Zootaxa, Genus- and family-group names in the order Embioptera

Zootaxa 2055: 1 – 34 (2009) www.mapress.com

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Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

Genus- and family-group names in the order Embioptera (Insecta)

KELLY B. MILLER

Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131, USA.

E-mail: kbmiller@unm.edu

Abstract

All available genus- and family-group names in the Embioptera (Embiidina, Embiodea) are listed along with the original citation, original and current status, and the type taxon and method of designation. The early literature is reviewed, and the following taxonomic acts are established: 1) Embia brasiliensis Gray in Griffith and Pidgeon 1832 is fixed as the type species of Olyntha Gray in Griffith and Pidgeon 1832 (junior homonym, replaced by Embolyntha Davis) using the

First Revisor Principle, 2) Embia savignii Westwood 1837 is fixed as type species of Euembia Verhoeff 1904, and 3)

Saussurembia davisi Ross 1992 is synonomized with Saussurembia ruficollis Davis 1939. Numerous other taxonomic problems are examined and clarified and the various ordinal names briefly summarized.

Key words: Webspinner, Embiidina, Embiodea, classification, catalog, taxonomy.

Introduction

Embioptera (webspinners) are among the most poorly known insects. Less than 400 valid species are currently recognized, but estimates by one prominent embiopteran researcher suggests there may also be as many as 1500 undocumented species already in collections (Ross 1991). The most recent comprehensive treatments of Embioptera family- and genus-group names in single documents were by Enderlein (1912) and

Krauss (1911). Since then there have been a large number of names added, and no modern synopsis of supraspecific names exists. Such a resource would be invaluable given the relatively large numbers of taxa remaining to be described in this interesting group. A suitable first step in establishing a framework for future species-level taxonomic work on embiopterans is to catalog the available supraspecific names within the group. Current problematic issues with genus- and family-group names include incorrect, unclear or invalid type designations, incorrect spellings, usages and author attributions, and unsolved problems with synonomy and priority. Uncorrected problems with Embioptera nomenclature will inhibit future work or inevitably will produce problems for future researchers that could be solved now. This catalog of Embioptera family- and genus-group names seeks to review and correct many of these problems and create a taxonomic framework for future work on the group.

Methods

Taxon name entries .

All available names at the genus and family rank referring to extant and extinct taxa (marked with † ) belonging to the order Embioptera are included. Families are presented alphabetically and genera are presented alphabetically within their currently assigned family. Synonymic lists are ordered by date of name publication and alphabetically within the same date. The literature survey is complete through September,

2008.

Accepted by P. Kerr: 2 Mar. 2009; published: 27 Mar. 2009 1

For taxon names the following are provided:

1) original citation

2) original status with original spelling

3) type taxon and method of designation (if the type taxon is a synonym then the name of its senior synonym is given in parenthesis)

4) current status based on recent literature

í if a name is a replacement, this is indicated along with the name it replaces and the reason for its replacement

í if a name is a subjective synonym, a reference is made to the first time such synonymy was proposed.

í the original usage and taxonomic placement (immediate parent taxon) of names are provided.

í changes to names based on ICZN Opinions are indicated.

Rules and provisions of the fourth edition of the International Code of Zoological Nomenclature (ICZN

1999) (hereafter referred to as “the Code”) are strictly followed. Reference to Articles, Recommendations and

Examples used hereafter refer to those items in the Code.

Nomenclatural acts .

A few new nomenclatural acts, such as new synonymies, designation of type species, etc., are introduced in this catalog (see Discussion). No new taxa are introduced.

Distribution.

Distribution of taxa are given by the following zoogeographical regions: Afrotropical (AF), Palearctic (PA),

Nearctic (NA), Neotropical (NE), Australian (AU), Oriental (OR) and Pacific (PA). Potentially problematic boundaries between these regions are the following: Afrotropical/Palaearctic: southern border of Morocco,

Algeria, Libya, Egypt; Ausralian/Pacific: Oceanic islands between 100º and 180ºW and 30ºS to 20ºN;

Nearctic/Neotropical: Border between Mexico and United States; Oriental/Palaearctic: border between India and Pakistan, Kashmir, Himachal Pradesh, Uttar Pradesh, Nepal, Sikkim and Darjeeling, Bhutan, Arunachal

Pradesh, border between China and Burma, Laos and Vietnam (Löbl & Smetana 2003;2004); Oriental/

Australian: Lydekker Line.

Order-group names

Most insect orders, such as Coleoptera and Diptera, have names that date back to pre-Linnaean times and were well-established at the time formal classification began. Webspinners, however, were more recently discovered and have had considerable variation in the order-group names applied to them. Though not regulated by the International Code of Zoological Nomenclature, a review of some order-group names and their history is warranted.

During the earliest studies of taxa now included in Embioptera, they were regarded as related to taxa now in Isoptera (Westwood 1837), and early treatment including both of these groups placed them together in the

Neuroptera. The first clear effort to recognize standardized insect orders was by Sharp (1895), but several of his names were unacceptable since they bore endings typical of families (– idae ). This was true of Embiidae, as recognized by Shipley (1904), who introduced the name Embioptera Shipley 1904:260. However, several other supra-familial group names were already in use for these insects, and others were introduced after this.

The following annotated list summarizes many of the suprafamilial-group names associated with Embioptera:

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· Zootaxa 2055 © 2009 Magnolia Press MILLER

Adenopoda Verhoeff 1904:196

Original usage; Suborder of order Isoptera including only the family Embiidae

Aetioptera Enderlein 1909:171

Original usage; Superorder of class Insecta including Isoptera and Embioptera

Embiaria Handirsch 1903:733

Original usage; Order of subclass Embioidea

Embidaria Handlirsch 1906:33

Original usage; Subclass of class Insecta

Embiidina Enderlein 1909:171

Original usage; Order of superorder Aetioptera

Notes . This is one of the more widely used names for this order. It is the more appropriate spelling of the name Embidina introduced by Hagen 1861 (and referred to by Enderlein (1903)), and the current use of this name might be most appropriately attributed to Hagen as it is by Ross (2000). The first use of this spelling, however, was by Enderlein (1909). Enderlein (1912) attributed the origin of the name (or this particular spelling of the name) to himself (Enderlein 1903), but in that document he also used the spelling Embidina. In addition, and more importantly, both Hagen (1861) and Enderlein (1903) used the name at what would now be regarded as the family rank, not as an order. Therefore, Embiidina as an ordinal name is properly attributed to Enderlein (1909) as first pointed out by Engel & Grimaldi (2006).

Embiodea Kusnezov 1903:208

Original usage; Suborder of order Neuroptera.

Embioidea Handlirsch 1903:733

Original usage; Subclass of class Pterygogenia.

Embiomorpha Ross 2007:576

Original usage; Infraorder of Embiidina.

Embioptera Shipley 1904:260

Original usage; Order of group Exopterygota

Euplatyptera Crampton 1916:252

Original usage; Order of class Insecta.

Euembiaria Tillyard 1937:251

Original usage; Suborder of order Embiaria

Notes . This taxon included extant embiopterans whereas the suborder Protembiaria Tillyard 1937 included extinct taxa. Protembiaria no longer refers to taxa considered to be embiopterans (see Discussion below under Names no longer associated with Embioptera ).

GENUS- AND FAMILY-GROUP NAMES IN EMBIOPTERA Zootaxa 2055 © 2009 Magnolia Press ·

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Euembioptera Davis 1940:535

Original usage; Suborder of order Embioptera, replacement name for Euembiaria Tillyard 1937 (Davis

1940a).

Notes . See under Euembiaria above.

Netica Navás 1918:88

Original usage; Suborder of Embioptera.

Notes . The order Embioptera sensu Navás (1918) included the family Cylindrachaetidae Giglio-Tos, which he placed in the suborder Oryttica Navás. Cylindrachaetidae is currently in the Orthoptera and Netica

Navás refers only to Embioptera as currently defined.

Neoembiodea Engel and Grimaldi 2006:4

Original usage; Suborder of order Embioptera.

Notes . Engel and Grimaldi (2006) established this suborder to include all Embioptera except

Clothodidae Enderlein which has been regarded as the sister to a group containing the remaining families

(Ross 1944, 2000; Szumik 1996). They did not establish an order name to include the (possibly paraphyletic)

Clothodidae.

Oligoneura Börner 1904:526

Original usage; Suborder of order Isoptera including Embioptera.

Notes . Rendered obsolete by Handlirsh (1904) by recognition of Isoptera and Embiaria as separate orders and sunk by him into Embiaria Handlirsch 1903.

Oligotomorpha Ross 2007:577


Paedembiamorpha Ross 2006a:785


Lately there has been some concern about standardizing the order name for these insects. The two most commonly used during recent times have been Embioptera Shipley 1904 and Embiidina Enderlein 1903.

Ross (2000) has emphasized the use of the name Embiidina (though attributing it to Hagen (1861), see under

Embiidina above) because it is an older name (when attributed to Hagen (1861), not Enderlein (1903; or

1909)). He also regards Embioptera as an “…awkward, inappropriate name” (Ross 2000:1). Grimaldi and

Engel (2005) and Engel and Grimaldi (2006) argue that Embioptera, meaning “lively wing” in Greek, is not particularly descriptive. They also point out that the suffix – ina is standard for the rank of subtribe making

Embiidina undesirable as well. Their solution is the use of the name Embiodea Kusnezov 1903 based, in part, on priority (Engel & Grimaldi 2006; Grimaldi & Engel 2005). Formalized names that refer to Embioptera at the ordinal rank appear in the literature in the following order:

Embiodea Kusnezov 1903:208

Embiaria Handirsch 1903:733

Embioidea Handlirsch 1903:733

Oligoneura Börner 1904:526

Embioptera Shipley 1904:260

Adenopoda Verhoeff 1904:196

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Embidaria Handlirsch 1906:33

Aetioptera Enderlein 1909:171

Embiidina Enderlein 1909:171

Euembiaria Tillyard 1937:251

Euembioptera Davis 1940a:677

Neoembiodea Engel and Grimaldi 2006:4

Paedembiamorpha Ross 2006:785

Oligotomorpha Ross 2007:577

Embiomorpha Ross 2007:576

I agree with Grimaldi and Engel (2005) that the use of Embiidina is undesirable given that the suffix – ina is a standard ending for a family-group rank in zoological nomenclature. Declining to use Embioptera, however, because it is not descriptive, and adoption of the rarely used name Embiodea (which has an ending similar to the standardized superfamily suffix – oidea ), does not seem desirable, either. Webspinners are, in fact, lively and active insects, so Embioptera is more descriptive than they suggest, even if they are not great fliers or even winged in many cases. And, as they themselves note (Engel & Grimaldi 2006), ordinal names need not be particularly meaningful. Perhaps more importantly, however, the name Embioptera is (next to

Embiidina) the most commonly used and is similar to the majority of other insect order names which end in various versions of the Greek word, – pter . Other than Engel and Grimaldi (2006) significant authorities on the group (e.g. Davis, Edgerly, Enderlein, Krauss, Navás, Rimsky-Korsakov, Ross, Stefani, Szumik, see references) have not used Embiodea for the past century, much less in modern history. For these reasons, and with due respect to the argument by Engel and Grimaldi (2006), I recommend continued use of Embioptera as the standardized name for this order.

Family- and genus-group available names in Embioptera

Family Andesembiidae Ross 2003 : NE.

Andesembiidae Ross 2003a:1; as family of Embiidina; type genus: Andesembia Ross 2003a.

Genus Andesembia Ross 2003 : NE.

Andesembia Ross 2003a:2; as genus of Andesembiidae Ross 2003a; type species: Andesembia cuencae

Ross 2003a by original designation.

Genus Bryonembia Ross 2003 : NE.

Bryonembia Ross 2003a:10; as genus of Andesembiidae Ross 2003a; type species: Bryonembia amplialata Ross 2003a by original designation.

Family Anisembiidae Davis 1940 : NA NE.

Anisembiidae Davis 1940c:537; as family of Embioptera; type genus: Anisembia Krauss 1911.

Anisembiidae Ross 1940b:642; as family of Embioptera; type genus: Anisembia Krauss 1911; preoccupied by Davis 1940c (see Discussion below).

Microembiinae Ross 2001:23; as subfamily of Embiidae Burmeister 1839; type genus: Microembia Ross

1944; synonymy by Szumik (2004).


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Subfamily Anisembiinae Davis 1940 : NA NE.


Tribe Anisembiini Davis 1940 : NA NE.


Genus Anisembia Krauss 1911 : NA NE.

Anisembia Krauss 1911:74; as genus of Embiidae Burmeister 1839; type species: Embia texana

Melander 1902 by original designation.

Genus Bulbocerca Ross 1940 : NE.

Bulbocerca Ross 1940b:654; as subgenus of Anisembia Krauss 1911, elevated to genus by Ross

(1984a); type species: Anisembia sini Chamberlin 1923 by original designation.

Genus Glyphembia Ross 2003 : NE.

Glyphembia Ross 2003b:19; as genus of Anisembiini Davis 1940c; type species: Mesembia chamulae

Ross 1984a by original designation.

Tribe Mesembiini Ross 1940 : NE.

Mesembiinae Ross 1940b:643; as subfamily of Anisembiidae Davis 1940c; type genus: Mesembia Ross

1940a.

Genus Ectyphocerca Ross 2003 : NE.

Ectyphocerca Ross 2003b:8; as genus of Anisembiinae Davis 1940c ( nec Stenembiini nomen nudum , see Discussion below); type species: Ectyphocerca aureata Ross 2003b by original designation.

Genus Mesembia Ross 1940 : NE.

Mesembia Ross 1940a:12; as genus of Embioptera; type species: Oligotoma hospes Myers 1928 by original designation.

Genus Phallosembia Ross 2003 : NE.

Phallosembia Ross 2003b:13; as genus of Anisembiinae Davis 1940c ( nec Stenembiini nomen nudum , see Discussion below); type species: Phallosembia andina Ross 2003b by original designation.

Genus Stenembia Ross 1972 : NE.

Stenembia Ross 1972:139; as genus of Anisembiidae Davis 1940c; type species: Stenembia perenensis

Ross 1972 by original designation.

Tribe Poinarembiini Ross 2003 : NE.

Poinarembiini Ross 2003b:37; as tribe of Anisembiinae Davis 1940c; type genus: Poinarembia Ross

2003b.

Genus Poinarembia Ross 2003 : NE.

Poinarembia Ross 2003b:37; as genus of Poinarembiini Ross 2003b; type species: Poinarembia rota

Ross 2003b by original designation.

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Anisembiinae, tribe incertae sedis (see Discussion below)

Genus Exochosembia Ross 2003 : NE.

Genus Exochosembia Ross 2003b:32; as genus of Anisembiinae Davis 1940c ( nec Exchosembiini nomen nudum , see Discussion below); type species: Exochosembia cavagnaroi Ross 2003b by original designation.

Genus Isosembia Ross 2003 : NE.

Genus Isosembia Ross 2003b:14; as genus of Anisembiinae Davis 1940c ( nec Isosembiini nomen nudum , see Discussion below); type species: Mesembia aequalis Ross 1944 by original designation.

Genus Pogonembia Ross 2003 : NE.

Genus Pogonembia Ross 2003b:35; as genus of Anisembiinae Davis 1940c ( nec Exchosembiini nomen nudum , see Discussion below); type species: Pogonembia motaguae Ross 2003b by original designation (see Discussion below).

Genus Saussurembia Davis 1940 : NE.

Saussurembia Davis 1940b:191; as replacement name for Saussurella Davis 1939e, type species:

Saussurella ruficollis Davis 1939e according to Article 11.10 (see Discussion below);.

Saussurella Davis 1939e:573; as genus of Embioptera (original family-group not indicated); type species: Saussurella ruficollis Davis 1939e by original designation; preoccupied by Bolivar 1887.

Subfamily Aporembiinae Ross 2003 : NE.

Aporembiinae Ross 2003b:42; as subfamily of Anisembiidae Davis 1940c; type genus: Aporembia Ross

2003b.

Genus Aporembia Ross 2003 : NE.

Aporembia Ross 2003b:43; as genus of Aporembiinae Ross 2003b; type species: Aporembia sturmi


Subfamily Chelicercinae Ross 1984 : NA NE.

Chelicercinae Ross 1984a:28; as subfamily of Anisembiidae Davis 1940c; type genus: Chelicerca Ross

1940b.

Genus Brasilembia Ross 2003 : NE.

Brasilembia Ross 2003b:58; as genus of Chelicercinae Ross 1984a; type species: Brasilembia beckeri


Genus Chelicerca Ross 1940 : NA NE.

Chelicerca Ross 1940b:656; as subgenus of Anisembia Krauss 1911, elevated to genus by Ross (1944); type species: Anisembia davisi Ross 1940b by original designation.

Protochelicerca Ross 1944:449; as subgenus of Chilicerca Ross 1940b; type species: Chelicerca dampfi Ross 1944 by monotypy; synonymy by Ross (1984a).


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Genus Dactylocerca Ross 1940 : NA NE.

Dactylocerca Ross 1940b:659; as subgenus of Anisembia Krauss 1911, elevated to genus by Ross

(1984b); type species: Anisembia rubra Ross 1940b by original designation.

Genus Oncosembia Ross 2003 : NE.

Oncosembia Ross 2003b:120; as genus of Chelicercinae Ross 1984a; type species: Oncosembia biarmata Ross 2003b by original designation.

Genus Pelorembia Ross 1984 : NE.

Pelorembia Ross 1984a:41; as genus of Anisembiinae Davis 1940c; type species: Pelorembia tumidiceps Ross 1984a by original designation.

Genus Schizembia Ross 1944 : NE.

Schizembia Ross 1944:440; as genus of Anisembiidae Davis 1940c; type species: Schizembia grandis


Subfamily Chorisembiinae Ross 2003 : NE.

Chorisembiinae Ross 2003b:44; as subfamily of Anisembiidae Davis 1940c; type genus: Chorisembia

Ross 2003b.

Genus Chorisembia Ross 2003 : NE.

Chorisembia Ross 2003b:45; as genus of Chorisembiinae Ross 2003b; type species: Chorisembia howdeni Ross 2003b by original designation.

Subfamily Cryptembiinae Ross 2003 : NE.

Cryptembiinae Ross 2003b:48; as subfamily of Anisembiidae Davis 1940c; type genus: Cryptembia Ross

2003b.

Genus Cryptembia Ross 2003 : NE.

Cryptembia Ross 2003b:49; as genus of Cryptembiinae Ross 2003b; type species: Cryptembia amazonica Ross 2003b by original designation.

Subfamily Platyembiinae Ross 2003 : NE.

Platyembiinae Ross 2003b:45; as subfamily of Anisembiidae Davis 1940c; type genus: Platyembia Ross

2003b.

Genus Platyembia Ross 2003 : NE.

Platyembia Ross 2003:46; as genus of Platyembiinae Ross 2003b; type species: Platyembia tessellata


Subfamily Scolembiinae Ross 2003 : NE.

Scolembiinae Ross 2003b:38; as subfamily of Anisembiidae Davis 1940c; type genus: Scolembia Ross

2003b.

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Genus Scolembia Ross 2003 : NE.

Scolembia Ross 2003b:39; as genus of Scolembiinae Ross 2003b; type species: Scolembia celata Ross

2003b by original designation.

Anisembiidae, subfamily incertae sedis .

Genus Microembia Ross 1944 : NE.

Microembia Ross 1944:416; as genus of Embiidae Burmeister 1839; type species: Microembia rugosifrons Ross 1944 by original designation; transferred to Anisembiidae Davis 1940c, subfamily incertae sedis by Szumik (2004).

Family Archembiidae Ross 2001 : AF NA NE.

Archembiinae Ross 2001:3; as subfamily of Embiidae Burmeister 1839, elevated to family rank by

Szumik (2004); type genus: Archembia Ross 1971.

Scelembiinae Ross 2001:24; as subfamily of Embiidae Burmeister 1839; type species: Scelembia Ross

1960 (= Rhagadochir Enderlein 1912); synonymy by Szumik (2004).

Pachylembiinae Ross 2001:81; as subfamily of Embiidae Burmeister 1839; type species: Pachylembia

Ross 1984a; synonymy by Szumik (2004).

Genus Ambonembia Ross 2001 : NE.

Ambonembia Ross 2001:66; as genus of Scelembiinae Ross 2001; type species: Ambonembia incae


Ischnosembia Ross 2001:32; as genus of Scelembiinae Ross 2001; type species: Ischnosembia amazonica Ross 2001 by original designation; synonymy by Szumik (2004).

Genus Archembia Ross 1971 : NE.

Archembia Ross 1971:30; as genus of Embioptera (original family-group not indicated); type species:

Archembia lacombea Ross 1971 (= Embia kotzbaueri Navás 1925) by original designation.

Genus Biguembia Szumik 1997 : NE.

Biguembia Szumik 1997:149; as genus of Embiidae Burmeister 1839; type species: Biguembia copo

Szumik 1997 by original designation.

Aphanembia Ross 2001:64; as genus of Scelembiinae Ross 2001; type species: Aphanembia obscura

Ross 2001 by original designation; synonymy by Szumik (2004).

Genus Calamoclostes Enderlein 1909 : NE.

Calamoclostes Enderlein 1909:188; as genus of Embiinae Burmeister 1839; type species:

Calamoclostes albistriolatus Enderlein 1909 by original designation.

Genus Conicercembia Ross 1984 : NE.

Conicercembia Ross 1984a:10; as genus of Embiidae Burmeister 1839; type species: Conicercembia tepicensis Ross 1984a by original designation.


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Genus Dolonembia Ross 2001 : NE.

Dolonembia Ross 2001:30; as genus of Scelembiinae Ross 2001; type species: Dolonembia tapirape


Genus Ecuadembia Szumik 2004 : NE.

Ecuadembia Szumik 2004:226; as genus of Archembiidae Ross 2001; type species: Archembia arida


Genus Embolyntha Davis 1940 : NE.

Olyntha Gray in Griffith and Pidgeon 1832:347; as subgenus of Embia Latreille 1825; type species:

Embia brasiliensis Gray in Griffith and Pidgeon 1832 by monotypy; preoccupied by Hübner 1818

(see Discussion below).

Embolyntha Davis 1940d:344; as replacement name for Olyntha Gray in Griffith and Pidgeon 1832; type species: Olyntha brasiliensis Gray in Griffith and Pidgeon 1832 according to Article 67.8.

Argocercembia Ross 2001:63; as genus of Scelembiinae Ross 2001; type species: Argocercembia guyana Ross 2001 by original designation; synonymy by Szumik (2004).

Genus Gibocercus Szumik 1997 : NE.

Gibocercus Szumik 1997:141; as genus of Embiidae Burmeister 1839; type species: Gibocercus chaco


Subgenus Amazonembia Ross 2001 : NE.

Amazonembia Ross 2001:39; as subgenus of Gibocercus Szumik 1997; type species: Amazonembia sandrae Ross 2001 by original designation.

Subgenus Gibocercus Szumik 1997 : NE.

Gibocercus Szumik 1997:141; as genus of Embiidae Burmeister 1839; type species: Gibocercus chaco


Genus Litosembia Ross 2001 : NE.

Litosembia Ross 2001:58; as genus of Scelembiinae Ross 2001; type species: Litosembia oligembioides


Genus Malacosembia Ross 2001 : NE.

Malacosembia Ross 2001:70; as genus of Scelembiinae Ross 2001; type species: Malacosembia tucumana Ross 2001 by original designation.

Genus Neorhagadochir Ross 1944 : NE.

Neorhagadochir Ross 1944:418; as genus of Embiidae Burmeister 1839; type species: Neorhagadochir inflata Ross 1944 by original designation.

Brachypterembia Ross 1984a:7; as genus of Embiidae Burmeister 1839; type species:

Brachypterembia moreliensis Ross 1984a by original designation; synonymy by Szumik (2004).

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Subgenus Drepanembia Ross 1984 : NE.

Drepanembia Ross 1984a:6; as subgenus of Neorhagadochir Ross 1944; type species: Embia salvini

McLachlan 1877 by original designation.

Subgenus Neorhagadochir Ross 1944 : NE.

Neorhagadochir Ross 1944:418; as genus of Embiidae Burmeister 1839; type species: Neorhagadochir inflata Ross 1944 by original designation.

Genus Ochrembia Ross 2001 : NE.

Ochrembia Ross 2001:73; as genus of Scelembiinae Ross 2001; type species: Embia wagneri Navás

1923 by original designation.

Genus Pachylembia Ross 1984 : NE.

Pachylembia Ross 1984a:13; as genus of Embiidae Burmeister 1839; type species: Pachylembia chapalae Ross 1984a by original designation.

Genus Pararhagadochir Davis 1940 : NE.

Pararhagadochir Davis 1940b:181; as genus of Embioptera (original family-group not indicated); type species: Embia trinitatis de Saussure 1896 by original designation.

Genus Rhagadochir Enderlein 1912 : AF.

Rhagadochir Enderlein 1912:54; as genus of Embiidinae Burmeister 1839; type species: Embia vosseleri Enderlein 1909 by original designation (see Discussion below).

Scelembia Ross 1960:300; as genus of Embioptera (original family-group not indicated); type species:

Rhagadochir malkini Ross 1952a by original designation; synonymy by Szumik (2004).

Genus Xiphosembia Ross 2001 : NE.

Xiphosembia Ross 2001:28; as genus of Scelembiinae Ross 2001; type species: Xiphosembia amapae


Family Australembiidae Ross 1963 : AU.

Australembiidae Ross 1963:124; as family of Embioptera; type genus: Australembia Ross 1963.

Genus Australembia Ross 1963 : AU.

Australembia Ross 1963:128; as genus of Australembiidae Ross 1963; type species: Australembia incompta Ross 1963 by original designation.

Genus Metoligotoma Davis 1936 : AU.

Metoligotoma Davis 1936:248; as genus of Oligotomidae Enderlein 1909; type species: Metoligotoma reducta Davis 1936 by original designation.

Family Clothodidae Enderlein 1909 : NE.

Clothodinae Enderlein 1909:175; as subfamily of Embiidae Burmeister 1839, elevated to family by Davis

(1940a); type genus: Clothoda Enderlein 1909.

Note . Erroneously proposed again as a new family under Clothodidae by Tillyard 1937:251, but invalidly as the type genus is the same.


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Genus Antipaluria Enderlein 1912 : NE.

Antipaluria Enderlein 1912:63; as genus of Embiidinae Burmeister 1839; type species: Antipaluria aequicercata Enderlein 1912 by original designation.

Genus Chromatoclothoda Ross 1987 : NE.

Chromatoclothoda Ross 1987:26; as genus of Clothodidae Enderlein 1909; type species:

Chromatoclothoda elegantula Ross 1987 by original designation.

Genus Clothoda Enderlein 1909 : NE.

Clothoda Enderlein 1909:176; as genus of Clothodinae Enderlein 1909; type species: Embia nobilis

Gerstäcker 1888 by original designation.

Genus Cryptoclothoda Ross 1987 : NE.

Cryptoclothoda Ross 1987:17; as genus of Clothodidae Enderlein 1909; type species: Cryptoclothoda spinula Ross 1987 by original designation.

Family Embiidae Burmeister 1839 : AF PA OR.

Embiidae Burmeister 1839:768, as family (“Embidae”) of Tribus Corrodentia; type genus: Embia Latreille

1925.

Genus Acrosembia Ross 2006 : AF.

Acrosembia Ross 2006b:307; as genus of Embiidae Burmeister 1839; type species: Acrosembia lichenophila Ross 2006 by original designation.

Genus Apterembia Ross 1957 : AF.

Apterembia Ross 1957:81; as replacement name for Dictyoploca Krauss 1911, nec Jordan 1911; type species: Dictyoploca cercocyrta Krauss 1911 by original designation under Rule (f), Article 30 (as of 1959).

Dictyoploca Krauss 1911:54; as genus of Embiidae Burmeister 1839; type species: Dictyoploca cercocyrta Krauss 1911 by original designation; suppressed in Opinion 545 of the International

Commission of Zoological Nomenclature (ICZN 1957).

Genus Arabembia Ross 1981 : PA.

Arabembia Ross 1981:203; as genus of Embiidae Burmeister 1839; type species: Arabembia biarmata


Genus Berlandembia Davis 1940 : AF.

Berlandembia Davis 1940b:191; as replacement name for Berlandiella Davis 1939e; type species:

Embia berlandi Navás 1922 according to Article 67.8.

Berlandiella Davis 1939e:567; as genus of Embioptera (original family-group not indicated); type species: Embia berlandi Navás 1922 by original designation; preoccupied by Mello Leitao 1929.

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Genus Chirembia Davis 1940 : AF.

Chirembia Davis 1940b:175; as genus of Embioptera (original family-group not indicated); type species: Embia xanthocera Navás 1930 by original designation.

Navasiella Davis 1940b:177; as genus of Embioptera (original family-group not indicated); type species: Oligotoma sulcata Navás 1923 by original designation; synonym by Ross (2006).

Genus Cleomia Stefani 1953 : PA.

Cleomia Stefani 1953:93; as genus of Embiidae Burmeister 1839; type species: Cleomia guareschi

Stefani 1953 by original designation.

Genus Dihybocercus Enderlein 1912 : AF.

Dihybocercus Enderlein 1912:109; as subgenus of Embia Latreille 1825, elevated to genus rank by

Navás (1931); type species: Embia severini Enderlein 1912 by original designation.

Genus Dinembia Davis 1939 : AF.

Dinembia Davis 1939e:559; as genus of Embioptera (original family-group not indicated); type species: Dinembia ferruginea Davis 1939e by original designation.

Genus Donaconethis Enderlein 1909 : AF.

Donaconethis Enderlein 1909:176; as genus of Embiinae Burmeister 1839; type species: Donaconethis abyssinica Enderlein 1909 by original designation.

† Genus Electroembia Ross 1956 : AF.

Electroembia Ross 1956:77; as genus of Embiidae Burmeister 1839; type species: Embia antiqua Pictet

1854 by original designation.

Genus Embia Latreille 1825 : AF PA OR.

Embia Latreille 1825:437; as group of Termitines.

Termitinae ; type species: Embia savignii Westwood

1837 by subsequent monotypy.

Euembia Verhoeff 1904:201; as subgenus of Embia Latreille 1825; type species: Embia savignii

Westwood 1837 by current designation (see Discussion below); synonymy by Davis (1940d).

Monotylota Enderlein 1909:188; as genus of Embiinae Burmeister 1839; type species: Embia ramburi

Westwood 1837 by original designation; synonymy by Davis (1940d).

Genus Enveja Navás 1916 : AF.

Enveja Navás 1916:23; as genus of Embiidae Burmeister 1839; type species: Enveja bequaerti Navás

1916 by monotypy.

Genus Leptembia Krauss 1911 : AF.

Leptembia Krauss 1911:71; as genus of Embiidae Burmeister 1839; type species: Leptembia hamifera

Krauss 1911 by original designation.

Genus Machadoembia Ross 1952 : AF.

Machadoembia Ross 1952a:43; as genus of Embiidae Burmeister 1839; type species: Machadoembia angolica Ross 1952a by original designation.


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Genus Macrembia Davis 1940 : AF.

Macrembia Davis 1940b:179; as genus of Embioptera (original family-group not indicated); type species: Embia lunaris Navás 1926 by original designation.

Genus Metembia Davis 1939 : PA.

Metembia Davis 1939d:474; as genus of Embioptera (original family-group not indicated); type species: Metembia ferox Davis 1939d by original designation.

Genus Odontembia Davis 1939 : AF.

Odontembia Davis 1939a:489; as genus of Embioptera (original family-group not indicated); type species: Dihybocercus spinosus Navás 1931 by original designation.

Genus Oedembia Ross 2007 : OR.

Oedembia Ross 2007:576; as genus of Embiidae; type species: Oedembia dilatamenta Ross 2007 by original designation.

Genus Parachirembia Davis 1940 : AF.

Parachirembia Davis 1940b:178; as genus of Embioptera (original family-group not indicated); type species: Embia apicata Silvestri 1921 by original designation.

Genus Parembia Davis 1939 : PA.

Parembia Davis 1939a:478; as genus of Embioptera (original family-group not indicated); type species: Parembia valida Davis 1939a by original designation (see Discussion below).

Genus Parthenembia Ross 1960 : AF.

Parthenembia Ross 1960:298; as genus of Embiidae Burmeister 1839; type species: Parthenembia reclusa Ross 1960 by original designation.

Genus Pseudembia Davis 1939 : PA.

Pseudembia Davis 1939d:476; as genus of Embioptera (original family-group not indicated); type species: Pseudembia paradoxa Davis 1939d by original designation.

Family Embonychidae Navás 1917 : OR.

Embonychidae Navás 1917:16; as family of Embioptera; type genus: Embonycha Navás 1917.

Genus Embonycha Navás 1917 : OR.

Embonycha Navás 1917:15; as genus of Embonychidae Navás 1917; type species: Embonycha interrupta Navás 1917 by original designation.

Family Notoligotomidae Davis 1940 : AU OR.

Notoligotomidae Davis 1940c:536; as family of Embioptera; type genus: Notoligotoma Davis 1936.

Subfamily Notoligotominae : AU OR.

Notoligotomidae Davis 1940c:536; as family of Embioptera; type genus: Notoligotoma Davis 1936.

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Genus Notoligotoma Davis 1936 : AU.

Notoligotoma Davis 1936:244; as genus of Oligotomidae Enderlein 1909; type species: Oligotoma hardyi Friederichs 1914 by original designation.

Genus Ptilocerembia Friederichs 1923 : OR.

Ptilocerembia Friederichs 1923:24; as genus of Embiidae Burmeister 1839; type species:

Ptilocerembia roepkei Friederichs 1923 by monotypy.

† Subfamily Burmitembiinae Engel and Grimaldi 2006 : OR.

Burmitembiinae Engel and Grimaldi 2006:7; as subfamily of Notoligotomidae Davis 1940c; type genus:

Burmitembia Cockerell 1919.

† Genus Burmitembia Cockerell 1919 : OR.

Burmitembia Cockerell 1919:194; as genus of Oligotominae Enderlein 1909; type species:

Burmitembia venosa Cockerell 1919 by monotypy.

Family Oligotomidae Enderlein 1909 : AU AF NA NE PA OR.

Oligotomidae Enderlein 1909:175; as family of Embiidina; type genus: Oligotoma Westwood 1837.

Genus Aposthonia Krauss 1911 : AU PA OR.

Aposthonia Krauss 1911:48; as genus of Oligotomidae Enderlein 1909, type species: Aposthonia vosseleri Krauss 1911 (= Oligotoma borneensis Hagen 1885) by original designation.

Genus Bulbosembia Ross 2007 : OR.

Bulbosembia Ross 2007:594; as genus of Oligotomidae Enderlein 1909, type species: Bulbosembia thailandica Ross 2007 by original designation.

Genus Eosembia Ross 2007 : OR.

Eosembia Ross 2007:578; as genus of Oligotomidae Enderlein 1909, type species: Eosembia nepalica


Genus Haploembia Verhoeff 1904 : NA PA.

Haploembia Verhoeff 1904:201; as subgenus of Embia Latreille 1825, elevated to genus by Enderlein

(1909); type species: Embia solieri Rambur 1842 by monotypy.

Dityle Friederichs 1907:272; as subgenus of Embia Latreille 1825; type species: Embia solieri Rambur

1842 by subsequent designation (type species subsequently fixed by Enderlein (1909) according to

Article 69.1.1); synonymy by Enderlein (1909).

Gynembia Ross 1940b:664; as genus incertae sedis with respect to family; type species: Gynembia tarsalis Ross 1940b (= Embia solieri Rambur 1842) by original designation; synonym by Stefani

(1955).

Genus Lobosembia Ross 2007 : OR.

Lobosembia Ross 2007:594; as genus of Oligotomidae Enderlein 1909, type species: Lobosembia mandibulata Ross 2007 by original designation.


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Genus Oligotoma Westwood 1837 : AU AF NA NE PA OR.

Oligotoma Westwood 1837:373; as subgenus of Embia Latreille 1825, elevated to genus by Burmeister

(1839); type species: Embia saundersii Westwood 1837 by monotypy.

Family Paedembiidae Ross 2006 : PA.

Paedembiidae Ross 2006a:786; as family of Paedembiamorpha; type genus: Paedembia Ross 2006a.

Genus Badkhyzembia Gorochov and Anisyutkin 2006 : PA.

Badkhyzembia Gorochov and Anisyutkin 2006:795; as genus of Paedembiidae Ross 2006a; type species: Badkhyzembia krivokhatskyi Gorochov and Anisyutkin 2006:795 by original designation.

Genus Paedembia Ross 2006 : PA.

Paedembia Ross 2006a:786; as genus of Paedembiidae Ross 2006a; type species: Paedembia afghanica Ross 2006a by original designation.

† Family Sorellembiidae Engel and Grimaldi 2006 : OR.

Sorellembiidae Engel and Grimaldi 2006:4; as family of Embiodea; type genus: Sorellembia Engel and

Grimaldi 2006.

† Genus Sorellembia Engel and Grimaldi 2006 : OR.

Sorellembia Engel and Grimaldi 2006:5; as genus of Sorellembiidae Engel and Grimaldi 2006; type species: Sorellembia estherae Engel and Grimaldi 2006 by original designation.

Family Teratembiidae Krauss 1911 : AF NA NE.

Teratembiidae Krauss 1911:33; as family of Embiidina; type genus: Teratembia Krauss 1911.

Oligembiidae Davis 1940c:536; as family of Embioptera; type genus: Oligembia Davis 1939b; synonymy by Ross (1952a).

Genus Diradius Freiderichs 1934 : NA NE.

Diradius Freiderichs 1934:419; as genus of Oligotominae Enderlein 1909; type species: Diradius pusillus Freiderichs 1934 by monotypy.

Dilobocerca Ross 1944:476; as subgenus of Oligembia Davis 1939b; type species: Oligembia lobatus

Ross 1944 by original designation; synonymy by Ross (1984b).

Genus Oligembia Davis 1939 : NA NE.

Oligembia Davis 1939b:217; as genus of Embioptera (original family-group not indicated); type species: Oligotoma hubbardi Hagen 1885 by original designation.

Genus Paroligembia Ross 1952 : AF.

Paroligembia Ross 1952a:50; as genus of Teratembiidae Krauss 1911; type species: Paroligembia angolica Ross 1952a by original designation.

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Genus Teratembia Krauss 1911 : NE.

Teratembia Krauss 1911:33; as genus of Tertembiidae Krauss 1911; type species: Teratembia geniculata Krauss 1911 by original designation.

Idioembia Ross 1944:455; as genus of Oligembiidae Davis 1940c; type species: Oligembia banksi

Davis 1939b by original designation; synonymy by Ross (1952b).

Family incertae sedis

Genus Condylopalama Sundevall 1847:255 : NE?.

Condylopalama Sundevall 1847:255; as genus of Dermaptera; type species: Condylopalama agilis

Sundevall 1847 by monotypy (see Discussion below).

Dicussion

Review of early literature and Embia Latreille 1825, date of description, spelling and type species

Early Embioptera literature presents some of the most complicated and problematic nomeclatural issues in the order and requires detailed discussion and careful interpretation. Although Davis (1940d) provides the most thorough account of some of the earliest literature and addresses, clarifies and solves several issues, there are still several problems to remedy and his treatment of them is inadequate in some cases.

Authorship and date of publication of Embia Latreille 1825 . The first published reference to include insects now regarded as Embioptera was two figures by Savigny (1809-1813) in the Description de l’Égypte and placed near Termes Linnaeus (Isoptera) (the preparation of which apparently caused Savigny to eventually go blind (Westwood 1837)). No taxonomic name was introduced or otherwise applied to the illustrated insect. In 1825, Latreille published the name Embie for an insect similar to Termes , “… mais à antennes différentes ” (Latreille 1825). There were no illustrations, references to illustrations or specific names included in this publication either, but it seems likely that Latreille relied on Savigny’s illustrations as evidence for these insects. Audoin (1827) subsequently used the name Embie directly in reference to

Savigny’s figure. The same year, Berthold (1827) used a Latinized form of the name, “ Embium .”

Subsequently, Latreille (1829) published a more clear reference to Savigny’s illustrations together with some description and observations about the insects with a Latinized form ( Embia ) of the original name. Unlike authors contemporary with Latreille (e.g. Westwood 1837) Davis (1940d) attributes the authorship and date of publication of Embie (and Embia , see below) to Latreille (1829), not Latreille (1825). Davis (1940d) regarded

Latreille’s (1825) original name, Embie , as vernacular, and, therefore, without standing. However, according to Article 11.3, this does not, by itself, invalidate Latreille’s name. Article 1.3.5 (which excludes names from the provisions of the Code intended “…as means of temporary reference and not for formal taxonomic use as scientific names in zoological nomenclature”) might possibly be invoked, though given Latreille’s explicit reference to the (italicized) name on par with Termes Linnaeus, an available type genus name for Termitidae

(Isoptera), this does not seem to be a valid explanation. Even apart from questions of validity of vernacular names, Davis (1940d) also regarded the name as without description or proper indication, and, therefore, unavailable (Article 12.1). He apparently believed Latreille’s statement separating Embie from Termes on the basis of “different antennae” was insufficient to count as a description or definition, but this seems to be a subjective judgement. After all, a taxon explicitly regarded as the same as Termes (and included together in the description of Termitines.

Termitinae ) but with the antennae not the same as in that taxon does constitute a description in a way that mere inclusion of a name in a list or index does not. If Latreille (1825) is not the proper source of the name, then Audoin (1827), by reference to the figure in Savigny (1809-1813) should be the author of the name by indication (again, the use of the vernacular notwithstanding). But I disagree with

Davis’s (1940d) interpretation and consider that Latreille (1825) first made available the name Embie (for correct spelling, see below).


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Original spelling of Embia Latreille 1825 . A noteworthy problem, however, is the correct original spelling of the generic name introduced as the vernacular Embie by Latreille (1825). The first Latinized form of the name was by Berthold (1827) who used Embium . The next was by Latreille (1829), who used Embia .

The next was by Gray in Griffith and Pidgeon (1832) (see discussion of date and authorship below), who used

Embius . The Code is noticeably vague about what to do in instances of variations in Latinization of originally vernacular names. Although the Code recommends emending a vernacular name to a Latinized form

(Recommendation 11A), it is not required. However, the Code also indicates that, “The original spelling of a name is the “correct original spelling”, unless it is demonstrably incorrect as provided in Article 32.5.”

Article 32.5 maintains that inadvertent errors (among other clearly non-applicable items) must be corrected, but that, “Incorrect transliteration or Latinization…” is not to be considered an inadvertent error. Article 32.3

states that “The correct original spelling of a name is to be preserved unaltered, except where it is mandatory to change the suffix or the gender ending under Article 34,” but mandatory changes (Article 34) apply only to family-group (suffix) and species-group (to match genus gender) names. The alteration of the original Embie does not appear to be an emendation according to Article 33.2 which requires an explicit statement of intention, citation of each name combined with selection of the alternative spelling or similar treatment of other names. In this case the first two criteria do not appear to apply, and the third is difficult to demonstrate.

At a minimum, there is some doubt as to whether these are emendations or incorrect subsequent spellings, suggesting that Article 33.5 applies, “In any case of doubt whether a different subsequent spelling is an emendation or an incorrect subsequent spelling, it is to be treated as an incorrect subsequent spelling (and therefore unavailable), and not as an emendation.” In any case, this would seem to suggest that the correct original spelling of the genus is Embie , and not the subsequent alterations ( Embium , Embius , Embia ) which are each attempts at Latinizations. However, according to Article 33.2.3.1, “…when an unjustified emendation is in prevailing usage and is attributed to the original author and date it is deemed to be a justified emendation.” Alternatively, Article 33.3.1 states that the prevailing usage, even if incorrect, is to be considered the correct original spelling. Given the current state of knowledge, it would appear that this second article applies; Embia has been in prevailing usage for well over a century, and generally (though not always) attributed to Latreille 1825 (e.g. Ross 1966). Therefore, I conclude that Embia is the correct original spelling according to Article 33.3.1 (or, alternatively, a justified emendation by appeal to Article 33.2.3.1).

It should be noted that Davis (1940d) regarded Embius Gray in Griffiths and Pidgeon (see discussion of authorship below) as a homonym of Embia Latreille by appeal to Opinion 115 of the International

Commission of Zoological Nomenclature (ICZN 1931). It is not, technically, a homonym according to either

Articles 33.2.3 or 33.3.1. Therefore, Embius Gray in Griffiths and Pidgeon, and Embium Berthold are each incorrect subsequent spellings, and nomina nuda .

Type species of Embia Latreille 1825 . Up to and including Latreille (1829), no species names yet existed in combination with Embia (or Embioptera). This is not problematic for availability of the uninomial genus name (Article 69). The first instance of a species name associated with Embioptera was in the two volumes of

Cuvier’s The Animal Kingdom (Griffith & Pidgeon 1831;1832). In the second insect volume (volume 15) the species Embia ( Olyntha ) brasiliensis Gray in Griffith and Pidgeon was established (see below for discussion of authorship). Also in this publication, a plate is bound with the caption reading “ Embius? brasiliensis G.R.

Gray.” This plate potentially initiates a problem for nomenclature relating to the correct type species of

Embia . In reference to the plate (Plate 72, which is bound opposite page 337 in volume two; L. Stewart,

Mann Library, Cornell University, personal communication) Embius (= Embia ) brasiliensis became available by indication (Article 12.2.7) and is therefore the type species for Embia by subsequent monotypy (Article

69.3). In reference to the text, the name erected for the species in question is Embia ( Olyntha ) brasiliensis and this species would, therefore, be the type for Olyntha , not Embia . Based on this, it would appear that the same type species was simultaneously designated for both “ Embius ” (= Embia Latreille, see above) and Embia

( Olyntha ) Gray in Griffith and Pidgeon (= Embolyntha Davis). Two additional problems implied by the plate caption and the text are discussed in detail below.

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The caption to the figure on plate 72 reads “ Embius? brasiliensis G.R. Gray.” Davis (1940d:323) proposed that the question mark likely referred to uncertainty of the correct Latin ending for Latreille’s (1825)

“ Embie ,” and an alternative to Latreille’s (1829) Embia rather than any uncertainty about identification given that there was no other genus with which to confuse it (in 1831, the year he believed the plate published, see below). This would have no change on interpreting brasiliensis as the type of Embia Latreille (see above).

However, an alternative interpretation is that Gray, the author of the new taxon depicted on the plate (see below, Cowan 1969; Evenhuis 1997), believed the species to belong to a different genus than the known one described, a scenario supported by its subsequent placement in Embia ( Olyntha ) (Griffith & Pidgeon 1832)).

Because of this, it might be interpreted that “ Embius ?

brasiliensis ” indicated doubtful placement of the species in “ Embius ”, possibly disqualifying it as type species of Embia as not an originally included species

(Article 67.2.5). However, this appears contradicted by the criterion of availability of species names according to Article 11.9.3.4 which indicates that, “The generic combination, although it must be unambiguous, can be tentative.” It seems most likely that the plate was prepared (and possibly published, see

Davis (1940d) and below) prior to the text, and the species name was unambiguously, if tentatively, placed in the genus, despite a later change of opinion about its generic placement. Thus, it appears that the nominal species became available within a context in which there was only a single nominal genus, regardless of whether the ambiguity (as indicated by the question mark) was due to uncertainty about the Latinized ending or the placement of the species in “ Embius ”.

This scenario is further complicated by Davis’s (1940d) assertion that the plate (drawn by Westwood

(Davis 1940d:323) but with the name credited to Gray (Cowan 1969; Evenhuis 1997; Griffith & Pidgeon

1832)) was issued in 1831. Of the two volumes on insects, the first (volume 14) was apparently published serially in three parts in 1831 and the second (volume 15) in four parts in 1832 (Cowan 1969; Evenhuis 1997).

It is not clear why Davis (1940d) believed the plate to be issued in 1831, rather than along with the text in

1832, but he is emphatic about this, and he does regard “ Embius Griffith and Pidgeon 1831 [sic]” as a homonym of Embia Latreille (see above). He (Davis 1940d:323) comments also that the name was corrected in the second volume by Griffith and Pidgeon (1832) to Embia , again implying that the plate originated in

1831. He then (Davis 1940d:324) further comments that, “It would be misleading to accept brasiliensis

Griffith and Pidgeon 1831 [sic] as the genotype (by monotypy) of Embius ( Embia homonym)…” Thus, it would appear that Davis (1940d) in fact regarded “ Embius ?

brasiliensis ” as having been published in the year prior to Gray’s (in Griffith & Pidgeon 1832) proposing the name Embia ( Olyntha ) brasiliensis . Given that the currently rare two volumes were issued in a series and subsequently bound, it is certainly possible that plate

72 was issued in 1831 rather than 1832, with which it was eventually bound. As evidence for this, Evenhuis

(1997) and Cowan (1969; 1971) show that the bound version of the first volume, although labeled as published in “1832” is known to have been disseminated in 1831. It certainly seems unlikely that the authors or editors (Gray, Griffiths and Pidgeon) would have published the figure caption simultaneously with the text without the figure caption properly reflecting the text. Even so, other than Davis’s (1940d) comments

(however unambiguous) there is little evidence of this, but he may have had evidence that we no longer have available. In any case, it does not appear to change the fact that “ brasiliensis ” is the type species of Embia

Latreille by either subsequent monotypy (Article 69.3) in 1831 (despite possibly being “misleading” as suggested by Davis (1940d:324)) or during simultaneous designation of it as the type of Olyntha Gray in

Griffith and Pidgeon in 1832.

If the plate was published prior to the text as promoted by Davis (1940d), this act of typification has precedence over any subsequent act. This situation would have serious ramifications for the nomenclature of

Embioptera families since with brasiliensis as its type species, Embia would refer to different taxa than currently thought. Instead, the name would refer to taxa until now circumscribed by Embolyntha Davis


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(replacement name for Olyntha Gray in Griffith and Pidgeon nec Hübner; type species E. brasiliensis Gray in

Griffith and Pidgeon, see above).

Embolyntha would become a junior objective synonym of Embia , and the species until now referred to Embia would require a new name. Also, since Embia is the type genus for

Embiidae Burmeister, Archembiidae Ross (the taxon currently including Embolyntha ) would become a junior synonym of Embiidae and a new nominal taxon would be required for those taxa currently classified in

Embiidae.

In summary, several issues seem relevant in determining the types of Embia and Olyntha

(= Embolyntha ): 1) there is ambiguity about whether the plate and the text were published separately with the plate earlier and the text later, or were published simultaneously, 2) there is also ambiguity about the meaning of the question mark in the plate caption which could indicate doubtful placement of the species in Embius

(= Embia ), thereby disqualifying it as type, and finally, 3) if there was simultaneous designation of the type species for Embia and Olyntha (= Embolyntha ), a decision between them may be required. According to the

Principle of the First Reviser (Article 24.2.1), “When the precedence between… nomenclatural acts cannot be objectively determined, the precedence is fixed by the action of the first author citing… those… acts and selecting from them.” This appears to apply to issues 1 and 2. According to Article 24.2.2, “If… two or more nomenclatural acts… are published on the same date… the precedence of the… acts is fixed by the First

Reviser…” This appears to apply to issue 3.

The only author specifically addressing problems with type species in Embia was Davis (1940d).

However, he did not recognize the ambiguity implied in issues 1 or 2 above, and could not address issue 3 since he did not believe there was a simultaneous designation of types. Therefore, he does not appear to be the First Reviser. Given this, it seems appropriate to invoke the First Reviser Principle here in the face of ambiguity and/or uncertainty and choose (according to Recommendation 24A) the act that promotes the greatest stability. To that end, it is determined that brasiliensis is the type of Olyntha Gray in Griffith and

Pidgeon (= Embolyntha Davis) not Embius (= Embia Latreille). This is the historically recognized circumstance, and does not affect current classification.

This still leaves open the question of which is the type species of Embia Latreille. As outlined by

Davis (1940d), the first (unambiguous) association of a species name with Embia was by Westwood (1837) who erected Embia savignii Westwood, and this is the type species of Embia by subsequent monotypy

(Article 69.3).

Embia brasiliensis Gray in Griffith and Pidgeon 1832 and Olyntha Gray in Griffith and Pidgeon 1832, authorship and date of publication

The correct authorship of new insect names in Griffith and Pidgeon (Griffith & Pidgeon 1831;1832) is properly attributed to G.R. Gray, as specifically indicated by Griffith and Pidgeon (Griffith 1832 #74:784, and see Cowan 1969; Evenhuis 1997) and according to Article 50.1.1. The date of publication of Embia brasiliensis is either 1831 or 1832, but is equivocal. Based on the best data available today the year of publication is 1832 (see above under discussion of Embia ).

Euembia Verhoeff 1904, type species.

The name Euembia was erected as a subgenus of Embia Latreille 1929 to include those species of

Embia with males bearing wings (at that time excluding Embia ( Haploembia ) solieri Rambur 1942) (Verhoeff

1904). Verhoeff did not, however, designate a type species. He included 12 species of Embioptera in

Euembia including those from numerous different currently recognized genera.

Euembia has been regarded as a subjective synonym of Embia since soon after it was first recognized (Enderlein 1909;1912). Therefore, in the interest of selecting a type species best reflecting the historical interpretation of Embia Latreille 1925, I

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hereby designate as type species of Euembia Verhoeff 1904 the species Embia savignii Westwood 1837 ( new type designation ).

Australembiidae Ross 1963, status

Engel and Grimaldi (2006) recently placed Australembiidae as a subfamily in Notoligotomidae Davis.

However, in a recent paper dealing with Metoligotoma Davis, Australembiidae was returned to family status

(Miller & Edgerly 2008).

Anisembiidae Davis 1940, homonymy and authorship.

Ross (1940b) and Davis (1940c) proposed the name Anisembiidae at the family rank the same year based on the same type genus, Anisembia Krauss 1911. The date on Ross (1940b) is “December, 1940” whereas the date on Davis (1940c) is “Read 27th November, 1940.” A note by Ross (1944:434) reads, “Davis's 15-day prior use of the name Anisembiidae (1940e) [sic] was due to an unexpectedly earlier mailing date of his publication.” Nevertheless, because of this, Anisembiidae Davis, 1940c has priority. In some recent cases, however, (e.g.Ross 2003b) Anisembiidae is recognized at the family rank attributed to Davis (1940c) but at the subfamily rank attributed to Ross (Ross 1940b). According to the Principle of Coordination described in

Article 36.1, Anisembiidae Davis 1940c has priority over Anisembiidae Ross 1940b at all family-group ranks and Anisembiidae Ross 1940b is a junior homonym of Anisembiidae Davis 1940c (and likewise

Anisembiinae Ross 1940b is a junior homonym of Anisembiidae Davis 1940c).

Olynthidae Krauss 1911, status

Krauss (1911) apparently based this family on a misidentification of the species Embolyntha brasiliensis

(Gray in Griffith and Pidgeon 1832) (Davis 1940a). Ordinarily (if the name were in use) this would require action according to Article 65.2. However, according to Article 39, “the name of a family-group taxon is invalid if the name of its type genus is a junior homonym…” This name is based on the junior homonym

Olyntha Gray in Griffith and Pidgeon 1832 (= Embolyntha Davis 1940d) and cannot be used as a valid name and no action is required regarding the misidentification of the type.

Anisembiinae Davis 1940, status of tribes

Ross (2003b) attempted to establish a tribal classification of the subfamily Anisembiinae, but (as explained by Engel and Grimaldi (2006)) most of his new tribal names are not available under the Code. To be available, a new family-group name; 1) must be explicitly indicated as new (Article 16.1), 2) must cite the name of the type genus (Article 16.2), and 3) must diagnose the taxon in words, cite a reference to a published description, or be expressly proposed as a new replacement name (Articles 13.1.1-3). Most of the new tribe names proposed by Ross (2003b) fail to satisfy one or more of these conditions. As a result, they are unavailable. Within Anismebiinae, two tribal names were historically available and used in the classification by Ross (2003b), Mesembiini Ross and Anisembiini Davis. The only new tribe name proposed by Ross

(2003b) that satisfies conditions for availability set by the code is Poinarembiini Ross. The following names are each nomina nuda :

Exochosembiini Ross 2003b, nomen nudum

Isosembiini Ross 2003b, nomen nudum

Saussurembiini Ross 2003b and Saussurembini Ross 2003b, nomen nudum and nomen incorrectum since it is inappropriately formed from an available genus or is based on a non-existent genus

“ Saussurembus ”

Stenembiini Ross 2003b, nomen nudum


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Saussurembia Davis 1939 (= Saussurella Davis 1939, homonym), type species.

Saussurella Davis 1939e was recognized as a preoccupied name by Davis (1940b) and replaced with

Saussurembia Davis 1940b. According to Article 67.8, the two names have the same type species. Davis

(1939e) described Saussurella based on the species Embia ruficollis de Saussure 1896, but this was a misidentification (Ross 1992;2003b). Davis (1939e) based his new genus expressly on a specimen examined by him and identified by Navás (1924) as Oligotoma ruficollis (de Saussure).

Embia ruficollis de Saussure

1896 is actually a species now placed in Chelicerca (Ross 1992;2003b). However, when an author employs the name of a species as the type of a new genus, but expressly in the sense of a misidentification, then the author is deemed to have established a new nominal species bearing the specific name in combination with the new nominal genus (Article 11.10 of the 2000 Code, Article 11i of the 1985 Code), and that species is the type of the new genus. In other words, by basing the new genus Saussurella on an express misidentification of

Embia ruficollis ( sensu Navás (1924)) Davis (1939e) established the new species Saussurella ruficollis Davis

1939e. Ross (1992) did not adhere to Article 11.10 (Article 11i) in dealing with this issue. Instead, he erected a new name, Saussurembia davisi Ross 1992, for the species misidentified by Davis (1939e) and Navás

(1924) based on specimens collected by him in Costa Rica. Because of this, Saussurembia davisi Ross 1992 is recognized here as a junior subjective synonym of Saussurembia ruficollis (Davis 1939e) ( new synonym ).

This assumes, of course, that Ross (1992) correctly identified his holotype specimen as the same species as that misidentified by Davis (1939e), but he indicated that the specimen examined by Davis (1939e) (which is the holotype of Saussurella ruficollis Davis 1939e) was also examined by him (1992).

Oligotoma valida attributed to Hagen (1885) by Davis (1939d) and the type species of Parembia Davis

1939a.

In Hagen’s (1885) monograph of the Embioptera (“Embidina”) he mentions the name “ Oligotoma valida ” in reference to a large female Embioptera specimen. However, he makes it clear that although he had considered naming a new species, he instead regarded the specimen as a member of O. michaeli McLachlan writing, “The female, of which I possess a specimen, seems rather gigantic compared with the described male, so that I had provisionally described is as O. valida . But as McLachlan [1883:259] by the words in brackets,

‘apparently O. Michaeli [sic] McLachl.,’ accepts this determination, I have followed him not to encumber synonymy” (Hagen 1885:150).

Hagen’s statement clearly indicates his intention to regard the specimen as

O. michaeli and not a new species, O. valida . Later in the document he states that “…the supposition that the large wingless female belongs to O. Michaeli [sic] needs the support of further careful observation,” (Hagen

1885:224) clarifying, again, that he regarded the specimen to be O. michaeli and not the basis of a new species, O. valida . This is further reinforced by Hagen’s (1885) own summary of his work when he states that he recognizes 17 species and that “…I have been purposely as careful as possible in making new species. I could have enumerated six species more, against which with our actual knowledge little objection could have been made.” Since O. valida is not one of the 17 he recognizes, it is clear that it is among the six he did not enumerate. According to Article 11.5, a name must be used as valid when proposed unless it was first published as a junior synonym and subsequently made available. Under this scenario, O. valida is a nomen nudum and is (ironically) invalid. Davis (1939d) regarded Oligotoma valida Hagen 1885 as a valid name

(with the synonym Embia major Imms 1913) and used it as the type species of a new genus Parembia Davis

1939a. Ross (1950) regarded O. valida Hagen as a valid name and believed it was proposed “inadvertently.”

As Hagen (1885) made it clear that he did not intend the name to be valid, each of these interpretations is incorrect.

Davis (1939d) believed Hagen (1885) proposed the name O. valida , but simultaneously used it as a synonym of O. michaeli . According to Article 11.6.1 a name proposed as a junior synonym but treated before

1961 as an available name and adopted as the name of a taxon is thereby made available and dates from the

22


first publication as a synonym (i.e. Hagen (1885)). Under this scenario, O. valida Hagen 1885 may have been made available by the actions of Davis (1939d). However, this still requires the original name to be explicitly proposed as the name of a taxon, whereas Hagen (1885 ) explicitly stated just the opposite, that his name did not apply to a taxon. Under this scenario, in my opinion, O. valida Hagen 1885 remains a nomen nudum .

It is also possible that one could regard O. valida as a name published conditionally. According to Article

15.1, a new name proposed conditionally before 1961 may be available. There is little reason to believe, however, that Hagen (1885) even proposed his name conditionally or with doubt or reservation. Instead, it seems clear that he had no intention of publishing the name as applied to any taxon at all. Under each of these scenarios it seems clear that Oligotoma valida Hagen 1885 is a nomen nudum .

This still leaves the problem of what the type species is of Parembia Davis 1939a and the status and authorship of Parembia valida . Given that Davis (1939d) recognized the combination Parembia valida and referred to descriptions by Imms (1913) (for Embia major Imms) and Hagen (1885 ), this name in this combination is an available name for the species attributable to Davis (1939d). However, it is also a synonym of Embia major Imms according to Davis (1939d) and Ross (1950). Therefore, the valid name of the species is Parembia major (Imms 1913) with the invalid subjective synonym Parembia valida Davis 1939a. The type species of Parembia is P. valida Davis 1939a (= Parembia major Imms 1913).

Pogonembia Ross 2003, spelling of type species.

Two spellings of the type species of Pogonembia Ross 2003b were presented in the original descriptions of the genus and species (Ross 2003b), P. motaguae (in the description of the species) and P. motaquae (in the type species designation of the genus). It seems clear that the intended spelling was P. motaguae since it was named for the Rio Motagua Valley, Guatemala. For this reason, P. motaguae is the correct original spelling and P. motaquae is an incorrect original spelling according to Articles 19.3 and 24.2.3.

Rhagadochir Endlerlein 1912, spelling of type species

Enderlein (1909) described the species, Embia vossleri Enderlein, apparently naming it in honor of the collector (“Gesammelt von Prof. Dr. Vossler” (Enderlein 1909:182). The correct name of the collector was, however “Vosseler,” not “Vossler,” as evident in the change of spelling of the species name (“ E. vosseleri ”) and collector (“Vosseler”) by Krauss (1911) and Enderlein (1912). The change in spelling was accepted by

Davis (1940b) who explained Enderlein’s (1909) original spelling as a lapsis calami . Davis’s (1940b) interpretation may be correct, since the clear intent by Enderlein was to name the species after a misspelling of

Vosseler, which was an inadvertent error (Article 32.5.1). In this case, the spelling must be emended, which was done by Krauss (1911). Regardless, whether the emendation was justified or unjustified, the emended name E. vosseleri has been in prevailing usage attributed to Enderlein (1909) for the last century (e.g. Davis

1940b; Enderlein 1912; Krauss 1911; Szumik 2004), and can be deemed a justified emendation according to

Article 33.2.3.1. The correct species name is E. vosseleri Enderlein 1909.

Lithembia Ross 1984, status

This name is a nomen nudum since it was not made properly available (see Engel & Grimaldi 2006).

Condylopalama Sundevall 1847, status

Sundevall (1847) described Condylopalama agilis as a new genus and species of Dermaptera from an apparent Brazilian specimen based on a possible embiopteran nymph (Krauss 1899) or wingless adult (Ross

1944). The genus was subsequently placed in tentative synonymy with Olyntha Gray in Griffith and Pidgeon

(Krauss 1899) and then in synonymy with Embia Latreille, after Olyntha was synonymized with that genus

(Enderlein 1909;1912) (not followed by Krauss (1911)). Ross (1944) regarded the genus and species as incertae sedis with respect to family, where it is currently placed.


23

Genus- and Family-group names no longer associated with Embioptera.

† Clothonopsis Hong and Wang 1987:258

Original usage; Genus of Clothodidae Tillyard 1937

Notes . Transferred to Bibionidae (Diptera) by Zhang (1993).

† Epiembiodea Zalessky 1938:62

Original usage; Suborder of Embiodea.

Notes . Included Tillyardembiidae Zalessky 1938 (see above).

† Megagnathidae Tillyard 1917:128, status.

Original usage; Family of Embioptera.

Notes . Megagnathidae was described based on a species, Megagnatha odonatiformis Bolton 1917, from Late Carboniferous fossils from central France. The species was originally placed in the Perlidae

Latreille (Plecoptera), but was subsequently given its own family in the Embioptera (Tillyard 1917). Based on the illustration of the fossil, which appears to indicate a lack of expanded protarsi, and the origin of the specimen in the Carboniferous, well before an expected origin of Embioptera, it is unlikely that this species belongs in the Embioptera. According to M. Engel (personal communication) Megagnathidae is currently considered “Protorthoptera.”

† Protembiaria Tillyard 1937:243

Original usage; Suborder of Embiaria

Notes . This taxon (with a replacement name, Protembioptera Davis 1940a) is no longer regarded as

Embioptera.

† Protembiidae Tillyard 1937:243

Original usage; Family of Protembiaria (see above)

Notes . Transferred to Protorthoptera by Carpenter (1950).

† Protembioptera Davis 1940:677

Original usage; Suborder of Embioptera, replacement name for Protembiaria Tillyard 1937

Notes . No longer regarded as Embioptera (see under Protembiaria above).

† Sheimiidae Martynova 1958

Original usage; Family of Embiidina

Notes . Transferred to Protorthoptera by Carpenter (1976).

† Tillyardembiidae Zalessky 1938:64

Original usage; Family of Epiembiodea (see below)

Notes . Included species transferred to Protorthoptera by Carpenter (1950).

Alphabetical list of Embioptera genus- and family-group names

The following list includes all available Embioptera genus- and family-group names by alphabet and their current status. Nomonotypical taxa are not included.

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Acrosembia Ross 2006b, valid genus of Embiidae Burmeister 1839.

Amazonembia Ross 2001, valid subgenus of Gibocercus Szumik 1997.

Ambonembia Ross 2001, valid genus of Archembiidae Ross 2001.

Andesembia Ross 2003a, valid genus of Andesembiidae Ross 2003.

Andesembiidae Ross 2003a, valid family of Embioptera.

Anisembia Krauss 1911, valid genus of Anisembiini Davis 1940c.

Anisembiidae Davis 1940c, valid family of Embioptera.

Anisembiidae Davis 1940c, valid subfamily of Anisembiidae Davis 1940c.

Anisembiidae Ross 1940b, junior objective homonym of Anisembiidae Davis 1940c.

Anisembiini Davis 1940c, valid tribe of Anisembiinae Davis 1940c.

Antipaluria Enderlein 1912, valid genus of Clothodidae Enderlein 1909.

Aphanembia Ross 2001, junior subjective synonym of Biguembia Szumik 1997.

Aporembia Ross 2003b, valid genus of Aporembiinae Ross 2003.

Aporembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Aposthonia Krauss 1911, valid genus of Oligotomidae Enderlein 1909.

Apterembia Ross 1957, valid genus of Embiidae Burmeister 1839.

Arabembia Ross 1981, valid genus of Embiidae Burmeister 1839.

Archembia Ross 1971, valid genus of Archembiidae Ross 2001.

Archembiidae Ross 2001, valid family of Embioptera.

Argocercembia Ross 2001, junior subjective synonym of Argocercembia Ross 2001.

Australembia Ross 1963, valid genus of Australembiidae Ross 1963.

Australembiidae Ross 1963, valid family of Embioptera.

Badkhyzembia Gorochov and Anisyutkin 2006, valid genus of Paedembiidae Ross 2006a.

Berlandembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Berlandiella Davis 1939e, junior homonym replaced by Berlandembia Davis 1940b.

Biguembia Szumik 1997, valid genus of Archembiidae Ross 2001.

Brachypterembia Ross 1984a, junior subjective synonym of Neorhagadochir Ross 1944.

Brasilembia Ross 2003b, valid genus of Chelicercinae Ross 1984a.

Bryonembia Ross 2003a, valid genus of Andesembiidae Ross 2003.

Bulbocerca Ross 1940b, valid genus of Anisembiini Davis 1940c.

Bulbosembia Ross 2007, valid genus of Oligotomidae Enderlein 1909.

† Burmitembia Cockerell 1919, valid genus of Burmitembiinae Engel and Grimaldi 2006.

† Burmitembiinae Engel and Grimaldi 2006, valid subfamily of Notoligotomidae Davis 1940c.

Calamoclostes Enderlein 1909, valid genus of Archembiidae Ross 2001.

Chelicerca Ross 1940b, valid genus of Chelicercinae Ross 1984a.

Chelicercinae Ross 1984a, valid subfamily of Anisembiidae Davis 1940c.

Chirembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Chorisembia Ross 2003b, valid genus of Chorisembiinae Ross 2003b.

Chorisembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Chromatoclothoda Ross 1987, valid genus of Clothodidae Enderlein 1909.

Cleomia Stefani 1953, valid genus of Embiidae Burmeister 1839.

Clothoda Enderlein 1909, valid genus of Clothodidae Enderlein 1909.

Clothodidae Enderlein 1909, valid family of Embioptera.

Clothodidae Tillyard 1937, junior homonym of Clothodidae Enderlein 1909.

Condylopalama Sundevall 1847, valid genus of Embioptera.

Conicercembia Ross 1984a, valid genus of Archembiidae Ross 2001.

Cryptembia Ross 2003b, valid genus of Cryptembiinae Ross 2003b.

Cryptembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.


25

Cryptoclothoda Ross 1987, valid genus of Clothodidae Enderlein 1909.

Dactylocerca Ross 1940b, valid genus of Chelicercinae Ross 1984a.

Dictyoploca Krauss 1911, suppressed senior homonym replaced by Apterembia Ross 1947.

Dihybocercus Enderlein 1912, valid genus of Embiidae Burmeister 1839.

Dilobocerca Ross 1944, junior subjective synonym of Diradius Freiderichs 1934.

Dinembia Davis 1939e, valid genus of Embiidae Burmeister 1839.

Diradius Freiderichs 1934, valid genus of Teratembiidae Krauss 1911.

Dityle Friederichs 1907, junior objective synonym of Haploembia Verhoeff 1904.

Dolonembia Ross 2001, valid genus of Archembiidae Ross 2001.

Donaconethis Enderlein 1909, valid genus of Embiidae Burmeister 1839.

Drepanembia Ross 1984a, valid subgenus of Neorhagadochir Ross 1944.

Ectyphocerca Ross 2003b, valid genus of Mesembiini Ross 1940b.

Ecuadembia Szumik 2004, valid genus of Archembiidae Ross 2001.

† Electroembia Ross 1956, valid genus of Embiidae Burmeister 1839.

Embia Latreille 1825, valid genus of Embiidae Burmeister 1839.

Embiidae Burmeister 1839, valid family of Embioptera.

Embolyntha Davis 1940d, valid genus of Archembiidae Ross 2001.

Embonycha Navás 1917, valid genus of Embonychidae Navás 1917.

Embonychidae Navás 1917, valid family of Embioptera.

Enveja Navás 1916, valid genus of Embiidae Burmeister 1839.

Eosembia Ross 2007, valid genus of Oligotomidae Enderlein 1909.

Euembia Verhoeff 1904, junior objective synonym of Embia Latreille 1825.

Exochosembia Ross 2003b, valid genus of Anisembiinae Davis 1940c.

Gibocercus Szumik 1997, valid genus of Archembiidae Ross 2001.

Glyphembia Ross 2003b, valid genus of Anisembiini Davis 1940c.

Gynembia Ross 1940b, junior objective synonym of Haploembia Verhoeff 1904.

Haploembia Verhoeff 1904, valid genus of Oligotomidae Enderlein 1909.

Idioembia Ross 1944, junior subjective synonym of Teratembia Krauss 1911.

Ischnosembia Ross 2001, junior synonym of Ambonembia Ross 2001.

Isosembia Ross 2003b, valid genus of Anisembiinae Davis 1940c.

Lobosembia Ross 2007, valid genus of Oligotomidae Enderlein 1909.

Leptembia Krauss 1911, valid genus of Embiidae Burmeister 1839.

Litosembia Ross 2001, valid genus of Archembiidae Ross 2001.

Machadoembia Ross 1952a, valid genus of Embiidae Burmeister 1839.

Macrembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Malacosembia Ross 2001, valid genus of Archembiidae Ross 2001.

Mesembia Ross 1940a, valid genus of Mesembiini Ross 1940b.

Mesembiini Ross 1940b, valid tribe of Anisembiinae Davis 1940c.

Metembia Davis 1939d, valid genus of Embiidae Burmeister 1839.

Metoligotoma Davis 1936, valid genus of Australembiidae Ross 1963.

Microembia Ross 1944, valid subfamily of Anisembiidae Davis 1940c.

Microembiinae Ross 2001, junior synonym of Anisembiidae Davis 1940c.

Monotylota Enderlein 1909, junior subjective synonym of Embia Latreille 1825.

Navasiella Davis 1940b, junior subjective synonym of Chirembia Davis 1940b.

Neorhagadochir Ross 1944, valid genus of Archembiidae Ross 2001.

Notoligotoma Davis 1936, valid genus of Notoligotominae Davis 1940c.

Notoligotomidae Davis 1940c, valid family of Embioptera.

Ochrembia Ross 2001, valid genus of Archembiidae Ross 2001.

Odontembia Davis 1939a, valid genus of Embiidae Burmeister 1839.

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Oligembia Davis 1939b, valid genus of Teratembiidae Krauss 1911.

Oligembiidae Davis 1940c, junior subjective synonym of Teratembiidae Krauss 1911.

Oligotoma Westwood 1837, valid genus of Oligotomidae Enderlein 1909.

Oligotomidae Enderlein 1909, valid family of Embioptera.

Olyntha Gray in Griffith and Pidgeon 1832, junior homonym replaced by Embolyntha Davis 1940d.

Olynthidae Krauss 1911, invalid name due to homonymy of its type genus.

Oncosembia Ross 2003b, valid genus of Chelicercinae Ross 1984a.

Pachylembia Ross 1984a, valid genus of Archembiidae Ross 2001.

Pachylembiinae Ross 2001, junior synonym of Archembiidae Ross 2001.

Paedembia Ross 2006a, valid genus of Paedembiidae Ross 2006a.

Paedembiidae Ross 2006a, valid family of Embioptera.

Parachirembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Pararhagadochir Davis 1940b, valid genus of Archembiidae Ross 2001.

Parembia Davis 1939a, valid genus of Embiidae Burmeister 1839.

Paroligembia Ross 1952a, valid genus of Teratembiidae Krauss 1911.

Parthenembia Ross 1960, valid genus of Embiidae Burmeister 1839.

Pelorembia Ross 1984a, valid genus of Chelicercinae Ross 1984a.

Phallosembia Ross 2003b, valid genus of Mesembiini Ross 1940b.

Platyembia Ross 2003b, valid genus of Platyembiinae Ross 2003b.

Platyembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Pogonembia Ross 2003b, valid genus of Anisembiinae Davis 1940c.

Poinarembia Ross 2003b, valid genus of Poinarembiini Ross 2003b.

Poinarembiini Ross 2003b, valid tribe of Anisembiinae Davis 1940c.

Protochelicerca Ross 1944, junior synonym of Chelicerca Ross 1940b.

Pseudembia Davis 1939d, valid genus of Embiidae Burmeister 1839.

Ptilocerembia Friederichs 1923, valid genus of Notoligotominae Davis 1940c.

Rhagadochir Enderlein 1912, valid genus of Archembiidae Ross 2001.

Saussurella Davis 1939e, junior homonym replaced by Saussurembia Davis 1940b.

Saussurembia Davis 1940b, valid genus of Anisembiinae Davis 1940c.

Scelembia Ross 1960, junior subjective synonym of Rhagadochir Enderlein 1912.

Scelembiinae Ross 2001, junior synonym of Archembiidae Ross 2001.

Schizembia Ross 1944, valid genus of Chelicercinae Ross 1984a.

Scolembia Ross 2003b, valid genus of Scolembiinae Ross 2003b.

Scolembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

† Sorellembia Engel and Grimaldi 2006, valid genus of Sorellembiidae Engel and Grimaldi 2006.

† Sorellembiidae Engel and Grimaldi 2006, valid family of Embioptera.

Stenembia Ross 1972, valid genus of Mesembiini Ross 1940b.

Teratembia Krauss 1911, valid genus of Teratembiidae Krauss 1911.

Teratembiidae Krauss 1911, valid family of Embioptera.

Xiphosembia Ross 2001, valid genus of Archembiidae Ross 2001.

List of Embioptera family-group names in order of priority

The following list includes all available Embioptera family-group names by date of publication and their current status. Names are listed alphabetically within the same year.

Embiidae Burmeister 1839, valid family of Embioptera.


27

Clothodidae Enderlein 1909, valid family of Embioptera.

Oligotomidae Enderlein 1909, valid family of Embioptera.

Olynthidae Krauss 1911, invalid name due to homonymy of its type genus.

Teratembiidae Krauss 1911, valid family of Embioptera.

Embonychidae Navás 1917, valid family of Embioptera.

Clothodidae Tillyard 1937, junior homonym of Clothodidae Enderlein 1909.

Anisembiidae Davis 1940c, valid family of Embioptera.

Anisembiinae Davis 1940c, valid subfamily of Anisembiidae Davis 1940c.

Anisembiidae Ross 1940b, junior objective homonym of Anisembiidae Davis 1940c.

Anisembiini Davis 1940c, valid tribe of Anisembiinae Davis 1940c.

Mesembiini Ross 1940b, valid tribe of Anisembiinae Davis 1940c.

Notoligotomidae Davis 1940c, valid family of Embioptera.

Oligembiidae Davis 1940c, junior subjective synonym of Teratembiidae Krauss 1911.

Australembiidae Ross 1963, valid family of Embioptera.

Chelicercinae Ross 1984a, valid subfamily of Anisembiidae Davis 1940c.

Archembiidae Ross 2001, valid family of Embioptera.

Microembiinae Ross 2001, junior synonym of Anisembiidae Davis 1940c.

Pachylembiinae Ross 2001, junior synonym of Archembiidae Ross 2001.

Scelembiinae Ross 2001, junior synonym of Archembiidae Ross 2001.

Andesembiidae Ross 2003a, valid family of Embioptera.

Aporembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Chorisembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Cryptembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Platyembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Poinarembiini Ross 2003b, valid tribe of Anisembiinae Davis 1940c.

Scolembiinae Ross 2003b, valid subfamily of Anisembiidae Davis 1940c.

Paedembiidae Ross 2006a, valid family of Embioptera.

† Burmitembiinae Engel and Grimaldi 2006, valid subfamily of Notoligotomidae Davis 1940c.

† Sorellembiidae Engel and Grimaldi 2006, valid family of Embioptera.

List of Embioptera genus-group names in order of priority

The following list includes all available Embioptera genus-group names by date of publication and their current status. Names are listed alphabetically within the same year.

Embia Latreille 1825, valid genus of Embiidae Burmeister 1839.

Olyntha Gray in Griffith and Pidgeon 1832, junior homonym replaced by Embolyntha Davis 1940d.

Oligotoma Westwood 1837, valid genus of Oligotomidae Enderlein 1909.

Condylopalama Sundevall 1847, valid genus of Embioptera.

Euembia Verhoeff 1904, junior objective synonym of Embia Latreille 1825.

Haploembia Verhoeff 1904, valid genus of Oligotomidae Enderlein 1909.

Dityle Friederichs 1907, junior objective synonym of Haploembia Verhoeff 1904.

Calamoclostes Enderlein 1909, valid genus of Archembiidae Ross 2001.

Clothoda Enderlein 1909, valid genus of Clothodidae Enderlein 1909.

Donaconethis Enderlein 1909, valid genus of Embiidae Burmeister 1839.

Monotylota Enderlein 1909, junior subjective synonym of Embia Latreille 1825.

Anisembia Krauss 1911, valid genus of Anisembiini Davis 1940c.

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Aposthonia Krauss 1911, valid genus of Oligotomidae Enderlein 1909.

Dictyoploca Krauss 1911, suppressed senior homonym replaced by Apterembia Ross 1947.

Leptembia Krauss 1911, valid genus of Embiidae Burmeister 1839.

Teratembia Krauss 1911, valid genus of Teratembiidae Krauss 1911.

Antipaluria Enderlein 1912, valid genus of Clothodidae Enderlein 1909.

Dihybocercus Enderlein 1912, valid genus of Embiidae Burmeister 1839.

Rhagadochir Enderlein 1912, valid genus of Archembiidae Ross 2001.

Enveja Navás 1916, valid genus of Embiidae Burmeister 1839.

Embonycha Navás 1917, valid genus of Embonychidae Navás 1917.

† Burmitembia Cockerell 1919, valid genus of Burmitembiinae Engel and Grimaldi 2006.

Ptilocerembia Friederichs 1923, valid genus of Notoligotominae Davis 1940c.

Diradius Freiderichs 1934, valid genus of Teratembiidae Krauss 1911.

Metoligotoma Davis 1936, valid genus of Australembiidae Ross 1963.

Notoligotoma Davis 1936, valid genus of Notoligotominae Davis 1940c.

Berlandiella Davis 1939e, junior homonym replaced by Berlandembia Davis 1940b.

Dinembia Davis 1939e, valid genus of Embiidae Burmeister 1839.

Metembia Davis 1939d, valid genus of Embiidae Burmeister 1839.

Odontembia Davis 1939a, valid genus of Embiidae Burmeister 1839.

Oligembia Davis 1939b, valid genus of Teratembiidae Krauss 1911.

Parembia Davis 1939a, valid genus of Embiidae Burmeister 1839.

Pseudembia Davis 1939d, valid genus of Embiidae Burmeister 1839.

Saussurella Davis 1939e, junior homonym replaced by Saussurembia Davis 1940b.

Berlandembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Bulbocerca Ross 1940b, valid genus of Anisembiini Davis 1940c.

Chelicerca Ross 1940b, valid genus of Chelicercinae Ross 1984a.

Chirembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Dactylocerca Ross 1940b, valid genus of Chelicercinae Ross 1984a.

Embolyntha Davis 1940d, valid genus of Archembiidae Ross 2001.

Gynembia Ross 1940b, junior objective synonym of Haploembia Verhoeff 1904.

Macrembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Mesembia Ross 1940a, valid genus of Mesembiini Ross 1940b.

Navasiella Davis 1940b, junior subjective synonym of Chirembia Davis 1940b.

Parachirembia Davis 1940b, valid genus of Embiidae Burmeister 1839.

Pararhagadochir Davis 1940b, valid genus of Archembiidae Ross 2001.

Saussurembia Davis 1940b, valid genus of Anisembiinae Davis 1940c.

Dilobocerca Ross 1944, junior subjective synonym of Diradius Freiderichs 1934.

Idioembia Ross 1944, junior subjective synonym of Teratembia Krauss 1911.

Microembia Ross 1944, valid subfamily of Anisembiidae Davis 1940c.

Neorhagadochir Ross 1944, valid genus of Archembiidae Ross 2001.

Protochelicerca Ross 1944, junior synonym of Chelicerca Ross 1940b.

Schizembia Ross 1944, valid genus of Chelicercinae Ross 1984a.

Machadoembia Ross 1952a, valid genus of Embiidae Burmeister 1839.

Paroligembia Ross 1952a, valid genus of Teratembiidae Krauss 1911.

Cleomia Stefani 1953, valid genus of Embiidae Burmeister 1839.

† Electroembia Ross 1956, valid genus of Embiidae Burmeister 1839.

Apterembia Ross 1957, valid genus of Embiidae Burmeister 1839.

Parthenembia Ross 1960, valid genus of Embiidae Burmeister 1839.

Scelembia Ross 1960, junior subjective synonym of Rhagadochir Enderlein 1912.


29

Australembia Ross 1963, valid genus of Australembiidae Ross 1963.

Archembia Ross 1971, valid genus of Archembiidae Ross 2001.

Stenembia Ross 1972, valid genus of Mesembiini Ross 1940b.

Arabembia Ross 1981, valid genus of Embiidae Burmeister 1839.

Brachypterembia Ross 1984a, junior subjective synonym of Neorhagadochir Ross 1944.

Conicercembia Ross 1984a, valid genus of Archembiidae Ross 2001.

Drepanembia Ross 1984a, valid subgenus of Neorhagadochir Ross 1944.

Pachylembia Ross 1984a, valid genus of Archembiidae Ross 2001.

Pelorembia Ross 1984a, valid genus of Chelicercinae Ross 1984a.

Cryptoclothoda Ross 1987, valid genus of Clothodidae Enderlein 1909.

Chromatoclothoda Ross 1987, valid genus of Clothodidae Enderlein 1909.

Biguembia Szumik 1997, valid genus of Archembiidae Ross 2001.

Gibocercus Szumik 1997, valid genus of Archembiidae Ross 2001.

Amazonembia Ross 2001, valid subgenus of Gibocercus Szumik 1997.

Ambonembia Ross 2001, valid genus of Archembiidae Ross 2001.

Aphanembia Ross 2001, junior subjective synonym of Biguembia Szumik 1997.

Argocercembia Ross 2001, junior subjective synonym of Argocercembia Ross 2001.

Dolonembia Ross 2001, valid genus of Archembiidae Ross 2001.

Ischnosembia Ross 2001, junior synonym of Ambonembia Ross 2001.

Litosembia Ross 2001, valid genus of Archembiidae Ross 2001.

Malacosembia Ross 2001, valid genus of Archembiidae Ross 2001.

Ochrembia Ross 2001, valid genus of Archembiidae Ross 2001.

Xiphosembia Ross 2001, valid genus of Archembiidae Ross 2001.

Andesembia Ross 2003, valid genus of Andesembiidae Ross 2003.

Aporembia Ross 2003b, valid genus of Aporembiinae Ross 2003.

Brasilembia Ross 2003b, valid genus of Chelicercinae Ross 1984a.

Bryonembia Ross 2003a, valid genus of Andesembiidae Ross 2003.

Chorisembia Ross 2003b, valid genus of Chorisembiinae Ross 2003b.

Cryptembia Ross 2003b, valid genus of Cryptembiinae Ross 2003b.

Ectyphocerca Ross 2003b, valid genus of Mesembiini Ross 1940b.

Ecuadembia Szumik 2004, valid genus of Archembiidae Ross 2001.

Exochosembia Ross 2003b, valid genus of Anisembiinae Davis 1940c.

Glyphembia Ross 2003b, valid genus of Anisembiini Davis 1940c.

Isosembia Ross 2003b, valid genus of Anisembiinae Davis 1940c.

Oncosembia Ross 2003b, valid genus of Chelicercinae Ross 1984a.

Phallosembia Ross 2003b, valid genus of Mesembiini Ross 1940b.

Platyembia Ross 2003b, valid genus of Platyembiinae Ross 2003b.

Pogonembia Ross 2003b, valid genus of Anisembiinae Davis 1940c.

Poinarembia Ross 2003b, valid genus of Poinarembiini Ross 2003b.

Scolembia Ross 2003b, valid genus of Scolembiinae Ross 2003b.

Acrosembia Ross 2006b, valid genus of Embiidae Burmeister 1839.

Badkhyzembia Gorochov and Anisyutkin 2006, valid genus of Paedembiidae Ross 2006a.

Paedembia Ross 2006a, valid genus of Paedembiidae Ross 2006a.

† Sorellembia Engel and Grimaldi 2006, valid genus of Sorellembiidae Engel and Grimaldi 2006.

Bulbosembia Ross 2007, valid genus of Oligotomidae Enderlein 1909.

Eosembia Ross 2007, valid genus of Oligotomidae Enderlein 1909.

Lobosembia Ross 2007, valid genus of Oligotomidae Enderlein 1909.

30


Acknowledgements

Special thanks to the interlibrary loan staff at Brigham Young University and the University of New Mexico.

Also, thanks to L. Stewart and M. Schlabach of the Cornell University Library for information regarding pieces of old literature. Thanks to M. Engel for extremely productive comments. External support for this project came from NSF grant #DEB-0515924.

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