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September 1991

Notes On Larvae 1989-1991

George C. Wheeler Florida State Collection of , Florida Department of Agriculture and Consumer Services, Gainesville, FL

Jeanette Wheeler Florida State Collection of Arthropods, Florida Department of Agriculture and Consumer Services, Gainesville, FL

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Wheeler, George C. and Wheeler, Jeanette, "Notes On Ant Larvae 1989-1991" (1991). Insecta Mundi. 427. https://digitalcommons.unl.edu/insectamundi/427

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Vol. 5, No. 3-4, September-December 1991 167

Notes On Ant Larvae 1989-1991

George C. Wheeler' and Jeanette Wheeler Research Associates, Florida Collection of Arthropods Gainesville, Florida 3358 NE 58th Avenue Silver Springs, Florida 32688

Introduction studied based solely upon the head. The 44 In 1976 we published "Ant Larvae: Review and drawings of the whole head are in anterior (=full- Synthesis," which we regarded as a summary of face) view and of the mouth parts enlarged. The our life-time of research. During the following drawings are quite good except that the diameters years so many new larvae were added to our of the hairs are greatly exaggerated. collection and so much was published about ant Pierce 1987. This author has written a thor- larvae that we decided another general treatment ough review of the relations between and was desirable; so in 1986 we published a "Ten-Year lycaenid larvae, but unfortunately has not named Supplement" to include additions and revisions to many of the ant species involved. The larvae of our Memoir. In 1987 and 1988 additional material several lycaenid species feed on ant brood. warranted a supplement to the "lo-year Supple- Wheeler and Wheeler 1989a. The printer failed ment" in 1989a. This article is the third supple- to heed many of the corrections we made on the ment to the Memoir. galley proof although our manuscript had been In the supplement, two genera are charac- corrected. This was especially true of Literature terized, one generic characterization is revised, and Cited. Therefore, we have prepared corrected copies references in the literature are increased by 43. of this part, which we will be glad to mail upon Changes in our Memoir (1976) and its supplements request. In addition the references on page 458 to (1986, 1989a) are noted. and Mystrium should have been deleted. Family Formicidae Hijlldobler and Wilson 1990. These authors Subfamily have cited many references to our publications on Body profile myrmecioid. Body hairs short.and ant larvae. We cite some of these in appropriate moderately abundant. Head hairs smooth and places here And hereafter. unbranched. Mandible large, pogonomyrmecoid. On page 22 is our figure showing our classifica- Labium with dorsal transverse welt, which is tion of the body profiles of ant larvae (1976: Fig. 3). densely beset with coarse spinules; palp a slight On page 21 they state: "Ant larvae have been elevation bearing 5-7 sensilla; opening of sericteries systematically described by Wheeler and Wheeler wide and salient. (1951-1986, syntheses 1976 and 1979), with a supplementary analysis supplied by Picquet Fabricius (1958)." This implies that Picquet's article sup- Body profile myrmecioid. Body hairs short; plements our publications, but a casual glance moderately numerous, uniformly distributed, shows that this is incorrect as to its contents, while unbranched, smooth or more or less denticulate, Picquet's article was published in 1958; and a sup- rarely uncinate. Head subpyriform in anterior plement is something added later. view. Head hairs few, unbranched, smooth or Picquet 1958. The larvae of 18 species are denticulate. Labrum small, short and bilobed, described and figured. The author's conclusions posterior surface with large isolated spinules and from this study were expressed by a key to the sensilla of various sizes. Mandible large, pogono-

George C. Wheeler died on February 19, 1991. Insecta Mundi myrmecoid and heavily sclerotized; basal half Platythyrea schultzi Fore1 usually bearing isolated spinules. Maxilla and Wheeler and Wheeler 1989b:51. Description. labium spinulose with many large isolated spin- ules. Genus PROBOLOMYRMEX Mayr

Myrmecia sp. Probolomyrmex angusticeps M. R. Smith Hijlldobler and Wilson 1990:347, Fig. 8-48. Taylor 1965:348. Description of mature larvae Photograph of larvae. based on pharate with figures on p. 348. Taylor said: "The Probolomyrmex is distin- Genus Clark guished from those of all other known ponerine Body profile myrmecioid. Body hairs short, ants by the shape of the body and the unique moderately abundant and unbranched; of 2- types: posterodorsal suspensory organ, which is analagous (1) rather stout, slightly curved and with minute (but clearly not homologus) with the dorsal 'door- denticles near the apex, on all somites; (2) shorter, knob' tubercles found in some genera of the tribe sharply bent and denticulate, on venter of anterior (see G. C. and J. Wheeler, 1952, 1964)." somites. Head subcircular in anterior view. Head But we have to challange the above: body shape is hairs moderately numerous and moderately long, the one character that cannot be determined from smooth and unbranched. Labrum short, moderate- the prepupa. sized. Mandible large, pogonomyrmecoid. Maxilla Holldobler and Wilson 1990:348. "Workers... coarsely spinulose. Labium with a dorsal trans- carry the larvae by the [posterodorsall tubercles. verse welt, which is densely beset with coarse They also attach the larvae to the roof of the nest spinules. Hyopharynx with a few minute spinules. with the apparently adhesive outer surface of the knob-like endings." Nothomyrmecia macrops Clark Holldobler and Wilson (1990:165, Fig.3-19). A Tribe Proceratiini photograph of living larvae. Genus Roger

Subfamily Proceratium watasei, P. itoi, P. japonicum Tribe Amblyoponini Masuko 1986. Queens cut holes in the integu- Genus AMBLYOPONE Dalla Torre ment of worker larvae and feed on the exuding . Amblyopone silvestri Wheeler Masuko 1986. Queens cut holes in the integu- Tribe Ponerini ment of mature or nearly mature larvae and feed Genus HYPOPONERA Santschi upon the exuding larval hemolymph. Wheeler and Wheeler 1989b:52. Correction in Masuko 1990. The external anatomy of five characterization. instars is described and illustrated by SEM. Wheeler and Wheeler 1989a:458. Headings and Genus PLECTROCTENA F. Smith references should have been deleted by printer. Genus MYSTRIUM Roger Plectroctena conjugata Santschi Wheeler and Wheeler 1989b:52. Description. Wheeler and Wheeler 1989a:458. Headings and references should have been deleted by the printer. Genus Latreille Tribe Platythyreini Genus PLA NTHYREA Roger Ponera coarctata Latreille Picquet 1958:22, Figs. 24-27. Platythyrea lamellosa Roger Genus SIMOPELTA Mayr Wheeler and Wheeler 1989b:51. Description. Wheeler and Wheeler 1989b:52. Characteriza- tion. Vol. 5, No. 3-4, September-December 1991

Genus STREBLOGNATHUS Mayr numerous, short, unbranched, smooth. Mouth parts Wheeler and Wheeler 1989b:53. Characteriza- small. Mandible tetraponeroid. tion. Tetraponera (=Pachysima) latifrons Emery Streblognathus aethiopicus (F. Smith) Holldobler and Wilson 1990:166, Figs. 3-20 Wheeler and Wheeler 1989b:53. Characteriza- repeat Wheeler 1918, Fig. 9 and our 1956, Text tion. Fig. 5. Subfamily Cerapachyinae Subfamily Genus CERAPACHYS F. Smith Tribe Myrmicini Genus Latreille Cerapachys turneri Fore1 Holldobler and Wilson 1990:569. Workers of Myrmica laevinodis Nylander species in this group store, after stinging, the Picquet 1958:30, Figs. 38-42. living larvae of . Myrmica ruginodis Nylander Subfamily Picquet 1958:31, Fig. 43. Genus NEIVAMYRMEX Borgmeier Tribe Pheidolini Neivamyrmex pilosus (F. Smith) Genus APHAENOGASTER Mayr Akre and Torgerson 1968. The staphylinid beetle Diploeciton nevermanni Reichensperger is Aphaenogaster cockerelli (E. Andr6) predaceous on the ant brood. Holldobler and Wilson 1990:239, Fig 7-21. Photograph of larvae. Subfamily Genus Emery Aphaenogaster gib bosa Latreille Picquet 1958:25, Figs. 32-35. Baroni-Urbani Aphaenogaster subterranea Latreille Masuko 1987. Unique among ant larvae is a Picquet 1958:24, Figs. 28-31. pair of hemolymph feeding pores. Through these Holldobler and Wilson 1990:68, Fig 3-2. A good the queen feeds on the hemolymph of the larvae. photograph of larvae placed on food by workers. Masuko also found that when a worker moves a larva it grasps with its posterior mouth parts the peculiar anteroventral projection from the pro- Genus PHEIDOLE Westwood thorax of the larva. Wheeler and Wheeler 1988:185-189. Descrip- Pheidole bicornis Fore1 tion of young larvae. Letourneau 1990. Larvae of a clerid beetle Phyllobaenus sp. feed upon the Pheidole brood. Subfamily Genus TETRAPONERA F. Smith Tribe REVISED CHARACTERIZATION. Body profile Genus SOLENOPSIS Westwood crematogastroid. Trophothylax well developed. Body hairs numerous. Of 3 types: (1) minute to Solenopsis fugax Latreille short, unbranched, smooth, on all somites; (2) Picquet 1958:26, Fig 36. moderately long, 2-10 on dorsal and lateral surfac- es of most somites; (3) long, with sinuous shaft and Tribe Tetramoriini uncinate tip, several in a row across dorsum of Genus TETRAMORIUM Mayr each thoracic and several abdominal somites. Head subquadrangular. Antennae minute. Head hairs Insecta Mundi

Tetramorium caespitum Linnaeus Lasius niger Linnaeus Picquet 1958:28, Fig. 37. Picquet 1958:16, Fig. 17. Tribe Blepharidattini Tribe Camponotini Genus BLEPHARIDATTA Wheeler Genus CAMPONOTUS Mayr

Blepharidatta brasiliensis Wheeler Camponotus aethiops Latreille Wheeler and Wheeler 1991: 134-135. Descrip- Picquet 1958:10, Figs. 7-8. tion. Camponotus floridanus (Buckley) Subfamily Davis and Jouvenaz 1990. Another eucharitid, Genus Mayr Obeza floridana (Ashmead), is parasitoid on this species. Iridomyrmex humilis Mayr Camponotus lateralis Olivier Wheeler and Wheeler 1989a:465. Description. Picquet 1958:15, Fig. 16.

Genus TAPINOMA Foerster Camponotus ligniperdus Larteille Picquet 1958:12, Figs. 9-12. Tapinoma erraticurn Latreille Picquet 1958:32, Fig. 44. Camponotus modoc Wheeler Hansen and Akre 1985: Figs. 19 and 35. Photo- Genus TECHNOMYRMEX Mayr graphs of larvae.

Technomyrmex sp. Camponotus vagans Scopoli Wheeler and Wheeler 1989a:467. Description of Picquet 1958:13, Figs. 13-15. young larva. Genus F. Smith Subfamily Holldobler and Wilson 1990:485. A syrphid fly Tribe Formcini Trichopsomyia is a parasitoid on the brood. Genus FORMICA Linnaeus Wheeler and Wheeler 1990:753-767.The larvae of 11 species are described. Holldobler and Wilson 1990:484. The apterous females of the phorid flies Aenigmatias lay on Polyrhachis arachne Emery and P. hodgsoni Fore1 host larvae (Formica). Dorow and Maschwitz (1990: 77-78). The colony is housed in a "pavilion" contructed of silk secreted Formica fusca Linnaeus by the larvae, which are manipulated by the Picquet 1958:9, Fig. 6. workers. The underside of a bamboo leaf contrib- utes the ceiling of the pavilion, which houses the Formica rufa Linnaeus nest. This ceiling is covered with a thin layer of Picquet 1958:4, Figs. 1-5. silk. "The brood was fixed longitudinally to this layer with a few threads of silk." Fig. 2 shows a Genus LASIUS (Fabricius) weaving larva of P. hodgsoni.

Lasius brunneus Latreille CHANGES IN OUR 1976 MEMOIR (and in our 10- Picquet 1958:18, Figs. 18-19. year supplement [I9861 and in our 1989a:457-473)

Lasius emarginatus Olivier GEOGRAPICAL DISTRIBUTION (1976:2) Picquet 1958:19, Figs. 20-22. ADD AFRICA--Natal, Zimbabwe; SOUTH AMERI- CA-- Lasius flavus Fabricius Picquet 1958:20, Fig. 23. Vol. 5, No. 3-4, September-December 1991

BODY SHAPES (1976:B) which will have all the characters of the last instar DELETE except body shape. But such favorable specimens From pogonomyrmecoid--Ponerinae: Cryptopone, are rarely found; hence we like to get a large Ponera. Myrmicinae: Leptothorax (Mychothorax and number of larvae from the same nest. Nesomyrmex), Formicinae: Gesomyrmex, Plagio- In our Memoir (1976:81) we mentioned the lepis. number of instars reported in the literature. In our From pheidoloid--Myrmicinae: Megalomyrmex, 10-year Supplement (1986:695) we brought it up to Trigonogaster. date. Holldobler and Wilson (1990:170) have updated it again citing authorities. Masuko (1990) ADD gives in a table the number of instars in 25 species pogonomymecoid--Formicinae:Anoplolepis, Dendro- and a 26th is cited in a postscript. After adding our myrmex. recent contributions (1987) for Veromessor pergan- pheidoloid--Myrmicinae:Leptothorax (Mychothorax dei, (198913) for Neivamyrmex opacithorax and and Nesomyrmex). Formicinae: Gesomyrmex. (1991) for Blepharidatta the whole list of reported attoid--Myrmicinae: Blepharidatta, Myrmicaria. instars looks like this: myrmecioid--Ponerinae: Ponera. 3 instars: Cataglyphis cursor, aphaenogastroid--Myrmicinae: Adelomyrmex, scutellaris, C. stadelmanni, C. striatula, For- Megalomyrmex. mica japonica, Messor rubra, leptanilloid--Ponerinae: Cryptopone, Myopopone. minimum, M. pharaonis (workers and queen), Myrmicinae: Trigonogaster. Myrmica ruginodis, Tetramorium caespitum. oecophylloid--Cerapachyinae: Simopone. Formic- 3 or 4 instars: Oecophylla longinoda, Pheidole inae : Acropyga. pallidula. 4 instars: Acromyrmex octospinosa, Brachyponera HAIRS (1976:34) chinensis, Formica polyctena, Paratrechina ADD flavipes, Pheidole bicarinata, Polyrhachis G. Short, stout, with narrow base and wider top, lamellidens, Solenopsis invicta, Zacryptocerus which has a flat pebbly surface. Known only in minutus. Platythyrea lamellosa; see our 198913: Fig. 1. 5 instars: Acantholepis frauenfeldi, , Camponotus aethwps,Eciton burchel- MANDIBLE SHAPES (1976:39) li, E. hamatum, Neivamyrmex opacithorax, DELETE Polyrhachis armata, P. bihamata, P. mulleri, P. 5. amblyoponoid--Cerapachyinae:Lioponera, Phyra- scissa, P. ypsilon, Veromessor pergandei. caces. 6 instars: Camponotus aethiops queen. 6. pristomyrmecoid--Myrmicinae: Mascramisch- oides. CARE (1976:82) 10. attoid--Myrmicinae: . Holldobler and Wilson 1990:168. A good para- graph on feeding. ADD 1. ectatommoid--Myrmicinae: Acanthomyrmex, TAXONOMIC BIBLIOGRAPHY OF OUR PUBLI- Macromischoides. Formicinae: Proformica. CATIONS ON ANT LARVAE (1976:93, 1986:697 3. dolichoderoid-Xerapachyinae: Lioponera. and 1989a:672) 5. amblyoponoid--Myrmicinae: Blepharidatta, Eight subfamilies 1989. Notes on ant larvae. Sericomyrmex, Trachymyrmex. Trans. Amer. Ent. Soc. 115:457-473. 7. pheidoloid--Myrmicinae: Macromischoides. Ponerinae. 1989. Notes on ant larvae: Ponerinae. J. New York Ent. Soc. 97:50-55. LIFE CYCLE (1976:80; 1986:694) Leptanillinae. 1988. The larva of Leptanilla japoni- When we describe immature larvae we do not ca, with notes on the genus. Psyche 93:185-189. like to call them instars unless certain conditions Formicinae. 1990. Larvae of the formicine ant are fblfilled; an ready to hatch will reveal the genus Polyrhachis. Trans. Amer. Ent. Soc. characters of the first instar; a first instar ready to 116:753-767. moult will contain a fully formed second instar; Myrmicinae. 1991. The larva of Blepharidatta. J. etc.; etc.; until we find a semipupa (=prepupa) New York Ent. Soc. 99:132-137. Insecta Mundi

MATERIAL STUDIED (1976:96 and 1986:698) IMPORTANCE OF LARVAE (1986:696) Holldobler and Wilson 1990:392. "According to ADD Davison (19821, colonies of Monomorium (=Chelan- Ponerinae er) in arid New South Wales are sensitive to 2. Plaththyreini. Platythyrea: lamellosa Roger, fluctuations in the supply of seeds on which they schultzi Forel. depend. In times of scarcity the number of larvae 7. Ponerini. Plectroctena conjugata Santschi. declines. Because the adult workers depend on Streblognathus aethiopicus (F. Smith). larval secretions for food (the larvae consume and metabolize the raw seeds for the colony), the Pseudomyrmecinae worker population also declines." malignus Wheeler. Holldobler and Wilson 1990:164-168. Brood care and larval reciprocation. Myrmecinae 11. Myrrnecinini. Acanthomyrmex: ferox Emery, Literature Cited notabilis F. Smith. Akre, R. D., and R. L. Torgerson. 1968. The 13. Leptothoracini. Lachnomyrmex scrobicula- behavior of Diploeciton nevermanni, a staphy- tus Wheeler. Leptothorax: diuersipilosus Smith, linid beetle associated with army ants. Psyche longispinosus Roger, quebecensis (Francoeur), 75:211-215. whe.eteri Smith. Rogeria: belti Mann, blanda F. Smith. Davis, L. R., and D. P. Jouvenaz. 1990. Obeza 15. Tetramoriini. Tetramorium: blochmani floridana, a parasitoid of Camponotus abdomi- Forel, lobulicornis Forel. nalis floridanus from Florida. Florida Ent. 21. Blepharidattini. Blepharidatta brasiliensis 73:335-337. Wheeler. Dorow, W. H. 0. and U. Maschwitz. 1990. The Dolichoderinae arachne-group of Polyrhachis: weaver ants 3. . Iridomyrmex humilis (Mayr). cultivating Homoptera on bamboo. Insectes SOC.37:73-89. Formicinae 12. Camponotini. Polyrhachis, 11 species. Hansen, L. D., and R. D. Akre. 1985. Biology of carpenter ants in Washngton state. Melanderia Change 4311-62. Cerapachyinae Change all Phyracaces to Cerapachys. Holldobler, B., and E. 0. Wilson. 1990. . Harvard Univ. Press, Cambridge. 732 p., Delete 24 pl. col. 9. Solenopsidini. Delete all references to the genus Vollenhovia. Letourneau, Deborah K. 1990. Code of ant-plant mutualism broken by a parasite. Science ENEMIES OF ANT LARVAE (1976:102) 248:215-217. Order Diptera Syrphidae: Trichopsomyia brood parasitoid on Masuko, Keiichi. 1986. Larval hemolymph feed- Polyrhachis. Microdon--many genera. ing: a nondestructive parental in the primitive ant Amblyopone silvestri Wheeler. Order Behav. Ecol. Sociobiol. 19:249-255. Eucharitidae: Add Pheidotoxenus. Add to hosts: Ectatomma, Messor. Formicidae: Phyracaces turneri feeds upon larvae of Pheidole. Vol. 5, No. 3-4, September-December 1991 173

Masuko, Keiichi. 1987. Leptanilla japonica: the Wheeler, G. C., and Jeanette Wheeler. 1987. first bionomic information on the enigmatic ant Young larvae of Veromessor pergandei. Psyche subfamily Leptanillinae. Pages 597-598 in 94~303-307. "Chemistry and Biology of Social " J. Eder and H. Rambold, eds. Proc. 10th Intern. Wheeler, G. C., and Jeanette Wheeler. 1988. Congr. I. U. S. S. I. 1986, Verlag Peperny, The larva of Leptanilla japonica, with notes on Munich. the genus. Psyche 95:185-189.

Masuko, Keiichi. 1990. The instars of the ant Wheeler, G. C., and Jeanette Wheeler. 1989a. Amblyopone silvestri. Sociobiology 17:221-244. Notes on ant larvae. Trans. Amer. Ent. Soc. 115~457-473. Picquet, N. 1958. Contribution a l'etude des larves de Formicidae de la C8te d'Or. Travaux Wheeler, G. C., and Jeanette Wheeler. 1989b. du Laboratoire de Zoologie et de la Station Notes on ant larvae: Ponerinae. J. New York Aquicole Grimaldi de la Faculte des Sciences Ent. Soc. 97:50-55. de Dijon. No. 23, 48 p. and 44 figs. Wheeler, G. C., and Jeanette Wheeler. 1990. Pierce, Naomi E. 1987. The and bioge- Larvae of the formicine ant genus Polyrhachis. ography of associations between lycaenid Trans. Amer. Ent. Soc. 116:753-767. butterflies and ants. Pages 89-116 in "Oxford Surveys in Evolutionary Biology." P. H. Harvey Wheeler, G. C., and Jeanette Wheeler. 1991. and Linda Partridge eds. Oxford Univ. Press. The larva of Blepharidatta. J. New York Ent. SOC.99~132-137. Taylor, R. W. 1965. A monographic revision of the rare tropicopolitan ant genus Probolomyrmex Wheeler, W. M. 1910. Ants. Columbia Univ. Press, Mayr. Trans. R. Ent. Soc. London 117:345-365. New York. 663 p.

Wheeler, G. C., and Jeanette Wheeler. 1976. Ant Larvae: Review and Synthesis. Mem. Ent. Soc. Washington, No. 7, 102 p.

Wheeler, G. C., and Jeanette Wheeler. 1986. Ten-year supplement to: "Ant Larvae: Review and Synthesis." Proc. Ent. Soc. Washington 88:684-702.