Replacement names for junior homonyms in ants (Hymenoptera, Formicidae)

81 Replacement names for junior homonyms in ants (Hymenoptera, Formicidae) Brian L. Fisher1 1 Department of Entomology, California Academy of Sciences, San Francisco, California, USA Corresponding author: Brian L. Fisher ([email protected]) Copyright: © Brian L. Fisher. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Replacement names are proposed for 40 unresolved junior species group homonyms in Formicidae (Hymenoptera) as required by ICZN Articles 23 and 60. This effort aims to enhance taxonomic stability and facilitate digital management for ant names. In each case, the junior homonym is considered acceptable and in use. To avoid the creation of unnecessary replacement names, names were not proposed for unresolved junior homonyms in cases where 1) the status of the junior homonym is a junior or senior synonym or unidentifiable; 2) the primary homonym pair have not been regarded as congeneric since 1899; or 3) the congeneric placement of the junior secondary homonym is unreliable. The following new names are introduced to replace existing homonyms: Acromyrmex aurina nom. nov., Camponotus chucki nom. nov., Camponotus dorex nom. nov., Camponotus durnyx nom. nov., Camponotites flarex nom. nov., Camponotus robecchii clyne nom. nov., Camponotus substitutus cylrix nom. nov., Colobopsis breva nom. nov., Crematogaster scutellaris gynia nom. nov., Dolichoderus thoracicus jexina nom. nov., Ectomomyrmex pylixy nom. nov., Ectomomyrmex pylor nom. nov., Euponera tynara nom. nov., Formica lavateri liora nom. nov., Formica mytara nom. nov., Hypoponera rylin nom. nov., Hypoponera veltan nom. nov., Lepisiota vynia nom. nov., Temnothorax flavispinus novira nom. nov., Messor minor nubina nom. nov., Messor nuvia nom. nov., Pheidole arnoldi qindra nom. nov., Pseudoneoponera sublaevis quorra nom. nov., Strumigenys wylixa nom. nov., Camponotus foleyi cybira nom. nov., Camponotus maculatus brexyl nom. nov., Camponotus nixra nom. nov., Camponotus planus brina nom. nov., Carebara dazia nom. nov., Carebara xynera nom. nov., Colobopsis vitrea yevira nom. nov., Formica draven nom. nov., Messor draxil nom. nov., Pheidole singaporensis zolara nom. nov., Pseudolasius zynia nom. nov., Tetramorium drunex nom. nov., Tetramorium fenix nom. nov., Tetramorium flinex nom. nov., Temnothorax fynor nom. nov., and Tetraponera vantix nom. nov. In addition, Estevia nom. nov. is proposed for the preoccupied genus Wilsonia Hong, 2002. Additional taxonomic changes are presented, including the reinstatement of Tetramorium schultzei Forel, 1910a stat. nov. and the following new combinations: Bothroponera jonesii (Forel, 1891), comb. nov.; Diacamma unicolor (Smith, 1860a), comb. nov.; Pseudoneoponera vidua (Smith, 1857), and Diacamma solitaria (Smith, 1860b), comb. nov. Key words: AntCat, Formicidae, homonymy, ICZN, nomenclature, nomen novum, replacement names, taxonomy Academic editor: Sebastian Salata Received: 19 June 2025 Accepted: 3 September 2025 Published: 16 October 2025 ZooBank: https://zoobank. org/022D48B6-D029-470F-907D96134B37CE80 Citation: Fisher BL (2025) Replacement names for junior homonyms in ants (Hymenoptera, Formicidae). ZooKeys 1256: 81–113. https://doi.org/10.3897/ zookeys.1256.162607 ZooKeys 1256: 81–113 (2025) DOI: 10.3897/zookeys.1256.162607 82 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Introduction A primary purpose of zoological nomenclature is to ensure that each taxon at every rank has a unique and stable name. In accordance with the International Code of Zoological Nomenclature (ICZN 1999), homonyms — identical names for different taxa at the same rank, whether within the species group, genus group, or family group — must be resolved to maintain this stability. When two taxa share the same name, the later (junior) homonym must either be replaced by an available synonym or, if no appropriate synonym exists, by a new replacement name (nomen novum) (ICZN Article 60). Homonymy is particularly problematic in ants (Formicidae), a group with a long history of taxonomic revisions and a high diversity of described species. Currently, 16,881 valid species and subspecies of ants are recognized worldwide (AntCat 2025), reflecting more than 250 years of continuous taxonomic work beginning with Linnaeus (1758). The recurrent use of common descriptive epithets (e.g., niger, rufus, elongatus), compounded by limited historical access to prior literature, has led to the creation of multiple junior homonyms. Additionally, generic boundaries that have shifted over time have resulted in secondary homonyms as species are moved between genera. The consequences of unresolved homonymy go beyond nomenclatural confusion. Unresolved homonymy can lead to misinterpretations in biodiversity data integration and conservation planning, especially in the era of big data (Morrison et al. 2009; Patterson et al. 2010; Sánchez-González 2020). This review seeks to provide clarity by examining ICZN guidelines and illustrating their application through historical and modern examples within Formicidae. In doing so, I hope to provide guidelines and historical context for ant taxonomists. Overall, this work takes a practical approach that emphasizes avoiding unnecessary nomenclatural changes while ensuring that valid taxa are properly named and recognized. Simplified overview of ICZN rules for homonymy The ICZN Code outlines strict rules to address homonymy. Two main categories are distinguished. Primary homonymy occurs when two taxa are originally described under the same genus using the same species-group name. For example, if two species were named “Formica rufa” by different authors, the latter-named one is a junior primary homonym and must be replaced. According to Article 57.2, junior primary homonyms are permanently invalid and cannot be used, regardless of later taxonomic changes. Secondary homonymy arises when species described in different genera are later placed in the same genus, resulting in identical combinations. In such cases, the junior name becomes invalid under Article 59.1, but only if the taxa are considered congeneric. If subsequent revisions treat the species as belonging to separate genera and they have not been regarded as congeneric since 1899, the junior name may be maintained (Article 23.9.5). A replacement name (nomen novum) should only be proposed if the junior homonym is an accepted name in use and no available synonym exists (ICZN Articles 23.3.5, 60.3); this name will then compete for priority with any synonym recognized later. If a junior homonym was already replaced by a nomen novum 83 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) prior to 1961 and that replacement name is still in use, the replacement name retains priority, even if the act was unnecessary (ICZN Article 59.3). These provisions are intended to prevent the proliferation of superfluous names while promoting clarity and stability in nomenclature. Practical considerations Addressing homonymy requires careful attention to historical taxonomy, synonymy, and the ICZN rules. Before introducing a new name, taxonomists should undertake a thorough review of the nomenclatural history of both the senior and junior homonyms. This includes verifying type specimens (AntWeb 2025), consulting original descriptions, and reviewing synonymies. Modern tools such as AntCat.org (https://antcat.org/) and ZooBank (https:// zoobank.org/) provide critical resources for checking name availability and synonym status. However, even with digital tools, homonym detection and resolution remain challenging, particularly in groups with extensive historical literature. As emphasized by Gardner and Hayssen (2004), replacement names should be introduced with precision and only when truly necessary. Thorough historical research and careful verification of original sources are required to avoid introducing unnecessary new names, which can further complicate taxonomic databases and scientific communication. When in doubt, authors should err on the side of nomenclatural stability and avoid coining new names if an existing synonym or classification solution is available. Neubauer et al. (2014) illustrate this restrained approach in proposing replacement of homonyms within European freshwater gastropods. Those authors documented six cases of species-group homonyms that were unambiguously considered junior synonyms, and thus they chose not to introduce new names for those taxa. This approach follows the spirit of stability. If a name is not needed (because the taxon is not valid or in use), introducing a new name would only add clutter. As those authors noted, the Code has no explicit instruction for disused homonyms, but “following the intent expressed in Article 23.9.5, which seems to discourage the proposal of unnecessary replacement names” they refrained from naming new replacements. In fact, the Code explicitly provides that such cases may be referred to the Commission for a ruling under its plenary power, and that until such a ruling is made, prevailing usage of both names should be maintained (ICZN 1999: Art. 82). This nomenclatural review establishes criteria for proposing replacement names for homonyms, with an emphasis on stability and alignment with the ICZN Code. Replacement names are proposed only for junior homonyms (primary or secondary) that are accepted, currently in use, and lack available synonyms. For primary homonyms, a replacement is introduced only when the taxa have been regarded as congeneric since 1899 (Art. 23.9.5). For secondary homonyms, a replacement is proposed only when the taxon occupies a stable generic placement supported by molecular or strong taxonomic evidence, consistent with the Code’s emphasis on nomenclatural stability (Art. 23.9.5). Although all junior primary homonyms are technically invalid under the Code, replacement names are introduced only for those that are currently accepted taxa, not for names that are junior synonyms or otherwise not in use. For example, Suppl. material 1 lists 155 unresolved junior primary homonyms among ant taxa that are treated as synonyms or are taxonomically indeterminate; for 84 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) these, no replacement names are introduced, since doing so would not contribute to stability. In all cases, a complete historical review of taxonomic combinations is essential before any replacement is made. Replacement names proposed in this work were selected to ensure uniqueness and compliance with the ICZN, with priority given to names that are easy to use and phonetically distinct. While some replacement names in taxonomy preserve etymological intent, reference morphology, geography, or honor individuals, such semantic continuity is not required by the Code, and the primary goal in this work was to provide clear, stable, and unambiguous names. Type specimen, authorship, and date Stability in nomenclature depends not only on name priority but also on accurately linking type specimens to accepted species-group names. Even in the absence of homonymy, this linkage can be complicated by the lack of standardized unique specimen identifiers in older taxonomic works. Homonyms exacerbate this problem, as illustrated below. A nomen novum (replacement name) is attributed to the author who proposes it and dated to that publication. The original author of the junior homonym is not credited as the author of the replacement name. However, the nomen novum inherits the name-bearing type specimen of the replaced junior homonym. Thus, although the authorship is new, the type specimen of the nomen novum is that of the junior homonym it replaces. As a result, the nomen novum becomes a senior objective synonym of the replaced name, since it is based on the same type (ICZN Article 60.1). A change in type is permitted only under exceptional circumstances, such as when the original type is inadequate or non-diagnostic following Recommendation 60A (e.g., as described in Article 75.5 or, in the case of a genus-group homonym, the type species is poorly defined). If the junior homonym is replaced not with a nomen novum but with an existing synonym, then the type specimen(s) of that synonym becomes the name-bearing type(s) of the accepted taxon. Nevertheless, the type specimen of the junior homonym remains important, as synonymy is reversible. Should taxonomic revisions restore the junior homonym to validity, its type specimen will again carry nomenclatural weight. This creates a rare scenario where two distinct type specimens, those of the accepted synonym and of the suppressed homonym, must be tracked and curated. To reduce future instances of homonymy and their attendant complexities, Formicidae taxonomists are strongly encouraged to propose species-group names that are unique across the family. While the ICZN permits identical species epithets in different genera, the practical burden of resolving homonyms is considerable for modern taxonomic databases and digital biodiversity platforms. Avoiding homonyms from the outset enhances long-term nomenclatural clarity and database integrity. Case studies in Formicidae Within Formicidae, 97 instances were identified where previously proposed replacement names are now unnecessary. In most cases, these names were rendered superfluous when a secondary junior homonym was no longer con- 85 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) sidered congeneric with its senior homonym, thus eliminating the homonymy. In other cases, replacement names were unnecessary because an available synonym already existed and could serve as the valid name. Finally, several new names were introduced due to oversight, even though valid replacement names had already been established. The following case studies illustrate common pitfalls and resolutions in the management of homonymy within ants. By highlighting these patterns, I aim to reduce the future creation of homonyms and to guide best practices when addressing unresolved cases, particularly by avoiding unnecessary nomina nova. Each case is presented in the context of the relevant ICZN Articles, underscoring the importance of a careful, context-aware taxonomic approach. All names in the case studies are provided in both their original combination (protonym) and, where applicable, their current valid combination. Temporary nature of secondary homonymy Secondary homonyms may be temporary due to shifting genus boundaries. Proposing replacement names in such cases has often proved premature. It is preferable not to propose a replacement name until generic limits stabilize. This is especially relevant in ants, where molecular phylogenetics and, more recently, phylogenomic datasets (e.g., Ward et al. 2015; Branstetter et al. 2017; Blaimer et al. 2018; Doré et al. 2025) have led to substantial redefinitions of genera and higher-level relationships. While these studies have greatly improved our understanding of evolutionary lineages, they also highlight the fluidity of generic boundaries in some groups. Until phylogenomic frameworks are more fully integrated and stabilized across all Formicidae genera, restraint in proposing replacement names for secondary homonyms is warranted. Case 1 Micromyrma pygmaea Dufour, 1857: 61 (current combination Tapinoma pygmaeum) was combined in Tapinoma in 1859. Formica pygmaea Latreille, 1798: 45 was combined in Tapinoma in 1852 and then Plagiolepis in 1861. Tapinoma dufouri Donisthorpe, 1943: 662 was proposed as a replacement name for Tapinoma pygmaeum Dufour, 1857: 61, but is unnecessary as Tapinoma pygmaeum Dufour, 1857: 61 was no longer a secondary junior homonym. Case 2 Meanwhile, Micromyrma dufourii Perris, 1878: 382 (current combination Plagiolepis dufourii) was combined in Tapinoma in 1983 but moved to Plagiolepis in 1925. But that did not stop Tapinoma confusum Smith, 1951: 837 being proposed as a replacement name for Tapinoma dufouri Donisthorpe, 1943: 662. Case 3 When described, Ponera apicalis Smith, 1857: 66 (current combination Ectomomyrmex apicalis) was a junior secondary homonym of Formica apicalis 86 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Latreille, 1802: 204 (current combination Neoponera apicalis) which was combined in Ponera by Latreille, 1809: 128. However, Mayr (1863: 439) combined the taxa in Pachycondyla in 1863. Thus, Ponera apicalis Smith, 1857: 66 was a junior secondary homonym in Ponera from 1857–1863. By the time Ectomomyrmex terminalis Smith, 1871: 321 was proposed as a replacement name, the name was no longer needed. Ectomomyrmex terminalis Smith, 1871: 321 is an unnecessary replacement name and an objective synonym of Ponera apicalis Smith, 1857: 66. For Ponera apicalis Smith, 1857: 66, the secondary homonym story continues. Ponera apicalis Smith, 1857: 66 was combined in Pachycondyla by Brown in 1995 (Brown 1995: 302), joining Formica apicalis Latreille, 1802: 204. Thus, once again, Ponera apicalis Smith, 1857: 66 was a junior secondary homonym. In 2010, homonym hunter Özdikmen created the replacement name Ectomomyrmex terminalis Özdikmen, 2010: 993 for Ponera apicalis Smith, 1857: 66. However, in 2014, both Ponera apicalis Smith, 1857: 66 and Formica apicalis Latreille, 1802: 204 left Pachycondyla for different genera (Schmidt and Shattuck 2014). This shift made Ectomomyrmex terminalis Özdikmen, 2010: 993 an unnecessary name in 2014 and an objective synonym of Ponera apicalis Smith, 1857: 66. This sequence illustrates the risk of proposing new names based on transient classifications. Another issue is Özdikmen’s (2010) use of the same species epithet proposed by Smith, 1871. Replacement names, even when unnecessary, still need to be recorded and managed, the proposal of an identical name to replace a homonym does not help. Identical names should never be proposed. Case 4 Wheeleria santschii Forel, 1905: 171 (current combination Monomorium santschii). In 1906, Forel (1906: 64) noted that the genus-group name Wheeleria was already in use and thus a junior homonym of Wheeleria Tutt, 1905: 37 (Lepidoptera). Forel was correct to provide a genus replacement name Santschia Forel, 1906: 65. However, Forel also unnecessarily proposed a replacement name for the species Santschia intrudens Forel, 1906: 65 [perhaps to avoid having the genus and species names to be the nearly identical Santschia santschii (Forel 1905: 171).] However, the replacement name is unnecessary, and thus, Monomorium intrudens Forel, 1906: 65, is now an objective junior synonym of Monomorium santschii Forel, 1905: 171. In addition, the proposed name is a junior secondary homonym of Monomorium intrudens Smith, 1874: 406. Because there is an existing, valid synonym for Monomorium intrudens Forel, 1906: 65, the replacement name for Monomorium intrudens Forel, 1906: 65 is Monomorium santschii Forel, 1905: 171. Junior secondary homonym replaced before 1961 If a junior secondary homonym was replaced before 1961, and the substitute name remains in use, it retains priority under ICZN Article 59.3. The following case illustrates this problem: 87 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Syllophopsis jonesi Arnold, 1952: 465 was proposed as a replacement name for Syllophopsis arnoldi Santschi, 1921: 120; a junior secondary homonym of Monomorium arnoldi Forel, 1913a: 137. The replacement name Syllophopsis jonesi Arnold, 1952: 465 was proposed before 1961 (when both arnoldi taxa were included in Monomorium). Article 59.3 stipulates that valid pre-1961 replacement names retain priority, even if the original homonymy is later resolved. In 2015, Ward et al. combined arnoldi Santschi, 1921: 120 in Syllophopsis and mistakenly proposed that jonesi was an unnecessary replacement name. The priority awarded by Ward et al. (2015: 73) is incorrect and Syllophopsis jonesi Arnold, 1952: 465 has priority. Complete taxonomic histories required Confusion also arises when earlier replacements are overlooked; the following case illustrates this problem. Anochetus jacobsoni Menozzi, 1939: 178 is a junior primary homonym of Anochetus jacobsoni Forel, 1911b: 193. Anochetus menozzii Donisthorpe, 1941b: 237 was proposed as a replacement name. This new name was overlooked and a second and unnecessary replacement name was proposed: Anochetus ineditus Baroni Urbani, 1971: 360. Use of available synonyms If an available synonym exists for the junior secondary homonym, the synonym should be used instead of proposing a new name. However, a replacement name has priority if proposed before a junior synonym. Case 1 Ponera simillima Smith, 1860b: 105 (current combination Prionopelta simillima) is a junior primary homonym of Leptogenys simillima Smith, 1860b: 104 (original combination: Ponera simillima). [The same species epithet was used one page later to describe a different species]. Prionopelta nominata Smith, 1871: 323 was proposed as a replacement name for the junior homonym but this name was overlooked, and a second replacement name was proposed, Prionopelta poultoni Donisthorpe, 1932: 462. Subsequently Brown (1953: 12) synonymized three species, including Prionopelta majuscula Emery, 1897: 595 with Ponera simillima Smith, 1860b: 105, and thought the oldest synonym Prionopelta majuscula Emery, 1897: 595 would take priority as a replacement name for Ponera simillima Smith, 1860b: 105. However, nom. nov. proposed before the synonymy take priority, and thus Prionopelta nominata Smith, 1871: 323, nom. restit. has priority as the replacement name. Case 2 Here is another example of an overlooked replacement name and the use of a synonym that occurred after the nomen novum replacement name. All species are currently in Anochetus, but I use original combinations to help keep the story straight. 88 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Odontomachus tyrannicus Smith, 1861: 44 is a junior primary homonym of Odontomachus tyrannicus Smith, 1859: 144. As with the Ponera simillima example above, we have name duplication by the same author within the same genus. The situation is further complicated: Smith had shared specimens with colleagues and labeled them “gladiator” (see Brown 1978: 574, suggesting that gladiator was the intended species epithet for Odontomachus tyrannicus Smith, 1861: 44). In 1862, Mayr included one of the specimens labeled gladiator in the description of a new genus for the taxon and used gladiator as the species name, providing an unintentional but formal description of the new species Stenomyrmex gladiator Mayr, 1862: 712. In 1863, Roger proposed Odontomachus smithii Roger, 1863: 21 as a replacement name for Odontomachus tyrannicus Smith, 1861: 44. In 1871, Odontomachus gladiator Smith, 1871: 320 was proposed as a replacement name. In 1893, Stenomyrmex gladiator Mayr, 1862: 712 was proposed as a junior synonym of Odontomachus tyrannicus Smith, 1861: 44 by Dalla Torre 1893: 48. In 1932, Odontomachus gladiator Donisthorpe, 1932: 467 was proposed as a replacement name for Odontomachus tyrannicus Smith, 1861: 44. Brown (1978: 556) mistakenly prioritizes Stenomyrmex gladiator Mayr, 1862: 712 as the replacement name, although Odontomachus smithii Roger, 1863: 21 nom. restit. actually has priority. Results In the homonym lists below, collections are referred to by the following acronyms. When the type depository is not known and not given in the original description, it is recorded here as “unspecified”. AMNH American Museum of Natural History, New York, USA; ANIC Australian National Insect Collection, Canberra, Australia; ATPC Alberto Tinaut personal collection, Universidad de Granada, Granada, Spain; CASC California Academy of Sciences, San Francisco, USA; DEIB Deutsches Entomologisches Institut, Berlin-Dahlem, Germany; HNHM Hungarian Natural History Museum, Budapest, Hungary; IEUB Istituto di Entomologia “Guido Grandi”, Università degli Studi di Bologna, Bologna, Italy; LACM Los Angeles County Museum, Los Angeles, USA; LKCNHM Lee Kong Chian Natural History Museum, Singapore; MCZC Museum of Comparative Zoology, Harvard University, Cambridge, USA; MHNG Muséum d’histoire naturelle, Geneva; MNHN Muséum National d’Histoire Naturelle, Paris; MNHU Museum für Naturkunde an der Humboldt-Universität zu Berlin (Zoologisches Museum), Berlin, Germany; MSNG Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy; MVMA Museum of Victoria, Melbourne, Victoria, Australia; NHMB Naturhistorisches Museum Basel, Basel, Switzerland; NHMUK British Museum of Natural History, now Natural History Museum London, London, England; 89 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) NHMW Naturhistorisches Museum Wien, Vienna, Austria; NUYC Laboratory of Insect Specimens, Agricultural College, Ningxia University, Yinchuan, China; OXUM Oxford University Museum of Natural History, Oxford, England; SAMC Iziko Museums of Cape Town (= South African Museum), Cape Town, South Africa; SIZK Schmalhausen Institute of Zoology, Kyiv, Ukraine; SMNG Senckenberg Museum für Naturkunde (Staatliches Museum für Naturkunde), Görlitz, Germany; SWFU Faculty of Conservation Biology, Southwest Forestry College [University], Kunming, Yunnan, China; WUIK Wonkwang University, Division of Biological Sciences, Iksan (= Iri City), North Cholla Province, Republic of Korea. A. Genus-group name homonyms Estevia Fisher, nom. nov. †Wilsonia Hong, 2002: 608 (Insecta, Hymenoptera). Preoccupied by Wilsonia Bonaparte, 1838: 23 (Aves). Type species. †Wilsonia megagastrosa Hong, 2002: 608, by original designation. Etymology. The genus name is proposed in honor of Flávia Esteves, a Brazilian myrmecologist and taxonomist, recognized for her significant contributions to the study of ant morphology, diversity, and systematics. Gender: feminine. Estevia megagastrosa Hong, 2002, comb. nov. †Wilsonia megagastrosa Hong, 2002: 608, figs 565–573 (w.) China (Liaoning, Fushun Amber, Eocene). B. Primary homonyms Below are the primary homonyms that satisfy the following criteria for the creation of a new replacement name (nomen novum): the junior primary homonym is an accepted taxon, that is, in use as a valid species. In one case, the senior homonym is not valid (a junior synonym of another species) but a new name is still proposed since the junior homonym is in use. Replacement names are not established when the junior primary homonym is a disused junior synonym (and already treated as a synonym), when the junior primary homonym’s status is highly doubtful, nor when the two homonyms have not been congeneric for over 100 years. All such long-separated homonym pairs are listed in Section C (ICZN Article 23.9.5), where no replacement names are introduced and prevailing usage is maintained. Homonyms are presented first with the original combination and then the current combination is given if different than the original combination. Homonyms are listed by the current combination of the junior primary homonym. 96 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) mary homonym of Formica cordata Holl, 1829: 140 (combination in Pheidole by Mayr (1868: 17), Boudinot et al. (2024: 148), current combination Pheidole cordata). Type locality. Indonesia: Aru Islands. Type material. Syntype workers. Type depository. OXUM. Note. Primary homonyms have not been congeneric since 1868. C3. No replacement name for Formica nigerrima Nylander, 1856: 71 (combination in Tapinoma by Mayr (1861: 41), current combination Tapinoma nigerrimum), junior primary homonym of Formica nigerrima Christ, 1791: 513 (combination in Lasius by Emery (1892a: 162), current combination Lasius nigerrimus). Type locality. France: Prades-le-Lez. Type material. Neotype worker. Type depository. SMNG. Note. Primary homonyms have not been congeneric since 1861. C4. No replacement name for Formica nitida Smith, 1859: 138 (combination in Camponotus by Roger (1863: 4), current combination Camponotus nitidus), junior primary homonym of Formica nitida Razoumowsky, 1789: 300. Type locality. Indonesia: Aru. Type material. Holotype worker. Type depository. OXUM. Note. Primary homonyms have not been congeneric since 1863. C5. No replacement name for Formica pallida Smith, 1857: 57 (combination in Camponotus by Mayr (1862: 656), current combination Camponotus irritans pallidus), junior primary homonym of Formica pallida Latreille, 1798: 41 (combination in Lasius by Emery (1893: 183), current combination Lasius pallida), junior synonym of Lasius brunneus (Latreille, 1798: 41). Type locality. Malaysia: Borneo, Sarawak. Type material. Syntype workers. Type depository. NHMUK. Note. Primary homonyms have not been congeneric since 1862. C6. No replacement name for Formica timida Jerdon, 1851: 122 (combination in Camponotus by Roger (1863: 3), current combination Camponotus timidus), junior primary homonym of Formica timida Foerster, 1850: 35 (combination in Lasius by Nylander (1856: 68), current combination Lasius timida), junior synonym of Lasius brunneus (Latreille, 1798: 41). Type locality. India: Malabar coast. Type material. Syntype workers, syntype queen. Type depository. Unspecified. Note. Primary homonyms have not been congeneric since 1862. C7. No replacement name for Ponera sulcata Mayr, 1867: 441 (combination in Pachycondyla (Bothroponera) by Emery (1901a: 46), current combination Bothroponera sulcata), unresolved junior primary homonym of Ponera sulcata Smith, 1858: 99 (combination in Ectatomma (Gnamptogenys) by Emery (1893: 26), current combination Gnamptogenys sulcata). 97 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Type locality. India. [Emery, 1911: 78 gives India as type locality, possibly based on the collection data in Forel 1900b: 326, and Bingham 1903: 99.] Type material. Syntype workers. Type depository. NHMW. Current subspecies (nominal subspecies plus the following): B. sulcata fossulata (Forel, 1900b: 323) B. sulcata sulcatotesserinoda (Forel, 1900b: 323). Note. Primary homonyms have not been congeneric since 1893. D. Secondary homonyms Under ICZN Article 59.1, a junior secondary homonym must be replaced if both names are in use and are congeneric. The following cases satisfy the criteria for the creation of a new replacement name (nomen novum): the junior secondary homonym is an accepted taxon, that is, in use as a valid species. Replacement names are not established when the junior secondary homonym is a disused junior synonym (and treated as a synonym already), when the junior secondary homonym’s status is highly doubtful, nor when the secondary homonym generic placement is not certain (see section E for a list of these exceptions). D1. Camponotus chucki Fisher, nom. nov. Note. Replacement name for Camponotus kugleri Mackay, 2025: 453, junior secondary homonym of Camponotus kugleri Ionescu-Hirsch, 2010: 78. Type locality. Colombia: Zona Buenos Aires. Type material. Holotype minor worker. Type depository. MCZC. Etymology. The name honors Charles (“Chuck”) Kugler, myrmecologist and Peace Corps colleague, who collected the holotype and numerous specimens of Camponotus in Colombia. The epithet is treated as a noun in apposition. D2. Camponotus foleyi cybira Fisher, nom. nov. Note. Replacement name for Camponotus (Tanaemyrmex) foleyi subsp. rufescens Bernard, 1953: 190, junior secondary homonym of Dendromyrmex fabricii var. rufescens Wheeler, 1916: 13 (current combination Camponotus fabricii rufescens), junior synonym of Camponotus nidulans (Smith, 1860c: 69). Type locality. Libya: Fezzân, El Abiod. Type material. Syntype major workers. Type depository. MNHN. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D3. Camponotus maculatus brexyl Fisher, nom. nov. Note. Replacement name for Camponotus (Myrmoturba) maculatus r. foveolatus Stitz, 1925: 125, junior secondary homonym of Formica foveolata Mayr, 98 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) 1853: 277 (current combination Camponotus lateralis foveolatus, junior synonym of Camponotus piceus (Leach, 1825: 292). Type locality. Philippines. Type material. Syntype workers. Type depository. MNHU. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D4. Camponotus nixra Fisher, nom. nov. Note. Replacement name for Camponotus (Karavaievia) nigripes Dumpert, 1995: 96, junior secondary homonym of Dendromyrmex nidulans var. nigripes Wheeler, 1916: 13 (current combination Camponotus nidulans nigripes), junior synonym of Camponotus nidulans (Smith, 1860c: 69). Type locality. Malaysia: Perak, Belum. Type material. Holotype worker. Type depository. NHMB. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D5. Camponotus planus brina Fisher, nom. nov. Note. Replacement name for Camponotus (Myrmorhachis) planus var. indefessus Wheeler, 1919: 294, junior secondary homonym of Formica indefessa Sykes, 1835: 104 (current combination Camponotus indefessus), junior synonym of Camponotus compressus (Fabricius, 1787: 307). Type locality. Ecuador: Galapagos Is, Indefatigable I. (= Santa Cruz Is.). Type material. Syntype workers. Type depository. MCZC. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D6. Carebara dazia Fisher, nom. nov. Note. Replacement name for Carebara laeviceps Liu & Zhong, 2024: 17, junior secondary homonym of Oligomyrmex (Hendecatella) capreolus subsp. laeviceps Wheeler, 1928: 24 (current combination Carebara capreola laeviceps). Type locality. China: Sichuan Province, Kaijiang County, Dazhou City. Type material. Holotype worker. Type depository. SWFU. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D7. Carebara xynera Fisher, nom. nov. Note. Replacement name for Aneleus politus Santschi, 1914a: 79 (current combination Carebara polita), junior secondary homonym of Myrmica polita Smith, 1860b: 108 (current combination Carebara polita), junior synonym of Carebara diversa (Jerdon, 1851: 109). 99 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Type locality. Kenya (“Afrique orientale anglaise”): Blue Post Hotel, Kikuyu territory. Type material. Syntype workers. Type depository. NHMB. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D8. Colobopsis vitrea yevira Fisher, nom. nov. Note. Replacement name for Camponotus (Colobopsis) vitreus r. carinata Stitz, 1938: 116 (current combination Colobopsis vitrea carinata), junior secondary homonym of Colobopsis carinata Mayr, 1870: 943, homonym replaced by Colobopsis polynesica (Emery, 1896: 374). Type locality. Papua New Guinea (“German New Guinea”): Aprilfluss. Type material. Syntype queens. Type depository. MNHU. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D9. Formica draven Fisher, nom. nov. Note. Replacement name for Myrmecocystus melliger subsp. californicus Creighton, 1950: 445 (current combination Formica californica), junior secondary homonym of Formica microgyna subsp. californica Wheeler, 1917: 543, junior synonym of Formica adamsi whymperi Wheeler, 1917: 544. Type locality. USA: California, Weed. Type material. Holotype worker. Type depository. LACM. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D10. Messor draxil Fisher, nom. nov. Note. Replacement name for Messor sculpturatus Carpenter, 1930: 32, junior secondary homonym of Cratomyrmex sculpturatus Stitz, 1916: 377 (current combination Messor regalis sculpturatus) junior synonym of Messor regalis (Emery, 1892b: 572). Type locality. USA: Colorado (Florissant Fossil Beds) Eocene. Type material. Fossil. fossil imprints (compression fossils). Holotype ♀ No. 2920, paratypes Nos. 2921 (MCZC), 10,032 (Peabody Museum), 7850 (Princeton University), 17,017a (University of Colorado), 78,804 (USNM), 11 (Carnegie Museum). Type depository. MCZC. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D11. Pheidole singaporensis zolara Fisher, nom. nov. Note. Replacement name for Aphaenogaster (Ischnomyrmex) longipes var. continentis Forel, 1911a: 24 (current combination Pheidole singaporensis continen- 100 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) tis), junior secondary homonym of Pheidole tasmaniensis var. continentis Forel, 1902c: 437. Type locality. Myanmar (“Burma”). Type material. Syntype minor workers. Type depository. MHNG. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D12. Pseudolasius zynia Fisher, nom. nov. Note. Replacement name for Rhizomyrma emeryi Forel, 1915: 347 (current combination Pseudolasius emeryi), junior secondary homonym of Pseudolasius emeryi Forel, 1911c: 286. Type locality. Papua New Guinea: Hansemann Mts. Type material. Holotype worker. Type depository. MHNG. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D13. Tetramorium drunex Fisher, nom. nov. Note. Replacement name for Tetramorium flavum Chang & He, 2001: 2, junior secondary homonym of Xiphomyrmex gambogecus var. flavus Donisthorpe, 1941a: 58 (current combination Tetramorium gambogecum flavum), junior synonym of Tetramorium gambogecum (Donisthorpe, 1941a: 57). Type locality. China: Pengyang, Ningxia. Type material. Holotype worker. Type depository. NUYC. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D14. Tetramorium fenix Fisher, nom. nov. Note. Replacement name for Tetramorium intermedium Hita Garcia et al., 2010: 48, junior secondary homonym of Rhoptromyrmex rothneyi var. intermedia Forel, 1913b: 80 (current combination Tetramorium rothneyi intermedium), junior synonym of Tetramorium wroughtonii (Forel, 1902a: 231). Type locality. Ghana: Aiyeola Forest Reserve, Kade. Type material. Holotype worker. Type depository. NHMUK. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D15. Tetramorium flinex Fisher, nom. nov. Note. Replacement name for Teleutomyrmex kutteri Tinaut, 1990: 202 (current combination Tetramorium kutteri), junior secondary homonym of Tetramorium semilaeve var. kutteri Santschi, 1927a: 57, junior synonym of Tetramorium indocile Santschi, 1927a: 53. 101 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Type locality. Spain: Granada, Sierra Nevada, “Prados de Otero”. Type material. Holotype queen. Type depository. ATPC. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D16. Temnothorax fynor Fisher, nom. nov. Note. Replacement name for Temnothorax similis Csősz et al., 2015: 23, junior secondary homonym of Leptothorax (Macromischa) similis Baroni Urbani, 1978: 501 (current combination Temnothorax similis), junior synonym of Temnothorax laetus (Wheeler, 1937: 456). Type locality. Turkey: Nur Daglari. Type material. Holotype worker. Type depository. HNHM. Etymology. Arbitrary combination of letters, treated as a noun in apposition. D17. Tetraponera vantix Fisher, nom. nov. Note. Replacement name for Tetraponera insularis Ward, 2022: 49, junior secondary homonym of Sima nigra var. insularis Emery, 1901b: 113 (current combination Tetraponera nigra insularis), junior synonym of Tetraponera nigra (Jerdon, 1851: 112). Type locality. Madagascar: Mahajanga, Forêt Anabohazo. Type material. Holotype worker. Type depository. CASC. Etymology. Arbitrary combination of letters, treated as a noun in apposition. E. Junior homonyms In accordance with ICZN Articles 57.3 and 59.1, the following junior secondary homonyms are not replaced. In these cases, either (i) the senior homonym is a junior synonym and no longer valid, thus eliminating the conflict, or (ii) the taxonomic status is unresolved and a replacement would be premature. Replacement names are proposed only when required to resolve homonymy between valid, congeneric names in current use. E1. No replacement name for Monomorium bidentatum Mayr, 1887: 616 (current combination Monomorium bidentatum), junior secondary homonym of Myrmica bidentata Smith, 1858: 124 (current combination Monomorium bidentata). The combination of Monomorium bidentata Smith, 1858 in Monomorium is unconvincing but even if correct, then a junior synonym of Monomorium bidentatum Mayr, 1887 exists (Monomorium bidentatum piceonigrum Borgmeier, 1948: 468). Type locality. Chile: Valdivia. Type material. Syntype workers, syntype queens (numbers not stated). Type depository. NHMW. 102 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) E2. See Suppl. material 1 of 155 species and subspecies where unresolved junior homonyms are not replaced because the status of the junior homonym is unidentifiable (24 species, 2 subspecies) or because the junior homonym is treated as a junior synonym of another valid taxon (55 species, 74 subspecies). F. Supplementary taxonomic notes F1. Reinstatement of Tetramorium schultzei Forel, 1910a, stat. nov. Examination of the holotype of Tetramorium caespitum subsp. schultzei Forel, 1910a: 19 (worker), together with a review of its taxonomic history, indicates that this taxon is not conspecific with T. mossamedense. Consequently, T. schultzei is here reinstated from synonymy and elevated to species rank (stat. nov.). Type locality. Botswana: Kalahari, Kgokong-Kang, no. 966 (L. Schultze). Type material. Holotype worker (AntWeb unique specimen identifier: FoCol2071 https://www.antweb.org/specimen/FoCol2071). Type depository. MNHU. The type was presumed lost by Bolton, 1980: 314; but B. Bolton (pers. comm. 17 April 2025) has noted that the schultzei holotype is not conspecific with mossamedense. Tetramorium schultzei is here reinstated and raised to species rank. Tetramorium schultzei Forel, 1910a, stat. nov. Subspecies of caespitum: Santschi 1914b: 367; Arnold 1917: 331; Wheeler, W.M. 1922a: 894; Emery 1924: 278. Junior synonym of mossamedense: Bolton 1980: 314 (mistake); Bolton 1995: 414 (mistake). F2. New genus combinations from male-based reevaluations. Based on a detailed morphological reexamination of male specimens that remained in Pachycondyla following the treatment of Schmidt and Shattuck (2014), the following new genus combinations are proposed. F2.1 Bothroponera jonesii (Forel, 1891), comb. nov. Lobopelta jonesii Forel, 1891 219 (m.) Madagascar. Type locality. Madagascar: Forêt d’Andrangoloaka (F. Sikora). Type material. Holotype male (AntWeb unique specimen identifier: CASENT0101709). Type depository. MHNG. Combination in Leptogenys by Emery (1911: 102). Combination in Pachycondyla by Bolton (1975: 298). Incertae sedis in Pachycondyla by Schmidt and Shattuck (2014: 155). F2.2 Diacamma unicolor (Smith, 1860a), comb. nov. Ponera unicolor Smith, 1860a 73 (m.) Indonesia (Sulawesi). Type locality. Indonesia: Indonesia: Sulawesi (“Celebes”, Makassar, “Mak.” (A.R. Wallace). 103 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) Type material. Holotype male (apex of gaster missing) (AntWeb unique specimen identifier: CASENT0901368). Type depository. OXUM. Donisthorpe (1932: 458) cites 1 male in OXUM (see CASENT0901368). 2) There is a second male in OXUM, from Tondano (Sulawesi), apparently of the same species and complete (Bolton unpublished notes, 1978). Combination in Pachycondyla (Bothroponera) by Donisthorpe (1932: 458). Combination in Bothroponera by Joma and Mackay (2013: 2). Incertae sedis in Pachycondyla by Schmidt and Shattuck (2014: 155). F2.3 Pseudoneoponera vidua (Smith, 1857), comb. nov. Ponera vidua Smith, 1857 68 (m.) Borneo (East Malaysia: Sarawak). Type locality. Malaysia: Borneo, Sarawak, “SAR.” (A.R. Wallace). Type material. Holotype male (gaster missing) (AntWeb unique specimen identifier: CASENT0901367). Type depository. OXUM. Donisthorpe (1932: 448) cites 1 male in OXUM (see CASENT0901367). Combination in Pachycondyla by Donisthorpe (1932: 448). Incertae sedis in Pachycondyla by Schmidt and Shattuck (2014: 155). F2.4 Diacamma solitaria (Smith, 1860b), comb. nov. Ponera solitaria Smith, 1860b 103 (m.) Indonesia (Bacan). Type locality. Indonesia: Bachian (= Bacan I.), “Bac.” (A.R. Wallace). Type material. Holotype male (head detached, glued separately onto the stage card) (AntWeb unique specimen identifier: CASENT0901366). Type depository. OXUM. Type notes. Donisthorpe, 1932: 461, cites 1 male in OXUM (see CASENT0901366). Combination in Pachycondyla (Bothroponera) by Donisthorpe (1932: 461). Combination in Bothroponera by Wilson (1958: 362); Joma and Mackay (2013: 2). Combination in Pachycondyla by Brown (1995: 309). Incertae sedis in Pachycondyla by Schmidt and Shattuck (2014: 155). Acknowledgements I thank Barry Bolton, Bradley Reynolds, and Michele Esposito for their valuable feedback on earlier drafts of this manuscript. I am also grateful to the many curators and collection managers who provided access to type specimens and historical material, particularly at NHMUK, MNHU, MHNG, CASC, and MCZC. Special thanks to the AntCat and AntWeb community for maintaining the nomenclatural infrastructure that made this review possible. I also thank my colleagues in the global ant taxonomy network for ongoing discussions that continue to illuminate difficult cases in the application of the ICZN Code. This work benefited from the digital resources of AntWeb and from decades 104 ZooKeys 1256: 81–113 (2025), DOI: 10.3897/zookeys.1256.162607 Brian L. Fisher: Replacement names for junior homonyms in Ants (Hymenoptera: Formicidae) of foundational literature and historical insights shared by generations of ant taxonomists. I am grateful to Rodrigo Feitosa and an anonymous reviewer, as well as to the subject editor Sebastian Salata, for their careful reviews of the manuscript, which greatly improved its clarity and presentation. All errors remain my own. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement Not applicable (nomenclatural work). Use of AI AI tools build into MS Word were used for language editing and formatting assistance under the author’s supervision. Funding AntCat.org and AntWeb.org was supported in part by NSF DEB-1932467 (AoW) and NSF DEB-1856400 (CanBe) Author contributions BLF conceived, researched, and wrote the manuscript. Author ORCIDs Brian L. Fisher https://orcid.org/0000-0002-4653-3270 Data availability All of the data that support the findings of this study are available in the main text or Supplementary Information. References AntCat (2025) AntCat: An Online Catalog of the Ants of the World. https://antcat.org/ [accessed 22 April 2025] AntWeb (2025) AntWeb, California Academy of Sciences, San Francisco, California, USA. http://www.antweb.org [accessed 22 April 2025] Arnold G (1917) A monograph of the Formicidae of South Africa. Part III. Myrmicinae. Annals of the South African Museum 14: 271–402. 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Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d’Agricoltura 22: 3–38. Wheeler WM (1937) Ants mostly from the mountains of Cuba. Bulletin of the Museum of Comparative Zoology 81: 439–465. Wilson EO (1958) Studies on the ant fauna of Melanesia. III. Rhytidoponera in western Melanesia and the Moluccas. IV. The tribe Ponerini. Bulletin of the Museum of Comparative Zoology 119: 303–371. Zhang J, Sun B, Zhang X (1994) Miocene insects and spiders from Shanwang, Shandong. [In Chinese.]. Science Press, Beijing, v + 298 pp. Supplementary material 1 The 155 species-group names where unresolved junior homonyms are not replaced because the status of the junior homonym is unidentifiable (24 species, 2 subspecies) or a synonym (55 species, 74 subspecies). Author: Brian L. Fisher Data type: docx Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1256.162607.suppl1