W. M. Wheeler.
The Subfamilies of Formicidae, and other Taxonomic Notes.
Psyche 27(2-3):46-55, 1920.

This article at Hindawi Publishing: https://doi.org/10.1155/1920/14961
CEC's scan of this article: http://psyche.entclub.org/pdf/27/27-046.pdf, 888K
This landing page: http://psyche.entclub.org/27/27-046.html

The following unprocessed text is extracted automatically from the PDF file, and is likely to be both incomplete and full of errors. Please consult the PDF file for the complete article.

46 Psyche [April-June
THE SUBFAMILIES OF FORMICID.E, AND OTHER TAXONOMIC N0TES.l
A comparison of the seventh volume of Dalla Torre's " Catalogus Hymen~pterorum,~' which summarizes what was known of the classification of the Formicidse down to 1890, with any very recent monograph of these insects, gives the impression that there has been no change in expert opinion concerning the limits of the family and its subfamilies during the past thirty years. Dalla Torre recognizes five subfamilies, the Dorylinse, Ponerinse, Myrmi- cinse, Dolichoderinse and Camponotinse and the same groups are retained in Emery's contributions to the "Genera Insectorumyy (1910-'13), so far as published, and in his recent sketch of the classification of the Myrmicinse (1914). Between the appearance of the " Catalogus " and the works just mentioned, however, Emery, who has shown greater interest than other myrmecologists in the definition of taxonomic categoriesabove the rank of the genus, proposed an additional subfamily, the Pseudomyrminse in 1899, and in 1895 transferred a group of genera, comprising the tribe Cerapachyini, from the Ponerinse, where it had been placed by Fore1 in 1893, to the Dorylinse. After Fore1 and I had objected to this proceeding, Emery, in the "Genera In~ectorurn'~ (1913) returned the Cerapachyini to the Ponerinse, but gave them the rank of a section, the Prodorylinse. He had long since reunited the Pseudomyrminse with the Myrmicinae. In his most recent sketch of the classification of this subfamily (1914) he unites the tribes Metaponini and Pseudomyrmini as the first section, the Promyrminse, and places all the other tribes in a second section, the Eumyrmicinse. Thus in 1920 the five subfamilies have again acquired the limits which they had in 1890. During the past year a study of ant-larvse, representing more than a hundred genera and many subgenera of all five subfamilies, has convinced me that Emery was right in 1899, when he regarded the Pseudomyrminse as constituting an independent subfamily. I am also of the opinion that the Cerapachyini should be removed IContributions from the Entomological Laboratory of the Bussey Institution, Harvard University. No. 169.
Pu&e 27:46-55 (1920). hup ttpsychu einclub orgt27/27-M html



================================================================================

19201 WheelerÌÔSubfamiIie of Formicidae and Other Taxonomic Notes 47 from the Ponerinae and raised to the rank of an independent sub- family, between the Dorylinse and Ponerinse. A number of reasons may be adduced for making these changes. In 1899 Emery, after a comparative study of the larvae of several Formicid genera, concluded that "Those of Sima and Pseudomyrma, besides their extremely hypocephalic development, exhibit a very special character in the presence of rudiments of antennae. I believe that this very noteworthy fact, together with the well- known peculiar characters of the head of the imagines, will justify the separation of these genera from the remainder of the Myrmi- cine, to form the new subfamily of the Pseudomyrminae." My study of numerous species of this group, which now embraces four genera, Tetraponera Smith ( = Sima Roger), Pachysima Emery and Viticicola Wheeler of the Old and Pseudomyrma Lund of the New World, shows that Emery was far from realizing the full import of their larval characters. Not only have the larvae peculiar long, straight, cylindrical, distinctly segmented bodies with blunt ante- rior and posterior ends, a large, usually subquadrate head, ventrally placed and with rudiments of antennae (which are also present in the larvae of many other ants, notably in the Ponerinae), but the thoracic and first abdominal segments are furnished with peculiar exudatory papillae (exudatoria), which form a cluster around the mouth. I have described and figured these organs in Viticicola and Paslysima (1918b) and have shown that they have the form of extraordinary appendages in the first larval stage (trophidium) of the two known species of the latter genus, and that the swollen ventral portion of the first abdominal segment, just behind the mouth, forms a pocket in which the workers place a pellet of food. The exudatoria, the pocket, which I call the trophothylax, and the unusual method of feeding are characteristic of all four genera and no distinct traces of such conditions have been found in any other ant-larvae.
More recent study has added two very interesting facts, which, in advance of a complete account to be published in collaboration with my colleague, Prof. I. W. Bailey, may be briefly considered in this place. The food pellet proves to be merely the small pellet ("corpuscule enroulb," or c'corpuscule de nettoyage" of Janet) which the worker ant moulds
in its own infrabuccal pocket
and consists of the solid food-particles from which the juices are



================================================================================

48 Psyche [April-June
sucked, plus the various particles collected by the ant by means of the strigils of the fore tibiae from the surfaces of the antennae and other parts of the body and carried into the infrabuccal pocket after being wiped off by the maxillae. Other ants eventually spit out the pellet which is commonly a moulded, subspherical conglomer- ate of diverse particles, such as small pieces of insects, fragments of plant tissue, fungus spores and hyphse, pollen grains, etc., and cast it away as refuse, but the worker nurses of thePseudomyrm- inse place it as pabulum in the trophothylax of the larva! Even this, however, is not the whole story. An examination of
the mouth of the larva reveals a singular structure, evidently used for reducing the food pellet to such a finely divided state that it can, when acted on by the digestive juices of the mesenteron, yield a certain amount of nutriment, which the worker ant could not extract from it while it was in the infrabuccal pocket. This larval structure, which may be called the trophorhinium, consists of two flat, opposable plates, the dorsal and ventral surfaces of the buccal cavity, each furnished with very fine, parallel, transverse striae or welts, which, under a high magnification are seen to be made up of minute chitinous projections or spinules. The ventral usually has *more numerous rows of spinules than the dorsal sur- face. The two surfaces are evidently rubbed on one another and thus triturate the substance of the food pellet, only small portions of which are ingested at a time from the trophothylax. In all Pseudomyrmine larvae and in many larvae of the other subfamilies, except the Dorylinse and Cerapachyinse, the trophorhinium is beautifully developed, although in many ants (Ponerinse) it may be used for comminuting parts of insects given directly to the larvae by the workers. A detailed description of the organ and of its extraordinary variations of structure in the various genera of Formicidse is reserved for future publication. In its development the trophorhinium bears a strange resem- blance to the stridulatory organs of the petiole and postpetiole of many adult Ponerinse and Myrmicinse. It may, in fact, function also as a stridulatory organ, when the food supply is exhausted, and thus apprise the worker nurses of the larva's hunger. Many ant-larvse, notably those of the Ectatommiine Ponerinse and of most genera of Camponotinse (Formicinse) , also have elaborate but coarser stridulatory surfaces on the mandibles, so that the larva



================================================================================

19201 Wheeler-Subfamilies of Formicidae and Other Taxonomic Notes 49 may be able to produce a variety of sounds and therefore apprise the nurses of more than one need or craving. The adult Pseudomyrminse are so peculiar in structure that Emery, Ashmead (1905) and others have been led to separate them sharply from all other Myrmicinse. The shape of the head in the worker and female and especially of the clypeus and frontal carinse is unique, the eyes are very large and there is a strong tendency to development of ocelli in the workers, the conformation of the pet- iole, postpetiole and tibia1 spurs is peculiar, and as I have recently shown (1919b), the number of antenna1 joints (12) is the same in the male as in the worker and female in all four genera. Fig. 1.
a, Ingluvies, or "crop," b, calyx of proventriculus, or "gizzard," and c, ventriculus, or "stomach," of Pachysima aethiops Fabr.; d, proventriculus seen
from the front under a higher magnification. Little study has been devoted to the structure of the proventri- culus, or "gizzard" in the Myrmicinse, but Meinert, Fore1 and Emery have described and figured it as simple and tubular in most genera and of a very primitive type compared with the conditions in the Dolichoderinse and Camponotinse. I find, however, that the proventriculus of all four genera of the Pseudomyrminse is much more specialized, being anteriorly developed as an apple- or quince-shaped ball, covered with longitudinal and circular muscles and with four distinct, connate sepals, bluntly rounded and finely hairy at their tips, and posteriorly as a very short, tubular, con-



================================================================================

50 Psyche [April-June
stricted portion which projects as a button into the cavity of the ventriculus (Figs. 1 and 2).
The peculiarities mentioned seem to
me to justify us in returning to Emery's contention of 1899 that the Pseudomyrminse constitute an independent subfamily. I have endeavored to show in a recent paper (1919a) that neither the larval nor the imaginal Metaponini can be regarded as at all closely related to the Pseudomyrminse. Emery's section Pro- myrmicinse should therefore be abandoned and his term Eumyrmi- cinse may be regarded as merely synonymous with Myrmicinse. Fig. 2. Viticicola tessmanni Stitz; a, sagittal section through part of the ali- mentary tract, including a, the ingluvies, or "crop" (much contracted); b, calyx of proventriculus, or "gizzard," x, its cylindrical portion, and c, anterior portion of ventriculus, or "stomach."
A study of the larvae of the Cerapachyini shows that they are extremely like the larvae of the Dorylinse. This was noticed by
Emery in his observations on the larva of Acanthostichus serratulus (1899). The mandibles are, small, narrow, pointed and rather feebly chitinized, and I have failed to find a trophorhinium in either group. Apparently the young are fed only on soft food. That the foraging habits of certain Cerapachyini (Phyracaces) resemble those of the Dorylinse was shown in my paper on the Australian species (1918a). We know nothing of the pupae, but they are probably not enclosed in cocoons as in the Ponerinse. Although the worker of the Cerapachyini has a Ponerine habitus, the characters of the female in the various genera are peculiarly diverse. In some cases (Phyracaces), this caste is winged and not unlike the females of certain Ponerinse, in others (Parasyscia, Eusphinctus) the female is wingless and ergatomorphic and in still others (Acanthostichus, Nothosphinctus) the female is so much like the corresponding caste in the Dorylinse, that it might be regarded



================================================================================

19201 Wheeler-Subfamilies of Formicidae and Other Taxonomic Notes 51 as a dichthadiigyne. A similar diversity is seen in the males of the Cerapachyini. The male of Acanthostichus afflictus, recently discovered by Gallardo (1919) in Argentina, is so much like an Eciton or Dorylus male that even an expert myrmecologist would not hesitate to place it among the Dorylinse. The males of other genera (Lioponera, Phyracaces, Cerapachys, Eusphinctus) on the ' other hand, though lacking the cerci, have a decidedly Ponerine habitus. It would seem, therefore, that the Cerapachyini are intermediate between the Dorylinae and Ponerinse, as Emery has contended, and that we might unite them with either. I should prefer, however, to separate them out as an independent sub- family, which may be ascribed to Forel, who in 1893 first recog- nized the "Cerapachysii" as a natural tribe. Of course, the name Prodorylinse Emery cannot be used for the subfamily, because there is no genus Prodorylus.
For many years I have deemed it necessary to introduce another nomenclatorial change, namely that of the subfamily name Camp- onotinse to Formicinse. Forel, in his study of the poison apparatus and anal glands of ants, published in 1878, divided the subfamily Formicidse Mayr (1855) into two subfamilies, which he called Camponotidse and Dolichoderidse. This was unjustifiable accord- ing to our present rules of nomenclature, for Mayr's name should have been retained and restricted to the group containing the genus Formica. At that time, which antedated the use of ince as a subfamily suffix, Fore1 justified his course on the ground that C 6
Formicidse" was already in use as a family name. Owing to the fact that definite rules and conventions in regard to the suffixes of family and especially of subfamily names in Zoology have been stabilized only within recent decades, there is considerable confusion concerning the authors to whom our modern names in idee and ince are to be attributed. It seems to be custom- ary to accredit a family or subfamily name to the author who first recognized the group as supergeneric and gave it a Latin or Greek name based on that of one of its genera. If this is done in the case of the Formicidse the authorities cited in the literature require revision. Frederick Smith (l85l), Westwood (l84O), Shuckard (1840) and Stephens (1829) all attribute Formicidse as a family name to Leach. They appear to refer to his article published in the Edinburgh Encyclopaedia in 1815, where he used the term



================================================================================

52 Psyche [April-June
Formicarides, or to some later work which I have not seen. La- treille, however, as early as 1810, used Formicarii as a family name, and it would seem to be permissible to cite him as the author of Formicidse. The subfamily Dorylinse is attributed by Emery and others to Shuckard (1840), but this author says: "Mr. Haliday has first raised them to a family equivalent to the whole of the social Ants, etc." and at p. 195 he definitively attributes the Dorylidse to Haliday. This may have been based on correspond- ence as I find no mention of the term in such published writings of Haliday as I have seen. But the matter is of little moment because Leach, in the 1815 paper referred to above, created a family Dorylida, so that, unless there is an earlier authority, the subfamily Dorylinse should be accredited to this early British entomologist. Fore1 attributes the subfamilies Ponerinse and Myrmicinse to Lepeletier, but Dalla Torre gives Mayr as the author of the latter and Donisthorpe refers the Ponerinse also to Mayr. Smith regarded himself as the authority for Poneridse and Myrmi- DORYLINAE
CERAPACHYINAE
ANCESTORS
Fig. 3.
Phylogenetic relationships of the seven subfamilies of Formicidae.



================================================================================

19201 WheelerÌÔSubfamilie of Formiddae and Other Taxonomic Notes 53 cidse. It is clear, nevertheless, that not only the Ponerinse and Myrmicinse but also the Formicinse are to be referred to Lepeletier (1836), who called them respectively the tribes Ponbrites, Myrmi- cites and Formicites, the last, like Mayr's subfamily Formicidse, being made to include both the modern Dolichoderinse Fore1 and Formicinse (Camponotinse Forel).
The phylogenetic relations of the seven subfamilies, as under- stood at the present time, are indicated in the accompanying dia- gram (Fig. 3). For taxonomic purposes they may be most con- veniently arranged in the following linear sequence: Family Formicidae Latreille (1 9 10).
Subfamily 1. Dorylinse (Leach 1815)
2. Cerapachyinse (Fore1 1893)
3. Ponerinse (Lepeletier 1836)
4. Pseudomyrminse (Emery 1899)
5. Myrmicinae (Lepeletier 1836)
6. Dolichoderinae (Fore1 1878)
7. Formicinse (Lepeletier 1836)
In conclusion I may add that while working on the ants of the Belgian Congo and constructing dichotomic keys for the identi- fication of the genera and subgenera of the world, I have been led to adopt the following new names based on previously described species :
Phrynoponera gen. nov. (Genotype: Bothroponera gabonensis Ern. Andre)
Viticicola gen. nov. (Genotype: Sima tessmanni Stitz) Macromischoides gen. nov. (Genotype: Macromischa aculeata Mayr)
Hypocryptocerus subgen. nov. (Subgenotype: Formica hc~morrhoi- dalis Latreille)
Heteromyrmex gen. nov. (Genotype: Vollenhovia rufiventris Forel.) Diodontolepis gen. nov. (Genotype: Melophorus spinisquamis Ern. Andre)
Pseudaphomomyrmex gen. nov. (Genotype : Aphomomyrmex emeryi Ashmead)
Cladom yrma gen. nov. (Genotype : Aphomom yrmex hewitti Wheeler).



================================================================================

54 Psyche [April-June
1905.
ASHMEAD, W. H. A Skeleton of a New Arrangement of the Families, Subfamilies, Tribes and Genera of the Ants, or the Superfamily Formicoidea. Canad. Ent. 1905, pp. 381-384.
1877. EMERY, C. Saggio di un Ordinamento Naturale dei Myrmicidei e Considerazioni sulla Filogenesi delle Formiche. Bull. Soc. Ent. Ital. 9, 1877, pp. 67-81, 1 pi. 1895. EMERY, C. Die Gattung Dorylus Fabr. und die systema- tische Eintheilung der Formiciden. Zool. Jahrb. Abt. Syst. 8, 1895, pp. 685-778, 3 pis., 41 text-figs. 1899. EMERY, C. Intorno alle Larve di Alcune Formiche. Mem. R. Accad. Sc. 1st. Bologna 8, 1899, pp. 3-10,2 pis. 1909. EMERY, C. Beitrage zur Monographic der Formiciden des palaarktischen Faunengebietes. Deutsch. Ent. Zeit- schr. Teil VIII Ponerinffi, 1909, pp. 355-376. 1910-12. EMERY, C. Dorylinse (l9lO), Ponerinse (1911) and Dolichoderinse (1913) in Wytsman's " Genera Insec- torum."
19 14. EMERY, C. Intorno alla Classificazione dei " Myrmicinse." Rend. R. Accad. Sc. 1st. Bologna 1914, pp. 29-42. 1878. FOREL, A. Der Giftapparat und die Analdriisen der Ameisen. Zeitschr. wiss. 2001. 30, 1878, pp. 28-66, 2 pis. 1893. FOREL, A. Sur la Classification de la Famille des Formi- cides avec Remarques Synonymiques.
Ann. Soc. Ent.
Belg. 37, 1893, pp. 161-167.
19 19. GALLARDO, A. Una nueva Prodorilina Acanthostichus afflictus.
Anal. Mus. Nac. Hist. Nat. Buenos Aires 30, 1919, pp. 237-242, 3 figs.
1815. LEACH, W. E. Article "Entomology" in Brewster's Edinburgh Encyclopaedia, 1815, pp. 57-172. 1836. LEPELETIER, A. DE ST. FARGEAU. Histoire Naturelle des Insectes. Hymhopt2res, 1, 1836. Paris, de Roret. 1855. MAYR, G. Formicina Austriaca. Verh. zool. bot. Ver. Wien 5, 1855, pp. 373-478, 1 pi.
1877. MAYR, G. Gruppirung der Myrmiciden. Sitzb. zool. bot. Ges. Wien, 27, 1877, 4 pp.




================================================================================

19201 Wheeler-Subf amilies of Form-icidae and Other Taxonomic Notes' 55 1840. SHUCKARD, W. E. Monograph of the Dorylidse, a Family of Hymenoptera Heterogyna. Ann. Nat. Hist. 5, 1840, pp. 188-231.
1851.
SMITH, F.
List of the British Animals in the Collection of the British Museum. 4, Hymenoptera ~cuieata. London
1851, 134 pp.
1829. STEPHENS, J. F. A Systematic Catalogue of British Insects.
London, Baldwin and Cradock, 1829.
1840. WESTWOOD, J. 0. Introduction to the Modern Classifica- tion of Insects. Vol. 2, London, 1840.
1918a. WHEELER, W. M. The Australian Ants of the Ponerine Tribe Cerapachyini. Proc. Amer. Acad. Arts Sc. 53, 1918, pp. 215-265,17 figs.
1918b. WHEELER, W. M. A study of Some Ant Larvae, with a Consideration of the Origin and Meaning of the Social Habit among Insects. Proc. Amer. Phil. Soc. 57, 1918, pp. 293-343,152 figs.
1919a. WHEELER, W. M. The Ants of the Genus Metapone Forel. Ann. Ent. Soc. Amer. 12, 1919, pp. 173-191, 7 figs.
1919b. WHEELER, W. M. A Singular Neotropical Ant (Pseudo- myrma filiformis Fabricius) . Psyche 26, 1919, pp. 124- 131, 3 figs.
ODONATA OF CHATHAM, MASSACHUSETTS.
BY R. HEBER HOWE, JR.,
Thoreau Museum of Natural History, Concord, Mass. The following list of Odonata includes material collected last summer at Chatham, and also that taken on various excursions to the surrounding towns. Mr. C. W. Johnson had collected a few species at Eastham of which I make mention, and other species have been recorded from Provincetown, Cotuit, Hyannisport, Woods Hole, Martha's Vineyard, the Elizabeth Islands, beside those listed by the author from Nantucket (May, 1919, report Maria Mitchell Association), and from Wareham (Psyche 26 : June, 1919). Specimens of all recorded material are in the author's collection.




================================================================================