ZooKeys 148: 171–184, doi: 10.3897/zookeys.148.1682
Family-group names for termites (Isoptera), redux
Michael S. Engel 1
1 Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66049-2811, USA

Corresponding author: Michael S. Engel (msengel@ku.edu)

Academic editor: D. Grimaldi

received 8 June 2011 | accepted 29 August 2011 | Published 21 November 2011


(C) 2011 Michael S. Engel. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.


For reference, use of the paginated PDF or printed version of this article is recommended.

Abstract

Forty-eight family-group names are identified for insects among the Isoptera (termites), representing a nearly 19% increase since the last compilation less than 10 years ago. Accordingly, these names are newly catalogued, including various updates from the original summary. The name Reticulitermitidae is recognized as a nomen nudum while Caatingatermitinae is newly considered a nomen invalidum, and neither is available in zoological nomenclature. A catalogue of the suprafamilial names for Isoptera is appended. The name Xylophagodea is formally proposed for the Isoptera + Cryptocercidae clade.

Keywords

Isoptera, termites, nomenclature, classification, family-group names, type genera

Introduction

During the last 25 years numerous efforts have been undertaken to document family-group names for insects and to provide their correct authorship, date, type genus, combining stem, and availability or validity (e.g., Michener 1986; Wahl and Mason 1995; Menke 1997; Sabrosky 1999; Speidel and Naumann 2004; Engel 2005; Engel and Haas 2007; Menke et al. 2008; Miller 2009; Bouchard et al. 2011). Seven years ago such an exercise was completed for the termites (Engel and Krishna 2004a) and it is remarkable that the number of names has grown significantly for this small insect lineage, of approximately 3500 species, such that a full 19% of the names known today were not included in that original account. Accordingly, I provide an updated list, in order of priority, for all family-group names proposed for termites through to the present day. The list is an updated and corrected version of that provided by Engel and Krishna (2004a), incorporating the various new names and edits of subsequent works (e.g., Engel and Krishna 2004b, 2007; Cancello and DeSouza 2005; Engel et al. 2009). As in the earlier summary, all names are presented in their original forms, regardless of present day rank or suffix, with type genus and correct combining stem provided for each. Daggers (†) indicate names proposed for fossil taxa. The format generally follows that of Engel and Krishna (2004a).

In addition I have included here for the first time a summary of all names applied for suprafamilial groups of termites (i.e., names above the family-group ranks and not regulated by the ICZN).

Catalog of Family-group names

1. Termitina Latreille, 1802: 293. Type genus: Termes Linnaeus, 1758. Combining stem: Termit–. Note: Latreille (1805, 1810) subsequently changed the name of his “famille” to Termitinae but the name was made first available in 1802 (ICZN 1999: Art. 11.7).

2. Calotermitinae Froggatt, 1897: 516 [recte Kalotermitinae; in accordance with ICZN (1999) Art. 29.1 the name is automatically emended to Kalotermitinae Froggatt, 1897]. Type genus: Kalotermes Hagen, 1853 [Calotermes Hagen, 1858 is an unjustified emendation: vide Engel and Krishna 2001a; ICZN 2002]. Combining stem: Kalotermit–.

3. Glyptotermitinae Froggatt, 1897: 518. Type genus: Glyptotermes Froggatt, 1897. Combining stem: Glyptotermit–.

4. Rhinotermitinae Froggatt, 1897: 518. Type genus: Rhinotermes Hagen, 1858. Combining stem: Rhinotermit–.

5. Heterotermitinae Froggatt, 1897: 550. Type genus: Heterotermes Froggatt, 1897. Combining stem: Heterotermit–.

6. Mastotermitinae Desneux, 1904a: 284. Type genus: Mastotermes Froggatt, 1897. Combining stem: Mastotermit–.

7. Hodotermitini Desneux, 1904c: 284. Type genus: Hodotermes Hagen, 1853. Combining stem: Hodotermit–.

8. Stolotermitinae Holmgren, 1910a: 285. Type genus: Stolotermes Hagen, 1858. Combining stem: Stolotermit–. Note: Stolotermitinae has often been included in an expanded Termopsidae for which the latter name has been used for the combined grouping despite the priority of the name based on Stolotermes. A petition was submitted (Engel et al. 2003; Engel and Krishna 2004c) and approved (ICZN 2005) reversing priority in favor of Termopsidae whenever Termopsis and Stolotermes are placed within the same family group. However, in the present classification (Engel et al. 2009 and table 1 herein) stolotermitines are segregated into their own family.

9. Leucotermitinae Holmgren, 1910a: 285. Type genus: Leucotermes Silvestri, 1901. Combining stem: Leucotermit–.

10. Coptotermitinae Holmgren, 1910a: 285. Type genus: Coptotermes Wasmann, 1896. Combining stem: Coptotermit–. Note: Proposed again as new in Holmgren (1910b).

11. Serritermitinae Holmgren, 1910a: 285. Type genus: Serritermes Wasmann, 1897. Combining stem: Serritermit–.

12. Termitogetoninae Holmgren, 1910a: 286. Type genus: Termitogeton Desneux, 1904b. Combining stem: Termitogeton–.

13. Microcerotermitinae Holmgren, 1910b: 145. Type genus: Microcerotermes Silvestri, 1901. Combining stem: Microcerotermit–.

14. Eutermitinae Holmgren, 1910b: 146. Type genus: Eutermes Heer, 1849. Combining stem: Eutermit–. Note: Given the uncertainty in the application of the name Eutermes, and thereby the family-group name Eutermitinae, a petition to suppress proactively the name was submitted to the ICZN for consideration (Engel and Krishna 2005a) which, after discussion (Roisin 2005; Engel and Krishna 2005b), was not approved (ICZN 2007). The name is presently considered incertae sedis.

15. Termopsinae Holmgren, 1911: 35. Type genus: Termopsis Heer, 1849. Combining stem: Termops–. Note: Refer to comments provided for Stolotermitinae (vide supra).

16. Psammotermitinae Holmgren, 1911: 64. Type genus: Psammotermes Desneux, 1902. Combining stem: Psammotermit–.

17. Pseudomicrotermitinae Holmgren, 1912: 5. Type genus: Pseudomicrotermes Holmgren, 1912. Combining stem: Pseudomicrotermit–.

18. Foraminitermitinae Holmgren, 1912: 5. Type genus: Foraminitermes Holmgren, 1912. Combining stem: Foraminitermit–.

19. Stylotermitinae Holmgren and Holmgren, 1917: 141. Type genus: Stylotermes Holmgren and Holmgren, 1917. Combining stem: Stylotermit–.

20. †Pliotermitinae Pongrácz 1917: 28. Type genus: †Pliotermes Pongrácz, 1917. Combining stem: Pliotermit–.

21. Arrhinotermitinae Sjöstedt, 1926: 8. Type genus: Arrhinotermes Wasmann, 1902. Combining stem: Arrhinotermit–.

22. Acanthotermitinae Sjöstedt, 1926: 8. Type genus: Acanthotermes Sjöstedt, 1900. Combining stem: Acanthotermit–. Note: This name has priority over Macrotermitinae; however, a petition was submitted to conserve the usage of Macrotermitinae (Engel and Krishna 2001b) and was approved by the ICZN (2003). Macrotermitinae is to be used whenever Macrotermes and Acanthotermes are placed into the same family-group taxon.

23. †Miotermitinae Pongrácz, 1926: 29 [chart]. Type genus: †Miotermes Rosen, 1913. Combining stem: Miotermit–.

24. Macrotermitinae Kemner, 1934: 69. Type genus: Macrotermes Holmgren, 1909. Combining stem: Macrotermit–. Note: Refer to comments provided for Acanthotermitinae (vide supra).

25. Amitermitinae Kemner, 1934: 110. Type genus: Amitermes Silvestri, 1901. Combining stem: Amitermit–.

26. Miro-capritermitinae Kemner, 1934: 166. Type genus: Mirocapritermes Holmgren, 1914. Combining stem: Mirocapritermit–. Note: Although Kemner (1934) hyphenated the name in its original spelling, the ICZN (1999) does not allow hyphenation and the family-group name must be considered a single word.

27. Nasutitermitinae Hare, 1937. Type genus: Nasutitermes Dudley, 1890. Combining stem: Nasutitermit–.

28. †Electrotermitinae Emerson, 1942: 10. Type genus: †Electrotermes Rosen, 1913. Combining stem: Electrotermit–.

29. Porotermitinae Emerson, 1942: 10. Type genus: Porotermes Hagen, 1858. Combining stem: Porotermit–.

30. Apicotermitinae Grassé and Noirot, 1954 [1955]: 360. Type genus: Apicotermes Holmgren, 1912. Combining stem: Apicotermit–.

31. Odontotermitini Weidner, 1956: 82. Type genus: Odontotermes Holmgren, 1910a. Combining stem: Odontotermit–.

32. Cubitermitini Weidner, 1956: 99. Type genus: Cubitermes Wasmann, 1906. Combining stem: Cubitermit–.

33. Mirotermitini Weidner, 1956: 99. Type genus: Mirotermes Wasmann, 1897. Combining stem: Mirotermit–.

34. Capritermitini Weidner, 1956: 100. Type genus: Capritermes Wasmann, 1897. Combining stem: Capritermit–.

35. Indotermitidae Roonwal and Sen-Sarma In Roonwal, 1958: 81. Type genus: Indotermes Roonwal and Sen-Sarma In Roonwal, 1958. Stem: Indotermit–. Note: Proposed as new again in Roonwal and Sen-Sarma (1960).

36. †Cretatermitinae Emerson, 1967 [1968]: 278. Type genus: †Cretatermes Emerson, 1967 [1968]. Combining stem: Cretatermit–.

37. Prorhinotermitinae Quennedey and Deligne, 1975: 265. Type genus: Prorhinotermes Silvestri, 1909. Combining stem: Prorhinotermit–.

38. †Lutetiatermitinae Schlüter, 1989: 61. Type genus: †Lutetiatermes Schlüter, 1989. Combining stem: Lutetiatermit–.

39. †Carinatermitinae Krishna and Grimaldi, 2000: 134. Type genus: †Carinatermes Krishna and Grimaldi, 2000. Combining stem: Carinatermit–.

40. †Archeorhinotermitinae Krishna and Grimaldi, 2003: 2. Type genus: †Archeorhinotermes Krishna and Grimaldi, 2003. Combining stem: Archeorhinotermit–.

41. Reticulitermatidae Szalanski, Austin, and Owen 2003: 1514, nomen imperfectum [recte Reticulitermitidae] et nomen nudum. Note: This name has appeared in several publications (e.g., Sobti et al. 2009) but has not been formally established. It would represent simply a junior synonym of Heterotermitinae.

42. Syntermitinae Engel and Krishna, 2004a: 6. Type genus: Syntermes Holmgren, 1909. Combining stem: Syntermit–.

43. Sphaerotermitinae Engel and Krishna, 2004a: 6. Type genus: Sphaerotermes Holmgren, 1912. Combining stem: Sphaerotermit–.

44. Cornitermitinae Ensaf et al., 2004: 284, nomen nudum. Type genus: Cornitermes Wasmann, 1897. Combining stem: Cornitermit–.

45. Glossotermitinae Cancello and DeSouza, 2005: 35. Type genus: Glossotermes Emerson, 1950. Combining stem: Glossotermit–.

46. †Caatingatermitinae Martins-Neto, Ribeiro-Júnior, and Prezoto, 2006: 127, nomen invalidum. Note: I consider this name to be unavailable as the type genus was not explicitly indicated. ICZN (1999) Art. 16.2 requires that after 1999 all new family-group names must have the type genus precisely identified, not implied through the formation of the name. Martins-Neto et al. (2006) nowhere mention the type genus for their new subfamily and include two genera and two species within their grouping. Accordingly, this name fails to meet all of the criteria for availability.

47. †Cratomastotermitidae Engel, Grimaldi, and Krishna, 2009: 9. Type genus: †Cratomastotermes Bechly, 2007. Combining stem: Cratomastotermit–.

48. Archotermopsidae Engel, Grimaldi, and Krishna, 2009: 11. Type genus: Archotermopsis Desneux, 1904d. Combining stem: Archotermops–.

Table 1.

Hierarchical and synonymic outline of termite classification (modified after Engel et al. 2009). Nomina nuda and nomina invalida omitted.

Infraorder Isoptera Brullé, 1832
  Family †Cratomastotermitidae Engel et al., 2009
  Parvorder Euisoptera Engel et al., 2009
  Cretatermitinae Emerson, 1967 [1968]
  Lutetiatermitinae Schlüter, 1989
  Carinatermitinae Krishna & Grimaldi, 2000
  Family Mastotermitidae Desneux, 1904a
  = Pliotermitinae Pongrácz, 1917
  = Miotermitinae Pongrácz, 1926
  Family †Termopsidae Holmgren, 1911
  Family Hodotermitidae Desneux, 1904c
  Family Archotermopsidae Engel et al., 2009
  Family Stolotermitidae Holmgren, 1910a
  Subfamily Porotermitinae Emerson, 1942
  Subfamily Stolotermitinae Holmgren, 1910a
  Family Kalotermitidae Froggatt, 1897
  = Glyptotermitinae Froggatt, 1897
  = Electrotermitinae Emerson, 1942
  Nanorder Neoisoptera Engel et al., 2009
  Family †Archeorhinotermitidae Krishna & Grimaldi, 2003
  Family Stylotermitidae Holmgren & Holmgren, 1917
  Family Rhinotermitidae Froggatt, 1897
  Subfamily Coptotermitinae Holmgren, 1910a
  = Arrhinotermitinae Sjöstedt, 1926
  Subfamily Heterotermitinae Froggatt, 1897
  = Leucotermitinae Holmgren, 1910a
  Subfamily Prorhinotermitinae Quennedey & Deligne, 1975
  Subfamily Psammotermitinae Holmgren, 1911
  Subfamily Termitogetoninae Holmgren, 1910a
  Subfamily Rhinotermitinae Froggatt, 1897
  Family Serritermitidae Holmgren, 1910a
  = Glossotermitinae Cancello and DeSouza, 2005
  Family Termitidae Latreille, 1802
  Subfamily Apicotermitinae Grassé & Noirot, 1954 [1955]
  = Indotermitidae Roonwal & Sen Sarma in Roonwal, 1958
  Subfamily Foraminitermitinae Holmgren, 1912
  = Pseudomicrotermitinae Holmgren, 1912
  Subfamily Sphaerotermitinae Engel & Krishna, 2004a
  Subfamily Macrotermitinae Kemner, 1934, nomen protectum [ICZN 2003]
  = Acanthotermitinae Sjöstedt, 1926, nomen rejiciendum [ICZN 2003]
  = Odontotermitini Weidner, 1956
  Subfamily Syntermitinae Engel & Krishna, 2004a
  = Cornitermitinae Ensaf et al., 2004, nomen nudum
  Subfamily Nasutitermitinae Hare, 1937
  Subfamily Cubitermitinae Weidner, 1956
  Subfamily Termitinae Latreille, 1802
  = Microcerotermitinae Holmgren, 1910b
  = Amitermitinae Kemner, 1934
  = Mirocapritermitinae Kemner, 1934
  = Mirotermitini Weidner, 1956
  = Capritermitini Weidner, 1956
  Incertae Sedis
          Eutermitinae Holmgren, 1910b
Catalog of Names above the Family Group

Here I provide a brief checklist of those names applied to termites above the family-group category. While I have included those supraordinal names which combined termites within an expanded taxon alongside one other group of insects (e.g., Aetioptera Enderlein, 1909), I have not listed those older names which united Isoptera with what are today recognized as numerous other orders (e.g., Platyptera Packard, 1883, for Isoptera, Embiodea, Plecoptera, and Psocoptera). In older literature it is often challenging to determine at what rank a particular name was intended or to what categorical level such a name might be equivalent to today. When it has appeared that a name was intended as a family or category loosely equivalent to today’s family group ranks, I have not included it here. For example, the ‘Termitida’ of Haeckel (1866) was as a family of his order Tocoptera, suborder Pseudoneuroptera, section Corrodentia, despite it having a termination reminiscent of that used in other literature as an ordinal, or other suprafamilial, suffix. Accordingly, I do not consider the Termitida of Haeckel (1866) to be the same as the Termitina or Termitida of Krausse (1906a, 1906b) and Krausse and Wolff (1919), since the former was clearly a family-group name [and thereby a nomen translatum, whether intentional or not, of Latreille’s (1802) Termitina], while the latter two were explicitly employed as ordinal names. Lastly, despite the considerable biological significance of, and increasing reference in the literature to, the combined Isoptera + Cryptocercidae clade, a name has not formally been proposed for this lineage. Herein I offer the name Xylophagodea for this important biological grouping.

1. Isoptères Brullé, 1832: 66 [Latinized by Brauer 1885].

2. Orthoptera socialia Gerstaecker, 1863: 40. Originally deemed a ‘guild’ or ‘fraternity’ (“Zunft”, conceptually equivalent in his system to a suborder); equivalent to Isoptera.

3. Socialia Börner, 1904: 526. Originally deemed a suborder; equivalent to Isoptera of today [Note: Börner’s ‘Isoptera’ included both Embiodea (as suborder Oligoneura Börner, 1904) and Isoptera (as Socialia auctorum)].

4. Termiten Krausse, 1906a: 116. Originally deemed an order; equivalent to Isoptera.

5. Aetioptera Enderlein, 1909: 171. Originally deemed a superorder; equivalent to Isoptera + Embiodea [as Embiidina].

6. Cryptoclidoptera Enderlein, 1909: 171. Originally deemed a suborder; equivalent to all Isoptera excluding Mastotermitidae.

7. Hemiclidoptera Enderlein, 1909: 172. Originally deemed a suborder; equivalent to Mastotermitidae.

8. Termitida Krausse and Wolff, 1919: 159 [vide etiam Rohdendorf 1977]. Equivalent to Isoptera, a form simultaneously used and preferred by the authors.

9. Termitodea Kevan, 1977: 12. Originally deemed a suborder; equivalent to Isoptera + Puknoblattinidea Kevan, 1977 [Note: The latter was simultaneously deemed an infraorder for “Puknoblattinidae Sellards, 1908”, although Sellards (1908) never established a family-group for his genus Puknoblattina. In considering this genus as the sister group to Isoptera, Kevan (1977) was apparently following the notion of Tillyard (1936).].

10. Termitidea Kevan, 1977: 12. Originally deemed an infraorder; equivalent to Isoptera.

11. Isopterodea Boudreaux, 1979: 217. Originally deemed a superorder; equivalent to Isoptera as it was the only included order.

12. Afontanella Myles, 1998: 334. Originally deemed a suborder; equivalent to Mastotermitidae, Termopsidae s.l., Hodotermitidae, and Kalotermitidae (obviously paraphyletic).

13. Fontanella Myles, 1998: 334. Originally deemed a suborder; equivalent to Rhinotermitidae, Serritermitidae, and Termitidae.

14. Octatubula Myles, 1998: 334. Originally deemed an infraorder; equivalent to Rhinotermitidae and Serritermitidae.

15. Quadritubula Myles, 1998: 334. Originally deemed an infraorder; equivalent to Termitidae.

16. Euisoptera Engel, Grimaldi, and Krishna, 2009: 3. Originally rankless; equivalent to all Isoptera exclusive of Cratomastotermitidae and Mastotermitidae.

17. Neoisoptera Engel, Grimaldi, and Krishna, 2009: 9. Originally rankless; equivalent to clade comprising Stylotermitidae, Rhinotermitidae, Serritermitidae, and Termitidae.

18. Xylophagodea, herein. Originally rankless; equivalent to clade comprising Isoptera and Cryptocercidae.

Acknowledgements

I am grateful for numerous years of fruitful and enjoyable collaboration with Prof. Kumar Krishna. Dr. David A. Grimaldi and an anonymous reviewer provided valuable input on the manuscript. This is a contribution of the Division of Entomology, University of Kansas Natural History Museum.

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