Ecology and SocietyEcology and Society
 E&S Home > Vol. 21, No. 3 > Art. 19
The following is the established format for referencing this article:
Pauli, N., L. K. Abbott, S. Negrete-Yankelevich, and P. Andrés. 2016. Farmers’ knowledge and use of soil fauna in agriculture: a worldwide review. Ecology and Society 21(3):19.

Farmers’ knowledge and use of soil fauna in agriculture: a worldwide review

1School of Earth and Environment, The University of Western Australia, Crawley, Australia, 2Red de Ecología Funcional, Instituto de Ecología A. C. (INECOL), Veracruz, Mexico, 3CREAF, Cerdanyola del Vallès, Spain


General knowledge of the small, invisible, or hidden organisms that make soil one of the most biodiverse habitats on Earth is thought to be scarce, despite their importance in food systems and agricultural production. We provide the first worldwide review of high-quality research that reports on farmers’ knowledge of soil organisms in agriculture. The depth of farmers’ knowledge varied; some farming communities held detailed local taxonomies and observations of soil biota, or used soil biological activity as indicators of soil fertility, while others were largely unaware of soil fauna. Elicitation of soil biota knowledge was often incidental to the main research goal in many of the reviewed studies. Farmers are rarely deliberately or deeply consulted by researchers on their existing knowledge of soil biota, soil ecology, or soil ecological processes. Deeper understanding of how farmers use and value soil life can lead to more effective development of collaborative extension programs, policies, and management initiatives directed at maintaining healthy, living soils.
Key words: agriculture; ethnoecology; ethnopedology; farmer knowledge; local knowledge; soil biota


The sustainability of our food systems depends on the maintenance of healthy soils. In recognition of the crucial importance of soils, 2015 was proclaimed by the United Nations as the International Year of Soils. The organisms that inhabit soils are responsible for many ecosystem services (Bardgett and van der Putten 2014). The soil system is likely to harbor the greatest concentration of terrestrial biodiversity, although the vast majority of species are undescribed (Decaëns 2010, Jeffery et al. 2010). Worldwide, there are thought to be 900,000 species of mites, 200,000 species of soil-dwelling protozoa, and upward of 1,000,000 species of soil-dwelling fungi, compared with an estimated 300,000 species of vascular plants (Barrios 2007). The soil habitat is complex and opaque, which presents substantial challenges to scientists and farmers interested in understanding soil ecology and biology.

Arguably, until recently soil biology has lived a niche existence (Wall et al. 2010), with little influence on policy and limited appreciation among the wider public of the value and diversity of soil biota (Breure et al. 2012). Soil biodiversity is seldom addressed in national policy (GSBI 2012), and the UN Convention on Biological Diversity (1992) only adopted a cross-cutting theme on soil biodiversity in 2006 (CBD COP 8 Decision VIII/23). The European Union’s withdrawn proposed Soil Framework Directive stated that although soil biodiversity loss was one of the eight major degrading processes affecting European soils, scientific knowledge was “too limited to allow for specific provisions...aiming at its protection” (Commission of the European Communities 2006:10). There are suggestions of growing awareness in policy circles of the importance of soil organisms in attaining broader goals in agriculture, food security, and global change (Wall et al. 2010, Bardgett and van der Putten 2014), but few studies have provided specific recommendations on how to incorporate soil organisms and soil biological processes into management, planning, and policy frameworks.

Although most policy makers may rarely give a thought to soil life, there is one diverse group of people who might well value and hold detailed knowledge about soil organisms: farmers who make their living from the land. We propose that greater understanding of how farmers view soil life can help in the development of extension programs, policies, and management initiatives directed at maintaining healthy soils. The literature reporting on how farmers value and understand soil organisms in an agricultural context has not yet been systematically examined at a worldwide scale. This literature is diffuse and dispersed, belonging to several disciplines that rarely intersect. Two published reviews discuss local knowledge on soil biology for the African region for termites (Sileshi et al. 2009) and pests and pathogens (Sekamatte and Okwakol 2007). Related topics where reviews have proven insightful include: entomophagy (insect-eating; Gahukar 2011); environmental manipulation for insect procurement (Van Itterbeeck and Van Huis 2012); traditional pest management (Morales 2002); entomotherapy (medicinal uses of insects; Costa-Neto 2005); and local ecological knowledge of fungi (de Roman 2010), insects (Posey 1986), and soils (ethnopedology; Barrera-Bassols and Zinck 2003, WinklerPrins and Barrera-Bassols 2004). Much has been written and reviewed on local knowledge of soil physical and chemical properties as well as soil, land, and water management (see Barrera-Bassols and Zinck 2000), but soil biological knowledge is far less widely reported.

We present a worldwide synthesis of peer-reviewed journal articles and high-quality grey literature research on local farmer knowledge of soil biota in agriculture, encompassing a wide range of agricultural systems and cultural contexts. We note that what has been published in the available literature presents only a small fraction of what is actually known by farmers. Our review is limited to visible fauna; although our original intention was to include all soil organisms, we found few papers that addressed farmer knowledge of fungi, rhizobia, or soil microbes in an agricultural context (Romig et al. 1995, Sillitoe 1995, Grossman 2003, Lobry de Bruyn and Abbey 2003, Kelly et al. 2009, Miyagawa et al. 2011), and these papers also discussed visible soil fauna. The aims of the review are the following: (1) to identify patterns in geographic regions, farming systems, and groups of organisms represented in the studies; (2) to ascertain the common themes addressed, and the primary research motivations; (3) to apply a conceptual framework used in ethnoecology to explore how farmers perceive, value, and use soil biota; and (4) to set an agenda to guide future work in extension, management, policy, and research.

Local, hybrid, and scientific ecological knowledge

Scientific research is not the only means by which people develop a meaningful understanding of their surroundings. Local, traditional, indigenous, experiential, and tacit are terms commonly used to describe knowledge systems drawn from sources other than the formal scientific method. Hybrid knowledge can be viewed as the fusion of local knowledge and new expert or technical knowledge gained from external sources, such as agronomists or scientists (Barrios et al. 2006, Reid et al. 2011), although Raymond et al. (2010) caution against overly simplistic categorization of environmental knowledge along a spectrum from local to hybrid to scientific. By combining local experience with global perspectives, the integration of different forms of knowledge can lead to insights into sustainable management, and reduce the risks associated with sustaining livelihoods in marginal environments, or during periods of rapid environmental change (Oberthür et al. 2004, Reed et al. 2007). From a resilience perspective, where local knowledge is seen as influencing the adaptability of social-ecological systems, integration of diverse sources of knowledge is thought to aid management of complexity and uncertainty (Folke et al. 2005), although empirical evidence remains scarce (Bohensky and Maru 2011).

Given the subtleties around what constitutes local ecological knowledge, we have adopted a broad definition that allows our analysis to encompass a wide range of different knowledges, geographic regions, agroecosystems, and socioeconomic contexts. Here, “local knowledge” comprises knowledge gained by indigenous people, farmers, and other resource users based on interactions with their environment, society, and culture over time. Similarly, a broad definition of “farmer” is used to encompass any person practicing cultivation of annual or perennial crops and/or rearing of livestock, for subsistence, exchange, or sale outside the household.

The process by which people incorporate observations on biological interactions and ecological processes into natural resource management and their worldview is known as the corpus-praxis-kosmos complex (or “knowledge-practice-belief”) in ethnoecology (Berkes et al. 2000, Barrera-Bassols and Toledo 2005). The corpus or body includes people’s observations on climate, soils, plants, animals, and vegetation, which may be gained individually or collectively over generations. Praxis encompasses activities that use the body of environmental knowledge to harness resources, and includes agriculture, horticulture, hunting, fishing, beekeeping, agroforestry, livestock, and resource extraction. Kosmos includes culturally important concepts and constructs such as sacred spaces, rituals, myths, and elements of the belief system and moral code. The three domains overlap, and at their centre lies the “Ethnoscape,” which views a landscape as a socio-cultural construct rather than a purely biophysical one (Barrera-Bassols and Toledo 2005). We use this conceptual framework, with its explicit recognition that local environmental knowledge reaches beyond simply identifying or labeling particular features, to review the contributions and research gaps of studies on farmer knowledge of soil fauna in agriculture.


The body of case studies in this review were published in peer-reviewed journals and in high-quality grey literature to December 2015. Systematic keyword searches of online journal databases using combinations and contractions of relevant terms including “knowledge,” “farmer,” “local,” “traditional,” “indigenous,” “ecological,” “agriculture,” “soil,” “fauna,” “biota,” “biology,” and “organisms” were conducted periodically to add new case studies, as well as cross-referencing citations of and within qualifying articles. High quality grey literature (including PhD and MSc theses) was sought via searches of the following: (i) the ProQuest Dissertations and Theses Database; (ii) online publication databases of relevant, renowned sources, Centro Internacional de Agricultura Tropical (CIAT), the Food and Agriculture Organization of the United Nations (FAO), and the World Agroforestry Centre (ICRAF); and (iii) a subject-indexed annotated bibliography of ethnopedological studies (Barrera-Bassols and Zinck 2000). The search was conducted in English, Spanish, Portuguese, and French. Case studies had to include an agricultural context; studies on topics such as geophagy (soil eating; e.g., Rowland 2002), entomophagy, edible wild fungi, and local knowledge of soil biota in isolation from agriculture, e.g., Brazilian studies on myriapods (Costa-Neto 2006) and giant earthworms (Drumond et al. 2015), were not included.

We found 60 studies that met our search criteria (see Table A1.1). Of these, around 47% had a substantial focus on soil biology or invertebrates; in the others, soil biota were often mentioned only briefly. The relatively small number of studies is likely a reflection of the minimal overlap between soil biology and the social sciences. Further, invertebrates are less well-studied than vertebrates in ethnobiology (Meyer-Rochow and Changkija 1997, Ratcliffe 2006) and in the conservation literature (Clark and May 2002), despite being orders of magnitude more abundant. Finally, there may be additional research on this topic in less accessible grey literature, such as local reports and dissertations that are not publicly available. Indeed, some of the grey literature accessed in this review contained extremely detailed insights from farmers on soil biological knowledge (e.g., Dix 1997, Nyeko and Olubayo 2005, Pincus 2015).


Geographic, thematic, and taxonomic coverage

Geographically, research on farmer knowledge of soil fauna in agriculture is most prevalent in East Africa, Central America, and South and Southeast Asia (Fig. 1). One recent study was uncovered in Europe, with three from the USA and two from Australia. The geographic distribution may partially reflect the perceived importance of soil fauna in agriculture. The majority of the systems studied (n = 51) were exclusively smallholder agricultural systems (Table A1.1), where the management of soil biological fertility may be an important base for productivity. Local knowledge, joint investigation, and participatory research may be taken as a more legitimate path of agricultural enquiry by funding bodies in developing country contexts than in high-income countries. The mapped distribution may also reflect the location of institutions such as the Tropical Soil Biology and Fertility Institute (TSBF), with headquarters in Nairobi, Kenya, and field sites for global projects such as the Belowground Biodiversity Project (BGBD), including Brazil, India, Indonesia, Côte d’Ivoire, Uganda, Kenya, and Mexico.

Examining the primary research motivations (Table 1), half the studies were exploratory, conceptual, or methodological contributions, i.e., not applied research. Nearly one quarter of the studies, including four from high-income countries, were focused on developing locally relevant (often farmer-friendly) soil assessment or management tools. Thirteen studies were motivated by a desire to improve agricultural training, extension, or research. Two papers examined the potential for local soil knowledge to be formally recognized in national databases and agricultural development programs.

The majority of publications in the study (n = 35) included attention to multiple soil fauna taxa, while the remainder covered one taxonomic group (such as earthworms or termites; Fig. 2). Earthworms were a focal taxon in around 60% of studies, while termites figured prominently in a third of studies. Publications where a diverse range of soil invertebrates were explored (at least four taxonomic groups) were typically studies of ethnoecology (e.g., Sillitoe 1995, Gurung 2003), integrated pest management (e.g., Morales and Perfecto 2000, Mugerwa et al. 2011), or concerned with farmer views of all soil fauna (e.g., Grossman 2003, Pauli et al. 2012, Kipkorir 2015). Geographically, studies on termites were predominately from Africa, highlighting their importance in this region (Sileshi et al. 2009), studies on beetles and beetle larvae were concentrated in Central and South America, because of the economic importance of soil-dwelling scarab beetle larvae as crop pests (Dix 1997), while other taxa were more evenly distributed across global regions (Fig. 2). Farmers nominated the focal taxa in 62% of studies, researchers defined the taxa in 18% of the papers, and the remaining studies included joint definition (12%), or consisted of observations without consultation (8%).

Exploring themes using the corpus-praxis-kosmos complex

A number of distinct themes emerged from the reviewed research (Table 1). We explore these themes first in terms of the corpus-praxis-kosmos (c-p-k) complex, and second in relation to knowledge integration. The reviewed studies were scored according to whether any of the three domains in the c-p-k complex were examined, following Barrera-Bassols and Toledo’s (2005) categorization. Few of the reviewed publications explicitly used an ethnoecological approach; the presence of each of the c-p-k domains within the studies was scored according to our interpretation of the reported findings. The numbers of studies in each category are presented in a Venn diagram based on Barrera-Bassols and Toledo’s (2005) conceptualization (Fig. 3). Our analysis is based only on reported material; it remains possible that local knowledge of soil fauna encompassed other dimensions that were unexplored or undocumented by the authors of the studies.

The body of knowledge (corpus) held by farmers on soil biota was the most commonly elicited domain, with all but three studies documenting some element of corpus. Examples of knowledge held by farmers include local taxonomies of invertebrates; the level of detail elicited varied widely. The Nepalese Tharu people named 95 varieties of invertebrates from varying habitats (Gurung 2003), the Wola people of Papua New Guinea had 82 different names for invertebrates (Sillitoe 1995), and a study of Honduran folk entomology identified around 140 commonly known invertebrate taxa (Bentley and Rodríguez 2001), whereas other authors suggest that the farmers they worked with had limited traditional knowledge of soil biota (Pincus 2015), or had to be specifically prompted to divulge any information about soil fauna. Although farmers in some regions may have detailed knowledge of invertebrates and soil biota, deep knowledge may extend only to aspects of the soil biota that are easy to observe, or that are culturally or agronomically important (Bentley and Rodríguez 2001).

Another common theme within corpus was the presence of soil fauna as an indicator of soil fertility status (n = 39), or as a component of soil classification (n = 6; Table 1). The most common indicator taxa included earthworms, beetle larvae, termites, and ants. Often, the presence of earthworms, earthworm casts, and/or beetle larvae was widely thought to indicate productive land (e.g., Murage et al. 2000, Barrios and Trejo 2003, Saleque et al. 2008), but in other cases some or all farmers in particular locations believed earthworms to have deleterious or neutral effects on soils and crops (e.g., Sillitoe 1995, Ortiz et al. 1999, Birang et al. 2003, Saïdou et al. 2008). In some regions, farmers use the activity or abundance of particular taxa to classify soil types. For example, earthworms and termites are used by the Béte people of Côte d’Ivoire to aid in identification of subsoils because the soil fauna carry soil particles to the surface (Birmingham 2003). Several authors elicited detailed local observations of species (n = 18; Table 1), such as farmers’ knowledge of life cycles, preferred habitats, and seasonal abundance (e.g., Dix 1997), mound morphology of different termite species (Malaret and Ngoru 1989, Nyeko and Olubayo 2005), and ecological interactions with tree species (Pauli et al. 2012, Kipkorir 2015).

In around one-third of cases reviewed, researchers elicited information on how farmers use information on the abundance or distribution of soil fauna for agricultural decisions (such as where or when to plant particular crops, or how to manipulate soil fauna for agricultural purposes); these were scored as examples of praxis. Eighteen studies documented both corpus and praxis; the three studies that described only praxis were observational studies that showed deliberate consideration of termite mounds in the spatial arrangement of smallholder fields in Zimbabwe (Carter and Murwiwa 1995) and Tanzania (Mielke and Mielke 1982), and the use of termites to rehabilitate degraded soils in Burkina Faso (Roose et al. 1999). Examples of how farmers might use soil fauna information in decision making include assessing whether soil management strategies are working over the short term (Desbiez et al. 2004), and using soil fauna abundance or community composition to determine when soil fertility is sufficient to commence cropping (Black and Okwakol 1997, Dawoe et al. 2012). Soil from termite mounds is used by Lao rice farmers as fertilizer (Miyagawa et al. 2011), a practice also reported from several locations on the African continent (Sileshi et al. 2009).

Deliberate use of soil fauna to improve soils has been documented in Africa and South America. Zaï practice from semiarid West Africa relies on the action of termites to dig galleries in degraded, crusted soil (attracted by organic matter placed in small pits by farmers), allowing water to infiltrate and providing nutrients for plants through the decomposition of organic matter (Roose et al. 1999). Variants of this traditional system have been examined experimentally by soil scientists, highlighting its effectiveness in soil rehabilitation (e.g. Mando et al. 1996, 1999). The Kayapó people of the Brazilian Amazon basin add soil from termite mounds and ant nests, along with live termites and ants, to mounds of mulch placed in shallow depressions. These mounds are tended and slowly evolve to become forest “islands” (apêtê) in the surrounding savanna over the course of decades, which are highly valued as refuges, sources of food, firewood, poisons, medicines, and materials for daily life (Posey 1985).

Several studies gave insight into strategies employed by farmers to reduce the severity of pest attacks by soil invertebrates (examples of praxis). Farmers in Honduras used crop rotation, ash application, and reliance on natural predators to deter white grub infestation (Pauli et al. 2012); similar practices were documented by Morales and Perfecto (2000) and Wyckhuys and O’Neil (2007) in central America. A variety of techniques to discourage termites from attacking tree crops were elicited from farmers in Malawi, Mozambique, and Zambia, including planting cuttings of a plant believed to attract termites in termite-infested areas, digging up the mound, and destroying the queen, applying wood ash in planting holes, and applying meat to attract predatory ants (Sileshi et al. 2008), while farmers in the rangelands of Uganda had a detailed understanding of the links between overgrazing, ecosystem deterioration, and heightened termite damage of pasture vegetation (Mugerwa et al. 2011).

Although seldom described in the reviewed studies, soil invertebrates can figure in local people’s belief and spiritual systems (kosmos). For example, termites feature prominently in iconography in San rock art in Southern Africa (Mguni 2006), the “Honey Ant Dreaming” mural painted at Papunya in 1971 was the catalyst that started the famous Western Desert Australian Aboriginal Art movement (Carmichael and Kohen 2013), and scarab beetles were widespread in religion and cosmology in ancient Egypt (Ratcliffe 2006). The cosmological significance of invertebrates was touched on in seven reviewed studies. The Cakchiquel Maya of Guatemala believe they should share their corn with the animals, and for this reason they do not believe in killing invertebrate “pests” that may attack their sacred crop (Morales and Perfecto 2000). Elsewhere in Mesoamerica, the soil is conceptualized as a living being (Barrera-Bassols 2003, Barrera-Bassols et al. 2006), and earthworms figure in beliefs and myths as a “symbolic bridge of fertility and health between man and nature” (Ortiz et al. 1999:246). The only reviewed study of nonindigenous people to consider elements of kosmos indicated that Michigan farmers’ worldview influenced their management strategies (organic or nonorganic) and the regard attached to “living soil” (Atwood 2010). Some cultures hold negative views of invertebrates stemming from overt or covert beliefs. The Tharu of Nepal believe “small living things,” including insects, are a mistake in God’s creation, while the Wola of Papua New Guinea attribute painful sores to earthworm bites (Sillitoe 1995). People’s belief systems may have a clear link to perceptions of and values attached to soil fauna, which could have an impact on the uptake (or otherwise) of management strategies designed to foster improved soil health through greater biological activity.

Hybrid knowledge: comparing, validating, and integrating

Nearly half of the reviewed studies included some element of comparing or integrating different types of knowledge held on soil biota or soils (Table 1). Early papers in the field of ethnopedology tended to view knowledge gained by the scientific method as correct, with an emphasis on validating whether local knowledge reflected or correlated with scientific understanding, and could therefore be proven (Barrera-Bassols and Zinck 2003). More recent work acknowledges that an integrated approach, where multiple forms of enquiry are pursued, collaboration with local people is actively sought, and no particular type of knowledge is privileged as superior, is required to better understand the role that local cultural, social, and economic processes play in agricultural management (Barrera-Bassols and Zinck 2003). Most of the studies sampled for this review reflect the latter trend, which is perhaps due to the relative novelty of research on local knowledge of soil biota. Rationale for comparing or integrating knowledge included collaborative development of local indicators of soil fertility (Rousseau et al. 2013; see also Barrios et al. 2012); creation of locally appropriate soil maps through integrated knowledge (e.g., Saleque et al. 2008, Tesfahunegn et al. 2011); joint investigation of the life cycle and distribution of poorly understood soil-dwelling crop pests (Dix 1997); and assessing the similarities and differences among local and scientific understanding of soil biota in pest management and soil fertility (e.g., Price 2001, Ericksen and Ardon 2003, Saïdou et al. 2008).

Several papers examined farmers’ understanding of the role of soil organisms in soil processes within the context of developing agricultural extension. In the Ashanti region of western Ghana, a large majority of interviewed farmers understood that soil fauna assist in the physical breakdown of organic matter and through this contribute to soil fertility, but a much smaller proportion appreciate their role in gas and water exchange (Dawoe et al. 2012). Arguably, comminution of organic matter is visible, whereas physical activity in the soil profile is not, reflecting Bentley and Rodríguez’ (2001) assertion that deeper knowledge extends to soil-dwelling species that are easily observed. Similarly, Grossman (2003) found that although organic coffee farmers in Chiapas (Mexico) had a thorough understanding of organic matter decomposition, some important knowledge gaps existed in processes that farmers could not see, including nitrogen fixation, soil microbial activity, and mineralization. These studies highlight the importance of developing collaborative approaches to agricultural extension, where knowledge from a range of different sources and social-ecological contexts is seen as valuable for the development of sustainable agriculture.

A topic on which little research has been published is the use of soil biota as a potentially rich talking point around which to build knowledge interchange between farmers and researchers. Visible soil biota can give farmers immediate feedback on how their land management practices are working, while the use of narratives and guided use of appropriate technology can make the invisible visible, and facilitate the process of integrating knowledge. Recent research highlights this trend. For example, the L’Observatoire Agricole de la Biodiversité in France provides interested farmers with training on how to quantify elements of agricultural biodiversity (including litter and soil invertebrates) that relate to farm management (Deschamps and Demeulenaere 2015). In Uganda, farmers interviewed by Pincus (2015) were initially largely unaware of the role earthworms play in agriculture, but after attending training and participating in soil testing, over 80% of farmers viewed earthworm presence as an indicator of soil fertility. In the following paragraphs, we report several as-yet unpublished examples encompassing a diverse range of agroecosystems and cultural contexts to illustrate how this can work.

In Mexico, researchers have developed illustrated narrative booklets to discuss the consequences of different management strategies for vital plant symbionts including mycorrhizal fungi in roots and nitrogen fixing bacteria (Fig. 4). Land degradation is a serious problem in mountainous areas in Mexico that has resulted in decreased maize productivity and food insecurity. At the center of this problem is the loss of traditional crop diversity (intra- and interspecific) after the ill-informed adoption of technological packages including maize hybrids and chemical fertilization. Research suggests that the loss of locally developed crops and pulses of nutrients have diminished the diversity of well-adapted mycorrhizal fungi and nitrogen-fixing bacteria symbionts that developed with millennia of crop domestication by local Popoluca people (López-López et al. 2013, Sangabriel-Conde et al. 2014). The BioPop project (lead by author S. N.-Y.) developed a strategy to open discussion with farmers about this problem. A pair of illustrated publications including hybrid knowledge was handed to producers: a short story (“Don Erasmo’s milpa”; Fig. 4A) and a triptych (“What is happening to the milpa?”; Fig. 4B). The short story is a first-person narrative of what happened to a farmer’s soil, traditional knowledge, and food availability since technological packages arrived. The triptych is a symptom (“have you noticed that...?”)-awareness (“what has happened is...”), that attempts to draw the links between traditional crop diversity, microbial symbiont conservation, nutrient use efficiency, and food security.

The Western Australian Wheatbelt region is an ancient, weathered landscape within a global biodiversity hotspot (Myers et al. 2000; Fig. 5). Broadacre farming of grain and livestock is the major land use. Author L. K. A. has been conducting workshops with farmers throughout the region for many years, most recently with the On-Farm Soil Health Monitoring project (Wheatbelt NRM et al. 2013). The goal of these workshops is to introduce farmers and landowners to the diversity of organisms in their soils, through on-farm soil monitoring methods such as extraction of soil mesofauna, and staining root samples to detect the presence of mycorrhizal fungi (Mahdi et al. 2016). Farmers are empowered to do their own experiments and analyses to support adaptive management. In the nutrient-poor soils of the Wheatbelt, mycorrhizal fungi can be important for crop growth. During workshops, farmer-friendly techniques for determining the presence or absence of mycorrhizal fungi in roots are demonstrated. Although these methods may not compare with the precision afforded by research laboratory images (Fig. 5), they are sufficient to help answer farmers’ questions.

In the tropical dry forests of Nicaragua, soil arthropods were identified as an important local indicator of soil quality as part of collaborative, participatory research on land degradation. Between 2005 and 2010, author P. A. led an integrated planning process for pasturelands within the Nature Reserve Mesas de Moropotente in Nicaragua (Fig. 6). Land degradation due to overgrazing had caused economic losses in an area already affected by poverty. Stakeholders were brought together for a social multicriteria evaluation, with the intention of generating a constructive dialogue between local and scientific knowledge of the situation. Early on, improving soil quality emerged as a priority. There were substantial differences in the way that researchers and producers sought to describe and understand soil quality. Producers tended to aggregate different soil characteristics together into one complex soil quality indicator, while researchers focused on a series of independent, measurable soil parameters. Soil arthropods were identified by producers but were not initially associated specifically with soil. The presence of fauna at the soil surface was associated with healthy pastures, crops, and forests, and conveyed a sense of an ideal, pristine ecosystem, when there were no weeds and pastures were richly colored. As part of the dialogue, soil arthropods were eventually identified as indicators of soil fertility, through the effects of their faeces on soil aggregation. Producers granted access to their lands for soil sampling to quantify the diversity of epigeic fauna during the wet and dry season. The willingness of producers to support scientific sampling to evaluate indicators developed from discussion forums indicates the strength of local support for the process and research project.


There is a potentially rich body of local knowledge on soil life, but one that is seldom tapped and often eclipsed by a focus on (a) other elements of the biota, or (b) soil physical and chemical properties. The lack of attention to this topic is particularly noticeable for high-income countries. Although farming systems in these countries may depart widely from the largely low-input, subsistence systems covered in this review, there is growing interest in biological farming and in more holistic views of soil health; recent work in Austria shows that farmers see soil as a key part of their identity, and many value “soil life” (Wahlhütter et al. 2016). Researchers investigating local soil knowledge and management should give consideration to the biological component of soil. In particular, researchers should direct attention not just to observations of soil biota (corpus), but also to how these organisms are considered in agricultural activities (praxis) and to the belief systems that influence agricultural practices and perceptions of soil life (kosmos). Further, there are few published data on local knowledge of the agricultural role of symbiotic microorganisms such as rhizobia and mycorrhizal fungi, or of other “invisible” organisms that have direct influence on agricultural productivity and soil fertility. We encourage collaborative partnerships among social scientists, soil scientists, farmers, and extension workers to jointly investigate these issues.

Our review raises questions about the local knowledge that has been lost, or is in danger of being lost. With the advent of synthetic inputs, technological solutions to increase yield, and greater productivity, coupled with out-migration from rural areas to urban zones, long-standing knowledge of the biological component of soil fertility could be eroded. The indigenous Guatemalan farmers interviewed by Morales and Perfecto (2000) feared their children would not continue with traditional practices and their knowledge would be lost. At the other end of the spectrum, many of the studies conducted in high-income countries highlighted the value of empowering farmers by developing locally relevant soil health assessments, reducing reliance on costly outside expertise. Although we did not explore gender as an influence on soil biological knowledge in this review, several authors noted gender-related differences (Saïdou et al. 2008, Sileshi et al. 2008, Zúñiga et al. 2013). The trend toward increasing feminization of agriculture in some global regions (Deere 2005) may also influence the knowledge that is retained, transmitted, and used in agriculture. Future work should consider how local soil knowledge may change over time in relation to socio-cultural and demographic drivers, as well as changes in land use and agricultural production systems.

In the last decades, science has made great strides in understanding the diversity and importance of soil life. However, general public awareness is said to be low (Wall et al. 2010), and interest from decision makers and government agencies is similarly subdued (Kust 2013). Our review shows that there are groups within the community who do value and understand soil life. However, aside from these few notable and fascinating exceptions, farmers are rarely deliberately or deeply consulted on their knowledge of soil organisms or soil biological processes, and research is rarely published in the peer-reviewed literature on the understanding or uptake of practices designed to enhance soil biological activity. A clear theme in many of the reviewed studies was that understanding and respecting how farmers view soil and soil life can help improve agricultural extension programs, soil management initiatives, and training in integrated pest management. Indeed, extension programs and farmer-led activities that incorporate soil biota exist (such as “microscope clubs” among grower groups in Australia), but they are rarely documented in the peer-reviewed literature. Collaborating with farmers, documenting their knowledge through participatory research, and presenting their views as equally important as those of soil scientists may also help to bridge the science-policy divide on this topic and add legitimacy to efforts to include soil organisms within broader legal and policy frameworks on soils.

Because of the sparse literature and the diverse, often site-specific investigative techniques used, much remains unknown about the depth of farmers’ knowledge of soil biology. The integration of locally relevant knowledge with globally relevant scientific principles may help reduce risks associated with farming in marginal environments, or aid in adaptation to rapid environmental change (Oberthür et al. 2004). To aid adaptation to environmental and socioeconomic change, we urge researchers in this field to seek a clearer understanding of how famers value and perceive soil biota in agricultural production and sustainable land management. Properly applied, this knowledge will help deliver improved extension programs and management toolkits that are locally appropriate and tailored to farmers’ needs.


Responses to this article are invited. If accepted for publication, your response will be hyperlinked to the article. To submit a response, follow this link. To read responses already accepted, follow this link.


The BioPop project was financed by FOMIX (94427, CONACYT-Veracruz); “Don Erasmo’s milpa” and “What is happening to the milpa?” were illustrated by Rafael Ruiz Moreno and coauthored by Jimena Mejía Alemán and Simoneta Negrete-Yankelevich. Funding for the Monitoring Soil Science On-Farm project was provided by the Australian Federal Government through Caring for Our Country to Wheatbelt Natural Resource Management Inc, Western Australia; partner organizations were The University of Western Australia (UWA), South West Catchments Council and the SPICE program at the Centre for Learning Technology (UWA). Bede Mickan provided the laboratory image of a root with mycorrhizal fungi. The project “Social multicriteria evaluation for the sustainable management and conservation of the Miraflor-Moropotente Protected Terrestrial Landscape” was funded by the Catalan Agency for Development and Cooperation (ACCD) and the Autònoma Solidària Foundation (FAS) at the Autònomous Univeristy of Barcelona (Spain). Author NP wishes to acknowledge the valuable suggestions from participants in the first Global Soil Biodiversity Initiative (Dijon, France, 2-5 December 2014) and UWA colleagues M Tonts, S Prout, J Clifton, and F Haslam McKenzie for advice on previous versions of this paper. We are grateful for the constructive and insightful comments received on this manuscript from two anonymous reviewers.


Atwood, L. W. 2010. Interpreting the farm as a system: differences in worldviews among large-scale non-organic and organic farmers in Michigan’s Thumb region. Thesis, Michigan State University, Lansing, Michigan, USA.

Bardgett, R. D., and W. H. van der Putten. 2014. Belowground biodiversity and ecosystem functioning. Nature 515:505-511.

Barrera-Bassols, N. 2003. Symbolism, knowledge and management of soil and land resources in indigenous communities: ethnopedology at global, regional and local scales. Dissertation, Ghent University, Ghent, Belgium.

Barrera-Bassols, N., and V. M. Toledo. 2005. Ethnoecology of the Yucatec Maya: symbolism, knowledge and management of natural resources. Journal of Latin American Geography 4:9-41.

Barrera-Bassols, N., and J. A. Zinck. 2000. Ethnopedology in a worldwide perspective: an annotated bibliography. International Institute for Aerial Survey and Earth Sciences (ITC), Enschede, The Netherlands.

Barrera-Bassols, N., and J. A. Zinck. 2003. Ethnopedology: a worldwide view on the soil knowledge of local people. Geoderma 111:171-195.

Barrera-Bassols, N., J. A. Zinck, and E. Van Ranst. 2006. Symbolism, knowledge and management of soil and land resources in indigenous communities: ethnopedology at global, regional and local scales. Catena 65:118-137.

Barrios, E. 2007. Soil biota, ecosystem services and land productivity. Ecological Economics 64:269-285.

Barrios, E., H. L. C Coutinho, and C. A. B. Medeiros. 2012. InPaC-S: Participatory knowledge integration on indicators of soil quality - methodological guide. ICRAF, Embrapa, CIAT. World Agroforestry Centre, Nairobi, Kenya.

Barrios, E., R. J. Delve, M. Bekunda, J. Mowo, J. Agunda, J. Ramisch, M. T. Trejo, and R. J. Thomas. 2006. Indicators of soil quality: a south-south development of a methodological guide for linking local and technical knowledge. Geoderma 135:248-259.

Barrios, E., and M. T. Trejo. 2003. Implications of local soil knowledge for integrated soil management in Latin America. Geoderma 111:217-231.

Bentley, J. W., and G. Rodríguez. 2001. Honduran folk entomology. Current Anthropology 42:285-301.

Berkes, F., J. Colding, and C. Folke. 2000. Rediscovery of traditional ecological knowledge as adaptive management. Ecological Applications 10:1251-1262.[1251:ROTEKA]2.0.CO;2

Birang, M., S. Hauser, and D. L. Amougou. 2003. Farmers’ perception of the effects of earthworms on soil fertility and crop performance in southern Cameroon. Pedobiologia 47:819-824.

Birmingham, D. M. 2003. Local knowledge of soils: the case of contrast in Côte d’Ivoire. Geoderma 111:481-502.

Black, H. I. J., and M. J. N. Okwakol. 1997. Agricultural intensification, soil biodiversity and agroecosystem function in the tropics: the role of termites. Applied Soil Ecology 6:37-53.

Bohensky, E. L., and Y. Maru. 2011. Indigenous knowledge, science and resilience: What have we learned from a decade of international literature on “integration”? Ecology and Society 16(4):6.

Breure, A. M., G. B. De Deyn, E. Dominati, T. Eglin, K. Hedlund, J. Van Orshoven, and L. Posthuma. 2012. Ecosystem services: a useful concept for soil policy making! Current Opinion in Environmental Sustainability 4:578-585.

Carmichael, B., and A. Kohen. 2013. The forgotten Yuendumu Men’s Museum murals: shedding new light on the progenitors of the Western Desert Art Movement. Australian Aboriginal Studies 1:110-116.

Carter, S. E., and H. K. Murwiwa. 1995. Spatial variability in soil fertility management and crop response in Mutoko Communal Area, Zimbabwe. Ambio 24:77-84.

Clark, J. A., and R. M. May. 2002. Taxonomic bias in conservation research. Science 297:191-192.

Commission of the European Communities. 2006. Proposal for a Directive of the European Parliament and of the Council establishing a framework for the protection of soil and amending Directive 2004/35/EC. Commission of the European Communities, Brussels, Belgium.

Costa-Neto, E. M. 2005. Entomotherapy, or the medicinal use of insects. Journal of Ethnobiology 25:93-114.[93:EOTMUO]2.0.CO;2

Costa-Neto, E. M. 2006. “Piolha-de-cobra” (Arthropoda: Chilopoda: Geophilomorpha) na concepção dos moradores de Pedra Branca, Santa Terezinha, Estado de Bahia, Brasil. Acta Scientiarum Biological Sciences Maringá 28:143-148.

Dawoe, E. K., J. Quashie-Sam, M. E. Isaac, and S. K. Oppong. 2012. Exploring farmers’ local knowledge and perceptions of soil fertility and management in the Ashanti Region of Ghana. Geoderma 179-180:96-103.

Decaëns, T. 2010. Macroecological patterns in soil communities. Global Ecology and Biogeography 19:287-302.

Deere, C. D. 2005. The feminization of agriculture? Economic restructuring in rural Latin America. United Nations Research Institute for Social Development, Geneva, Switzerland.

de Roman, M. 2010. The contribution of wild fungi to diet, income and health: a world review. Pages 327-347 in M. Rai and G. Kövics, editors. Progress in mycology. Springer, Dordrecht, The Netherlands.

Desbiez, A., R. Matthews, B. Tripathi, and J. Ellis-Jones. 2004. Perceptions and assessment of soil fertility by farmers in the mid-hills of Nepal. Agriculture, Ecosystems and Environment 103:191-206.

Deschamps, S., and E. Demeulenaere. 2015. L’Observatoire Agricole de la Biodiversité. Vers un ré-ancrage des pratiques dans leur milieu. Etudes Rurales 195:109-126.

Dix, A. M. 1997. The biology and ecology of broccoli white grums (Coleoptera: Scarabaeidae) in the community of Chilasco, Baja Verapaz, Guatemala: an integrated approach to pest management. Dissertation, University of Georgia, Athens, Georgia, USA.

Drumond, M. A., A. Q. Guimarães, and R. H. P. da Silva. 2015. The role of local knowledge and traditional extraction practices in the management of giant earthworms in Brazil. PLoS ONE 10:e0123913.

Ericksen, P. J., and M. Ardon. 2003. Similarities and differences between farmer and scientist views on soil quality issues in central Honduras. Geoderma 111:233-248.

Folke, C., T. Hahn, P. Olsson, and J. Norberg. 2005. Adaptive governance of social-ecological systems. Annual Review of Environment and Resources 30:441-473.

Gahukar, R. T. 2011. Entomophagy and human food security. International Journal of Tropical Insect Science 31:129-144.

Global Soil Biodiversity Initiative (GSBI). 2012. White Paper on the first open meeting of the Global Soil Biodiversity Initiative (GSBI). GSBI, London, UK. [online] URL:

Grossman, J. 2003. Exploring farmer knowledge of soil processes in organic coffee systems of Chiapas Mexico. Geoderma 111:267-287.

Gurung, A. B. 2003. Insects - a mistake in God’s creation? Tharu farmers’ perception and knowledge of insects: a case study of Gobardiha Village Development Committee: Dang-Deukhuri, Nepal. Agriculture and Human Values 20:337-370.

Jeffery, S., C. Gardi, A. Jones, L. Montanarella, L. Marmo, L. Miko, K. Ritz, G. Peres, J. Römbke, and W. van der Putten, Editors. 2010. European atlas of soil biodiversity. European Commission, Publications Office of the European Union, Luxembourg.

Kelly, B., C. Allan and B. P. Wilson. 2009. Soil indicators and their use by farmers in the Billabong Catchment, southern New South Wales. Soil Research 47:234-242.

Kipkorir, L. D. 2015. Influence of indigenous trees on soil macrofauna and soil organic matter dynamics in tropical Miombo woodlands. Dissertation, University of Nairobi, Nairobi, Kenya.

Kust, G. 2013. Terminal evaluation of the UNEP/GEF Project “Conservation and Sustainable Management of Below Ground Biodiversity.” United Nations Environment Programme, Nairobi, Kenya.

Lobry de Bruyn, L. A., and J. A. Abbey. 2003. Characterisation of farmers’ soil sense and the implications for on-farm monitoring of soil health. Australian Journal of Experimental Agriculture 43:285-305.

López-López, A., S. Negrete-Yankelevich, M. A. Rogel, E. Ormeño-Orrillo, J. Martínez, and E. Martínez-Romero. 2013. Native bradyrhizobia from Los Tuxtlas in Mexico are symbionts of Phaseolus lunatus (Lima bean). Systematic and Applied Microbiology 36:33-38

Mahdi, J. E., L. K. Abbott, N. Pauli, and Z. M. Solaiman. 2016. Biological indicators for soil health: potential for development and use of on-farm tests. In A. Varma and A. K. Sharma, editors. Modern tools and techniques to understand microbes. Springer, Soil Biology Series in press.

Malaret, L., and F. N. Ngoru. 1989. Ethno-ecology: a tool for community based pest management farmer knowledge of termites in Machakos district, Kenya. Sociobiology 15:197-211.

Mando, A., L. Brussaard, and L. Stroosnijder. 1999. Termite- and mulch-mediated rehabilitation of vegetation on crusted soil in West Africa. Restoration Ecology 7:33-41.

Mando, A., L. Stroosnijder, and L. Brussaard. 1996. Effects of termites on infiltration into crusted soil. Geoderma 74:107-113.

Meyer-Rochow, V. B., and S. Changkija. 1997. Uses of insects as human food in Papua New Guinea, Australia, and North-East India: cross-cultural considerations and cautious conclusions. Ecology of Food and Nutrition 36:159-185.

Mguni, S. 2006. Iconography of termites’ nests and termites: symbolic nuances of formlings in southern African San rock art. Cambridge Archaeological Journal 16:53-71.

Mielke, H. W., and P. W. Mielke. 1982. Termite mounds and chitemene agriculture: a statistical analysis of their association in southwestern Tanzania. Journal of Biogeography 9:499-504.

Miyagawa, S., Y. Koyama, M. Kokubo, Y. Matsushita, Y. Adachi, S. Sivilay, N. Kawakubo, and S. Oba. 2011. Indigenous utilization of termite mounds and their sustainability in a rice growing village of the central plain of Laos. Journal of Ethnobiology and Ethnomedicine 7:24.

Morales, H. 2002. Pest management in traditional tropical agroecosystems: lessons for pest prevention research and extension. Integrated Pest Management Reviews 7:145-163.

Morales, H., and I. Perfecto. 2000. Traditional knowledge and pest management in the Guatemalan highlands. Agriculture and Human Values 17:49-63.

Mugerwa, S., M. Nyangito, J. Nderitu, C. Bakuneta, D. Mpairwe, and E. Zziwa. 2011. Farmers’ ethno-ecological knowledge of the termite problem in semi-arid Nakasongola. African Journal of Agricultural Research 6:3183-3191.

Murage, E. W., N. K. Karanja, P. C. Smithson, and P. L. Woomer. 2000. Diagnostic indicators of soil quality in productive and non-productive smallholders’ fields of Kenya’s Central Highlands. Agriculture, Ecosystems and Environment 79:1-8

Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. da Fonseca, and J. Kent. 2000. Biodiversity hotspots for conservation priorities. Nature 403:853-858.

Nyeko, P., and F. M. Olubayo. 2005. Participatory assessment of farmers’ experiences of termite problems in agroforestry in Tororo District, Uganda. Agricultural Research and Extension Network, London, UK.

Oberthür, T., E. Barrios, S. Cook, H. Usma, and G. Escobar. 2004. Increasing the relevance of scientific information in hillside environments through understanding of local soil management in a small watershed of the Colombian Andes. Soil Use and Management 20:23-31.

Ortiz, B., C. Fragoso, I. M’Boukou, B. Pashanasi, B. K. Senapati, and A. Contreras. 1999. Perception and use of earthworms in tropical farming systems. Pages 239-249 in P. Lavelle, L. Brussaard, and P. Hendrix, editors. Earthworm management in tropical agroecosystems. CABI, Oxford, UK.

Pauli, N., E. Barrios, A. J. Conacher, and T. Oberthür. 2012. Farmer knowledge of the relationships among soil macrofauna, soil quality and tree species in a smallholder agroforestry system of western Honduras. Geoderma 189-190:186-198.

Pincus, L. M. 2015. Increasing indigenous vegetable yield and nutritional quality through traditionally- and scientifically-informed soil fertility management. Horticulture and Agronomy. Dissertation, University of California Davis, Davis, California, USA.

Posey, D. A. 1985. Indigenous management of tropical forest ecosystems: the case of the Kayapó Indians of the Brazilian Amazon. Agroforestry Systems 3:139-158.

Posey, D. A. 1986. Topics and issues in ethnoentomology with some suggestions for the development of hypothesis-generation and testing in ethnobiology. Journal of Ethnobiology and Ethnomedicine 6:99-120.

Price, L. L. 2001. Demystifying farmers’ entomological and pest management knowledge: a methodology for assessing the impacts on knowledge from IPM-FFS and NES interventions. Agriculture and Human Values 18:153-176.

Ratcliffe, B. C. 2006. Scarab beetles in human culture. Coleopterists Society Monographs 60(sp5):85-101.[85:sbihc];2

Raymond, C. M., I. Fazey, M. S. Reed, L. C. Stringer, G. M. Robinson, and A. C. Evely. 2010. Integrating local and scientific knowledge for environmental management. Journal of Environmental Management 91:1766-1777.

Reed, M. S., A. J. Dougill, and M. J. Taylor. 2007. Integrating local and scientific knowledge for adaptation to land degradation: Kalahari Rangeland management options. Land Degradation and Development 18:249-268.

Reid, K. A., K. J. H. Williams, and M. S. Paine. 2011. Hybrid knowledge: place, practice and knowing in a volunteer ecological restoration project. Ecology and Society 16(3):19.

Romig, D. E., M. J. Garlynd, R. F. Harris, and K. McSweeney. 1995. How farmers assess soil health and quality. Journal of Soil and Water Conservation 50:229-236.

Roose, E., V. Kabore, and C. Guenat. 1999. Zaï practice: a West African traditional rehabilitation system for semiarid degraded lands, a case study in Burkina Faso. Arid Soil Research & Rehabilitation 13:343-355.

Rousseau, L., S. J. Fonte, O. Téllez, R. van der Hoek, and P. Lavelle. 2013. Soil macrofauna as indicators of soil quality and land use impacts in smallholder agroecosystems of western Nicaragua. Ecological Indicators 27:71-82.

Rowland, M. J. 2002. Geophagy: an assessment of implications for the development of Australian indigenous plant processing technologies. Australian Aboriginal Studies 2002:51-66.

Saïdou, A., D. Kossou, L. Brussaard, P. Richards, and T. W. Kuyper. 2008. Earthworm activities in cassava and egusi melon fields in the transitional zone of Benin: linking farmers’ perceptions with field studies. Wageningen Journal of Life Sciences 56:123-135.

Saleque, M. A., M. K. Uddin, A. K. M. Ferdous, and M. H. Rashid. 2008. Use of farmers’ empirical knowledge to delineate soil fertility-management zones and improved nutrient- management for lowland rice. Communications in Soil Science and Plant Analysis 39:25-45.

Sangabriel-Conde, W., S. Negrete-Yankelevich, I. E. Maldonado-Mendoza, and D. Trejo-Aguilar. 2014. Native maize landraces from Los Tuxtlas, Mexico show varying mycorrhizal dependency for P uptake. Biology and Fertility of Soils 50:405-414.

Sekamatte, M. B., and M. J. N. Okwakol. 2007. The present knowledge on soil pests and pathogens in Uganda. African Journal of Ecology 45:9-19.

Sileshi, G. W., E. Kuntashula, P. Matakala, and P. O. Nkunika. 2008. Farmers’ perceptions of tree mortality, pests and pest management practices in agroforestry in Malawi, Mozambique and Zambia. Agroforestry Systems 72:87-101.

Sileshi, G. W., P. Nyeko, P. O. Y. Nkunika, B. M. Sekematte, F. K. Akinnifesi, and O. C. Ayaji. 2009. Integrating ethno-ecological and scientific knowledge of termites for sustainable termite management and human welfare in Africa. Ecology and Society 14(1):48.

Sillitoe, P. 1995. Ethnoscientific observations on entomology and mycology in the southern highlands of Papua New Guinea. Science in New Guinea 21:3-26.

Tesfahunegn, G. B., L. Tamene, and P. L. G. Vlek. 2011. A participatory soil quality assessment in Northern Ethiopia’s Mai-Negus catchment. Catena 86:1-13.

Van Itterbeeck, J., and A. Van Huis. 2012. Environmental manipulation for edible insect procurement: a historical perspective. Journal of Ethnobiology and Ethnomedicine 8:3.

Wahlhütter, S., C. R. Vogl, and H. Eberhart. 2016. Soil as a key criteria in the construction of farmers’ identities: the example of farming in the Austrian province of Burgenland. Geoderma 269:39-53.

Wall, D. H., R. D. Bardgett, and E. Kelly. 2010. Biodiversity in the dark. Nature Geoscience 3:297-298.

Wheatbelt Natural Resource Management (NRM), The University of Western Australia, Southwest Catchments Council, and SPICE Centre for Learning Technology. 2013. On-farm soil monitoring handbook. Wheatbelt Natural Resource Management, Northam, Australia.

WinklerPrins, A. M. G. A., and N. Barrera-Bassols. 2004. Latin American ethnopedology: a vision of its past, present, and future. Agriculture and Human Values 21:139-156

Wyckhuys, K. A. G., and R. J. O’Neil. 2007. Local agro-ecological knowlede and its relationship to farmers’ pest management decision making in rural Honduras. Agriculture and Human Values 24:307-321.

Zúñiga, M. C., A. Feijoo, H. Quintero, N. J. Aldana, and A. F. Carvajal. 2013. Farmers’ perceptions of earthworms and their role in soil. Applied Soil Ecology 69:61-68.

Address of Correspondent:
Natasha Pauli
School of Earth and Environment
The University of Western Australia
35 Stirling Highway
Crawley 6009
Western Australia
Jump to top
Table1  | Figure1  | Figure2  | Figure3  | Figure4  | Figure5  | Figure6  | Appendix1