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Paratype

A paratype is a specimen cited in the original description, or protologue, of a new or infraspecific in biological , but it is neither the (the single name-bearing specimen) nor one of the syntypes (multiple name-bearing specimens when no holotype is designated). Unlike name-bearing types, paratypes do not serve to fix the scientific name of the but provide supplementary material to demonstrate morphological variation and diagnostic characters. In zoological nomenclature, the (ICZN) defines the type series as the specimens cited in the original publication to define the , excluding any expressly excluded or those identified as variants or doubtful; paratypes are the specimens in this type series other than the . 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. Similarly, in under the International Code of Nomenclature for algae, fungi, and plants (ICN), paratypes are additional cited specimens in the protologue that support the 's circumscription without nomenclatural priority. 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. These specimens are typically preserved and deposited in recognized scientific institutions, such as museums or , where they facilitate ongoing studies in , assessment, and phylogenetic analysis. The designation of paratypes dates back to the late , with the term first appearing in zoological literature around 1893 to standardize descriptive practices.

Fundamental Concepts

Definition of Paratype

In biological , a paratype is any specimen from the original type series of a or , excluding the , that supports the and diagnosis of the taxon. The type series comprises the collection of specimens on which the original is based, with paratypes serving as additional elements in this series but without the status of the , which is the single primary name-bearing specimen. Paratypes are typically duplicates or additional examples derived from the same population or closely related populations as the , providing supplementary morphological or genetic evidence to characterize the 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. The term "paratype" first appeared in zoological literature in , as part of early efforts to standardize type specimen practices in .

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 diagnosis and minimizing ambiguities in identification. This supplementary evidence helps anchor the concept over time, as a single may not fully capture intraspecific variation, and paratypes provide a broader empirical foundation for confirming boundaries without altering the name-bearing function. In species descriptions, paratypes contribute essential data on morphological, genetic, or ecological traits that complement the , particularly for populations exhibiting variability such as geographic or ontogenetic differences. 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. Paratypes deposited in collections are invaluable for ongoing research and efforts, supporting applications like to resolve phylogenetic relationships or mapping species distributions for biodiversity assessments. These specimens enable non-destructive analyses that inform ecological studies and priority-setting in , as they preserve historical context for evaluating trends and threats. Despite their utility, paratypes serve only a supplementary role and do not bear the scientific name if the is lost or destroyed, unlike neotypes, which can be designated from paratypes to stabilize if the is lost. Lectotypes are designated from syntype series when no was originally specified. This limitation underscores their supportive rather than definitive status in taxonomic practice.

Zoological Nomenclature

Provisions in the ICZN

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

Designation and Usage

In zoological , paratypes are designated by including them in the type series in the original description alongside the , with explicit details on their collection data and deposition to support the taxon's and intraspecific variation. These specimens are typically sourced from multiple individuals or localities to provide a representative sample of the ' morphology, , and , and they must be deposited in recognized public institutions such as museums to ensure accessibility. As per ICZN Recommendation 16C, paratypes should be preserved in collections available for study by other researchers. 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 , paratypes of insects like butterflies in the order often include specimens from diverse sites to capture wing pattern variations essential for distinguishing cryptic species. In , paratypes of such as Miocene sperm whales provide insights into skeletal morphology across populations. Similarly, in , paratypes of pangolins have supported genomic studies confirming evolutionary relationships. Contemporary approaches in emphasize the use of paratypes in molecular , where samples or extracts from these specimens aid in phylogenetic analyses and delimitation. Digital and of paratypes, such as for vertebrate skeletons, enhance global access via online databases like those of the , minimizing physical handling while allowing detailed examination of features like or scale patterns. However, challenges include ensuring non-destructive sampling for and standardizing digital protocols to preserve microscopic details across institutions.

Botanical Nomenclature

Provisions in the ICN

In the International Code of Nomenclature for , fungi, and (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 nor an isotype, nor one of the syntypes when two or more specimens were simultaneously indicated as syntypes in the protologue. This definition ensures that paratypes form part of the original material used to characterize the , distinct from the primary name-bearing elements. The type series under the ICN encompasses all specimens and illustrations cited in the protologue, including the (if designated), isotypes (duplicates of the ), syntypes (when no is selected), and paratypes. Paratypes may derive from the same gathering as the or from separate collections, provided they are explicitly cited in the protologue; however, if a is designated, any additional cited specimens are classified as paratypes rather than syntypes, excluding the latter from the type series. Paratypes hold no direct name-bearing function, as the nomenclatural application of the name is fixed solely by the ; yet, they support taxonomic stability by allowing selection as a lectotype if the becomes unavailable and no higher-priority (such as isotypes or syntypes) exists, such as through or destruction, in accordance with Articles 9.11 and 9.12. Since the 2012 Code, valid publication of new names—particularly for fungi—requires citation of type details in the protologue, with paratypes often included to fulfill descriptive requirements, though only indication is mandatory under Article 40.1. Designation of paratypes must occur explicitly in the protologue, where they are labeled as such or simply cited alongside the , ensuring their recognition as part of the original material. For fungi and , the provisions mirror those for , but include additional mandates for deposition: and often paratypes must be preserved in recognized herbaria or culture collections to enable verification, as stipulated in Article 40.

Designation and Usage

In botanical taxonomy, paratypes are designated by explicitly citing them in the protologue alongside the , 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 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 (ICN), a paratype is any specimen cited in the protologue that is neither the nor an isotype, nor one of the syntypes when multiple specimens are indicated as types simultaneously. Paratypes play a crucial role in practice by supporting the verification of morphological traits and enabling molecular analyses to confirm taxonomic boundaries. Deposited in , 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 shape and coloration, which are essential for distinguishing closely related taxa. 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. Contemporary approaches emphasize of paratypes to enable global access through online repositories, reducing wear on physical specimens and broadening collaborative opportunities in . 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 protocols across institutions.

Related Concepts and Variations

Allotype and Other Type Specimens

In zoological , an allotype is a designated specimen of the opposite to that of the , serving as part of the original type series to illustrate without possessing any name-bearing function or nomenclatural priority. This term, unregulated by the (ICZN), is typically selected from the same collection as the to provide comparative morphological data, particularly in where sexes differ significantly. Allotypes are especially prevalent in , where they are often paired with holotypes to document intraspecific variation between males and females, aiding in identification and taxonomic revisions. For instance, in descriptions of new insect , 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. 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. Other type specimens in include syntypes, which are multiple specimens collectively forming the name-bearing type when no has been designated, often from pre-1999 descriptions where selection was not required. Lectotypes are subsequently selected from syntypes to designate a single name-bearing specimen, stabilizing when original material is ambiguous. Neotypes, meanwhile, replace lost or destroyed s, lectotypes, or syntypes under strict ICZN conditions to preserve nomenclatural continuity. 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. 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 ; their primary lies in bolstering taxonomic evidence through additional exemplars. In under the International Code of Nomenclature for , 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.

Applications in Other Disciplines

In , under the International Code of Nomenclature of Prokaryotes (ICNP), paratypes refer to additional strains beyond the type strain (equivalent to a ) that are cited in the original description to support the 's characterization, particularly for and . 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 species, paratype strains like NCTC 10411 have been used to confirm enzymatic activities and morphological traits distinguishing the from related species. Similarly, for strains, paratypes contribute to genomic comparisons, enabling validation of phylogenetic placement and functional annotations through sequencing of multiple isolates. In , paratypes are integrated into nomenclature following the (ICZN) for fossil animals or the International Code of Nomenclature for algae, fungi, and plants (ICN) for plant fossils, where they supplement the by providing additional skeletal elements or morphological details essential for taxonomic . 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 . Mycology, governed by the ICN, employs paratypes for fungal taxa, where they include additional cultures or specimens cited in the protologue to illustrate variability in morphology, colony growth, or molecular markers. In , paratype usage is rare due to the International Committee on of Viruses (ICTV) framework, which emphasizes sequence-based demarcation rather than physical specimens; however, "type strains" or isolates function analogously to paratypes by providing supplementary genomic data for species delineation. 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.

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