George C. Wheeler and Jeanette Wheeler.
Supplementary Studies on Ant Larvae: Cerapachyinae, Pseudomyrmecinae and Myrmecinae.
Psyche 80(3):204-211, 1973.

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SUPPLEMENTARY STUDIES ON ANT LARVAE:
CERAPACHYINAE, PSEUDOMYRMECINAE
AND MYRMICINAE*
BY GEORGE C. WHEELER AND JEANETTE WHEELER Laboratory of Desert Biology, Desert Research Institute, University of Nevada System, Reno
89507
Subsequent to the publication of our first supplement on the ant larvae of the subfamily Cerapachyinae
(1964), our first paper on
Pseudomyrmecinae (1956) and several supplements on Myrrnicinae 1960, 1972, 1973) we have received from other myrmecologists so much additional material that it has now become necessary to pub- lish a supplement.
Genus PHYRACACES Emery
REVISION: The last sentence of our generic characterization ( I 964 : 69)
should read : Hypopharynx usually spinulose dorsally. Phyracaces elegans Wheeler (Fig. 2). Length (through spiracles) about 4.7 mm. Very similar to Ph. larvatus (1964: 69) except as follows. Body more slender. A pair of bosses on lateral surfaces of venter of AI-AVI. Spiracles small, A1 largest, diameter decreasing posteriorly. Integument densely spinulose, spinules in short to long, subtransverse to arcuate rows. Body hairs less numerous and shorter (0.025-0.05 mm long). Head hairs shorter (0.009-0.019 mm long). Posterior surface of labrum with a ventrally directed medial boss bearing 6 sensilla, about 5 sensilla on each lateral surface. Mandi- bles with narrower base. No spinules seen on hypopharynx. YOUNG LARVA: Length (through spiracles) about 2.1 mm. Simi- lar to mature larva above except as follows. Neck curved, abdomen with straight ventral profile and C-shaped dorsal profile. Body hairs shorter (0.01-0.033 mm long). Integument spinulose, spinules mi- nute, isolated laterally and in short rows dorsally and ventrally. Antennae less distinct. Maxillae lacking spinules; galea a slightly raised pair of sensilla.
No spinules seen on labium; opening of
sericteries a short slit.
VERY YOUNG LARVA:
Length (through spiracles) about I .5 mm. Entire body arcuate ventrally. Otherwise similar to young larva. Material studied: numerous larvae from New South Wales, courtesy of Rev. B. B. Lowery.
*Manuscript received by the editor September 28, 1973 Pu&e 80:204-211 (1973). http:llpsyche cnlclub ore/SWSOlM html



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WhecIer ‰â Wheeler - Ant Larvae
Fig. 1. Cerapachys (Sycia) australis: a, head in anterior view, X 95 ; b, left mandible in anterior view, X314; c, larva in side view, X22; d and e, two types of body hairs, X44-4. Fig. 2. Phyracaces elegans: head
in anterior view, X 74. Fig. 3. Tetraponera natalensis: left antenna in lateral view, X 339.
Phpcaces ficosus Wheeler. Length (through spiracles) about 4.4 mm.
Very similar to Ph. larvatus (1964: 69) except in the following details. Spiracles on first abdominal sornite slightly larger, remainder small and subequal. Body hairs shorter (0.013-0.063 mnl long). Antennae with 2 sensilla each. Head hairs shorter (0.008- 0.025 mm long). Galeae digitiform. (Material studied: 14 larvae from New South Wales, courtesy of Rev. B. B. Lowery.) Genus CERAPACHYS F. Smith
REVISION : Our generic characterization ( I 964 : 67) should be replaced with the following: Leg vestiges small paraboloidal papillae. Body hairs usually simple. Head (including mouth parts) subpyri- form in anterior view. Head hairs usually short. Mandibles long, slender and with median border erose. Maxillary palp short; galea long and digitiform.
Cerapachys ( Syscia) australis Fore1 ( Fig. I ) . Length ( through spiracles) about 3.2 mm.
Body long and subcylindrical; about 12




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206 Psyche [September
differentiated somites; head on anterior end; a small posteriorly projecting boss on AX. Anus ventral. Spiracles small. Entire integument densely spinulose, spinules minute and in short to long straight or arcuate rows. Body hairs short, uniformly distributed and moderately numerous. Of two types: ( I) 0.025-0.063 mm long, mostly bifid, sometimes with one or both branches rebranched, on all somites;
(2) 0.037-0.05 mm long, simple, a few on each somite. Cranium subhexagonal in anterior view, slightly longer than wide. Antennae large, each a low mound with 3 minute sensilla, each bearing a minute spinule.
Head hairs few, 0.025-0.05 mm long,
simple or bifid. Labrum subarcuate, about twice as wide as long; anterior surface with 8 sensilla on and near ventral border; posterior surface with about 6 sensilla ventromedially and with a few oblique arcuate rows of minute spinules. Mandibles narrowly subtriangular in anterior view; apex rather long, narrow and heavily sclerotized; medial border with 6-8 small denticles. Maxillae with apex para- boloidal and sparsely spinulose, spinules minute to short and in a few arcuate rows; palp a peg with 4 (2 encapsulated and 2 bearing a spinule each) apical and one lateral sensilla; galea digitiform with 2 apical sensilla, each bearing a minute spinule. Labium subtrape- zoidal, widest distally, anterior surface densely spinulose, spinules minute and in numerous short arcuate rows; palp a rounded elevation with 5 (2 encapsulated and 3 bearing a spinule each) sensilla; an isolated sensillum between each palp and opening of sericteries; the latter a slit in a shallow depression on anterior surface. Hypopharynx with minute spinules in long transverse sub-parallel rows. (Material studied : 10 larvae from Queensland, courtesy of Rev. B. B. Lowery.) We have never been able to key the genera of this subfamily. Except for head shape, where the difference in the species of Pachy- simu is greater than that between any two genera, some of the vari- ants of any character in any genus can be found in other genera. Bernard (1951 : 1053) included larval characters in his character- ization of the subfamily, which he called family Promyrmicidae. Sudd ( 1967 : 123) discussed the feeding of the larvae. He stated (erroneously) that the trophothylax was formed by the bases of the rudimentary legs; we have shown (1956: 375, 383) that it is "formed from the depressed ventral surface of the thorax and elab- oration of the first a,nd second abdominal somites."



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Wheeler &f Wheeler - Ant Larvae
Genus PSEUDOMYRMEX Lund
Janzen (1967 : 344). Beltian bodies are cut up by the workers and fed to the larvae.
The following species of Pseudomyrmex are compared with Ps. alliodorae 1956: 379) ; only differences are given here. Pseudomyrmex ahstus Borgmeier. Length (through spiracles) about 4.8 mm; straight length about 4.6 mm. Body hairs: (2)
0.05-0.25 mm long, longest of AI-AV; (3) 0.2-0.25 mm long, 2 only on each TI-3 and AI-AIII. Head hairs more numerous and 0.05-0.25 mm long. Posterior surface of labrum with a cluster of 3 sensilla in the middle of each half. (Material studied : 7 larvae from Brazil, courtesy of Dr. I<. Lenko.)
Pseudomyrmex belti fulvescens Emery (= Ps. ferrugineus I?. Smith). Janzen 1967 : Description p. 394; feeding of larvae p. 416- 417; handling of larvae p. 418. Similar information in Janzen I 966.
Pseudomyrmex elongatus Borgmeier. Length ( through spiracles) about 3.8 mm; straight length about 3.6 mm. Body hairs longer: ( I) 0.006-0.018 mm long; (2) 0.018-0.2 mm long; (3) 0.175-0.22 mm long, 4 in a row across the dorsum of each TI-3 and AI-AIV. Head hairs more numerous and slightly longer (0.0 I -0.05 mm long). (Material studied: 15 larvae from Brazil, courtesy of Dr. K. Lenko.) Pseudonzyrmex schupfii Forel. Length (through spiracles) about 5.9 mm; straight length about 5.7 mm. Largest spiracles on AI. Body hairs longer: (I) 0.013-0.025 mm long; (2) 0.038-0.25 mm long; (3) 0.25-0.33 mm long, 4 in a row across the dorsum of each TI-3 and AI-AIII. (Material studied: numerous larvae from Brazil, courtesy of Dr. K. Lenko.)
Pseudomyrmex subtilissimus Emery. Length (through spiracles) about 4.3 mm ; straight length about 3.9 mm. Body hairs ( I) 0.006- 0.018 mm long; (2) 0.018-0.15 mm long; (3) about 0.15 mm long, 4 in a row across the dorsum of each TI-3 and AI-AIV. Head hairs slightly longer (0.025-0.05 mm long).
( Material studied : 6 larvae
from Brazil, courtesy of Dr. K. Lenko.)
Pseudomyrmex termitarius F. Smith. Length (through spiracles) about 5.1 mm; straight length about 4.7 mm. Body stouter. Body hairs about twice as numerous: ( I ) 0.013-0.05 mm long; (2) 0.05- 0.275 mm long; (3) 0.25-0.35 mm long, 4 in a row across dorsum of each TI-3 and AI-AIV. Head hairs more numerous, longer (0.013-0.05 mm long) and finely denticulate. Labrum with width twice the length, with anterior lobes more prominent and with 2



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208 Psyche [September
minute hairs on anterior surface. Mandibles with teeth stouter and blunter ; lateral outline less curved ; denticles on anterior surface more numerous. Maxillary apex less constricted and with spinules longer and covering a greater portion of the surface. Labium with more numerous spinules. (Material studied: 9 larvae from Brazil, courtesy of Dr. K. Lenko.)
Genus TETRAPONERA I?. Smith
Tetraponera natalensis F. Smith (Fig. 3). Length (through spira- cles)
about 8.2 mm; straight length about 6.2 mm. Similar to T. aitkeni ( 1956 : 388) except as follows. Body slightly stouter at AV and AVI.
Integument of AIX and AX with minute spinules. Body hairs: ( I ) 0.008-0.075 mm long; (2) 0.025-0.15 mm long, longest with tip branched or denticulate;
(3) 0.175-0.3 mm long, 4 in a
row across the dorsum of each TI-3 and AI-AVI. Each antenna represented by 3 individually raised sensilla on a small base. Head hairs longer (0.013-0.1 I mm long) and less numerous, with or without alveolus and articular membrane, some with denticles near the tip. Labrum with breadth less than twice length; borders sinu- ate; anterior surface with 6 sensilla and 2 hairs on each half; pos- terior surface with 9 sensilla on each half; spinules as in T. aitkeni. Anteromedial surface of mandibles with large spinules, which are isolated or in short rows of 2 or 3. Maxillae with rather numerous long spinules in short arcuate rows; palp represented by a cluster of 5 sensilla on a slight elevation. (Material studied: numerous larvae from South Africa, courtesy of Dr. W. L. Brown.) Genus PACHYSIMA Emery
Pachysvna latifrons Emery : Bernard ( I 95 I : 1054-1057) de- scribed and figured the young (after W. M. Wheeler). Genus VITICICOLA Wheeler
Fiticicola tessmanni (Stitz) : Bernard ( 195 I : 1054) described and figured the larva (after W. M. Wheeler). SUBFAMILY MYRMICINAE
Ettershank ( 1966 :
I 61, 162) : "The larvae of the Forrnicidae have not been used to any extent in taxonomic studies, although numerous descriptions and figures of scattered genera and species occur in the literature. The only wide-scale comparative larval study



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19731 Wheeler ‰â Wheeler - Ant. Larvae 3-39 that has been attempted is the series of papers by G. C. Wheeler (later with J. Wheeler), which constitute a fundamental contribu- tion to the subject that will be used for a long time." "Reference to all the publications by the Wheelers on myrmicine ants are contained in a summary article (G. C. and J. Wheeler 1960). In this paper, the authors conclude that three characters are of major importance: body profile, mandible shape,
and setal form. They recognize 22
body profiles and 30 mandibular shape categories all of which are explained and illustrated."
Genus MESSOR Forel
Messor capitatus Latreille:
Delage ( 1968a) gave in a table the
sizes and abundance of larvae throughout the year. She stated that only small larvae overwinter. She (1968b) discussed larval enzymes and digestion.
Genus PHEIDOLE Westwood
Kempf (1972:
457) : Ph. vallifica is the host of the eucharitid Orasenza costaricensis Wheeler and Wheeler. Genus MELISSOTARSUS Emery
Delage-Darchen ( 1972a) : Hairs few, long, with bifid tips. Crude sketch of a larva on p. 219.
Genus CREMATOGASTER Lund
Delage-Darchen ( 1972b) found only three larval stages in C. (Nematocrema) stadelmanni Mayr. Fig. I hairs enlarged ; Fig. 2 and 3 larvae of various stages in side view; Fig. 4 head in anterior view. Pilosity is taxonomically worthless because of extreme vari- ation between colonies and even in the same colony. Genus MONOMORIUM Mayr
Cloudsley-Thompson ( I 962 : I 79) : The calliphorid flies Ben- galia peuhi Vil. and B. minior Malloch fed on the larvae of M. saZomonis (Linnaeus) in the central Sudan. Van Pelt and Van Pelt ( 1972 : 978) : Larvae of the syrphid Microdon baliopterus Loew fed upon the larvae of M. minimum ( Buckley ) .




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Psyche
[September
Genus SOLENOPSIS Westwood
Markin et al. (1972: 1053) : Life cycle of Solenopsis invicta Buren in an incipient colony: egg 6-8 days, la,rva 14-15 days, pupa 20-24 days.
Genus TETRAMORIUM Mayr
Tetramo?-ium caespitum (Linnaeus) . Donisthorpe ( 1927 : 197) : "The larvae were fed with disgorged liquid food as long as they were young and gathered together in groups, but when they grew older and were separated, the workers fed them with solid sub- stances." Many larvae were hung on to the plaster walls of the nest by their anchor-tipped hairs.
BERNARD, F.
1951. Super-famille des Formicoidea. Trait6 de Zoologic, Tome X, Fasc. 11: 907-1119, 1258-1263, 1272-1275. CLOUDSLEY-THOMPSON, J, L.
1962. A note on the association between Bengalia spp. (Dipt., Cal- liphoridae) and ants in the Sudan. Entomol. Monthly Mag. 98: 177-179.
DELAGE, BERNADETTE.
1968a.Recherches sur la fourmis moissoneuses du Bassin Aquitain: 6cologie et biologic. Bull. Biol. 100 : 315-367. 1968b.Recherches sur les fourrnis moissoneuses du Bassin Aquitain: kthologie. Physiologie de l'alimentation. Ann. Sci. Nat., 2001. Biol. Anim. (12) 10 : 197-265.
1972a. Une fourrni de Cote-dlIvoire: Melissotarsus titubans Del., n. sp. Insectes Sociaux 19 : 213-226.
1972b. Le polymorphisme larvaire chez les fourmis Nematocrema dlAfrique. Insectes Sociaux 19 : 257-277. DONXSTHORPE, H. ST. J. K.
1927. British ants. Geo. Routledge & Sons, London. 436 pp. ETTERSHANK, G.
1966. A generic revision of the world Myrmicinae related to Solenopsis and Pheidologeton. Australian J. 2001. 14: 73-171. JANZEN, D. H.
1966. Coevolution of mutualism between ants and acacias in Central America. Evolution 20 : 249-275.
1967. Interaction of the bull's-horn acacia (Acacia cornigera L.) with an ant inhabitant (Pseudornyrmex ferruginea F. Smith) in eastern Mexico. Univ. Kansas Sci. Bull. 47: 315-558. KEMPF, W. W.
1972. A study of some Neotropical ants of genus Pheidole Westwood. I. Studia Entomol. 15 : 449-464.




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19731 Wheeler &? Wheeler - Ant Larvae 21 I MARKIN, G. P., H. L. COLLINS AND J. H. DILLIER. 1972. Colony founding by queens of the red imported fire ant, Solenop- sis invicta. Ann. Entomol. Soc. Amer. 65: 1053-1058. SUDD, J. M.
1967.
An
introduction to the behavior of ants. St. Martin's Press, New York. 200 pp.
VAN PELT, A. F., AND S. A. VAN PELT.
1972. Microdon (Diptera: Syrphidae) in nests of Monomorium in Texas. Ann. Entomol. Soc. Amer. 65 : 977-979. WHEELER, G. C., AND JEANETTE WHEELER.
1956. The ant larvae of the subfamily Pseudomyrmecinae. Ann. Ento- mol. SOC. Amer. 49 : 374-398.
1960. Supplementary studies on the larvae of the Myrmicinae. Proc, Entomol. Soc. Washington 62 : 1-32.
1964. The ant larvae of the subfamily Cerapachyinae: supplement. Proc. Entomol. Soc. Washington 66 : 65-71. 1972. Ant larvae of the subfamily Myrmicinae: second supplement on tribes Myrmicini and Pheidolini. J. Georgia Entomol. Soc. 7: 233-246.
1973a. The ant larvae of six tribes: second supplement. J. Georgia Entomol. SOC. 8: 27-39.
1973b. Ant larvae of four tribes: second supplement. Psyche 80: 70-82. 1973c.Ant larvae of the myrmicine tribe Attini: second supplement. Entomol. Soc. Washington. (In press.)
1973d. The ant larvae of the tribes Basicerotini and Dacetini: second supplement. Pan-Pacific Entomol. (In press.)



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