacks up the hypothesis that disturbed vegetationmay constitute a preferred habitat for collectors andusers of medicinal plants (Gavin 2009; Voeks1996).Moreover, Gavin (2009) estimates that land-scape that includes forests of different ages canmaximize the availability of medicinal plant species.However, the effect of different vegetation typesof landscape on the diversity of home gardens isan emerging focus. The only information avail-able in this regard has been outlined by Larios andcollaborators (2013), who found that the highestdiversity was recorded in home gardens where theneighboring forest had the least diversity and viceversa. The authors conclude that local people main-ly use their home gardens to manage plant speciesthat are not available in the wilderness close to theirtowns. This phenomenon has been named theBscarcity compensation effect^ (see Larios et al.2013).

Our research has evolved around the twodominant questions of how local people man-age plants in their home gardens and how thispractice can be related to the surrounding

151KUJAWSKA ET AL.: LANDSCAPE STRUCTURE AND HOME GARDEN PLANT RICHNESS2018]

landscape structure. Additionally, we wouldlike to ask whether home gardens may becomea convenient venue for the domestication ofnative medicinal plants and the pre-adaptionor acclimation of potentially invasive alien spe-cies (see Marco et al. 2008).In this paper, we propose an exploratory

approach to the integration of ethnobiologicalresearch and geospatial tools in order to studyaccess to medicinal plants by means of analysisof the landscape around home gardens. Weworked in the home gardens of Paraguayanmigrants who live in the Misiones provincein Argentina.Our hypothesis is presented as follows: given that

Paraguayan migrants live in a landscape that is socio-productively homogenous on a large scale, they culti-vate and use a similar diversity of medicinal species intheir home gardens. However, the landscape struc-ture close to home gardens influences the richnessand status of medicinal plants in them, due toaccess to patches of different types of vegetation,from which plant material is obtained.Our prediction is that home gardens surrounded

by a continuous area of native forest or mixed useareas (i.e., small farms with a mosaic of subsistencecrops, secondary vegetation, pastures and patches ofdegraded native forest), or located inside a hetero-geneous landscape, have less richness of native me-dicinally useful species in their home gardens com-pared to home gardens located in the vicinity ofproductive forests of introduced species (e.g., Pinusspp.), due to the scarcity compensation effect. Thuspeople living in areas with a high proportion ofnative forest and mixed use vegetation, or with easyaccess to them, can find their native plant resourcesin nearby areas instead of having to cultivate orprotect them in their home gardens. On the otherhand, we do not expect a clear pattern concerningthe richness of introduced species, because these arepreviously acclimatized and domesticated plants,easily available due to their widespread use in theregion, and which are chiefly obtained throughsocial networks, not from the landscape (Furlanet al. 2016).Therefore, our objectives are to (1) evaluate

the richness, status (native versus introduced)and management (cultivated, protected,transplanted, etc.) of medicinal plants foundin the gardens of the Paraguayan immigrantsin Misiones province; (2) describe the structureand characteristics of the nearby landscapewhere the home gardens of Paraguayan

migrants are found; (3) evaluate the potentialrelationship between the richness of native andintroduced medicinal species tended in homegardens and landscape variables (e.g., distanceto the native forest).

Methods

STUDY AREA

Our study area is the Misiones province locatedin the Argentinean Northeast. It is comprised ofextensive native forest cover, next to former forestland currently used by companies (pine monocul-tures and processing factories), as well as medium-size cash crop production (Ilex paraguariensis A. St -Hil., Citrus spp. or Camellia spp.) complementedwith animal husbandry (Hilgert et al. 2016;Mastrangelo et al. 2011).The native cover of the total study area is semi-

deciduous Atlantic Forest. The climate is subtropi-cal humid with no dry seasons, a mean annualrainfall of 2000 mm and a mean annual tempera-ture of 20 °C (Campanello et al. 2009). The forestis characterized by the presence of three well-defined canopy strata including more than 70tree species, usually covered with numerouslianas and epiphytes, and mixed with shrubs,bamboos, and grasses (Campanello et al. 2009;Chediack 2008). Currently nearly 40% of the orig-inal forest cover is preserved in Argentina, in con-trast to Brazil and Paraguay, where it comprisesapproximately 8% (Galindo-Leal and Câmara2003). However, native forest cover is decreasing,due to an increasing population size in the regionand the increasing demand for wood productscoupled with the reduced economic value of theremaining forests—the result of uncontrolled selec-tive logging (Campanello et al. 2009). Forest plan-tations, mainly Pinus and Eucalyptus species, havegrown in area from 80,000 to 370,000 ha, anincrease of 1 to 11% of the provincial territorybetween 1973 and 2006 (Izquierdo et al. 2008).These forest plantations are implemented and man-aged by large forestry companies and to a lesserextent by small and medium producers (Hilgertet al. 2016). Concurrently, the remaining naturalforest persists principally within protected and un-cultivated areas (e.g., high slopes or bad quality soil).The native vegetation also persists as degraded pri-mary forest (capueras) and secondary forest along-side the farmland (Holz et al. 2009).

152 ECONOMIC BOTANY [VOL 72

ETHNOGRAPHIC SETTING AND DATA

COLLECTION

Since the 19th century, mestizos (criollos) havebeen coming to Misiones province from the southof Argentina (Corrientes) and from neighboringBrazil and Paraguay. Since 1947, Paraguayans havebeen the most populous migrant community inArgentina. As of 2001, they have accounted forapproximately 550,713 people, representing30.5% of all immigrants in the country, and nearly8.6% of the total population of Paraguay (INDEC2010; Torres 2014). The push and pull factors formigration have been of both economic and politicalcharacter (i.e., the Chaco War in 1936, the Para-guayan Civil War in 1947, and long years of mili-tary dictatorship ending in 1989), leading Paraguay-an people to seek agricultural employment oppor-tunities in the production of cotton, sugar cane, tea,and tobacco in the Argentinian northeast (Benencia2007). Paraguayan migrants exhibit significant cul-tural cohesion in the study area, unified bylanguage—most speak the native Guarani, inter-marry, and have cultural institutions that keep themtogether (e.g., groupings of family residences, cul-tural associations, e.g., in Puerto Piray).

A total of 60 home gardens were surveyed alongthe study area. Home gardens are understood hereas a front and backyard (Spanish: el patio y el huerto)adjacent to a home. In the case of Paraguayan mi-grants’ home gardens, the backyard can be partly orfully fenced, with some herbs (e.g., oregano, chive)and vegetables cultivated in it. Trees, ornamental,and medicinal plants usually grow in both the frontand backyard (based on field observations).

Home garden inventories of medicinal plantswere performed among 44 women (aged 30–95)and 16 men (aged 45–90). Adults participated vol-untarily, recruited by snowball technique. Forty-three of the study participants hail from the ruralareas of Eastern Paraguay, which belongs to thesame ecoregion of the Atlantic Forest, while 17 wereborn in Argentina. Their parents were of Paraguay-an origin and spoke Guarani, and all but two couldspeak Spanish. Fifty-two participants (87%) de-clared not having completed primary school, whileonly six have completed primary and two havecompleted secondary school. Forty-three partici-pants claimed to have health insurance, while 17declared that they did not have any. In generalterms, the study participants can be described assmall scale farmers working on 1–2 ha plots(chacras), where they cultivate staple crops such as

manioc and maize, often intercropped with pea-nuts. Additionally, male participants usually findoff-farm employment, mainly in the forest industry.

In most cases, the first author collected medicinalplants in the company of study participants. All ofthe documented botanical taxa are supported withherbarium specimens identified by the first authorand stored in the CITS herbarium in the Institutode Botánica del Nordeste (IBONE), Corrientesprovince, Argentina. Prior to each interview, studyparticipants gave oral informed consent for theirparticipation. Although there were no requirementsfor the project to pass any ethical commission,either in Argentinean or Polish institutions, the firstauthor obtained permission from the Ministry ofEcology in Posadas, capital of Misiones, to collectherbarium specimens and for their transfer to theherbarium in Corrientes.

SELECTION OF STUDY SITES AND LANDSCAPE

ANALYSIS

In order to achieve the greatest variability of theParaguayans’ home gardens while also concentratingon work-sampling effort, we worked in and aroundthree sites along the coast of the Parana River whereParaguayan migrants live (Fig. 1): (1) North of thestudy area within and around the municipal town ofColonia Wanda; (2) Center of the study area, calledPiray Km 18, within the Puerto Piray municipality,which is a small settlement composed of farmhousesand small cultivated plots, and (3) South of thestudy area, near Puerto Leoni, a municipal town,the smallest of the three localities. These three sitesare representative of the socio-productive landscapeof the area and have a high concentration of Para-guayan migrants. The home gardens are located in arural and semi-rural environment, near small towns,which have populations ranging from 2600 to15,911 inhabitants (INDEC 2010).

In order to evaluate and quantify the landscapestructure that surrounds the study home gardens, weused the land cover map ofMisiones created by Zuletaet al. (2015). Land covers were re-classified into 10classes: (1) Urban (Ur = town and other built-upareas), (2)Native Forest (Nf = native forest), (3) Treecrops (Tc = Pinus spp., Eucalyptus spp.), (4) Shrubcrops (Sc = Ilex paraguarensis—yerba mate, Citrusspp., Camellia spp.—tea, and other perennialcrops), (5) Annual crops (Ac = Glycine max (L.)Merr.—soybean, Zea mays L.—corn, and otherherbaceous crop species), (6)Mixed use areas (Mx =small farms with a mosaic of subsistence crops,


Mixed use areas (Mx)

Fig. 1. The greater study area in Misiones Province, Northeast of Argentina. In detail, the three study sites of thehome gardens, each 25 km2 surface area showing the surrounding sampled land cover.


pasture, secondary vegetation, and patches of de-graded native forest, (7) Wetland (Wl), (8) Pasture(Ps), (9) Water bodies (Wt = rivers, streams, lagoonsand artificial water bodies), and (10)Others (naturaldeforested areas, bare soil, and not classified pixels).

Local people in the study area obtain medicinalplants from different vegetation units. Some of thespecies are exclusively obtained from the NativeForest (Keller 2008; Kujawska and Pardo deSantayana 2015), although mixed use vegetationhas been found to be a site where many ruderalspecies of medicinal interest are harvested(Kujawska et al. 2012). Medicinal plants are alsocollected from farmland where perennial shrubs andannual crops are cultivated, but in lesser proportionthan from forest and ruderal areas, due to use ofpesticides and weeding practices (Kujawska andPardo de Santayana 2015). Only a very limitednumber of medicinal species, e.g., Nasturtiumofficinale R.Br. grow in the wetlands. The sponta-neous vegetation of the tree crops areas contributesto the least favorable conditions for the growth ofmedicinal species as it is the product of intensivemanagement with the use of agrochemicals. As aresult, the soils are extremely acidic by the deposi-tion of acyclics, and this is also a very shady envi-ronment (Frank and Finckh 1997).

Beyond this classification, we characterized thelandscape structure into three surface samples of25 km2 (one per locality), including all homegardens (Fig. 1). A study of landscape features andcomputed descriptive statistics was conducted usingArcGis 10.1, Spatial Analysis tools and FRAGSTATSsoftware (McGarigal and Marks 1995). The selec-tion of metrics was based on their ability to charac-terize various aspects of fragmentation such as totalarea of fragments per class, mean size of patch (indexof fragmentation), number of patches (indicator ofseveral ecological processes and landscape heteroge-neity), total edge (indicator of patch complexity),average nearest-neighbor distance and mean prox-imity index (see McGarigal et al. 2002).

DATA ANALYSIS

In order to test the main hypotheses, we exploredthe relationship between the landscape variables (i.e.,distance to different types of vegetation where medic-inal plants can be found) and the richness of nativeand introduced species found in home gardens. Fig-ure 2 shows a schematic representation of ourmethods and a conceptual framework of our meth-odological proposal. Using the multiple minimum

distance tools from ArcGis 10.1, we calculated theshortest distance between home gardens and differ-ent land-use classes (i.e., Ur, Nf, Tc, etc.). Wemeasured distance between each home garden anddifferent vegetation classes exceptWetland, Pasture,and Water bodies because these are poorly repre-sented in the region and we used a new distancevariable called Accr (access to national and/or pro-vincial routes and secondary paved roads) to per-form an independent analysis.

To evaluate the influence of factors associated withthe landscape on the richness of native and introducedspecies in the home gardens, a partial least squaresregression (PLS) was performed. This analysis is par-ticularly appropriate when several predictors are corre-lated with each other and the number of cases isrelatively low (Haelein and Kaplan 2004). PLS,which generalizes and combines principal compo-nent analysis with linear regression, consists in find-ing axes or latent variables that maximize the varia-tion of the matrix of dependent variables, associatedwith the variation of the matrix of predictor vari-ables. After generating latent variables with greaterexplanatory power, simple linear regressions wereperformed between the latent variables and the sixpredictor variables of the model. This was done inorder to evaluate the degree of relationship betweenthese latent variables and predictor variables, andthus, we were able to identify the variables that bestexplain the variation in the richness of native andintroduced species of Paraguayans’ home gardens.All analyses were performed using the Infostat/Pv2010 program (Di Rienzo et al. 2010).

To evaluate the management and the role ofhome gardens as social ecological spaces where theprocesses of domestication of native plants mayoccur, we performed a classification of the speciesaccording to the type of acquisition managementthat they were subjected to (Casas et al. 1996).Consequently, we divided the plant managementinto two stages: initial management and furthermanagement. In the first stage, we distinguishedpractices, such as planting, sowing, and direct trans-plant from other habitats. In the second stage, wedistinguished two major practices, namely, cultiva-tion and protection (see: Blanckaert et al. 2004;Casas et al. 1996; Lins Neto et al. 2014).

In this study, we used the etic notion of nativeand introduced species. We did not consider theperception of resources as familiar or new by studyparticipants incorporated within living memory (seeHeckler 2007; Peroni et al. 2016). Therefore, in-troduced plants are exotic species to the local flora,


which arrived in Argentina with or without the helpof people from the area. The list of native andintroduced species was elaborated based on theFlora de Conosur (http://www.darwin.edu.ar/Proyectos/FloraArgentina/Familias.asp). With thisanalysis, we wanted to see whether migrants origi-nating from a similar environment (i.e., Paraguay)contribute to the maintenance of diversity of nativespecies in the host country (i.e., Argentina).

Results

RICHNESS AND THE STATUS OF MEDICINAL

PLANTS IN HOME GARDENS

The overall richness of medicinal plants found in60 Paraguayan home gardens corresponds to a total

of 136 species with an average of 11.1 per homegarden, the maximum being 24 and the minimumbeing 3 species (Appendix 1, ElectronicSupplementary Material, ESM). The native taxa(82) outnumber the introduced ones (50). Fournaturalized species are cultivated, with an averageof 0.2 species per home garden. Native and intro-duced species are equally represented when analyz-ing the average per home garden (5.4 and 5.6,respectively). The home gardens that have moreintroduced species than natives represent 47% ofthe total, while those that have more native thanintroduced species, or the same proportion, reach35 and 11%, respectively. Notably, gardens withmore native species present a proportion up to threetimes greater than those that have a prevalence ofintroduced species. However, these are not

Fig. 2. Synthesis of methods used in the analysis of the environmental scarcity effect.


http://www.darwin.edu.ar/Proyectos/FloraArgentina/Familias.asp

http://www.darwin.edu.ar/Proyectos/FloraArgentina/Familias.asp

necessarily the most diverse gardens. For example, ahome garden with 5 native and 1 introduced speciesequals a proportion native/introduced of 5 (Fig. 3).

The species with the highest frequency of occur-rence was the native Lippia alba (Mill.) N.E.Br. exBritton & P.Wilson and was present in 32 homegardens, accounting for more than 50% of all thestudied home gardens. However, in general terms,introduced species exhibit greater frequency of oc-currence than natives, with six species present in atleast 1/3 of home gardens: Aloe maculata All.,Cymbopogon citratus (DC.) Stapf, Ruta chalepensisL., Rosmarinus officinalis L., Artemisia absinthiumL., and Mentha spp. Among the naturalized plants,Citrus x aurantium L. was mentioned with thegreatest frequency, but it was found in only sixgardens (Appendix 1, ESM).

LANDSCAPE STRUCTURE

The landscape configuration in proximity to thestudy home gardens (25 km2 surface area) differsaccording to the surrounding areas (Fig. 4a, b).Home gardens from the Northern study area areincluded in the largest urban area, but at the sametime are surrounded by large areas of non-fragmented native forest. The home gardens in thecenter study area are located within a homogeneousand monospecific landscape, due to a high propor-tion of non-fragmented tree crops cover. Nativeforest is very fragmented compared to the Northand South areas, which may be due to the presenceof tree crops and annual crops. The Southern areadisplays a more heterogeneous landscape. Thehome gardens within this area appear to be

Fig. 3. Examples of Paraguayan farmers’ home gardens and landscape, in which they are inserted. a A view onmaizeand cassava field and pine productive forest from fenced part of a home garden in Piray Km 18. b Householdsurroundings in Puerto Wanda. Fruit trees and other tree species are in the home garden. c Medicinal plant cornerin a home garden of PuertoWanda,Misiones:Cymbopogon citratus, Ruta chalepensis, Rosa sp.,Alternanthera brasiliana. dPart of a home garden with edible and medicinal plants in Puerto Leoni, Misiones.


Fig. 4. a Land class uses surrounding the home gardens in the three surveyed areas. b Different indices of spatialpatterns of landscape. Bars indicate cover use classes. Ur (Urban area); Nf (Native forest); Tc (Tree crops); Sc (Shrubscrops); Ac (Annual crops); Mx (Mixed use area). Wetlands and Pasture classes were not represented because theyconstituted a very small portion of the total analyzed area.


surrounded by the most transformed landscape, thearea having been very fragmented and transformedinto different productive units.

RELATIONSHIP BETWEEN RICHNESS, STATUS, AND

LANDSCAPE VARIABLES

The PLS analysis shows that the first two latentvariables explain 90.4% of the variation of therichness of native and introduced species of medic-inal plants in the home gardens of Paraguayanimmigrants in Misiones (Fig. 5). The first latentvariable (Y axis) explains 64.6% of the variation. Itis negatively associated with distance to the mono-cultures of tree crops (Tc) (R2 = 0.53; F = 64.68;P 0.001). It has a positive association with thedistance to two land cover classes: mixed use areas(Mx) (R2 = 0.42; F = 41.70; P 0.001) and shrubcrops (Sc) (R2 = 0.35; F = 30.56; P 0.001).

The second latent variable (X axis) comprises25.8% of the variation and is associated negativelywith shrub crops (Sc) (R2 = 0.42; F = 42.16;P 0.001), and positively to the distance to the

routes and roads (R2 = 0.42; F = 41.46; P 0.001)and to a lesser extent positively to the mixed useareas (Mx) (R2 = 0.19; F = 13.30; P = 0.001). Theobserved trend suggests that gardens closer to shrubcrops (Sc) have more introduced species than thosegardens further away, whereas introduced speciestend to increase as the gardens drift away from theroads. Finally, the first variable is positively andsignificantly associated with the native species vector(NF) although with a weak R-squared value (R2 =0.18; F = 13.08; P = 0.001).

MANAGEMENT AND DOMESTICATION OF

MEDICINAL PLANTS IN PARAGUAYANS’ HOME

GARDENS

Different methods of plant management wereobserved and registered in this study. They can bedivided into two kinds: those requiring initial man-agement and those that do not require planting.Within the first group of plants are those that needto be planted through cuttings and other plantpropagules, directly transplanted from other

Fig. 5. Tri-plot of the first two latent variables obtained in the partial least squares analysis (PLS) performed with thetwo dependent variables: richness of native and introduced medicinal species (green circles) from 60 Paraguayan homegardens (black circles). The dotted vectors (dependent variables) indicate the magnitude (length) and direction of thechange of the predictor variables distance to different land uses and access to national and/or provincial routes andsecondary paved roads.


habitats, or sown from seeds. These planted speciesaccount for 81% of all taxa identified in the homegardens. The planted, sown, and transplanted fromother habitats species may become cultivated—andthe further activity of a gardener is required, such aswatering, weeding, providing shade or sun exposi-tion, or creating barriers for domestic animals. Cul-tivated plants account for 69% of all medicinalspecies found in the home gardens of this study.The second group of plants in home gardens areprotected species (19%). These plants do not re-quire any initial management, they are sown byanimals or the wind, or are plants of early succes-sion, and therefore, it is the decision of a particulargardener to protect them or even convert them intoa cultivated species—by providing further tending.Protected species are mainly natives. There are someoverlaps between these two groups of cultivated andprotected plants—as some species are cultivated bysome and just protected by others. The full scope ofmanagement of 136 recorded species is presented inAppendix 1, ESM.

Discussion

LANDSCAPE STRUCTURE AFFECTS MEDICINAL

PLANT DIVERSITY IN HOME GARDENS

The results show an under-studied relationshipbetween home gardens and the surrounding land-scape, and provide novel findings to disciplines thathave until now been only loosely related, i.e., eth-nobiology and landscape ecology. In our study, anassumption of relationships between land use andbiodiversity has been demonstrated and extended tothe domestic sphere. Moreover, we contributedevidence to support the environmental scarcitycompensation hypothesis. This operates in the fol-lowing way in the medicinal context; gardeners wholive in the proximity of native vegetation have noneed to cultivate and protect these resources in theirhome gardens because they have easy access to themin the nearby landscape.As we could demonstrate, native species are bet-

ter represented in gardens situated farther frommixed use areas and are less represented in gardensfound farther from tree crops. This remains inagreement with the first prediction that a greaternumber of native species can be found in homegardens next to tree crops. Gardeners who live inrelative proximity to monoculture forest plantationsof pine or eucalyptus, and are inserted in a less

heterogeneous landscape, have lesser access to wildmedicinal resources. Therefore, their gardening ef-forts are concentrated on planting, transplanting,sowing and protecting native medicinal plantsthat cannot be found easily in their surround-ings. Previous research in our study area re-ported that 30% of medicinal species used inherbal mixtures with honey by mestizo peopleand Polish migrants originated from habitatscharacteristic of mixed use vegetation, beingsuperior to the medicinal species obtainedfrom the native forest (19%) (Kujawska et al.2012). This finding reinforces the relevance of sec-ondary vegetation in the provision of medicinalresources (see Gavin 2009), and thus explains therelatively weak relationship between the native for-est and the richness of medicinal plants in ouranalysis (Fig. 5).On the other hand, variable distance to shrub

crops and to roads contributes to the explanation ofthe presence of introduced species, although theseare not correlated to the second latent variable of themodel. Therefore, the richness of introduced speciesin home gardens would not be fully explained bylandscape variables associated with the availability ofmedicinal resources. At this point, we assume thatsocial webs of exchange of propagules and seeds mayhave prevalence over landscape factors (see Díaz-Reviriego et al. 2016). However, this aspect hasnot been studied in this research.Although with some exceptions (Peroni et al.

2016), a considerable number of studies have re-ported greater biodiversity richness in peri-urbanand urban home gardens than in those situatedfarther from urban centers (Bernholt et al. 2009;Heckler 2007; WinklerPrins 2002). This was ex-plained by a greater possibility of exchange ofknowledge and plant material among the homegardeners. However, those studies did not take intoaccount the possible effect of landscape on plantavailability and therefore access to these resources(see Gaoue et al. 2017). On the other hand, Wezeland Ohl (2005) explain that the daily travel offarmers to swiddens and cultivated plots mayreduce the need for diverse home gardens in ruralareas. Larios et al. (2013) found that the mostdiverse Nahuatl home gardens were located in areaswith less diverse vegetation, suggesting the impor-tance of the scarcity compensation effect. Similarresults were found by Poot-Pool and collaborators(2015), who observed that people also tend to growmedicinal resources, which are not directly availablein their surrounding landscape. Although these


studies contribute to understanding of the influenceof the landscape on the diversity of home gardens,our results are the first to provide conclusive data infavor of the scarcity compensation hypothesis onthe diversity of medicinal plants. While the pre-dominant view is that home gardens emulate natu-ral vegetation in their interior, as models of agrofor-estry systems (Larios et al. 2013; Peroni et al. 2016),according to our results home gardens can be con-sidered as a complement to what is not available inthe environment.

These findings provide new arguments in theethnobotanical debate concerning relevant hypoth-eses, such as the hypothesis of ecological appearanceor the plant use value hypothesis, among others(Albuquerque 2006; Gaoue et al. 2017; Gonçalveset al. 2016). Our results indicate that less abundantresources in the environment are not necessarily lessused, because a portion of these resources, probablyintensely used ones, are transferred to home gardenswhere they are readily available for use. Thus, thehypothesis of ecological appearance makes sensewhen it takes into account species occurring natu-rally in the landscape as well as tended in homegardens. As we could observe, the landscape acces-sibility of wild medicinal resources motivates man-agement practices that will finally affect the usepatterns of the species, regardless of whether theyare available or not in the vicinity of the familyresidence.

THE DOUBLE ROLE OF HOME GARDENS: THE

DOMESTICATION OF NATIVE SPECIES AND

ACCLIMATIZATION OF INTRODUCED ONES

The richness of native species in home gar-dens outnumbered the introduced ones. Thediversified management practices in Paraguayans’home gardens show another facet of human-plant relatedness, as well as the interaction be-tween home gardens and the surrounding vege-tation. Wild native plants that invade gardensare not removed by gardeners when they areconsidered useful. Other species are transplantedfrom natural habitats, planted from vegetativepropagules and sown from seeds, especially whengardeners observe their absence in their immedi-ate surroundings. These practices have been ob-served in other studies as well (Blanckaert et al.2004; Coomes 2010; Vogl-Lukasser and Vogl2010). Hence, home gardens become valuable sitesfor researching the domestication of plant popula-tions (Casas et al. 1996).

Of all the registered species in our study, fruittrees from the Myrtaceae family, such as Eugeniauniflora L., Eugenia involucrata DC., and Eugeniapyriformis Cambess. represent canonical cases ofplants that are in the process of domestication. Thisfinding remains in agreement with Clement’s(1999) affirmation about E. uniflora, but here ithas been broadened to others species of Myrtaceae.However, it can be presumed that the domestica-tion of these particular species is enhanced by theirdouble role as edible and medicinal resources (Par-aguayan people commonly utilize leaves as medicineand fruit as snacks). Other incipiently domesticatedplants analyzed by Clement, such as Garciniabrasiliensis Mart. and Acrocomia aculeata (Jacq.)Lodd. ex Mart. were also found in Paraguayans’home gardens. Incipiently domesticated plants arethose at an early stage of domestication with relativelylow phenotypic and genetic differentiation comparedto their wild relatives (Lins Neto et al. 2014).

Home gardens may also be perceived as placeswhere some introduced species may potentially be-come invasive after a process of adaptation/acclimation prior to invasion. Practices in homegardens not only can reduce risk of plant failuredue to protection from unsuitable environmentalconditions and sufficient time for them to adapt tolocal conditions, but also, because they can regulatethe location of seed sources and increase the prop-agule pressure (Hulme 2011; Mack 2000; Perglet al. 2016). The importance of several introducedplants in Misiones is closely related to the history ofthis multi-cultural region (Bartolomé and Schiavoni2008; Kujawska and Pardo de Santayana 2015).Overall, although several plants of foreign originare important medicinal resources in the home gar-dens of Paraguayan migrants, this fact does notcontradict the role of the Paraguayan diaspora inthe promotion and protection of native medicinalresources. Other studies document that native spe-cies richness was sustained with the introduction ofspecies (Akinnifesi et al. 2010). Here, we show thatthe home gardens of Paraguayan migrants displaythe double function of intensive cultivation andincipient management of several native plant re-sources. Moreover, the prevalence of native speciesin home gardens suggest that Paraguayan peoplewho hail from the ecoregion of the Atlantic Forestin Paraguay can continue their phytotherapy in anundisturbed way in Misiones. In these circum-stances, knowledge on native plants is transmittedand exercised in the host country. The assessment ofthese practices and the importance of native species


for the preservation of the natural environment inMisiones, has been, however, beyond the range ofour research.

Conclusions

The presented approach in research on me-dicinal plant richness in home gardens is abreakthrough in the study of home gardens.We outlined a novel methodology, which com-bines approaches and tools from ethnobiologyand landscape ecology. We applied coarse grainand fine grain analysis, which enabled us toestablish a relationship between the characteris-tic of the landscape and the plant diversity inhome gardens. We contributed evidence in sup-port of the environmental scarcity compensationhypothesis. We identified the effect of landscapevariables on the richness of native species, sug-gesting that access to, and availability of, theseresources influence the decisions of Paraguayanimmigrants regarding what is preserved andmanaged in the gardens. Finally, it is importantto point out the possible role of home gardens associal spaces where native plants are preserved,and where the process of plant domestication ismost likely to occur.

AcknowledgementsThe authors are grateful to the study participants

for their generosity in sharing their ethnobotanicalknowledge. The authors thank Gabriela Orofino,Lucía Cariola, Violeta Furlan, Ana Calviño, RamiroAguilar and Carlos De Angelo for useful referencesand comments. The project was funded by theNational Science Centre, Poland, on the basis ofdecision DEC-2013/09/N/HS3/02226.

Open Access This article is distributed under theterms of the Creative Commons Attribution 4.0I n t e r n a t i o n a l L i c e n s e ( h t t p : / /creativecommons.org/licenses/by/4.0/), which per-mits unrestricted use, distribution, and reproduc-tion in any medium, provided you give appropriatecredit to the original author(s) and the source, pro-vide a link to the Creative Commons license, andindicate if changes were made.

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