cretaceous insects fromlabrador new genus termite

CRETACEOUS INSECTS FROM LABRADOR3. A NEW GENUS AND SPECIES OF TERMITE.

(ISOPTERA: HODOTERMITIDAE)

BY ALFRED E. EIMERSONDepartment of Zoology, The University of Chicago., and Depart-

ment of Entomology, American Mus.eum of Natural History.

INTRODUCTIONThe origin and dispersal o.f numerous higher taxa of termites by

mid-Cretaceous times have been postulated from present geographicaldistributions and the existence of fairly advanced types in early tomiddle Tertiary deposits (Emerson, I955, NISS). With the excep-tion of a few extinct genera, the Oligocene termite fauna of Balticamber includes several genera that are characteristic of present warmtemperate climates of Europe and Asia. A recently discovered trop-ical termite fauna in Mexican amber o.f Oligocene-Miocene age isgenerically closely similar to the living fauna of the same region(Emerson, NISS). Heretofore, the ea.rliest known fossil termiteswere found in Eocene deposits of North America, Australia, andEurope. The few specimens of Eocene age belong to the Mastoter-mitidae (Emerson, I965) and Kalotermitidae (Emerson, MSS),but early Oligocene fossils from several deposits including Balticamber belong to. the Hodotermitidae, Mastotermitidae, Kaloter-mitidae, a.nd Rhinotermitidae. The author and others, have. consideredthe fauna of Baltic amber to be of upper Eocene age (Emerson,I955), but the general concensus now places this fauna in the louverOligocene. The lack of the most advanced Termitidae from earlyTertiary strata is probably due to the general scarcity or absence ofthis family in temperate climates in the northern hemisphere, and itis predicted that abundant fossils from tropical regions will provethe existence of numerous advanced genera of Termitidae in Creta-ceous times. Indeed the tropical fauna, of Mexican amber on theOligocene-iiocene border proves the presence of living genera ofvarious families that include the Termitidae at that time (Emer-son, MSS).The discovery of a fossil termite wing from late Early Cretaceous

or early Late Cretaceous (Cenomanian) associated with a, warmtemperate flora in Labrador is a considerable extension of accuratelydetermined specimens of the order Isoptera backward in geological

This investigation was supported by the National Science Foundation(Grant No. G-25146).

276

1967] Emerson Cretaceous Insects 277

time. Earlier reports o.f fossil termites in Paleozoic deposits are allincorrect because of the erroneous assignment of specimens of otherorders to the Isoptera. However, the author is of the opinion, ad-mittedly speculative, that the Isoptera may have originated in earlyMesozoic or even late Paleozoic times.The single forewing here described is without doubt a termite.

Only the Isoptera have a humeral suture at the base o the wing thatenables the wings to be easily shed at the end o the colonizingflight. The deciduous wings are associated with the strictly sociallife of these insects, and this Cretaceous fossil is the earliest knownsocial insect, inasmuch as the Labrador horizon is of an ea.rlier agethan the recently described fossil ants from Cretaceous amber of NewJersey.

Although the order may be determined accurately, a single fore-wing, although remarkably complete and well preserved, makes theassignment to other higher taxa somewhat more precarious. In thefir.st place, the variation in the wing venation of species and generao.f termites, particularly primitive termites, is great compared to therelative invariability characteristic o many other orders o insects.Wing venation alone cannot be used as con(idently for taxonomic andphylo.genetic interpretation of termites as it may be or other orders.Slight differences of wing venation, particularly of fossils, have oftenbeen interpreted as indications of species or generic differences thatare obviously false when the variability of comparable living speciesis investigated. In the second place, fossils are always fragmentaryand show a paucity of characters as compared to abundant series ofall castes co,mmo.nly collected from the living fauna.

In spite of the lack of other characters of the head, thorax, andabdomen, the author feels justified in assigning this forewing to thefamily Ho.do.termitidae, the subfamily Cretatermitinae, new sub-family, the genus Cretatermes, new genus, and the species C. car-penteri, new species. These taxa and their phylogenetic relationshipsmust be considered as tentative working hypotheses until more. abun-dant and extensive material is available for study and comparison.

FAMILY HODOTERMITIDAE DESNEUXThe present status of the Hodotermitidae is discussed by Emerson

(in press). The distribution of fossil and living genera is interpretedby Emerson (I955, pp. 473, 482, 484, 49o, 492, 498, 499, 5o2, 5o6,507, 509, 516, 517).The author has little doubt that Cretatermes carpenteri agrees with

the wing venation of the Hodotermitidae. The wide area occupied

278 Psyche [December

by the radial sector (Rs) in the o.uter fourth of the forewing withthe innermost branch joined to the inner margin near the wing tip(figs. I, 2) is found only in some of the subfamilies of the Hodoter-mitidae. The relatively short cubitus (Cu) and the reticulations inthe wing membrane between the veins are also in conformity withsome subfamilies of the. Hodotermitidae.

These combined characters ,of the frewing eliminate all otherknown families of termites. It should also, be pointed out that thiswing venation is associated with characteristic dentition of the imagomandibles, the lack o.f ocelli, more than two, articles, in the cerci, to-g’ether with numerous other characters, in more completely preservedfossils, and living species of the. Hodotermitidae. It is postulated(Emerson, in press) that an unknown ancestral group possessed theocelli of the Mastotermitidae, Rhinotermitidae, and Termitidae, thedentition of the imago mandible .clo.se to that of Archotermopsis a,ndthe cockroaches, and a still more ancestral group probably possessedthe anal lobe. of the hind wing of the Mastotermitidae. Withoutpreserved characters of the head, pronotum, legs, and abdomen ofCretatermes carpenteri, all that can be said at present is that thisspecies most probably belongs to the Hodotermitidae. It is antici-pated that thi,s taxonomic assignment will be verified with the dis-covery of more specimens from the same depo.sit or o.f related speciesfro,m other localities and strata.

Subfamily Cretaterrnitinae, new subfamilyIn the early stages of this investigation, the author was loath to

erect a new monotypic subfamily on the basis of a single fossil fo.re-wing, and he still has misgivings. However, he thinks that thetentative assignment to the new su’bfamily Cretatermitinae is thebest nomenclatural and bibliographic symbol to. handle the compara-tive data available. No other known subfamily o.f the Hodotermitidaehas the combination of diagnostic characters illustra.ted in figs. *, 2,and 3. These include: (I) the evenly curved humeral suture; (2)the grea.ter length of the wing from the inner end of the humeralsuture than from the costal end of the humeral suture; (3) the smallsize of the forewing; (4) the relatively wide wing in proportionto its length; (5) the gradual increase in the width of the areaoccupied by the radial sector (Rs) from the base to. the apical fourtho.f the wing; (6) the branching of the media (M) in the basalfourth of the wing; (7) the position o.f the media (M) about halfway between the radial sector (Rs) and the cubitus (Cu) and(8) the short cubitus (Cu) not reaching beyond the basal half of


the inner margin of the wing. Some of these characters are sharedwith some genera of other subfamilies, but the combination does notfit any presently known .subfamily of the Hodotermitidae.The Termopsinae (Emerson, 1933, figs. 13, 30, 40) has a pro-

portionately narrower area occupied by Rs in the. outer portion ofthe wing; the length of the wing is considerably larger in all kno.wnspecies; the length of the forewing from the costal end of the humeralsuture is about the same as or is shorter than the length o.f the fore-wing from the inner end of the humeral suture; Rs is roughlyparallel to the costal margin; M is closer to Cu than to. Rs in themiddle of the wing; and the reticulatio.ns enclose smaller and morenumerous spaces of the wing membrane.The Hodotermitinae (Emerson, in press) has well defined in-

ferio.r branches of Rs reaching the inner margin of the wing behindthe tip; the. humeral suture of the forewing is somewhat sinuate; thelength of the forewing from the costal end of the humeral sutureis about equal to the length from the inner end of the humeral suture.The chara.cteristic inferior branches of Rs and the width of the areaoccupied by Rs and its branches in the outer portion of the wingallow rather easy identification of this subfamily from the forewingsalone. It should be pointed out that there is only a little differencein the angles of the branches of Rs in designating a branch as in-ferior or superior, and in some individual cases the distinction issomewhat arbitrary. From an evolutionary point of view, the trans-ition from superior to inferior branches wo.uld seem to be only aslight step, and some other subfamilies such as the Termopsinae,Poro.termitinae, and Stolotermitinae have individuals that may haveone or more inferior branches of Rs.

The. subfamily Porotermitinae vith a single genus including threespecies distributed in temperate areas of southern Australia, SouthAfrica, and Chili, has a fairly straight humeral suture of the forewingwith a short curve near the costal end; all known species of Poro-termes are much larger than Cretatermes; the. wing is proportionatelylonger compared to its width in most species of Poroternes exceptP. adamsoni in which the proportions are similar to those in Creta-termes; Rs is closely parallel to the costal margin throughout thelength of the wing; the superior branches of Rs are thick and strongand are not secondarily branched as in the’ Cretatermitinae; the mainstem of Rs is as strong as the superior branches, but from o-3 in-ferior branches are as weak as M and reach the inner border of thewing belo.w the tip; M is about half way between Rs and Cu in the

28o Psyche December

middle o1: the wing or may be closer to Cu than to Rs; and Cu has1:rom 5-12 primary branches, the longest extending 1:rom about ha.11:to, 1:our-fi1:ths the length o1: the wing ro.m the base. The reticulatio.nso1: the wing membrane do not differ diagnostically 1:rom Cretater-mitinae, and the inner margin o1: the wing o1: the Porotermitinae islonger than the costal margin rom the suture in both sub1:amilies.

Sc, R1, R2, Ra, or R2+a vary in the Porotermitinae and maynot be diagnostically different rom this obscure region o1: thewing ia the single ossil specimen o1: Cretatermitinae. In the Poro-termitinae there may be as 1:ew as two. branches o1: t’he radius emergingfrom the suture, or there may be a 1:ull complement o.1: Sc, R1, R2, R,and Rs at the suture, thus illustrating the. extreme variability o1:these veins in some species and genera ot: primitive termites.

In an occasional rather exceptional wing, M may have. branchesthat invade the normal position o1: Rs, and in such cases the superiorbranches o.1: M may be as strong as those o1: Rs and may also reachthe costal margin o the outer portion o.1: the 1:orewing. Again, thisillustrates the. great variability o the venation o.1: these primitive ter-mites and the ditkiculty o.1: making definitive comparisons. In spiteo1: this degree o variation, however, the general pattern is sufficientlycorrelated with other .characters to indicate valid taxa rom the spe-cies to the 1:amily.The Stolotermitinae is a sub1:amily based upon a single, genus with

six described species, one 1:rom temperate South A1:rica, oneNew Zealand, one 1:rom Tasmania, one 1:rom temperate southeasternAustralia, and two (possibly only one) rom tropical eastern Aus-tralia.

The dentition o1: the mandibles o1: Stolotermes is o1: a very primitivetype with .strictly homolo.gous teeth to those o1: cockroaches (Ahmad,I95o), and is only slightly advanced in a 1:ew particulars 1:rom thato1: Archotermopsis (Yermopsinae) and Ulmeriella (Hodotermitinae).T’he reticulations o1: the 1:orewing are not diagnostically different(rom Cretatermes; the humeral suture o1: the 1:orewing is 1:airlystraight or convexly curved in contrast to the concave curve o1: thehumeral suture o.1: Cretatermes ((ig. 3); the humeral suture slantsbackward at a sharp angle 1:rom the costal to. the inner margin; theveins 1:ro,m Rs to Cu are bent or curved do,wn toward the innerborder near the suture (Hill, 1942, fig. ); the costal margin ofthe 1:orewing is much shorter than the inner margin from the humeralsuture to the tip; the, shape, of the wing may be similar to. that ofCretatermes or a little longer compared to width; the linear length

1967] Emerson Cretaceous Insects 28I

o the o.rewing i,s longer in a,ll species o Stolotermes except S.africanus Emerson in which it may be slightly shorter than in Creta-ternes carpenteri; Sc is short; R is present and is occasionallybranched; R2 and R are absent; Rs is gradually wider in the areaoccupied from the humeral suture to the outer portion o the wing,but is not so proportionately wide as in Cretatertnes (figs. I, 2);the superior branches o Rs are closer to each other with less widthbetween them than in Cretatermes; Rs may have rom 0-2 weakerinerior branches reaching the inner margin near the tip; M is long,not quite reaching the tip o the wing but with the. lower branchesreaching the inner margin about 1/3-3/5 he length o the. innermargin from the, suture; the irst branch o M may be close to thesuture as in Cretatermes; M is usually weaker than Rs and is usuallycloser to Cu than to Rs; Cu is usually short with 3-7 branchesclose together, the longest branch reaching t’he inner margin rom1/6 to, nearly 1/2 the. length o the wing (differs among species.).The arolium is absent in the. Stolotermitinae, Porotermitina.e, andsome Hodotermitinae and is present in the Termopsinae and someHodotermitinae.The major distinctions between the. orewings o Stolotermes and

Cretatermes are the shape and position o, the humeral suture and thewidth o the area occupied by Rs and its superior branches about 3/4the length o. the wing rom the. base.

It is the opinion o. the author that the orewing o Cretatermesis closer in more characters to the orewing o various species o.Stolotermes than to other subfamilies, and resembles Porotermes nextmost closely. However, the resemblances and differences o a singleorewing are insufticie.nt to, warrant any firm conclusion concerningthe phylogenetic relationship between the Cretatermitinae and eitherthe Stolotermitinae or the Porotermitinae other than to state thatCretatermes is advanced in its small size, the shape and position othe humeral suture, and in the relatively large spaces between thereticular thickenings o the wing membrane.

PHYLOGENETIC RELATIONS BETWEEN THE SUBFAMILIESOF THE H’ODOTERMITIDAE

There would seem to be little doubt that the more primitive genera(Termopsis, Archotermopsis) o the Termopsinae are the mostprimitive members o the Hodotermitidae, and in several particularsare the most primitive known termites (Emerson, I933).

Recently (Emerson, in press) a redescription o the ossil genusUlmeriella o the Hodotermitinae indicates that it is rela.ted to but


is slightly more advanced than Archotermopsis. The various char-acters o.f Ulneriella seem to be derived directly from primitiveTermopsinae.

The Stolotermitinae has a very primitive let mandible of theimago (Ahmad, 95o; Emerson, 942, 947), but is derivative innumerou.s or’her characters, particularly in the sharply slantinghumeral suture of the forewing. L. R. Cleveland discovered thatStolotermes is the only genus of termites that shares a genus of wood-ingesting intestinal flagellates with the wood-feeding cockroach,Cryptocercus (see Emerson, 96, for a discussion of the. evolutionaryimplications of this discovery). The. Stolotermitinae could havebranched off from primitive Termopsinae or possibly from primitiveHodotermitinae, but coulc[ not have the Porotermitinae in its an-cestry as that subfamily is now conceived, primarily because the Poro-termitinae has a much reduced second ma.rginal tooth of the leftmandible. The Porotermitinae could not be derived from the Stolo-termitinae because of the advanced type of humeral suture, in thelatter subfamily, but it could have branched off from the less. primitiveTermo.psinae or possibly from the Hodotermitinae although onlybefore the specialized harvester behavior had evolved.

Because of the less advanced type of humeral suture, the fore-wing of the Cretatermitinae could not have arisen from the Stolo-termitinae which it resembles, in some characters. The wing of theCretatermitinae might be related remotely to the Porotermitinae,but the data a.re insufficient to enable us clearly to. separate homolo-gous from analogous characters, and parallel reduction from homolo-gous reduction. With more characters of the species available, itmay be possible to find consistent patterns of phylogenetic impor-tance, but at present it is necessary to state that several alternativehypotheses of phylogenetic derivation could account for the meagerfacts. The small size of Cretatermes is reasonably certain to bederivative, but the small size of Stolotermes africanus could havearisen independently, each from more primitive groups o.f la.rger size.With our present information, the Hodotermitidae consist o.f fossil

and living relicts that are spotty and often discontinuous in theirdistribution. The Porotermitinae and Stolotermitinae, neither rep-resented by known fossils, are. fo.und only in the southern hemisphere,and the genera are probably Mesozoic in o.rigin and dispersal beca.useof their existence on different continental masses long iso.lated fromone another. The living and fossil Termopsinae are all found inthe north warm temperate zones with a gradation from the most

28 Psyche [December

primitive in Eura.sia and the derivative genera in Tertiary and presentNorth America. The only known fossil of the subfamily Hodoter-mitinae is Ulmeriella (Emerson, in press) ound in Tertiary tem-perate deposits of Europe, Asia, and North America, while the livinggenera (3) o the Hodotermitinae are the desert and steppe har-vester termites of temperate and tropical A.frica and Asia. T’hesedistributions in time and space would seem to indicate a very ancientorigin with considerable extinction of intergrades between the relicttypes of fossils and present survivors (Emerson, 1955, in press.).

Cretatermes now provides direct evidence of the existence o.f afairly advanced subfamily of the otherwise primitive family of Hodo-termitidae in mid-Cretaceous times, and indicates that the origin ofthis family of termites and the order Isoptera is much earlier thanthe Labrador deposits. The author suggests that the origin o.f theHodotermitidae may be Triassic and that the origin of the Isopte.ramay be late Paleozoic.

Genus (]retatermes, new genusTYPE SPECIES: Cretatermes carpenteri, new species.With only a single forewing representing a monotypic subfamily,

and with no. data on the varia.tion within the species or the commoncharacteristics o.f several species within the genus, the descriptionof the genus and the subfamily coincides for the present.From the knowledge o.f species and generic differences in the other

subfamilies of Hodotermitidae, however, it is possible to. indicatethe characters that might be co.nsidered of subfamily rank a.nd thosethat might be considered of generic rank.The forewings of various genera of Termopsinae show differences

in the shape of the humeral suture (Emerson, I933) with very fewdifferences in the wing venation beyond the suture.

Generic characters of the Hodotermitinae are found in the man-dibles and the tibial spines, while the wings show little distinction.The subfamilies o.f Porotermitinae and Stolotermitinae, each

known rom a single genus, show little variation of the humeralsuture within each genus, and also show numerous generic char-acters of the wing venation. Smaller degrees o.f difference in thewing venation indicate species distinctions.The species within the genera can ften be distinguished by the

shape of the wing, the relation of length to breadth, small differencesin the shape of the humeral suture, and the mean sizes of male andfemale forewings and other structures. On occasion, specific varia-tion in wing venation can be detected, although many wings must

1967-1 Emerson Cretaceous Insects :z85

Fig. 2. Drawing of visible veins and reticulations of holotype f.orewingof Cretatermes carpenteri, new genus, new species.Rs radial sector. M media. Cu Cubitus.

be examined because of the large degree of variation between individ-uals and even between the right and left wing of the same individual.For adequate species distinction, a set .of head, tho.racal, and abdom-inal characters that are correlated with the wing venation shouldbe described, but unfortunately this is not yet possible for Cretatermescarpenteri. The following description, therefore, contains charactersof various systematic categories from individual variation to subfam-ily distinctions, and we must await the discovery of more materialand a more thorough study of variation at va.rious related taxonomiclevels before we can make taxonomic distinctions and find commonqualities of each taxon. This does not mean that the author agreeswith those who would eliminate the co.ncepts of species and highertaxonomic categories. On the, contrary, he finds the taxa with theirshared homologous characters have distinctions that rest upon awealth of demonstrated biological principles resulting from investiga-tions in a variety of di.sciplines. Taxonomic and phylogenetic ordercan be discovered from abundant material subjected to a variety ofobservational and experimental techniques together with comparativecross-disciplinary studies (Emerson, 96, in print, MSS).

Cretatermes carpenteri, new speciesRIGHT FOREWING: General shape as in figs. , 2. The costal

margin of the basal o/3 of the wing is slightly convex; tip of wingrounded and about half way between the costal and inner margins;inner margin evenly convex, more curved than the costal margin.Numerous thickenings or reticulatio.ns (archidictyon) of the wing

membrane occur between the branches of Rs, between Rs and M,between M and Cu, and between the branches of both M and Cu.The enclosed spaces within the reticulum are large compared tothose of the Termopsinae and Hodotermitinae.

286 Psyche IDecember

Scale broken and the shape is not preserved. The costal marginthat oten shows important taxonomic distinction is broken off. Someveins seem to cross the humeral suture rom the scale to the wingbeyond the suture, but some strongly marked structures do not seemto fit the anatomy o the o,rewing scale and may be other structuresseparate rom the wing that impinge upon it in the preserved scale,o,r else are ma,rks in the rock.

Humeral suture (fig. 3) seems to be. reasonably well preserved.It is evenly curved and slants backward toward its junction with theinner margin. The inner portion o, the suture is not as clear as thecostal end, but the extension o the clear edges of the suture andthe inner margin seem to indicate the true sha,pe and position o theinner portion o, the suture. The suture is not angulate in the regiono Rs. The inner margin o the wing is somewhat concave whereit joins the suture.

Fig. 3. Drawing of base. of holotype forewing of Gretatermes carpenteri,new genus, new species, showing visible veins emerging t?rom thehumeral suture.

The veins that emerge from the suture at the costal margin (fig. 3)are somewhat obscure and the wing in this region is either brokenor folded. Sc is not clearly visible although it is probably present inwell preserved wings. What seems to, be the costal margin at thesuture, may possibly be R1 with the actual costal margin either brokenor folded under. There are indications of veins between the costaledge and Rs, and it is possible that R1 and R2+3 are present. These


veins vary considerably in ,some genera of Hodotermitidae. A veinjoins the costal margin about 1,22 mm. from the suture but it isnot clear whether this is the fir,st superior branch of Rs or the endof one of the other branches, of the radius. Clear superior branchesof Rs occur about 2.63 mm., 3.88 mm., and 5.08 mm. from thesuture, and secondary and tertiary branches occur in the apicalfourth of the wing. The main stem of Rs ends well below the tipof the wing. In summary, seven and possibly eight branches of Rsreach the costal margin and tip of the wing. The width of theradial sector (Rs) and its branches in the, outer fourth of the wing(figs. I, 2) and t,he gradual divergence of the main stem o,f theradial sector from the costal margin are interpreted as generic andsubfamily characters. The branches of Rs are more defined and arenarrower than those of M and Cu. M branches into an upper andlower branch about I.oI mm. from the suture, somewhat closer tothe suture than in most termites, but the taxonomic significancecannot be interpreted without many more specimens. Five branchesof M reach the lower margin in the apical half of the wing, but thischaracter is almost surely variable within the species. The generalposition of M about half way between Rs and Cu is probably ofgeneric importance and might even be of species importance. Cu iscomparatively short, its longest branch ending close to the middle ofthe inner margin. The length of Cu is probably of species or genericimportance. Cu and M emerge from the suture in a fan-like spreadof veins, in this particular strikingly different from the sharplycurved veins of 8tolotermes.

MEASUREMENTS OF THE FOREWING OF

CRETATERMES CARPENTERI NEW SPECIES

Length of suture from costal to, inner margin about 0.72 mm.Length of wing from costal end of suture 7.23 mm.Length of wing from inner end of ,suture 7.42 mm.Width o.f wing 2.23 mm.Widest width of radial sector 0.93 mm.Width between Rs and M near middle of wing 0.39 mm.Width between upper and lower branches of M in

middle of wing o.37 mm.Widest width between M and Cu about 1/3 length

of wing from suture o.44 mm.

SPECIMEN: Holotype, riffht forewing and counterpart impressions,o.f Cenomanian age, at the border between Lower and UpperCretaceous, in a bed of hard ferruginous argillite associated with


rubble ores at the Redmond No. deposit near Schefferville (lat-itude 5405o N., longitude 67000 W.) in the Knob Lake district,Labrador; collected by Professor Erling Dorf. The holotype isNo.. 87270, Invertebrate Paleontology collections, Princeton Uni-versity. From associated fossil plants, Dorf (1959) suggests thatthe climate at the time of preservation was warm temperate.

ACKNOWLEDGEMENTS

The author is indebted to Frank M. Carpenter of the BiologicalLabora.tories, Harvard University, for making the specimen avail-able to me and for ,his numerous critical comments through manyyears on various aspects o.f insect paleontology. The author alsowishes to thank his wife, Eleanor Fish Emerson, for her valued as-si,stance in the preparation and editing of the manuscript.

SUM:MARY

The earliest known specimen of a termite (Isoptera), that is alsothe earliest known tossil of a strictly social insect, is described andtgured trom a to.rewing trom mid-Cretaceous deposits near Schef-ferville, Labrador. The forewing is classified as Cretatermes car-penteri, new genus, new species, and included in the subtamilyCre.tatermitinae, new subfamily of the family Hodotermitidae. Sys-tematic and phylogenetic interpretations and compari.sons are included.

BIBLIOGRAPHY

AHM,AD, M.1950. The phylogeny of termite genera based on imago-worker mandi-

bles. Bull. Amer. Mus. Nat. Hist., 95: 37-86.DORF, ERLING

1959. Cretaceous flora from beds associated with iron-ore deposits inthe Labrador trough. Program of Meetings, Geol. Soc. Amer.,1959 33A.

EMERSON, A. E.1933. A revision of the genera of fossil and recent Termopsinae

(Isoptera). Univ. California Publ. Ent., 6" 165-196.1942. The relations of a relict South African termite (Isoptera, Hodo-

termitidae, Stolotermes). Amer. Mus. Novitates, 1187: 1-12.1947. The imago of Stolotermes africanus Emerson. Jour. Ent. Soe.

So. Africa, 9: 127-129.1955. Geographical origins and dispersions of termite genera. Field-

iana: Zool., 37: 465-521.196q. Vestigial characters of termites and processes of regressive evo-

lution. Evolution, 15: 115-131.


1965. A review of the Mastotermitidae (Isoptera), including a newfossil genus from Brazil. Amer. Mus. Novitates, 2236: 1-46.

(In press) A revision of the fossil genus Ulmeriella (Isoptera; Hodo-termitidae; Hodotermitinae). Amer. Mus. Novitates.

(In preparation) MSS on fossil Kalotermitidae, Rhinotermitidae, andTermitidae.

HILL, G. F.1942. Termites (Isoptera) from the Australian region. Melbourne,

Australia, Council for Scientific and Industrial Research, pp. 1-479.

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