W. L. Brown, Jr.
Synonymic and Other Notes on Formicidae (Hymenoptera).
Psyche 56(1):41-49, 1949.

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SYNONYMIC AND OTHER NOTES ON
FORMICIDB (HYMENOPTERA) *
BY WILLIAM L. BROWN? JR.
The Biological Laboratories, Harvard University In 1945 Dr, E. V. Enzmann published a paper entitled "Systematic Notes on the Genus Pse~domyrrna.'~' Since this contains more confused taxonomy per page than any other work on the Formicidae I have ever en- countered in twelve years of reading in the field7 I have considered it advisable to publish an account of some of the synonymy involved,
The worst7 but by no means the only7 category of er- rors lies in the series of forms of Pseudomy~ma de- scribed as new from the types which Wheeler had set up in his "Studies of Neotropical Ant-plants and Their ant^^'^ published posthumously in 1942' and overlooked by Enzmann.
Wheeler's types were labelled as types in the usual Museum manner, and each series bore Wheeler's clearly legible determination label, Enzmann copied these names and used them in his paper7 creating a series of synonym- homonyms7 but since he made several mistakes in tran- scribing the spelling, some of the species may be consid- ered synonymous but not strictly homonymous. Of the remainder of Enzmann's publication, much may be safely ignored by taxonomists7 including the erratic keys and the pseudophylogenetic separation into "branches" and "gro~ps.~' Some forms described as new are from sources other than the Wheeler type material; since the Enzmannian types have not been made available for study7 it will devolve upon the future reviser of Pseudo- my~ma, a genus well-scrambled even in pre-Enzmannian times7 to decide the fate of the species not treated here. The species are listed as Wheeler had them7 each with the corresponding Enzmannian form beneath it. To *Published with a grant from the Museum of Comparative Zoology at Harvard College.
1 Psyche, 51: 59-103, 3 pls. (1945).
2 Bull. Mus. Comp. Zool. Harvard, 90: 1-262, 56 pls. (1942). 41




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42 Ps yc7xe
[Mar.
shorten the task, I have given date and page references only ; plate and' figure riferences are om&ed. Refer-
ences to Wheeler's 1942 and Enzmann's 1945 papers are given in the preceding footnotes. The Museum of Com- parative Zoology type catalog numbers are contained in parentheses with the initials (MCZ).
Pseudomgrma alliodora Wheeler
Wheeler, 1942, pp. 157-158.
P. allidora [sic!] E. Enzmann, 1945, pp. 77-78 (MCZ 20533).
Pseudomyrma belti subsp. safordi Wheeler Wheeler, 1942, p. 162.
P. sabumica var. safordi E. Enzmann, 1945, p. 89 (MCZ , 20537).
The term "sabanica" is evidently a misspelling of Wheeler's specific name satawka (P. sataka Wheeler, 1942, pp, 174-175), of which Enzmann considered saf- fo~di a variety.
Pseudomgrma belti subsp. ve~ifica Wheeler Wheeler, 1942, pp. 162-163.
P. belti subsp. ve~ifica E. Enzmann, 1945, p. 81 (MCZI 20538).
Pseudomyrma belti subsp. bequaerti Wheeler Wheeler, 1942, p. 164.
P. belti subsp. bequaerti E. Enzmann, 1945, pp. 80-81 (MCZ 23139).
Pseudomyrma lati~oda var. coromata Wheeler Wheeler, 1942, pp. 167-168
P. lathoda var. coro~ata E. Enzmann, 1945, p. 88 (MCZ 20542).
Pseudomgrma latinoda subsp. b~adle yi Wheeler Wheeler, 1942, p. 169.
P. bradleyi E. Enzmann, 1945, p. 82 (MCZ 22864).



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1949 1 Brow-Notes 0% Formicida 43
Pseudomyrma sericea var. acaciarum Wheeler Wheeler, 1942, p. 176.
P. sericea var. acaciorum [sic!] E. Enzmann, 1945, p. 90 (MCZ 22865).
Pseudomyrma spimkola subsp. sclerosa Wheeler Wheeler, 1942, pp. 180-181.
P. spimola [sic!] var. iufernalis E. Enzmann, 1945, p. 91 (MCZ 20547).
Pseudomyrrna spiuicola subsp. sclerosa Wheeler Wheeler, 1942, pp. 181-182.
P. spinola [sic !I var. sclerosa E. Enzmann, 1945, pp. 91- 92 (MCZ 23145).
Pseudomyrrna triplaridis subsp. baileyi Wheeler Wheeler, 1942, pp. 185-186.
P. triplaridis subsp. biolleyi [sic!] E. Enzmann, 1945, pp. 93-94 (MCZ 20548).
Pseudomyrma triplaridis subsp. tigriua Wheeler Wheeler? 1942? p. 186.
P. triplaridis subsp. trigma [sic!] E. Enzmann, 1945, pp. 94-95 (MCZ 23147).
Pseudomyrma t~iplaridis subsp. bozi Wheeler Wheeler, 1942, p. 184.
P. triplaridis subsp. bozi E. Enzmann, 1945, p. 94 (MCZ 23146).
The two following forms which Wheeler saw, but re- frained from describing, are easily synonymized with common species of Pseudomyrwza.
Pseudomyrrna gracilis (Fabricius)
Fabricius, 1805, Syst. Piez., p. 405 (Formica). P. gracilis var. longi~oda E. Enzmann, 1945, p. 87 (MCZ 268121.




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44 Psyche [Mar.
Pseudom yrma triplari~a (Weddell)
Weddell, 1849, Ann. Sc. Nat. Bot. (3) 13: 40-113, 249-268 (Myrmica) .
P. arboris-sancta E,mery, 1894, Bull. Soc. Ent. Ital., 26 : 147.
P. arboris-sancta var. ecuadoriana E. Enzmann, 1945, pp. 79-80 (MCZ 26809).
The types of ecuadoriana are few, partially frag- mented, and accompanied by what appear to be Agteca workers glued to the card with the ecuadoriana. I can see no characters which distinguish them (ecuadoriana) from a series of triplarina workers from several South American localities in the Wheeler Collection. Following the publication of Dr. Enzmann's paper on Pseudomyrma, others were published by his daughter, Miss Jane Enzmann. All but one of the species de- scribed, however, appear to be synonyms of common Nearctic forms. Dr. William S. Creighton has discussed these forms with me, and I am grateful for his opinions on several obscure cases. His forthcoming book, which amounts to a revision of North American ants, will also carry notes on the synonymy of these forms, but techni- cal difficulties prevent him from dealing with them at any length. Most species treated below involve Enzmannian names, but several other forms of older authors are changed in status as well.
Myrmecina americana Emery
M. latreillei subsp. americafia Emery, 1895, Zool. Jahrb. Syst., 8 : 271.
M. latreillei subsp. americana var. brevispi~osa Emery, idem.., p. 271.
M. g~am~nwola subsp. quadrispina J. Enzmann, 1946, Jour. N. Y. Ent. Soc., 54: 13-15, figs. I, 2, worker. In the manuscript of his work on North American ants, which he has kindly allowed me to examine, Dr. Creigh- ton has raised the form known for many years as Myrme- cina graminicola subsp. americana to the rank of species.



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19491 Brow-Notes on Forrnicidce 45
There appears little objection to this move, though the differences between the Palearctic .qramikola and the Nearctic form are very slight. he forms quadrispina and brevispinosa, however, cannot be considered valid forms.
The Enzmannian subspecies (quadrispina) was taken (holotype worker) on the south slope of the Blue Hills, a rather restricted elevated area just outside Boston, Massachusetts. Two colonies collected by me in this locality were confined for several months in artificial nests. Specimens killed at the time of collection and others examined after two months of rearing show a wide range of variation in size, sculpture and color. The larger workers, mostly those killed at the time of collec- tion, agree well with the description and figures, as well as my impressions, gained from a rather cursory exami- nation of the type, of quadrispina. These workers also agree with Emery's original description of arnericana and with specimens identified as arnericana by Wheeler and by Creighton.
My nests also produced, after a month or so of sub- starvation conditions, small light-colored workers corres- ponding well with published descriptions of brevispi~osa and with specimens determined as such in the Wheeler Collection. These workers were raised from small larv~ during a period in which the colonies refused all types of prepared foods, including bread and fats. When ripe seed-heads of timothy and some small herbaceous plants were later introduced, the colony eagerly accepted the seeds as food, but the workers which had previously hatched never became? even after four weeks? as fully colored as the workers reared in the wild. I conclude that the variant b~evispinosa is merely the stunted workers from either an incipient or poorly-nourished colony.
Both my nests were taken under large, well-embedded stones in a rbh, shady beech woods. Each colony occu- pied a small oval chamber in the soil, about three quar- ters of an inch in greatest diameter and less than a quar- ter inch deep, with the smooth lower surface of the stone



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46 Psyche [Mar.
forming the immediate roof.
The artificial nests were
set up on the evening of collection (June 10). A few
males also developed from the larvae taken with the nests, and these pupated during early August and de- veloped into adults in late August. All the males es- caped both nests through cracks during one night in early September, presumably on nuptial flight, since they had not previously attempted to leave the brood cham- ber which the ants constructed at the end of each nest from small particles of earth that had been scattered over the nest floor. These chambers were an almost exact replica of the ones found under the stones, open at the top and with a small passage at one side. The queens never left the brood except on the occasion of the introduction of the first grass seed, when all the workers and one queen left the brood and examined the seeds. The queen returned after a brief period and re- sumed her watch over the brood.
In studying various Myrmecina in the "Wheeler Collec- tion, I have seen other forms of very doubtful validity. All these are presently considered subspecies of grami- icola, under which Wheeler placed them in his original descriptions. Texana is supposed to differ from ameri- cana by its "scotch grain" shagreening of the first gas- tric segment. However, specimens from many localities in the states east of the Mississippi also possess this characteristic to a varying degree, and specimens from North Carolina and northern Ohio show much heavier sculpture of this type than do the texana types. The
texana types, however, do seem to differ slightly from americana in having a much less definitely longitudinal orientation to the rugulation of the head, with the longi- tudinal rug= having many prominent transverse spurs and branches. Other Texan specimens I have seen all belong to the typical americana, including a specimen identified by Wheeler as texana. Since sculpture ap- pears to be one of the several very unstable features of Holarctic Myrmeciha, I believe that further collecting in Texas and Mexico will show that this form is synonymous with americana.




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1949 I Brown-Notes on Formicida 47
Wheeler's two Oriental forms, grarninicola subsp. nipponica and graminicola subsp. sinensis, are also doubtful. The former has the anterior clypeal tubercles developed much as in graminicola, and seems hardly separable from that form. The latter has the clypeal tubercles reduced and seems scarcely distinguishable from americana. I should not be surprised if sinensis were to prove to be the same as sicula, from the southern Palearctic region; or if both of these (siczila and sinen- sis) were identical to americana. In fact, the entire ~olarctic Myrmecina fauna may end by being considered as one huge species dine in which the geographical races have not yet become sufficiently isolated to form distinct subspecies exclusively inhabiting a given area. Tetramorium cc~spitum (Linnaeus)
Linnaeus, 1785, Syst. Nat. (Ed. lo), 1 : 581 (Formica) Myrmica (Myrmica) brevinodis var. transversinodis J. Enzmann, 1946, Journ. N. Y. Ent. Soc., 54: 47-49, figs. 1, 2, worker.
Dr. Creighton and I are in complete agreement that this form (transversinodis) must be added to the long list of synonyms of the common pavement ant. Al- though I have not seen the type, the description, figures and notes on the habits leave little doubt of the correct placement. This ant should not be mistaken for Myr- mica lavinodis, listed under various names and possibly a subspecies of M. rubra, which is an introduced form quite common in the Boston area.
M. lavinodis some-
times enters houses, but then as
solitary individuals
probably brought in on clothing, as has been my frequent observation in Cambridge. This Myrmica possesses a very potent sting, the effects of which may last for sev- eral hours.
Crematogast er Uneolata (Say)
Say, 1836, Boston Jour. Nat. Hist., 1: 290, all castes (Myrmica) .
C. lineolata cerasi var. punctinodis J. Enzmann, 1946, Jour. N. Y. Ent. Soc., 54: 91-92, pi. 2, fig. 7, all castes.




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48 Psyche [Mar.
C. lineolata cerasi var. wizeldeni J. Enzmann, idem., p. 92, worker.
Dr. Creighton and I agree that these two forms either represent the typical lineolata or intergrade with what Dr. Creighton considers subsp. subopaca. Enzmann has raised cerasi Fitch to subspecific rank, but Dr. Creigh- ton's forthcoming book will show that this name must be dropped.
Crematogaster vermiculata Emery
Emery, 1895, Zool. Jahrb. Syst., 8: 286. Considered impossible of exact determination, but probably equivalent to vermiculata or an integrade be- tween vermiculata and a subspecies, are three forms de- scribed in a paper by Jane Enzmann in 1946.' These all have in common the name coachellai and the subgeneric classification as Crematogaster (Acrocodia), but here the consistency ends. The synonymous forms with page references to Miss. Enzmann's paper are as follows: C. lineolata subsp. coachellai " E. Enz. in lit., " p. 93, see. iii. C. sanguinea subsp. coachellai "E. Enzmann, in lit.," '
p. 95, couplet 19.
C. lineolata var. coachellai J. Enz-
mann, PI. 2 (p. 97), fig. 3.
The first of these three names is given in a grouped list with a superficial characterization of major sections only, the second appears in a dichotomous key, and the third appears in the legend to the plate. It is doubtful whether or not the authorship should be ascribed to E. Enzmann for the first two of these, even though it seems clear that such was intended. The types of these forms have not been made available to me for study, so I con- sider the form coachellai unrecognizable in the absence of a proper description.
In still another paper by Jane Enzrnann4 the tribe ApJzcenogastrini is set up, a category which is untenable. The genera included in this "tribe" have numerous in- tergrades with other groups of the Pheid~lini,~ to which 3 Jour. N. Y. Ent. Soc., 54: 91-92 (1946). 4 Journ. N. Y. Ent. Soe., 55: 147-152, (1947). 5 Pheidolini Emery, Rend. Acad. Sc. Bologna, (1913-14).



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Aphanogaster and Novomessor clearly belong. One as- tounding error is the appearance of Lobognathus as a sub-genus in the key on page 152. This appeared to be miscopy of a large label earlier placed by Dr. Creighton on an unidentified specimen of Veromessor: Creighton's label in the Wheeler Collection reads "lobognatkus new subspecies." The name must be considered a lapsus and a synonym of Verome~sor.~
Two of the species described in this paper are minor workers, probably from incipient nests, of two well- known North American ants, which are listed below. Novomessor albisetosus (Mayr)
Mayr, 1886, Verh. Zo01.-bot. Ges. Wien, 36: 443-446, (Aphcenogast er ) .
N. cockerelli var. minor J. Enzmann, 1947, pp. 147-148, PI. 8, top.
Aphcenogaster fulva Roger
Roger, 1863, Berl. Ent. Zeitschrift, 7: 190. A'phcenogaster fulva var. rubida J. Enzmann, 1947, pp. 147-148, PI. 8, bottom.
I have not considered other Enzmannian forms be- cause of my unfamiliarity with the groups concerned and because of my lack of time and taste for the task. The publications considered above should certainly sug- gest to all who examine them the need for some means of formal nullification of the published extremes of such irresponsible taxonomy.
6 Veromessor Forel, 1917, Bull. Soc. Vaud. Sc. Nat., 51: 235 (described as subgenus of Novomessor) .




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