Family-group names for all taxa of earwigs (living and extinct) are listed with dates and sources indicated; in total 85 entries are recorded along with a single entry of dubious taxonomic identity (i.e., Ocelliidae, nomen dubium, a name apparently applied to a fossil earwig nymph of uncertain status and identity). This survey revealed two instances in which currently accepted names must be changed owing to priority by an older name: Platylabiinae and Cosmiellinae must be replaced by Palicinae and Skendylinae, respectively, as the family-group names remain valid despite synonymy of their type genera. The type genus of Verhoeff's Gonolabididae (as Gonolabidae) is Gonolabis, not Gonolabina (which is the type genus of Gonolabininae, Gonolabiinae auctorum) as asserted by various authors. In addition, the generally asserted authorship and dates of numerous names are found to be incorrect and are therefore revised herein. Most notably, the name Carcinophorinae as used by Hincks was a nomen nudum and therefore unavailable (the name was first made available by Popham). The widely employed spellings (all incorrectly formed) of Anataelinae, Challinae, Protolabinae, Anophthalmolabiinae, Titanolabinae, Gonolabinae, Brachylabinae, Isolabinae, Antisolabinae, Parisolabinae, Chaetospanini, Irdexinae, Eudohrninae, Rudraxinae, Gonolabinae, Rhyacolabinae, and Kinellinae are corrected to Anataeliinae, Challiinae, Protolabidinae ( = Echinosomatinae), Anophthalmolabidinae, Titanolabidinae, Gonolabidinae ( = Anisolabidinae), Brachylabidinae, Isolabidinae, Antisolabidinae, Parisolabidinae, Chaetospaniini, Irdicinae ( = Spongophorinae), Eudohrniinae ( = Neolobophorinae), Rudracinae, Gonolabininae, Rhyacolabidinae ( = Ancistrogastrinae), and Kinesinae, respectively. The following nomenclatural changes are proposed: Acrania, reinstated as genus; Epicranopygia, new synonym of Acrania; Acrania angulata, new combination; A. constricta, new combination; A. eximia, reinstated combination; A. fletcheri, new combination; A. picta, reinstated combination; A. triangulata, new combination; A. vittipennis, new combination; Pyge, reinstated as genus; Paracranopygia, new synonym of Pyge; Pyge assamensis, new combination; P. bakeri, new combination; P. burmensis, new combination; P. comata, new combination; P. formosa, new combination; P. maculipes, new combination; P. meghalayana, new combination; P. modesta, reinstated combination; P. pallidipennis, new combination; P. proxima, new combination; P. semenovi, new combination; P. siamensis, new combination; P. similis, new combination; P. tonkinensis, new combination; P. variegata, new combination; P. vicina, new combination; Paradiplatys, reinstated as genus; Lobodiplatys, Heterodiplatys, and Epidiplatys, reduced to subgenera of Paradiplatys (and their included species reinstated in combination with Paradiplatys); Paradiplatys (Lobodiplatys) coriaceus, reinstated combination; P. (Paradiplatys) conradti, reinstated combination; P. (P.) lamottei, reinstated combination; P. (P.) pectinatus, reinstated combination; P. (P.) salvazae, reinstated combination; P. (P.) spinulosus, reinstated combination; P. (Heterodiplatys) bicolor, reinstated combination; P. (H.) bihamatus, reinstated combination; P. (H.) burri, reinstated combination; P. (H.) rotundicollis, reinstated combination; P. (H.) schoutedeni, reinstated combination; P. (Epidiplatys) gladiator, reinstated combination; Cretolabiinae, new subfamily (Anisolabididae); Paratitanolabis, new synonym of Titanolabis; Titanolabis bormansi, new combination; and Titanolabis myanmarensis, new name.
Introduction
Nomenclature is the set of pragmatic principles that stabilize the names employed in biology. Although not often recognized, taxonomy is by its nature a dynamic field of study, particularly in the revolutionized world of cladistic phylogenetics. Rules of nomenclature serve to maximize the continuity in the usage of names and, more importantly, the communication of volumes of data associated with the taxa to which we give these names. All intellectual endeavors related to the biological sciences depend on effective transmittal of information, and vital to this is the correct application of scientific names. While a tremendous amount of effort in Zoology is spent on ensuring the correct usage of species-group and genus-group names, family-group names are often employed with relatively little consideration and, as a result, may frequently be incorrect. Herein we have attempted to catalog for the first time the family-group names for earwigs (Dermaptera), correctly identifying their type genus, combining stem, and their precedence in competitions of nomenclatural priority. Although there have been several, comprehensive overviews of Dermaptera classification to the level of genera and species (e.g., Popham, 1965, 1968a, 1968b, 1968c; Popham and Brindle, 1966a, 1966b, 1966c, 1967a, 1967b, 1968, 1969; Steinmann, 1986, 1989a, 1989b, 1990, 1993), these accounts were found to have numerous errors in the application of family-group names, prompting this catalog and overview. Names above the family group are not regulated in zoological nomenclature and for this reason the numerous ordinal, subordinal, &c. names proposed for earwigs are not considered, although they are briefly summarized. As in any historical endeavor some obscure references may have been missed and earlier name usages may have been overlooked. However, we believe this list to be relatively complete and to correctly indicate for the first time the author and date by which each name became taxonomically available. As such, a few changes to earwig classification owing to priority are noted.
The Principle of Coordination (ICZN, 1999: Art. 36) has, unfortunately, not generally been applied in the Dermaptera, with different authors and dates applied to names based on the same type genus but at different ranks. This principle simply states that once a name has been validly established for a particular type genus, regardless of its rank, it is simultaneously validated at all other ranks in the family group with the same authorship and date. For example, although the family Chelisochidae is frequently attributed to Burr (1907), since he was the first to use this name at the family rank (e.g., Steinmann, 1975, 1989b, 1993; Sakai, 1982; Engel, 2003), the correct authorship is Verhoeff (1902a), since he was the first author to propose a name in the family group based on the type genus Chelisoches. Changes in rank within the family group (e.g., elevating a tribe to family) do not affect authorship or date of any name based on the same type genus. The corrected authorship and date of all names is provided in the catalog presented herein and summarized in table 1.
Table 1
Hierarchical Classification of Dermaptera
In some instances the name currently in use for a subfamily is incorrect, perhaps owing to the general disregard for the Principle of Coordination, described above. This disregard or misunderstanding has led to a failure to correctly identify instances of priority. The most serious failure to follow the Principle of Priority (ICZN, 1999: Art. 23) is with the name Labiinae, which is, in fact, a junior synonym of Isolabellinae in the current system (e.g., Steinmann, 1989b, 1990). Thus, the valid name for this group is Isolabellinae. However, owing to the confusion created by such a required change, a petition to grant priority under plenary powers to Labiinae was submitted to and approved by the Commission (Engel, 2004; ICZN, 2005). Another required change under the Code is the usage of Skendylinae Burr, 1907, in place of Cosmiellinae Steinmann, 1975. Skendylinae is the only valid name for this subfamily and, because changing the name back to its taxonomically correct epithet is not so damaging, no petition was submitted to conserve Cosmiellinae. The synonymy of Skendyle Burr, 1907, with Cosmiella Verhoeff, 1902a, apparently led to the confusion that Skendylinae, although older (i.e., having priority), should be a synonym of Cosmiellinae. This is not correct, and Skendylinae is indeed valid despite the synonymy of its type genus (ICZN, 1999: Art. 40.1), since the synonymy of Skendyle was not done until 1989 (Steinmann, 1989b; ICZN, 1999: Art. 40.2). A similar situation is true for Platylabiinae versus Palicinae and no action was taken to conserve the junior name.
In the following list, names are listed in order by priority according to date of establishment and in their form as originally proposed (even as originally misspelled: corrected spellings of all names are in table 1). The format generally follows that used for other family-group name summaries (e.g., Engel and Krishna, 2004; Engel, 2005). Daggers (†) indicate names for fossil lineages. The author and date are given for the first usage of a family-group name based on the indicated type genus. Finally, the correct orthography of the family-group name is provided. This stem is to be used when adding family-group suffices (e.g., –oidea, –idae, –inae, –ini, –ina) to the name. There is understandable difficulty in determining the appropriate termination for family-group names, particularly for those based on generic names of third-declension Greek nouns, e.g., labis, meaning “forceps”. Family-group names are to be based on the genitive singular form (ICZN, 1999: Art. 29.3.1). For the example given, the genitive singular form is labidos. Thus, names based on labis have the stem labid– while names based on labia (Latin, meaning “lips”) have the stem labi–. Thus, Anisolabididae (Anisolabidinae, Anisolabidini, &c.) is the correct spelling for a family-group name based on Anisolabis while Labiidae (Labiinae, Labiini, &c.) is the correct spelling for a family-group name based on Labia.
Various Latin terms and phrases are employed for taxonomic events herein. A listing of this terminology and their English equivalents is summarized as follows: pro (for, or, in place of), nec (not), nomen conservandum (a conserved name), nomen correctum (a corrected name, i.e., the stem has required a correction in spelling), nomen dubium (a dubious name), nomen illegitimum (an illegitimate name), nomen imperfectum (an imperfect name), nomen invalidum (an invalid name), nomen novum (a replacement name), nomen nudum (a bare name, i.e., a name without an associated description), nomen praeoccupatum (a preoccupied name), nomen rejiciendum (a rejected name), nomen translatum (an altered name, i.e., altered in suffix); recte (corrected as).
To all of this is appended several nomenclatural changes for earwigs that were identified in connection with the production of the present work, including the description of a new, extinct subfamily (see appendix).
Classification of Dermaptera
Table 1 briefly summarizes the current hierarchical classification of Dermaptera, with a particular emphasis on the application of taxonomic names. This summary is not meant to be a definitive classification of the order. Higher earwig classification is in the process of being cladistically revised (e.g., Haas, 1995; Haas and Kukalová-Peck, 2001; Haas and Klass, 2003; Colgan et al., 2003; Grimaldi and Engel, 2005; Jarvis et al., 2005), and the current outline is meant solely to provide a preliminary foundation from which to correctly apply family-group names throughout the order as such work progresses.
A still difficult question, which has strong influence on the rank and circumscription of particular groups, is the phylogenetic position of the epizoic Hemimeridae (on African hamster rats) and the Arixeniidae (on oriental bats). Both lineages have acquired a highly derived morphology and natural history (e.g., both are oviparous, both are largely paedomorphic) and were thus often regarded as separate suborders of the Dermaptera (as Hemimerina and Arixeniina). The remaining earwigs were thus classified into a third suborder, the Forficulina. However, there are indications that the epizoic lineages are indeed derived from within the main lineage of nonparasitic Dermaptera (i.e., the former suborder Forficulina; e.g., Haas and Klass, 2003) and should consequently be demoted in rank. Inclusion of these paedomorphic, epizoic parasites also has implications for the monophyly and definition of the Dermaptera and its constituent groups, issues that deserve further investigation as phylogenetic studies are expanded.
In table 1, names presently considered synonymous are italicized under the valid name. All names are presented with their corrected authorship, date, and spelling of their combining stem (when the stem, not the suffix, has required correction, then the name is labeled as “nomen correctum”, see above).
Family-group Names
In the following summary of family-group names in Dermaptera we have ordered the names by their nomenclatural priority, i.e., by date from which the name became available (ICZN, 1999: Art. 23), following the format of similar family-group name accounts (e.g., Engel, 2005; Engel and Krishna, 2004). For ease of use, the names are also summarized alphabetically in table 2. Readers wishing to locate a name based on a particular type genus but who are unfamiliar with its correct authorship and date can locate the name in table 2 and then find the appropriate place in the text (note that many authorships and dates employed in earwig taxonomy are nomenclaturally incorrect, e.g., Steinmann, 1989b).
Table 2
Alphabetical Arrangement of Family-Groups Names in Dermaptera (all names appear in their original form except nomina imperfecta are corrected)
1. Forficulariae Latreille, 1810: 244, 246. Type genus: Forficula Linnaeus, 1758. Combining stem: Forficul–. Note: Authorship of this name is often given to Stephens (1829a: 299; 1829b: 25; e.g., Steinmann, 1989b, 1993; Engel, 2003). However, Stephens was not the first to propose a family-group name based on Forficula. Latreille proposed the Forficulariae as a family for Forficula in Orthoptera. Article 11.7.1.3 (ICZN, 1999) states that “a family-group name of which the family-group name suffix [Art. 29.2] is incorrect is available with its original authorship and date, but with a corrected suffix [Arts. 29, 32.5.3].” Indeed, even the example given on the same page is for a Latreille family-group name employing his typical family-group suffix (–ariae). The example states, “Latreille … established a family Tipulariae, based on Tipula Linnaeus, 1758. The suffix –ariae is corrected to –idae; Tipulidae is attributed to Latreille, not to the author who first corrected the spelling.” The same applies for Latreille's Forficulariae, and despite Stephens's later action as the first author to employ a family-group name with what we today consider a correct suffix, Forficulidae (and its other forms) must date from 1810 with Latreille as author.
2. Hemimeridae Sharp, 1895: 217. Type genus: Hemimerus Walker, 1871. Combining stem: Hemimer–. Note: These paedomorphic, epizoic earwigs on hamster rats in sub-Saharan Africa are highly autapomorphic and as such have at times been considered a separate order or suborder under names such as Diploglossata, Dermodermaptera, and Hemimerina (e.g., de Saussure, 1879; Verhoeff, 1902c; Burr, 1911c; Popham, 1961; Nakata and Maa, 1974; see “Names above the Family Group”, below).
3. Karschiellidae Verhoeff, 1902a: 183. Type genus: Karschiella Verhoeff, 1902a. Combining stem: Karschiell–. Note: This group is variously grouped as a subfamily of Pygidicranidae but is now considered a separate, basal family (see table 1).
4. Anisolabidae Verhoeff, 1902a: 185, nomen imperfectum [recte Anisolabididae]. Type genus: Anisolabis Fieber, 1853. Combining stem: Anisolabid–. Note: In many modern works the terms Anisolabididae and Carcinophoridae are used interchangeably.
5. Gonolabidae Verhoeff, 1902a: 186, nomen imperfectum [recte Gonolabididae]. Type genus: Gonolabis Burr, 1900a. Combining stem: Gonolabid–. Note: Steinmann (1989a, 1989b) erroneously cites Gonolabina Verhoeff, 1902b, as the type genus for this name and thereby asserts that it is identical to the Gonolabiinae of Popham and Brindle (1966c). Verhoeff (1902a) clearly includes only Gonolabis in his “Gonolabidae” (in fact, he did not describe and add Gonolabina until Verhoeff, 1902b) and thus Gonolabis must be considered the type genus of this name (indeed, the family-group name is grammatically derived from Gonolabis and not Gonolabina), thereby making this name synonymous with Anisolabidinae ( = Carcinophoridae Popham, 1965) in the current system of Dermaptera.
6. Cheliduridae Verhoeff, 1902a: 186. Type genus: Chelidura Latreille, 1825. Combining stem: Chelidur–.
7. Diplatyidae Verhoeff, 1902a: 187. Type genus: Diplatys Audinet-Serville, 1831. Combining stem: Diplaty–. Note: This group is variously grouped as a subfamily of Pygidicranidae but is now considered a separate, basal family (see table 1).
8. Pygidicranidae Verhoeff, 1902a: 188. Type genus: Pygidicrana Audinet-Serville, 1831. Combining stem: Pygidicran–.
9. Pyragrinae Verhoeff, 1902a: 189. Type genus: Pyragra Audinet-Serville, 1831. Combining stem: Pyragr–. Note: The circumscription of the Pygidicranidae in taxonomic works is unstable, with various genera, and Karschiellidae as well as Diplatyidae, moved in and out of the taxon.
10. Labiduridae Verhoeff, 1902a: 189. Type genus: Labidura Leach, 1815. Combining stem: Labidur–. Note: The circumscription of the Labiduridae in taxonomic works is unstable, with various genera moved in and out of the taxon.
11. Nesogastrinae Verhoeff, 1902a: 191. Type genus: Nesogaster Verhoeff, 1902a. Combining stem: Nesogastr–.
12. Ancistrogastrinae Verhoeff, 1902a: 193. Type genus: Ancistrogaster Stål, 1855. Combining stem: Ancistrogastr–.
13. Spongiphorinae Verhoeff, 1902a: 193. Type genus: Spongiphora Audinet-Serville, 1831. Combining stem: Spongiphor–.
14. Allodahliinae Verhoeff, 1902a: 194. Type genus: Allodahlia Verhoeff, 1902a. Combining stem: Allodahli–.
15. Opisthocosmiinae Verhoeff, 1902a: 195. Type genus: Opisthocosmia Dohrn, 1865. Combining stem: Opisthocosmi–.
16. Chelisochini Verhoeff, 1902a: 196. Type genus: Chelisoches Scudder, 1876. Combining stem: Chelisoch–.
17. Anechurini Verhoeff, 1902a: 196. Type genus: Anechura Scudder, 1876. Combining stem: Anechur–.
18. Apterygidini Verhoeff, 1902a: 196. Type genus: Apterygida Westwood, 1839. Combining stem: Apterygid–.
19. Sparattinae Verhoeff, 1902a: 198. Type genus: Sparatta Audinet-Serville, 1839. Combining stem: Sparatt–. Note: Steinmann (1990) established a new tribe based on the genus Sparatta Audinet-Serville, 1839. However, according to the Principle of Coordination (ICZN, 1999: Art. 36) any family-group name based on Sparatta is to take its authorship and date from its first available usage, in this case, Verhoeff (1902a).
20. Apachyidae Verhoeff, 1902a: 200. Type genus: Apachyus Audinet-Serville, 1831. Combining stem: Apachy–.
21. Isolabidae Verhoeff, 1902b: 10, nomen imperfectum [recte Isolabididae]. Type genus: Isolabis Verhoeff, 1902b. Combining stem: Isolabid–.
22. Isolabellinae Verhoeff, 1902b: 15. Type genus: Isolabella Verhoeff, 1902b. Combining stem: Isolabell–. Note: This name is to be suppressed in favor of Labiidae Burr, 1909b (see below) whenever the two names are considered synonyms (Engel, 2004; ICZN, 2005).
23. Allostethidae Verhoeff, 1904: 70. Type genus: Allostethus Verhoeff, 1904. Combining stem: Allosteth–.
24. Parisolabinae Verhoeff, 1904: 119, nomen imperfectum [recte Parisolabidinae]. Type genus: Parisolabis Verhoeff, 1904. Combining stem: Parisolabid–.
25. Eudohrninae Burr, 1907: 97, nomen imperfectum [recte Eudohrniinae]. Type genus: Eudohrnia Burr, 1907. Combining stem: Eudohrni–. Note: This name is a junior synonym of Neolobophorinae Burr, 1907.
26. Diaperasticinae Burr, 1907: 97. Type genus: Diaperasticus Burr, 1907. Combining stem: Diaperastic–.
27. Skendylinae Burr, 1907: 117. Type genus: Skendyle Burr, 1907. Combining stem: Skendyl–.
28. Neolobophorinae Burr, 1907: 118. Type genus: Neolobophora Scudder, 1875. Combining stem: Neolobophor–.
29. Eparchinae Burr, 1907: 120. Type genus: Eparchus Burr, 1907. Combining stem: Eparch–. Note: This name is a junior synonym of Opisthocosmiinae Verhoeff, 1902a.
30. Doratinae Burr, 1907: 123. Type genus: Doru Burr, 1907. Combining stem: Dorat–. Note: This name is a junior synonym of Forficulinae Latreille, 1810.
31. Brachylabinae Burr, 1908a: 247, nomen imperfectum [recte Brachylabidinae]. Type genus: Brachylabis Dohrn, 1864. Combining stem: Brachylabid–.
32. Arixeniidae Jordan, 1909: 323. Type genus: Arixenia Jordan, 1909. Combining stem: Arixeni–. Note: The erroneous spelling Arixenidae is unfortunately common for these epizoic species on bats from Borneo and the Philippines. In the past the family was placed in a separate suborder (see “Names above the Family Group”, below).
33. Esphalmeninae Burr, 1909a: 250. Type genus: Esphalmenus Burr, 1909a. Combining stem: Esphalmen–.
34. Anataelinae Burr, 1909b: 322, nomen imperfectum [recte Anataeliinae]. Type genus: Anataelia Bolivar, 1899. Combining stem: Anataeli–.
35. Labiidae Burr, 1909b: 323. Type genus: Labia Leach, 1815. Combining stem: Labi–. Note: This name is to be preserved in favor of Isolabellinae Verhoeff, 1902b (see below) whenever the two names are considered synonyms (Engel, 2004; ICZN, 2005). In many modern works the terms Labiidae and Spongiphoridae are used interchangeably.
36. Psalinae Burr, 1909b: 325. Type genus: Psalis Audinet-Serville, 1831. Combining stem: Psal–. Note: This name is a junior synonym of Anisolabidinae Verhoeff, 1902a.
37. Auchenominae Burr, 1909b: 326. Type genus: Auchenomus Karsch, 1886. Combining stem: Auchenom–.
38. Echinosomatinae Burr, 1910a: 67. Type genus: Echinosoma Audinet-Serville, 1839. Combining stem: Echinosomat–.
39. Palicinae Burr, 1910a: 67. Type genus: Palex Burr, 1910a. Combining stem: Palic–.
40. Pericominae Burr, 1911a: 59. Type genus: Pericomus Burr, 1911a. Combining stem: Pericom–.
41. Vandicinae Burr, 1911a: 59. Type genus: Vandex Burr, 1911a. Combining stem: Vandic–.
42. Strongylopsalinae, Burr, 1911a: 59. Type genus: Strongylopsalis Burr, 1900b. Combining stem: Strongylopsal–.
43. Platylabiinae Burr, 1911c: 43. Type genus: Platylabia Dohrn, 1867. Combining stem: Platylabi–. Note: This name is a junior synonym of Palicinae Burr, 1910a.
44. Blandicinae Burr, 1915: 425. Type genus: Blandex Burr, 1912. Combining stem: Blandic–.
45. Landicinae Burr, 1915: 445. Type genus: Landex Burr, 1915. Combining stem: Landic–. Note: This name is a junior synonym of Anisolabidinae Verhoeff, 1902a.
46. †Protodiplatyidae Martynov, 1925a: 573. Type genus: †Protodiplatys Martynov, 1925a. Combining stem: Protodiplaty–.
47. Cylindrogastrinae Maccagno, 1929: 7. Type genus: Cylindrogaster Stål, 1855. Combining stem: Cylindrogastr–.
48. †Ocelliidae Ewing, 1942: 95, nomen dubium. Type genus: †Ocellia Olfers, 1907. Combining stem: Ocelli–. Note: This name was proposed for an enigmatic fossil in Baltic amber that was originally believed to be an apterous hexapod of the Diplura (Ewing, 1942). The fossil is, however, actually an immature earwig of uncertain identity. The name Ocelliidae is, therefore, an available name within the Dermaptera but of uncertain status. It is likely a synonym of Forficulidae since species of this family are apparently the most common form in Baltic amber (Burr, 1911d; Wappler et al., 2005), although nymphs of Labiduridae and Pygidicranidae are also known in Baltic amber, so it is possible that Ocelliidae is instead synonymous with one of them. Olfers' (1907) original material must be located and examined in order to clarify the status of Ocelliidae, something best done in the context of a revision of all Baltic amber Dermaptera.
49. Isopyginae Hincks, 1951: 12. Type genus: Isopyge Borelli, 1931. Combining stem: Isopyg–.
50. Diplatymorphinae Boeseman, 1954: 5. Type genus: Diplatymorpha Boeseman, 1954. Combining stem: Diplatymorph–.
51. Protolabinae Bey-Bienko, 1959a: 598, nomen imperfectum [recte Protolabidinae] et nomen illegitimum. Type genus: Protolabis Bey-Bienko, 1959a [nomen praeoccupatum, nec Protolabis Cope, 1876 (Mammalia): see Prolabiscinae, below]. Combining stem: Protolabid–. Note: This family-group name is invalid since its type genus is a junior homonym (ICZN, 1999: Art. 39).
52. Prolabiscinae Bey-Bienko, 1959b: 943. Type genus: Prolabisca Bey-Bienko, 1959b [nomen novum pro Protolabis Bey-Bienko, 1959a]. Combining stem: Prolabisc–. Note: A replacement for Protolabidinae (see also Protolabinae, above).
53. Carcinophorinae Popham, 1965: 132. Type genus: Carcinophora Scudder, 1876. Combining stem: Carcinophor–. Note: This name is traditionally attributed to Hincks (1954: p. 5) who first used the name as a subfamily of Labiduridae. However, Hincks (1954) failed to provide characters to differentiate the subfamily from other taxa or descriptive details of any kind for this group, an explicit requirement for availability for all family-group names established after 1930 (ICZN, 1999: Arts. 13.1, 13.2). The name Carcinophorinae in 1954 as used by Hincks was a lic>nomen nudum and is unavailable. The first author to make a family-group name based on Carcinophora available was Popham (1965) who used the name as a family, Carcinophoridae, of Labiodea. The name must therefore take its date and authorship from Popham (1965). In many modern works the terms Carcinophoridae and Anisolabididae are used interchangeably.
54. Gonolabinae Popham and Brindle, 1966c: 277, nomen imperfectum [recte Gonolabininae]. Type genus: Gonolabina Verhoeff, 1902b. Combining stem: Gonolabin–.
55. Physogastrinae Ramamurthi, 1967: 237, nomen illegitimum. Type genus: Physogaster Ramamurthi, 1967 [nomen praeoccupatum, nec Physogaster Lacordaire, 1830 (Coleoptera), and others: see Ramamurthiinae, below]. Combining stem: Physogastr–. Note: This family-group name is invalid since its type genus is a junior homonym (ICZN, 1999: Art. 39).
56. Geracinae Brindle, 1971: 158. Type genus: Gerax Hebard, 1917. Combining stem: Gerac–.
57. Challinae Steinmann, 1973: 388, nomen imperfectum [recte Challiinae]. Type genus: Challia Burr, 1904. Combining stem: Challi–.
58. Anophthalmolabiinae Steinmann, 1975: 205, nomen imperfectum [recte Anophthalmolabidinae]. Type genus: Anophthalmolabis Brindle, 1968. Combining stem: Anophthalmolabid–.
59. Idolopsalinae Steinmann, 1975: 206. Type genus: Idolopsalis Borelli, 1910. Combining stem: Idolopsal–.
60. Nalinae Steinmann, 1975: 207. Type genus: Nala Zacher, 1910. Combining stem: Nal–.
61. Ramamurthiinae Steinmann, 1975: 210. Type genus: Ramamurthia Steinmann, 1975 [nomen novum pro Physogaster Ramamurthi, 1967]. Combining stem: Ramamurthi–. Note: A replacement name for Physogastrinae (see above).
62. Chelisochellinae Steinmann, 1975: 214. Type genus: Chelisochella Verhoeff, 1902a. Combining stem: Chelisochell–.
63. Sarcinatricinae Steinmann, 1975: 216. Type genus: Sarcinatrix Rehn, 1903. Combining stem: Sarcinatric–.
64. Cosmiellinae Steinmann, 1975: 217. Type genus: Cosmiella Verhoeff, 1902a. Combining stem: Cosmiell–.
65. Rhyacolabinae Steinmann, 1975: 219, nomen imperfectum [recte Rhyacolabidinae]. Type genus: Rhyacolabis Rehn, 1921. Combining stem: Rhyacolabid–.
66. Antisolabiinae Brindle, 1978a: 18, nomen imperfectum [recte Antisolabidinae]. Type genus: Antisolabis Burr, 1911b. Combining stem: Antisolabid–.
67. Isolaboidinae Brindle, 1978b: 204. Type genus: Isolaboides Hincks, 1958. Combining stem: Isolaboid–.
68. †Dermapterinae Vishniakova, 1980: 87. Type genus: †Dermapteron Martynov, 1925a. Combining stem: Dermapter–.
69. †Semenoviolinae Vishniakova, 1980: 90. Type genus: †Semenoviola Martynov, 1925b. Combining stem: Semenoviol–.
70. Cosmogeracinae Brindle, 1982: 35. Type genus: Cosmogerax Hebard, 1933. Combining stem: Cosmogerac–.
71. Titanolabinae Srivastava, 1982a: 97, nomen imperfectum [recte Titanolabidinae]. Type genus: Titanolabis Burr, 1910b. Combining stem: Titanolabid–.
72. Homotaginae Srivastava, 1985a: 206. Type genus: Homotages Burr, 1909b. Combining stem: Homotag–.
73. Irdexinae Srivastava, 1985a: 206, nomen imperfectum [recte Irdicinae]. Type genus: Irdex Burr, 1911a. Combining stem: Irdic–. Note: This name is a junior synonym of Spongiphorinae Verhoeff, 1902a.
74. Brindlensiinae Srivastava, 1985b: 45. Type genus: Brindlensia Srivastava, 1985b. Combining stem: Brindlensi–.
75. Genitalatinae Steinmann, 1987: 114. Type genus: Genitalata Kapoor, 1974. Combining stem: Genitalat–.
76. Caecolabiinae Steinmann, 1990: 70. Type genus: Caecolabia Brindle, 1975. Combining stem: Caecolabi–.
77. Chaetospanini Steinmann, 1990: 173, nomen imperfectum [recte Chaetospaniini]. Type genus: Chaetospania Karsch, 1886. Combining stem: Chaetospani–.
78. †Longicerciatidae Zhang, 1994: 231. Type genus: †Longicerciata Zhang, 1994. Combining stem: Longicerciat–. Note: This name is a junior synonym of †Protodiplatyidae Martynov, 1925a.
79. Rudraxinae Srivastava, 1995: 76, nomen imperfectum [recte Rudracinae]. Type genus: Rudrax Srivastava, 1995. Combining stem: Rudrac–.
80. Placolabidinae Srivastava, 1999: 80. Type genus: Placolabis Bey-Bienko, 1959a. Combining stem: Placolabid–. Note: This name is a junior synonym of Anisolabidinae Verhoeff, 1902a.
81. †Sinopalaeodermatidae Zhang, 2002: 351. Type genus: †Sinopalaeodermata Zhang, 2002. Combining stem: Sinopalaeodermat–. Note: This name is a junior synonym of †Dermapterinae Vishniakova, 1980.
82. †Turanoviidae Engel, 2003: 116. Type genus: †Turanovia Vishniakova, 1980. Combining stem: Turanovi–.
83. †Turanodermidae Engel, 2003: 116, nomen imperfectum [recte †Turanodermatidae; rectified by Engel and Grimaldi, 2004]. Type genus: †Turanoderma Vishniakova, 1980. Combining stem: Turanodermat–.
84. Kinellinae Srivastava, 2003: 160, nomen imperfectum [recte Kinesinae]. Type genus: Kinesis Burr, 1907. Combining stem: Kines–.
85. †Burmapygiinae Engel and Grimaldi, 2004: 1018. Type genus: †Burmapygia Engel and Grimaldi, 2004. Combining stem: Burmapygi–.
86. †Cretolabiinae Engel and Haas, herein (see appendix). Type genus: †Cretolabia Popham, 1990. Combining stem: Cretolabi–.
Names Above The Family Group
Presently zoological nomenclature does not regulate names applied above the family group (i.e., those names above superfamily). As such, the application and stabilization of suprafamilial names has largely come about by convention rather than regulation (e.g., principles like priority have no authority for such names). Table 3 provides a brief listing of names applied to lineages of earwigs above the family group (e.g., suborders, infraorders, parvorders). We have not included those names pertaining to the extinct, likely paraphyletic, order †Protelytroptera Tillyard, 1931 (in part comprising Paleozoic, stem-group Dermaptera: see Grimaldi and Engel, 2005). †Protelytroptera and Dermaptera together comprise the superorder Dermapterida Boudreaux, 1979 (see hierarchical classification of Polyneoptera in Arillo and Engel, 2006). For a similar grouping of some taxa of the †Protelytroptera and the Dermaptera, the typified name Forficulida is used by some (e.g., Shcherbakov, 2002). Like †Protelytroptera we have also not included those names applicable to groupings above the level of order (i.e., above Dermaptera). Dermapteroidea Jeannel (In Grassé, 1949) is equivalent to Dermapterida but should be avoided owing to confusion with the standard suffix for the superfamily rank, i.e., –oidea. In general we recommend avoiding names for higher categories that terminate with suffixes identical to those employed within the family group as they only serve to confuse.
Table 3
Names Above the Family Group in Dermaptera (names are ordered by date)
Our list includes only those names equivalent to Dermaptera or groupings of families within Dermaptera. Certainly taxonomic and phylogenetic concepts change through time and some names were originally formulated to include taxa not consistent with today's usages or notions (just as some modern assertions will, in time, similarly prove untenable). Even some of our most familiar and widely employed names fall into this category. For example, Eudermaptera and even Dermaptera were established by their original authors for amalgamations of taxa differing from the taxon membership we today ascribe to them (e.g., Eudermaptera in the sense of Verhoeff included all modern earwigs except Arixeniidae, Hemimeridae, and Apachyidae; Dermaptera as conceived by de Geer included not only earwigs but mantises, roaches, and many orthopterans). Another relevant example would be the “Orthoptera” which originally included earwigs, roaches, grasshoppers (and their kin), and occasionally dragonflies as well. To today's standards such an application is more a loose ecological grouping than anything else.
Nonetheless, we believe it is most conservative to maintain current usage of such names rather than create confusion by reverting to archaic names more closely matching our modern concepts of taxonomic composition (e.g., given the universal usage of Dermaptera for earwigs it is preferable for taxonomic stability to retain this name rather than to revert to something like Labidoura, Euplekoptera, or Dermatoptera simply because they were originally established strictly for earwigs or a group of earwigs).
Acknowledgments
We are grateful to Drs. Torsten Wappler and Vincent Perrichot for carefully checking the manuscript. Partial support for this work was provided by NSF DEB-0542909 and a Guggenheim Fellowship from the John Simon Guggenheim Memorial Foundation (to MSE). This is contribution No. 3446 of the Division of Entomology, University of Kansas Natural History Museum.
References
Appendices
Appendix
Nomenclatural Notes for Earwigs
In the process of studying the taxonomy and specimens of Dermaptera in the preparation of this catalog, a few nomenclatural difficulties or changes for modern earwigs were identified. These are attended to herein. In addition, the Early Cretaceous genera †Cretolabia and †Kotejalabis are placed in a new subfamily of Anisolabididae.
Family Pygidicranidae Verhoeff
Genus Acrania Burr, reinstated
Acrania Burr, 1915: 436. Type species: Pygidicrania picta Guérin-Méneville, 1838, by original designation.
Epicranopygia Steinmann, 1986: 269. Type species: Pygidicrania picta Guérin-Méneville, 1838, by original designation. New synonymy.
Comments
Steinmann (1989b) erroneously considered Epicranopygia the valid name of this genus since Acrania had been considered a synonym of Cranopygia by earlier authors. Earlier synonymies do not render the name Acrania invalid, and it remains an available name for the group containing its type species, thereby making it the oldest available name for the present genus. Acrania is here reinstated, and Epicranopygia (based on the same type species) becomes a junior, objective synonym. The following nomenclatural changes are thus required: Acrania angulata (Srivastava), new combination; A. constricta (Hincks), new combination; A. eximia (Dohrn), reinstated combination; A. fletcheri (Bharadwaj and Kapoor), new combination; A. picta (Guérin-Méneville), reinstated combination; A. triangulata (Ramamurthi), new combination; A. vittipennis (Hincks), new combination.
Genus Pyge Burr, reinstated
Pyge Burr, 1908b: 390. Type species: Pygidicrania modesta de Bormans, 1894.
Paracranopygia Steinmann, 1986: 277. Type species: Forficula pallidipennis de Haan, 1842. New synonymy.
Comments
Steinmann (1989b), for the same erroneous reasons discussed for Acrania (see above), provided a new name for this genus and relegated Pyge to synonymy. Pyge remains an available genus-group name for any group containing its type species. Thus, it becomes the valid name for the group appearing in the literature as Paracranopygia. The following combinations result: Pyge assamensis (Hincks), new combination; P. bakeri (Borelli), new combination; P. burmensis (Hincks), new combination; P. comata (Hincks), new combination; P. formosa (Hincks), new combination; P. maculipes (Hincks), new combination; P. meghalayana (Biswas et al.), new combination; P. modesta (de Bormans), reinstated combination; P. pallidipennis (de Haan), new combination; P. proxima (Hincks), new combination; P. semenovi (Burr), new combination; P. siamensis (Dohrn), new combination; P. similis (Zacher), new combination; P. tonkinensis (Hincks), new combination; P. variegata (Brindle), new combination; P. vicina (Hincks), new combination.
Family Diplatyidae Verhoeff
Genus Paradiplatys Zacher, reinstated
Comments
In recent monographic treatments this genus has been erroneously cited as Lobodiplatys Steinmann, 1974 (Steinmann, 1986, 1989b). These usages, however, are in violation of the Principle of Priority (ICZN, 1999: Art. 23), and the entire genus must bear the name of the oldest, included, available genus-group name (whether or not it was equivalent at its original proposal to what is today merely a subgenus of a larger taxonomic entity). Accordingly, the valid name for Lobodiplatys is Paradiplatys. The usage of these names in the literature is infrequent enough that there is no reason to petition the ICZN for suppression of Paradiplatys under the plenary powers. The resultant nomenclatural changes in the four subgenera are (the utility of the subgeneric classification should be critically re-examined as Steinmann's subgenera likely render Paradiplatys s. str. paraphyletic):
Paradiplatys (Lobodiplatys), new subgeneric placement: Paradiplatys (Lobodiplatys) coriaceus (Kirby), reinstated combination.
Paradiplatys (Paradiplatys): Paradiplatys (Paradiplatys) conradti (Burr), reinstated combination; P. (P.) lamottei (Hincks), reinstated combination; P. (P.) pectinatus (Hincks), reinstated combination; P. (P.) salvazae (Burr), reinstated combination; P. (P.) spinulosus (Hincks), reinstated combination.
Paradiplatys (Heterodiplatys), new subgeneric placement: Paradiplatys (Heterodiplatys) bicolor (Dubrony), reinstated combination; P. (H.) bihamatus (Hincks), reinstated combination; P. (H.) burri (Hincks), reinstated combination; P. (H.) rotundicollis (Hincks), reinstated combination; P. (H.) schoutedeni (Hincks), reinstated combination.
Paradiplatys (Epidiplatys), new subgeneric placement: Paradiplatys (Epidiplatys) gladiator (Burr), reinstated combination.
Family Anisolabididae Verhoeff
†Cretolabiinae, new subfamily
Diagnosis
Earwigs of average size (ca. 7–11 mm in length), not flattened. Compound eyes well developed. Scape short, much shorter than distance between antennae; antennae with 20 articles (known only for †Kotejalabis). Tegmina present and well developed. Probasisternum cordiform, lateral margins tapering posteriorly between procoxae; mesosternum roughly quadrate, posterior margin straight (not convex as in many Anisolabidinae). Abdomen ovoid (sides in †Kotejalabis more elongate, but still somewhat convex). Pygidium distinct, apparently bent ventrally, not fused to apicalmost tergum.
Comments
Among anisolabidids †Cretolabiinae is similar to the subfamily Gonolabininae (as Gonolabidinae in Popham, 1990) in that both have cordiform probasisternum, albeit more strongly so in the gonolabinines. The probasisternum in other neodermapterans is typically constricted posteriorly near the procoxae but then expands along its posterior margin. The cordiform shape is likely an apomorphic feature given its distribution across Neodermaptera and thus might indicate a relationship between the southern South American Gonolabininae (today occurring in Chile and Peru) and the fossil lineage. †Cretolabiinae can be differentiated from Gonolabininae by the more ovoid abdomen (roughly parallel-sided in gonolabinines), by the presence of wings (absent in gonolabinines), and by the separate apical abdominal tergum and pygidium (fused in gonolabinines), all notably plesiomorphic features.
The genus †Kotejalabis, from the same deposits as †Cretolabia, is tentatively included in †Cretolabiinae owing to a similar cordiform probasisternum (Engel and Chatzimanolis, 2005: fig. 4). Unlike †Cretolabia, †Kotejalabis has an elongate and prominent pygidium, shorter thorax, and differences in cercal form (Engel and Chatzimanolis, 2005). Although not discussed by Engel and Chatzimanolis (2005), †Kotejalabis also has an enlarged profurcasternum ( = prosternellum) in comparison to †Cretolabia (cf. Popham, 1990: fig. 1; Engel and Chatzimanolis, 2005: fig. 4). In †Cretolabia the profurcasternum is a small, transverse sclerite (its width more than four times is length) tightly bordering the posterior margin of the probasisternum and no wider than the posterior width of the probasisternum. By contrast, in †Kotejalabis the profurcasternum, while also transverse, is longer, approximately twice as wide as long, and is slightly wider than the posterior margin of the probasisternum. In both genera the mesosternum is roughly quadrate with rounded corners, although in †Cretolabia the posterior margin is slightly narrower than the anterior margin while in †Kotejalabis the anterior and posterior margins are apparently equal.
Subfamily Titanolabidinae Srivastava
Genus Titanolabis Burr
Titanolabis Burr, 1910b: 168. Type species: Forcinella colossea Dohrn, 1864.
Paratitanolabis Srivastava, 1982a: 98. Type species: Paratitanolabis bormansi Srivastava, 1982a, monobasic and original designation. New synonymy.
Comments
The characters purported to differentiate Titanolabis from Paratitanolabis are relatively minor changes in the relative lengths of the male parameres (scarcely worthy subgeneric distinction and assuredly rendering Titanolabis paraphyletic). It is most conservative to recognize a single genus for this homogeneous group of Asian earwigs. The following two taxonomic changes are required:
Titanolabis bormansi (Srivastava), new combination
Paratitanolabis bormansi Srivastava, 1982a: 99.
Titanolabis myanmarensis, new name
Titanolabis bormansi Srivastava, 1982b: 379. Nomen praeoccupatum, nec Titanolabis bormansi Srivastava, 1982a.
