TaxonImprint v1.5 (Beta Release)
Eikoniki, 2025

TaxonImprint is an evidentiary metadata system designed to formalise the 
explicit evidentiary basis of species-level taxonomy. The v1.5 Beta Release 
provides a stable lexicon and extraction protocol for public testing and 
evaluation. This release is intended to allow users to assess reproducibility, 
clarity, and suitability across a wide range of modern taxonomic works.

Users may apply the protocol manually or through a large language model. 
Reproducibility can be tested independently on any species-level taxonomic 
paper. No copyrighted text is included in this release.

All materials in v1.5 remain subject to refinement based on community feedback 
and will be version-controlled in subsequent releases. The definitive technical 
description of TaxonImprint will be published in Eikoniki.

    '
  domains:
    IMAG:
      description: Visual documentation used as evidentiary support, regardless of subject.
      tokens:
        IMAG.PHOTO:
          definition: Photographic images (including macro, micro, stacked, or in situ) used as taxonomic
            evidence.
          applies_when: 'Applied when photographic figures are cited or interpreted to show diagnostic
            characters, variation, or species differences.

            '
          examples:
            generic: Photographs are used to highlight the distinctive shell sculpture of the new species.
            gastropod: Stacked macro-photos illustrate the protoconch and teleoconch transition in a
              volutid.
          boundary_notes: 'Images mentioned only as illustrative, without explicit diagnostic or comparative
            use, do not trigger this token.

            '
        IMAG.SEM:
          definition: Scanning electron micrographs used as taxonomic evidence.
          applies_when: 'Applied when SEM images are used to display microstructure, radula, protoconch
            sculpture, or anatomical surfaces in the taxonomic argument.

            '
          examples:
            generic: SEM micrographs reveal micropitted sculpture on the cuticle.
            gastropod: SEM images of the radula are used to diagnose cusp arrangement in Conidae.
          boundary_notes: 'Morphological content revealed by SEM is recorded in MORPH.MICRO, MORPH.RAD,
            or ANAT as appropriate; IMAG.SEM captures only the imaging modality.

            '
        IMAG.ILLUST:
          definition: New scientific illustrations, diagrams, or line drawings used as taxonomic evidence.
          applies_when: 'Applied when newly created drawings or diagrams are cited or interpreted to show
            characters, structures, or comparative detail in support of a taxonomic conclusion.

            '
          examples:
            generic: Line drawings show the distinctive shape of the male genitalia.
            gastropod: Camera lucida drawings of the radula differentiate similar mitrid species.
          boundary_notes: 'Historical illustrations, if used comparatively, fall under COMP.LIT (if that
            were retained); here IMAG.ILLUST refers to new illustrations produced in the focal work.

            '
        IMAG.HIST:
          definition: Historical figures or plates reused or reinterpreted as taxonomic evidence.
          applies_when: 'Applied when historical images (old plates, lithographs, or earlier photographic
            figures) are referenced explicitly in the argument as evidentiary support.

            '
          examples:
            generic: The original plate from 1850 is re-examined to confirm the presence of a dorsal keel.
            gastropod: Reinterpretation of Reeve''s Cymbiola plates helps stabilise the identity of an
              old name.
          boundary_notes: 'If historical figures are cited only as part of a literature review without
            explicit evidentiary use, they do not trigger this token.

            '
        IMAG.3D:
          definition: Three-dimensional reconstructions (e.g. \u03bctomography, surface scans) used as
            evidence.
          applies_when: 'Applied when 3D reconstructions of shell, anatomy, or internal structures are
            used explicitly in taxonomic reasoning.

            '
          examples:
            generic: \u03bctomography reveals internal shell partitions diagnostic for the clade.
            gastropod: 3D reconstructions of the columellar canal distinguish closely related volutids.
          boundary_notes: 'Underlying anatomical or shell characters are also coded under PHYSIOL.MORPH
            or PHYSIOL.ANAT as appropriate.

            '
    GENE:
      description: 'DNA sequence evidence supporting taxonomy. GENE.MULTI refers to multi-locus datasets
        (Sanger, RAD, UCEs, nuclear+mito mixes). GENE.MITO refers to full or nearly-complete mitochondrial
        genomes.

        '
      tokens:
        GENE.MULTI:
          definition: Use of multi-locus datasets comprising two or more independent loci as taxonomic
            evidence.
          applies_when: 'Applied when the study uses multi-locus datasets (e.g. combined mitochondrial
            and nuclear markers, UCEs, RADseq, exon-capture) in taxonomic inference.

            '
          examples:
            generic: Phylogeny based on COI, 16S, and 28S reveals three lineages.
            gastropod: An exon-capture dataset of hundreds of loci resolves relationships among Cypraeidae.
          boundary_notes: 'Individual loci also trigger their specific GENE tokens; GENE.MULTI is recorded
            in addition for explicitly multi-locus datasets.

            '
        GENE.MITO:
          definition: Use of complete or near-complete mitochondrial genome data as taxonomic evidence.
          applies_when: 'Applied when mitochondrial genomes (partial or full) are analysed or compared
            directly in taxonomic reasoning.

            '
          examples:
            generic: Mitogenome organisation supports placement of the lineage in family X.
            gastropod: Volutid mitogenomes reveal rearranged tRNA clusters diagnostic for the clade.
          boundary_notes: 'Use of only short mitochondrial fragments does not trigger GENE.MITO and is
            captured by the relevant single-locus tokens.

            '
      subdomains:
        LOCI:
          description: Locus-specific mitochondrial and nuclear sequence evidence (e.g. COI, 16S, 28S)
            used explicitly in taxonomy.
          tokens:
            GENE.LOCI.COXI:
              definition: Use of mitochondrial COI sequence data as taxonomic evidence.
              applies_when: 'Applied when COI sequences are generated, cited, or used in comparisons,
                phylogenies, or delimitation decisions.

                '
              examples:
                generic: COI sequences differentiate the two lineages by 7.8% K2P.
                gastropod: Volutid species delimitation is based on COI barcode divergence.
              boundary_notes: 'Phylogenetic trees derived from COI fall under PHYL; the presence of COI
                data itself is recorded by GENE.COI.

                '
            GENE.LOCI.16S:
              definition: Use of mitochondrial 16S rRNA sequence data as taxonomic evidence.
              applies_when: 'Applied when 16S sequences are obtained, cited, and used in explicit comparisons
                or phylogenies for taxonomic purposes.

                '
              examples:
                generic: 16S sequences reveal two well-supported clades.
                gastropod: 16S divergence distinguishes sympatric rissooidean lineages.
              boundary_notes: 'Any phylogenetic interpretation based on 16S is recorded using PHYL tokens.

                '
            GENE.LOCI.28S:
              definition: Use of nuclear 28S rRNA sequence data as taxonomic evidence.
              applies_when: 'Applied when 28S sequences are included in the evidentiary basis for taxonomy,
                whether alone or in combination with other loci.

                '
              examples:
                generic: 28S alignment shows consistent diagnostic substitutions between species.
                gastropod: 28S sequences differentiate species within Turridae.
              boundary_notes: 'Nuclear ribosomal evidence is distinct from mitochondrial data, which is
                covered by GENE.COI, GENE.16S, and GENE.MITO.

                '
    PHYL:
      description: Phylogenetic trees and clade structure used as explicit evidence.
      tokens:
        PHYL.TREE:
          definition: Explicit use of phylogenetic trees (any method) to support taxonomic decisions.
          applies_when: 'Applied when phylogenetic trees are cited, interpreted, or invoked in diagnosis,
            delimitation, or synonymic reasoning.

            '
          examples:
            generic: The tree shows two reciprocally monophyletic clades.
            gastropod: The ML tree supports splitting the widespread turrid into three species.
          boundary_notes: 'Underlying sequence evidence is recorded under GENE; only the use of tree structure
            is captured here.

            '
        PHYL.CLADE:
          definition: Use of clade membership or monophyly as an evidentiary argument.
          applies_when: 'Applied when monophyly, sister-group relationships, or clade support are invoked
            explicitly as evidence for taxonomic decisions.

            '
          examples:
            generic: The species is placed in clade A, sister to clade B.
            gastropod: The new volute falls within a well-supported clade corresponding to subgenus X.
          boundary_notes: 'Often co-occurs with PHYL.TREE; recorded when clade concepts themselves are
            used as arguments.

            '
        PHYL.DATED:
          definition: Use of time-calibrated phylogenies or divergence dates as evidence.
          applies_when: 'Applied when dated trees, divergence-time estimates, or temporal splits are invoked
            to interpret species status, radiation timing, or biogeographic scenarios relevant to taxonomy.

            '
          examples:
            generic: Divergence dating places the split between the lineages in the Pliocene.
            gastropod: Dated phylogeny of cone snails suggests rapid Pleistocene diversification in the
              Indo-Pacific.
          boundary_notes: 'Purely palaeontological age assignments without phylogenetic context belong
            under GEOD.TIME.

            '
    ECO:
      description: Ecological traits, niches, and habitat features used explicitly in taxonomic arguments.
      tokens:
        ECO.HABITAT:
          definition: Habitat type (substrate, exposure, vegetation, etc.) used as taxonomic evidence.
          applies_when: 'Applied when habitat differences are invoked as relevant to species delimitation
            or diagnosis.

            '
          examples:
            generic: One species occurs in rocky intertidal zones while another is restricted to sandy
              flats.
            gastropod: Cymbiola A is confined to muddy slope habitats, while Cymbiola B inhabits shallow
              reefal sand.
          boundary_notes: 'Habitat mentioned only in passing or as background ecology does not trigger
            this token.

            '
        ECO.MICROHABITAT:
          definition: Fine-scale microhabitat differences used as evidence.
          applies_when: 'Applied when microhabitat (e.g. under stones, within specific host sponges, in
            seagrass beds) is invoked as a key difference among taxa.

            '
          examples:
            generic: The new species is found exclusively within dead coral heads.
            gastropod: A rissooidean snail restricted to mangrove pneumatophores is distinguished from
              a similar species on seagrass leaves.
          boundary_notes: 'Broader habitat-level distinctions should be captured under ECO.HABITAT; here
            the emphasis is on micro-scale ecological niches.

            '
        ECO.DIET:
          definition: Diet, prey specificity, or host association used as taxonomic evidence.
          applies_when: 'Applied when dietary differences or host specificity (e.g. parasitic or predatory
            associations) are used in delimiting or distinguishing taxa.

            '
          examples:
            generic: One species feeds exclusively on bivalves, the other on echinoderms.
            gastropod: Conus species with distinct prey preferences (fish vs. mollusc) are separated.
          boundary_notes: 'Stable isotopes or biochemical diet proxies fall under CHEM or BIOC, not ECO.DIET,
            unless specifically framed as ecological evidence.

            '
    CHEM:
      description: Chemical, geochemical, and biochemical evidence used explicitly as part of taxonomic reasoning.
      tokens:
        CHEM.ICPMS:
          definition: Bulk or micro-sample elemental or isotopic compositions (e.g. ICP-MS, LA-ICP-MS)
            used to differentiate or characterise taxa.
          applies_when: 'Applied when shell or tissue chemistry is measured and interpreted as evidence
            of distinct species, populations, or lineages.

            '
          examples:
            generic: ICP-MS data show discrete elemental profiles between populations.
            gastropod: Trace-element signatures in shells distinguish sympatric volute species.
          boundary_notes: 'Purely environmental chemical data without direct linkage to the specimens
            do not trigger CHEM.ICPMS.

            '
        CHEM.RAMAN:
          definition: Raman spectroscopy or similar vibrational techniques used as evidence.
          applies_when: 'Applied when Raman spectra of shell, pigments, or soft tissues are interpreted
            taxonomically.

            '
          examples:
            generic: Raman spectra indicate the presence of aragonite vs calcite in different species.
            gastropod: Raman analysis of shell pigments differentiates cryptic colour morphs.
          boundary_notes: 'If used solely to confirm standard mineralogy with no taxonomic argument, this
            token is not triggered.

            '
        CHEM.ISOTOPE:
          definition: Stable or radiogenic isotope data interpreted as taxonomic evidence.
          applies_when: 'Applied when isotopic signatures (C, O, Sr, etc.) are used to distinguish taxa,
            infer habitat partitioning, or support evolutionary scenarios central to taxonomy.

            '
          examples:
            generic: Oxygen isotope ratios distinguish cool-temperate and warm-temperate lineages.
            gastropod: Carbon isotopes reveal trophic differences supporting separation of species in
              a complex of mud snails.
          boundary_notes: 'Isotope data used only for palaeoenvironmental reconstruction, without explicit
            taxonomic import, do not trigger CHEM.ISOTOPE.

            '
      subdomains:
        BIOC:
          description: Biochemical, bio-optical, or biophysical evidence used in taxonomic interpretation.
          tokens:
            BIOC.VEN:
              definition: Biochemical or proteomic characterisation of venom (or other secretions) used as
                taxonomic evidence.
              applies_when: 'Applied when venom composition, proteomic profiles, or toxin repertoires are
                used to distinguish taxa or infer relationships.

                '
              examples:
                generic: Venom proteomes differ markedly between the two forms.
                gastropod: Conus venom peptides identify distinct toxin clades linked to species boundaries.
              boundary_notes: 'Purely anatomical venom apparatus traits are ANAT.TOX; genetic evidence for
                venom genes is GENE, not BIOC.VEN.

                '
            BIOC.LUM:
              definition: Bioluminescent or photobiological traits used explicitly as taxonomic evidence.
              applies_when: 'Applied when presence, absence, or properties of bioluminescence or bio-optical
                signals are invoked as diagnostic or comparative traits.

                '
              examples:
                generic: Only one of the two species exhibits flashing bioluminescence.
                gastropod: A deep-sea snail''s bioluminescent mantle spots are cited as unique within its
                  genus.
              boundary_notes: 'Underlying photoreceptor anatomy belongs to ANAT; light spectra may also
                trigger CHEM if chemically based.


            '
    STAT:
      description: 'Quantitative or model-based evidence used in species delimitation, morphometrics,
        or population structure.

        '
      tokens:
        STAT.MORPH:
          definition: Morphometric or shape-analytic methods used as taxonomic evidence.
          applies_when: 'Applied when statistical analysis of morphology (e.g. PCA, geometric morphometrics,
            ratio analyses) is used to assess species limits or differences.

            '
          examples:
            generic: PCA of shell shape separates the two lineages.
            gastropod: Geometric morphometrics show non-overlapping shell outlines in rissooidean snails.
          boundary_notes: 'Underlying morphology is also coded under MORPH tokens; STAT.MORPH captures
            the analytical method.

            '
        STAT.STRUCTURE:
          definition: Population-structure analyses (e.g. STRUCTURE, fineRADstructure, ADMIXTURE) used
            as evidence.
          applies_when: 'Applied when inferred population clusters, admixture patterns, or genetic partitions
            are used to support or refute species boundaries.

            '
          examples:
            generic: STRUCTURE reveals two non-admixed genetic clusters corresponding to morphotypes.
            gastropod: Genetic clustering analyses identify divergent populations of a mud snail complex.
          boundary_notes: 'Raw genetic loci fall under GENE; statistical clustering methods are captured
            here.

            '
        STAT.ABGD:
          definition: Use of the ABGD method as species-delimitation evidence.
          applies_when: 'Applied when ABGD output (number of groups, partitions) is invoked to justify
            or interpret species boundaries.

            '
          examples:
            generic: ABGD split the samples into three putative species.
            gastropod: ABGD partitions match protoconch-based morphotypes in rissooideans.
          boundary_notes: 'ABGD operates on genetic distances; the sequences themselves are captured under
            the appropriate GENE tokens.

            '
        STAT.GMYC:
          definition: Use of the GMYC method as species-delimitation evidence.
          applies_when: 'Applied when GMYC results are invoked to justify or interpret species limits.

            '
          examples:
            generic: GMYC recognised five independently evolving lineages.
            gastropod: GMYC split a widespread turrid complex into three deeply divergent entities.
          boundary_notes: 'GMYC is usually applied to ultrametric trees; the underlying tree is recorded
            under PHYL.TREE, and the GMYC analysis under STAT.GMYC.

            '
        STAT.bPTP:
          definition: Use of the (b)PTP method as species-delimitation evidence.
          applies_when: 'Applied when PTP or bPTP species partitions are cited or interpreted as supporting
            particular taxonomic decisions.

            '
          examples:
            generic: bPTP analysis supports recognition of two species.
            gastropod: PTP delimitation corresponds to shell-morphological differences in a conoidean
              complex.
          boundary_notes: 'The phylogeny on which PTP is run is recorded under PHYL.TREE; the delimitation
            method under STAT.bPTP.

            '
        STAT.BPP:
          definition: Use of Bayesian Phylogenetics and Phylogeography (BPP) as species-delimitation evidence.
          applies_when: 'Applied when BPP analyses and posterior probabilities are used to justify acceptance
            or rejection of candidate species.

            '
          examples:
            generic: BPP strongly supports three species rather than two.
            gastropod: BPP delimitation using multi-locus data identifies four well-supported volute
              lineages.
          boundary_notes: 'Underlying loci and trees are recorded under GENE and PHYL; STAT.BPP captures
            the BPP analysis itself.

            '
        STAT.ASAP:
          definition: Use of ASAP (Assemble Species by Automatic Partitioning) as species-delimitation
            evidence.
          applies_when: 'Applied when ASAP partitions are invoked explicitly to justify species boundaries
            or candidate species sets.

            '
          examples:
            generic: ASAP recognised six genetic clusters corresponding to morphological forms.
            gastropod: ASAP supports separation of sympatric cone-snail morphs into distinct species.
          boundary_notes: 'ASAP is a distance-based delimitation method; underlying data are captured
            by GENE tokens.

            '
    DEV:
      description: Developmental mode and life-history traits used as taxonomic evidence.
      tokens:
        DEV.MODE:
          definition: Developmental mode (e.g. planktotrophic vs lecithotrophic, direct vs indirect development)
            used explicitly in taxonomic reasoning.
          applies_when: 'Applied when developmental mode is inferred (e.g. from protoconch) or directly
            observed and used to support species boundaries or evolutionary interpretation.

            '
          examples:
            generic: One species has planktotrophic larvae, the other has direct development.
            gastropod: A paucispiral protoconch is used to infer non-planktotrophic development in a
              new volutid.
          boundary_notes: 'Inferences based on protoconch morphology also trigger MORPH.PROT; DEV.MODE
            captures the developmental interpretation.

            '
    LIFE:
      description: Life-history traits beyond development used as evidence.
      tokens:
        LIFE.PARASITIC:
          definition: Parasitic vs free-living lifestyle used as taxonomic evidence.
          applies_when: 'Applied when the presence, absence, or specificity of parasitism is invoked as
            a key distinction.

            '
          examples:
            generic: One lineage is parasitic on echinoderms, the other is free-living.
            gastropod: A parasitic eulimellid is distinguished from a non-parasitic relative on life-history
              grounds.
          boundary_notes: 'Host identity, if important, also triggers ECO.DIET or ECO.MICROHABITAT.

            '
    BEH:
      description: Behavioural traits used as explicit evidence.
      tokens:
        BEH.SPAWNING:
          definition: Spawning or mating behaviour used in taxonomic interpretation.
          applies_when: 'Applied when distinct spawning times, behaviours, or courtship patterns are invoked
            as evidence for separation of taxa.

            '
          examples:
            generic: Two taxa spawn in different months and show distinct courtship displays.
            gastropod: A nudibranch species with unique mating behaviour is distinguished from its congeners.
          boundary_notes: 'Behaviour observed but not used in taxonomic reasoning does not trigger this
            token.

            '
    PHYSIOL:
      description: Physiological evidence encompassing morphology and soft-part anatomy (hard parts and
        soft tissues) used explicitly in extant Gastropoda taxonomy.
      subdomains:
        MORPH:
          description: Shell, protoconch, operculum, radula, and microstructural morphology (hard parts).
          tokens:
            MORPH.SHELL:
              definition: Adult teleoconch morphology, including overall shell shape, proportions, visible
                sculpture, and colour pattern.
              applies_when: 'Applied when adult shell form or visible surface characters are explicitly
                used as diagnostic or comparative evidence.

                '
              examples:
                generic: Shell elongate-ovate with fine spiral cords.
                gastropod: The new volutid species has a tall spire and inflated last whorl.
              boundary_notes: 'Protoconch characters are recorded under MORPH.PROT, and SEM-only microstructure
                under MORPH.MICRO; internal soft-part features are ANAT.

                '
            MORPH.PROT:
              definition: Protoconch morphology, including whorl number, shape, and sculpture.
              applies_when: 'Applied when the protoconch is explicitly mentioned, figured, or compared
                in the taxonomic argument.

                '
              examples:
                generic: Protoconch multispiral with two smooth whorls.
                gastropod: The species has a paucispiral protoconch consistent with non-planktotrophy.
              boundary_notes: 'Developmental mode inferred from protoconch is recorded separately under
                DEV.MODE only if explicitly invoked as evidence.

                '
            MORPH.OPERC:
              definition: Operculum morphology, including shape, size, material, and nucleus position.
              applies_when: 'Applied when the operculum is described, figured, or compared as part of
                the evidentiary basis.

                '
              examples:
                generic: Operculum corneous, oval, with terminal nucleus.
                gastropod: The naticid has a thick, calcareous operculum unlike its congeners.
              boundary_notes: 'Soft-tissue attachments revealed only by dissection belong to ANAT.

                '
            MORPH.RAD:
              definition: Radula morphology, including tooth formula, cusp number, shape, and arrangement.
              applies_when: 'Applied when radular features are described or illustrated and used taxonomically.

                '
              examples:
                generic: Radula with formula 1-3-1 and tricuspid lateral teeth.
                gastropod: The cone snail has long, barbed radular teeth characteristic of piscivory.
              boundary_notes: 'Internal foregut structures and venom apparatus are ANAT.FOREGUT or ANAT.TOX,
                not MORPH.RAD.

                '
            MORPH.MICRO:
              definition: Shell or surface microstructure, including SEM-resolved sculpture.
              applies_when: 'Applied when microstructural details (e.g. micro-pits, granules, crossed-lamellar
                fabrics) are used as evidence.

                '
              examples:
                generic: SEM reveals a microgranulate periostracum.
                gastropod: The volutid shell has fine intritical spiral striae visible only under SEM.
              boundary_notes: 'IMAG.SEM captures the imaging method, while MORPH.MICRO captures the morphological
                content; internal anatomical microstructure belongs under ANAT.

                '
        ANAT:
          description: Soft-part anatomical traits requiring dissection or internal imaging.
          tokens:
            ANAT.ANIM:
              definition: External body anatomy (colour, papillae, mantle, tentacles, foot).
              applies_when: 'Applied when external soft tissues are described or compared and used as
                evidence.

                '
              examples:
                generic: Mantle edge with distinct tentacular processes.
                gastropod: The nudibranch has elongate dorsal papillae absent in related species.
              boundary_notes: 'Shell and operculum are MORPH; internal soft tissues belong to ANAT.FOREGUT,
                ANAT.DIGESTIVE, ANAT.ORG, ANAT.CIRC, or ANAT.TOX as appropriate.

                '
            ANAT.FOREGUT:
              definition: Foregut structures, including buccal mass, odontophore, proboscis, salivary
                glands, and Leiblein gland.
              applies_when: 'Applied when foregut anatomy is dissected, illustrated, or used explicitly
                in taxonomic comparison.

                '
              examples:
                generic: A muscular proboscis with large salivary glands.
                gastropod: The presence of a well-developed gland of Leiblein separates conoideans from
                  other neogastropods.
              boundary_notes: 'Radular morphology is MORPH.RAD; venom bulbs and ducts are ANAT.TOX.

                '
            ANAT.DIGESTIVE:
              definition: Digestive tract anatomy beyond the foregut, including stomach, intestine, digestive
                gland, and associated structures.
              applies_when: 'Applied when midgut or hindgut structures are used in taxonomic reasoning.

                '
              examples:
                generic: A looped intestinal tract characteristic of the group.
                gastropod: The presence of a caecal appendix on the stomach is diagnostic.
              boundary_notes: 'Foregut-specific traits remain ANAT.FOREGUT; histological details may also
                be covered by ANAT.ORG if organ-level.

                '
            ANAT.REPRO:
              definition: Reproductive system anatomy, including male and female ducts, glands, and accessory
                structures.
              applies_when: 'Applied when reproductive anatomy is dissected, described, or figured and
                used as taxonomic evidence.

                '
              examples:
                generic: A long, coiled vas deferens with a large prostate gland.
                gastropod: The volutid has a distinctive penial papilla and accessory gland complex.
              boundary_notes: 'Developmental mode (e.g. planktotrophic vs direct) is recorded under DEV.MODE.

                '
            ANAT.ORG:
              definition: Organ-level anatomical traits outside the foregut and reproductive system, including
                heart, kidney, ctenidium, osphradium, and mantle-cavity organs.
              applies_when: 'Applied when such organs are dissected and used as explicit taxonomic evidence.

                '
              examples:
                generic: The species has a single monopectinate ctenidium.
                gastropod: The volutid has a short osphradium with 6\u20137 lamellae.
              boundary_notes: 'Circulatory and hemal details can also be captured more specifically under
                ANAT.CIRC.

                '
            ANAT.CIRC:
              definition: Circulatory and hemal system anatomy, including heart chambers and hemocoelic
                structures.
              applies_when: 'Applied when circulatory or hemal structures are described and used in taxonomic
                comparison or diagnosis.

                '
              examples:
                generic: The auricle is small and positioned posteriorly to the ventricle.
                gastropod: The haliotid shows enlarged hemocoelic sinuses in the mantle cavity.
              boundary_notes: 'Biochemical properties of blood or pigments are BIOC or CHEM, not ANAT.

                '
            ANAT.TOX:
              definition: Venom apparatus anatomy, including venom duct, venom gland, venom bulb, and
                associated stylet or sac structures.
              applies_when: 'Applied when internal venom-delivery structures are described, illustrated,
                or compared in the taxonomic argument.

                '
              examples:
                generic: The venom bulb is elongate, with a narrow duct entering the buccal region.
                gastropod: The Conus species has a proportionally long venom duct running parallel to
                  the oesophagus.
              boundary_notes: 'Venom biochemical or proteomic profiles are BIOC.VEN; foregut glands such
                as the gland of Leiblein are ANAT.FOREGUT; radular tooth morphology remains MORPH.RAD.

                '
    SPATIAL:
      description: Spatial, bathymetric, altitudinal, temporal, and barrier evidence used explicitly in
        taxonomy.
      subdomains:
        GEOD:
          description: Spatial, bathymetric, altitudinal, temporal, and barrier evidence used explicitly
            in taxonomy.
          tokens:
            GEOD.LOC:
              definition: Locality-based or range-based evidence used in taxonomic interpretation.
              applies_when: 'Applied when specific localities, disjunct ranges, or geographic distances
                are invoked as part of the evidentiary argument.

                '
              examples:
                generic: The two forms occur on opposite sides of a 600 km arid barrier.
                gastropod: Indo-Pacific volutid populations isolated across the Sahul Shelf are treated
                  as distinct species.
              boundary_notes: 'Locality listed only as specimen metadata, without being used in argumentation,
                does not trigger GEOD.LOC.

                '
            GEOD.DEPTH:
              definition: Bathymetric evidence, including depth ranges and depth discontinuities, used
                in taxonomy.
              applies_when: 'Applied when depth differences or depth-restricted populations are invoked
                as relevant to species delimitation or comparison.

                '
              examples:
                generic: A shallow-water lineage (<10 m) is distinct from a deep-water lineage (150\u2013300
                  m).
                gastropod: Volutid morphotypes segregated by depth on the continental slope are used to
                  delimit species.
              boundary_notes: 'Depth recorded only as metadata does not trigger this token.

                '
            GEOD.TIME:
              definition: Geological-age evidence (formation, faunal stage, geological period) used in
                taxonomic interpretation.
              applies_when: 'Applied when geological age of occurrences, shells, or deposits is used to
                justify taxon boundaries, relationships, or historical interpretation.

                '
              examples:
                generic: Specimens from Miocene strata represent an ancestral species distinct from the
                  extant form.
                gastropod: Volutid material from the Late Oligocene is treated as a separate fossil species
                  from the living descendant.
              boundary_notes: 'Molecular divergence times inferred from phylogenies are recorded under
                PHYL.DATED.

                '
            GEOD.ALT:
              definition: Altitudinal evidence used in taxonomic reasoning.
              applies_when: 'Applied when altitude or altitudinal separation is used explicitly to distinguish
                taxa or lineages.

                '
              examples:
                generic: One species is restricted to lowland floodplains, another to montane slopes.
                gastropod: A freshwater snail restricted to high-altitude streams is separated from a
                  lowland congener.
              boundary_notes: 'Altitude mentioned only as part of locality metadata does not trigger GEOD.ALT.

                '
            GEOD.CURRENT:
              definition: Oceanographic barriers, currents, or biogeographic breaks used as taxonomic
                evidence.
              applies_when: 'Applied when ocean circulation patterns or major current systems are invoked
                to explain restricted dispersal, lineage isolation, or species limits.

                '
              examples:
                generic: The equatorial current system forms a barrier separating two sister lineages.
                gastropod: The East Australian Current is cited as a major barrier separating Australian
                  and New Caledonian volutes.
              boundary_notes: 'Currents mentioned only as ecological background do not trigger GEOD.CURRENT.

                '
    MODEL:
      description: Model-based structures (including species distribution models) used explicitly as taxonomic
        evidence.
      subdomains:
        DISTR:
          description: Species distribution modelling used explicitly as evidence.
          tokens:
            DISTR.ALG:
              definition: Use of SDM/ENM algorithms (e.g. Maxent, BIOCLIM, GARP) as taxonomic evidence.
              applies_when: 'Applied when modelled habitat suitability, predicted ranges, or SDM-derived
                clusters are used to support species boundaries or comparative interpretations.

                '
              examples:
                generic: Maxent models identified two non-overlapping climatic niches supporting recognition
                  of separate species.
                gastropod: SDM predictions for New Caledonian versus Australian volutes show distinct
                  ecological envelopes consistent with genetic divergence.
              boundary_notes: 'Maps of observed occurrences without modelling are recorded under GEOD.LOC;
                climate data without modelling, when used as evidence, belong to ECO.

                '
    VOUCHER:
      description: Formally designated voucher objects (including type material) examined or used explicitly
        as taxonomic evidence.
      subdomains:
        TYPE:
          description: Type material examined or used explicitly as taxonomic evidence.
          tokens:
            VOUCHER.TYPE.HOLO:
              definition: Use of the holotype (or name-bearing type such as a designated lectotype) as
                explicit evidence.
              applies_when: 'Applied when the holotype is examined, re-examined, figured, measured, or
                compared to support species delimitation, identity, or synonymy.

                '
              examples:
                generic: Examination of the holotype confirms the presence of three axial ribs on early
                  whorls.
                gastropod: The lectotype of Cymbiola cathcartiae stabilises interpretation of the SNG
                  lineage.
              boundary_notes: 'Reference to type locality alone, without examination of the specimen,
                does not trigger TYPE.HOLO.

                '
            VOUCHER.TYPE.PARA:
              definition: "Use of paratypes, paralectotypes, or non\u2013name-bearing syntypes as explicit\
                \ evidence."
              applies_when: 'Applied when paratype or related non\u2013name-bearing type material is examined,
                measured, or figured to assess variation or support species boundaries.

                '
              examples:
                generic: Paratypes show consistent shell sculpture across localities.
                gastropod: Paratypes of the new volutid exhibit identical protoconch morphology.
              boundary_notes: 'Literature-based mention of paratypes without indication of examination
                does not trigger this token.

                '
  token_string_model:
    description: >
      Ev1 tokens are expressed as a space-separated, uppercase dot-notation string for
      each species-level taxon. Each token string summarises the evidentiary basis of the
      taxon across its entire taxonomic history (original description + any redescriptions).
    syntax:
      basic: "MORPH.SHELL MORPH.PROT ANAT.REPRO GENE.COI PHYL.TREE"
      ordered: "Tokens may optionally be grouped or ordered by domain, but ordering conveys
        no semantic weight."
    rules:
      - "Tokens are deterministic: a token is either present (evidence used) or absent (not used)."
      - "Tokens represent evidence, not interpretations or conclusions."
      - "Tokens are non-evaluative: they do not encode quality, only presence/absence."
      - "Token strings may be extended as new evidence appears in later works."
  provenance_model:
    description: Optional mapping of tokens to the publication(s) in which the evidence appears.
    syntax:
      compact: TOKEN[Source1|Source2]
      structured:
        example:
          MORPH.SHELL:
            - TW1877: Tenison Woods 1877, figs 1–2
            - JK2024: Johnson & Korshunova 2024, fig. 5
    rules:
      - Sources must be citable publications.
      - Provenance lists may contain multiple sources.
      - Provenance is descriptive, not evaluative.
