Family-Group Names for Termites (Isoptera)

2004·0.15 MB·English

by  Michael S. Engel

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3432, 9 pp., 1 table February 27, 2004
Family-Group Names for Termites (Isoptera)
MICHAEL S. ENGEL1 AND KUMAR KRISHNA2
ABSTRACT
Thirty-nine available family-group names are identified within the insect order Isoptera
(termites).Forallnamesthecorrectauthor,date,typegenus,andcombiningstemareprovided
for the first time. This nomenclatural compilation is done to stabilize the usage of family-
group names in the Isoptera in advance of a world catalog. Several problems of priority are
identified and discussed. The little understood subfamily Foraminitermitinae is diagnosed;
while generally believed by many authors to be a new, unnamed subfamily, it was in fact
established by Holmgren nearly a century ago. The subfamilies Syntermitinae and Sphaer-
otermitinaearenewlyproposedforthemandibulategeneraofnasutetermitesandforSphaer-
otermes, respectively. The classification of Isoptera is briefly outlined.
INTRODUCTION lowed, thereby jeopardizing the stability of
termite classification. In fact, the numerous
It has long been recognized that the cur-
errors that appear in Snyder’s (1949) catalog
rentusageoffamily-groupnamesintheclas-
have been perpetuated by subsequent au-
sification of termites (Isoptera) isnotfullyin
thors, who, rather than checking the original
accord with the rules of zoological nomen-
clature (ICZN, 1999). Not only has the au- literature and applying the rules of nomen-
thorship and date of many names been in- clature, have lifted from Snyder or from oth-
correctly stated (e.g., by Snyder, 1949) and er catalogs which were in turn derived from
the Principle of Coordination (ICZN, 1999: Snyder (e.g., Roonwal and Chhotani, 1989;
Art. 36) ignored, but in many instancesmore Chhotani, 1997).
critical issues of priority have not been fol- In our preparation of a revised catalog of
1DepartmentofEcologyandEvolutionaryBiology,andDivisionofEntomology,NaturalHistoryMuseum,Snow
Hall,1460 JayhawkBoulevard,UniversityofKansas,Lawrence,Kansas66045–7523.e-mail:[email protected]
2DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory;CityCollegeandtheGraduateSchool,
City UniversityofNewYork.e-mail:[email protected]
Copyright(cid:113)AmericanMuseumofNaturalHistory2004 ISSN0003-0082
2 AMERICAN MUSEUM NOVITATES NO. 3432
the world termite fauna (Krishna and Engel, stem: Termit–. Note: Latreille (1805, 1810)
in prep.), we have gone to great lengths to subsequently changed the name of his ‘‘fam-
attend to these nomenclatural difficulties ille’’ to Termitinae but the name was made
(e.g., Engel and Krishna, 2001a, 2001b, in available in 1802 (ICZN, 1999: Art. 11.7).
press a & b; Engel et al., 2003). In order to 2. Calotermitinae Froggatt, 1897: 516.
bring the usage of family-group names into Type genus: Kalotermes Hagen, 1853 [Cal-
accord with ICZN (1999) rules, we provide otermes Hagen, 1858 is an unjustified emen-
herein a catalog of all such names that have dation: see EngelandKrishna,2001a;ICZN,
been applied in the Isoptera. This has been 2002]. Name emended to Kalotermitinae
done to correct the variouscirculatingerrors, Froggatt, 1897, in accordance with ICZN
to highlight the most egregious problems, (1999: Art. 29.1). Combining stem: Kaloter-
and to lay a standardized foundation for the mit–.
forthcoming catalog. In addition, we have 3. Glyptotermitinae Froggatt, 1897: 518.
provided systematic descriptions for three Type genus: Glyptotermes Froggatt, 1897.
subfamilies (two new), which are otherwise Combining stem: Glyptotermit–.
not employed by isopterists today but must
4. Rhinotermitinae Froggatt, 1897: 518.
be validated prior to the publication of the
Type genus: Rhinotermes Hagen, 1858.
catalog (Krishna and Engel, in prep.). We
Combining stem: Rhinotermit–.
have also included those published names
5. Heterotermitinae Froggatt, 1897: 550.
that are otherwise unavailable at the end of
Type genus: Heterotermes Froggatt, 1897.
thelistingofavailablenames.Itmustbenot-
Combining stem: Heterotermit–.
ed that additional unavailable names have
6. Mastotermitinae Desneux, 1904a: 284.
been used on internet sites and employed by
Type genus: Mastotermes Froggatt, 1897.
GenBank, but none of these have met the
Combining stem: Mastotermit–.
criteria of valid publication. While being un-
7. Hodotermitini Desneux, 1904a: 284.
available, like the three Holmgren names
Typegenus:HodotermesHagen,1853.Com-
consideredbelow,thesenameshavenotbeen
bining stem: Hodotermit–.
published in the sense defined by the ICZN
8. Stolotermitinae Holmgren, 1910a: 285.
(1999) (unlike Holmgren’s names which
Type genus: Stolotermes Hagen, 1858. Com-
were published in accordance with the
bining stem: Stolotermit–. Note: This name
ICZN) and are therefore excluded from this
unfortunately has priority over Termopsidae
study.
Holmgren, 1911, a fact discovered too late
TAXONOMIC CATALOG to be included in the petition to conserve
Termopsis and Termopsidae relative to Mas-
Below we list in order of priority all fam-
totermitidae (Engel et al., 2003). Therefore,
ily-group names proposed for termites. The
a second petition was prepared to the ICZN
names are presented in their original forms,
to conserve Termopsidae relativeto Stoloter-
regardless of present day rank or suffix. The
mitidae(EngelandKrishna,inpressa),since
typegenus isprovidedalongwiththecorrect
it would be destabilizing fornomenclatureto
combining stem for composing a family-
use Stolotermitidae in place of what is today
groupname.Daggers(†)indicatenamespro-
referred to as Termopsidae.
posed for fossil taxa. Some names were be-
lieved to have been nomina nuda by Snyder 9. LeucotermitinaeHolmgren,1910a:285.
(1949) when they were first proposed; how- Type genus: Leucotermes Silvestri, 1901.
ever,priorto1931family-groupnamescould Combining stem: Leucotermit–.
be made available by simple formation from 10. Coptotermitinae Holmgren, 1910a:
anavailablegenus-groupname(ICZN,1999: 285. Type genus: Coptotermes Wasmann,
Art. 12.2.4) and need not have had a formal 1896. Combining stem: Coptotermit–. Note:
description. ProposedagainasnewinHolmgren(1910b).
11. Serritermitinae Holmgren, 1910a:285.
AVAILABLE NAMES Type genus: Serritermes Wasmann, 1897.
1. TermitinaLatreille,1802:293.Typege- Combining stem: Serritermit–.
nus: Termes Linnaeus, 1758. Combining 12. Termitogetoninae Holmgren, 1910a:
2004 ENGEL AND KRISHNA: FAMILIES OF TERMITES 3
286. Type genus: Termitogeton Desneux, 22. Macrotermitinae Kemner, 1934: 69.
1904b. Combining stem: Termitogeton–. Type genus: Macrotermes Holmgren, 1909.
13. Microcerotermitinae Holmgren, Combining stem: Macrotermit–. Note: See
1910b: 145. Type genus: Microcerotermes comments for Acanthotermitinae (above).
Silvestri, 1901. Combining stem: Microcer- 23. Amitermitinae Kemner, 1934: 110.
otermit–. Type genus: Amitermes Silvestri, 1901.
14. Eutermitinae Holmgren, 1910b: 146. Combining stem: Amitermit–.
Type genus: Eutermes Heer, 1849. Combin- 24. Miro-capritermitinae Kemner, 1934:
ing stem: Eutermit–. Note: This name has 166. Type genus: Mirocapritermes Holm-
priority over Nasutitermitinae, despite the gren, 1914. Combining stem: Mirocapriter-
synonymy of its type genus with Nasutiter- mit–. Note: Although Kemner (1934) hy-
mes (ICZN, 1999: Art. 40.1). Eutermes phenatedthenameinitsoriginalspelling,the
should also take precedence over Nasutiter- ICZN(1999)doesnotallowhyphenationand
mes,compoundingtheconfusion.UseofEu- the family-group name must be considered a
termitinae in place of Nasutitermitinae,how- single word.
ever, would be destabilizing for termite no- 25. Nasutitermitinae Hare, 1937. Type ge-
menclature. A petition to the ICZN has been nus: Nasutitermes Dudley, 1890. Combining
prepared to conserve Nasutitermes and Na- stem: Nasutitermit–. Note: See comments
sutitermitinae (Engel and Krishna, in press above on Eutermitinae.
b). Current usage should be maintained until 26. †Electrotermitinae Emerson, 1942:10.
an authoritative decision is rendered. Type genus: †Electrotermes Rosen, 1913.
15. Termopsinae Holmgren, 1911: 35. Combining stem: Electrotermit–.
Type genus: Termopsis Heer, 1849. Combin- 27. Porotermitinae Emerson, 1942: 10.
ing stem: Termops–. Note: See commentson Type genus: Porotermes Hagen, 1858. Com-
Stolotermitinae (above). bining stem: Porotermit–.
16. Psammotermitinae Holmgren, 1911: 28. Apicotermitinae Grasse´ and Noirot,
64. Type genus: Psammotermes Desneux, 1954 [1955]: 360. Type genus: Apicotermes
1902. Combining stem: Psammotermit–. Holmgren, 1912. Combining stem: Apicoter-
17. Pseudomicrotermitinae Holmgren, mit–.
1912: 5. Type genus: Pseudomicrotermes 29. Cubitermitini Weidner, 1956: 99. Type
Holmgren,1912.Combiningstem:Pseudom- genus:CubitermesWasmann,1906.Combin-
icrotermit–. ing stem: Cubitermit–.
18. Foraminitermitinae Holmgren, 1912: 30. Mirotermitini Weidner, 1956:99.Type
5. Type genus: Foraminitermes Holmgren, genus: Mirotermes Wasmann, 1897. Com-
1912. Combining stem: Foraminitermit–. bining stem: Mirotermit–.
19. Stylotermitinae Holmgren and Holm- 31. Capritermitini Weidner, 1956: 100.
gren, 1917: 141. Type genus: Stylotermes Type genus: Capritermes Wasmann, 1897.
Holmgren and Holmgren, 1917. Combining Combining stem: Capritermit–.
stem: Stylotermit–. 32. Indotermitidae Roonwal and Sen-Sar-
20. Arrhinotermitinae Sjo¨stedt, 1926: 8. ma In Roonwal, 1958: 81. Type genus: In-
Type genus: Arrhinotermes Wasmann, 1902. dotermes Roonwal and Sen-Sarma In Roon-
Combining stem: Arrhinotermit–. wal, 1958. Stem: Indotermit–.
21. Acanthotermitinae Sjo¨stedt, 1926: 8. 33. †Cretatermitinae Emerson, 1968: 278.
Type genus: Acanthotermes Sjo¨stedt, 1900. Type genus: †Cretatermes Emerson, 1968.
Combiningstem:Acanthotermit–.Note:This Combining stem: Cretatermit–.
name has priority over Macrotermitinae; 34. Prorhinotermitinae Quennedey and
however, a petition has been submitted to Deligne, 1975: 265. Type genus: Prorhino-
conserve the usage of Macrotermitinae (En- termes Silvestri, 1909. Combining stem:
gel and Krishna, 2001b), which has been ap- Prorhinotermit–.
provedbytheICZN(2003).Macrotermitinae 35. †Lutetiatermitinae Schlu¨ter, 1989: 61.
is to be used whenever Macrotermes and Type genus: †Lutetiatermes Schlu¨ter, 1989.
Acanthotermes are placedintothesamefam- Combining stem: Lutetiatermit–.
ily-group taxon. 36. †Carinatermitinae Krishna and Gri-
4 AMERICAN MUSEUM NOVITATES NO. 3432
maldi, 2000: 134. Type genus: †Carinater- 11.7.1.1). Note: This name was proposed as
mes Krishna and Grimaldi,2000.Combining new again in Holmgren (1910a) and Holm-
stem: Carinatermit–. gren (1910b). [A genus-group name, Proter-
37. †Archeorhinotermitinae Krishna and mes, was not proposed until 1910, at which
Grimaldi,2003:2.Typegenus:†Archeorhin- time Holmgren (1910a) did not include Pro-
otermes Krishna and Grimaldi, 2003. Com- termesinhisProtermitidae,placingitinstead
bining stem: Archeorhinotermit–. in Metatermitidae (in fact, Protermes was
38. Syntermitinae Engel and Krishna, never included in Protermitidae). Protermi-
herein (see below). Type genus: Syntermes tidae was used exclusively for what Holm-
Holmgren, 1909. Combining stem: Synter- gren considered to be the basal termites.
mit–. Thus, Protermitidae was not subsequently
39. Sphaerotermitinae Engel and Krishna, validated in the later publications of Holm-
herein (see below). Type genus: Sphaeroter- gren.]
mes Holmgren, 1912. Combining stem:
Sphaerotermit–. SYSTEMATIC DESCRIPTIONS
Below we provide descriptions for three
UNAVAILABLE NAMES little understood subfamilies of Termitidae,
one hitherto little understood and two new.
Someauthors(e.g.,Banks,1920;Roonwal
The Foraminitermitinae was originally pro-
and Chhotani, 1989)have erroneouslyequat-
posed by Holmgren (1912) but has not been
ed the following three names with some of
generally recognized. The genera Foramini-
the present families or subfamilies. All three
termes and Labritermes, however, have in-
are unavailable and therefore do not enter
creasingly been recognizedasdistinctiveand
into zoological nomenclature and should not
worthy of exclusion from Termitinae (e.g.,
be used.
Krishna, 1963; Noirot, 2001; Bitsch and
1. Metatermitidae Holmgren, 1909: 99.
Noirot, 2002), where they had been placed.
Unavailable, as it was not based on an avail-
The subfamily is characterized here to make
able genus-group name (ICZN, 1999: Art.
its diagnostic characters more widely under-
11.7.1.1).
stood. Similarly, the mandibulate genera of
2. Mesotermitidae Holmgren, 1909: 100.
the nasute termites are excluded from Na-
Unavailable, as it did not include Mesoter-
sutitermitinae and a new subfamilial name is
mesHaase,1890(ICZN,1999:Art.11.7.1.1)
required to accommodate this group (pro-
and was therefore not based on a genus-
posed as Syntermitinae, below). Lastly,
group name. Note: This name was proposed
Sphaerotermes is excluded from the Macro-
as new again in Holmgren (1910a) and
termitinae owing to a unique combination of
Holmgren (1910b). [Mesotermes was an
plesiomorphicand apomorphictraits(seebe-
available genus-group name at that time, but
low).
recognized by Handlirsch (1906) as a neu-
ropteran and not a termite. Holmgren (1909,
Subfamily FORAMINITERMITINAE Holmgren
1910a) clearly did not intend this name tobe
based on the fossil neuropteran Mesotermes Foraminitermitinae Holmgren, 1912: 5. Type ge-
but established it instead for a series of ex- nus: Foraminitermes Holmgren, 1912.
plicitly included, living termite genera (see Combining stem: Foraminitermit–.
Holmgren,1910a,1910b),whichhebelieved DIAGNOSIS [Derived fromKrishnaandAd-
to occupy an intermediate phylogenetic po- ams (1982)]: Imago. Head densely pilose;
sition in his scheme of termite relationships; fontanelle punctiform, situated at tip of a
thus, Mesotermes was not included by infer- cornical projection. Forecoxa with longitu-
ence of the generic stem of Mesotermitidae dinal ridge, without protuberance; tibial
(ICZN, 1999: Art. 11.7.1.1) and the name spurs3–2–2.Soldier.Leftmandiblewithup-
was never made available.] per, inner cutting edge finely serrated, and
3. Protermitidae Holmgren, 1909: 100. withbluntteethappearingascrenulationsbe-
Unavailable, as it was not based on an avail- low serrations; right mandible smooth, with-
able genus-group name (ICZN, 1999: Art. out teeth, base of molar region with one or
2004 ENGEL AND KRISHNA: FAMILIES OF TERMITES 5
TABLE1
Hierarchical Outline of Termite Classification
All family-group names indicated; synonymic names italicized.
6 AMERICAN MUSEUM NOVITATES NO. 3432
two very tiny, pointed, thornlike spines or the inner scaly membrane (except reducedin
projections. Labrum with hyaline tip. Work- Syntermes).
er. Gut with mixed segment absent; four INCLUDED GENERA: Syntermes Holmgren,
malpighian tubules separately attached at 1909; Cornitermes Wasmann, 1897; Procor-
transverse mesenteron-proctodeum junction, nitermes Emerson In Snyder, 1949; and La-
extending forward for short distance and biotermes Holmgren, 1912. Other mandibu-
then looping around toward hindgut; mal- late genera of Nasutitermitinae may eventu-
pighian tubules swollen basally; first proc- ally be included here as well.
todealsegment(P )tubularandnarrowprox-
1
imally, almost same diameter as midgut, and
SPHAEROTERMITINAE, new subfamily
dilated and saclike distally, with malpighian
tubules forming jumbled mass ((cid:53) pseudom- TYPE GENUS: Sphaerotermes Holmgren,
alpighian knot of Noirot, 2001; Bitsch and 1912.
Noirot, 2002). DIAGNOSIS: Imago. Head with light,small,
INCLUDED GENERA: ForaminitermesHolm- punctiform fontanelle. Labrum uniformly
gren,1912andLabritermesHolmgren,1914. sclerotized, transparent; postclypeus with
COMMENTS: This group was first recog- median line.Soldier.Headwithsidesround-
nized by Holmgren (1912), who had diffi- ed; inconspicuous fontanelle present in mid-
culty placing it with any other group of Ter- dleofhead.Labrumwithsidesrounded,with
mitinae (referring to it alsoasthe‘‘Foramin- hyaline point. Left mandible with a few ser-
itermes-reihe’’) (also recognized as distinc- rations and blunt tooth near base; right man-
tive by other authors; e.g., Krishna, 1963; dible with inner cutting-edge smooth. Pron-
Krishna and Adams, 1982; Noirot, 2001). otum much narrower than head; anterior and
Noirot(2001)suggestedthatForaminitermes lateral margins even, without projecting
belongedtoanewsubfamily,whileafamily- spines. Worker. Right mandible with 6–7
group name had already been proposed forit ridges on molar plate. Labrum uniformly
nearly a century ago. sclerotized (Sands, 1998: lacking the trans-
verse apical band of sclerotization seen in
SYNTERMITINAE, new subfamily Macrotermitinae,e.g.,Donovanetal.,2000);
postclypeus strongly convex, with distinct
TYPE GENUS: Syntermes Holmgren, 1909 median line. Gizzard with small pulvilli at-
(not 1910a as cited by Snyder, 1949; Con- tached posteriorly on columns, crenulated
stantino, 1995, 1998; and other authors). crests of columns I and II poorly developed;
DIAGNOSIS: Imago. Fontanelle generally setae of pulvilli poorly developed (Noirot,
large, round; slightly depressed or forming a 2001). Conspicuous, backward-orientedbris-
convex plate. Postclypeus moderately or tles at the P -P (paunch-colon) junction
3 4
strongly convex, with a median line. Fore- (Noirot, 2001) (unique among the Isoptera).
and hindwing with short R , sometimes INCLUDED GENERA: Includes only Sphaer-
1
R (cid:49)R joining costal margin very near the otermes Holmgren, 1912.
2 3
suture. Pronotum with anterior margins pro- COMMENTS:Thisisaremarkablesubfamily
jecting laterally into points in Syntermes; in sister to the Macrotermitinae (e.g., three
Procornitermes, Cornitermes and Labioter- nymphal instars, unique worker instar, male
mes anterolateral corners without points, workers larger than females) but unlike the
margins even. Arolium absent. Soldier. Na- macrotermitines, Sphaerotermes is not asso-
sus (frontal tube) well developed, tip flat; ciatedwithTermitomycesfungiandlacksthe
frontal pore wide and conspicuous; well-de- transverse band of sclerotization across the
veloped, functioning biting-mandibles; la- labrum of the worker and the imago (Sands,
brum simple, narrow, with hyaline tip. 1998). While these are indeed plesiomor-
Worker. Gut with mixed segment doubled, phies relative to Macrotermitinae, Sphaero-
dual lobes overlapping the first proctodeal termitinae has some remarkable apomorphic
segment (P ) (except reduction to single in traitsthatsupportitsrecognition[e.g.,unique
1
Procornitermes);entericvalvewithspinesin possession of conspicuous, backward-orient-
2004 ENGEL AND KRISHNA: FAMILIES OF TERMITES 7
ed bristles at the P -P (paunch-colon) junc- 25]: 1–52. Bruxelles [Brussels]: P. Wytsman.
3 4
tion: Noirot, 2001]. [publ. date: 15 November 1904]
Donovan, S.E., D.T. Jones, W.A. Sands, and P.
Eggleton. 2000. Morphological phylogenetics
ACKNOWLEDGMENTS
of termites (Isoptera). BiologicalJournalofthe
We are grateful to Valerie Krishna, David Linnean Society 70: 467–513.
Dudley,P.H.1890.ThetermitesoftheIsthmusof
Nickle, and Charles D. Michenerforreviews
Panama—PartII.TransactionsoftheNewYork
of the manuscript. Support was provided by
Academy of Sciences 9: 157–180. [publication
a National Science Foundation grant to cat-
date June 1890, not 28 April 1890 as cited by
alog the termites of the world (NSF DEB- Constantino (2002): results of 5 May 1890
9870097 to K. Krishna and D. Grimaldi). meeting are published on same page as Dud-
This is contribution number 3344 of the Di- ley’s article and date on top of facing page is
vision of Entomology, Natural History Mu- dateofpreviousAcademymeeting,notofpub-
seum and BiodiversityResearchCenter,Uni- lication;journalcoverindicatesanearliestpos-
versity of Kansas. sible publication date of June 1890]
Emerson, A.E. 1942. The relations of a relict
South African termite (Isoptera: Hodotermiti-
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