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  <front>
    <journal-meta>
      <journal-title-group>
        <journal-title>BioDiv</journal-title>
      </journal-title-group>
    </journal-meta>
    <article-meta>
      <title-group>
        <article-title>STWO: An Ontology for Soil Food Web Reconstruction</article-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author">
          <string-name>Nicolas Le Guillarme</string-name>
          <xref ref-type="aff" rid="aff1">1</xref>
        </contrib>
        <contrib contrib-type="author">
          <string-name>Mickaël Hedde</string-name>
          <xref ref-type="aff" rid="aff0">0</xref>
        </contrib>
        <contrib contrib-type="author">
          <string-name>Wilfried Thuiller</string-name>
          <xref ref-type="aff" rid="aff1">1</xref>
        </contrib>
        <aff id="aff0">
          <label>0</label>
          <institution>INRAE, UMR Eco &amp; Sols</institution>
          ,
          <addr-line>Montpellier</addr-line>
          ,
          <country country="FR">France</country>
        </aff>
        <aff id="aff1">
          <label>1</label>
          <institution>Univ. Grenoble Alpes, Univ. Savoie Mont Blanc</institution>
          ,
          <addr-line>CNRS, LECA</addr-line>
          ,
          <institution>Laboratoire d'Ecologie Alpine</institution>
          ,
          <addr-line>F-38000 Grenoble</addr-line>
          ,
          <country country="FR">France</country>
        </aff>
      </contrib-group>
      <pub-date>
        <year>2021</year>
      </pub-date>
      <volume>3</volume>
      <fpage>11</fpage>
      <lpage>18</lpage>
      <abstract>
        <p>While food webs are pivotal tools to understand the structure, dynamics and functioning of ecosystems, their reconstruction is not trivial since feeding relationships are not always known. To this end, soil ecologists often simplify the problem by either grouping morphologically similar organisms into trophic groups with known interactions or by assuming that feeding relationships are predictable from consumer diets (e.g. frugivore or bacterivore). Interestingly, the scientific community has collected a considerable amount of information on trophic interactions and feeding habits. However, the largescale exploitation of these data for food web reconstruction is hampered by the lack of standards for representing and reasoning upon trophic knowledge. The goal of our work is to propose an ontology that will support the automatic reconstruction of soil food webs from community composition data.</p>
      </abstract>
      <kwd-group>
        <kwd>eol&gt;soil ecology</kwd>
        <kwd>food web</kwd>
        <kwd>ontology development</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec id="sec-1">
      <title>1. Introduction</title>
      <p>
        Food webs (also called trophic webs, trophic interaction networks) encode both the composition
of ecological communities as well as the feeding relationships within the community. In soil
ecology, food webs are often used to understand the structure and dynamics of soil assemblages,
and their impact on decomposition processes and nutrient cycling [
        <xref ref-type="bibr" rid="ref1">1</xref>
        ]. Yet, reconstructing soil
food webs is not a straightforward task. The nature of soil as a black-box ecosystem makes
direct observations of most trophic interactions fairly impossible, and knowledge of resource
preferences of many taxonomic groups of soil fauna are not well known. These preferences
may be inferred from morphological similarities with species of known feeding habits or by
phylogenetic proximity. This results in a more or less fine categorization of soil flora, fauna,
fungi and microbes into a multitude of trophic groups (e.g. bacterivorous nematodes, arbuscular
mycorrhizal fungi or saprotrophic fungi...). These trophic groups allow reconstructing simplified
food webs that are expected to have the same structural properties as real soil food web. The
development of high-throughput species identification methods (e.g. eDNA metabarcoding),
and the availability of massive amounts of data about trophic interactions and feeding habits
collected by researchers over the past decades have paved the way for new knowledge-based
methods to automate the reconstruction of food webs on an unprecedented scale [
        <xref ref-type="bibr" rid="ref2">2</xref>
        ]. Still,
some problems need to be solved first, the most important of which is probably the lack of
consensus on what exactly a trophic group is and how to classify soil organisms. This absence
of formal description of trophic knowledge prevents the consistent mapping of consumers (e.g.
Acrobeloides sp.) to their trophic group(s) (e.g. bacterivore) and resources (e.g. bacteria belonging
to Alphaproteobacteria), which is needed to automatically assign a species to a trophic group
based on its known interactions, or conversely, to predict the species potential interactions
based on the trophic group(s) to which it belongs.
      </p>
      <p>To address this issue, we are developing the Soil Trophic Web Ontology, whose role is to
provide formal definitions of trophic groups that are consistent across all taxonomic groups
of soil organisms, together with an ontological structure that enables inference in support of
our main objective which is to automate the reconstruction of food webs from community
inventories (Fig. 1).</p>
    </sec>
    <sec id="sec-2">
      <title>2. The Soil Trophic Web Ontology</title>
      <p>The Soil Trophic Web Ontology (STWO) is a domain ontology which represents and maps
together knowledge about trophic interactions (consumer-resource relationships) and trophic
groups (feeding habits, diets). Following good practices of ontology development, STWO is built
by leveraging existing resources as much as possible. In particular, STWO extends a "trophic
subset" of the ECOCORE ontology of core ecological entities with additional classes for missing
trophic groups and resources. As much as possible, classes for resources are imported from
existing OBO ontologies (Fig. 2). STWO includes classes to represent trophic groups at diferent
resolutions (e.g. heterotroph, decomposer, saproxylophage). Trophic resources may be of diferent
types: an organism represented by a taxonomic unit (e.g. Bacteria, Fungi, Viridiplantae), an
anatomical part of an organism (e.g. leaf, mycelium, blood), and any type of environmental
material (e.g. carbon dioxyde, soil organic matter). STWO also reuses object properties from
the Relation Ontology (RO) to describe trophic interactions (e.g. eats, acquires nutrients from).
Finally, STWO provides logical definitions of trophic groups in the form of OWL equivalence
axioms, which makes it possible, using an OWL reasoner, to infer the trophic group(s) an
organism belongs to based on the resources it consumes, as well as to predict potential trophic
interactions from its feeding regime (Fig. 3).</p>
      <p>
        STWO development process is both collaborative and iterative. The initial version of the
ontology was created from a list of relevant terms (trophic groups and resources) and their
definitions created by a small specialized group of soil ecologists. A subset of these terms could
be mapped to existing resources using the Ontobee search engine. These resource identifiers
(URIs) were used as "seeds" for ROBOT’s MIREOT extraction method [
        <xref ref-type="bibr" rid="ref3">3</xref>
        ]. This method enables
to extract a subset of terms (a module) from an external ontology instead of importing the
whole ontology, while preserving the subclasses/subproperties hierarchy. The resulting modules
where merged to form the backbone of STWO. Missing classes and their logical definitions
where added manually using the Protégé editor. The whole development workflow (extraction,
merging, validation, release, versioning) is managed using the Ontology Development Kit [4].
Each new release of STWO is submitted to a group of &gt;20 international experts in soil ecology
to collect their feedback as well as suggestions for revisions and new terms. Debatable points
are discussed and agreed upon using collaborative decision-making tools. The ontology is thus
progressively corrected and enriched with each new iterations.
      </p>
    </sec>
    <sec id="sec-3">
      <title>3. Project Status and Future work</title>
      <p>STWO development is now in its second iteration. The first draft of the ontology includes over
60 newly-defined resource and trophic group classes. Experts are in the process of agreeing on
the revisions to be made to the current version. Once the terms and structure of the ontology
are stabilized, we plan to submit a request for new content and revision to the ECOCORE team,
so that STWO new classes and axioms for trophic groups and resources are made publicly
accessible as part of ECOCORE. It is our wish that the work of our team of soil ecology experts
benefits to a large community of users. We will also consider adding new properties to describe
potential trophic interactions, which would be useful to distinguish between documented and
inferred interactions (e.g. using trait matching).</p>
    </sec>
    <sec id="sec-4">
      <title>Acknowledgments</title>
      <p>The research received funding from the French Agence Nationale de la Recherche (ANR) through
the GlobNets (ANR-16-CE02-0009) project and through MIAI@Grenoble Alpes
(ANR-19-P3IA0003).
[4] N. Matentzoglu, INCATools/ontology-development-kit: June 2020 release (2021). doi:10.
5281/zenodo.4973944.</p>
    </sec>
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