E. V. Enzmann.
Systematic Notes on the Genus Pseudomyrma.
Psyche 51:59-103, 1944.

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PSYCHE
SYSTEMATIC NOTES ON THE GENUS
PSEUDOMYRMA1
BY E. V. ENZMANN
The Haskins Laboratory
The genus Pseudomyrma is one of the most difficult of the genera of the family Formicidse and few systematists have at- tempted to explore the intricacies of its classification. The genus was created by Latreille, 183 1 ; up to the middle of the past century only a few forms were known of which Fabricius described two and Gu6rin and Spinola each contrib- uted two more. The genus began to suffer from a sort of ab- normal growth due to the writings of I?. Smith who added some forty new forms, most of them badly characterized. In one case Smith illustrated the description of a Pseudomyrma with the drawing of an Eciton. The resultant chaos discouraged many later writers, a fact which can be readily seen by a perusal of the small list of later contributors. Thus, Mayr described 7 forms, Roger 4, Emery 15, Fore1 78, Wheeler 6 and Santschi 16. In addition, Buckley, Norton, Wasmann and Aquayo each de- scribed one; Mann and Sitz described two each, and Wheeler in collaboration with Mann added four more. Through the kindness of Professor Banks, Curator of the Entomological Collections of the Museum of Comparative Zoology, Harvard University, we are in the fortunate position of having access to the large collection, which is mainly built around Wheeler's material; many of the types and identified specimens are available here.
The Haskins Collection contains an additional number of Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College.
Wailing address: The Biological Laboratories, Harvard University.



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60 Psyche [Sept.-Dec.
types and identified specimens. The two collections between them contain specimens of nearly half of all the described spe- cies and varieties.
Following Forel's example we have disregarded a number of inadequately described forms, mostly those of F. Smith. Forel's viewpoint on Smith's work is given in the following quotation: "F. Smith hat eine grosse Anzahl Arten dieser Gattung (Pseudomyrma) moeglichst oberflaechlich, fast ausschliesslich nach der bei den verschiedenen Individuen ungemein variie- renden Farbe beschrieben. Es folgt daraus, dass die Mehrzahl seiner Arten unentwirrbar ist. Was bis jetzt entraetselt werden konnte ist von Mayr, Emery und mir klargestellt worden. Den Rest muss man ignorieren, oder man muss auf die Beschreibung und Neubenennung von Pseudomyrmaarten verzichten. Ich will nun das erstere tun."
Emery, 1895-1 897, Zool. Jb. Syst., expresses similar views. Fore1 however can not be absolved from adding to the diffi- culties of the systematics of Pseudomyrma. He described a very large number of new forms, mostly subspecies and varie- ties, based on minute differences, but failed to arrange his and the previously described forms and integrate them into a sys- tem. He was able to do this only because of his remarkable knowledge, which prevented him from making mistakes and duplications.
Emery in his catalogue (Gen. Ins. 192 1-23) made some at- tempt to arrange the known forms of Pseudomyrma which at his time of writing numbered nearly two hundred. He split the genus into three groups: gradis, tennis and divers, only one of which - gracilis - was characterized by him. In the present account we have divided Pseudomyrma into five branches and each of the branches into numerous groups wherever possible. The terms "branch" and "group" were chosen deliberately and are employed in the same sense in which Emery used the term group, to denote a classificatory category which can not be sharply characterized, i.e., where there are too many intergrades to permit the use of such terms as genus or subgenus.
This paper contains the descriptions of a number of new Pseudomyrma which the late W. M. Wheeler had named and labelled as types and cotypes, but whose descriptions have not been published. Professor Brues and Professor Banks have



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19441 Genus Pseudomyrma 61
advised us to include these descriptions in this paper. Other undescribed forms were found in the unidentified ma- terial of the collection of the Museum of Comparative Zoology; the remainder of the newly described forms are material col- lected by us and for us in Central and South America. Genus Pseudomyrma (Latreille) .
Pseudomyrma is the only American genus of Forel's Pro- myrmicinse (Promyrmicini of some writers), which includes the following tribes and genera :
I. Tribe Metaponini Fore1
1. Genus Metapone Fore1
11. Tribe Pseudomyrmini Fore1
1. Genus Sima Roger
2. Genus Pseudomyrma Latreille
Branch Ornatinoda. PI. 11, fig. 1, 2; PI. 111, fig. 1, 2,3. Characters : The petiole with angular or rounded projections above and behind; or deeply excavated above; or with longi- tudinal or transverse impressions. This branch corresponds roughly to Emery's group tenuis.
Type : Pseudomyrma tenuis (Fabricius) .
Groups : excavata, tenuis, picta-schuppi. Branch Clavanoda. PI. 11, fig. 3,4, 5 ; PI. 111, fig. 15 ; PI. IV, fig. 26-35.
Characters: The petiole is clubshaped, with a long thin pe- duncle in front, and is not or only weakly marginated at the sides. This branch corresponds to Emery's group gracilis. Type: Pseudomyrma graciUs (Fabricius).
Groups: Zcevigata, sol& mutica, gracilis. Branch Triangulinoda. PI. 11, fig. 6-1 2. Characters: The pedicel from above has approximately the shape of a triangle, usually with rounded corners and convex sides; the peduncle is much shorter than in the preceding branch. The node is margined or submargined in some of the forms.
Type: Pseudomyrma spinicola Emery.
Groups: dolichopsis, sabanica, championi, spinicola, elegans. 'An explanation of this term has been given on p. 60.



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62 Psyche [Sept.-Dec.
Branch Apedunculata. PI. 11, fig. 13-15. Characters: The petiole rises from the articulation with the epinotum in a straight line or in a convex line, i.e., the node extends clear to the articulation and the peduncle is not differ- entiated; the node is usually strongly margined and sometimes the lower face of the petiole in profile is concave. Type : Pseudomyrma sericea Mayr .
Groups: subtillissima, sericea, acanthobia, elongata, fili- formis, fiebrigi-brunnea.
Branch Latinoda. PI. 11, fig. 17-22.
Characters: The peduncle is very short and notably broad- ened; the node is short and broad, sometimes broader than long; it is usually not margined at the sides. Type : Pseudomyrma latinoda Mayr.
Groups: triplaridis-maligna, arboris-sanctce, damnosa-lati- noda, canescens.
Branches of Pseudomyrma
1. The pedicel is ornamented with spines or cones or carinae on the posterior face, or with longitudinal impression or suture ........ Ornathoda, PI. 11, fig. 1, 2, 3; PI. 111, fig. 1 ............................................ The petiole is not ornamented
2
2. Petiole usually very long, club shaped, not marginate or .........................
submarginate Clavanoda, PI. 11, fig. 4, 5 ........................................ Petiole not distinctly club shaped
3
3. Petiole from above triangular in outline; marginate or not marginate; usually with a peduncle. The species which have the petiole triangular but the peduncle very short and broad (canescens, Pl. 11, fig. 18), have been included ...................
in Latinoda.. .Triangulinoda, PI. 11, fig. 6-13 ......................................
Petiole from above not triangular
4
4. Petiole from above convex at the sides, usually broadest in the posterior third; sometimes subtriangular, but always lacking a differentiated peduncle, i.e., the anterior face *This key can not be used generally for the identification of the females and males, although it is often possible to place the former by means of it. No attempt was made at completeness, because the main purpose of the key is to indicate clearly the position of the newly described forms.



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19441 Genus Pseudomyma 63
............ of the petiole in profile is straight or even convex ....................................... .Apedunculata, PI. 11, fig. 13-15 Pedicel of different shape; with the node transversely oval, or trapezoidal, or globular, or triangular, but always ............................ with a very broad and short peduncle.. .............................................. .Lathda, PI. 11, fig. 17-22 Detailed keys to the groups and species
1. Pedicel in profile long and low, with the upper border al- most horizontal. PI. 111, fig. 3.. ......... .tenuissima Emery Petiole in profile higher.. ................................................... .2 2. Pedicel in profile with a high rounded node. PI. 111, fig. 1 .group excavata
................................................................... Petiole in profile with a sharp or blunt superior-posterior corner. PI. 111, fig. 2.
....
A. - Posterior face of the node straight. PI. 11, fig. 2 ................................................... .group tenuis ...........................
B. - Posterior face of the node convex.
............................................. group picta-schuppi The group picta-schuppi is transitional to the branch Latinoda and some of its forms are listed there. We include under Ornatinoda the following additional forms: 1. gebelli-which has a transverse suture on the petiolar node.
2. excissa - the node with a strong semilunar excava- tion.
3. schuppi var. geraensis - which has the node exca- vated on top.
The remaining forms of schuppi we have placed near belti, following Forel.
Group tenuis
1. tenuis, typical. Length 5 mm. Entirely yellow-ochrous ; sides of the head very convex; basal face of the epinotum with a longitudinal impression; node from above long and narrow ; prothorax strongly marginate. Maranho, Brazil. 2. var. pa~~nsis. Length 5 mm. Yellowish; mesonotum broader than long (as long as broad in the type) ; node with rounded angles instead of sharp corners; posterior face of node a little convex. Para, Brazil.




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64 Psyche [Sept.-Dec.
3. var. pittieri. Length 6 mm. Reddish brown; abdomen brownish. Costa Rica.
4. subsp. nigripes. Color ferruginous red; head black. Brazil, Peru.
5. var. andina var. nov. Length 5 mm. Reddish yellow; sides of the pronotum coarsely sculptured (finely punctate in the type). Peru.
6. var. guatemalensis var. nov. Length 6 mm. Reddish yel- low; head brown; prothorax and mesothoracic disc both broader than long; petiolar node without horns or cones, but sharply raised brown margins. Guatemala. 7. var. rufa (redescribed as P. ruf a by Mann, 19 16). Length 6 mm. Color ferruginous; meso- and epinotum infuscated. Para, Brazil.
............................................................ 1. Monocolored forms. .2
Bicolored forms.. ................................................................. .3 2. Yellow colored.. ............. group lcevigata-solisii (see below) Color brown to black.
A. unicolor (syn of the type and monochroa) ; black. B. alfari. Brownish yellow; length 4.1-4.8 mm.; pe- duncle short.
C. duckei. Length 3.5-4 mm.; peduncle short and in- distinct ; this form is transitional to Apedunculata. D. nigropilosa. Length 6 mm.; testaceous yellow with black hairs.
3. Larger forms ; 1 1 mm. or larger :
A. gracilis var. major. Yellowish red; head and abdo- men black.
B. mutilloides. Black, except the peduncle which is red. ..................................
Smaller forms ; usually under 9 mm..
.4
4. Thorax not impressed at the mesepinotal suture, which is very faint; length 5.6-6.3 mm.; head black, rest of the ........
body reddish brown. The color is however variable ................................................................................ salvini ....................................................... Thorax not impressed..
.5
5. Peduncle very short; length 6.5 mm. Pedicel and gaster ...........................
brown, rest of the body black..
.godmani
................................................................. Peduncle longer.. .6




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19441 Genus Pseudomyrma 65
Gaster light brown with dark brown spots, like freckles.. .... . .
................................................................ santschii, sp. nov. Gaster not spotted with brown ..................... gacilis group, (remaining forms).
Group gracilis
PI. IV, fig. 26-35
1. Petiolar node in profile with a very steep anterior face. PI. 11, fig. 32 ; length 9-10 mm. ; dark brown ........................ ................................................................ subsp. squamifera ........................................ Petiolar node of different shape
..2
2. Pronotum broader than long; length 8.5 mm.; color vari- ................................................................ able. subsp. agilis
Pronotum as long as broad or longer than broad. ............. ..3 3. Head and thorax smooth and shiny; length 6.4-6.8 mm.; epinotum with a semilunar impression; gaster without ............................................. pubescence.. .var. glabiventris
................................................... Not with these characters 4
4. Peduncle relatively short :
A. subsp. maculata. Peduncle shorter than in the type; pilosity more abundant. PI. 111, fig. 34. B. gracilis type. Color extremely variable; most of the named forms fall in fact under the definition of the type.
C. subsp. mexicana. More robust than the type; black with the prothorax red. PI. IV, fig. 29,33. D. subsp. carapuna. Smaller than agilis and more pilose than the type. PI. IV, fig. 27.
Peduncle relatively long, usually with a secondary dilata- tion at the level of the stigmata:
E. subsp. sericata. Thorax vividly red, head and gaster black. Pl. IV? fig. 2 6.
F. var. peruviana. Peduncle very long; mesepinotal disc very elongated. PI. IV, fig. 3 1.
G. var. longinoda var. nov. Peduncle very long as in the preceding form; shoulders very overhanging. (The species alternuns and faber are probably nothing more than color varieties of the species gracilis; we have not seen the types.)




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66 Psyche [Sept.-Dec.
Group Icemgata-soUsii
Zcevigata type. Length 8.5 mm.; with fine punctation, body very shiny; color testaceous yellow.
subsp. kitscheli. Length 9 mm. ; head less convex than in the type; epinotum not narrowed behind.
subsp. osura. Length 7.2 mm.; head longer than in the type, longer than broad; base of the epinotum more con- vex than in the type and in kitscheli; color reddish yellow. subsp. insularis subsp. nov. Smaller than the type and with abundant golden hairs (the pilosity in the type is sparser and black).
P. solisii. This species is intermediate between Clavanoda and Triangulinoda. We have put it into the latter, because of its short peduncle. It may be considered the extreme border case of this group (Icevigata).
flavidula. PI. 111, fig. 15. Entirely smooth and shiny; yel- low colored; head broader than in pallida; abdomen with a brown spot of variable size. Texas to Brazil. flavidula var. pallida (formerly pallida) ; very close to the preceding, but the abdomen is not spotted. Florida to Mexico.
flavidula var. jaumei. Length 4 mm.; close to the variety pazosi, but with the head shorter and less convex; gaster shiny and black; rest of the body dark red. Cuba. subsp. levivertex. Color reddish yellow; first petiolar node more sharply bordered than in the type; abdomen banded brown. Argentina.
subsp. lizeri. Length 6 mm.; abdomen spotted, thorax tinted brown; clypeus not toothed (toothed in levivertex) . BRANCH TRIANGULINODA
PI. 111, fig. 6-10
Pedicel in profile less than twice as long as high; peduncle barely indicated by a concave anterior border of the node. PI. Ill, fig. 4. ........................................................ 2 Pedicel in profile more than twice as long as high; peduncle well developed and long. PI. 111, fig. 7.. ...................... .4 Thorax without impressed mesepinotal suture; length 6.5 mm.; pronotum margined; epinotum and first node reddish, postpetiole and abdomen brown or black.



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Genus Pseudomym 67
Guatemala. (Differs from the related form antiguana by the shorter peduncle.) ................................................ stolli Mesepinotal suture impressed.. ........................................... .3 3. With a small antero-ventral tooth on the petiole. PI. 111, .................................................... fig. 8.. .group dolichopsis
A. Larger forms, 4.3-5 mm.:
I. dolichopsis. The node in the typical form is very broad and convex on the sides.
Colombia.
11. var. curacmis. Length 4.3-5 mm.; color testa- ceous; brown in the type. Curacao.
111. var. implicata. Length 4.3 mm.; head narrower than in the type; erect hairs absent on the femora. Amazonas.
B. Smaller forms: Length 3.6-3.9 mm. ; scapes longer than in dolichopsis; reddish yellow or brownish. Guatemala. ...................................................... peperi With larger antero ventral tooth and with a poster0 ventral tooth; the two teeth connected by a lamina. PI. 111, fig. 4. ............................................... .group championi-denticoZZis A. denticollis, type and varieties; length 5-7 mm.; uni- colored or not.
I. denticollis type. Length 6-7 mm.; reddish; Pa- raguay. PI. 111, fig. 5.
11. var. infuscata. Length 5-6.2 mm.; head brown, gaster testaceous, sometimes the whole body brown. S5o Paulo, Brazil.
B. goeldii. Length 3.2-3.5 mm.; pronoturn red, rest black. S5o Paulo.
C. rochai. Length 4-4.7 mm.; head bright red, rest of the body black.
D. championi and subspecies. PI. 111, fig. 4. Length 4.5-5.5 mm.; antero ventral tooth strong and curved backward.
I. championi type. Thorax and petiole red, rest blackish brown; length 4.7 mm.
11. var. haytiana. Same color as the type, but the antennae are bright yellow; postpetiole and gas- ter with reddish spots.
111. subsp. haytiana var. affinis. Head black, epi- notum with a black streak in the center; pedicel and gaster brown.




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Psyche [Sept .-Dec.
IV. subsp. haytiana var. torquata. Head to post- petiole black, except the pronotum, which is bright red.
V. subsp. haytiana var. paulina. Transitional to subsp. incurrens.
VI. subsp. incurrens. Length 4.6-5.3 mm.; brown as in torquata but the node is three times as long as broad.
4. Unicolored forms. PI. 11, fig. 6... ..... group sabanica-pallens A. depressa. Length 4.3 mm.; thorax strongly com- pressed; color brownish black; head as long as broad, with convex sides; smooth and shiny. Colombia. B. kuenckeli. PI. 11, fig. 10. The node from above forms a short triangle.
I. kuenckeli type. Color reddish testaceous, gaster brownish. Costa Rica.
11. var. bierigi. Length 5 mm.; darker than the type, nearly black; head f erruginous. Panama. 111. var. dichoa. Larger than the type and more robust; head yellowish red, thorax yellow in front, rest of the body brown. Colombia. IV. var. hondwiana. Head reddish brown, thorax dark reddish, gaster brown. Honduras.
(The varieties of P. kuenckeli are separated mainly by their color patterns; we have not enough material on hand to determine how real these differences are.) C. sabanica type. Color dark brown (PI. 11, fig. 6) and the variety saffordi length 3.4-4 mm.; color uni- formly brown, lighter than in the type.
D. pallens type. Length 5-6 mm.; color testaceous; pilosity sparse; pubescence moderate. Colombia. I. var. gibbinoda. Mesonotum very convex (in the type it is straight and forms an acute angle with the base) ; Siio Paulo.
11. var. landoldti. Length 6 mm.; reddish yellow; first node much broader than in the type. Co- lombia.
................................................................. Bicolored forms.. .5
5. Thorax without mesepinotal constriction. ............................. ................................................................. .antiguana sp. nov.



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Genus Pseudomyrma 69
I. antiguana type. Length 5 mm.; reddish brown, head and gaster slightly darker. Guatemala. 11. var. brunneipes var. nov. Length 6 mm.; sides of the thorax and petiole bright yellow contrasting sharply with the dark brown legs. Guatemala. Thorax with a pronounced mesepinotal constriction . . . . . . . . .6 6. Smaller forms. PI. 111, fig. 7. ........... ......... group spimcola A. edvardi type. Color black and the variety ccecilice. Black as the type; length 2.9-3 mm. Differs from the type by the straight posterior edge of the head (concave in the type).
B. ejecta type. Length 1 % lines. Color rufo-testaceous,