The Notropical Genus SYNTERMES

THE NEOTROPICAL GENUS SYNTERMES(ISOPTERA: TERMITIDAE)

THE NEOTROPICAL GENUS SYNTERMES(ISOPTERA: TERMITIDAE)

ALFRED E. EMERSON

Research Associate, Department of Insects and SpidersThe American Museum of Natural History

Professor of ZoologyThe University of Chicago

BULLETINOF THE

AMERICAN MUSEUM OF NATURAL HISTORY

VOLUME 83: ARTICLE 7 NEW YORK: 1945

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY

Volume 83, article 7, pages 427-472, text figures 1-12

Issued August 6, 1945

CONTENTS

INTRODUCTION .................. 433

Acknowledgments.. . ............... 433

General Ecology ...... . .. . . . ................... 433

SYSTEMATIC DESCRIPTION .................. 437

Genus Syntermes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 439

Key to the Species of Syntermes . . . . . . . . . . . . . . . . . . . . . . . 442

Syntermes aculeosus, new species . . . . . . . . . . . . . . . . . . . . . . 443

Syntermes obtusus Holmgren . . . . . . . . . . . . . . . . . . . . . . . . 444

Syntermes magnoculus Snyder . . . . . . . . . . . . . . . . . . . . . . . 445

Syntermes snyderi Emerson . . . . . . . . . . . . . . . . . . . . . . . . 446

Syntermes solidus, new species . . . . . . . . . . . . . . . . . . . . . . . 447

Syntermes chaquimayensis chaquimayensis (Holmgren) . . . . . . . . . . . . . 447

Syntermes chaquimayensis, subspecies emersoni Snyder . . . . . . . . . . . . . 449

Syntermes chaquimayensis, subspecies parvinasus, new subspecies . . . . . . . . . 450

Syntermes grandis (Rambur) . . . . . . . . . . . . . . . . . . . . . . . . 451

Syntermes lighti, new species . . . . . . . . . . . . . . . . . . . . . . . . 453

Syntermes hageni Holmgren . . . . . . . . . . . . . . . . . . . . . . . . 454

Syntermes wheeleri, new species . . . . . . . . . . . . . . . . . . . . . . . 455

Syntermes brevimalatus, new species . . . . . . . . . . . . . . . . . . . . . 457

Syntermes peruanus Holmgren . . . . . . . . . . . . . . . . . . . . . . . 458

Syntermes dirus (Burmeister) . . . . . . . . . . . . . . . . . . . . . . . . 459

Syntermes silvestrii Holmgren.. . . . . . . . . . . . . . . . . . . . . . . 462

Syntermes calvus, new species.. . . . . . . . . . . . . . . . . . . . . . . 463

Syntermes territus Emerson.. . . . . . . . . . . . . . . . . . . . . . . 464

Syntermes parallelus Silvestri.. . . . . . . . . . . . . . . . . . . . . . . 466

Syntermes molestus (Burmeister).. . . . . . . . . . . . . . . . . . . . . 467

Other Records of Syntermes.. . . . . . . . . . . . . . . . . . . . . . . 470

BIBLIOGRAPHY ............................... . 470

431

INTRODUCTION

THE GENUS Syntermes includes a number oflarge species of termites confined to theSouth American continent and generally rec-ognized as the most primitive group of thesubfamily Nasutitermitinae of the familyTermitidae. In spite of the large size, whichhas attracted the general collector, the sub-terranean habitat and the tendency of theworkers and soldiers to retire under groundwhen disturbed have resulted in meager col-lections. This study attempts to bring thetaxonomy of the genus up to date and tostimulate further collecting and ecological in-vestigations.

ACKNOWLEDGMENTSI am indebted to Messrs. P. J. Anduze,

R. L. Araujo, N. Banks, J. C. Bradley, E. A.Chapin, P. J. Darlington, Jr., W. J. Gerhard,N. Holmgren, H. Kahl, F. E. Lutz, J. V.Pinheiro, A. Reyne, A. Roman, C. H. Seevers,F. Silvestri, T. E. Snyder, G. Vivas-Berthier,and N. A. Weber for specimens and for theopportunity to examine collections. MissLaura B. Churchill prepared the drawingof the map, and I was assisted in some of thedrawings of the species by Mr. Rupert L.Wenzel. During the course of the study Ivisited the Reale Scuola Agraria, Portici,Italy; Staatsammlung, Munich, Germany;Museum fur Naturkunde, Berlin, Germany;Naturhistoriska Riksmuseets, Stockholm,Sweden; United States National Museum,Washington, D. C.; the American Museum ofNatural History, New York; Museum ofComparative Zoology, Cambridge, Massa-chusetts; Carnegie Museum, Pittsburgh,Pennsylvania; and the Chicago Natural His-tory Museum, Chicago, Illinois. Portions ofthe study were made during my tenure of aJohn Simon Guggenheim Memorial Founda-tion Fellowship in 1926-1927. Recently Ihave received financial aid from the Dr. Wal-lace C. and Clara A. Abbott Memorial Fundof the University of Chicago.

GENERAL ECOLOGYSyntermes is confined to the tropical savan-

nas and rain forests of South America (fig.1). Although this genus or its immediate

generic ancestor must have been in existencebefore the Cretaceous dispersal of the de-rived genera Nasutitermes and Subulitermes,it seems likely that biotic barriers preventedextension of its range. It is also noteworthythat the Pliocene invasion of Cornitermes andArmitermes into Central America was not ac-companied by Syntermes.The species of the genus usually have their

nests entirely under the ground, but some-times they make mounds above the ground.The mounds of S. snyderi found at Kartabo,British Guiana, were composed of excavateddirt brought to the surface by the workersand, aside from the openings, did not containgalleries (Emerson, 1938, p. 258). In one casethe surface mound of loose dirt was 2 feethigh, 3' feet long, and 2 feet wide. One largeopening was located near the top of themound. The chambers extended below thesurface of the ground about 4 feet andcovered an area with a diameter of about 12feet. The galleries were large and seemed tobe simple excavations in the sandy clay soil,although there were small lumps lining thegalleries in places which might have been ex-cretions of the termites. In another case thenest was underneath a large Mora tree. Thetermites had deposited the dirt between thebuttressed roots of the tree so that the moundsloped down from the base of the roots andstood 3 feet from the ground level in places.Holes were found at the top of each slopenext to the trunk of the tree.No nests of S. territus were found at Kar-

tabo. Digging 1I feet below entrance holeson the ground surface failed to disclose nest-ing galleries. Soldiers and workers were col-lected from an open trail near small moundswith openings at the top which were prob-ably connected with the underground nest. Atanother time I saw S. territus soldiers andworkers emerge from a hole under leaf debrisnear the roots of a large dead stump. On oneoccasion S. territus workers and soldiers werefound in a short open trail under leaves on theforest floor at Kartabo, and a queen of fairsize was found under a leaf with a few smallnymphs, some carried by workers. When dis-turbed, the workers and a soldier tapped their

433

BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

GS SERIES OF BASE MAPSNwLa nu"

FIG. 1. Distribution of the species of the genus Syntermes showing the approximate border of thetropical savanna and rain forest combined. S. wheeleri from Sao Paulo (23°44'S., 46°38' W.), Sao Paulo,Brazil, was recorded after the map was prepared.

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EMERSON: THE NEOTROPICAL GENUS SYNTERMES

heads rapidly on the dried leaves, making adistinct noise. The soldier tapping was some-what slower than that of the workers. Gradu-ally all disappeared inside a decayed log ofabout 4 or 5 inches diameter which was thenbrought back to the laboratory. The king waslater found inside this log. Near the regionwhere I found these termites I had seen anumber of workers earlier coming from ahole in the ground. No mound was seen, andit is difficult to understand what the repro-ductives were doing above the ground. Directsunlight used when we took moving picturesof the termites killed all of the castes veryquickly. Ants of three species were foundpreying upon the workers in the open trailand were later determined by Dr. W. M.Wheeler as Neoponera apicalis Latreille,Ectatomma quadridens Fabricius, and Phei-dole biconstricta Mayr.

Dr. N. A. Weber's field notes concerningthe collection of S. aculeosus and S. brevi-malatus from 02°42' N., on the OronoqueRiver in British Guiana, state: "Nesting inclay at base of tree. Openings one centimeterin diameter leading directly down for a fewcentimeters to horizontal galleries in top tento thirty centimeters of soil." These specieswere found together in the same vial, but thedescription of the nest probably refers to thatof S. brevimalatus because more specimens ofthis species were in the vial.

Dr. Charles Seevers sent me the followingfield notes: "A large colony of Syntermesmolestus was located beneath a log in an openfield near Villavicencio, Colombia. Thephysogastric queen and the king occupied acell on the surface of the ground. When thelog was removed, the soldiers very quicklyaligned themselves in defensive positionswith their heads directed outwards. Theworkers began retiring into the subterraneangalleries, taking with them the immatureforms. The king and queen also withdrewvery slowly into these galleries but were cap-tured before retirement had been completed.The soldiers, which had been drawn up inclose rank around and throughout the colony,completed the withdrawal of the colony aftera time."

Dr. J. C. Bradley of Cornell University re-ports a large mound of S. chaquimayensisfrom the Putamayo region of Peru with a

colony of stingless bees of the genus Trigonaestablished in the mound.

All the species of Syntermes, as far as isknown, collect portions of leaves which theycut out of the dead leaves or pieces of grassand store them in their nests. In the case ofS. snyderi, I found circular leaf pieces about1 centimeter in diameter stored in chambersin their nest. I once saw a worker of S. ter-ritus emerge from a hole in the ground, pickup a fragment of a dead leaf about one-halfinch long and return to the hole with the leafpiece held vertically in its mandibles. Be-quaert (1925, p. 294) records the leaf-collect-ing activities of S. grandis and S. molestus(S. brasiliensis synonymous). Silvestri (1903)states that it is his belief that the leaf frag-ments are used for growing fungus uponwhich the termites feed. He observed fungusgrowing on pieces of grass stored by S. moles-tus. During my observations at Kartabo,however, I never saw any sign of fungus-growing behavior by these termites eventhough I dug some distance into their nests.Fungus gardens of the type found among allspecies of the Macrotermitinae have neverbeen observed in Syntermes nests nor associ-ated with any other South American termite.I am inclined to believe that the fungus grow-ing on the stored grass noted by Silvestri wasaccidental. The hypothesis that Syntermes isnot a wood eater but is largely confined tograss and leaves for food conforms to knownobservations. Thi's enables the genus to livein both rain forests and treeless savannas(fig. 1).The workers show a distinct dimorphism in

every species of the genus I have examined.One type of worker has a large yellowishhead, and the other has a distinctly smaller,lighter head. Silvestri (1903) states that thesmall form stays inside the nest and is adaptedfor functions in the nests. Both forms ofworkers, however, were found in the opentrails of S. territus in the day time at Kar-tabo. The smaller worker may possibly be anearlier instar of the worker caste.The soldiers of Syntermes have very strong

mandibles with which they are able to drawblood by cutting through'the skin of the hand.In addition to this means of defense, thefrontal gland of the soldiers is probably ableto emit a defensive fluid and is the most

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primitive in the phylogenetic series whichends in the nasute soldiers with degeneratemandibles. The soldier of S. snyderi had adistinctive odor when placed in a closed vialfor a time. I was unable to witness much de-fensive action during normal predation, how-ever. Ponerine ants were able to sting thesoldiers so effectively that they could offer noresistance. I also failed to observe any de-fensive effect of the thoracic spines of thesoldiers and workers.

Wheeler (1936) has published a rather ex-tended account of the taxonomy and ecologyof the ant Termitopone (Syntermitopone) com-mutata (Roger), including my field observa-tions taken in the rain forest of British Gui-ana. This interesting ponerine ant seems tobe a termite raider specializing on the speciesof Syntermes. I have witnessed these ants at-tacking colonies or transporting workers andsoldiers of S. territus, S. snyderi, and S.calvus at Kartabo, British Guiana. Dr.W. T. M. Forbes collected this ant carrying aworker of S. chaquimayensis at El Cam-pamiento, Perene, Peru. Bequaert (1925,1926) observed raids on S. grandis and S.molestus (S. brasiliensis is considered a syno-nym) on the savanna of Vista Alegre, RioBranco, Brazil. Dr. A. Roman observed T.commutata transporting workers and soldiersof S. territus at Cururuzinho, Rio Autaz,Amazonas, Brazil, on October 22, 1914.Wheeler (1936) lists records of 25 colony raidsinvolving many hundreds of individuals at-tacking various species of Syntermes. Thegeographical distribution of T. commutata co-incides quite well with the known distributionof Syntermes. It seems reasonable to supposethat further observations will verify thepredatory relationship of this species of antand this genus of termites.

While excavating a nest of Syntermessnyderi at Kartabo, I picked up specimens ofthe ant Carebara uwnifredae Wheeler, whichWheeler later (1922, p. 4) named after mywife, Winifred Jelliffe Emerson. Wheeler

(1936, pp. 176-177, 197-202) discussed thedistribution and relationships of the speciesof Carebara which are found in the Indo-Malayan, Ethiopian, and Neotropical regionsin the nests of Acanthotermes, Macrotermes,Odontotermes, and Syntermes. His record of S.dirus in British Guiana (p. 176) is in realityS. snyderi. Wheeler classifies the relations ofCarebara to their host termites under the term"termitolesty." The relationships of Syn-termes to these genera of the Macrotermitinaeare discussed later. Here we find a correlationbetween taxonomic, ecological, and zoogeo-graphical data, although more informationconcerning the ecological relationship be-tween Carebara and its hosts needs to begathered before we become convinced of areciprocal evolution.

Seevers (1941) described a genus and spe-cies of phorid flies, Syntermophora micro-phthalma, from the nests of Syntermes mo-lestus-at Villavicencio, Colombia. He also de-scribed a new species, Cryptophora colombiae,from nests of S. molestus. The genotype spe-cies of Cryptophora, C. coeca Borgmeier, wascollected from a marching colony of Syn-termes sp. at Santa Cruz, Goyaz, Brazil.These two genera of flies have been foundonly with Syntermes, but the data are toomeager for a reciprocal evolution to be as-sumed.

Borgmeier (1930) has described a histeridbeetle, Scapolister sternalis, from the nest ofSyntermes molestus (brasiliensis synonymous)from Campinas, Goyaz, Brazil. Reichen-sperger (1936) described another histerid,Cossyphodister schwarzmaieri, from the nestof Syntermes sp. from Santa Cruz, Goyaz,Brazil.Wasmann (1894) described the acantho-

cerid beetle A canthocerus termiticola from thenest of Syntermws dirus from Rio de Janeiro,Brazil. This genus of beetles has been foundin other termite nests (Constrictotermes cavi-frons at Kartabo, British Guiana), and alsofrom habitats dissociated from termites.

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SYSTEMATIC DESCRIPTION

EARLIER AUTHORS, including Hagen (1858a,pp. 107, 108; 1858b, p. 16; 1862, p. 59),Silvestri (1903, p. 47), and Holmgren (1906,p. 544), included species of the genus under thegeneric name Termes. In the post-Linnaeancentury, Termes was virtually synonymouswith our present concept of the order Isop-tera. Starting with Hagen (1853, 1858a) theold genus Termes was subdivided, and gradu-ally the modern generic groups were recog-nized and named mainly in the works ofWasmann, Silvestri, and Holmgren. Thegeneric concept of the genus Syntermes hasnot been changed since 1910 when it wasnamed by Holmgren (1910, p. 285) with S.dirus as the generitype species. Further de-scriptions and the addition of new speciesand castes have only substantiated and clari-fied the concept. The genus would seem to bea natural group of related species, and no ne-cessity has arisen for subdividing it into twoor more genera or even into subgenera.

Because the nomenclatural system in tax-onomy seems to give an impression of finalityto many non-taxonomists, it seems wise topoint out from time to time that, as in othersciences, the nomenclature is no better thanthe correlations of the gathered data it sym-bolizes and that it changes as new knowledgeis gained, particularly if the tentative con-cepts are based upon meager information.The taxonomy of the genus Syntermes has byno means been completed. It was in a com-paratively unsatisfactory condition prior tothe new studies contained in this paper, andno more can be said than that a little moreprogress has been made toward the definitionof the various species, their geographic distri-bution, and phases of their ecological relation-ships. I maintain, however, that this littleprogress has scientific value and is necessaryfor the more accurate elucidation of certainbroad generalizations of interest to biologistswho may never be called upon to determinea species of the genus or who may never seea living specimen in its natural habitat.There has been some question in the minds

of various biologists concerning the reality ofthe higher taxonomic categories. Kinsey(1936, p. 5) states: "All of the higher cate-

gories are artificial conventions useful forcataloging biologic data, but hardly realeither in manner of origin or in their intrinsicqualities. The evolutionary pattern is notthat of a 'tree of life' in which the mainbranches represent ancestral stocks whichdisappeared as they gave rise by radiateevolution to the species at the ends of thetree.... Higher categories are not necessar-ily groups of similar units (e.g. genera are notgroups of similar species), and higher cate-gories are not groups of units with a commonorigin."

In the following diagnosis of the genusSyntermes, I have conformed to the conven-tional taxonomic tradition of describing thecharacters which are common to the variousspecies which I and others have includedwithin this genus. These characters are con-sidered homologous, by which we mean thatthey not only have similar visible patternsbut also exhibit in common subtle and com-plex growth activities during their ontoge-netic development-developmental processeswhich were initiated and influenced by a set-of genetic factors or genes highly similar inthe various species due to their evolutionfrom a common ancestry. This brief sum-mary of the basic assumptions of taxonomichomology rests upon extensive researchesin the fields of genetics, embryology, andphysiology which are applicable to this par-ticular case of the genus Syntermes, notbecause of special knowledge of the geneticsand physiological development of these ter-mites, but because of the correlation of thedata with similar information on organismsmore thoroughly studied by geneticists andembryologists. If these basic assumptions arevalid, one would expect other types of evi-dence to point toward the same general con-clusions concerning homology. Geographicaldistribution (fig. 1), ecology, and behaviorshould be related to phylogeny. The ecologi-cal relationships of species of Syntermes to theant Termitopone (Syntermitopone) commutata(Roger), described in the preceding pages,may be cited as an example. Such observa-tions tend to substantiate the objectivity oftaxonomic categories. If the genus Syntermes

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is an "artificial convention useful for cata-loguing biologic data, but hardly real eitherin manner of origin or in intrinsic qualities,"Termitopone commutata must have extra-sensory perception in reading the minds oftermite taxonomists.Whether the species of this group might be

classified further into subgenera or genera, orwhether they might be incorporated into alarger genus including the species now as-signed to Cornitermes depends upon com-parative patterns of homologies and the gapsdue to extinction. To a certain extent thesedivisions are arbitrary, and Kinsey is right inemphasizing a degree of subjectivity, but thedata still point to the objective reality ofgenetic and phylogenetic relationships, thesymbol of which is the genus or other highercategory. It should also be mentioned thatthe evolution of generic groups and highertaxonomic categories gives us an understand-ing of the long-term action of evolutionaryfactors which are valuable in the analysis ofthe dynamics of evolutiQn, even though spe-ciation still remains the key study of phy-logeny.The arrangement of the species is based

upon morphological similarities. Theoreti-cally one should be able to construct a phylo-genetic tree of the species within the genus,but in this case the data are not adequate,and such evolutionary associations are merelysuggested in the linear order which, however,does not symbolize the multidimensional rela-tionships.The specific distinctions of the imagoes are

more difficult to discover than those of thesoldiers, a fact in harmony with the general-ization that the imago is a more conservativecaste than the soldier. Conversely, the imagomay be very useful for the diagnosis of highertaxonomic categories. The result, however, isthat the key to the species of imagoes ismuch less satisfactory for determinationsthan the key to the soldiers, and it is expectedthat the discovery of the imagoes of speciesnow described only from soldiers will 'overlapwith some of the specific characters used atpresent and that larger series will probablyshow overlaps that are not now included. Thekey is based upon observed series and is use-ful only as a preliminary guide for specificdetermination. I have in my collection two

vials of imagoes with characters that do notfit the key descriptions. They may be newspecies, imagoes of species known only fromsoldiers, or even varieties of described spe-cies. It seems best to await further specimensbefore attempting to form a taxonomic judg-ment. Many of the species based upon thesoldier caste seem satisfactory, althoughgreater series and new species will doubtlessmake it necessary to hunt for new charactersand better quantitative data. In some casesthe taxonomy must remain quite tentativeuntil more information is gathered.

Size may be fairly characteristic of a spe-cies or subspecies, particularly in the Termi-tidae, and it is deemed important to giveexact measurements of the specimens studied.When sufficient numbers are available, itwould be wise to express the variation interms of standard deviation from the mean.However, the small numbers of Syntermesavailable for study make this refinement oflittle value, and I have given only the rangeand the mean. It is to be expected that fur-ther data will extend the ranges so far notedand that different means may be found. Light(1927) has expressed proportions by means ofvarious indices. In some cases such data mayprove quite valuable, but I have not foundthem of sufficient importance in this genus towarrant the extra time involved in measure-ment. Larger collections may ultimately bestudied from this point of view with profit.The male imagoes are somewhat smaller thanthe females, but I have grouped the measure-ments in most of the cases. All measurementsare in millimeters and are taken by means ofan ocular micrometer. Inasmuch as differentstudents have different systems of measure-ments, I have explained below the pointswhich need clarification.

TERMS1. The length of the head with mandibles

includes the extended mandibles.2. The length of head to side base of man-

dibles is the greatest length of the head cap-sule at the side.

3. The width of the head of the imago in-cludes the eyes.

4. The diameter of the eye is the longestdiameter and includes the diameter of theocular suture.

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5. The length of the mandible includes therounded side condyle projection. This canusually be measured without removing themandible.

6. The length of the postmentum is themedian length of the sclerotized portion. Inearlier works this sclerite has often been re-ferred to as the "gula."

7. The length of the pronotum is thelength of the middle line and, therefore, isnot the maximum length if there are emargin-ated front or hind borders.

8. The width of the thoracic nota includesthe spines when present.

9. The length of the wing from the sutureand the length of the scale are taken from thecostal border of the suture. If the suture isnot mentioned, the wing length includes thescale.

Unless otherwise stated, the determina-tions of the listed specimens were made bythe author and conform to the nomenclatureused in this paper. The specimens are de-posited in the collection of the AmericanMuseum of Natural History unless anotherrepository is given. Other numbers listed,if a repository is not indicated, are the fieldnumbers of the collectors. The author ex-amined all the listed specimens, and the de-scriptions are based upon this material.

GENUS SYNTERMES<Genus Termes HAGEN, 1858a, pp. 107, 108.<Genus Termes HAGEN, 1858b, p. 16.<Genus Termes HAGEN, 1862, p. 59.<Genus Termes SILVESTRI, 1903, p. 47.<Genus Termes HOLMGREN, 1906, p. 544.-Genus Syntermes HOLMGREN, 1910, p. 285.-Genus Syntermes HOLMGREN, 1911, p. 545.= Genus Syntermes HOLMGREN, 1912, pp. 11, 45,

46.-Genus Syntermes SNYDER, 1924, p. 21.-Genus Syntermes EMERSON, 1925, p. 357.-Genus Syntermes EMERSON, 1928, p. 406.=Genus Syntermes WHEELER, 1936, p. 176.-Genus Syntermes HARE, 1937, pp. 462, 474,

475.

IMAGO (FIGS. 2, 3): Color yellowish brownto dark brown, wing membranes hyaline orbrown. Head, pronotum, and wing scaleswith fairly long, scattered bristles, wing mem-brane and veins with short hairs and nochitinous punctations. Size large (smallestspecimens measure about 27 mm. long with

FIG. 2. Head and pronotum of imago ofSyntermes dirus (Burmeister), cotype.

the wings). Head oval or egg-shaped, a fairlysharp ridge running diagonally from the frontof the eye to the lower base of the mandibles.Eyes relatively small or medium-sized, prom-

FIG. 3. Mandibles of imago of Syntermesdirus (Burmeister), cotype.

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inent. Ocelli medium size (smallest 0.27 mm.long), at least one and one-third times theirlength from the eyes. FQntanelle large (small-est 0.18 mm. long), conspicuous, roundish,lighter or about the same color as the head,depressed or forming a convex plate, with asmall light spot immediately anterior to thefontanelle. Arms of the Y-suture visible infavorable light, and sometimes the mediansuture is also visible. Antennae with 19 to 21articles. Postclypeus flatly arched with amedian longitudinal line, less than half aslong as wide. Anteclypeus white, with a V-shaped, darker, sclerotized mark in the mid-dle, the arms curving toward the front lateralmargin. Labrum hatchet-shaped (fig. 2) witha thin white anterior lip. The left mandible(fig. 3) with the apical and first marginaltooth roughly similar in size and a long, con-tinuous, even, slightly curved or nearlystraight cutting edge between the first mar-ginal tooth and the small second marginaltooth, a blunt notch posterior to the secondmarginal tooth. (Note: The above homologiesof the teeth seem to be a fair assumption froma comparison with the mandibles of theMacrotermitinae which have an acute notchbetween the first and second marginal teeth.)The right mandible (fig. 3) with an apicaltooth slightly larger or nearly the same sizeas the first marginal tooth, and the secondmarginal tooth smaller than the first marginaltooth, all notches sharper than a right angle.Pronotum somewhat saddle-shaped, propor-tionately wide (narrowest about seven-eighths width of head), either somewhat nar-rower or wider than the head, a frontal lobewith an anterior vertical surface and flatlateral projections which may be sharplypointed or somewhat rounded. Hind marginconspicuously emarginate. Hind margin ofmesonotum widely concave with fairly sharplateral angles (occasionally rounded). Hindmargin of metanotum widely concave withfairly sharp or rounded lateral angles. Fore-legs with three tibial spines near the apex, andmiddle and hind legs with two tibial spinesnear the apex (3:2:2). Tarsi with four arti-cles. Arolium absent. Scales of wings not over-lapping the posterior border of the notum.Shape of wings elongate with slightly curvedcostal and inner margins; with some reticula-tions between the costal border and the

radius near the tip. Wing suture nearlystraight with small undulations. Fore andhind wings with a short R1 and sometimesR2+3 joining the costal border a little beyondthe wing suture. R4+5 running singly to thetip of the wing where it may be branched. Mweak, running roughly medianly between Rand Cu, joining R4+5 beyond the suture in thehind wing. Cu weak with 14 to 17 branches.Cerci short with two articles. Male and fe-male with styli absent.SOLDIER (FIGS. 4-12): Color yellow brown,

including base of mandibles. Outer portionsof mandibles black. Head with many or fewhairs or bristles. Head large (width 2.4-7.1mm.), sides straight or a little curved, paral-lel or converging toward the front, a longi-tudinal lateral ridge on each side on the undersurface. Frontal tube short, no longer thanthe diameter of the antennal socket andsometimes hardly projecting above the sur-face of the head. Two small visible structuresor spots making a triangular arrangementwith the opening of the frontal gland. Post-mentum somewhat constricted in the middle,width of narrowest part over half the widthof the widest part. Antennae with 19 to 21articles. Penultimate article distinctly morethan twice as long as wide (5:2). Postclypeusfused with the front, the suture still visiblein a favorable light. Anteclypeus white, lessthan half as long as wide, with two sclero-tized, yellowish, diagonal, curved stripes con-verging at the posterior margin. Labrumthree-pointed, with white median tip, andlateral points which are either slightly morethan a right angle or sharper (to 600 angle).Mandibles large and powerful, curved orsomewhat hooked near the tips, basal lightportion quite sharply separated from outerblack portion and showing a small but dis-tinct notch at the junction on the outermargin. Left mandible with a marginal toothfrom a little less than half to more than halfthe length of the mandible from the base.Right mandible with a marginal and usuallyconspicuous tooth which may be almost ves-tigial (S. molestus and parallelus). Thoracicnota with sharp lateral spines, except in S.molestus in which the nota have somewhatangular sides. Pronotum with median emar-gination or entire front margin, anterior lobelarge and sharply marked from the rest of the

VOL. 83AA0


pronotum by angular notches on the marginsand in profile rising at a fairly sharp anglefrom the middle portion. Forelegs with threetibial spines near the apex, and middle andhind legs with two tibial spines near the apex(3:2:2).WORKER: Head yellow brown or light yel-

lowish white, covered with a few or numerousbristles. Fontanelle fairly large, round andwhite, as large as, or a little smaller than, theantennal sockets. Y-suture present. Antennaewith 19 to 20 articles. Anteclypeus as in thesoldier. Postclypeus a little shorter than halfits width, with a median line. Labrumhatchet-shaped with a thin white tip. Man-dibles similar to those of the imago, uniformin general tooth pattern in all species (seefig. 3). Thoracic nota with sharp lateral spinesor without such spines in S. molestus.

RELATIONSHIPS: Syntermes is the mostprimitive genus of the Nasutitermitinae(Termitidae) and is closest to the genusCornitermes. The relative primitiveness is in-dicated by the large size of the imago andsoldier, the relatively small frontal tube of thesoldier, the 19 to 21 articles in the antennaeof the imago and soldier, the proportionatelylarge mandibles of the soldier, the lateralspines or pointed sides of the thoracic nota ofthe soldier, the three apical spurs on the tibiaeof the forelegs, and the short R1 joining thecostal border a little beyond the wing suture.The closest relative outside of the subfam-

ily Nasutitermitinae would seem to be thegenus Acanthotermes of the Macrotermitinae.The similarities of Syntermes and Acantho-termes include what seem to be homologouslateral spines on the soldier mesothorax andmetathorax, vestiges of lateral spines on thesides of the prothorax of Acanthotermes (thesespines are very small on the worker of A can-thotermes but are present on most of the spe-cies of Syntermes), the white tip on the la-brum of the soldier in both genera (the tipof the imago labrum has a white lip in Syn-termes and Pseudacanthotermes), the post-mentum of the soldier in both genera is notparticularly wide in the middle in contrastto the wide postmentum in the higher Macro-termitinae, and both Syntermes and Acan-thotermes possess three spines near the tip ofthe tibiae of the foreleg and two spines onthe tibiae of the other legs (3:2:2).

The more important differences betweenAcanthotermes and Syntermes may be sum-marized as follows:

1. Syntermes has no frontal spines on thepronotum in either the imago or the soldier.

2. The major soldier of Acanthotermes hasno lateral projections of the labrum, althoughthe minor soldier has fleshly lateral projec-tions not sclerotized as in Syntermes.

3. Acanthotermes has a frontal spine nearthe opening of the frontal gland and does notpossess the frontal tube which is slightly de-veloped in Syn,termes.

4. Acanthotermes possesses trimorphic sol-diers compared to the monomorphic sol-diers of Syntermes. (Note: At least one andprobably two of the five types of Acantho-termes soldiers described and figured bySj6stedt [1926] are modifications caused byparasitic fly larvae in the heads.)

5. The left mandible of Acanthotermesworkers has what appears to be a vestigialsecond marginal tooth homologous with thesecond marginal tooth of the imago-workermandible of the Rhinotermitidae. (Thiswould indicate that the conspicuous secondmarginal tooth of the left mandible of theworker of Acanthotermes is homologous withthe third marginal tooth in the Rhinotermi-tidae; the left worker mandible of Acantho-termes is definitely more primitive than thatof Syntermes.)

6. The reduced teeth on the mandible ofthe soldier of Acanthotermes must be consid-ered more specialized than the toothed man-dibles of Syntermes.

7. The styli on the male imago of Pseuda-canthotermes must be considered a moreprimitive character than their absence inSyntermes.

8. Acanthotermes is a fungus-growing ter-mite in common with all the genera of theMacrotermitinae, while Syntermes does notcultivate fungi in common with all genera ofthe Nasutitermitinae and other termites.Inasmuch as each of these primitive genera

of the Macrotermitinae and Nasutitermitinaehas generalized characters not found in theother subfamily, we must suppose thatneither developed directly from the other,but that both are descended from an extinctgroup possessing the primitive characters ofboth of these subfamilies.

4411945


KEY TO THE SPECIES OF SyntermesIMAGOES

1. Wing membrane brown . . . . . . . . . 2Wing membrane hyaline . . . . . . . . 5

2. Pronotum distinctly wider than the head . 3Pronotum distinctly narrower than the head. 4

3. Pronotum 4.27-4.77 mm. wide . . S. snyderiPronotum 3.72-4.27 mm. wide (fig. 2) . S. dirusPronotum 2.99-3.6 mm. wide. . . S. territus

4. Pronotum 3.6-3.72 mm. wide, 1.58 mm. long(color of wing not known) . . S. peruanus

Pronotum 2.99-3.6 mm. wide, 1.46-1.67 mm.long............ . S. territus

Pronotum 2.74-3 mm. wide, 1.38-1.41 mm.long........... . S. parallelus

S. Pronotum 4.25 mm. wide, diameter of eye1.19 mm. ..... . . . . . S. obtusus

Pronotum 3.88-4.08 mm. wide, diameter ofeye 1.03-1.15 mm. .... S. magnoculus

Pronotum 3.62-3.72 mm. wide, diameter ofeye 0.97-1.03 mm. . . . . . . S. grandis

Pronotum 3.6-3.72 mm. wide, diameter of eye0.85 mm. (color of wing not known).. . ......... . S. peruanus

Pronotum 2.41-3.03 mm. wide, diameter of eye0.71-0.88 mm........ . S. molestus

SOLDIERS

1. Tooth of right mandible comparatively large;tooth of left mandible definitely projectingwell beyond the apical margin . . . . 2

Tooth of right mandible comparatively smallor vestigial; tooth of left mandible not pro-jecting much beyond the apical margin.................. . 12

2. Notal spines very large; pronotum 6.61 mm.wide, nearly as wide as width of head (6.96mm.); head covered with curly hair (fig. 4) .............. . S. aculeosus

Notal spines of medium to short length. . 33. Length of left mandible more than half the

width of the head . . . . . . . . . . 4Length of left mandible about half the width

of the head (fig. 11) . . . S. brevimalatus4. Cutting edge of left mandible beyond the

notch not distinctly sigmoid. . . . . . 5Cutting edge of left mandible beyond the

notch distinctly sigmoid . . . . . . . 65. Larger species (width of head 6.58-7.1 mm.;

width of pronotum 5.08-5.79 mm.)............... . S. snyderi

Smaller species (width of head 6.02 mm.;width of pronotum 4.13 mm.) (fig. 5)............. . S. magnoculus

6. Profile of frontal tube showing a shallow anglewith the line of the top of the head; openingof gland vertical; frontal tube compara-tively large; sides of head strongly con-

verging toward the front; head coveredabundantly with straight bristles (fig. 6) .

... . . . . . . . . . . . . . S. solidusProfile of frontal tube showing a shallow angle

with the line of the top of the head; openingof gland somewhat upward; head coveredrather abundantly with straight bristles . 7

Profile of frontal tube not showing any anglewith the line of the top of the head or show-ing an extremely shallow angle; opening ofgland somewhat upward; head covered withfew to numerous straight bristles; length ofhead to side base of mandibles over 5.6 mm... . . . . . . . . . . . . . . . . 10

7. Large or medium-sized species; width of headover 5.6 mm. . . . . . . . . . . . . 8

Small species; width of head 4.94-5 mm. .

...... . . . . . . . . S. peruanus8. Cutting edge of apical tooth of left mandible

not or slightly sigmoid; tip to point of tooth1.19-1.59 mm.; head thickly covered withbristles; width of head 6.58-7.1 mm.; lengthof left mandible 3.9-4.23 mm.; frontal tubeprominent . . . . . . . . . S. snyderi

Cutting edge of apical tooth of left mandibledistinctly sigmoid . . . . . . . . . . 9

9. Length of left mandible 4.14-4.37 mm.; widthof head 6.2-7.05 mm.; tip to point of toothof left mandible 1.21-1.32 mm.; head witha number of bristles but not thickly cov-ered, frontal tube not prominent (fig. 7)

S. chaquimayensis subspecies parvinasusLength of left mandible 3.66 mm.; width of

head 6.64 mm.; angle between frontal tubeand top of head in profile comparativelysharp (1350)..............

S.. chaquimayensis subspecies emersoniLength of left mandible 3.47-3.78 mm.; width

of head 5.67-6.32 mm.; angle between fron-tal tube and top of head in profile flat (160°)... . . . . . . . . S. c. chaquimayensis

10. Sides of head straight, converging only slightlytoward the front; width of head 5.37-5.79mm.. . . . . . . . . . . . . . . .11

Sides of head straight, converging markedlytoward the front; width of head 5.61-6.34 mm.; length of head to side base ofmandibles 5.67-6.39 mm.. . . S. grandis

Sides of head a little curved, converging alittle toward the front; width of head 5.92-6.14 mm.; length of head to side base ofmandibles 6.58-6.77 mm. (fig. 8). S. lighti

11. Length of left mandible 3.66 mm. (fig. 9) .

... . . . . . . . . . . . . . S. hageniLength of left mandible 3.23-3.47 mm. (fig. 10).S.. . . . . . . . . . . . . S. wheeleri

12. Notal spines medium-sized .... . . . 13Notal spines short, sides of head parallel or

442 VOL. 83


slightly converging. . . . . . . . . .15Notal spines absent . . . . . . S. molestus

13. Sides of head slightly converging toward thefront . . . . . . . . . . . . . . . 14

Sides of head strongly converging toward thefront and nearly straight; opening of frontaltube forward or a little upward with an an-gle at the junction of the frontal tube andvertex; width of pronotum 3.63-3.84 mm.(fig. f2) . . . . . . . . . . . S. calvus

14. Frontal tube elevated above the head at an-

curved; width of pronotum 3.17-3.23 mm. .. . . . . . . . . . . .... S. peruanus

Frontal tube hardly elevated above head atanterior margin, posterior portion continu-ous with the top of head without angle;opening of frontal tube upward at angle ofabout 450 to the long axis of the head; sidesof head nearly straight; width of pronotum3.66-4.16 mm.. . . . . . . . S. silvestrii

15. Width of head 4.21-5.24 mm. . . S. territusWidth of head 2.97-3.4 mm. . S. parallelus

FIG. 4. Head, pronotum, and postmentum of soldier of Syntermes aculeosus, new species, holotype.

terior margin, opening of frontal tube for-ward and nearly vertical to the axis of thehead or slightly upward at an angle of 45°or greater; sides of head a little curved;width of pronotum 3.54-4.17 mm.. S. dirus

Frontal tube hardly elevated above the headat the anterior margin; opening of the fron-tal tube slightly upward at an angle to theaxis of the head; sides of head a little

Syntermes aculeosus, new speciesSOLDIER (FIG. 4): Head, labrum, base of

mandibles, postmentum, thoracic nota, legs,sternites, and posterior tergites thickly cov-ered with hair, many of the hairs rather deli-cate and curved, giving a fuzzy appearance.Head stocky, widest behind, sides straightand converging toward the front; frontal

1945 443


tube wrinkled, forming a flat angle with thevertex, opening forward and slightly upward.Antennae incomplete in specimen, one with19 articles, subapical articles about 0.18-0.21mm. wide and 0.47 mm. long. Labrum three-pointed with white tip, all points a little moreobtuse than a right angle. Postmentumstocky, narrowest portion about two-thirdsthe width of the widest portion. Left mandi-ble with a large tooth forming an angle withthe apical portion somewhat less than a rightangle, apical cutting edge not sigmoid; rightmandible also with large tooth which has abroken tip in the specimen. Front margin ofpronotum emarginate; lateral spines veryrobust, thick, and conical. Spines on the meso-nota and metanota also robust and conicalsimilar to the prothoracic spines but directeda little posteriorly.Total lengthLength of head with mandiblesLength of head to side base of mandi-

blesWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

20.00 mm.10.34

7.146.964.323.761.771.062.076.616.237.337.33

LARGER WORKIER: I judge this worker tobe conspecific with the soldier because of thepilosity, but I cannot be sure of the asso-ciation. Color similar to soldier; antennaebrownish. Pilosity a little thicker than S.brevimalatus. Thoracic spines a little thickerthan S. brevimalktus.Width of head 3.00 mm.Width of pronotum 2.59COMPARISONS: Syntermes snyderi soldiers

have shorter and straighter bristles. The an-tennae are less robust, with the subapical ar-ticles about 0.17 mm. wide. The mandiblesare similar, but the inside edge of the apicalportion in S. snyderi is sometimes more sig-moid, with a sharper angle at the tooth in theleft mandible, and the tip of the left mandibleis more hooked. The lateral spines of thethoracic nota are much less robust in S.snyderi.

Syntermes magnoculus has a similar-shapedmandible. The size is smaller (head width6.02 mm.). The notal spines are much smaller(width of pronotum 4.13 mm.). The pilosityis not so abundant or so curly.

Syntermes aculeosus is unique among thedescribed species of Syntermes in the type ofpilosity and in the enormous size of the tho-racic spines. It is closest to S. snyderi and S.magnoculus in the shape of its mandibles.Although I have only a single specimen, Ihave no hesitation in describing the indi-vidual as belonging to a new species.

DISTRIBUTION (FIG. 1): Oronoque River (02°42'N., 57025' W.), British Guiana (type locality),1 soldier (holotype), 1 worker, coll. N. A. Weber,22.VII.1936, No. 584. The single soldier andworker were found in a vial with S. brevimalatus.

Syntermes obtusu HolmgrenTermes grandis SILVESTRI, 1901, p. 4.Termes grandis SILVESTRI, 1903, p. 49 (imago,

workers), p. 116 (biology), pl. 2, figs. 79-83(imago, worker).Syntermes obtusus HOLMGREN, 1911, pp. 546,

547 (imago).Syntermes obtusus SNYDER, 1924, p. 22 (imago).

IMAGO: Antennae with 21 articles. Wingshyaline with yellow brown costal border andradial veins. Membrane hairs absent near thesuture on the inner portion, long on the outermargin giving a ciliated appearance and al-most disappear on the outer portion.Length with wingsLength without wingsLength of head to side base of mandi-

blesWidth of headDiameter of eyeEye from lower margin of headLength of ocelliWidth of ocelliOcelli from eyesLength of fontanelleLength of left mandibleLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wingWidth of anterior wing

40.00 mm.19.50

3.424.581 .190.320.480.290.650.522.122.174.256.33

31.477.10

COMPARISONS: Syntermes magnoculus issmaller but is very close and could conceiv-ably belong to the same species. The imago is

VOL. 83444


somewhat larger than S. grandis and haslarger eyes. These species are quite closelyrelated. Males were compared to males toavoid sexual differences.

DISTRIBUTION (FIG. 1): Villa Rica (250551 S.,560271 W.), Paraguay (type locality), 2 imagoes(types), workers, det. N. Holmgren, det. F. Sil-vestri as T. grandis, coll. F. Silvestri, 8.X.1900,Silvestri Coll., Portici.

Syntermes magnoculus SnyderSyntermes magnoculus SNYDER, 1924, pp. 22, 23

(imago, soldier), pl. 4, fig. 22 (soldier).

IMAGO: Head dark brown. Wing membranehyaline. Fontanelle yellow, the same color asthe ocelli. Front and top of the head with afew long hairs. Fontanelle round, about thesame size as the ocelli, in a depression. An-tennae with 20 articles. Pronotum with an-terior angles not very sharp, angle at the tipgreater than a right angle. Pronotum strik-ingly saddle-shaped with a depressed linenear the anterior margin and both sides, butnot in the middle.

Length with wingsLength of head with mandi-

blesWidth of headDiameter of eyeEye from lower marginLength of ocellusWidth of ocellusOcelli from eyeOcelli from fontanelleLength of fontanelleWidth of fontanelleLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wingfrom suture

Width of anterior wing

and converging a little toward the front. An.terior edge of frontal tube raised a little abovethe rest of the head, posterior edge making aflat angle with the vertex, the gland openingforward but not quite vertical to the longi-tudinal axis of the head. Antennae with 20articles. Lateral angles of labrum aboutequal to a right angle, tip white. Left mandi-

MALE FEMALE39.50 40.00 mm.

4.894.141.030.290.410.230.63

0.440.351.883.886.86

4.391.150.290.470.350.640.82

2.014.08

31.006.77

COMPARISONS: Syntermes obtusus is closeand might be the same species. However, thetype of S. obtusus has a head width of 4.58mm. and a pronotum width of 4.25 mm.

Syntermes graniis is very close. The pro-

notal angles are similar, but the frontal lobeof the pronotum of S. grandis rises at a

sharper angle from the main portion.SOLDIER (FIG. 5): Head with a few scat-

tered bristles. Sides of head fairly straight

FIG. 5. Left mandible of soldier of Syntermesmagnoculus Snyder, topotype.

ble with curved apical cutting edge not sig-moid, with a large first marginal tooth mak-ing an angle of about 600 with the apicalcutting edge. Front margin of the pronotumemarginate. Notal spines comparatively ofmedium length.Length of head to side base of mandi-

bles 6.22 mm.Width of head 6.02Length of left mandible 3.90Widest width of postmentum 1.50Narrowest width of postmentum 0.96Length of pronotum 1.89Width of pronotum 4.13Width of mesonotum 3.17Width of metanotum 4.08

COMPARISONS: The closest species seems tobe S. snyderi which is distinctly larger, has

4451945


more bristles on the head, and a more sig-moid apical cutting edge of the left mandible.

DISTRIBUTION (FIG. 1): Chapada (14010' S.,590491 W.), Matto Grosso, Brazil (type locality),1 female (holotype), det. T. E. Snyder, coll. H. H.Smith, X, type No. 14510, M.C.Z.; 1 male (para-type), det. T. E. Snyder, coll. H. H. Smith, X,U.S.N.M.; 1 soldier (topotype), det. T. E. Snyder,VII, M.C.Z.

Syntermes snyderi Emerson? Termes dirus WALKER, 1853, p. 510 (imago,

soldier, worker).Syntermes snyderi EMERSON, in Snyder, 1924,

p. 21 (imago), p. 22 (soldier).Syntermes dirus SNYDER, 1924, pp. 27, 28

(soldier from Kartabo, British Guiana, only).Syntermes snyderi EMERSON, 1925, pp. 294, 310,

358 (imago, soldier), text fig. 44 (imago, soldier).Syntermes snyderi SNYDER, 1926, pl. 2, fig. 7

(soldier).Syntermes dirus WHEELER, 1936, p. 176 (biol-

ogy, British Guiana only).Syntermes snyderi WHEELER, 1936, pp. 174, 176,

221 (biology).Syntermes snyderi HARE, 1937, p. 468 (sys-

tematics), fig. 28 (worker and soldier mandibles).Syntermes snyderi EMERSON, 1938, p. 258 (nest).IMAGO: Head, pronotum, and sternites

orange yellow brown, wings and tergitesbrown contrasting quite strongly with yellow-ish regions. Head and pronotum with a fewbristles. Wing scales with numerous bristles.Fontanelle round, light, larger than ocelli.Eyes small, prominent, about one-third theirlength from the lower margin. Ocelli of me-dium size, over twice their length from theeyes and about midway between the eyes andfontanelle. Antennae with 19 or 20 articles.Pronotum wider than head, anterior anglesquite sharp and outturned, the tip aboutequal to a right angle.Length with wingsLength of head with mandiblesLength of head to side base of

mandiblesWidth of headDiameter of eyeEye from lower marginLength of ocelliWidth of ocelliOcelli from eyesOcelli from fontanelleLength of fontanelleWidth of fontanelle

35.00-42.50 mm.4.76- 4.88

2.93- 3.113.57- 3.920.71- 0.75

0.260.320.230.710.710.410.41

Width of labrumLength of postclypeusWidth of postclypeusLength of left mandibleLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wing from

sutureWidth of anterior wing

1.610.651.762.25

2.01- 2.104.27- 4.776.78- 7.09

30.837.33- 7.80

COMPARISONS: The closest species seems tobe S. dirus which, however, has a narrowerpronotum in all the specimens examined. Thecoloration seems to be quite similar.

SOLDIER: Head, labrum, base of mandibles,postmentum, pronotum, and abdomen cov-ered with a great many straight bristles. Headvery large, sides straight and convergingsomewhat toward the front. Frontal tubewith a flat angle where it joins the vertex anddistinctly elevated above the head in front,opening nearly vertical to about 450 to theaxis of the head. Postmentum stocky, narrow-est width about two-thirds of widest width,Antennae with 19 or 20 articles. Left mandi-ble with apical cutting edge somewhat orslightly sigmoid forming an angle with thelarge prominent tooth of about 600 or a littlewider (450 in the Venezuelan specimens).Right mandible with a large marginal toothforming an angle with the apical cutting edgeof about 750. Front margin of pronotum emar-ginate or evenly curved. Spines fairly largeand of about equal size on the three nota.

Length of head to sidebase of mandibles

Width of headLength of left mandi-

bleLength of left mandi-

ble from tip to tipof tooth

Length of postmen-tum

Widest width of post-mentum

Narrowest width ofpostmentum

Length of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

No. RANGE

2 6.11-6.5814 6.58-7.10

13 3.90-4.23

MEAN

6.35 mm.6.81

4.13

10 1.19-1.59 1.42

1 3.63 -

1 1.76

1 1.121 1.88

14 5.08-5.791 4.791 5.643 6.49-6.74

5.39

6.60

446 VOL. 83

447EMERSON: THE NEOTROPICAL GENUS SYNTERMES

COMPARISONS: Syntermes snyderi seems tobe closest to S. solidus which has a larger,more prominent frontal tube, a more stronglysigmoid apical cutting edge of the left mandi-ble, and a less projecting marginal tooth.The Venezuelan form is close to S. chaqui-

mayensts, subspecies parvinasus which hasfew bristles and a less prominent frontal tubeand a distinct sigmoid apical cutting edge ofthe left mandible.The species referred to S. dirus from Kar-

tabo, British Guiana, by Snyder (1924) andby Wheeler (1936, p. 176) belongs to S.snyderi.

Walker (1853, p. 510) records T. dirus fromBritish Guiana. It would seem best to con-sider this record as a synonym of S. snyderiuntil Walker's specimens are reexamined.

DISTRIBUTION (FIG. 1): Kartabo (06023' N.,58042' W.), British Guiana (type locality), 3imagoes (morphotype and paratypes), 8 soldiers(holotype and paratypes), coll. A. Emerson,V.1919, No. 52; 1 male, 1 female (topotypes),coll. A. Emerson. 2.VI.1924, No. 24.162g and un-numbered vial; 2 soldiers (paratypes), coll.A. Emerson, 5.VIII.1920, M.C.Z. Indians broughtme alates at Kartabo which they said were flyingon 4.V.1919. Flying alates were observed on2.VI.1924 at about 6 P.M. Some were observed100 to 150 feet up in the air.

St. Edward's Mission on the Mazaruni River2 miles above Kartabo, British Guiana, imagoes,soldiers, workers in stomach contents of a toad,Bufo typhonius, coll. by Indians, 6.VI.1924, No.24.170.

Surinam, 1 soldier, coll. A. Reyne, No. 10;1 soldier, coll. A. Reyne, No. 13. These soldiershave fewer bristles on the top of the head than thetype colony, but portions of the head and prono-tum have similar pilosity.

Venezuela, 3 soldiers, from Chicago M.N.H.These Venezuelan forms have a somewhat shorterleft mandible than the types, but the variationseems too slight for taxonomic distinction. Theangle formed by the tooth in the left mandible isnarrower. Future data may indicate subspecificdistinction between these forms.

Syntermes solidus, new speciesSOLDIER (FIG. 6): Color of specimen black

due to preservation. Portions indicate thatoriginal color was yellow brown. Head, la-brum, base of mandibles, postmentum, tho-racic nota, legs, tergites, and sternites ratherthickly covered with straight and fairly long

bristles. Head widest behind, sides straightand converging toward the front. Frontaltube comparatively large, not in much of adepression, but front concave and wrinkled,opening of gland vertical to long axis of head.Antennae broken in specimen. Tip of labruma little more obtuse than a right angle, lat-eral angles blunt and about equal to rightangles. Narrowest portion of postmentumover half the width of the widest portion. Leftmandible with a prominent marginal tooth,the angle with the apical tooth less than 450,the cutting edge of the apical tooth sigmoid.Right mandible with a fairly large toothforming an angle with the apical portion alittle less than a right angle, apical cuttingedge only barely suggesting a sigmoid shape.Front margin of pronotum emarginate, lat-eral spines fairly robust and conical. Meso-notal and metanotal spines also fairly robustand a little smaller than pronotal spines.Total length 18.00 mm.Length of head with mandibles 9.21Length of head to side base of mandi-

blesWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

6.716.733.903.541.470.902.075.003.804.636.34

COMPARISONS: The species is most closelyrelated to S. chaquimayensis from which itdiffers in its larger frontal tube, more con-vergent sides of the head, and sharper angleof the notch of the left mandible.

Syntermes snyderi is quite close but hassomewhat longer left mandibles, and thetooth of the left mandible extends farther andforms a wide-angled notch.

DISTRIBUTION (FIG. 1): St. Jean (05022' N.,5406' W.), Moroni River, French Guiana (typelocality), 1 soldier removed from pin (holotype),coll. Le Moult.

Syntermes chaquimayensis chaquimayensis(Holmgren)

Termes chaquimayensis HOLMGREN, 1906, p. 547(soldier, workers).

1945


FIG. 6. Dorsal view of head and pronotum and ventral view of head of soldier of Syntermes solidusnew species, holotype.

Syntermes chaquimayensis HOLMGREN, 191 1, pp.546-S548 (soldier).

Syntermes chaquimayensis SNYDER, 1924, pp. 22,26 (soldier).

Syntermes chaquimayensis SNYDER, 1926, p. 17(locality), pl. 2, fig. 4 (soldier).Syntermes chaquimayensis WHEELER, 1936, pp.

173, 176 (biology).

SOLDIER: Head, base of mandibles, post-mentum, thoracic nota, tergites, and stern-ites covered rather abundantly with bristles,about equal in pilosity to subspecies par-vinasus (fig. 7). Sides of head fairly straight,converging a little toward the front. Frontaltube lying in a slight depression, openingfacing slightly upward at an angle, posteriorjunction of tube with the head forming a flatangle (about 1600). Antennae with 19 or 20articles, penultimate article 0.16 mm. wideand 0.41 mm. long. Tip of labrum sharperthan a right angle, side points about equalto right angles or slightly more blunt. Nar-rowest width of postmentum about two-

thirds the widest width. Left mandible witha prominent marginal tooth extending some-what beyond the apical cutting edge andforming an angle of about 450 to 600 with theapical cutting edge which is sigmoid. Rightmandible with a prominent marginal tooth.Front margin of pronotum emarginate or notemarginate. Lateral spines on the thoracicnota comparatively of medium size and aboutequal in length and thickness on the threesegments.

I have included the total length given byHolmgren (1911, p. 548).LARGE WORKER: Antennae with 20 arti-

cles. Width of head 3.41 mm.COMPARISONS: This species may tentatively

be subdivided into three subspecies. There isno doubt that these subspecies are quiteclosely related, and further samples of thepopulations from various localities as well asthe collection of the reproductives are neces-sary for the validation or refutation of thesuggested categories. The typical subspecies is

448 VOL. 83


SOLDIERTotal lengthLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

very close to S. chaquimayensis emersoni, butis a little smaller and has a flatter angle(1600) at the junction of the frontal tube andvertex. S. chaquimityensis parvinasus is larger,and the frontal tube is proportionatelysmaller.

DISTRIBUTION (FIG. 1): Chaquimayo (13044' S.,70039' W.), Rio San Gaban Valley, Prov. deCarabaya, Peru (type locality). I have no speci-mens surely from the type locality but have3 soldiers (probably cotypes), det. N. Holmgren,locality unknown.

Llinquipata (13°49' S., 70°38' W.), Peru, 2 sol-diers (cotype), det. and coll. N. Holmgren,XI.1904.

Perene (10°53' S., 75°12' W.), Peru, 5 soldiers(metatypes), workers, coll. J. C. Bradley, 26.VI.1920, No. 377.Huachi t15036' S., 67007' W.), Bolivia, 11 sol-

diers (metatypes), workers, det. T. E. Snyder,coll. W. M. Mann.

Near Philadelphia (17053' S., 41019' W.),Brazil, 1 soldier, coll. Hartt and Copeland, ThayerExpedition, M.C.Z. This specimen from easternBrazil is geographically removed from the Peru-vian and Bolivian series and might conceivablybe different. Aside from somewhat narrowerthoracic nota, however, I am unable to makeany distinction and think it best to consider itconspecific until more specimens are available.The species is also recorded from El Campami-

ento (Col. Perene), Peru; and near Buturu(140171 S., 68035' W.) [Tuiche Valley], CocosValley, between Rio Azata and Rio Tavara in theTambopata Valley (Prov. Caupolican), Bolivia.

Syntermes chaquimayensis, subspeciesemersoni Snyder

Syntermes emersoni SNYDER, 1924, pp. 22, 26(soldier), pl. 4, fig. 20 (soldier).SOLDIER: Pilosity close to S. c. chaqui-

mayensis. There seem to be fewer hairs on the

No. RANGE5 15.00-18.5028

21745537554

8.83-5.73-5.67-3.47-3.20-1.34-1.01-1.83-4.25-3.62-4.08-5.55-

9.216.346.323.783.661.711.091.954.884.395.005.85

MEAN16.70 mm.9.026.085.893.613.491.541.051.894.613.964.595.64

base of the mandibles than in S. c. chaqui-mayensis. There are fewer hairs on the headin places, but this difference may be due torubbing. The hairs are stiff and straight as inS. c. chaquimayensis. The angle between theposterior portion of the frontal tube and thevertex is about 1350 in profile while this angleis flatter in S. c. chaquimayensis. The generalproportions of the head and the mandibledentation are virtually the same as in S. c.chaquimayensis.Length of head with mandibles (not

extended)Estimated length of head with man-

dibles extendedLength of head to tip of labrumWidth of headLength of left mandibleLength of left mandible from tip to

point of toothLength of right mandible from tip to

point of toothLength of pronotumWidth of pronotumLength of hind tibia

9.78 mm.

10.158.086.643.66

1.34

1.382.015.306.52

COMPARISONS: This form was originally de-scribed by Snyder (1924) as a separate spe-cies, but the morphology is very close to S.chaquimayensis. The larger size, slight differ-ence in pilosity, and small difference in theangle between the frontal tube and the vertexmay not be characteristic of the populations,but I am tentatively including the form as asubspecies of S. chaquimayensis on admit-tedly very inadequate data.

DISTRIBUTION (FIG. 1): Iguaripe (possiblyAguaripe, 13°08' S., 380551 W., Bahia), Brazil(type locality), 1 soldier (holotype), det. T. E.Snyder, type No. 25748, U.S.N.M.

1945 449


Syntermes chaquimayensis, subspeciesparvinasus, new subspecies

SOLDIER (FIG. 7): Head, labrum, base ofmandibles, postmentum, thoracic nota, legs,sternites, and tergites covered rather abun-dantly with straight bristles. Head converg-ing somewhat in front, sides straight. Frontaltube comparatively very small, raised only alittle above the vertex at the posterior edge,anterior edge level with the front, opening

the cutting edge of the apical tooth distinctlysigmoid, cutting edge posterior to marginaltooth undulating, a small notch anterior tothe basal tooth at the junction of the light anddark portions of the mandible. Right mandi-ble with fairly large marginal tooth whichmakes an angle with the apical portion alittle less than a right angle; the cutting edgeof the apical tooth is very slightly convex ornearly straight near the notch. Front margin

FIG. 7. Dorsal view of head and pronotum and ventral view of head of soldier of Syntermes chaqui-mayensis, subspecies parvinasus, new subspecies, holotype.

pointing up at an angle of about 45°. An-tennae broken in all specimens examined. Tipof labrum about equal to a right angle orslightly greater or less than a right angle,lateral points about equal to right angles.Narrowest portion of postmentum over halfas wide as the widest portion. Left mandiblewith a large tooth forming an angle with theapical portion of about 450 or slightly less,

of the pronotum emarginate. Lateral spinesfairly large, hind margin nearly straight. Lat-eral spines of mesonota and metanota nearlyas large as the pronotal spines.COMPARISONS: Close to S. solidus, but the

frontal tube is not so conspicuous and thenotch of the left mandible anterior to thefirst marginal tooth is wider than in S. solidus.Close to S. snyderi, but the frontal tube is not

VOL. 83450

451EMERSON: THE NEOTROPICAL GENUS SYNTERMES

SOLDIERTotal lengthLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of left mandible from tip to point of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

so conspicuous and the head bristles are fewerin S. c. parvinasus. The morphological rela-tionship of this form to S. c. chaquimayensisis so close that I am tentatively placing it ina subspecies until more data are available.,S. c. chaquimayensis is smaller in a number ofmeasurements and has a proportionatelylarger frontal tube. S. c. emersoni has a shorterleft mandible, and the posterior edge of thefrontal tube rises in a steeper angle from thevertex,

DISTRIBUTION (FIG. 1): Rio Waupes (oo00'o-02001' N., 67000-72°38' W.), Brazil or possiblyColombia (type locality), 5 soldiers (holotype andparatypes), 190a, Box No. 23.

Ireng River (05016' N., 60°22'-60054' W.) toRoraima, Brazil, 3 soldiers (paratypes), VIII.1911,M.C.Z., A.M.N.H. S. grandis was in the same vial.

Syntermes grandis (Rambur)Although all the museum specimens as-

signed to this species have not been examinedby the author (specimens reexamined by meare marked with an asterisk), there is noreason for not assuming that the followingsynonymy is accurate. Latreille incorrectlydetermined his species as Perla fusca DeGeer and placed it in the genus Ternmes.Perla fusca DeGeer is a synonym of Copto-termes testaceus (Linnaeus), and thus Ram-bur's subsequent name is valid.

Termesfuscum LATREILLE, 1805a, p. 68 (imago).Termesfuscum LATREILLE, 1805b, p. 60 (imago).Termes grandis RAMBUR, 1842, p. 306 (imago).*Termes decumanus ERICHSON, 1848, p. 582

(imago).Termes costatus WALKER, 1853, p. 518 (imago).Termes grandis WALKER, 1853, p. 519 (locality).

No. RANGE4 18.50-20.004 9.87-10.346 6.78- 7.478 6.20- 7.054 4.14- 4.372 1.21- 1.326 3.95- 4.276 1.59- 1.766 0.94- 1.126 1.88- 1.998 4.93- 5.476 4.27- 4.576 5.00- 5.244 4.76- 6.22

MEAN19.00 mm.10.157.056.644.211.264.021.631.011.945.174.435.145.63

*Termes grandis HAGEN, 1858a, p. 157 (imago,soldier), pl. 2, fig. 10, pl. 3, fig. 18 (imago).

*Termes grandis HAGEN, 1858b, p. 22 (imago).*Syntermes grandis HOLMGREN, 1911, pp. 546-

548 (imago, soldier).Termes grandis DESNEUX, 1915, p. 8 (location of

type).*Syntermes grandis SNYDER, 1924, pp. 22, 24, 27

(imago, soldier), pl. 4, fig. 23 (soldier).*Syntermes grandis EMERSON, 1925, p. 294

(locality).*Syntrmes grandis BEQUAERT, 1925, p. 294

(biology).*Syntermes grandis WHEELER, 1936, pp. 174,

176 (biology).IMAGO: Head fairly dark brown with lighter

yellowish postclypeus, labrum, and pro-notum. Wings hyaline and transparent exceptfor a yellow brown costal border and radialregion. Head, pronotum, and wing scales withnumerous long bristles. Fontanelle about as.long as, or a little shorter than, the length ofthe ocellus, round, yellowish. Eyes relativelysmall, oval, and prominent. Ocelli of mediumsize, closer-to the eyes than to the fontanelle.Antennae with 20 articles, the third longerthan the second and the second longer thanthe fourth. Pronotum narrower than thehead, somewhat saddle-shaped, anterior an-gles about equal to, or slightly more than, a.right angle, sides curved, hind margin con-spicuously emarginate.Length with wingsLength without wingsLength of head with mandiblesLength of head to side base of

mandiblesWidth of headDiameter of eye

39.30 mm.18.004.63

3.114.08- 4.290.97- 1.03

1945


Eye from lower marginLength of ocelliWidth of ocelliOcelli from eyesOcelli from fontanelleLength of fontanelleWidth of fontanelleWidth of labrumWidth of postclypeusLength of postclypeusLength of left mandibleLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wing scaleLength of anterior wing from

sutureWidth of anterior wing

0.18- 0.290.35- 0.390.23- 0.320.59- 0.610.68- 0.820.29- 0.410.23- 0.41

1.531.590.652.03

1.95- 2.013.62- 3.72

6.252.19

29 .89-30 .246.96- 7.34

COMPARISONS: Syntermes dirus has brownerwing membranes, somewhat larger fontanelle,smaller eye, ocelli in a more midway positionbetween the fontanelle and eyes, pronotum

SOLDIERLength of head to side base of mandiblesWidth of headLength of left mandibleLength of left mandible from tip to tip of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

wider than head, and sharper anterior anglesof the pronotum.

Syntermes magnoculus is quite close with alittle larger fontanelle, pronotum angles thesame, but has less of a frontal lobe rising at aflatter angle from the main part of the pro-notum.

SOLDIER: Head, base of mandibles, post-mentum, pronotum, and abdomen coveredwith numerous bristles. Head large, sidesconverging toward the front, fairly straight.Frontal tube virtually without an anglewhere it joins the vertex and front marginhardly elevated above the head, frontal open-ing slightly upward from the vertical. An-tennae with 20 articles, penultimate article0.15 mm. wide and 0.38 mm. long. Lateral

angles of labrum blunt, about equal to aright angle. Narrowest portion of postmen-tum four-sevenths to three-fourths width ofwidest portion. Left mandible with apicalcutting edge sigmoid, marginal tooth project-ing well beyond apical cutting edge and form-ing a notch with an angle of about 450 or alittle wider. Right mandible with a prominentmarginal tooth forming a notch slightly lessthan a right angle. Front margin of pronotumslightly emarginate. Thoracic nota withrather short spines.

COMPARISONS: Syntermes hageni is quiteclose, but the head is narrower, the sides aremore parallel, the capsule at the ventralmandibular condyle is less rounded, and theprothoracic spines are not so blunt. The tho-racic spines of the holotype are a little longerthan in the morphotype of S. grandis.

Syntermes lighti is quite close, but the headis longer and the sides are more parallel, the

No. RANGE8 5.67-6.39

11 5.61-6.349 3.18-3.668 0.91-1.122 3.29-3.417 1.23-1.407 0.81-0.927 1.46-1.71

11 3.29-4.108 2.74-3.479 3.23-4.477 5.61-5.88

MEAN6.07 mm.5.933.451.043.351.330.861.603.633.103.795.71

mandibles are a little less hooked, the tho-racic spines are a little longer, the pronotalspines are sharper, the frontal tube is similar,and the antennal articles are a little longerand thicker.

DISTRIBUTION (FIG. 1): Cayenne (type locality).Rambur erroneously recorded the species fromSenegal.Unknown locality, 1 soldier (morphotype), det.

N. Holmgren, det. H. Hagen as T. grandis,labeled "Hagen revid, 1856"; 1 soldier, det. T. E.Snyder, coll. P. Rambur, M.C.Z.; 1 soldier, un-labeled but associated with the Erichson imagofrom British Guiana, Berlin Mus.

British Guiana, 1 imago (cotype of decuma-nus), det. H. Hagen as T. grandis, det. W. F.Erichson as T. decumanus, coll. R. Schomburgk,No. 2764, Berlin Mus.; 1 female (cotype of de-

VOL. 83452


cumanus), det. H. Hagen as T. grandis, det. W. F.Erichson as T. decumanus, coll. R. Schomburgk,M.C.Z.Arabupu (05°00' N., 60045' W.), Mt. Roraima,

Brazil, soldiers, coll. G. H. H. Tate, XII.1927,4200 feet, G.87, Acc. 29780, A.M.N.H.

Ireng River (05016' N., 60°22'-60°54' W.) toRoraima, Brazil, soldiers, VIII.1911, M.C.Z.,A.M.N.H.

Tapajos River (02°27'-07°27' S., 54421-57055'W.), Brazil, soldiers, VIII, Thayer Expedition,M.C.Z., A.M.N.H.

Vista Alegre (01°35' N., 61009' W.), Rio Branco,Brazil, 1 soldier, det. T. E. Snyder, coll. J. Be-quaert, foraging in daytime.Santarem (03030' S., 54020' W.), Brazil, sol-

diers, det. T. E. Snyder, coll. Thayer Expedition.Congonhas (20054' S., 45047' W.), Brazil, 1 sol-

dier, det. H. A. Hagen as T. dirus, coll. H. Bur-meister, M.C.Z.

Bahia (13010' S., 38050' W.), Brazil, 1 male, det.N. Holmgren, 22.11.1903.Akuriman (not found on map but located in

upper Caroni River basin [05°-07° N., 61°30'-

63020' W.], Bolivar), Venezuela, soldiers, coll.P. J. Anduze, XI. 1940.

Syntermes lighti, new species

Syntermes peruanus SNYDER, 1926, p. 15, pl. 2,fig. 8 (soldier).SOLDIER (FIG. 8): Head, labrum, base of

mandibles, postmentum, thoracic nota, legs,tergites, and sternites covered with bristlesbut comparatively sparse on the head. Headlarge, widest behind, sides slightly curved andconverging somewhat toward the front.Frontal tube very short, posterior edge join-ing the vertex with an even curve in profile,opening facing upward at an angle. Antennaebroken in the specimens before me. Lateralangles of labrum about rectangular, someslightly more and some slightly less than aright angle. Narrowest portion of post-mentum about two-thirds width of widestportion. Left mandible with large tooth join-

/

FIG. 8. Dorsal view of head and pronotum and ventral view of head of soldier of Syntermes lighti,new species, holotype.

1945 453


ing apical cutting edge at an angle of about450, apical cutting edge sigmoid. Right man-dible with large tooth joining apical cuttingedge at an angle somewhat less than a rightangle. Margin of basal portion of mandibleon outer edge about a third of the length ofthe outer edge from base. Front margin ofpronotum slightly emarginate. Lateral tho-racic spines comparatively rather small.

SOLDIERTotal lengthLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of left mandible from tip to point of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

COMPARISONS: The closest species is S.hageni, which differs especially in the smaller-sized head and the straighter sides of thehead. When more collections have beenmade, it is possible that gradations betweenthese two named species may indicate thatthey should be placed in subspecific or vari-etal rank.

Syntermes grandis is also close, but thesides of the head converge more toward thefront, and the pronotal spines are less long.

DISTRIBUTION (FIG. 1): Villa Bella (10027' S.,65028' W.), Bolivia (type locality), 4 soldiers(holotype and paratypes) removed from pins, det.T. E. Snyder, 1926, p. 15, as S. peruanus, coll.J. D. Haseman, 9.X.1909, Acc. 4043, CarnegieMus., A.M.N.H.

Caigara (03014' S., 64049' W.), Brazil, 2 soldiers(paratypes), coll. Natterer, from Vienna Mus.,M.C.Z.

REMARKS: I have named this species inhonor of Prof. S. F. Light of the University ofCalifornia whose work has greatly advancedour knowledge of both the taxonomy andphysiology of termites.

Syntermes hageni HolmgrenSyntermes dirus, form hageni HOLMGREN, 1911,

pp. 547, 548 (soldier).

SOLDIER (FIG. 9): Head, labrum, post-mentum, thoracic nota, legs, tergites, andsternites with bristles, rather sparse on thehead and postmentum, short and few on thebase of the mandibles. Head fairly large, sidesslightly curved and converging only a littletoward the front. Frontal tube short, poste-rior margin joined in even, shallow curve withvertex in profile, opening facing upward at an

No. RANGE2 17.00-17.504 9.39- 9.775 6.58- 6.775 5.92- 6.146 3.63- 3.724 1.09- 1.235 3.38- 3.966 1.47- 1.596 0.91- 1.065 1.71- 1.886 3.96- 4.326 3.17- 3.415 3.78- 4.214 6.10- 6.58

MEAN17.25 mm.9.596.706.053.691.183.741.510.971.794.093.344.056.31

angle of about 45°. Antennae with 20 articles,penultimate article 0.44mm. longand 0.16mm.wide. Lateral angles of labrum about equal toa right angle. Narrowest portion of postmen-tum a little less than two-thirds the width ofthe widest portion. Left mandible with mar-ginal tooth extending well beyond the apicalcutting edge and making an angle of about450, apical cutting edge sigmoid. Right man-dible with prominent marginal tooth makingan angle with apical cutting edge a little lessthan a right angle. Front margin of pronotumemarginate. Lateral spines of medium sizeand about of equal size in the three thoracicnota.Length of head with mandiblesLength of head to side base of mandi-

blesWidth of headLength of left mandibleLength of left mandible from tip to

point of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

9.30 mm.

6.205.373.66

1.223.781.380.821.683.963.183.846.28

454 VOL. 83


FIG. 9. Dorsal view of head and pronotum and ventral view of head of soldierof Syntermes hageni Holmgren, holotype.

COMPARISONS: This species was described,as far as can be determined, from a uniquesoldier originally from Hagen's collection.This specimen seems to belong to a dis-tinct species. Other specimens determined asthis species in the literature belong to otherspecies.

Syntermes wheeleri is close, but the headand mandibles are shorter in proportion tothe width of the head.

Syntermes grandis is close, but the headwidth is greater, the sides of the head con-verge more toward the front, the bump at theventral mandibular condyle is less sharp, thepronotal spines are a little less long and aremore blunt. The frontal tube and mandibulartype are similar.

Syntermes lighti is close, but the head islarger and the sides are more curved. Frontal

tube, mandibular type, and thoracic spinesare quite similar.

DISTRIBUTION: Brazil (type locality), 1 soldier(holotype) removed from pin, det. N. Holmgrenas "T. Hageni n. sp. revid. Hagen 1856, Brazil."

Syntermes wheeleri, new speciesSOLDIER (FIG. 10): Head with fairly numer-

ous scattered bristles. Labrum, postmentum,base of mandibles, thoracic nota, tergites, andsternites with numerous bristles. Head withsides slightly curved and converging only alittle toward the front. Frontal tube short,posterior margin joined in an even curve withthe vertex in profile, anterior portion onlyvery slightly elevated above the head, open-ing facing upward at an angle of about 45°-600 to the main axis of the head. Antennaewith 20 articles, penultimate article 0.15-

4551945


0.17 mm. wide. Lateral angles of labrumslightly less to slightly greater than a rightangle. Narrowest portion of postmentumabout four-sevenths to nine-fourteenths thewidth of widest portion. Mandibles compara-tively somewhat short in relation to width ofhead, hook at tip comparatively short. Leftmandible with a prominent marginal toothprojecting well beyond the border of theapical cutting edge and forming an angle of

about 700 with the apical cutting edge whichis distinctly sigmoid. Right mandible with aprominent marginal tooth (0.2 mm. widefrom the notch to the tip) making an angleabout equal to a right angle with the apicalcutting edge. Front margin of pronotumslightly emarginate. Lateral spines some-what short, a little larger than the mesonotaland metanotal spines.LARGE WORKER: Color of head similar but

a little more variegated than soldier. Head,pronotum, and tergites with numerous bris-tles. Fontanelle white, about the same size asthe antennae sockets. Antennae with 20articles. Front margin of pronotum slightlyor not emarginate. Thoracic nota with sharplateral spines.Width of headWidth of pronotum

FIG. 10. Head and pronotum of soldier ofSyntermes wheeleri, new species, holotype.

SOLDIERLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of left mandible from tip to point of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

3.72-3.96 mm.2 .35-2 .78

COMPARISONS: I regret the paucity of thematerial of this form, and the determinationas a new species must be considered tenta-tive. The specimens are closer to S. hagenithan to any other species. I am more confi-dent of the specific distinction of the otherspecies named from few specimens than I amin this case. In view of the range of variationin other forms, however, I think the mostprobable hypothesis is that the specimensrepresent a new species or subspecies.

$yntermes hageni differs in having a longerhead and mandibles, and sparser distribu-tion of hairs on the head and postmentum.The frontal tube and general mandibularcharacters are similar in the two species.

DISTRIBUTION (FIG. 1): Rio (probably Rio deJaneiro, 23°18' S., 430 18' W.), Brazil (type local-ity), 1 pinned soldier (holotype), det. H. Bur-

No. RANGE4 8.36-8.655 5.73-6.105 5.37-5.794 3.23-3.474 0.97-1.104 3.17-3.545 1.44-1.585 0.82-1.105 1.50-1.645 3.45-4.035 2.64-3.705 3.25-4.335 4.82-5.30

MEAN8.52 mm.5.865.603.361.053.321.490.941.613.893.374.035.08

456 VOL. 83


meister and H. Hagen as T. dirus, type No. 25707, scattered bristles. Labrum, legs, and sternitesM.C.Z. with bristles, and sternites also with shortSao Paulo (Ibirapu6ra) (23044' S., 46038' W.), hairs. Head stocky, nearly quadrangular,

Sao Paulo, Brazil (not included in map which was sides only slightly converging toward theprepared before this record), 4 soldiers (para- ftypes) with workers and nymphs, coll. R. L. front and slightly curved; frontal tube short.Araujo, lO.VIII.1944, No. 2322, hill-like nest. Antennae with 19 articles, penultimate arti-

7TI7FIG. 11. Dorsal view of head and pronotum and ventral view of head of soldier of Syntermes brevimaltus,

new species, paratype.

REMARKS: The holotype specimen seems tohave been part of the original series of T.dirus determined by Burmeister. The othertwo soldiers of the series conform to otherS. dirus material.

I have named the species in honor of thelate W. M. Wheeler whose personality andachievements were a source of great inspira-tion to me.

Syntermes brevimalatus, new speciesSOLDIER (FIG. 11): Antennae brown and

contrasting with yellowish head. Head, baseof mandibles, postmentum, thoracic nota,and tergites almost bare with only a few

cle about 0.15 mm. wide. Tip of labrum alittle greater than a right angle, side anglesabout equal to a right angle. Postmentumstocky, only a little constricted in the middle,narrowest portion more than two-thirdswidth of widest portion. Mandibles compara-tively short for this genus, curved. Left man-dible with a large marginal tooth, fairlyblunt point about equal to a right angle,basal anterior angle slightly greater than aright angle; second marginal tooth quiteblunt. Right mandible with a fairly largemarginal tooth, the tip greater than a rightangle and not forming a sharp angle with theapical tooth, but cutting edge fairly evenly

4571945


SOLDIERTotal lengthLength of head with mandiblesLength of head to side base of mandibleWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumLength of hind tibia

curved in contrast to the somewhat straightcutting edge of the left mandible just pos-terior to the hooked tip. Front margin of thepronotum emarginate, hind margin nearlystraight or slightly concave, lateral spines ofmedium size and length. Lateral spines ofmesonota and metanota not so large as thoseof the pronotum, directed laterally or slightlyposteriorly.LARGE WORKER: Color similar to soldier,

including the brown contrasting antennae.Head, thorax, and tergites with few scatteredbristles. Fontanelle white, smaller than an-tennae sockets. Antennae with 19 articles.Labrum hatchet-shaped, anterior marginevenly curved. Front margin of pronotumslightly emarginate. Pronotum with a smallround or elongate depression in center.Thoracic nota with sharp lateral spines.Width of headWidth of pronotum

3.35-3.45 mm.2.40-2.48

COMPARISONS: The soldiers of this specieshave the shortest mandibles in relation to thewidth of the head of any described species ofSyntermes. Also the head is about as bare ofhairs and bristles as any species of the genus.The species is so distinct from the others thatI do not know which is the most closely re-lated. Possibly when the imago is discoveredwe may have sufficient characters to form ajudgment.

DISTRIBUTION (FIG. 1): On Oronoque River(020421 N., 57025' W.), British Guiana (type local-ity), 3 soldiers (holotype and paratypes) withworkers, coll. N. A. Weber, 22.VII.1936, No. 584.Found in a vial with S. aculeosus. Weber's fieldnotes are included under general ecology of thegenus.

Syntermes peruanus HolmgrenTermes dirus HOLMGREN, 1906, p. 545 (imago,

soldier, workers).

No. RANGE2 18.003 9.68-9.963 7.33-7.433 6.393 3.17-3.232 4.32-4.332 1.44-1.473 1.00-1.123 1.58-1.643 4.39-4.632 4.57-4.63

MEAN18.00 mm.9.847.366.393.214.331.461.061.614.494.60

Syntermes dirus HOLMGREN, 1909, p. 85 (anat-omy).

Syntermes peruanus HOLMGREN, 1911, pp. 546,548 (imago, soldier).

? Syntermes bolivianus HOLMGREN, 1911, pp.547, 548 (soldier).

? Syntermes bolivianus HOLMGREN, 1912, p. 47(locality).

? Syntermes bolivianus SNYDER, 1924, p. 23 (sol-dier).

Syntermes peruanus SNYDER, 1924, p. 22 (imago,soldier).

IMAGO: Head with scattered long bristles.Fontanelle oval, about as long as the ocelli,light colored and depressed. Eyes relativelysmall and prominent. Ocelli of medium size,about midway between the eyes and fon-tanelle. Antennae with 20 articles, the thirdlonger than the second, the second aboutequal to the fourth. Pronotum narrower thanthe head, front margin not emarginate, frontlateral angles nearly round, very slightly an-gular, the angle being more obtuse than aright angle, sides curved, hind margin emar-ginate.

Length with wings (Holmgren,1911, p. 548)

Length of head to side base ofmandibles

Width of headDiameter of eyeEye from lower marginLength of ocelliWidth of ocelliOcelli from eyesOcelli from fontanelleLength of fontanelleWidth of fontanelleLength of postclypeusWidth of postclypeusLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wing scale

34.50-40.00 mm.

2.653.840.850.240.380.240.590.590.360.320.531.471.58

3.60- 3.726.102.07

VOL. 83458


COMPARISONS: Syntermes dirus has thepronotum wider than the head, lateral anglesof the pronotum sharper, and eyes smallerwith ocelli farther from the eyes. S. grandishas a longer pronotum and larger eyes, andthe pronotum has sharper lateral angles. S.magnoculus has a larger eye, proportionatelylonger pronotum, and in general is of largersize. S. molestus is smaller with the ocellicloser to the eyes than to the fontanelle. S.obtusus has a larger eye and is generallylarger. S. parallelus is a smaller species withsharper pronotal angles. S. snyderi has thepronotum wider than the head, and the pro-notal angles are much sharper. S. territus hasa smaller eye, proportionately longer pro-notum, and sharper pronotal angles.

SOLDIER: Head, pronotum, and tergiteswith a number of straight bristles. Head withsomewhat convex sides converging slightlytoward the front. Profile of head stronglyconvex. Frontal tube with a very slight anglewhere it joins the vertex, anterior marginhardly elevated above the head, frontal open-ing facing upward at an angle of 45-60° fromthe long axis of the head. Antennae with 20articles, the fourth barely separated from thethird in the two specimens. Lateral angles oflabrum about equal to a right angle or alittle more obtuse, tip sharper than a rightangle. Narrowest portion of postmentumabout two-thirds the width of widest portion.Left mandible with first marginal tooth ex-tending somewhat beyond the continuationof the apical cutting edge (less so than inS. lighti and more so than in S. calvus), apicalcutting edge sigmoid, the notch forming anangle of about 45°. Right mandible with a dis-tinct marginal tooth which is not projectingvery far from the apical cutting edge (about0.13 mm.). Pronotum with front marginslightly or distinctly emarginate, lateralspines of medium size and about equal in sizeto those of the mesonota and metanota.

Length of head to side base ofmandibles

Width of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotum

5.06-5.24 mm.4.94-5.00

3.233.05

1 .23-1 .290.88

1.40-1.533.17-3.232.56-2.71

Width of metanotum 3.29-3.54Length of hind tibia 4.69

COMPARISONS: No type specimens of S.bolivianus were found in the Holmgren col-lection in Stockholm. I cannot determineS. bolivianus from the meager descriptionsand am tentatively placing this species insynonymy with S. peruanus. Holmgren sep-arates S. bolivianus from S. peruanus by theweaker dentation in the mandibles of S.bolivianus.

Syntermes dirus (specimen determined byHagen) is larger, with more bristles on thehead; angle of junction between the frontaltube and the vertex is much sharper, but themandibles are very similar in type. I thinkthat S. dirus is the closest relative to S.peruanus. S. chaquimayensis (cotype) islarge, with more abundant bristles on thehead, proportionately larger and thickerthoracic spines, and with a proportionatelyshorter marginal tooth on the right mandible.S. silvestrii (autotype from San Bernardino,Paraguay) has proportionately shorter mar-ginal tooth on left mandible, is a larger spe-cies, and has no angle between the vertex andfrontal tube.

DISTRIBUTION (FIG. 1): Mojos (14°36' S., 68°56'W.), Bolivia, 2 imagoes (white), 1 soldier andworkers (cotypes), det. N. Holmgren as S. dirus,coll. N. Holmgren; 1 soldier (cotype) labeled asabove.The species is also recorded from Juan del Oro,

Tambopata Valley (13°22'-14°58' S., 69°00'-69°55' XW.), Peru (type locality); Santa Cruz(17037' S., 63016' W.), Bolivia; and Pata (14°34'S., 68°43' W.), Caupolican, Bolivia.

REMARKS: Holmgren originally recordedhis S. dirus from the above localities. S.boliviaknus was later recorded only from southBolivia. He described S. bolivianus from sol-diers only, so it is presumed that the vialcontaining imagoes, a soldier, workers, andnymphs in my collection from Mojos, Bolivia,are cotypes of S. peruanus.

Syntermes dirus (Burmeister)Klug is usually considered as the author of

this species, but no published descriptionfrom his pen has been found, and conse-quently the species should be assigned toBurmeister. It would be necessary to examineall the original material in various collectionsbefore the following synonymy could be veri-

4591945


fied. In some cases I have been able to exam-ine some old collections including specimensdetermined by Burmeister, Perty, Rambur,and Hagen, but I am by no means sure ofother portions of the synonymy. Hagen exam-ined many of these old determined specimens,and he was a careful and meticulous tax-onomist who laid an excellent foundation forour present knowledge of the classification oftermites. However, he made a few errors, andall his material needs further examinationand comparison in the light of recent investi-gations. Latreille's type of Termes spinosumis reported present in the Selys Longchampscollection in Brussels. If this finally proves tobelong to S. dirus, the name of the speciesshould revert to the earlier name. However, Ihesitate to use Latreille's name without re-examining his specimens.

In the following synonymy an asterisk de-notes specimens reexamined by me:

? Termes spinosum LATREILLE, 1805a, p. 70(soldier).

? Termes spinosum LATREILLE, 1805b, p. 63(soldier).Termes fatale PERTY, 1830, p. 127 (imago),

pl. 25, fig. 8 (imago).* Termes flavicolle PERTY, 1830, p. 128 (imago),

pl. 25, fig. 11 (imago), fig. 13 (soldier).* Termes dirus BURMEISTER, 1839, p. 766

(imago).

IMAGOLength with wingsLength without wingsLength of head with mandiblesLength of head to side base of mandiblesWidth of headDiameter of eyeEye from lower marginLength of ocelliWidth of ocelliOcelli from eyesOcelli from fontanelleLength of fontanelleWidth of fontanelleLength of labrumWidth of labrumLength of postclypeusWidth of postclypeusLength of left mandibleLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wing scaleLength of anterior wing from sutureWidth of anterior wing

Termes from Brazil, WESTWOOD, 1840, p. 12(imago), fig. 58, 1-11 (imago).

Termes obscurum BLANCHARD, 1840, pl. 47,fig. 1.

* Termes costatus RAMBUR, 1842, p. 305(imago).

* Termes dirus RAMBUR, 1842, p. 307 (imago).? Termes cephalotes RAMBUR, 1842, p. 309 (sol-

dier).? Termes dirus WALKER, 1853, p. 510 (imago,

synonymy).? Termes dubius WALKER, 1853, p. 521 (imago).? Termes cephalotes WALKER, 1853, p. 521

(imago).* Termes dirus HAGEN, 1858a, p. 151 (imago,

soldier, worker), pl. 2, figs. 16, 17 (imago), pl. 3,fig. 17 (imago). [Part.]

* Termes dirus HAGEN, 1858b, p. 21 (imago).* Termes dirus HAGEN, 1862, pp. 83-85 (imago,

soldier, worker), pl. 4, figs. 1-8 (imago).Termes spinosus WASMANN, 1897, p. 139 (syn-

onymy).Termes spinosus DESNEUX, 1904, p. 38 (syn-

onymy), pl. 2, fig. 8, 8a, 8b (imago, soldier,worker).

* Syntermes dirus HOLMGREN, 1911, pp. 545-548 (imago, soldier).

Termes spinosus DESNEUX, 1915, p. 8 (synon-ymy).

Syntermes dirus SNYDER, 1924, p. 21 (imago),p. 22 (soldier), p. 24 (imago), p. 27 (soldier fromBrazil only).

No. RANGE3 39.00-43.001 16.501 4.765 2.76- 3.11

17 3.45- 3.888 0.71- 0.744 0.29- 0.355 0.23- 0.354 0.19- 0.236 0.61- 0.716 0.56- 0.745 0.37- 0.535 0.38- 0.532 0.90- 1.212 1.45- 1.563 0.71- 0.793 1.53- 1.621 2.12

11 1.87- 2.0617 3.72- 4.273 5.98- 7.193 2.32- 2.357 28.58-34.228 6.96- 8.08

MEAN40.33 mm.

2.993.700.720.310.300.210.660.660.440.441.051.500.751.58

1.923.996.462.34

31.867.58

460 VOL. 83


* Syntermes hageni SNYDER, 1924, p. 22 (sol-dier), p. 28 (soldier), pl. 4, fig. 21 (soldier).

Termes spinosus HANDLIRSCH, 1930, p. 847, fig.894 (soldier).

IMAGO (FIGS. 2, 3): Head yellow brown;postclypeus and labrum slightly lighter thanthe head; pronotum slightly lighter than thehead, sometimes with dark markings; wingsbrown. Head with a few scattered bristles;pronotum with bristles on the anterior andposterior margins and a few on other parts;wing scales with medium long bristles. Fon-tanelle larger than the ocelli, round, white.Eyes relatively small, prominent. Ocelli ofmedium size, about midway between the eyesand fontanelle. Antennae with 20 articles, thethird as long or longer than the second, thesecond longer than the fourth. Pronotumwider than the head, somewhat saddle-shapedwith a vertical front surface in the middlecovered with short bristles, anterior anglesabout equal to, or slightly sharper than, aright angle, sides curved and hind marginconspicuously emarginate.COMPARISONS: The closest imago is that of

S. snyderi which is somewhat larger (pro-notum width 4.27-4.77 mm.).

SOLDIER: Head, base of mandibles, post-mentum, and pronotum covered rather abun-dantly with bristles. Head large, sides fairlystraight, converging a little toward the front.Frontal tube raised above the rest of thehead, the anterior and posterior portionsabout equally long, the opening nearly verti-cal or pointed upward at an angle of about450 to the axis of the head, an angle betweenthe vertex and frontal tube in profile. An-tennae with 20 articles. Lateral angles of thelabrum more obtuse than a right angle.

SOLDIERLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of left mandible from tip to point of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

Narrowest width of postmentum about three-fourths of the widest width. Left mandiblewith prominent marginal tooth a little widerthan the apical cutting edge. Apical cuttingedge distinctly sigmoid. Angle between toothand apical cutting edge about 30-45'. Rightmandible with a relatively small prominentmarginal tooth, about 0.15-0.18 mm. widefrom the base of the notch. Front margin ofpronotum emarginate or fairly evenly curved.Lateral spines of medium length. Lateralspines of mesonota and metanota about equalin size to the pronotal spines.COMPARISONS: In no case am I sure that

the soldiers examined were found with re-productives. An imago and a soldier fromBurmeister's material in the Museum of Com-parative Zoology have the same label andmay have been collected from the same col-ony. There is some variation among the exam-ined soldiers in the size of the pronotal spinesand in the teeth of the mandibles, which mayprove to be characteristic of subspecies, butfor the present I am not attempting to sub-divide the species on the basis of the fewspecimens available for study.

Syntermes peruanus cotypes are smaller;the frontal tube is not so large, particularlythe anterior projecting portion; and the headhas fewer bristles.

Syntermes silvestrii has shorter marginalteeth in both mandibles, a shorter frontaltube, and shorter thoracic spines.

Syntermes calvus has more strongly con-verging sides toward the front, somewhatfewer bristles, and with proportionatelylonger apical teeth in both mandibles.These four species seem to be quite closely

related.

No. RANGE3 8.05-8.659 5.43-6.1614 5.00-6.6111 3.06-3.797 1.04-1.354 3.03-3.4110 1.29-1.538 0.82-1.008 1.58-1.83

13 3.54-4.1711 2.93-3.4911 3.66-4.397 4.51-5.79

MEAN8.32 mm.5.755.483.341.153.251.400.911.723.973.104.095.30

1945 461


DISTRIBUTION (FIG. 1): Rio (probably Rio deJaneiro, 23018' S., 430 18' W.), Brazil (type local-ity), 1 imago (cotype), det. H. Burmeister andH. Hagen as T. dirus, coll. v. Olfers, No. 2763,Berlin Mus.; 4 imagoes (cotypes and probably theoriginal Klug material), No. 3628 from the BerlinMus.; 1 female (cotype), det. H. Burmeister andH. Hagen as T. dirus, M.C.Z.; 2 soldiers (probablyfrom the type series, although Burmeister did notdescribe the soldier caste), det. H. Burmeister andH. Hagen as T. dirus, M.C.Z.

Brazil (no other locality mentioned), 1 imago,det. M. Perty as T. flavicolle, redet. N. Holmgrenas S. dirus, Munich Mus.; 1 male, 1 female, 1 sol-dier, det. H. Hagen as T. dirus, coll. Winthem,type No. 204 (not sure these are Burmeistertypes), M.C.Z.; 1 female (possibly the type of T.costatus, although Rambur, 1842, records the spe-cies from Cayenne), det. P. Rambur as T. costa-tus, det. H. Hagen as T. dirus, coll. P. Rambur,type No. 419, M.C.Z.; 1 male, det. P. Rambur asT. dirus, coll. P. Rambur, M.C.Z.Unknown locality, 1 imago (metatype), det.

N. Holmgren, coll. Tschudi, 1863; 1 soldier, det.N. Holmgren, det. H. Hagen as T. dirus, labeled"Hagen revid. 1856"; 1 soldier, det. H. Hagen asT. dirus, from Berlin Mus.; 2 soldiers, Perty'sColl., Munich Mus. One of these seems to be atypical dirus, but the other is larger with a largertooth in the right mandible and may possibly notbe conspecific, although I have included it inthe table of measurements.Corcovado (not located on map), Rio, Brazil,

1 soldier, det. H. Hagen as T. dirus, M.C.Z.Entre Rios (22008' S., 43015' W.), Rio de

Janeiro, Brazil, 1 soldier, det. T. E. Snyder (1924,p. 28) as S. hageni, coll. J. D. Haseman, 4.VI.1908,Carnegie Mus. Acc. No. 3564, A.M.N.H.Mang (possibly Manga, 14045' S., 43056' W.),

central Brazil, 1 imago (metatype), det. N. Holm-gren, coll. R. Fischer, 3.XI.1913, from KaiserWilhelm Inst., Berlin-Dahlem.Barro Alto (not located on map but possibly

Barra, 15040' S., 46°35' W., or Barra, 17005' S.,

SOLDIERLength of head to side base of mandibleWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

44040' W.), Minas Geraes, Brazil, 1 soldier,coll. Jos6 Blaser, 31.XI., M.C.Z.

Iguassu (probably Ituassu, 13024' S., 43010'W.), Bahia, Brazil, 3 soldiers, coll. A. Roman,29.VII.1924 (1), 31.VII.1924 (2).Manaos (03006' S., 60°00' W.), Amazonas,

Brazil, 1 imago (metatype), coll. A. Roman,9.XI.1923, flying; 1 imago, coll. T. C. Fletcher,received 19.I.1863, from M.C.Z.

Surinam, 1 male, det. H. Hagen as T. dirus,coll. Thorey, M.C.Z. The wings seem a little morehyaline than the other S. dirus on pins. Thisspecimen could conceivably be the imago of S.calvus, but I am classifying it as S. dirus for thepresent.Walker (1853, p. 510) records the species

from Para (01035' S., 48°30' W.), Rio de Janeiro(23018' S., 43018' W.), and the interior of Brazil.Hagen (1858a, p. 151) records the species fromMinas Geraes, Caigara (03014' S., 64049' W.), Rio,Congonhas (20054' S., 45047' W.), and Lagoa Santa(19041' S., 43057' W.), Brazil, and also fromGuiana. Desneux (1915, p. 8) records Termesspinosus from Tijucca (probably Tijucas, 27015'S., 48°38' W.), Santa Catharina, Brazil.

Syntermes silvestrii HolmgrenTermes dirus SILVESTRI, 1901, p. 4.Termes dirus SILVESTRI, 1903, p. 48 (soldier,

workers), p. 115 (biology), p1. 2, figs. 75-78 (sol-dier, workers), pl. 6, fig. 298 (biology).

Syntermes silvestrii HOLMGREN, 1911, pp. 546,548 (soldier).

Syntermes silvestrii HOLMGREN, 1912, pp. 19, 47,text fig. 4 (soldier mandibles), pl. 2, fig. 14(soldier).

Syntermes silvestrii SNYDER, 1924, p. 23 (sol-dier).

SOLDIER: Head, base of mandibles, la-brum, postmentum, thorax, and abdomenwith numerous bristles. Sides of head con-verging only a little toward the front, fairlystraight. Frontal tube very short, without an

No. RANGE6 5.85-6.496 5.45-5.955 3.17-3.662 3.17-3.474 1.35-1.614 0.85-1.006 1.60-1.926 3.66-4.166 2.76-3.676 3.47-4.336 5.06-5.75

MEAN6.08 mm.5.773.423.321.490.941.733.953.133.795.29

462 VOL. 83


angle between the vertex and the frontal tubeanterior margin hardly elevated above thehead, opening upward at about an angle of450. Antennae with 20 to 21 articles. Lateralangles of labrum blunter than a right angle.Narrowest portion of postmentum abouttwo-thirds the width of the widest portion.Left mandible with marginal tooth extendingabout to the continuation of the apical cut-ting edge which is sigmoid, angle of notchless than 45°. Right mandible with blunt andrather short marginal tooth. Pronotum withfront margin slightly or not emarginate.Thoracic spines medium size.

COMPARISONS: Originally described bySilvestri as Termes dirus, this species was rec-ognized and described by Holmgren. S. sil-vestrii is closest to S. dirus, but there is noangle between the vertex and the frontal tu-be.S. peruanus is smaller and has a slight anglebetween the frontal tube and the vertex inprofile, and the left mandible has a propor-tionately longer marginal tooth. S. territusis smaller, the sides of the head are more

parallel, the mesonotal and metanotal spinesare shorter, the dentation of the mandibles issimilar, but the tips of the mandibles aremore curved and the head is a little lesshairy.

DISTRIBUTION (FIG. 1): Coxipo (15°30' S.,56°00' W.), Cuyaba, Matto Grosso, Brazil (typelocality), 1 soldier (cotype), det. F. Silvestri asT. dirus, redet. N. Holmgren as S. silvestrii, coll.F. Silvestri, 6.IX.1900, Silvestri Coll., Portici.San Bernardino (25°19' S., 57°16' W.), Para-

guay, 3 soldiers (autotypes), det. N. Holmgren,coll. K. Fiebrig.

Paraguay, 1 soldier (autotype), det. N. Holm-gren, coll. K. Fiebrig, No. 26.

S. Sofia (not located on map), Argentina, 1 sol-dier, coll. Reimoser, 20.V.1908, from Vienna Mus.

Syntermes calvus, new speciesSOLDIER (FIG. 12): Antennae not much

darker than head. Head, thoracic nota, andtergites with a few bristles. Head with sidesfairly straight, converging toward the front,hind margin slightly angular. Frontal tubeshort, in a small depression in the head, open-

FIG. 12. Dorsal view of head and pronotum and ventral view of head of soldier of Syntermes calvus,new species, holotype.

4631945


ing forward or a little upward with an angleat the junction of the frontal tube and vertex.Antennae with 19 to 20 articles, subapical ar-ticles about 0.15 mm. wide. Tip of labrumsharper than a right angle, side angles greaterthan a right angle. Postmentum curved incross section, narrowest portion more thantwo-thirds the width of the widest portion.Mandibles proportionately long, somewhatcurved and hooked a little near the tip. Leftmandible with a notch just in front of the firstmarginal tooth, the angle of the notch about450, the tip of the tooth sharper than a rightangle, another smaller and wider notch about

SOLDIERTotal lengthLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of left mandible from tip to point of toothLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumLength of hind tibia

halfway between the first marginal tooth andthe base and a fairly sharp tooth-like projec-tion just anterior to the molar plate. Rightmandible with anterior angle of marginaltooth slightly greater than a right angle, thetip of the tooth about equal to a right angleor less. Front margin of pronotum slightlyemarginate, hind margin nearly straight incenter with evenly curved angles. Lateralspines not very long, tip 0.76 mm. from theangle at the junction with the anterior lobe.Lateral spines of mesonota and metanota nar-rower than the pronotal spines.LARGE WORKER: Color a little lighter than

the soldier. Head, pronotum, and tergiteswith a few bristles. Fontanelle white, smallerthan antennae sockets. Antennae with 20 ar-ticles. Labrum hatchet-shaped with curvedmargins and rounded angles. Front marginof pronotum very slightly emarginate, anelongated pit in the center of the pronotum.Thoracic nota with sharp lateral spines.Width of head 3.60 mm.Width of pronotum 2.74

COMPARISONS: The soldier is closer to S.dirus than to any other described species.S. dirus has more bristles on the head, thesides are not so strongly converging, and thefrontal tube is more prominent and is notsunk in as much of a depression. The mandi-bles and notal spines are quite similar.

DISTRIBUTION (FIG. 1): Kartabo (06'23' N.,58°42'W.), British Guiana (type locality), 2 sol-diers (holotype and paratype), coll. A. Emerson,19.IV.1924, colony under log; 1 soldier, workers(paratype), 29.III.1924, No. 24.83, under log onground; 1 soldier (paratype), 13.VII.1924, foundin jaws of worker ant, Termitopone commutata

No. RANGE4 17.50-19.004 7.99- 8.274 5.24- 5.615 4.94- 5.305 3.25- 3.405 1.15- 1.355 3.05- 3.295 1.41- 1.535 0.98- 1.075 1.64- 1.775 3.63- 3.844 5.00- 5.24

MEAN18.25 mm.8.115.415.103.301.273.151.451.021.723.775.15

(Roger), No. 24.235; 1 soldier (paratype),6.IX.1920, found in jaws of worker T. commutata,No. 316, paratype No. 25708, M.C.Z.; 1 soldier(paratype), workers, 18.VI.1924, No. 24.178, fromstomach contents of the lizard, Ameiva ameiva.

Syntermes ternitus EmersonSyntermes territus EMERSON, in Snyder, 1924,

p. 22 (imago), p. 23 (soldier).Syntermes territus EMERSON, 1925, pp. 310, 359

(imago, soldier), text fig. 45 (imago, soldier).Syntermes territus WHEELER, 1936, pp. 173, 174,

176, 189 (biology).IMAGO: Head dark brown with yellow

brown streaks, postclypeus and labrum yel-low brown, pronotum similar color to head,portions of wing scales darker than generalpronotal color, wings brown, tergites darkbrown, sternites yellow brown. Head andpronotum with a few scattered bristles. Wingscaies with a number of bristles. Fontanelleyellowish, depressed, round, smaller thanantennal sockets. Eyes small, less than halftheir diameter from the lower margin. Ocelliof medium size, midway between the eyes and

464 VOL. 83


fontanelle. Antennae with 19 to 20 articles.Pronotum about as wide as, or slightly nar-rower or wider than, the head, with slightlyemarginate front margin, lateral anglesbluntly pointed forming an angle equal to,or greater than, a right angle, sides rounded,posterior margin emarginate.

IMAGOLength with wingsLength of head with mandiblesLength of head to side base of mandiblesWidth of headDiameter of eyeEye from lower marginLength of ocelliWidth of ocelliOcelli from eyesOcelli from fontanelleLength of fontanelleWidth of fontanelleWidth of labrumLength of postclypeusWidth of postclypeusLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wing from sutureWidth of anterior wing

COMPARISONS: Syntermes snyderi is larger,with proportionately wider pronotum. S.peruanus is larger and has a larger eye. S.parallelus is close, the only difference beingthe slightly smaller size and a pronotum1.38-1.41 mm. long. S. obtusus is larger andhas hyaline wings. S. molestus has hyalinewings, the eye is a little larger, and the ocelliare farther from the fontanelle than from theeyes. S. magnoculus is larger, with a muchlarger eye. S. grandis is larger and has hyaline

SOLDIERLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

wings. S. dirus is larger, with proportionatelywider pronotum.

SOLDIER: Head, base of mandibles, post-mentum, and pronotum moderately coveredwith numerous bristles. Head with sidesslightly convex or straight, converging only alittle toward the front or parallel. Frontal

No. RANGE4 32.00-35.003 3.75- 3.901 2.56

17 3.05- 3.507 0.53- 0.683 0.21- 0.291 0.291 0.192 0.53- 0.622 0.53- 0.622 0.292 0.29- 0.351 1.291 0.471 1.49

31 1.47- 1.6717 2.99- 3.602 4.49- 4.762 24.44-25.383 6.00- 6.29

MEAN33.00 mm.3.81

3.260.620.24

0.580.58

0.32

1.543.264.6324.916.16

tube forming a flat angle with the vertex,opening nearly vertical to the long axis of thehead. Postmentum with narrowest portionabout three-fifths the width of the widestportion, narrowest portion with straight orslightly concave sides. Tip of labrum forminga right angle, and lateral angles somewhatless than right angles. Antennae with 19articles, penultimate article 0.14 by 0.32 mm.Left mandible with first marginal tooth pro-jecting only slightly beyond the apical cutting

No. RANGE1 8.274 5.18-6.2213 4.21-5.245 2.80-3.191 3.541 1.261 0.761 1.52

13 2.99-3.681 2.861 3.473 4.15-4.37

MEAN- mm.5.794.813.00

3.44

4.22

4651945


edge which is sigmoid, angle of notch about45°. Right mandible with small, blunt margi-nal tooth. Thoracic nota with small but sharplateral spines. Front margin of pronotumemarginate.

COMPARISONS: Syntermes dirus has margi-nal teeth in the mandibles more prominent.S. parallelus has more slender mandiblesand is smaller.

DISTRIBUTION (FIG. 1): Kartabo (06023' N.,58°42' W.), British Guiana (type locality), 1imago (morphotype), soldiers (holotype and para-types), coll. A. Emerson, 1919; king, queen, sol-diers (topotypes), 5.IV.1924, No. 24.93a; 2 deal-ates, 2.VI.1924; 1 imago, 2.VI.1924, No. 24.162c;imagoes, 2.VI.1924, No. 24.162d; soldiers (para-types), 23.VIII.1920, No. 257; 1 soldier (para-type), 18.X.1920, No. 428, M.C.Z., from colonyraided by Termitopone commutata recorded byWheeler, 1936. Alates with workers and soldierswere brought in by Indians on 29.IV.1919 and4.V.1919. Flying alates were collected 14.VI.1919and 2.VI.1924.

Cururuzinho, Rio Autaz (03025' S., 58°50'-60005' W.), Amazonas, Brazil, 1 soldier, workers,coll. A. Roman, 22.X.1914, with T. commutata inthe same vial. This soldier has somewhat longerspines on the thorax than the British Guiana typesand topotypes but is otherwise very close.

Syntermes parallelus SilvestriSyntermes parallelus SILVESTRI, 1923, p. 318

(imago, soldier, worker), pl. 15, figs. 1-10 (imago,soldier, worker).

Syntermes parallelus SNYDER, 1924, p. 22(imago), p. 23 (soldier).Syntermes colombianus SNYDER, 1924, pp. 23, 29

(soldier), pl. 4, fig. 24 (soldier).Syntermes parallelus EMERSON, 1925, pp. 310,

361 (imago, soldier), text fig. 46 (imago, soldier).

IMAGO: Head and pronotum dark brown.Wings brown. Ocelli proportionately small,twice their length removed from the eyes.Fontanelle yellowish, depressed and round.Antennae with 19 articles. Pronotum nearlyas wide as the head with blunt lateral anglesmore than a right angle, side margin roundedand hind margin emarginate.

Length with wingsLength of head to side base of

mandiblesWidth of headDiameter of eye

Length of ocelli

RANGE27.50-29.00 mm.

2.292.89- 3.150.57- 0.630.27- 0.29

Ocelli from fontanelleOcelli from eyesLength of fontanelleWidth of fontanelleLength of left mandibleLength of pronotumWidth of pronotumLength of hind tibia (after Sil-

vestri)Length of anterior wingWidth of anterior wing

0.56- 0.570.53- 0.560.29- 0.350.23- 0.33

1.681.38- 1.412.74- 3.00

4.0020.88

5.42- 5.80

COMPARISONS: The imago is very close toS. territus but is a little smaller.

SOLDIER: Head brownish yellow, coveredwith short hairs, sides straight and almostparallel. Frontal tube short. Antennae with19 articles. Left mandible with first marginaltooth about even with an extension of theapical cutting edge. Right mandible with asmall obtuse tooth about two-fifths the lengthof the mandible from the base. Notal spinesrather short.

Length of head with mandiblesLength of head to side base of

mandiblesWidth of headLength of left mandibleLength of pronotumWidth of pronotumLength of hind tibia

RANGE6.34-6.80 mm.

4.08-4.282.97-3.402.44-2.621.10-1.222.19-2.362.93-3.50

COMPARISONS: There seems to be littledoubt that S. colombianus Snyder is a syno-nym of S. parallelus. I can detect no differ-ences of specific importance. The cotype ofS. paralklus has rather concave sides of thehead, while all other specimens from BritishGuiana have slightly convex sides as in thetype specimen of S. colombianus. The meas-urements all seem within the expected limitsof variation. The drawing of S. colombianusin Snyder, 1924, is quite accurate except thatthe angles between the lateral and centralpoints of the labrum are not so sharp in thespecimen as in the drawing and the right man-dible is not so curved at the tip as in thedrawing.

This species differs from most of the otherspecies in its small size and comparativelyparallel straight sides of the head of the sol-dier. The closest relatives are probably S.dirus or S. territus, from which it differs mark-edly.

466 VOL. 83


DISTRIBUTION (FIG. 1): Canister Falls (04'54'N., 58030' W.), Demerara River, British Guiana(type locality), 1 soldier (cotype), 1 worker, det.F. Silvestri, coll. A. A. Abraham (Cattle Trail Sur-vey).

Kartabo (06023' N., 58042' W.), British Guiana,1 imago, coll. A. Emerson, 2.VI.1924, No. 24.162c,flying; parts of 6 imagoes, 9 soldiers, workers,6.VI.1924, coll. A. Emerson, from the stomach of atoad, Bufo typhonius, No. 24.169.Oronoque River (02042' N., 57°18'-57°26' W.),

British Guiana, 1 soldier, coll. N. A. Weber,21.VII.1936, No. 583, in the jaws of a ponerineant, Pachycondyla crassinoda (Latreille).

Colombia, 1 soldier (holotype of S. colombianus),det. H. A. Hagen as T. dirus, det. T. E. Snyder asS. colombianus, coll. Winthem, type No. 14511,M.C.Z.

BIOLOGICAL NOTES: The ant Pachycon-dyla crassinoda was found by Neal Webercarrying a soldier of S. parallelus. This antwas observed by me at Kartabo collectingworkers of a species of Termes (formerlycalled Mirotermes) from a disturbed nest. Theants took many termites and stuffed them intheir mandibles. On April 11, 1924 (No.24.229), I also found this ant carrying aroach in its mandibles. Thus there is evi-dence that this species of ant does not confineits diet to termites.

Syntermes molestus (Burmeister)Termes mokestus BURMEISTER, 1839, p. 766

(imago).IMAGO

Length with wingsLength of head with mandiblesLength of head to side base of mandiblesWidth of headDiameter of eyeEye from lower marginLength of ocelliWidth of ocelliOcelli from eyesOcelli from fontanelleLength of fontanelleWidth of fontanelleLength of postclypeusWidth of postclypeusWidth of labrumLength of pronotumWidth of pronotumLength of hind tibiaLength of anterior wing scaleLength of anterior wing from sutureWidth of anterior wing

Termes molestus WALKER, 1853, p. 513 (imago).Termes molestus HAGEN, 1858a, p. 159, pl. 3,

fig. 19 (imago).Termes molestus HAGEN, 1858b, p. 22 (imago).Termes molestus SILVESTRI, 1901, p. 4 (locality).Termes molestus SILVESTRI, 1903, pp. 51, 116

(imago, soldier, workers, biology), text fig. 10(wings), p1. 2, figs. 84-85 (imago), figs. 86-88 (sol-dier), p1. 6, fig. 299 (food).

Syntermes molestus HOLMGREN, 1911, pp. 547,548 (imago, soldier).Syntermes brasiliensis HOLMGREN, 1911, pp. 547,

548 (soldier).Syntermes molestus SNYDER, 1924, pp. 22, 24

(imago), p. 23 (soldier).Syntermes braziliensis SNYDER, 1924, p. 23 (sol-

dier).Syntermes MS species of Emerson, SNYDER,

1924, p. 22 (imago).Syntermes braziliensis BEQUAERT, 1925, p. 294

(biology).Syntermes molestus SNYDER, 1926, pp. 14, 15,

pl. 2, fig. 5 (soldier).Syntermes braziliensis SNYDER, 1926, p. 13, pl. 2,

fig. 9 (soldier).Syntermes brasiliensis WHEELER, 1936, pp. 174,

176 (biology).

IMAGO: Head, pronotum, and tergites yel-low brown. Labrum, postclypeus, and stern-ites more yellowish. Wings hyaline and trans-parent except for yellow brown costal borderand radial veins. Head, pronotum, and wingscales covered sparsely with fairly longstraight bristles. Ocelli above the level of the

No. RANGE6 26.00-33.002 3.41- 3.665 1.94- 2.449 2.82- 3.35

11 0.71- 0.888 0.10- 0.238 0.29- 0.358 0.21- 0.268 0.35- 0.487 0.47- 0.597 0.18- 0.477 0.15- 0.421 0.501 1.231 1.158 1.26- 1.508 2.41- 3.036 3.66- 4.471 1.697 21.19-27.077 5.51- 6.96

MEAN29.92 mm.3.542.122.990.750.150.330.230.420.540.310.27

1.392.724.18

25.136.29

1945 467


top of the head in profile. Fontanelle yellow-ish white to brownish yellow, about the samelength as the ocelli, round and depressed orwith a convex plate. Eyes relatively of me-dium size and comparatively close to lowermargin of head. Ocelli of medium size, closerto the eyes than to the fontanelle, abouttwice the width of the ocelli from the eyes.Antennae with 20 articles, the third longerthan the second, the second longer than thefourth. Pronotum a little narrower than thehead, front margin entire, anterior anglesrounded, with a median anterior lobe, andsides slightly curved and converging towardthe hind margin which is emarginate in themiddle. Side posterior angles of mesonotumsharp or rounded and of metanotum rounded.

COMPARISONS: These imagoes differ fromeach other somewhat in size and in possessingeither a fontanelle depression or a fontanelleconvex plate. The variation in the fontanellemay occur within the same vial as in the caseof imagoes from Ciudad Bolivar, Venezuela.I am unable to discover consistent differenceswhich would require subdivision into two ormore species. The species was described fromthe imago alone from Bahia. Soldiers fromBahia in my collection conform closely to thedescription of S. brasiliensis which I considera synonym.

SOLDIERLength of head with mandiblesLength of head to side base of mandiblesWidth of headLength of left mandibleLength of postmentumWidest width of postmentumNarrowest width of postmentumLength of pronotumWidth of pronotumWidth of mesonotumWidth of metanotumLength of hind tibia

SOLDIER: Head, postmentum, thoracicnota, and abdomen with a number ofstraight bristles. Head converging stronglyor slightly toward the front, the sides fairlystraight or slightly concave in the middle.Frontal tube extremely short, hardly morethan a tiny ridge around the opening of thefrontal gland. Narrowest portion of the post-

mentum about two-thirds to three-fourthsthe width of the widest portion. Antennaewith 19 or 20 articles, penultimate article0.11-0.15 mm. wide and 0.23-0.35 mm. long.All points of labrum less than a right angleor lateral points more than right angle. Man-dibles long or fairly long, somewhat hookedto strongly hooked at the tip, in some casesthe tip pointing at right angles to the axisof the mandible. Left mandible with a smallnotch anterior to the first marginal toothforming about a 22-45° angle, and notch pos-terior to the first marginal tooth forming aright angle or slightly greater than a rightangle. First marginal tooth projecting to aline continuous with the apical cutting edge.Right mandible with marginal tooth barelyvisible. Pronotum with a comparatively largefrontal lobe, front margin slightly emarginate.Lateral angles of nota not prolonged intospines, but usually angular. Mesonotumsometimes with rounded sides.

COMPARISONS: The variation of the sol-diers in this species is quite great, particu-larly in the length of the curved hooks in themandibles in different localities. It may wellbe that more extensive collections will indi-cate subspecific populations or even specificdifferences. However, the gradations in thespecimens before me are so slight from one

No.488746668766

RANGE4.88-6.343.06-4.392 .42-3 .721.76-2.702.13-2.970.71-0.850.53-0.620.88-1.171.39-2.021.24-2.031.65-2.412.65-3.72

MEAN5.61 mm.3.572.872.252.670.780.551.021.641.632.093.07

colony to another that I feel it is best atpresent to treat them all as a single species.The characters are consistent within coloniesand in large series, although different col-onies from the same locality are not always aperfect match.The cotype soldiers of S. brasiliensis

Holmgren seem closely related to the Bom

468 VOL. 83

EMERSON: THE NEOTROPICAL GENUS SYNTERMES4

Fim, Bahia, specimens of S. mokstus in mycollection, and I regard these forms as vari-ants of the same species, although this hy-pothesis may be disproved as larger series areobtained. Specimens from Villavicencio, Co-lombia, seem to be almost identical with thecotypes of S. brasiliensis; with the Bom Fim,Bahia, specimens; with the Vista Alegre, RioBranco, Brazil, specimens; with the Tumu-pasa, Bolivia, specimens; and with the Akuri-man, Venezuela, specimens. The Villavicen-do specimens differ from those from Para-catu, Minas Geraes, in having a shorter an-terior portion of the postmentum in front ofthe widest part and in having the notch infront of the first marginal tooth of the leftmandible forming an angle of 45-60°. Alsothe tip of the labrum of the Villavicencio formis wider, the point forming a right angle orslightly less. Otherwise large series from thesetwo colonies match quite well. Soldiers fromIndependencia, Parahyba, Brazil, are a littlesmaller than the Villavicencio specimens andare without concave sides, the frontal tube isa little more prominent, but the mandiblesare similar. The labrum of the Independenciasoldier has the middle point narrower at thebase and is similar in proportions but smallerthan the Paracatu soldiers. A single soldierfrom Cuyab., Brazil, has slightly convexsides of the head and is similar in shape to thesoldiers from Independencia, but the mandi-bles have longer hooks. Soldiers from Mte.Sociedad, Paraguay, and Guarany, SaoPaulo, are very close to Barro Alto, MinasGeraes, specimens but are much smaller thanthe Villavicencio soldiers, the sides of thehead are only slightly converging and aresomewhat convex, the mandibles have com-paratively short hooks, and the labrum is sim-ilar to that of the Independencia soldiers.Soldiers from Corumbi, Matto Grosso, aresmaller than soldiers from Independencia, thetips of the mandibles are proportionately lesscurved, and the side angles of the metanotumare less sharp. The Corumbf specimens arevery close to those from Mte. Sociedad. Thesesmaller soldiers from southern localities withstraighter mandibles seem to agree withHolmgren's description of S. molestus, whilethe larger soldiers with more curved man-dibles from the region of the type locality ofS. molestus agree with his description of S.

brasiliensis. If the species is ultimately di-vided into subspecies, S. brasiliensis wouldstill seem to be a synonym of the typical sub-species.

DISTRIBUTION (FIG. 1): Bahia (13010' S., 38050'W.), Brazil (type locality), 1 imago (cotype), det.H. Burmeister and H. A. Hagen as T. molestus,coll. Gomez, No. 2765, Berlin Mus.; 1 male (topo-type), det. H. Holmgren, coll. Fruhstorfer.Bom Fim (10025' S., 40°10' W.), Bahia, Brazil,

1 soldier, coll. J. D. Haseman, 20.XI.1907, Car-negie Mus. Acc. No. 3441, A.M.N.H.Amazonas, Brazil, I male, det. H. A. Hagen, as

T. molestus, coll. Saunders, XII, M.C.Z.Unknown locality, 1 soldier (probably cotype of

S. brasiliensis), det. N. Holmgren as "Syntermesn. sp.," from Holmgren's Coll.

Vista Alegre (01035' N., 61009' W.), Rio Branco,Brazil, soldiers, workers, det. T. E. Snyder as S.brasiliensis, coll. J. Bequaert, 6.IX.1924, "forag-ing in daytime-savannah."

Paracatu (17005' S., 46055' W.), Minas Geraes,Brazil, 1 female, 1 soldier, det. R. G. Temple asS. brasiliensis, coll. T. Ivanauskas, 1931, from theBrit. Mus.

Barro Alto (not located on map), Minas Geraes,Brazil, 1 soldier, coll. Jos6 Blaser, XI.1931, M.C.Z.

Lassance (17043' S., 44035' W.), Minas Geraes,Brazil, 4 imagoes, coll. J. C. Bradley and Harris,12.XI.1919, 13.XI.1919, 9-19.XI.1919.

Independencia (06048' S., 35033' W.), Para-hyba, Brazil, soldiers, workers, det. T. E. Snyder,coll. W. M. Mann.Corumb& (19007' S., 570501 W.), Matto Grosso,

Brazil, 1 imago, 5 soldiers, det. F. Silvestri as T.molestus, coll. F. Silvestri, 27.IX.1900, SilvestriColl., Portici.Cuyabi (13035' S., 56010' W.), Brazil, 1 soldier,

workers, det. F. Silvestri as T. molestus, coll.F. Silvestri, VII-IX.1900. The species is also re-corded in the literature from Coxipo (15030' S.,56°00' W.), Cuyaba, Brazil.Guarany (21025' S., 48006' W.), Sao Paulo,

Brazil, 9 soldiers, workers, coll. Jayme V. Pin-heiro, 11.1943 (dealates in same vial belong to anundetermined species of Syntermes).

Mte. Sociedad (not located on map), Paraguay,soldiers, workers, coll. Ternetz, 1895, from BaselMus.Tumupasa (13059' S., 67048' W.), Bolivia, sol-

diers, workers, det. T. E. Snyder as S. brasiliensis,coll. W. M. Mann, XII.1921.

Villavicencio (04017' N., 73043' W.), Meta,Colombia, king, queen, soldiers, workers, coll.C. H. Seevers, 16.VII.1938, No. 14; soldiers,workers, coll. C. H. Seevers, 24.VII.1938, No. 28.

1945 469


Ciudad Bolivar (0807' N., 63056' W.), Vene-zuela, imagoes, coll. P. J. Anduze, 22.VII.1940.Akuriman (not located on map but in upper

Caroni River basin [05o-07' N., 61°30'-63°20'W.], Bolivar), Venezuela, soldiers, workers (2vials), coll. P. J. Anduze, XI.1940.

Borgmeier (1930) records the species from Cam-pinas (16045' S., 49015' W.), Goyaz, Brazil. Thespecimens were determined by T. E. Snyder as S.brasiliensis.

OTHER RECORDS OF SYNTERMESSyntermes sp. Reichensperger (1936, p. 230),

from Santa Cruz (17021' S., 48018' W.), Goyaz,Brazil.

Syntermes sp. Dealates, Guarany (21025' S.,48006' W.), Sao Paulo, Brazil, coll. Jayme V.Pinheiro, IL.1943 (soldiers and workers in the samevial were determined as S. molestus, but these

dealates do not conform to the description of thisspecies).

Syntermes sp. Wings, workers, Urucum (19013'S., 570351 W.), Matto Grosso, Brazil, 19.VIII.1926,coll. K. P. Schmidt, "large brown earthen nest,"Emerson Coll.

Syntermes sp. near grandis. Imagoes, Lassance(170431 S., 57035' W.), Minas Geraes, Brazil,12.XI.1919, coll. Harris, Emerson Coll.

Syntermes sp. A single pinned soldier in poorcondition in the M.C.Z. labeled "Maruriu. April."(Locality not located on map.) This species isprobably new but is in such poor condition thatit should not be named. The hairs on the head arethick and curly as in S. aculeosus, but the spines,although large, are not so large as in S. aculeosus,and the mandibles are quite different.

Syntermes sp. Workers only, Villavicencio,Meta, Colombia, coll. C. H. Seevers.

Syntermes sp. Workers only, Madeira Mamore(090431 S., 65022' W.), Peru.

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1945 471

l

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