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Incertae sedis

Incertae sedis (Latin for "of uncertain position") is a taxonomic term used in biological classification to designate groups or taxa whose phylogenetic relationships or exact placement within a hierarchical system remain unresolved due to insufficient evidence.^[1]^[2] This designation acknowledges the validity of the taxon while highlighting the ambiguity in its systematic position, often applied when morphological, molecular, or other data are inadequate for precise assignment to a family, order, or higher rank.^[3] The term is formally recognized in the International Code of Zoological Nomenclature (ICZN) as indicating uncertain taxonomic position and is similarly employed in botanical nomenclature under the International Code of Nomenclature for algae, fungi, and plants (ICN), though not explicitly defined in the latter.^[1]^[4] In practice, incertae sedis taxa are frequently placed provisionally under broader categories, such as orders or classes, to maintain organizational structure without implying resolved affinities.^[2] This approach facilitates ongoing research and accommodates revisions as new phylogenetic analyses—often incorporating DNA sequencing or fossil evidence—clarify relationships.^[3] Common in fields like paleontology, microbiology, and invertebrate zoology, incertae sedis highlights gaps in biodiversity knowledge and underscores the dynamic nature of taxonomy.^[5] For instance, many extinct genera or microbial lineages are classified this way pending further study, preventing premature or erroneous assignments that could distort evolutionary understanding.^[6] The abbreviation "inc. sed." is standard in scientific literature to denote such uncertainty efficiently.^[7]

Definition and Etymology

Literal Meaning

The Latin phrase incertae sedis directly translates to "of uncertain seat" or "of uncertain position," reflecting its use to denote ambiguity in placement. The component "incertae" derives from the genitive singular feminine form of the adjective incertus, which means "uncertain," "doubtful," or "unsettled."^[8] Similarly, "sedis" is the genitive singular form of the noun sedes, signifying "seat," "abode," "place," or "position."^[9] In biological taxonomy, incertae sedis specifically indicates a taxon or group whose broader relationships or exact position within a hierarchical classification remain unclear or undefined, often due to limited data or unresolved phylogenetic affinities.^[10] This interpretation aligns with its application in formal nomenclature codes, where it serves as a provisional status without implying invalidity. The phrase originated in scientific literature in the late 18th century, with the earliest documented use in botany by Antoine-Laurent de Jussieu in his 1789 work Genera Plantarum, where he listed several genera as plantae incertae sedis.^[11] An example of its application in entomology appears in the 1903 establishment of the moth genus Callambulyx within Smerinthinae incertae sedis.^[12] By the 1920s, it had become a common convention in botanical and zoological works to handle taxa with ambiguous affiliations.^[13]

Historical Usage

The term incertae sedis, derived from classical Latin meaning "of uncertain seat" or "position," was first adopted into biological taxonomy in the late 18th century, originating in botany with Antoine-Laurent de Jussieu's 1789 Genera Plantarum.^[11] It was subsequently borrowed more widely during the 19th century by systematists building on Carl Linnaeus's framework to address taxa whose placement within the hierarchical system remained ambiguous.^[7] This adoption reflected the growing complexity of classifying newly discovered species, particularly in botany and zoology, where Linnaean successors like Augustin Pyramus de Candolle employed Latin phrases to denote provisional or unresolved categorizations without violating strict binomial nomenclature rules.^[14] The term gained prominence in the early 20th century within paleontological literature for describing fossil taxa with unclear affinities, particularly amid discoveries like those from the Burgess Shale. Systematists such as Charles Doolittle Walcott, who described forms like chancelloriids in 1920, highlighted uncertainties in early metazoan evolution, with explicit incertae sedis placements becoming more routine in subsequent decades for such problematic fossils.^[15] Formal integration of incertae sedis into international nomenclatural codes emerged in the mid-20th century, aligning with revisions to the International Code of Botanical Nomenclature (ICBN, later ICN) and the International Code of Zoological Nomenclature (ICZN); by the 1950s, it was acknowledged in ICBN guidelines as a non-ranked status for taxa pending further revision, without conferring formal taxonomic validity.^[16] The ICZN similarly incorporated it in its 1964 edition, defining it in the glossary as denoting uncertain higher-level relationships, which standardized its use across disciplines.^[17] Usage of incertae sedis surged after the 1950s amid cladistic debates, as Willi Hennig's 1950 publication Grundzüge einer Theorie der phylogenetischen Systematik shifted focus to monophyly and sister-group relationships, often leaving basal or fragmentary taxa unresolved in early analyses.^[18] This trend peaked in the 1980s and 1990s during the molecular era, when DNA sequencing revealed discrepancies between morphological and genetic phylogenies, resulting in elevated placements of genera as incertae sedis—for example, numerous fish families in the 1990s percomorph radiation were so designated pending comprehensive molecular datasets.^[6] Such increases underscored the term's role in accommodating taxonomic flux without premature reclassification.^[3]

Applications in Biological Nomenclature

General Principles

In biological nomenclature, the designation of a taxon as incertae sedis serves as a provisional placement indicating that its exact position within the taxonomic hierarchy cannot be determined with current evidence, thereby maintaining the integrity of classifications while acknowledging uncertainty. This practice aligns with the core principles of stability and universality in nomenclature, allowing taxonomists to organize biodiversity without forcing unsubstantiated affiliations.^[19]^[20] The International Code of Zoological Nomenclature (ICZN, 4th edition) defines incertae sedis as a term denoting "of uncertain taxonomic position" and incorporates it into provisions for type species eligibility, where a nominal species cited as incertae sedis is not considered originally included in a genus, ensuring objective nomenclatural application.^[21] Similarly, the International Code of Nomenclature for algae, fungi, and plants (ICN, Madrid Code) permits such placements explicitly for higher ranks, stating that provisions on rank assignment do not preclude designating taxa as incertae sedis relative to families or above, thus supporting flexible yet regulated hierarchical structures.^[22] Both codes emphasize that taxonomic judgments, including incertae sedis assignments, fall outside strict nomenclatural regulation, prioritizing authorial responsibility for evidence-based decisions.^[19]^[20] To formally assign incertae sedis status in publications, taxonomists must include a clear justification, detailing the diagnostic characters, comparative analyses, or data gaps that render placement ambiguous, often as part of open nomenclature (ON) qualifiers like "inc. sed." This is typically documented in peer-reviewed systematic works, where the taxon is appended to an appropriate higher group (e.g., a family or order) with the uncertainty noted, facilitating future revisions without invalidating the name. Provisional ranking may involve temporary subordination, but the name's availability and priority remain governed by standard code requirements, such as publication in a scientific outlet and adherence to typification rules.^[3]^[21] In hierarchical classification, incertae sedis enables structured organization by positioning unresolved taxa at genus, family, or species levels within a broader clade, preserving monophyly where possible and avoiding artificial paraphyletic groups. For instance, a genus may be listed as incertae sedis in a family if subfamily affiliation is unclear, or a species within a genus if subgeneric ties are indeterminate, thereby supporting comprehensive catalogs like those in major biodiversity databases. This approach underscores the codes' commitment to nomenclatural stability over rigid taxonomy, allowing ongoing research to refine placements without retrospective disruptions.^[3]

Botanical Contexts

In botanical nomenclature, the term incertae sedis is applied under the International Code of Nomenclature for algae, fungi, and plants (ICN) to indicate taxa whose precise placement within a hierarchy remains uncertain due to insufficient evidence or ongoing debate.^[20] Article 11 of the ICN addresses priority among names and permits the use of separate names for fossil-taxa that may represent different parts, life-history stages, or preservational states of a single organism, facilitating incertae sedis designations for such ambiguous material without violating priority rules.^[23] Similarly, Article 63 governs the rejection of superfluous names, allowing provisional retention of names for taxa with uncertain affinities until further clarification, thereby supporting temporary incertae sedis status to maintain nomenclatural stability.^[24] These provisions enable botanists to describe and name entities without forcing premature assignments that could later require extensive emendations. Common scenarios for incertae sedis placements in botany include fossil plants, where fragmentary remains often preclude definitive family or higher-rank assignments due to morphological ambiguity. For instance, the fossil genus Paradinandra suecica, from Late Cretaceous deposits in southern Sweden, was placed incertae sedis within the order Ericales because its affinities could not be resolved to a specific family despite clear ericalean characteristics.^[25] Another frequent case involves cryptic species, which are morphologically indistinguishable but genetically distinct, leading to uncertain taxonomic boundaries; in such situations, incertae sedis is used pending molecular or ecological resolution to avoid synonymy errors.^[3] In families like Orchidaceae, incertae sedis is commonly invoked for genera with unresolved subtribal or tribal positions amid the group's rapid diversification and morphological convergence. The genus Devogelia (Epidendroideae), for example, has been designated incertae sedis due to its unique combination of traits that do not align clearly with established clades, as revealed by phylogenetic analyses.^[26] Likewise, Oncidium loxense was placed incertae sedis in the Cyrtochilum alliance owing to ambiguous floral and vegetative features overlapping multiple genera.^[27] These examples highlight how incertae sedis preserves nomenclatural flexibility in complex orchid systematics. The 2025 Madrid Code, superseding the 2018 Shenzhen Code, maintains and refines provisions affecting incertae sedis handling, particularly for hybrid taxa and fungal names, by clarifying rank designations (e.g., Article 3.2 explicitly referencing incertae sedis for uncertain hybrid placements) and electronic publication rules to accelerate descriptions of provisional taxa.^[20] These updates enhance stability for uncertain placements by integrating one-fungus-one-name principles, reducing dual nomenclature ambiguities that previously complicated fungal incertae sedis assignments.^[28]

Zoological Contexts

In zoological nomenclature, the term incertae sedis denotes taxa of uncertain taxonomic position, allowing for provisional classification while maintaining nomenclatural stability under the International Code of Zoological Nomenclature (ICZN).^[1] The ICZN, which governs the naming of animals, prioritizes stability to ensure consistent scientific communication, even when phylogenetic relationships remain unresolved.^[17] This approach contrasts with stricter rejection criteria in other contexts, as the Code explicitly states that a name should not be invalidated solely because its application is uncertain.^[29] Key provisions addressing nomina dubia—names of doubtful or unknown application—and uncertain placements are embedded in the ICZN's framework for availability and validity. Article 11 outlines requirements for name availability, including that genus-group names must function as nouns in the nominative singular, but it does not permit rejection based on uncertainty alone; instead, such names retain availability pending clarification.^[30] Similarly, Article 40 governs family-group nomenclature by ensuring that the validity of a name is unaffected by synonymy in its type genus, thereby supporting stable classifications for taxa potentially placed incertae sedis at higher levels.^[31] These articles underscore the ICZN's emphasis on nomenclatural stability, where uncertain taxonomic positions do not disrupt the priority and availability of names, allowing zoologists to continue using them without immediate reclassification.^[21] The use of incertae sedis is particularly prevalent in invertebrate taxonomy, especially among arthropods, where morphological complexity and fragmentary evidence often hinder precise placement. For instance, numerous fossil arthropods from Paleozoic deposits are assigned incertae sedis due to ambiguous traits that prevent confident allocation to orders or families.^[32] In cases of extinct species, the loss or inadequacy of type specimens frequently results in incertae sedis status, as seen in various Cambrian and Ordovician arthropod genera where holotypes have deteriorated or been destroyed, precluding modern re-examination.^[33] This practice preserves nomenclatural continuity while encouraging further research, such as through new fossil discoveries or molecular analogs in extant forms.^[34] The fourth edition of the ICZN, published in 1999, reinforced these principles by clarifying rules to minimize incertae sedis designations for well-supported clades, promoting the integration of phylogenetic data into stable nomenclature.^[21] Amendments in this edition, including those on publication criteria, aimed to reduce uncertainty by standardizing how evidence is evaluated, though incertae sedis remains essential for taxa lacking robust cladistic support.^[29] This balance has facilitated ongoing revisions in arthropod systematics, where incertae sedis placements serve as placeholders amid advancing paleontological techniques.^[3]

Reasons for Incertae Sedis Placement

Inadequate Taxonomic Description

Inadequate taxonomic descriptions frequently lead to the placement of taxa as incertae sedis due to insufficient diagnostic information that prevents confident assignment to a higher taxonomic rank or clade. Core criteria for inadequacy encompass the lack of a properly designated type specimen, such as a holotype, which serves as the nomenclatural anchor and reference for future comparisons; ambiguous or overly subjective morphological traits, often described in vague terms like "large size" without quantitative measurements or illustrations; and limited comparative analysis against established taxa, resulting in unresolved character states that obscure affinities. These issues are particularly prevalent in older descriptions, where minimal textual accounts or absent type localities violate foundational principles of the International Code of Zoological Nomenclature (ICZN) and International Code of Nomenclature for algae, fungi, and plants (ICN), rendering the taxon unavailable or indeterminable.^[35] The repercussions of such deficiencies are profound, as affected taxa languish in taxonomic limbo, unintegrated into systematic revisions and impeding progress in fields like conservation and evolutionary biology. Without clear placement, these taxa cannot be reliably mapped to biodiversity databases, potentially excluding 5–50% of pertinent records during data synthesis due to synonymy ambiguities or provisional listings. Prior to 2000, inadequate descriptions contributed to taxonomic instability, with approximately 45% of taxa experiencing concept revisions over time as a result of initial shortcomings in circumscription and character definition. This historical burden underscores the challenges in stabilizing nomenclature for the estimated 15,000–20,000 new species described annually, many of which risk similar fates without rigorous standards.^[36] Addressing incertae sedis status stemming from descriptive inadequacies requires targeted redescription to furnish robust, verifiable diagnostics while adhering to formal nomenclatural procedures. Contemporary strategies emphasize supplementing original material with modern tools, including high-resolution imaging (e.g., micro-CT scans) for non-destructive morphological documentation and DNA sequencing for phylogenetic context, thereby enabling precise character delineation without supplanting the type. A notable case is the genus Apodibius (Tardigrada: Eutardigrada), previously incertae sedis due to sparse morphological data; its position was clarified in 2014 through molecular sequencing (18S and 28S rRNA) that integrated it into the family Necopinatidae. These integrative redescriptions not only resolve uncertainty but also enhance reproducibility, as advocated in ongoing efforts to refine global species lists.^[37]^[36]

Omission from Phylogenetic Studies

Taxa are often designated as incertae sedis when they are excluded from phylogenetic studies due to the rarity of available specimens, which limits the collection of sufficient data for analysis.^[38] For instance, in cases involving rare or extinct species preserved only in historical museum collections, the scarcity of material hampers efforts to generate molecular sequences or detailed morphological comparisons needed for inclusion in cladistic analyses.^[39] Similarly, logistical barriers such as remote field locations, ethical restrictions on sampling endangered species, or challenges in accessing deep-sea or polar environments contribute to these omissions, leaving many taxa unexamined in large-scale phylogenies.^[40] The focus on model organisms in phylogenetic research further exacerbates this issue, as studies frequently prioritize well-studied, easily accessible species that yield robust datasets, sidelining less tractable groups.^[41] This selective inclusion results in "orphan" taxa—lineages positioned outside resolved cladograms—creating gaps in comprehensive trees of life.^[42] In the Open Tree of Life project, for example, numerous incertae sedis taxa in bony fishes remain unplaced due to incomplete sampling in synthetic phylogenies, highlighting how such omissions perpetuate taxonomic instability.^[43] These exclusions have significant impacts on classification, as orphan taxa cannot be reliably integrated into hierarchical frameworks, often requiring provisional placements that do not reflect evolutionary relationships.^[44] In protist phylogenetics, many orphan lineages previously labeled incertae sedis have been identified as major eukaryotic supergroups only after targeted inclusion, underscoring the distortion caused by methodological gaps.^[45] Since 2010, advancements in genomics have driven a notable trend toward greater inclusion of previously omitted taxa, reducing the overall rate of incertae sedis designations through culture-free sequencing and phylogenomic approaches.^[46] For example, genome-scale analyses have resolved positions for symbiotic fungi and other understudied lineages once excluded due to sampling limitations, enabling their integration into broader trees.^[47] This shift, facilitated by declining sequencing costs and improved bioinformatics, has progressively diminished the proportion of orphan taxa in fields like mycology and invertebrate systematics.^[48]

Debates and Controversies

One major type of debate leading to incertae sedis designations involves distinguishing between convergent evolution and true homology in morphological traits, where similar features in distantly related taxa create uncertainty about phylogenetic relationships. For instance, in sponge taxonomy (Porifera: Demospongiae), convergent evolution of spicule shapes has caused incongruences between molecular phylogenies and traditional classifications, resulting in several lineages being placed as incertae sedis until genomic data resolved them.^[49] Similarly, in marsupial evolution, bunodont dental specializations have arisen convergently across unrelated groups, complicating placements and leading to incertae sedis status for fossil taxa like those in the Yalkaparidontidae family during early assessments.^[50] Rival classifications further exacerbate these issues, particularly in diverse groups like mammals, where competing hypotheses on ordinal boundaries persist due to conflicting evidence from morphology, fossils, and genetics. In mammalian taxonomy, early 20th-century debates over groups such as Eutriconodonta—considered crown-Mammalia but with unresolved interrelationships—have maintained incertae sedis placements amid ongoing rival schemes that differ on whether they align closer to therians or other basal clades. These disputes highlight how interpretive differences in evolutionary history can temporarily suspend taxa in uncertain positions pending integrative evidence. Notable cases from mid-20th-century avian taxonomy illustrate the intensity of such controversies, with unresolved positions for families like Opisthocomidae (hoatzin) debated between affinities to galliforms, cuculiforms, or a monotypic order due to mosaic traits suggesting convergence. Erwin Stresemann's 1959 review of avian systematics emphasized these "unsolved problems," noting that radical proposals, such as René Verheyen's quantitative morphological analyses reclassifying gruiforms and anseriforms, clashed with traditional syrinx-based systems, prolonging incertae sedis for taxa like Rostratula amid critiques of unreliable parasitological correlations.^[51] The role of peer review in these debates often extends uncertainty, as rigorous scrutiny of novel classifications—such as those challenging Ratitae monophyly—requires extensive validation, delaying consensus in journals like The Auk.^[52] Outcomes of these controversies frequently involve resolutions through consensus papers that synthesize multidisciplinary data, as seen in modern avian phylogenomics where large-scale molecular studies have clarified positions once held incertae sedis, such as the integration of hoatzin into a basal neoavian clade.^[53] In broader biology, efforts like the 2023 survey on a global species list demonstrate how community-driven consensus mechanisms can settle disputes, reducing incertae sedis designations by establishing shared governance for taxonomic updates.^[54]

Notable Examples and Case Studies

Prominent Taxa

One prominent example of a taxon historically classified as incertae sedis is the genus Hallucigenia, a small, worm-like fossil from Cambrian deposits such as the Burgess Shale, known for its bizarre morphology including paired dorsal spines and lobopodian appendages. Its phylogenetic position remained debated for decades due to interpretive challenges in fossil orientation and anatomy, placing it among early panarthropods of uncertain affinity until cladistic analyses in the 2010s resolved it as a stem-group member of the ecdysozoans, closely related to modern arthropods and velvet worms.^[55] In the phylum Bryozoa, numerous genera and species within the order Cheilostomatida are assigned to incertae sedis categories owing to incomplete morphological data or ambiguous colonial structures that hinder family-level placement.^[56] This includes fossil and recent forms where skeletal features like autozooid arrangements do not align clearly with established families, reflecting ongoing taxonomic revisions in this diverse group of colonial invertebrates; as of 2024, WoRMS lists 23 genera in Cheilostomatida incertae sedis.^[56] The use of incertae sedis spans diverse kingdoms, illustrating its broad application in taxonomy; for instance, in protists, the free-living Meteora sporadica from deep-sea sediments long represented an extraordinary filose amoeboflagellate of uncertain phylogenetic affiliation due to its unique rowing motility and lack of close relatives among known eukaryotic lineages, but phylogenomic analyses in 2024 resolved it as sister to Hemimastigophora in a novel clade basal to Opisthokonta.^[57] Among vertebrates, the Paleozoic fossil Tullimonstrum gregarium, the Tully monster from Illinois, was long considered incertae sedis because its soft-bodied preservation obscured affinities; a 2016 study using synchrotron X-ray computed tomography proposed it as a vertebrate related to modern lampreys, but this has been challenged by subsequent analyses (2017, 2023) favoring an invertebrate affinity, leaving its exact position unresolved.^[58]^[59]^[60] Status updates via cladistics have clarified several such placements since 2000, including some basal dinosaur groups like early theropods whose fragmentary remains were once of uncertain position within Dinosauria but have been integrated into phylogenetic frameworks through matrix-based analyses.

Obelisk (biology)

An obelisk is a microscopic genetic element that consists of a type of infectious agent composed of RNA. Described as "viroid-like elements," obelisks consist of RNA in a circular rod shape without any protein shell coating. Obelisks were identified in 2024 by Ivan Zheludev^[61] and colleagues through computational analysis of vast genetic datasets. Their RNA sequences are entirely novel, and their placement within the tree of life remains uncertain as they do not appear to have a shared ancestry with any other life form, virus, or viroid. Obelisks are currently classified as an enigmatic taxon, forming a distinct phylogenetic group.^[61]

Specific Case Analyses

The genus Opisthopatus, comprising South African velvet worms in the family Peripatopsidae, exemplifies incertae sedis placement arising from morphological conservatism within Onychophora. Historically, intergeneric relationships in Peripatopsidae remained unresolved due to limited morphological disparity among extant species, such as similarities in leg number, skin texture, and reproductive structures, which often required scanning electron microscopy for differentiation and were sometimes confounded by fixation artifacts.^[62] These ambiguities contributed to taxonomic uncertainty, as early classifications relied on gross morphology that failed to distinguish deep phylogenetic signals, leaving Opisthopatus in a provisional position relative to genera like Peripatopsis and Metaperipatus.^[62] Resolution came through phylogenomic analysis in the early 2020s, utilizing transcriptomic data from multiple loci to recover a fully supported phylogeny for Peripatopsidae. This approach, employing nucleotide-based models, placed Opisthopatus as monophyletic within a South African clade, distinct from Chilean lineages, and highlighted Gondwanan vicariance patterns, overcoming challenges like the group's large genomes and low GC content that had hindered prior molecular efforts.^[62] The contributing factors—morphological convergence and sparse sampling—underscore how traditional traits alone inadequately capture evolutionary history in morphologically stagnant lineages. In botanical nomenclature under the International Code of Nomenclature for algae, fungi, and plants (ICN), certain green algal groups, particularly prasinophytes, were designated incertae sedis during the 1990s amid debates over fossil versus extant interpretations. Classic morphological systems, informed by fossil records of simple flagellated forms, positioned prasinophytes as a basal or separate class bridging Chlorophyta and Streptophyta, but this clashed with ultrastructural and limited molecular data from extant taxa suggesting paraphyly within core Chlorophyta. For instance, genera like Pycnococcus and Mantoniella exhibited variable thallus organization and flagellar traits that aligned ambiguously with fossil evidence of early algal diversification, fueling controversy over whether such groups represented primitive relics or derived forms. Molecular phylogenies based on SSU rDNA and rbcL sequences in the late 1990s and early 2000s resolved these uncertainties by demonstrating prasinophytes as polyphyletic, with multiple independent lineages nested within Chlorophyta, thus invalidating fossil-driven monophyly hypotheses. This shift emphasized the limitations of fossil morphology, often preserved incompletely, in reconstructing extant relationships, and prompted reclassification under ICN to reflect clade-based groupings rather than form-based ones. These cases illustrate key best practices in addressing incertae sedis status: prioritizing multi-omics integration to complement morphology, as seen in the transition from 1990s rDNA analyses resolving algal debates to 2020s transcriptomics clarifying velvet worm phylogenies and 2024 phylogenomics for protists like Meteora sporadica. Timelines reveal a pattern of prolonged uncertainty—decades for Opisthopatus from morphological descriptions in the early [20th century](/page/20th century) to 2021 resolution, and for prasinophytes from 1980s-1990s fossil-morphology conflicts to post-2000 molecular consensus—highlighting the need for ongoing genomic sampling to preempt nomenclatural instability and enhance predictive classification.^[62]^[57]

Conceptual Distinctions and Implications

Phylogenetic vs. Nomenclatural Uncertainty

Phylogenetic uncertainty in taxonomy refers to ambiguities in the evolutionary relationships among taxa, often arising from incomplete phylogenetic trees, limited sampling of genetic or morphological data, or conflicting signals in analyses. This type of uncertainty is commonly quantified using bootstrap resampling methods, where values below 70% typically indicate weak support for specific branches or clades, reflecting potential instability in inferred relationships. For instance, in molecular phylogenetics, low bootstrap support may stem from rapid evolutionary radiations or insufficient sequence data, leading to placements labeled as incertae sedis until further evidence clarifies the position. Such uncertainty is inherently tied to biological processes and can be mitigated through the accumulation of additional empirical data, such as expanded genomic datasets or fossil records, which enhance resolution in tree-building algorithms like maximum likelihood or Bayesian inference. In contrast, nomenclatural uncertainty pertains to issues in the formal naming of taxa under codes like the International Code of Zoological Nomenclature (ICZN) or the International Code of Nomenclature for algae, fungi, and plants (ICN), independent of underlying biology. It often results from non-compliance with rules, such as the absence of a designated type specimen, homonymy (duplicate names), or publication without adequate description, rendering names invalid (e.g., nomina nuda or junior synonyms). This form of uncertainty does not imply doubt about a taxon's evolutionary affinities but rather challenges to its legitimacy as a stable identifier, potentially placing it in incertae sedis for nomenclatural reasons alone. Resolution typically involves administrative actions, such as rulings by nomenclature commissions (e.g., ICZN opinions) or emendations to establish priority and validity, ensuring consistency without altering phylogenetic interpretations. The primary distinction lies in their foundations and remedies: phylogenetic uncertainty is empirical and data-driven, evolving with scientific advances to refine evolutionary hypotheses, whereas nomenclatural uncertainty is procedural and rule-bound, addressed through governance to maintain nomenclatural stability. While they are largely orthogonal—nomenclature serving as a neutral framework for taxonomy—overlap occurs in cases where inadequate original descriptions contribute to both invalid names and unresolved placements, complicating incertae sedis designations.

Impacts on Modern Classification

In major taxonomic databases such as the Global Biodiversity Information Facility (GBIF) and the Integrated Taxonomic Information System (ITIS), incertae sedis taxa are typically managed as provisional or unclassified entries to reflect their uncertain phylogenetic placement while maintaining data accessibility. In GBIF, these taxa appear as dedicated ranks or placeholders, such as "incertae sedis" under higher-level categories like Biota, allowing for indexing of occurrence records but often resulting in incomplete metadata that complicates cross-database queries and species distribution modeling.^[63]^[64] Similarly, ITIS incorporates incertae sedis status through taxonomic comments or unaccepted name fields, where experts note unresolved affinities, though this can lead to search challenges as users must navigate synonyms or provisional hierarchies without resolved lineages.^[65] These approaches ensure that uncertain taxa remain visible in biodiversity informatics pipelines but highlight persistent issues in data integration, such as fragmented records that hinder automated analyses.^[66] The presence of incertae sedis designations significantly impedes biodiversity assessments by introducing uncertainty into metrics like species richness, endemism, and conservation status evaluations. Taxonomic uncertainty propagates errors in diversification rate estimates and trend analyses, potentially underestimating or overestimating biodiversity hotspots and threatening accurate monitoring of global change impacts.^[67]^[68] For instance, unresolved placements can conflate trends across protected and invasive species in ecological models, reducing the reliability of policy-relevant assessments.^[69] This uncertainty has spurred increased funding for taxonomic revisions, particularly following the adoption of the Aichi Biodiversity Targets in 2010, with Target 19 emphasizing investments in taxonomy to enhance knowledge transfer and application by 2020.^[70] Efforts under the UN's Strategic Plan for Biodiversity (2011-2020) and the subsequent Kunming-Montreal Global Biodiversity Framework (adopted 2022) have channeled resources into global taxonomy initiatives, including through Targets 20 and 21, driving revisions to support sustainable development goals and reduce knowledge gaps in underrepresented taxa. The Kunming-Montreal Framework's Targets 20 (scientific research and monitoring) and 21 (accessible knowledge and data) further prioritize scientific knowledge and data accessibility, sustaining investments in taxonomy to resolve uncertainties like incertae sedis placements.^[71]^[72] Looking ahead, advancements in artificial intelligence (AI) and big data are poised to mitigate incertae sedis challenges by automating phylogenetic inference and resolving ambiguous classifications at scale. Machine learning models, such as random forest classifiers trained on protein family profiles, have demonstrated high accuracy (up to 99.6% at ordinal levels) in assigning giant virus genomes to taxonomic groups, offering a blueprint for broader application in eukaryotic and prokaryotic lineages.^[73] Similarly, hierarchical models leveraging large datasets like UNITE for fungal internal transcribed spacer sequences enable rapid curation and placement of uncertain taxa, potentially accelerating revisions in understudied groups.^[74] Big data integration from sources like metagenomic surveys further supports this by providing empirical evidence to test hypotheses, fostering a shift toward dynamic, evidence-based taxonomies that could substantially decrease the proportion of incertae sedis entries over time.^[75]

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International Code of Zoological Nomenclature
See under name. incertae sedis. A Latin term meaning "of uncertain taxonomic position". index (pl. indexes), n. A list arranged in a particular order ...
[20]
International Code of Nomenclature for algae, fungi, and plants
The International Code of Nomenclature for algae, fungi, and plants is the set of rules and recommendations that govern the scientific naming of all organismsHow to cite the Code · Provision 4 nomenclature... · Art. 17. Orders and subordersMissing: Incertae sedis
[21]
[PDF] International Code of Zoological Nomenclature - Bio-Nica.info
incertae sedis. A Latin term meaning "of uncertain taxonomic position". index (pl. indexes), n. A list arranged in a particular order (usually alphabetical) ...
[22]
https://www.iapt-taxon.org/nomen/main.php?page=art3
[23]
Article 11
The provisions of Art. 11 determine priority between different names applicable to the same taxon; they do not concern homonymy.Missing: incertae sedis
[24]
The International Code of Nomenclature for algae, fungi, and plants
The congress debated 447 proposals to amend the Code and accepted important new rules, including a mechanism for voluntary registration of plant and algal names ...
[25]
Fossil plants from Mississippi - Palaeontologia Electronica
We examine fossil plant reproductive material from two localities, the Bovay and Bolden clay pits, in northern Mississippi, USA.
[26]
updated classification of Orchidaceae - Oxford Academic
Another genus, Devogelia, takes the place of Thaia as incertae sedis in Epidendroideae. Gastrodieae are clearly placed among the tribes in the neottioid ...Missing: examples | Show results with:examples
[27]
Taxonomy of Cyrtochilum-alliance (Orchidaceae) in the light of ...
Jan 13, 2017 · Incertae sedis. Oncidium loxense Lindl. is similar to Cyrtochilum Kunth with which it shares a similar habit, creeping rhizome, long ...
[28]
Shenzhen Code (2018)
This Code aims at the provision of a stable method of naming taxonomic groups, avoiding and rejecting the use of names that may cause error or ambiguity.
[29]
Fungal nomenclature evolving: changes adopted by the 19th ...
This article summarizes the key changes in the rules relating to the nomenclature of fungi made at the XIX International Botanical Congress in Shenzhen, ...
[30]
The Code Online | International Commission on Zoological ...
See under name. incertae sedis. A Latin term meaning "of uncertain taxonomic ... The provisions of this Code supersede those of the previous editions ...
[31]
Article 11 - International Code of Zoological Nomenclature
See under name. incertae sedis. A Latin term meaning "of uncertain taxonomic position". index (pl. indexes), n. A list arranged in a particular order ...
[32]
Article 40. Synonymy of the type genus
See under name. incertae sedis. A Latin term meaning "of uncertain taxonomic position". index (pl. indexes), n. A list arranged in a particular order ...
[33]
Codex Eurypterida: A Revised Taxonomy Based on Concordant ...
Jun 11, 2025 · Eurypterids, also known as sea scorpions, were aquatic chelicerate arthropods that were important components of Paleozoic marine and ...
[34]
[PDF] Is fossil insect taxonomy compatible with database-based research?
Jun 27, 2025 · We submit that the field must recognize that a specimen identified to, at some level, incertae sedis, can be as valuable to science as those ...
[35]
[PDF] a decade of experience in the World Register of Marine Species
Dec 28, 2017 · We 1) explain the categories of names included; 2) list standard reasons used to explain why a name is considered 'unaccepted' or 'uncertain'; 3 ...Missing: provisional | Show results with:provisional
[36]
Best practices for the naming of species - PMC - NIH
Nov 14, 2024 · Duobrachium sparksae (incertae sedis Ctenophora Tentaculata Cydippida): a new genus and species of benthopelagic ctenophore seen at 3,910 m ...
[37]
Towards a global list of accepted species V. The devil is in the detail
Oct 22, 2021 · Such species could be included on the list as incertae sedis within a higher-rank name, or they could be provisionally included within the ...
[38]
Integrative taxonomy elucidates phylogenetic position of a clawless ...
Oct 3, 2025 · Thus, the clawless genus Apodibius was classified as incertae sedis until 2014, when the first DNA sequences of Apo. confusus were published, ...
[39]
DNA barcodes from over-a-century-old type specimens shed light on ...
Jul 17, 2024 · DNA barcodes from over-a-century-old type specimens shed light on the taxonomy of a group of rare ... incertae sedis until further evidence to ...
[40]
Historical DNA phylogenomics clarifies the systematics of rare dung ...
Dec 31, 2024 · The resulting phylogeny propounded the reclassification of Onychothecus (previously incertae sedis) within the tribe Coprini. Our approach ...
[41]
Species evolution: cryptic species and phenotypic noise with a ...
Feb 3, 2025 · Cryptic species are common and can be found in any kingdom of the Tree of Life (McGuire, 2018). However, cryptic plant species are comparatively ...
[42]
Problematica old and new - PMC - PubMed Central - NIH
We address Problematica in the context of pre-cladistic phylogenetics, numerical morphological cladistics and molecular phylogenetics, and focus on general ...
[43]
Proposal for a revised classification of the Demospongiae (Porifera)
Merliida incertae sedis — but in the SP it was included in the ...
[44]
The Tree of Life and a New Classification of Bony Fishes - PMC
Apr 18, 2013 · It is likely to include involuntary errors and omissions in addition to the many unexamined, sedis mutabilis, and incertae sedis taxa.
[45]
Synthesis of phylogeny and taxonomy into a comprehensive tree of life
2.8 has 2,722,024 OTUs without descendants and includes 382,564 higher taxa; 585,081 of the names are classified as nonphylogenetic units (e.g., incertae sedis) ...
[46]
New Phylogenomic Analysis of the Enigmatic Phylum Telonemia ...
Interestingly, phylogenomics suggested that many of these “orphan” taxa are themselves supergroup-level lineages, likely representing early diverging ...
[47]
New Phylogenomic Analysis of the Enigmatic Phylum Telonemia ...
Jan 22, 2019 · With the maturation of culture-free genomic solutions, we expect that inaccuracy due to poor taxon sampling will continue to reduce in ...New Phylogenomic Analysis Of... · Results · Materials And Methods<|separator|>
[48]
Genome-level analyses resolve an ancient lineage of symbiotic ...
Dec 5, 2022 · We augmented this sampling by targeting several other fungal lineages treated as “incertae sedis” or previously undersampled in genome-scale ...
[49]
Genomics-guided refinement of butterfly taxonomy - PMC
Genome-scale DNA analysis opens a new dimension in exploration of butterfly taxonomy and offers a promise of more objective and internally consistent ...
[50]
Molecular Phylogeny of the Astrophorida (Porifera, Demospongiaep ...
Incongruences between our molecular tree and the current classification can be explained by the banality of convergent evolution and secondary loss in spicule ...<|control11|><|separator|>
[51]
A Bizarre New Family of Marsupialia (Incertae sedis) from the Early ...
Jul 1, 2008 · In the absence of more nearly complete specimens that might clarify its relationships, we refer Numbigilga to Marsupialia incertae sedis. We ...
[52]
Mammal classification - Wikipedia
†Eutriconodonta - incertae sedis. Triconodonts appear to be a member of the crown-Mammalia clade, but their relationships within it are unknown. It is also ...
[53]
[PDF] The Status of Avian Systematics and Its Unsolved Problems
groups "incertae sedis," those elements from Wetmore's structure which they removed as incongruous. However, like Lowe, they thought that that would "tell ...
[54]
In the 21st century, taxonomic decisions in herpetology are ...
Aug 6, 2025 · In the 21st century, taxonomic decisions in herpetology are acceptable only when supported by a body of evidence and published via peer review.
[55]
Higher‐order phylogeny of modern birds (Theropoda, Aves ...
Jan 24, 2007 · Avian systematics in the late 20th century has been marked by a trough in morphological phylogenetics (Fautin & Watling, 1999; Jenner, 2004a) ...
[56]
Widespread support for a global species list with a formal ... - PNAS
Oct 30, 2023 · The survey suggests that consensus on a mechanism to create, manage, and govern a single widely accepted list of all the world's species is achievable.
[57]
World Register of Marine Species - Cheilostomatida incertae sedis
Apr 19, 2010 · Cheilostomatida incertae sedis ; Animalia (Kingdom) ; Bryozoa (Phylum) ; Gymnolaemata (Class) ; Cheilostomatida (Order) ; Cheilostomatida incertae ...
[58]
https://www.nature.com/articles/nature16992
[59]
incertae sedis - GBIF
incertae sedis. Dataset; GBIF Backbone Taxonomy: Rank; KINGDOM. Classification. kingdom; incertae sedis. Name. Homonyms: incertae sedis · What is GBIF?Missing: ITIS | Show results with:ITIS
[60]
How Does Sharing Data from Research Institutions on Global ...
It should be noted that a large group of specimens remains classified as incertae sedis (351,083), awaiting further classification.
[61]
ITIS.gov | Integrated Taxonomic Information System (ITIS)
### Summary of ITIS Handling of Incertae Sedis or Taxa of Uncertain Placement
[62]
13.5 DNA derived data | The OBIS Manual
For unknown sequences it is required to populate the scientificName field with “Biota incertae sedis”, or the lowest taxonomic information if available. For ...
[63]
Global Patterns of Taxonomic Uncertainty and its Impacts on ...
Feb 15, 2025 · Our study demonstrates that taxonomic uncertainty can significantly impact diversification rate estimates, thereby affecting our understanding ...
[64]
Revealing uncertainty in the status of biodiversity change - Nature
Mar 27, 2024 · This emphasizes how little is known about biodiversity change across vast spatial and taxonomic scales. Despite this uncertainty at vast scales, ...
[65]
Revealing hidden sources of uncertainty in biodiversity trend ...
Mar 6, 2025 · Aggregation to higher taxonomic ranks was a significant source of uncertainty, leading to conflation of trends among protected and invasive ...
[66]
[PDF] Quick guide to the - Aichi Biodiversity Targets
Reaching this target will require substantial investment in global and national biodiversity observation networks, implementation of the Global Taxonomy ...
[67]
Aichi Biodiversity Targets - Convention on Biological Diversity
Sep 18, 2020 · Strategic Goal A: Address the underlying causes of biodiversity loss by mainstreaming biodiversity across government and society
[68]
Automated classification of giant virus genomes using a random ...
Mar 8, 2024 · A machine learning-based approach to predict the taxonomic classification of novel giant virus MAGs based on profiles of protein family content.Missing: projections | Show results with:projections
[69]
HFTC: a hierarchical fungal taxonomic classification model for ITS ...
Oct 3, 2025 · First, to mitigate the effects of data imbalance and noise, we rigorously curated the UNITE database to construct a high-quality ITS dataset.
[70]
The Sordariomycetes: an expanding resource with Big Data for ...
In this review, we examine recently published studies and data on the Sordariomycetes that have contributed novel insights to the field of fungal evolution.
[71]
Viroid-like colonists of human microbiomes
Peer-reviewed article by Tom O. Delmont et al. in Cell, describing the discovery and characteristics of obelisks as viroid-like RNA elements in human microbiomes, published November 2024.