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Paratype of Lepidothrix vilasboasi (Sick, 1959) in Museum für Naturkunde, Berlin
Paratype of Cadurcotherium nouleti – MHNT

In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Often there is more than one paratype. Paratypes are usually held in museum research collections.

The exact meaning of the term paratype when it is used in zoology is not the same as the meaning when it is used in botany. In both cases however, this term is used in conjunction with holotype.

Zoology

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In zoological nomenclature, a paratype is officially defined as "Each specimen of a type series other than the holotype."[1]

In turn, this definition relies on the definition of a "type series". A type series is the material (specimens of organisms) that was cited in the original publication of the new species or subspecies, and was not excluded from being type material by the author (this exclusion can be implicit, e.g., if an author mentions "paratypes" and then subsequently mentions "other material examined", the latter are not included in the type series), nor referred to as a variant, or only dubiously included in the taxon (e.g., a statement such as "I have before me a specimen which agrees in most respects with the remainder of the type series, though it may yet prove to be distinct" would exclude this specimen from the type series).

Thus, in a type series of five specimens, if one is the holotype, the other four will be paratypes.

A paratype may originate from a different locality than the holotype. A paratype cannot become a lectotype, though it is eligible (and often desirable) for designation as a neotype.

The International Code of Zoological Nomenclature (ICZN) has not always required a type specimen, but any species or subspecies newly described after the end of 1999 must have a designated holotype or syntypes.

A related term is allotype, a term that indicates a specimen that exemplifies the opposite sex of the holotype,[1] and is almost without exception designated in the original description, and, accordingly, part of the type series, and thus a paratype; in such cases, it is functionally no different from any other paratype. It has no nomenclatural standing whatsoever, and although the practice of designating an allotype is recognized by the ICZN, it is not a "name-bearing type" and there are no formal rules controlling how one is designated. Apart from species exhibiting strong sexual dimorphism, relatively few authors take the trouble to designate such a specimen. It is not uncommon for an allotype to be a member of an entirely different species from the holotype, because of an incorrect association by the original author.

Botany

[edit]

In botanical nomenclature, a paratype is a specimen cited in the original description that may not have been said to be a type. It is not the holotype nor an isotype (duplicate of the holotype).

  • If no types were specified, then all specimens cited are syntypes.
  • If more than one specimen was cited as the type, then they are all syntypes, but specimens cited but not listed as types are paratypes.[2] (Articles 9.5 and 9.6).

Like other types, a paratype may be specified for taxa at the rank of family or below (Article 7).[2]

A paratype may be designated as a lectotype if no holotype, isotype, syntype, or isosyntype (duplicate of a syntype) is extant (Article 9.12).[2]

See also

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References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Paratype

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A paratype is a specimen cited in the original description, or protologue, of a new species or infraspecific taxon in biological nomenclature, but it is neither the holotype (the single name-bearing specimen) nor one of the syntypes (multiple name-bearing specimens when no holotype is designated).[1] Unlike name-bearing types, paratypes do not serve to fix the scientific name of the taxon but provide supplementary material to demonstrate morphological variation and diagnostic characters.[2] In zoological nomenclature, the International Code of Zoological Nomenclature (ICZN) defines the type series as the specimens cited in the original publication to define the taxon, excluding any expressly excluded or those identified as variants or doubtful; paratypes are the specimens in this type series other than the holotype.[3] They lack name-bearing status and cannot be selected as lectotypes if the holotype is lost, but may be selected as neotypes when they meet the relevant criteria.[3] Similarly, in botanical nomenclature under the International Code of Nomenclature for algae, fungi, and plants (ICN), paratypes are additional cited specimens in the protologue that support the taxon's circumscription without nomenclatural priority.[1] Paratypes play a crucial role in ensuring the stability and precision of taxonomic classifications by offering multiple reference points for future researchers to verify species boundaries and evolutionary relationships.[1] These specimens are typically preserved and deposited in recognized scientific institutions, such as museums or herbaria, where they facilitate ongoing studies in systematics, biodiversity assessment, and phylogenetic analysis.[2] The designation of paratypes dates back to the late 19th century, with the term first appearing in zoological literature around 1893 to standardize descriptive practices.[4]

Fundamental Concepts

Definition of Paratype

In biological nomenclature, a paratype is any specimen from the original type series of a species or subspecies, excluding the holotype, that supports the description and diagnosis of the taxon. The type series comprises the collection of specimens on which the original description is based, with paratypes serving as additional elements in this series but without the status of the holotype, which is the single primary name-bearing specimen.[3][5] Paratypes are typically duplicates or additional examples derived from the same population or closely related populations as the holotype, providing supplementary morphological or genetic evidence to characterize the taxon while lacking independent nomenclatural priority. Unlike name-bearing types, paratypes do not fix the application of the name if the primary type is lost or ambiguous, though they may inform subsequent type designations such as neotypes.[6] The term "paratype" first appeared in zoological literature in 1893, as part of early efforts to standardize type specimen practices in taxonomy.[4]

Role in Biological Nomenclature

Paratypes play a crucial role in maintaining taxonomic stability by allowing subsequent researchers to examine additional specimens from the original type series, thereby verifying the morphological and other characteristics described in the initial species diagnosis and minimizing ambiguities in identification.[7] This supplementary evidence helps anchor the species concept over time, as a single holotype may not fully capture intraspecific variation, and paratypes provide a broader empirical foundation for confirming boundaries without altering the name-bearing function.[8] In species descriptions, paratypes contribute essential data on morphological, genetic, or ecological traits that complement the holotype, particularly for populations exhibiting variability such as geographic or ontogenetic differences.[1] By offering multiple exemplars, they enable more robust diagnoses, facilitating accurate comparisons in future taxonomic revisions and reducing the risk of misinterpretation based solely on one specimen.[7] Paratypes deposited in natural history collections are invaluable for ongoing research and conservation efforts, supporting applications like DNA barcoding to resolve phylogenetic relationships or mapping species distributions for biodiversity assessments.[9] These specimens enable non-destructive analyses that inform ecological studies and priority-setting in conservation, as they preserve historical context for evaluating population trends and threats.[10] Despite their utility, paratypes serve only a supplementary role and do not bear the scientific name if the holotype is lost or destroyed, unlike neotypes, which can be designated from paratypes to stabilize nomenclature if the holotype is lost. Lectotypes are designated from syntype series when no holotype was originally specified.[3][11] This limitation underscores their supportive rather than definitive status in taxonomic practice.[8]

Zoological Nomenclature

Provisions in the ICZN

In the International Code of Zoological Nomenclature (ICZN), paratypes are defined as specimens that belong to the type series of a nominal species-group taxon but are neither the holotype nor syntypes, nor part of a type series from which a lectotype or neotype has been designated.[3] This definition is outlined in Article 72.4.5, which specifies that when a holotype is designated under Article 73.1, the remaining specimens in the type series automatically become paratypes.[3] The type series encompasses all specimens upon which the author of a new nominal species-group taxon has based the taxon, excluding any expressly excluded by that author; for works published after 1999, all such specimens must be explicitly specified in the original publication.[3] Paratypes thus form part of this series but exclude any specimens merely mentioned in the description without inclusion in the type series.[3] Paratypes possess no name-bearing function, meaning they do not serve to fix the name of the taxon, unlike holotypes or syntypes; however, they are crucial for the validity of the original description as supporting material.[3] The loss of all paratypes does not affect the nomenclatural validity of the taxon, provided a holotype or syntypes exist to bear the name.[3] Paratypes are automatically designated as the remaining specimens in the type series when a holotype is fixed, though authors typically list them explicitly in the original publication with details such as locality, date, collector, and repository. While ideally consisting of specimens of the named species, the type series may include variants, immature stages, or other material if cited in the original description, per ICZN provisions.[3] The formalization of paratype roles in the ICZN stems from the fourth edition published in 1999, which introduced mandatory designation of a holotype or syntypes for new taxa, thereby clarifying the status of remaining type series specimens as paratypes, building on provisions from prior editions.[12]

Designation and Usage

In zoological nomenclature, paratypes are designated by including them in the type series in the original description alongside the holotype, with explicit details on their collection data and deposition to support the taxon's diagnosis and intraspecific variation. These specimens are typically sourced from multiple individuals or localities to provide a representative sample of the species' morphology, geography, and ecology, and they must be deposited in recognized public institutions such as natural history museums to ensure accessibility. As per ICZN Recommendation 16C, paratypes should be preserved in collections available for study by other researchers.[3][13] Paratypes play a crucial role in practice by enabling verification of diagnostic characters and facilitating integrative taxonomic studies, including morphological comparisons and molecular analyses to delineate species boundaries. Deposited in museums, they serve as reference material for future taxonomists to assess new collections against established types, promoting nomenclatural stability. For example, in entomology, paratypes of insects like butterflies in the order Lepidoptera often include specimens from diverse sites to capture wing pattern variations essential for distinguishing cryptic species. In ichthyology, paratypes of fish such as Miocene sperm whales provide insights into skeletal morphology across populations. Similarly, in mammalogy, paratypes of pangolins have supported genomic studies confirming evolutionary relationships.[13][14] Contemporary approaches in zoology emphasize the use of paratypes in molecular taxonomy, where tissue samples or DNA extracts from these specimens aid in phylogenetic analyses and species delimitation. Digital imaging and 3D modeling of paratypes, such as for vertebrate skeletons, enhance global access via online databases like those of the Smithsonian Institution, minimizing physical handling while allowing detailed examination of features like dentition or scale patterns. However, challenges include ensuring non-destructive sampling for DNA and standardizing digital protocols to preserve microscopic details across institutions.[13][15]

Botanical Nomenclature

Provisions in the ICN

In the International Code of Nomenclature for algae, fungi, and plants (ICN; Madrid Code, 2025), paratypes are defined under Article 9.7 as any specimen cited in the protologue—the original publication of the name—that is neither the holotype nor an isotype, nor one of the syntypes when two or more specimens were simultaneously indicated as syntypes in the protologue.[5] This definition ensures that paratypes form part of the original material used to characterize the taxon, distinct from the primary name-bearing elements. The type series under the ICN encompasses all specimens and illustrations cited in the protologue, including the holotype (if designated), isotypes (duplicates of the holotype), syntypes (when no holotype is selected), and paratypes. Paratypes may derive from the same gathering as the holotype or from separate collections, provided they are explicitly cited in the protologue; however, if a holotype is designated, any additional cited specimens are classified as paratypes rather than syntypes, excluding the latter from the type series.[5] Paratypes hold no direct name-bearing function, as the nomenclatural application of the name is fixed solely by the holotype; yet, they support taxonomic stability by allowing selection as a lectotype if the holotype becomes unavailable and no higher-priority material (such as isotypes or syntypes) exists, such as through loss or destruction, in accordance with Articles 9.11 and 9.12. Since the 2012 Melbourne Code, valid publication of new names—particularly for fungi—requires citation of type material details in the protologue, with paratypes often included to fulfill descriptive requirements, though only holotype indication is mandatory under Article 40.1.[5][16][17] Designation of paratypes must occur explicitly in the protologue, where they are labeled as such or simply cited alongside the holotype, ensuring their recognition as part of the original material. For fungi and algae, the provisions mirror those for plants, but include additional mandates for deposition: holotypes and often paratypes must be preserved in recognized herbaria or culture collections to enable verification, as stipulated in Article 40.[16]

Designation and Usage

In botanical taxonomy, paratypes are designated by explicitly citing them in the protologue alongside the holotype, ensuring they represent additional material from the same or comparable populations to illustrate intraspecific variation. These specimens are typically sourced from multiple collections to provide a broader sample of the taxon's morphology and distribution, and they must be clearly labeled as paratypes to comply with nomenclatural standards. As defined in Article 9 of the International Code of Nomenclature for algae, fungi, and plants (ICN), a paratype is any specimen cited in the protologue that is neither the holotype nor an isotype, nor one of the syntypes when multiple specimens are indicated as types simultaneously.[5][18] Paratypes play a crucial role in practice by supporting the verification of morphological traits and enabling molecular analyses to confirm taxonomic boundaries. Deposited in herbaria, they serve as reference material for subsequent researchers to compare against new collections, ensuring stability in species circumscription. For instance, in vascular plants such as orchids, paratypes facilitate the examination of subtle variations in floral structures, like petal shape and coloration, which are essential for distinguishing closely related taxa.[19][2] In mycology, paratypes often include associated documentation such as spore prints, which capture critical features like spore color and ornamentation to aid in species delimitation among morphologically similar fungi. In phycology, paratypes of highly variable algae, such as species in the genus Ulva, assist in delineating ecological niches by providing evidence of habitat-specific adaptations, informing studies on bloom dynamics and environmental responses.[20] Contemporary approaches emphasize digital imaging of paratypes to enable global access through online repositories, reducing wear on physical specimens and broadening collaborative opportunities in taxonomy. However, challenges persist, including the need for high-resolution captures that reveal microscopic details without compromising specimen integrity, as well as logistical issues in standardizing imaging protocols across institutions.[21][22]

Related Concepts and Variations

Allotype and Other Type Specimens

In zoological nomenclature, an allotype is a designated specimen of the sex opposite to that of the holotype, serving as part of the original type series to illustrate sexual dimorphism without possessing any name-bearing function or nomenclatural priority.[3][23] This term, unregulated by the International Code of Zoological Nomenclature (ICZN), is typically selected from the same collection as the holotype to provide comparative morphological data, particularly in species where sexes differ significantly.[24][25] Allotypes are especially prevalent in entomology, where they are often paired with holotypes to document intraspecific variation between males and females, aiding in species identification and taxonomic revisions.[26][13] For instance, in descriptions of new insect species, an allotype may be explicitly labeled to highlight dimorphic traits like antennal structure or wing patterns, though its designation is optional and carries no formal stability under the ICZN.[27] This practice enhances descriptive completeness but is not mandatory, and recommendations emphasize restricting allotypes to specimens from the original type series to maintain contextual integrity.[28] Other type specimens in zoology include syntypes, which are multiple specimens collectively forming the name-bearing type when no holotype has been designated, often from pre-1999 descriptions where holotype selection was not required.[3][24] Lectotypes are subsequently selected from syntypes to designate a single name-bearing specimen, stabilizing nomenclature when original material is ambiguous.[29] Neotypes, meanwhile, replace lost or destroyed holotypes, lectotypes, or syntypes under strict ICZN conditions to preserve nomenclatural continuity.[11] Unlike paratypes, which supplement the holotype as non-name-bearing originals, these types (syntypes, lectotypes, neotypes) directly bear the species name and establish priority, whereas allotypes and paratypes solely support morphological characterization without such authority.[3][6] Both allotypes and paratypes lack nomenclatural standing, meaning they do not fix the application of a name and can be numerous or even absent without invalidating a description; their primary value lies in bolstering taxonomic evidence through additional exemplars.[23][6] In botanical nomenclature under the International Code of Nomenclature for algae, fungi, and plants (ICN), no equivalent to the allotype exists, as the code focuses on holotypes, syntypes, lectotypes, and neotypes without provisions for sex-specific designations, reflecting differences in how dimorphism is typically addressed in plants.[30]

Applications in Other Disciplines

In microbiology, under the International Code of Nomenclature of Prokaryotes (ICNP), paratypes refer to additional strains beyond the type strain (equivalent to a holotype) that are cited in the original description to support the taxon's characterization, particularly for bacteria and archaea.[31] These paratype strains serve as supplementary material for phenotypic, genotypic, and chemotaxonomic validation, often deposited in culture collections for accessibility. For instance, in descriptions of Mycobacterium species, paratype strains like NCTC 10411 have been used to confirm enzymatic activities and morphological traits distinguishing the taxon from related species.[31] Similarly, for Escherichia coli strains, paratypes contribute to genomic comparisons, enabling validation of phylogenetic placement and functional annotations through sequencing of multiple isolates.[32] In paleontology, paratypes are integrated into nomenclature following the International Code of Zoological Nomenclature (ICZN) for fossil animals or the International Code of Nomenclature for algae, fungi, and plants (ICN) for plant fossils, where they supplement the holotype by providing additional skeletal elements or morphological details essential for taxonomic diagnosis.[3] These specimens aid in reconstructing incomplete holotypes and assessing intraspecific variation in extinct taxa. A notable example is the titanosaurian sauropod Dreadnoughtus schrani, where three paratypes (MPEF-PV 3399, 3401, 3402) consist of partial skeletons that corroborate the holotype's features, such as vertebral morphology, facilitating a more robust understanding of the species' size and anatomy in Late Cretaceous Patagonia.[33] Mycology, governed by the ICN, employs paratypes for fungal taxa, where they include additional cultures or herbarium specimens cited in the protologue to illustrate variability in spore morphology, colony growth, or molecular markers.[34] In virology, paratype usage is rare due to the International Committee on Taxonomy of Viruses (ICTV) framework, which emphasizes sequence-based demarcation rather than physical specimens; however, "type strains" or reference isolates function analogously to paratypes by providing supplementary genomic data for species delineation.[35] Paratype series in microbiology often comprise fewer specimens than in macroorganismal taxonomy, reflecting challenges in culturing viable strains and the emphasis on accessible type strains under ICNP rules. In bioinformatics, digital paratypes—such as high-resolution images, 3D models, or DNA sequences from physical specimens—enable virtual access and analysis, enhancing taxonomic stability without physical transfer; for example, online exsiccates serve as digital paratypes for plant and fungal taxa, supporting remote morphological and phylogenetic studies.[36]

References

  1. About Type Specimens in FLAS - Florida Museum of Natural History
  2. Type specimens - Purdue University - College of Agriculture
  3. Article 72. General provisions
  4. https://www.oed.com/dictionary/paratype_n?tl=true
  5. Article 9 - International Association for Plant Taxonomy (IAPT)
  6. [PDF] A Compendium of Zoological Type Nomenclature: a Reference Source
  7. International Code of Zoological Nomenclature
  8. epitype - International Code of Botanical Nomenclature
  9. Annotation of Type Specimens: Recommendations
  10. Art. 9 - International Code of Nomenclature for algae, fungi, and plants
  11. A taxonomic assessment of novel and remarkable fungal species in ...
  12. Envisaging a global infrastructure to exploit the potential of digitised ...
  13. The effectiveness and limitations of digital images for taxonomic ...
  14. Article 40
  15. International Code of Nomenclature for algae, fungi, and plants
  16. International Code of Nomenclature for algae, fungi, and plants
  17. [PDF] International Code of Zoological Nomenclature - Bio-Nica.info
  18. Types in International Code of Zoological Nomenclature
  19. (PDF) Allotypes should be from the type series: a position paper for ...
  20. [PDF] Allotypes should be from the type series - Smithsonian Institution
  21. How to describe a new species in zoology and avoid mistakes
  22. Type Collection - UBC Zoology
  23. 74.5. Lectotype designations before 2000
  24. Article 75. Neotypes - International Code of Zoological Nomenclature
  25. <i>Mycobacterium rhodesiae</i> sp. nov. - Wiley Online Library
  26. Phylogenetic and Chemotaxonomic Studies Confirm the Affinities of ...
  27. A Gigantic, Exceptionally Complete Titanosaurian Sauropod ...
  28. Art. 9 - International Code of Nomenclature for algae, fungi, and plants
  29. TAXONOMY, CLASSIFICATION AND NOMENCLATURE OF VIRUSES
  30. (PDF) Eleven new species of Begonia L. (Begoniaceae) from Laos ...
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