AMERICAN MUSEUM

Nov tta tes PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2570 MARCH 14, 1975

ALFRED E. EMERSON AND KUMAR KRISHNA The Termite Family Serritermitidae (Isoptera) ~~~~~s.u.,wsjx-9uw r¶ua-48u .wwIy, yPm r. tw CTt0F:

It AMERICAN MUSEUM

Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2570, pp. 1-31, figs. 1-6, tables 1, 2 March 14, 1975

The Termite Family Serritermitidae (Isoptera)1

ALFRED E. EMERSON2 AND KUMAR KRISHNA3

ABSTRACT The monotypic termite family Serritermitidae ogy, nests, food, and the coevolution of termites Holmgren and Serritermes serrifer (Hagen and and intestinal zooflagellates are discussed. Also Bates) from Brazil are described and figured, included are hypotheses on the phylogeny of all with synonymies and bibliography. The imago, termite families including Serritermitidae, genetic soldier, worker, and nymph are compared with persistence and change in Isoptera during 200 all other families of Isoptera and related genera. million years, and the natural selection of unitary New characters of Glosso termes Emerson and social populations. Hypotheses on the Mesozoic Macrorhinotermes Holmgren are given. Primitive origin of termite families are presented, based and derivative, homologous and analogous char- upon paleogeography, continental drift, and acters and their use for phylogenetic and evolu- comparative morphology of living and fossil tionary inference are discussed in detail. External genera. anatomy, glands, intestines, biogeography, ecol- INTRODUCTION The monotypic termite family Serritermitidae HISTORY includes only a single, small, rare, bizarre species, Serritermes sermfer (Hagen and Bates), known Specimens of soldiers and workers were first from three localities in Brazil. The purpose of the collected from two colonies by Bates at San- present paper is to redescribe this family and tarem on the Amazon River, Para, Brazil, in 1854. species, together with its known nutrition, nests, Bates named the species T. [= Termes] serrifer in and ecology, to compare it with related families his field notes, which were later expanded into and genera, and to draw phylogenetic inferences. his classic book (Bates, 1863, 1892). Hagen

' Study supported by a John Simon Guggenheim Foundation fellowship to Alfred E. Emerson 1926-1927 and a National Science Foundation Grant no. GB-20684 to Kumar Krishna 1970-1973. 2Research Associate, Department of Entomology, the American Museum of Natural History; Professor Emeritus, Department of Biology, The University of Chicago. 3Research Associate, Department of Entomology, the American Museum of Natural History; Professor, Department of Biology, City College, City University of New York.

Copyright i The American Museum of Natural History 1975 ISSN 0003-0082/Price $1.70 2 AMERICAN MUSEUM NOVITATES NO. 2570

(1858a) transferred the species to Calotermes, known species in the monotypic family Serri- described the soldier and nymph, and used termitidae. Bates's name and his notes, translated into German. The mention of the small size of the FAMILY SERRITERMITIDAE HOLMGREN soldier, the presence of the eye, and the name < Family Termitina: Hagen, 1858a, p. 4. denoting serrated mandibles are deemed suffi- < Subfamily Termitinae: Silvestri, 1903, p. 37. cient to warrant junior authorship to Bates. < Subfamily Calotermitinae: Silvestri, 1901, p. 1. Hagen (1858a) provided a fuller description and Desneux, 1904, pp. 9, 11, 24. a figure of the soldier and technically therefore is < Family Mesotermitidae Holmgren, 1909, p. the senior author of the species name, according 100; 1910a, p. 285; 1910b, p. 143; 191 la, pp. to modern rules of nomenclature. Wasmann 13,61; 191 lb, pp. 206, 209. (1897) placed the species in the new subgenus genus Calotermes. Silvestri 129, 254. (1901, 1903) gave full generic status to Serri- < Family Rhinotermitidae: Light, 1934, p. 112. termes and described the imago, soldier, and Grasse, 1949, p. 533. < Family Termitidae: Silvestri, 1901, p. 1. worker of S. serrifer from his own collections Snyder, 1949, pp. 5, 99. Ahmad, 1950, p. 56. from Coxipo, Mato Grosso, adding a few obser- Emerson, 1955, pp. 495, 497, 505, 506, 507, vations on the nest and food. Holmgren (1909, 510, 517. Harris, 1961, pp. 53, 64. Krishna, 191 Oa, 191 Ob, 1911a) recognized the affinity of 1961, p. 332. this species to his newly created but misnamed = Subfamily Serritermitinae Holmgren, 191 Oa, p. Mesotermitidae, a family that in large part forms 285; 1910b, p. 143; 1911a, pp. 62, 63, 82. the current Rhinotermitidae (Emerson, 1971). Synder, 1949, pp. 99, 100. Grasse, 1949, p. Holmgren gave subfamily status to the Serri- 534. Ahmad, 1950, pp. 56, 75. Emerson, termitinae within the Mesotermitidae. Light 1955, pp. 495, 497, 505, 506, 515, 516, 517. (1934) and Grasse (1949) included Serritermes in Weidner, 1955, pp. 10, 41. Tu, 1955, pp. 33, 34, 35. Harris, 1961, pp. 53, 64; 1962, p. 195. the Rhinotermitidae. Snyder (1949), Ahmad Krishna, 1961, p. 332. Springhetti, 1963, p. (1950), Emerson (1955), and Harris (1961) 213. Sen-Sarma, 1968, p. 7. placed the Serritermitinae in the Termitidae. = Family Serritermitidae: Emerson, 1965, pp. 17, Emerson (1965) raised the subfamily to family 38; 1968b, p. 4; 1971, p. 248. Krishna, 1969, status. Krishna (1969, 1970) described key char- pp. 3, 4; 1970, pp. 127, 130, 131, 137, 139, acters of the Serritermitidae. Araujo (1970, 140. Weesner, 1969a, p. 28. Araujo, 1970, pp. 1972) recorded its geography and ecology. 527, 531, 533, 545; 1972, pp. 67-70. Wilson, The confusion in the taxonomy of Serritermes 1971, p. 105. serrifer for more than a century was due to Type Genus. Serritermes Wasmann (Holm- changing theories of taxa and relationships gren, 1910a, p. 285). No other genus is included within Isoptera, to nomenclatural errors, and to a in the family. lack of thorough examination and comparison of Diagnostic Characters. The imago, worker, the few specimens available. The species pos- and nymph differ from all other termite families sesses some comparatively primitive characters by the proportionally elongated sharp apical and some strikingly derivative ones and has no tooth of each mandible separated from a single very close relative that might indicate gradations marginal tooth near the ridged molar plate by a in its phylogeny. widely curved inner margin (fig. 2A). There is no exact definition of the limits of a The soldier has long, relatively straight apical subfamily or a family; the category is determined portions of both mandibles, each with numerous by the grouped relationships of the included proportionally coarse, large serrations on the genera and the extent of the gap between inner apical margins behind the tips (figs. 4, 5). roughly equivalent higher systematic categories The pronotum is proportionally narrow, elon- (see Chatterjee and Thakur, 1964). We consider gated, and both front and hind margins are the gap sufficiently wide to include the single conspicuously bilobed, each with a relatively 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 3 deeply curved concave emargination in the peus slightly lighter than head; pronotum same middle (fig. 4). color as head or a little lighter; wing hyaline except for opaque white costal margin and radial GENUS SERRITERMES WASMANN sector (Rs). Head, postclypeus, labrum, and pronotum

FIG. 1. Imago of Serritermes serrifer (Hagen and Bates) from Coxipo, Mato Grosso, Brazil, Silvestri Collection, Portici, Italy. Head, pronotum, and hind mar- gins of mesonotum and metanotum from above; head and pronotum from side. Drawn by A. E. Emerson. margin and radial sector (Rs), straight from Rs to heavy and parallel to costal margin almost to tip inner margin. Wing membranes (fig. 3) with of wing. Media (M) weak, in one wing almost irregular vertical and slanting chitinizations, giv- disappearing at base, with about seven to nine ing a reticulated appearance to whole wing branches; in another wing lacking as a distinct between Rs and inner margin except at base near vein and possibly coalesced with the cubitus suture and a few cross reticulations between (Cu). Cu weak with eight to nine branches and costal margin and Rs in apical fourth. Forewing extending half length of wing. Hind wing scale with subcosta (Sc) and firsc radius (RO); second smaller than forewing scale, not extending nearly plus third radius (R2+3) absent. Radial sector (Rs) to margin of metanotum; humeral suture straight 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 5

A

FIG. 2. Mandibles of worker of Serritermes serrifer (Hagen and Bates) from Coxipo, Mato Grosso, Brazil, collected and determined by F. Silvestri. A. Left and right mandible from above. B. Left mandible from below. C. Right mandible from below at angle to show molar area. D. Left mandible molar area from below at angle. Drawn by K. Krishna. 6 AMERICAN MUSEUM NOVITATES NO. 2570

(fig. 3). Hind wing with costal margin and Rs noticeable depression in middle at junction with similar to forewing. M joined to Rs near base, postclypeus. Fontanelle small, oval, about mid- with 10-11 branches reaching tip and apical half way between front of postclypeus and rear of of inner margin; in one hind wing (fig. 3) first head, flat without projection. Frontal gland branch of M sharply turned and joined to Rs in moderately small, oval. Middle longitudinal line about middle of wing. (M is variable and details in rear, Y-suture absent otherwise. Postmentum should not be used for specific or generic diagnosis.) Cu separate at suture with about eight branches, usually not extending beyond middle TABLE 1 of wing (2.17 mm. from costal suture in one Measurements (in Millimeters) of Two Female wing) unless it coalesces with M. Anal vein (A) Imagoes of Semitermes serifer (Hagen and Bates) absent. Spiracles on lower edge of pleural portion of tergites, separated from tergites by light Range bands. Seventh sternite of female with bluntly Length with wings 6.12 pointed hind lobe. Styli absent in both male Length without wings 4.16 (Holmgren, 191 la) and female. Cerci short with Length of head to tip of labrum 0.89, 0.97 two articles. Width of head 0.81, 0.82 Soldier (figs. 4, 5). General size very small Diameter of eye 0.23 (table 2). Length of ocellus 0.08 Head and pronotum straw yellow; labrum Width of ocellus 0.05 mandibles Ocellus from eye 0.02, 0.03 yellowish white; outer portion of Length of antenna 1.39 reddish brown with light bases. Length of postclypeus 0.23 Top of head, postmentum, postclypeus, and Width of postclypeus 0.51 labrum sparsely covered with short or micro- Length of pronotum 0.34, 0.35 scopic hairs without long bristles; tip of labrum Width of pronotum 0.66, 0.68 with denser group of longer hairs; pronotum with Length of hind tibia 0.58, 0.68 numerous short hairs near margins but bare in Length of forewing scale costal margin 0.41 middle. Length of forewing from suture 4.41 Sides of head nearly straight, converging Width of forewing 1.35 toward front; head capsule wider than length of Width of forewing at suture 0.37 head to base of anteclypeus, thick in proportion Length of hind wing scale costal margin 0.28 to length and width; hind margin flatly rounded; Length of hind wing from suture 3.92 Width of hind wing 1.25 profile of short top flat in middle; profile of 0.23 front convex with extremely slight, hardly Width of hind wing at suture

FIG. 3. Left hind wing of Serritermes serrifer (Hagen and Bates) from Coxipo, Mato Grosso, Brazil, collected and determined by F. Silvestri. Drawn by K. Krishna. 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 7

FIG. 4. Soldier of Serritermes serrifer (Hagen and Bates) from Coxipo, Mato Grosso, Brazil, Silvestri Collection, Portici, Italy. Head and pronotum from above; head from below; antenna. Drawn by A. E. Emerson.

(fig. 4) short, wide, its sides somewhat convex in comparatively large, slightly convex, oval, white, middle, sutures slightly constricted near rear, almost continuous with head, slightly larger profile strongly convex, bulging. Vestigial eyes than antennal fossa; irregularities of surface 8 AMERICAN MUSEUM NOVITATES NO. 2570

possibly due to much reduced indistinct facets; one in each mandible with obscure homologies. ocular suture distinct in some lights. Vestigial Left mandible (figs. 2A, 2B, 2D) with one ocellus about two-thirds diameter of eye above relatively large marginal tooth close to molar eye on each side. Antenna with 12 or 13 articles, region, possibly homologous with third marginal third shortest, articles 8 to 11 constricted in tooth of Coptotermes because of its position and middle forming bulbous tips (fig. 4). Both backward pointed direction; slight hump on mandibles with outer margins concave near bases, cutting edge immediately anterior to marginal nearly straight in middle, and convex near apical tooth, possibly greatly reduced second marginal ends; blades about equal width from bases to tooth. (First marginal tooth of more primitive close to tips, with tips hardly hooked, each with Rhinotermitidae proportionally smaller than sec- numerous coarse inner serrations on apical third, ond marginal tooth, and consequently possibly and bases without molar ridges. Left mandible lost before reduction or modification of second with smaller forward pointing serrations reaching and third marginal teeth of Serritermes.) Molar almost to base, slight blunt hump in front of plate of left mandible with eight distinct molar base, small, sharp (about 60°), toothlike projec- ridges and grooves indicating some molar func- tion on basal portion. Right mandible with about tion. Lower edge of molar plate bounded by six smaller serrations posterior to three or four ridge with sharp projecting toothlike prolonga- coarse apical serrations, but not reaching so close tion at anterior edge conforming in position to to base as on left mandible; slight, pointed hump interpretation of homology of third marginal on inner margin near base (possibly vestigial tooth in Coptotermes. (Holmgren [ 19 lla, p. 82, tooth but too obscure to be sure of homology); small sharp toothlike projection on base. Postcly- peus short, with front concave, base straight, obscure shallow longitudinal depression in middle, suture at junction with front of head barely visible, profile flatly convex. Labrum much longer than wide, with sides evenly convex, narrowed anteriorly; tip white, obtusely pointed, narrowly convex, and slightly constricted behind; profile convex. Pronotum (fig. 4) from above elongate, with deeply indented front and hind margins; frontal lobes large and strongly convex, middle longitudinal line faint or obscure, sides moderately straight and converging toward rear; profile with large front lobes raised above straight hind portion. Mesonotum wider than pronotum, hind margin evenly curved. Meta- notum wider than mesonotum, hind margin evenly curved. Coxae without conspicuous sharp ridge, right and left close but bases not touching in middle. Femora normally thicker than tibiae, not markedly swollen. Each tibia with two apical spurs (2/2/2). Tarsi with four articles, fourth longer than all others combined. Styli absent. Cerci short with two articles, first much wider than second. Nymph and Worker. Antenna with 13 articles. Mandibles (fig. 2) most specialized of any known FIG. 5. Soldier mandibles of Serritermes ser- termite; apical portion proportionally prolonged, rifer (Hagen and Bates) from Coxipo, Mato ending in sharp point; marginal teeth reduced to Grosso, Brazil. 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 9 fig. 6] showed ridges on molar surface not shown TABLE 2 by Ahmad [1950, p. 74, fig. 17].) Measurements (in Millimeters) of Two Soldiers of Right mandible of S. serrifer (figs. 2A, 2C) Serritermes sermifer (Hagen and Bates) also with reduction of marginal teeth to one, and reduction of molar ridges to five. Single marginal Range tooth possibly homologous with first marginal Length of head with mandibles 1.54, 1.61 tooth of Coptotermes because faint hump ante- Length of head to middle front of rior to tooth possibly being reduction of subsidi- postclypeus 0.75, 0.77 ary tooth of primitive Hodotermitidae and all Width of head 0.77, 0.79 genera of Rhinotermitidae. (In all other cases of Maximum thickness of head without reduced marginal teeth, first present in right postmentum 0.48, 0.49 mandible.) Sharp toothlike projection above Maximum thickness of head with molar plate possibly extension of upper side of postmentum 0.61 molar plate on different level than marginal Length of postmentum in middle 0.38 tooth (fig. 2A), perhaps adaptation for interlock- Width of postmentum 0.24, 0.26 ing and fitting right mandible with left mandible. Maximum length of eye 0.13 Length of antenna 1.13, 1.22 In anterior intestine, crop not voluminous, Maximum length of fossal suture 0.10 probably not clearly separated from esophagus, Length of left mandible 0.87, 0.94 lightly asymmetrical. Gizzard with very reduced Length of right mandible 0.89 armature, 24 longitudinal folds, slightly promi- Median maximum length of labrum 0.32, 0.36 nent, not chitinized. Middle intestine generalized, Width of labrum 0.28 contact with hind intestine forming transverse Length of pronotum in middle 0.36 circle or ring. Malpighian tubules at least six in Maximum length of pronotum with lobes 0.52, 0.55 number, but probably eight in better preserved Width of pronotum 0.56, 0.58 material, according to Holmgren (1909, pp. 169, Length of hind tibia 0.58, 0.59 170, fig. 53), emerging exactly at junction of middle and hind intestine. Hind intestine preced- structure with normal claws at the tip. Of a total ing enteric valve very short and cylindrical. of two winged imagoes, two soldiers, 13 workers, Enteric valve not much differentiated, with and 10 nymphs with wing pads-one soldier, five posterior border projecting into dilated paunch, workers, and one nymph have coalesced tibiae six lightly chitinized folds corresponding to six and tarsi of the left hind legs; one soldier and six bands of longitudinal muscles, no distinct workers have coalesced right tibiae and tarsi; two rounded swellings in lumen. Paunch very volumi- imagoes, no soldiers, two workers, and nine nous, containing flagellates. Colon and rectum nymphs have normal right and left legs. simple and of normal length.' The senior author first thought this abnormal- Sternal Glands. Noirot (personal commun.) ity might be a Mendelian mutation analogous to stated that the sternal glands are at the anterior some mutations of legs or antennae in Drosophila border of the fifth sternite. (Diptera), but he now interprets the abnormal Seminal Vesicles. Springhetti (1963, p. 213) legs as regenerations of injured legs in early described the seminal vesicle. development, and therefore epigenetic rather Abnormal Legs. Numerous individuals in a than genetic. It is well known that early injured single vial (collected by F. Silvestri, Coxipo, structures may undergo partial regeneration in Cuyaba, 30.VIII.1900) have abnormal hind legs. insects and other Arthropoda. The femur, tibia, and tarsus are more or less Specimens and Collections. One of the origi- coalesced, often with the appearance of a single nal soldiers collected by H. W. Bates and described by Hagen (1858a) as "Calotermes 'Charles Noirot examined the intestines of three serrifer" is labeled "T. Bates TYPE" in workers and fixed the alcoholic specimens in Bouin's serrifer solution. The preservation was inadequate in several the Hagen Collection, Museum of Comparative respects, but Noirot has given us permission to use his Zoology, Harvard University. This soldier was observations and comments. collected at Santarem, Para, Brazil, in 1854. 10 AMERICAN MUSEUM NOVITATES NO. 2570

Hagen (1858a, pp. 72-73) examined five soldiers from this collection are included in the foregoing and three nymphs from two separate colonies. description. Three worker abdomens were sent to According to W. A. Sands (personal commun.), C. Noirot for a study of intestinal structures. specimens collected by Bates (1855, pp. 4551, Araujo (1972) collected two colonies at Cur- 4552) and identified by Hagen were sold to the velo, Mato Grosso, from which he compared British Museum (Natural History), London, Eng- soldiers with the Bates Collection in the British land. Bates sent his specimens to Samuel Stevens, Museum (Natural History). These soldiers should a dealer, auctioneer, and entomologist, with a be labeled "homolectotype determined, com- letter suggesting sale to the British Museum. pared and collected by R. L. Araujo." They were purchased from Stevens and are Geographical Distribution. Araujo (1972) labeled "Santarem on the Amazon, Bates, BM listed all known localities in Brazil (fig. 6) as Accession No. 1865-19." We are here designating follows: Santar6m, Para (type locality), at the as lectotype a soldier of Serritermes serrifer mouth of Rio Tapaj6s, 20 24' 54" S; 540 39' W; (Hagen and Bates) from the British Museum alt. 36 m., collected by H. W. Bates. Coxipo da collection, which has been selected by W. A. Ponte, Mato Grosso, about 5 km. SE of Cuiabai Sands and labeled "lectotype Soldier Serritermes (= Cuyaba), 150 35' 38" S; 560 05' 54" W; alt. serrifer (Hagen and Bates) Coll. H. W. Bates 150 m., collected by F. Silvestri. Curvelo, Minas 19.iv.1854." Therefore, the soldier labeled TYPE Gerais, 1.5 km. N of 18° 45' 40" S; 440 24' 46" at the Museum of Comparative Zoology should W; alt. 633 m., collected by R. L. Araujo. be labeled paralectotype. Hagen (1858a) revised Ecological Distribution. The vegetation in the the termites in the British Museum and retained vicinity of all the localities is described as one soldier in his personal collection now in the tropical savanna forest, campos or cerrado, with Museum of Comparative Zoology. small, scattered, low single trees or with wooded Specimens of imagoes, soldiers, and workers patches dotted with mound nests of Cornitermes, were collected by F. Silvestri, 30.VIII.1900, at and sometimes degraded by cultivation, felling of Coxipo da Ponte, about 5 km. southeast of trees or cattle grazing (Araujo, 1972). Bates Cuiaba, Mato Grosso. Silvestri (1901, 1903) specifically stated that the vicinity of Santar6m identified and described these specimens under on the Amazon River does not have dense high the name Serritermes semfer and deposited them forest (rain forest). The few collections of S. in alcohol in his collection in the Scuola Agraria, serrifer have always been associated with other Portici, Italy, from which he exchanged a few termites of various genera and species, particu- specimens. Silvestri sent specimens to N. Holm- larly mounds of Cornitermes spp. (C. cumulans gren in Stockholm, Sweden, who gave a number determined by Silvestri and Hagen needs reidenti- of termites from around the world to A. E. fication). Termites of different species are com- Emerson in 1927, among them one imago, one monly in close proximity and often in separate soldier, and workers of S. serifer, all formerly galleries within the nests of other species. With dried and shriveled. Emerson compared a soldier few exceptions [i.e., Termes inquilinus Emerson from Coxipo with the pinned paralectotype in the stored humus-like material in arboreal soldier at the Museum of Comparative Zoology, nests of Constrictotermes cavifrons (Holmgren) and found them specifically identical. The single in Guayana], there is seldom any species-specific soldier from Holmgren now in the American nest association. It is doubtful whether Serri- Museum of Natural History labeled homoparalec- termes serrifer is associated wholly with any totype is included in the present article. Draw- other species or genus of termite, although a ings (figs. 1, 3) and descriptions of an imago and small species of this type may find the micro- soldier were made in Portici in 1927 when climatic conditions of termite mounds such as Emerson was studying Silvestri's collection. those of Comitermes attractive for establishing Krishna obtained exchange specimens from the its own colonies. Silvestri Collection in Portici, Italy, and drew the Nests. Meager notes on the nests have been nymph-worker and soldier mandibles (figs. 2, 5), published by Hagen (1858a), Bates (1892), Sil- and one soldier and several nymphs or workers vestri (1903), and Araujo (1972). Serritermes 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 11 occupies a small chamber measuring 3 to 4 ence of particles of mineral matter in the square inches, subdivided into smaller cells, all contents of the intestines examined under polar- distinctly separated from the surrounding galler- ized light. The worker mandibles show the most ies of Cornitermes in mounds also occupied by extreme reduction of marginal teeth and propor- several other species of termites. The mounds, of tional elongation of the apical portion known course, were built by the large workers of among worker termites, but the mandibles pos- Cornitermes. The five colonies so far discovered sess basal molar ridges that are absent in numer- are insufficient to conclude that an inquilinous ous species of the Termitidae that feed on humus association always occurs between Serritermes alone. The intestinal armature described by and Cornitermes, although attention of future Noirot (personal commun.) indicated a reduction collectors should be focused on this possibility, in comparison with a wood-eating genus like and also upon the source of the food of Reticulitermes. William A. Sands (personal com- Serritermes. mun.) suggested that the worker mandibles might Food. Bates (noted in Hagen, 1858a) recorded be adapted to a predatory or seminivorous the nest within carton material. Silvestri (1901, (seed-eating) diet, or just possibly to a diet of 1903) thought the food was vegetable detritus. soft Cornitermes carton. Sands is sure Serri- Noirot (personal commun.) described the pres- termes is not a soil feeder and that the gut is too

FIG. 6. Distribution of Serritermes serrifer (Hagen and Bates) from Brazil. 12 AMERICAN MUSEUM NOVITATES NO. 2570 short for a humivore. Capritermes and related termes is closer to the head pilosity of the more genera are thought to feed on humus and soil, primitive families. Termitogeton has many hairs and in these genera the basal ridges of the and bristles. Parrhinotermes has numerous long mandibles are reduced to three, four, or five in bristles and a few microscopic hairs. Prorhino- both mandibles. It remains an open question termes has a few long hairs or bristles with short what the exact nature of the food ofSerritermes hairs nearly absent on top, and Macrorhino- is, and what the nutritional role of the intestinal termes and Schedorhinotermes have a few short flagellates may be. bristles only, in front of the eyes. The pronotum of Macrorhinotermes has a number of long bris- COMPARISONS OF FAMILIES AN) tles on the margins and a few short bristles on GENERA the disc, whereas the tergites and sternites are Descriptions, figures, and discussions of re- thickly covered with long bristles of similar motely related families and genera are contained length above and below. Acorhinotermes has in the literature cited of the present article; long bristles on the front part of the head and no among the most useful are Silvestri (1909); short hairs. Emerson (1933, 1950, 1965, 1968b); Krishna The wing pilosity of Mastotermes, Archo- (1961); and Chatterjee and Thakur (1964). Char- termopsis, and Zootermopsis is the most primi- acters indicating taxonomic and phylogenetic re- tive, with few hairs on the costal margins and an lations were discussed by Emerson (1971, fig. 1, occasional hair on the strong veins, and without pp. 249-254, 265, 272-273, 277, 287-289, 291, hairs on the weaker veins, inner margins, and 295). We did not examine every species in every membranes. Parastylotermes robustus (Rosen) is genus, so the ranges of variation may be greater equally primitive. Coptotermes, Heterotermes, than those given below. Termitogeton, and other relatively primitive rhinotermitids have numerous hairs on the costal and inner wing margins, veins, and membranes- IMAGO in Heterotermes, varying in abundance from spe- Size. The very small size of Serritermes is de- cies to species. The absence of hairs on the costal rivative compared with the moderate or large size margins, inner margins, membranes, and strong of primitive species of the Mastotermitidae, veins beyond the humeral suture in Stylotermes, Kalotermitidae, Hodotermitidae, Rhinotermiti- Prorhinotermes, Rhinotermes, Dolichorhino- dae, and Termitidae (Emerson, 1933, 1965, termes, and Serritermes is now considered deriva- 1969, 1971). The fossil Reticulitermes minimus tive in contrast to the earlier interpretation by Snyder is about as small as Serritermes. Parrhino- Emerson (1971, pp.252,253, tables 1, 2). termes and Termitogeton are small, but not so Head Capsule. The comparatively wide oval small as Serritermes. In a few cases, notably head of Coptotermes is considered primitive among the Termitidae, there is a rare evolution among the Rhinotermitidae. The slightly nar- toward larger size, but the far more obvious rower and less widely oval head of Serritermes is trend is from large to small. hardly of phylogenetic significance. The wide, Pilosity. Mastotermes and Archotermopsis angular, short and small head of Termitogeton is have short hairs on the head and pronotum. The unique among the Rhinotermitidae. The sides of longer and thicker hairs and bristles of Copto- the head of Prorhinotermes and Parrhinotermes termes have been considered primitive among the are straighter near the eyes. The head of Macro- Rhinotermitidae (Emerson, 1971), but the trend rhinotermes is proportionally wider, the front is toward many short hairs and bristles and the arched in profile from the side and rear, and is trend toward long bristles and few short hairs similar to the profile of Schedorhinotermes ex- may both be derivative. Sequences in allied gen- cept for the clypeus. Parrhinotermes, Macrorhi- era indicate evolutionary trends. Heterotermes notermes, Rhinotermes, Dolichorhinotermes, and may be more primitive with less abundance of Acorhinotermes have several small ridges or hairs and bristles than Coptotermes and more wrinkles behind and at the sides of the fon- abundance than Serritermes, although Serri- tanelle, and a shallow groove extending from the 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 13 fontanelle through the middle of the post- termes is moderately large and flatter than that clypeus. Heterotermes and Reticulitermes have of Coptotermes. The eye of Parrhinotermes is proportionally narrower heads with straighter round, convex, and relatively large. The eye of sides than in Serritermes. Macrorhinotermes is large, oval, and propor- The fontanelle of Coptotermes, Termitogeton, tionally similar to Serritermes although the size Stylotermes, and Prorhinotermes is small, oval, of the head is much larger (width of head with and more primitive than the thicker portions of eyes 2.74 mm.). The proportions and convexity the fontanelle of Serritermes. The fontanelle and of the eye of Rhinotermes (except R. nasutus), associated frontal gland originated with the Rhi- Dolichorhinotermes, and Acorhinotermes are notermitidae and are lacking in the Mastotermiti- similar to Coptotermes. The proportionally dae, Kalotermitidae, and Hodotermitidae. The smaller, triangular, flat eyes of Heterotermes and frontal gland is large and derivative in Prorhino- Reticulitermes are derivative. The trend is from a termes. The position of the fontanelle of Copto- large, primitive eye toward a small, derivative termes, Termitogeton, and Serritermes in back of eye, but there are numerous exceptions. the level of the antennal fossae and ocelli is prim- Ocelli. The Hodotermitidae lack ocelli-a de- itive compared to the forward position in the rivative character compared with their presence more derivative genera of the Rhinotermitinae. in the Blattodea, Mastotermitidae, Kalotermiti- The fontanelle of Parrhinotermes, Macrorhino- dae, Rhinotermitidae, Serritermitidae, and Ter- termes, Rhinotermes, Dolichorhinotermes, and mitidae. Ocellus-like spots in some Hodotermiti- Acorhinotermes is close to the level of the ocelli dae are not near the tips of the branches of the and is surrounded by a shallow depression Y-suture and are probably muscle insertions. The slightly concave in profile. The more complete ocellus is large, distinct, and close to the eye in and distinct the epicranial suture (Y-suture), the Coptotermes, Termitogeton, Prorhinotermes, and more primitive the genera and species. Masto- Parrhinotermes. In Serritermes the position is termes and Archotermopsis in particular have a similar but it faces forward and sideways com- primitive, complete Y-suture (Emerson, 1933). pared with Coptotermes. A distinct, small projec- Some species of Coptotermes have a distinct tion is present on the dorsal margin of the ocellus forked fontanelle indicating the vestigial arms of in Termitogeton. The ocellus is proportionally the Y-suture, and the stem is distinct, but C. tes- farther from the eye in Psammotermes allocerus, taceus lacks both the fork and the stem. The dis- Stylotermes, Parastylotermes, Macrorhinotermes, tinct arms of the Y-suture of Termitogeton form Rhinotermes, Dolichorhinotermes, and Aco- an angle greater than a right angle. Stylotermes rhinotermes. The distance of the ocellus from the chakratensis Mathur and Thapa (= Sarvaritermes eye often varies from species to species within a faveolus Chatterjee and Thakur) has a complete single genus. Some species of Heterotermes have Y-suture, whereas Stylotermes fletcheri and Para- the ocelli reduced or absent. stylotermes robustus do not. The stem and arms Antennae. In any sequence of related genera of the Y-suture are barely distinguishable inPro- or species, the more primitive taxa have the rhinotermes. The arms are distinct in Parrhino- largest number of antennal articles, although re- termes, but the stem is indistinct or absent. The duction may be analogous rather than homol- Y-suture of Serritermes is partially regressed. ogous. Coptotermes has 18-22, Psammotermes Compound Eye. The lenticular flatly convex has 15 or 16, Heterotermes has 13-19, Reticuli- eyes of Archotermopsis and Ulmeriella are blat- termes has 13-20, Termitogeton has 17, Stylo- toid in shape and are the most primitive among termes has 20-22, Prorhinotermes has 18, Macro- termites (Emerson, 1933, 1968b). The rounded rhinotermes has 20, Rhinotermes has 20, eye of Serritermes is proportionally larger than Dolichorhinotermes has 20, and Serritermes has that of Coptotermes, Stylotermes, Termitogeton, 13 or 14. Nearly all species of the Rhinotermiti- and Psammotermes, and the convex projecting dae have more antennal articles than Serritermes, eye of these genera and Mastotermes is probably but a rare exception is the fossil Reticulitermes primitive, although the flat eye of Archo- minimus with 13 or 14 (Emerson, 1971, p. 280). termopsis is more primitive. The eye of Prorhino- .Mastotermes has 30 or 31, Ulmeriella has 27-29, 14 AMERICAN MUSEUM NOVITATES NO. 2570 and Archotermopsis has 23-27. In some cases plate has about 15-18 ridges and a rounded ridge there is a correlation between the number of an- above the molar plate without a toothlike projec- tennal articles and general size, but in several gen- tion. The proportions of the marginal teeth of era the number does not vary with the relative the left mandible of Coptotermes, Psammo- size of the head. termes, Heterotermes, Termitogeton, and Pro- The third article is shorter than or equal to the rhinotermes are similar, but there are small varia- fourth in Mastotermes, Archotermopsis, Ulmeri- tions in the number of molar ridges (seven or ella, Coptotermes,Psammotermes, Heterotermes, eight in Heterotermes). The first marginal tooth Reticulitermes, Termitogeton (some specimens), of the left mandible is larger than the second in Stylotermes, and Parastyloternes. The third arti- Stylotennes, Parrhinotermes, Macrorhinotermes, cle of Serriternes is equally primitive. The third Schedorhinotermes, Rhinotermes, Dolichorhino- article is longer than the fourth in Ternitogeton termes, and Acorhinotermes. The smaller number (some specimens), Prorhinotermes, Parrhino- of molar ridges is derivative in Serritermes, but termes, Macrorhinotermes, Schedorhinotermes, the presence of a few molar ridges is primitive Rhinotermes, Dolichorhinotermes, and Aco- compared with some genera of the Termitidae, rhinotermes-a derived character. Since the third particularly in the Subulitermes branch of the article is the one that divides, variation in its rela- Nasutitermitinae, in which they are regressed in tive length often relates to the number of articles association with a humivorous diet. in individuals of the same species or genus. Postclypeus. The short, flat postclypeus of Mandibles. The most archaic termite mandi- Mastotermes and Archotermopsis is more primi- bles are those of Archotermopsis, with three dis- tive than that of any genus of the Rhinotermiti- tinct marginal teeth and 10 molar ridges on the dae. The relatively long-arched postclypeus not left mandible and two distinct marginal teeth, a projecting over the base of the labrum is deriva- subsidiary tooth on the front base of the first tive in Serritermes compared with the relatively marginal tooth, and 12 molar ridges on the right short (length less than half width) convex post- mandible. All genera of the Rhinotermitidae have clypeus of Archotermopsis, Coptotermes, Psam- dentition strikingly homologous to that of motermes, Heterotermes, Reticulitermes, Termi- Archotermopsis (Emerson, 1933, 1971; Ahmad, togeton, Stylotermes, and Parastylotermes. In 1950). The first and second marginal teeth of the Prorhinotermes and Parrhinotermes the post- left mandible are approximately equal in Archo- clypeus is wider than long, but is long compared termopsis and Reticulitermes, but these propor- with Coptotermes. It has a distinct convex longi- tions may not be of phylogenetic significance in tudinal median line in Heterotermes, Prorhino- Reticulitermes, which is not primitive among the termes, Parrhinotennes, Schedorhinotermes, Rhinotermitidae. Because of the numerous primi- Rhinotermes, Dolichorhinotermes, and Aco- tive characters of Coptotennes, its mandibles are rhinotermes. The anteclypeus does not project also considered primitive among the Rhino- over the base of the labrum in Prorhinotennes termitidae and are the type from which the man- and Parrhinotermes in which the profile is in line dibles of Serritermes probably were derived. The with the top of the head. The sclerotized post- left mandible of Coptotermes has three marginal clypeus of Macrorhinotermes is slightly arched in teeth, the first smaller than the second and the profi'le and joins the front of the head with second smaller than the third. The molar plate hardly an indentation in the middle which has a has 13 or 14 ridges and is bordered below by a shallow groove and a short longitudinal line. The sharp ridge extending to the base of the third front is nearly straight, and the front of the marginal tooth, with a blunt angle (close to a anteclypeus is rounded and projects over the base right angle) below the front end of the molar of the labrum. The postclypeus is longer than the ridges. The right mandible has a large, sharp first membranous and softer anteclypeus. The angle be- marginal tooth, a subsidiary tooth or hump on tween the anteclypeus and labrum is slightly less the front margin near its base, and a blunt second than a right angle in profile. The anteclypeus pro- marginal tooth with a long posterior cutting edge jects over the base of the labrum much more in between its rounded tip and the base. The molar Schedorhinotermes, Rhinotermes, Dolichorhino- 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 15 termes, and Acorhinotermes. The profile of the prothoracic, mesothoracic, and metathoracic junction of the postclypeus with the front of the spurs are 3/4/34 in Mastotermes, 3-5/3/2-3 in head is without an angle in Archotermopsis, Archotermopsis, and 3/2/2 in Coptotermes, shows a small or slight angle in Coptotermes, Psammotermes, Heterotermes, Reticulitermes, Psammotermes, Heterotennes, Reticulitermes, Stylotermes, Prorhinotermes (some species), Stylotermes, Parastylotermes, and Serritermes, Macrorhinotermes, and some specimens of Par- and is without an angle in Prorhinotermes, Par- rhinotermes. The outer spur is shorter than either rhinotermes, Macrorhinotermes, and Schedo- of the inner spurs in Prorhinotermes and Par- rhinotermes. The distinct angle in Rhinotermes, rhinotermes (outer spur not seen in all speci- Dolichorhinotermes, and Acorhinotermes is mens). The spurs are 2/2/2 in Termitogeton, secondarily derivative. Parastylotermes, and some species of Prorhino- Pronotum. The relatively wide, flatly arched termes, and in Rhinotermes, Dolichorhinotermes, pronotum of the Blattodea, Mastotermitidae, and Acorhinotermes, and Serritermes. The 3/2/2 for- Kalotermitidae is more primitive than that of the mula is considered primitive in the Rhinotermiti- Hodotermitidae, Rhinotermitidae, Serritermiti- dae, and is also primitive in the Termitidae. dae, and Termitidae. The most primitive pro- Tarsi. The tarsus has four articles in all but notum among the Rhinotermitidae is that of two genera of the Rhinotermitidae; five articles Coptotermes, where it is occasionally wider than in the Mastotermitidae, the primitive Termopsi- the head but usually narrower. The width in nae, and Ulmeriella of the Hodotermitidae; four Stylotermes, Prorhinotermes, and Serritermes is articles in the Serritermitidae and Termitidae; only slightly less primitive compared with Copto- and only three articles in Stylotermes and Para- termes. The widely concave front margin and the stylotermes of the Rhinotermitidae; and rarely in rounded sides of the pronotum of Mastotermes the Termitidae (Indotermes). The three-jointed and the Kalotermitidae is the most primitive tarsus is used as evidence of family rank by some among termites. Archotermopsis has a pronotum authors, but does not warrant such a category narrower than the head, sides somewhat convex unless associated with many other sharply defined and angular, and a low ridge behind an indented characters without known intergradations. and slightly lobed front margin-probably more A projecting arolium between the tarsal claws derivative than either Mastotermes, Ulmeriella, or is a primitive character of Archotermopsis, Mas- Coptotermes. In Coptotermes the front margin is totermes, and numerous genera and species of slightly indented as in Heterotermes and Pro- the Kalotermitidae, but is absent in the Rhino- rhinotermes, the frontal lobes are not so promi- termitidae, Serritermitidae, and Termitidae. nent as in Serritermes, there is a low ridge behind Wing Scales. The base of the wing behind the the front margin, and the sides are rounded from humeral suture is retained in all termites when above as in Prorhinotermes, Parrhinotermes, and the wings are shed after the colonizing flight. The Serritermes. Heterotermes has more prominent forewing scale of Mastotermes, Kalotermitidae, rounded frontal lobes. The front margin has a and Ulmeriella conspicuously overlaps the hind median projection in Termitogeton and an anal- margin of the mesonotum and the base of the ogous, convex slightly lobed front margin in hind wing scale. The forewing scales of Archo- Rhinotermes, Dolichorhinotermes, and Aco- termopsis are large, but do not overlap the base rhinotermes. The hind margin is straight without of the hind wing scales, whereas in Coptotermes an indention in Prorhinotermes, is shallowly con- the forewing scales slightly overlap the base of cave in Parrhinotermes and Rhinotermes, and is the hind wing scales and extend beyond the hind shallowly indented in Serritermes, but no obvi- margin of the mesonotum. In some species of ous phylogenetic sequence is apparent. Heterotermes, they overlap the base of the hind Tibial Spurs. The larger number of tibial spurs scales, but not the hind margin of the mesono- are often associated with the inner and outer lat- tum. They overlap one-fourth to one-half the eral tibial spines in the Mastotermitidae, Kaloter- length of the hind scales in Prorhinotermes, Par- mitidae, and Hodotermitidae that antedate the rhinotermes, Macrorhinotermes, Schedorhinoter- loss of lateral spines in the Rhinotermitidae. The mes, Rhinotermes, Dolichorhinotermes, and 16 AMERICAN MUSEUM NOVITATES NO. 2570

Acorhinotermes. The forewing scale is propor- togeton, Parrhinotermes, some species of tionally short in Psammotermes, Reticulitermes, Dolichorhinotermes, and Serrtermes lack Sc be- Termitogeton, and all the Termitidae. The pro- yond the humeral suture, doubtless indepen- portions in Serritermes are intermediate com- dently reduced in the different phylogenetic pared with the genera of the Rhinotermitidae. branches. The proportional reduction of the size of the The radius (R) and radial sector (Rs) have forewing scales is always derivative, but is also a numerous branches joining the costal margin in parallel analogy in several independent branches the Mastotermitidae (Spargotermes and Masto- of the Rhinotermitidae. termes), nearly all genera of the Kalotermitidae, Wing Membrane. Archotermopsis, Masto- and in the Termopsinae. An unbranched Rs is termes, Prorhinotermes, the advanced genera of close and parallel to the costal margin in all the the Rhinotermitinae, and Serrtermes lack punc- Rhinotermitidae and Serritermitidae, although tations on the wing membrane. Coptotermes, there is a little more separation in Stylotermes Psammotermes, Heterotermes, Termitogeton, and a slight divergence in the fossil Parastylo- Stylotermes, and Parastylotermes have rounded termes. punctations or micrasters differing in density, The media (M) is a distinct, separated vein at size, and darkness in different species. Although the humeral suture of the forewing and is joined indicating relationships between genera, puncta- to the Rs a short distance beyond the suture in tions are probably derivative compared with the the hind wing. The position in the middle of the ancestry of the Rhinotermitidae, but are associ- wing tends to be closer to the Rs or about half- ated with other primitive characters in the most way between the Rs and Cu in Mastotermes, primitive genera of the Rhinotermitidae. Heterotermes, and Reticulitermes but is closer to Chitinized vertical, parallel, or irregular reticu- the Cu in Archotermopsis, Coptotermes, and lations between the veins in the apical seven- other genera of the Rhinotermitidae. The M is eighths of the wing are characteristic of primitive more branched in the primitive genera, the and advanced genera, including Mastotermes, branches usually ending near the tip or on the Archotermopsis, Psammotermes, Stylotermes, inner margin in the outer half of the wing, but in Parastylotermes, Dolichorhinotermes, and Seri- some cases the branches end in the membrane or termes (fig. 3). Pseudo-cross veins or veinlets be- occasionally join the Rs or Cu in some speci- tween the branches of the radial sector (Rs) and mens. The M is more uniform in Coptotermes, the costal margin in the apical fourth of the wing Heterotermes, and Reticulitermes, but is variable are found in the primitive and advanced genera, and sometimes absent as a distinct vein in some and often in those genera in which the reticula- specimens of Psammotermes, Stylotermes, Pro- tions are found only in the apical two-thirds of rhinotermes, and Seritennes. The M is absent in the wing, such as Reticulitermes and Prorhino- Ternnitogeton. Variability is derivative compared termes, or are reduced as in Coptotermes, Het- with more uniformity in some genera. erotermes, Termitogeton, and Schedorhinoter- The Mastotermitidae have an anterior cubitus mes. (CuA, or Cu in higher families) and a posterior Wing Venation. The subcosta (Sc) occurs in cubitus (CuP) that is lost in other families. The the forewing of the Hodotermitidae, Mastotermi- Cu is unique in Termitogeton in having costal tidae, and Kalotermitidae, and also may be pres- branches that fill the area occupied by the M in ent in the hind wing of the most primitive Masto- other genera. termitidae (Spargotermes, Emerson, 1965) and The anal vein (A1) has numerous short inner Hodotermitidae (Archotermopsis and Zooter- branches in the Mastotermitidae, is reduced to mopsis, Emerson, 1933). The more primitive gen- two branches or a single vein (A) in the Termop- era of the Rhinotermitidae such as Stylotermes, sinae, and is a short single vein in the hind wings Parastylotermes, and some species of Prorhino- of all except one genus of the Kalotermitidae and termes, Rhinotermes, and Dolichorhinotermes in Reticulitermes, Stylotermes, Parastylotermes, possess a short Sc in the forewing only. Copto- Parrhinotermes, and Schedorhinotermes. The termes, Psammotermes, Heterotermes, Termi- hind wing of Rhinotermes has a diffuse chitin- 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 17 ized area that may be a vestigial anal vein. The Soldier Castes. Monomorphic soldiers with A is absent or indistinct in the hind wings of biting mandibles are found in all primitive fami- Coptotermes, Psammotermes, Heterotermes, Pro- lies, although gradation of sizes occurs in Archo- rhinotermes, Dolichorhinotermes (similar to Rhi- termopsis and Psammotermes and the more prim- notennes), and Serritermes. The anal veins in the itive genera of the Rhinotermitidae. A sharp anal lobe are branched and radiating in the hind dimorphism with a major and minor soldier oc- wing in some Blattodea, Spargotermes, and curs in Schedorhinotermes with a sequence of Mastotermes. reduced mandibles in the minor soldier from Styli. The presence of styli in the male is a Schedorhinotermes through Rhinotermes, Doli- primitive character-the more distinct and long chorhinotermes, and Acorhinotermes, the the more primitive (Archotermopsis, Emerson, latter genus having a monomorphic minor soldier 1933, p. 174, fig. 16; Mastotennes, Silvestri, only with the absence of the biting major soldier. 1909, pl. 16, fig. 13). Coptotermes, Psam- Serritermes, with a monomorphic biting soldier, motermes, Heterotermes, Reticulitermes, and is apparently most closely related to Glosso- Prorhinotermes have short styli in the male. termes in this particular, although few specimens Termitogeton, Stylotermes, Parrhinotermes, Rhi- have been collected. notermes, Dolichorhinotermes, and Serritermes Pilosity. Scattered short hairs on the heads of lack styli-probably an analogous regression in Archotermopsis, Mastotermes, and Serritermes relatively independent phylogenetic branches. are considered primitive. Glossotermes and Stylo- Cerci. Cerci with more than two articles are termes have scattered, contrasting bristles, which found only in the most primitive termites, like are lacking in Serritermes, and the short hairs are Mastotermes and Archotennopsis. Two articles- reduced. Glossotermes has microscopic hairs in an apical, and a bulbous basal article-are found the middle of the labrum and two conspicuous in the Rhinotermitidae, Serritermitidae, and short bristles at the tip. Psammotermes has Termitidae. microscopic hairs on the top of the head, with relatively long bristles on the ventral side and postmentum. Heterotermes, Prorhinotermes, and SOLDIER Parrhinotermes have numerous long bristles on Size. Large relative soldier size is characteristic the head with few microscopic hairs. Termito- of nearly all primitive genera in all families of geton has many hairs and numerous bristles. termites. Mastotermes is moderately large. Rhinotermes, Dolichorhinotermes, and Aco- Archotermopsis is very large. Psammotermes al- rhinotermes have long bristles only. The evolu- locerus has robust, large soldiers with a gradation tion of bristles and reduction of short hairs is of sizes within the same colony. Psammotermes considered derivative, although not necessarily hybostoma has a relatively thicker head than homologous in widely separated genera. does P. allocerus, but the general size of the Head Capsule. The relatively thick elongated largest soldiers is not so great as in P. allocerus. heads of Mastotermes, Glossotermes, Hetero- Rhinotermes has moderately large major soldiers. termes, Reticulitermes, Prorhinotermes, and The size is more uniform and relatively smaller in some species of Parrhinotermes are considered Heterotermes (length of head to side base of primitive. The head is relatively wide in Prorhino- mandibles, 1.15-2.72 mm.; width of head, termes, Rhinotermes, and Dolichorhinotermes. 0.68-1.40 mm.), Stylotermes, Prorhinotermes, The flattened heads of Archotermopsis, some Parrhinotermes, and the major soldier of species of Psammotermes, Termitogeton, and Dolichorhinotermes. The elongated head of Glos- some species of Parrhinotermes, that in extreme sotennes is moderately small (thickness of head cases may be lobed at the sides in the rear, are without postmentum, 0.76 mm.; with postmen- derivative and usually convergent. Both Glosso- tum, 0.82 mm.). In Heterotermes the thickness termes and Serritermes have convex fronts of the of the head with postmentum ranges from 0.66 heads in profile, with a shallow concave junction mm. to 1.27 mm. Serritermes has a compara- with the vertex and also with the postclypeus. tively small, short, thick head. The flatter or very slightly convex front in 18 AMERICAN MUSEUM NOVITATES NO. 2570

Archotermopsis and Mastotermes is considered far forward position of the fontanelle in the primitive, and increased convexity, ridges, minor soldiers of Rhinotermes, Dolichorhino- grooves, and lateral humps are deemed derivative termes, and Acorhinotermes in which the gland in Psammotennes, Heterotermes, Reticulitermes, extends into the abdomen and the secretion Prorhinotennes, Schedorhinotermes, Rhino- functions as a gaseous repellent defense with as- termes, and Dolichorhinotermes. Within the sociated frontal, clypeal, and labral grooves, dis- Rhinotermitidae and Serritermitidae, the irregu- persion hairs, and reduced mandibles. lar, flat front of Psammotennes and Termito- The Y-suture of the soldiers is distinct and geton and the slightly to more strongly convex entire in Mastotermes and Archotennopsis. It front and vertex of Glossotennes, Stylotermes, tends toward reduction in part or in whole in the Prorhinotermes, and Serritermes are primitive, Rhinotermitidae with gradations of arms and but derivative compared with the Mastotermiti- stem forming evolutionary sequences. In Ter- dae and Hodotermitidae. The shallow longitudi- mitogeton the arms are distinct, but the stem is nal groove in the front and convex humps on the absent. The median stem is distinct in the major sides are more developed in Glossotermes than in soldier of Dolichorhinotermes, but the anns are Serritennes and even more developed in Hetero- absent (some species). The vestige of the stem at termes and Reticulitennes. The head above the the rear of the head of Serritermes (fig. 4) is antennal fossae is more convex and bluntly pro- more primitive than the abscence of the Y-suture jecting in Glossotermes than in Psammotermes. in such genera as Glossotermes, Heterotermes, Conspicuous protuberances are present above the Prorhinotermes, Rhinotermes, and Dolichorhino- fossae of Archotermopsis (Emerson, 1933, fig. termes (some species). The reduction of the 17). Low ridges above the fossae occur in Masto- Y-suture is probably convergent in several genera. termes, Psammotermes, Termitogeton, and Stylo- Postmentum. The proportionally more elon- termes with some reduction in many other gen- gated postmentum of Glossotermes (Emerson, era of the Rhinotermitidae. 1950, fig. 1), Heterotermes, Termitogeton, Pro- The frontal gland and its fontanelle opening rhinotermes, Parrhinotermes, major soldier of are absent in the Mastotermitidae, Kalotermiti- Schedorhinotermes, Rhinotermes, and Dolicho- dae, and Hodotermitidae. The frontal gland and rhinotermes, constricted from below and less fontanelle are homologous in the Rhinotermiti- bulbous in profile, is considered more primitive dae, Serritermitidae, and Termitidae, with a defen- than that of Serritermes (fig. 4). The shape of the sive function of the glandular secretion in the postmentum is probably correlated with the rela- soldier caste (see Quennedey, 1973). A relatively tive length of the head. small, pear-shaped or oval gland and a fontanelle Compound Eye. Eye spots narrower than the placed to the rear of the level of the antennal antennal fossae, but with visible ocular sutures, fossae with a angle close to a right angle between are found in Mastotermes, Archotermopsis, the fontanelle and the fossae are characteristic of Psammotermes, Glossotermes (length of eye 0.13 the more primitive genera such as Psammo- mm., width of eye 0.10 mm., eye from fossa termes, Glossotermes (size of gland unknown), 0.74 mm., length of fossa 0.15 mm., width of Heterotermes, Termitogeton, and Stylotermes. fossa 0.13 mm.), Heterotermes (most species), The frontal gland of Termitogeton is pear-shaped Termitogeton, Stylotermes, and Prorhinotermes. but proportionally wider and longer than that of In Parrhinotermes the eye spot is present, but Serritermes. Seritermes is related to this group of irregular and indistinct. In Rhino termes and Doli- genera, although the fontanelle ofPsammotermes chorhinotermes both major and minor soldiers is slightly larger with a dark edge. The forward have very small, light eye spots. The mono- position of the large fontanelle and the enlarged morphic minor soldier of Acorhinotermes has a frontal gland extending into the abdomen are distinct, small eye spot. Distinctive pigmented, unique and analogously derived in Coptotermes. faceted eyes are present and possibly functional The fontanelle is placed in a more forward posi- in the Hodotermitinae. Pigmentation and facets tion starting with Prorhinotermes and ending are regressed in Mastotermes and Archo- with the extreme enlargement of the gland and termopsis. Pigmentation and facets show grada- 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 19 tions of reduction in the Kalotermitidae, with mitidae, Hodotermitidae, primitive genera of the pigmented facets in Pterotermes occidentis that Rhinotermitidae such as Coptotermes and Heter- are more reduced in Neotermes mona and N. otermes, and in Serritermes. In Glossotermes the jouteli. One undescribed species ofHeterotermes third article is about equal to the second and is from New Guinea has a small, pigmented eye slightly longer than the fourth. In Termitogeton spot without a distinct ocular suture and may be the third may be shorter or longer than the primitive in this respect. Facets are vestigial in all fourth. The third article is conspicuously longer the Rhinotermitidae, although some irregularities than the fourth in Psammotermes, Stylotermes, of the surface may be the last indications of the major and minor soldier ofRhinotermes, the regressed facets. In general, the proportionally major and minor soldier of Dolichorhinotermes, larger oval eye with an ocular suture in Serri- with equal length in some minor soldiers, and the termes is more primitive than in any genus of the minor soldier of Acorhinotermes. The antenna is Rhinotermitidae. moniliform in termites with the attachments of Ocelli. The presence of vestigial ocelli in Glos- the articles thicker toward the base than in the sotermes, some species of Stylotermes and Pro- more flexible outer portion. In Serritermes (fig. rhinotermes, the major soldier of Dolichorhino- 4), the bulging end of the middle articles is termes, and in Serritermes is a primitive character unique. compared with their absence in Psammotermes, Postclypeus. The postclypeus is separated Termitogeton, Prorhinotermes inopinatus, Par- from the front of the head by a complete, more rhinotermes, major and minor soldiers ofRhino- or less distinct suture in Mastotermes, Archo- termes, Dolichorhinotermes (some species), and termopsis, Coptotermes, Glossotermes, Psam- the minor soldier ofAcorhinotermes. motermes hybostoma, Heterotermes, Stylo- Antennae. The number of antennal articles is termes, Prorhinotermes, and Serritermes. In greater in primitive genera and reduced in deriva- Psammotermes allocerus, the suture with the tive genera. Without exhaustively counting all front of the head is distinct at the sides but indis- species of each genus, the approximate ranges are tinct in the middle. In Termitogeton the post- Mastotermes 23 or 24, Archotermopsis 22-27, clypeus is short, but fused with the front. In Par- Coptotermes 14-16, Psammotermes 12 or 13, rhinotermes, the postclypeus is fused with the Glossotermes 13, Heterotermes 12-18, Ter- front of the head. The shape is short and flat in mitogeton 13 or 14, Stylotermes 13-19 Pro- Mastotermes, Archotermopsis, Coptotermes, and rhinotermes 16-18, Parrhinotermes 13, Schedo- Serritermes, relatively short and more or less con- rhinotermes 15 or 16, Rhinotermes major soldier vex in Psammotermes and Glossotermes, and pro- 15-17 and minor soldier 15 or 16, Dolichorhino- portionally longer in Heterotermes, Stylotermes, termes major soldier 16 and minor soldier 14-16 Prorhinotermes, and Schedorhinotermes. A (young colony 12), Acorhinotermes 15, and Ser- shallow longitudinal groove occurs in the middle ritermes 12 or 13. The comparative reduction of in Archotermopsis, Psammotermes, and Glosso- antennal articles in imagoes is correlated with the termes. The groove is more distinct, deeper, nar- soldiers that invariably have fewer articles than rower, and continuous from the fontanelle to the imagoes of the same genus or species. There is tip of the labrum in Parrhinotermes, Schedo- some correlation between the size of the soldier rhinotermes, the major soldier of Rhinotermes, and the number of antennal articles, but in gen- the major soldier ofDolichorhinotermes and par- eral the trend is toward evolutionary reduction, ticularly in the minor soldier of Rhinotermes, with only a few exceptions in genera of the Ter- Dolichorhinotermes, and Acorhinotermes, in mitidae. It is fairly obvious that reduction has which the groove is in an elevated ridge and func- occurred independently in several phylogenetic tions for the dispersion of the frontal gland secre- branches, but the small number in Psammo- tion. This groove is absent in Coptotermes, termes, Glossotermes, and Serritermes possibly is Termitogeton, and Serritermes, and the front of homologous. the postclypeus is emarginate (fig. 4) in contrast The third article is either the shortest or about to the straight or slightly convex front margin in equal to the short fourth article in the Mastoter- Psammotermes, Glossotermes, Heterotermes, 20 AMERICAN MUSEUM NOVITATES NO. 2570

Stylotermes, Prorhinotermes, and in the major archaic imago mandibles. The left mandible of and minor soldiers of Schedorhinotermes. The the soldier of Mastotermes and the primitive postclypeus and front of the head of Copto- Kalotermitidae has three marginal teeth in con- termes is highly derivative in association with the trast to the fused first and second of the imago. forward position of the large fontanelle and the We assume that the soldier mandibles of the enlargement of the frontal gland. The post- ancestral Rhinotermitidae had three marginal clypeus of Glossotermes appears to be the most teeth on the left mandible and two distinct mar- primitive among the Rhinotermitidae, is slightly ginal teeth on the right mandible. The major less primitive in Psammotermes, and in somewhat soldier of Dolichorhinotermes has two distinct different characteristics is less primitive in Copto- marginal teeth on both right and left mandibles termes. Serritermes is most closely related to and a small toothlike projection on the base of that of Glossotennes. each mandible, but no molar ridges. Two distinct Labrum. The labrum of Mastotermes and marginal teeth on the left mandible and one on Archotermopsis has a wide, moderately straight the right mandible are seen in Parrhinotermes, front margin, with somewhat convex sides. The Schedorhinotermes, and Rhinotermes (a small labrum of Prorhinotermes and Stylotermes is hump close to the base may be a vestigial third tongue-shaped, with a broad, convex front mar- marginal on the left mandible, and a minute gin without a white lip. It is derivative in vestige of the second marginal occurs on the right Glossotermes (Emerson, 1950, fig. 1), particu- mandible), but other genera show varying degrees larly in the broadly convex, whitish anterior lip, of reduction of marginal teeth, probably analo- the horizontal, brownish yellow band in the gous in separate subfamilies. The basal portion of middle, and the slight constriction of the sides the mandible is distinct, with a notch on the behind the tip. The much more constricted sides, outer margin at the junction with the apical por- the distinctive smaller tip, and the wider middle tion in the more primitive genera and in Par- portion in Psammotermes indicate relationship to rhinotermes, but less distinct in various derivative Glossotermes. The tip of the labrum of Termito- genera. Molar ridges are present in Archo- geton has a small, pointed hyaline tip, with a termopsis and Parrhinotermes, but are lacking in slight constriction behind. Although derivative all other rhinotermitids and in Serritermes. The and unique in its shape, the labrum of Serri- left mandible of Glossotermes has a sharp (600 termes (fig. 4) may be remotely related to that of angle) basal, toothlike projection that is bluntly Glossotermes and possibly to Termitogeton. The convex in Stylotermes and may be homologous elongated narrowly convex or pointed tip of with the smaller, sharper projection in Serriter- Coptotermes and Heterotermes is probably mes. The left mandible of Termitogeton has a analogous to Serritermes. The concave or bilobed finger-like projection basal to a slight hump that tip of the elongated labra of Parrhinotermes, may be a vestigial marginal tooth. The left man- Schedorhinotermes, Rhinotermes, and Dolicho- dible of Prorhinotermes resembles Stylotermes rhinotermes indicates divergent evolution in a with a reduced third marginal and other teeth separate phylogenetic branch. The major soldier minute or absent. The right mandible of Ter- ofRhinotermes has a wide labrum with a convex, mitogeton has minute inner serrations, three slightly bilobed front margin, whereas the labrum slightly more visible than others. The right man- is narrow and distinctly bilobed in the Dolicho- dible of Prorhinotermes has a minute, pointed rhinotermes major mandibulate soldier and basal projection and vestigial first and second extremely narrow, elongate, and bilobed in the marginal teeth near the base. The blades of both minor soldier with vestigial mandibles in Rhino- mandibles curve inward, more so near the tip, termes, Dolichorhinotermes, and Acorhino- and taper to a sharp pointed tip not markedly termes. hooked. Serritermes shares a series of serrations Mandibles. Archotermopsis has the most with Glossotermes and Psammotermes with dif- primitive mandibles among living genera ferent numbers, proportions, and positions in (Emerson, 1933, fig. 20). The marginal teeth and each genus. Conspicuous serrations are derivative, molar plates are easily homologized with the contrary to Emerson (1971, p. 254, table 2). '5 EMERSON AND KRISHNA: SERRITERMITIDAE 21 all undulations of the inner margin near the tum ofArchotermopsis are both probably deriva- ,e in some rhinotermitids may be reduced tive from the wider, somewhat arched and flatter rginal teeth rather than serrations. Glosso- pronotum with rounded sides characteristic of mes and Psammotermes have tapering blades, several genera of the Kalotermitidae, and with hooked at the tips in contrast to Serritermes some resemblances to the pronota of many cock- J. 5). A tooth near the base of the right man- roaches. Among the Rhinotermitidae, the most ile of Glossotermes, together with a small, low, primitive pronotum is that of Coptotermes and ;htly pointed hump between the tip of the Prorhinotermes (Silvestri, 1909, fig. 59) with an ge tooth and the cutting edge of the blade (not indented front margin, slightly convex frontal wn by Emerson, 1950, fig. 1) may be homol- lobes, slightly convex sides, and slightly concave )us with the marginal teeth ofParrhinotermes. hind margin. The pronotum of Heterotermes is e outer margins of the base of each mandible narrow, with variations from rounded to straight Glossoternes and Psammotermes are convex. front lobes, and its sides and hind margin are relatively deep notch basal to the serrations on sometimes emarginate. The pronotum of Ter- left mandible of Glossotermes possibly indi- mitogeton has a bluntly angular, somewhat raised ,es the separation of the base from the toothed median projection. The pronotum of Stylotermes tting edge seen in Parrhinotermes. The 10 to is similar to that of Prorhinotermes, but propor- serrations on the basal two-thirds of the left tionally longer. Those of Glossotermes and Par- ndible of Glossotermes (Emerson, 1950, fig. rhinotermes are proportionally narrower than the are not readily homologized with the marginal head. In Glossotermes (Emerson, 1950, fig. 1), th of other genera; the inner margin of the the frontal lobes are proportionally narrower and ide is concave from the notch to the slightly more convex than in Prorhinotermes, and the oked and upwardly curved tip. The outer mar- sides are convex, evenly joining the slightly in- t of the right mandible of Glossotermes has a dented hind margin. The pronotum of Parrhino- trp but wide notch at the junction of the blade termes is strongly convex in front with a small i base and an evenly convex outer edge behind median indentation, its sides flatly convex, and a slightly hooked tip. The lower edge of the hind margin straight, with no indentation. In the ;e of the right mandible has a large, blunt, major soldier of Rhinotermes and Dolichorhino- )thlike projection that is probably homologous termes, the middle of the front margin is con- th the sharp, minute tooth on the base of Ser- spicuously raised with a convex lobe which is 'rmes (fig. 5). A small, slightly convex hump narrower and more sharply convex in the minor the apical portion of the large basal projection soldier of Rhinotermes, Dolichorhinotermes, and Glossotermes is absent in Serrtermes. In Glos- Acorhinotermes. The pronotum of Psammo- ermes, a sharp, deep notch with an angle termes allocerus is wide and relatively long, with ler than a right angle is between the inner mar- a small notch in the middle of the front margin, of the blade and the base of the right man- and somewhat concave in the middle of the hind dle. The serrations of Psammotermes are larger margin. It is about as wide as the head, and the I coarser than in Glossotermes, but in both frontal lobes are raised and overlap the back of iera the serrations diminish in size from the the flat head. The sides are moderately convex in ,e to the apex, whereas the reverse is true for front, concave or almost straight in the middle, -ritermes (fig. 5). The bases of the mandibles and evenly convex as they join the hind margin. Psammotermes are proportionally large and The pronotum of P. hybostoma is flatter than ivative. The outer margins have less distinct that of P. allocerus, with less prolonged frontal Ales at the junctions of the bases and blades in lobes. Serritermes has a unique and highly !mmotermes hybostoma compared with those derived pronotum (fig. 4), but possibly is re- P. allocerus. The concave area between the motely related to that of Glossotermes. ier base and upper ridge of P. hybostoma is Legs. The close proximity of the right and left t as marked or sharp as in P. allocerus. coxae is primitive in termite soldiers, including Pronotum. The wide, saddle-shaped pronotum Mastotermes, and Archotermopsis. The striking Mastotermes and the narrower, angular prono- bladelike protuberance on the front coxa ofMas- 22 AMERICAN MUSEUM NOVITATES NO. 2570 totermes is unique and probably derivative. The Styli. The presence of styli in the soldiers is separation of the right and left coxae of Termito- primitive, the longer and more conspicuous the geton is derivative. The normal relative sizes of more primitive. Styli are present in Mastotermes, the coxae, femora, and tibiae ofArchotermopsis Archotermopsis, Psammotermes, Heterotermes are primitive and similar to those of Glosso- (some species), and Prorhinotermes, and absent termes and Serritermes. The coxae of Prorhino- in Termitogeton, Stylotermes, Parrhinotermes, termes are moderately swollen, the prothoracic Rhinotermes, Dolichorhinotermes, Acorhino- and mesothoracic coxae are close, but the meta- termes, Glossotermes, and Serritermes. Loss of thoracic coxae are a little farther apart. The styli in separate branches of the phylogenetic coxae of Parrhinotermes are similar to those of tree is probably analogous, but may be homolo- Prorhinotermes except that they are not so swol- gous in closely allied genera within the same sub- len. The coxae of all thoracic segments are more family. separated in Rhinotermes. The front coxae of Cerci. The cercus has five to seven articles in Acorhinotermes are close, but the mesothoracic Archotermopsis and is twice the length of the and metathoracic coxae are more separated. The stylus. It is moderately long, with five articles in coxae of Glossotermes and Stylotermes are closer Mastotermes, the length about the same as the together than those of Serritermes. Those of stylus. In the Rhinotermitidae, Serritermitidae, Glossotermes are proportionally larger and more and Termitidae the cercus is reduced to two swollen than those of Serritermes. The markedly articles, the basal one swollen in comparison to swollen femora or tibiae of Psammotermes, Ter- the thinner terminal article or joint. mitogeton, and Stylotermes are derivative and possibly an adaptation to fossorial burrowing. WORKERS AND NYMPHS The tibiae of Glossotermes are more slender, less wide, and with far less convex margins than the Intestines. We include comparisons with other robust tibiae ofPsammotermes. termites described by Noirot and Noirot- The inner and outer spines on the tibiae of the Timothee (1965, 1969), with comments based primitive genera Mastotermes and Archotermop- on information supplied by Charles Noirot (per- sis are similar to those of the imago caste. Secon- sonal commun.). The comparisons are confined dary rows of short spines on the front tibiae of to the lower families that show relationships to some genera of the Rhinotermitidae and Termi- the Serritermitidae. The intestines of the Ter- tidae are derivative and often more developed in mitidae are much more varied, doubtless due to the soldier and worker than in the imago. The the more varied substances ingested and also tibial spurs on the legs of the prothorax, meso- probably to more varied digestive physiology. thorax, and metathorax have the same formulae as Few genera of the Rhinotermitidae have been in the imago, and show the same regressions in examined for their intestinal structures, and the evolutionary sequences discussed under the few that have are uniform, without much generic imago caste. The tibial spurs are 2/2/2 in the variation. soldier of Glossotermes, characteristics that may The crop is symmetrical in the Rhino- be a regression homologous with those of Serri- termitidae, but is slightly asymmetrical in Ser- termes, but as in the imago, reduction of num- ritermes. bers may be analogous on separated branches of The armature of the gizzard of Coptotermes the phylogenetic tree. The front tibial spurs of and Reticulitermes consists of sclerotized sur- Psammotermes hybostoma are thicker than those faces of 24 folds divided into three types or of Glossotermes, but not so dark, thick, and ro- orders (Noirot and Noirot-Timothee, 1969, pp. bust as those of Psammotermes allocerus. Psam- 54-55, fig. 2). Twenty-four small folds of the motermes is confined to arid steppe and desert fourth order do not have sclerotized cuticles. In soils, and the legs may be derivative adjustments the anterior region of the gizzard, the six folds of to digging. the first order and six folds of the second order The tarsal articles are the same in the soldier have thick cuticles and partial sclerotization, but as in the imago and follow the same evolutionary in the posterior region, the first order folds are trends toward reduction. prolonged, and have a thick but unsclerotized 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 23 cuticle. The 12 folds of the third order protrude possibly an analogous reduction in the Kaloter- less in the posterior region than in the anterior mitidae, and a homologous reduction in the region, but their cuticle is sclerotized in both Termopsinae and the more advanced families. regions. The folds of the gizzard of Serritermes The comparative ultrastructure and histochemis- are reduced to 24, but project very little and try of a related series of genera may provide have no sclerotization. Those present have very further evidence of systematic relationships (see reduced cuticular armature in comparison with Quennedey, 1969,1971b, 1972). Reticulitermes, but possibly are homologous Seminal Vesicles. The presence of seminal with the first, second, and third order folds, vesicles in the lower families including the whereas the 24 fourth-order folds are absent. The Serritermitidae and their absence in the Ter- reduction of the armature has no equivalent mitidae (Weesner, 1969b, p. 153) indicate an among the lower families of termites. evolutionary reduction, but the details of the The middle intestine shows no differences structure in related series are insufficient for between the Rhinotermitidae and Serriter- adequate comparison between taxa. mitidae. The Mastotermitidae has 12-15 Malpighian PHYLOGENETIC INFERENCES AND tubules, whereas the Termopsinae (Hodo- INTERPRETATIONS termitidae), Anacanthotermes, (Hodotermitinae, Hodotermitidae), Kalotermes (Kalotermitidae), The principles of phylogenetic inference from and Coptotermes, Reticulitermes, and Rhino- taxonomic data are summarized for the Kalo- termes (Rhinotermitidae) have eight. The Serri- termitidae by Krishna (1961, pp. 309-3 10, fig. termitidae have eight, whereas the known genera 81) and for the Rhinotermitidae by Emerson of the Termitidae have two or four, usually four. (1971, pp. 248-254, 265-266, tables 1, 2, fig. 1). The hind intestine is similar to that of the In the present attempt to postulate the ancestry lower termites below the Termitidae, although of the Serritermitidae, a few points need empha- the enteric valve differs. sis. In the opinion of Charles Noirot (personal Nearly every individual organism combines commun.), the characters of the digestive tube ancient homologous monophyletic primitive by themselves do not warrant family status, but characters with derived characters of compar- are not incompatible with family rank. The atively more recent origin. Homologous charac- discovery of flagellates in the intestines by ters in compared organisms seldom have 100 Noirot is consistent with a relationship to the percent identity. Derived characters may be lower families of termites (Krishna, 1970, p. homologous, but are often analogues without 139). Emerson (1971) has discussed the correla- shared genetic origin. In termites, derived charac- tions of the taxonomy of the flagellates and their ters are more often regressions or losses, but hosts. Alcoholic preservation does not allow some are progressive adaptations, particularly the exact description and classification of the defensive functions of the soldiers or the nutri- flagellates, but R. L. Araujo (personal commun.) tive and nest-building functions of the worker- has preserved specimens of Serritermes in like nymphs or the true sterile workers. Since the Schaudinn's fluid, and it is hoped that this genome is usually identical through equational material will enable accurate determinations and mitosis in a single individual organism, the comparisons of genera and species of flagellates. explanation of differences in rates of evolution Sternal Glands. The sternal glands are on the of primitive and derived characters is found in third, fourth, and fifth sternites of Mastotermes the ontogenetic expression of gene systems (Noirot and Noirot-Timoth6e, 1965). They are under different physiological, developmental, or on the fourth sternite only in Stolotermes, environmental conditions. Of course the onto- Porotermes, and the Hodotermitinae. They are at genetic lability may be based upon homologous the anterior border of the fifth sternite only in genetic capacities to react to different stimuli. the Termopsinae (Hodotermitidae), Kaloter- The castes in the same colony are epigenetic mitidae, Rhinotermitidae, Serritermitidae, and modifications resulting from highly similar Termitidae. The position on the fifth sternite is genomes from the same king and queen. Deriva- 24 AMERICAN MUSEUM NOVITATES NO. 2570 tions of multiple gene epigenetic characters The retention of primitive characters by gradually evolve with many gradations. A homol- several taxa allows the postulation of the ogous derivative character can be considered unknown ancestral stem, whereas derivative char- primitive when compared with even more derived acters indicate the branching of the phylogenetic types. Comparative indications of the phylogeny tree. Primitive characters may be assumed to be of the reproductive imago caste and the sterile homologous (Emerson, 1961), but derivative soldier caste provide fine examples of the genetic characters may be either homologous or analo- evolution of epigenetic enhancement or inhibi- gous. It is often difficult to determine whether tion during development. (In the most primitive they are monophyletic or polyphyletic. Difficul- termites, soldiers occasionally lay unfertilized ties of interpretation result from the extinction eggs which may hatch into parthenogenetic of intergrades without many known fossils, the nymphs.) few samples examined for many important and In the case of the frontal gland of the higher unrecorded characters, and the complicated mix- families, the defensive function is largely con- ture of homology and analogy in derived charac- fined to the soldier so far as is known, although ters. However, once a primitive character is lost poorly developed glandular tissues opening or reduced, it is almost never regained or through distinct fontanelles are found in the redeveloped in later more advanced taxa (Dollo's imagoes and workers including those of the Rule). As we know that the genetic capacity for soldier-less genera of the Termitidae. More histo- the development of a compound eye is unim- logical and biochemical study of possible glandu- paired in the reproductive imagoes of all ter- lar defensive functions of imagoes and workers is mites, one might expect that the regression of needed. It is probable that the defensive frontal the soldier eye could be reversed by relatively gland is selected primarily in the soldier, but is simple ontogenetic changes in phyletic genetically transmitted only in the reproductive sequences, but no such case of reappearance is caste. This is a strong indication of the selection known, and many other comparative regressions of the population unit as a whole, thus illus- follow a similar unidirectional evolution. The trating a connecting principle between the most plausible explanation is that there are genetic unity within an individual and a genetic multiple gene effects upon both regressive inhibi- unity of a social population of individuals, a tion and complex progressive functional adapta- principle that is basic to the concept of the social tions, although regression often is more simple supraorganism. The significance of the above genetically than progression (Emerson, 1961). homology and the much more common analogy Evidence is increasingly convincing that effec- between an individual and a population has been tive parts of gene systems have a high order of negated, attacked, or severely criticized by identity over periods of time encompassed by the numerous distinguished biologists, but upheld by evolution of the Isoptera from ancestral cock- others who may, however, use different terms for roaches. This genetic identity does not negate a social unitary systems. great deal of genetic change, and it is the relation It is obvious that the structures of the soldier between identity and change that we attempt to are much farther removed from the ancestral elucidate in this study of the ancestry of the blattoid insect than are those of the imago. For Serritermitidae. this reason, soldier traits more often are deriva- tive regressions or are adaptively progressive with THE ISOPTERAN STEM a greater possibility that similarities between different taxa are parallel or convergent rather By combining the primitive characters in the than genetically homologous. Nevertheless, com- Mastotermitidae and Hodotermitidae, one can parative soldier structures can be postulated to postulate the characters of the unknown ances- be primitive or derivative, and the consistent tral termite which arose from a blattoid ancestor constellation of many characters often is corre- closely similar to the living cockroach Crypto- lated with phylogenetic inferences drawn from cercus punctulatus Scudder of North America. the imago caste. Like all living primitive forms, Cryptocercus has 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 25 some derivative characters, in this case exempli- dentition easily homologized with that of the fied by the loss of wings. imago and resembling that of Archotermopsis, The hypothetical ancestral isopteran imago with many molar ridges on the bases; a flatly was of large size; had the head and wing pilosity arched pronotum; the left and right coxae nearly of Mastotermes and Archotermopsis; a complete touching; normal unswollen proportions of the epicranial suture (Y-suture); the large lenticular femora and tibiae; numerous tibial spines and eye of Archotermopsis and Umeriella (Emerson, spurs; a five-jointed tarsus without arolium; long 1968b); the two ocelli ofMastotermes; antennae styli; many-jointed cerci; and reduced organs of with 30 or more articles and a short third article; reproduction. A true sterile worker caste was mandible dentition close to Cryptocercus and absent, but the pseudergate nymph had at least Archotermopsis; a short flat postclypeus; a wide 12-15 Malpighian tubules, sternal glands on the pronotum with the concave front margin and third, fourth, and fifth sternites, seminal vesicles, rounded sides of Mastotermes; up to five spurs and the hind gut filled with many genera of on some of the tibiae; two rows of lateral inner wood-ingesting zooflagellates. This archaic ances- and outer spines on the middle tibiae; a five- tor probably lived in wet logs, used its excrement jointed tarsus with an arolium between the to build partitions between the excavated gal- claws; humeral sutures separating the basal scales leries in logs, and inhabited temperate latitudinal from the apical portions of both wings (weaker or altitudinal climates similar to the ecology of in the hind wing); the forewing scale large and Cryptocercus, Archotermopsis, and Porotermes overlapping the hind wing scale; no punctations on the continuous land mass of Pangaea before on the wing membrane; chitinous reticulations the split in Permian or lower Triassic times into covering seven-eighths of each wing between the southern and northern continental masses. veins; wing venation resembling the forewing of Of the 64 independently variable characters Mastotermes (Silvestri, 1909, figs. 10, 12) with compared in all castes and nymphs of Masto- the Sc, R1, R2, R3, Rs, M, and Cu separated termes and Archotermopsis, 18 are more primi- beyond the suture; the hind wing resembling that tive in Mastotermes, 34 are about equally primi- of Spargotermes (Emerson, 1965, fig. 4), with tive in both genera, four are equally derivative separated Sc, R1, R2, R3, Rs, the M joined with compared with the blattoids and the Hodo- the Rs close to but beyond the humeral suture, termitinae, and eight are more primitive in anterior Cu, posterior Cu, first anal (A1), radi- Archotermopsis. Not only do these two genera ating branches of the anal field with many show many primitive characters in the living secondary branches in a large anal lobe; long styli species, but each has a long history possibly as in the male and also probably in the female; far back as the Triassic. Mastotermes fossils occur many-jointed cerci with at least nine articles; and in the Eocene, Archotermopsis in Baltic amber of blattoid-like male and female reproductive organs late Eocene or early Oligocene age (Emerson, and appendages (Browman, 1935). A large mono- 1933, 1965), and an advanced hodotermitid, morphic soldier was the only nonreproductive Cretatermes, is known from the mid-Cretaceous caste, resembling Archotermopsis (Emerson, (Emerson, 1968a). 1933, figs. 17-20) more closely than any other living termite, with short scattered hairs on the THE head; large size; a thick head without hind lobes, MASTOTERMITIDAE-KALOTERMITIDAE a pigmented and faceted, reduced eye, resem- STEM bling that of the soldiers of the Hodotermitinae; The possible homologous derivative characters a vestigial ocellus, resembling that of Masto- of the Mastotermitidae-Kalotermitidae branch termes, primitive rhinotermitids, and Serritermes, from the primitive isopteran stem are a fused but lost in the Kalotermitidae, higher Rhino- first and second marginal tooth of the left imago termitidae, and Termitidae; 27 or more antennal mandible, a rounded eye with a flat anterior articles, with the third short; a short postclypeus; margin, and the position of the media vein (M) a tongue-shaped labrum with a straight or slightly which ends close to the tip of the wing and tends convex front margin; elongated mandibles with to be halfway between the radial sector (Rs) and 26 AMERICAN MUSEUM NOVITATES NO. 2570 the cubitus (Cu) or closer to the Rs than to the However, it also had the following derivative Cu in the middle of the wing. A number of characters: a reduction of the radius (R), loss of homologous characters (at least six), including a the posterior cubitus (CuP) and the anal lobe of wide, flatly arched pronotum with a concave the hind wing, the media (M) closer to the Cu front margin in the Mastotermitidae and Kalo- than to the Rs in the middle of the wing, and the termitidae, differ from the derived characters in sternal glands on the third stemite possibly lost. the Hodotermitidae-Rhinotermitidae branch (S1l- An alternative hypothesis is the independent vestri, 1909; Krishna, 1961; Emerson, 1969, radiation of the four more primitive families pp. 4-7, 10, figs. 1, 2). from the primitive isopteran stem. As the Kalotermitidae branched from the As the Hodotermitidae branched from the common stem of the Mastotermitidae-Kalo- Hodotermitidae-Rhinotermitidae stem, the ocelli termitidae, eight regressions in the imago, anal- were lost, the pronotum became narrower, the ogous to similar reductions in the advanced front margin became less concave, and the genera of the Hodotermitidae-Rhinotermitidae Malpighian tubules were reduced to eight (possi- branch, occurred. These include a lesser number bly independently of the reduction to eight in of antennal articles (24-26 in Prokalotermes, the Kalotermitidae). Emerson, 1969, p. 12), a four-jointed tarsus, An early splitting of the Hodotermitidae consolidation of portions of the wing venation, probably occurred, as the Stolotermitinae, Poro- loss of the posterior cubitus (CuP), loss of the termitinae, and Hodotermitinae retained sternal lobe of the hind wing with its branched anal glands on the fourth stemite only, whereas the veins, a two-jointed cercus, eight Malpighian Termopsinae retained sternal glands on the fifth tubules, and the sternal glands confined to the sternate only, a character that also occurs in the fifth sternite. The soldier of the Kalotermitidae Rhinotermitidae-possibly a homologous reduc- lost the ocellus spot, but in some genera retained tion. If so, the loss of the ocelli in all subfamilies three marginal teeth in the left mandible, which of the Hodotermitidae may not be homologous in can also be detected in Mastotermes. This is a every case. rare case of a character more primitive in the soldier than in the imago of the same species. THE RHINOTERMITIDAE STEM Where the reduction resulted in a similarity of derived characters in other branches from the A number of primitive characters were primitive isopteran stem, these characters are retained in the ancestral Rhinotermitidae as it guessed to be analogous, but it should be separated from the Hodotermitidae-Rhinoter- emphasized that such postulates rest upon mitidae stem, notably the head shape of Mas- meager data with alternative hypotheses possible totermes, the archaic blattoid dentition of the if homology and analogy have been confused. An left imago mandible, the subsidiary tooth on alternative hypothesis is the separate radiation of the front base of the first marginal tooth of the the Mastotermitidae and Kalotermitidae from the right mandible seen in Archotermopsis and all primitive isopteran stem with the retention of the Rhinotermitidae, the two ocelli of the many homologous primitive characters and paral- Mastotermitidae, a short third antennal article, lel reduction of numerous analogous derivative the head and wing pilosity close to Archoter- characters. mopsis, a short, flat postclypeus, a wide pro- notum with rounded sides, a large forewing scale overlapping the hind wing scale, chitinous reticu- THE HODOTERMITIDAE-RHINOTERMITIDAE lations covering seven-eighths of the length of the STEM wing, a monomorphic thick-headed soldier, eye The hypothetical ancestor of the Hodoter- and ocellus spots in the soldier, sternal glands on mitidae and Rhinotermitidae possessed most of the fifth sternite of the worker, seminal vesicles, the characters of the primitive isopteran stem and wood-ingesting zooflagellates. Symbiotic cel- including the blattoid lenticular eye, two ocelli, lulose-digesting flagellates are found in all fami- and blattoid dentition of the left mandible. lies of termites except the Termitidae, and 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 27 coevolve with their hosts (Emerson, 1971). termitidae branch. No living or fossil genus The derivative characters of the imago (Emer- of the Rhinotermitidae possesses all the postu- son, 1971, pp. 243-254, table 2) have included a lated primitive characters, and each subfamily trend toward some reduction of the Y-suture; a has derived characters not shared with other rounded eye; some reduction in the number of subfamilies (Emerson, 1971, pp. 251, 254, 256, antennal articles (22 or less); a straighter front fig. 1). The relative degree of primitiveness or margin of the pronotum with a small median derivativeness of living subfamilies or genera is indentation; the loss of all lateral tibial spines; statistical. Relations between some genera are reduction of the spurs to 3/2/2; reduction of better understood than others, and in one case tarsal articles to four; loss of the arolium; (Schedorhinotermes) the living genus is very shortening of the male styli; strengthening of the close to the ancestor of another (Rhinotermes). humeral suture of the hind wing; reduction of the Sc in the hind wing; and loss of RI, R2, and R3 in the forewing. The facets of the eye of the THE SERRITERMITIDAE BRANCH soldier have become vestigial, the third marginal Of 63 characters of the ancestral rhino- tooth of the left mandible has been reduced, and termitid and the castes of Serritermes compared, the soldier styli shortened. In addition to these 24 are more primitive in the ancestral rhino- regressive reductions and losses, the soldier termitid; 32 are equally primitive in both fami- acquired a defensive frontal gland and fontanelle lies, although several of these are derivative opening at the junction of the arms of the compared with the lower families; no characters Y-suture. There is also some indication that the are more derivative in the ancestral rhinotermitid queen became larger with a greater reproductive than in Serritermes, although the soldier eye is capacity of her ovaries, the social life of the proportionally larger in Serritermes than in any colony became more complex, the worker caste known genus of living Rhinotermitidae and was more differentiated from the pseudergate as large presumably in the ancestral rhino- nymphs (a true worker caste seems to have termitid; and seven compared characters do not evolved independently in the Hodotermitinae), show obvious or clear phyletic trends. excrement was used more for nest and tunnel Of the 43 compared characters of the imago, building, and the moist tropical soil and soil- workers, and nymphs of Coptotermes and Serri- connected wood were invaded; this group was termes, 13 are more primitive in Coptotermes, 17 later to be superseded to a large degree by its are equally primitive in both genera, two are more advanced descendants, the Termitidae. equally derivative in both genera but not neces- The closest living hodotermitid with many sarily homologous, five are more primitive in features of the ancestral rhinotermitid is Archo- Serritermes, and the trend is not clear in six termopsis, which, however, evolved some inde- characters. Of the 24 compared characters of the pendent derivative characters. Comparison of the soldier, five are more primitive in Coptotermes, 53 independently variable characters of all castes five are equally primitive in both genera, three in Archotermopsis with those in the hypothetical are equally derivative in both, five are more primitive rhinotermitid, shows 17 to be clearly primitive in Serritermes, and the trend is not more primitive in Archotermopsis, 23 equiv- clear in six characters. Coptotermes, combining alently primitive in both (four of which are 67 characters of all castes, has a generally more equally derivative compared with Mastotermes), primitive imago, but a markedly derivative seven more clearly derivative in Archotermopsis soldier that is not closely related to any other and more primitive in the ancestral rhinoter- rhinotermitid genus. It could not possibly be mitid, and six characters to have inconclusive very close to the ancestor of Serritermes. phyletic trends. Of the 56 compared characters of the imago It is tentatively postulated that the ancestral and soldier of Psammotermes and Serritermes, 12 rhinotermitid branched off from the primitive are more primitive in Psammotermes, 16 are isopteran stem or the more advanced hypo- equally primitive in both genera, nine are equally thetical ancestor of the Hodotermitidae-Rhino- derivative in both genera of which six are 28 AMERICAN MUSEUM NOVITATES NO. 2570 possibly homologous, 11 are more primitive in highly derived Serritermes could not have Serritermes, and the trend is unknown or not evolved directly from Glossotermes, and the gap clear in eight characters. Psammotermes and is sufficient to warrant different family status. Serntermes seem to be remotely related, but The Serritermitidae could have originated from neither could have evolved directly from the an unknown genus related to Glossotermes and other. The imago of Psammotermes in general also related to the common ancestor of Glosso- has more derivative characters than Coptotermes, termes, Psammotermes, and possibly to the but more primitive characters than Serritermes. highly derived Termitogeton. When the imago of The imago of the Heterotermitinae (Hetero- Glossotermes is discovered and compared, the termes and Reticulitermes) has fewer primitive relations with Serritermes and other genera will characters and more derivative characters than be better understood. either the Coptotermitinae or Psammotermitinae We draw the tentative conclusion that the Ser- and, with the soldier characters added, is less ritermitidae can be traced backward to an origin close to the ancestry of the Serritermitidae than from the base of the Psammotermitinae, and in to the Psammotermitinae (Emerson, 1971, pp. sequence back to the primitive rhinotermitid 265, 272). stem, to the hodotermitid-rhinotermitid stem, or Among the rhinotermitid genera, the soldiers to the primitive isopteran stem that arose from of Psammotermes and Glossotermes are most primitive blattoids possibly as early as Permian closely related, each with distinctions in both times. primitive and derivative characters, but both A large number of characters, including the included in the subfamily Psam-motermitinae. dentition of the imago and soldier mandibles, the Unfortunately, the imago of Glossotermes is marked differences in the intestinal structures, unknown. Of the 45 compared soldier characters the reduction of the Malpighian tubules to four of Glossotermes and Serritermes, 16 are more and in some cases to two, the loss of xy- primitive in Glossotermes, 11 are equally primi- lophagous flagellates, and the retention of the tive in both genera, seven are equally derivative frontal gland and sternal glands on the fifth in both genera and all are possibly homologous sternite, strongly indicate the separate origin of regressions, seven are more primitive in Serri- the Termitidae from the Rhinotermitidae inde- termes, and the trend is unknown (size of frontal pendently of the Serritermitidae. gland of Glossotermes) or not clear in four characters. Comparisons of soldiers are not so indicative TIME OF ORIGIN OF TERMITE FAMILIES of phyletic trends as are comparisons of imagoes with additional worker-nymph characters. How- The constellation of primitive and derivative ever, Glossotermes is more closely related to characters of each family, the coevolution of Serritermes than to the soldier of any other flagellates and termitophilous beetles, and the rhinotermitid genus. Because of this, it is possible geographical distribution related to continental to divide some characters into independent vari- drift provide some evidence for the time of origin ables (labrum, mandibles, and pronotum in par- of each known family (Emerson, 1971, pp. ticular). The Neotropical distribution of both 255-257). The Mastotermitidae possibly genera may not be a coincidence. The serrated branched off early in the Mesozoic. The Hodo- mandibles of Psammotermes (Ethiopian, Mala- termitidae probably appeared by Triassic or early gasy, Oriental, and Palaearctic), Glossotermes Jurassic times before the breakup of the southern (Neotropical), and Serritermes (Neotropical) are continents from Gondwana. The Kalotermitidae considered derivative compared with mandibles probably originated before Africa and South having larger and more distinct marginal teeth. America were separated in mid-Jurassic or Lower The mandibles of Glossotermes are more primi- Cretaceous times. The Rhinotermitidae probably tive than those ofPsammotermes. The mandibles appeared in late Jurassic or Early Cretaceous of Serritermes may have evolved from an ances- (Emerson, 1971, fig. 1), and the Termitidae in tral genus remotely related to Glossotermes. The the Cretaceous when northern tropical connec- EMERSON AND KRISHNA: SERRITERMITIDAE 29 tions between Asia and the Americas existed. fran9aises. Paris, vol. 1, no. 4, pp. The primitive Rhinotermitidae originated in trop- 125-365. ical Asia, and ultimately gave rise to the Neotrop- Browman, L. G. ical Serritermitidae, but family distinctions may 1935. The chitinous structures in the poste- not have evolved until the Tertiary. The most rior abdominal segments of certain female termites. Jour. Morph., vol. 57, specialized Termitidae of the subfamilies Ter- pp. 113-129. mitinae and Nasutitermitinae had arisen by mid- Chatterjee, P. N., and M. L. Thakur Cretaceous before the Old World and New World 1964. Sarvaritermes faveolus gen. et. sp. nov. tropics Were separated by sea barriers. The data from Kulu Valley (Punjab: India) are insufficient for more than speculation about [Isoptera], with a discussion on the the time of origin of each family, but the geo- systematic position and relationship of graphic patterns of related taxa make some the family Stylotermitidae. Zool. Anz., postulates more probable than others, and vol. 173, pp. 149-162. increasing information should provide some tests Desneux, J. of alternative hypotheses. 1904. Isoptera. Fam. Termitidae. In Wyts- man, P. A. G., Genera insectorum. Brussels, fasc. 25, pp. 1-52. Emerson, A. E. LITERATURE CITED 1928. Termites of the Belgian Congo and the Ahmad, M. Cameroon. Bull. Amer. Mus. Nat. Hist., 1950. The phylogeny of termite genera based vol. 57, pp. 401-574. on imago-worker mandibles. Bull. 1933. A revision of the genera of fossil and Amer. Mus. Nat. Hist., vol. 95, pp. recent Termopsinae (Isoptera). Univ. 37-86. California Publ. Ent., vol. 6, pp. 1958. Key to the Indomalayan termites. Bio- 165-196. logia (Lahore), vol. 4, pp. 33-198, i-xii. 1950. Five new genera of termites from South Araujo, R. L. America and Madagascar (Isoptera, 1970. Termites of the Neotropical region. In Rhinotermitidae, Termitidae). Amer. Krishna, K., and Frances M. Weesner, Mus. Novitates, no. 1444, pp. 1-15. Biology of termites. New York and 1952. The Neotropical genera Procornitermes London, Academic Press, vol. 2, pp. and Cornitermes (Isoptera, Termitidae). 527-576. Bull. Amer. Mus. Nat. Hist., vol. 99, pp. 1972. Notes on the geographical distribution 475-540. of Serritermes (Isoptera). Rev. Brasi- 1955. Geographical origins and dispersions of leira Ent., vol. 19, no. 9, pp. 67-70. termite genera. Fieldiana: Zool., vol. Armbruster, L. 37, pp. 465-521. 1941. Ober Insektenstaaten der Vorwelt. 1. 1961. Vestigial characters of termites and Miocine Randeker Termiten. Arch. f. processes of regressive evolution. Evolu- Bienenkunde. Leipzig und Berlin, vol. tion, vol. 15, pp. 115-131. 22, pp. 343. 1965. A review of the Mastotermitidae (Isop- Bates, H. W. tera), including a new fossil genus 1855. Proceedings of Natural-History Collec- from Brazil. Amer. Mus. Novitates, no. tors in Foreign Countries [Communi- 2236, pp. 1-46. cated by Mr. S. (Samuel) Stevens]. 1968a. Cretaceous insects from Labrador 3. A Zoologist, vol. 13, ser. 1, pp. new genus and species of termite (Isop- 45494553. tera: Hodotermitidae). Psyche, vol. 74, 1863. The naturalist on the River Amazons. pp. 276-289. London, John Murray, 2 vols. 1968b. A revision of the fossil genus Ulmeriella 1892. The naturalist on the River Amazons. (Isoptera, Hodotermitidae, Hodoter- London, John Murray, 2 vols. (reprint). mitinae). Amer. Mus. Novitates, no. Bathellier, J. 2332, pp. 1-22. 1927. Contribution 'a l'etude systematique et 1969. A revision of the Tertiary fossil species biologique des termites de l'Indochine. of the Kalotermitidae (Isoptera). Ibid., In Gruvel, A., Faune des colonies no. 2359, pp. 1-57. 30 AMERICAN MUSEUM NOVITATES NO. 2570

1971. Tertiary fossil species of the Rhinoter- mites. New York and London, Aca- mitidae (Isoptera), phylogeny of demic Press, vol. 1, pp. 1-17. genera, and reciprocal phylogeny of 1970. Taxonomy, phylogeny, and distribu- associated Flagellata (Protozoa) and the tion of termites. In Krishna, K., and Staphylinidae (Coleoptera). Bull. Amer. Frances M. Weesner, ibid. New York Mus. Nat. Hist., vol. 146, pp. 243-303. and London, Academic Press, vol. 2, Grasse, P. -P pp. 127-152. 1949. Ordre des isopteres ou termites. In Light, S. F. Grasse, P. -P., Trait' de zoologie. Paris, 1934. A world view of termites. In Kofoid, Masson et Cie., vol. 9, pp. 408-544. C. A., et al., Termites and termite Hagen, H. A. control. Second edition. Berkeley Uni- 1858a. Monographie der Termiten. Part 2. versity of California Press, chap. 10, Linnaea Ent., vol. 12, pp. 1-342, 459. pp. 108-117. 1858b. Catalogue of the specimens of neurop- Mathur, R. N., and 0. B. Chhotani terous insects in the collection of the 1959. Revision of Stylotermes Holmgren and British Museum. Part 1. Termitina. Holmgren (Isoptera: Rhinotermitidae: London, Taylor and Francis, pp. 1-34. Stylotermitinae). Zool. Anz., vol. 163, Harris, W. V. pp. 40-5 3. 1961. Termites. Their recognition and con- Noirot, C., and C. Noirot-Timothee trol. London, Longmans, Green and 1965. La glande sternale dans l'evolution des Co., 187 pp. termites. Insectes Sociaux, vol. 12, pp. 1962. Classification of the phytophagous 265-272. Isoptera. Symposia Genetica et Biologi- 1969. The digestive system. In Krishna, K., ca Italica. Atti Congresso De l'Union and Frances M. Weesner, Biology of ter- Internationale L'Etude des Insectes mites. New York and London, Aca- Sociaux, Pavia, 9-14 Settembre 1961, demic Press, vol. 1, pp. 49-88. vol. ll, pp. 193-201. Quennedey, A. Holmgren, N. 1969. Innervation de type neurosecreteur 1909. Termitenstudien. 1. Anatomische Unter- dans la glande sternale de Kalotermes suchungen. K. Svenska Vetensk.-Akad. flavicollis (Isoptera). Etude ultrastruc- Handl., vol. 44, no. 3, pp. 1-215, 76 turale. Jour. Insect Physiol., vol. 15, figs., 3 pls. pp. 1807-1814. 1910a. Das System der Termiten. Zool. Anz., 1971a. Les glandes exocrines des termites I. vol. 35, pp. 284-286. Etude histochimique et ultrastructurale 191 Ob. The Percy Sladen Trust Expedition to de la glande sternale de Kalotermes the Indian Ocean in 1905. No. 8, Isop- flavicollis Fab. (Isoptera, Kalotermiti- tera. Trans. Linnean Soc. London, dae). Zeitschr. Zellforsch., vol. 121, pp. Zool., ser. 2, vol. 14, pt. 1, pp. 27-49. 135-148. 1971 b. Les glandes exocrines des termites II. 191 la. Termitenstudien. 2. Systematik der Organisation de la glande sternale des Termiten. Die Familien Mastoter- Rhinotermitidae. Etude ultrastructurale mitidae, Protermitidae und Mesoter- preliminaire. Compt. Rendus Sceances mitidae. K. Svenska Vetensk.-Akad. Acad. Sci. (Paris), vol. 273, pp. 376-379. Handl., vol. 46, no. 6, pp. 1-88. 1972. Les glandes exocrines des termites III. 1911 b. Ceylon-Termiten. In Escherich, K., Ter- Structure fine de la glande sternale de mitenleben auf Ceylon. Jena, G. Trinervitermes geminatus Wasmann Fischer, pp. 185-212. (Termitidae, Nasutitermitinae). Zeit- Krishna, K. schr. Zellforsch., vol. 130, pp. 205-218. 1961. A generic revision and phylogenetic 1973. La glande frontale des soldats de study of the family Kalotermitidae Schedorhinotermes putorius (Isoptera): (Isoptera). Bull. Amer. Mus. Nat. Hist., Analyse chimique et fonctionnement. vol. 122, pp. 303-408. Insect Biochem., vol. 3, pp. 67-74. 1969. Introduction. In Krishna, K., and Sen-Sarma, P. K. Frances M. Weesner, Biology of ter- 1968. Phylogenetic relationship of the termite 1975 EMERSON AND KRISHNA: SERRITERMITIDAE 31

genera of the subfamily Nasutiter- Wasmann, E. mitinae (Isoptera, Termitidae). Oriental 1897. Termiten von Madagaskar und Ost- Insects, vol. 2, pt. 1, pp. 1-34. afrika. Abhandl. Senckenbergischen Silvestri, F. Nat. Gesell., vol. 21, pp. 137-182. 1901. Nota preliminare sui Termitidi sud- Weesner, Frances M. americani. Boll. Mus. Zool. Anat. 1969a. External anatomy. In Krishna, K., and Comp. Torino, vol. 16, no. 389, pp. Frances M. Weesner, Biology of ter- 1-8. mites. New York and London, Aca- 1903. Contribuzione alla conoscenza dei Ter- demic Press, vol. 1, pp. 19-47. mitidi e Termitofili dell-America mer- 1969b. The reproductive system. In Krishna, idionale. Redia, vol. 1, pp. 1-234. K., and Frances M. Weesner, ibid. New 1909. Isoptera. In Michaelsen, W., and R. York and London, Academic Press, vol. Hartmeyer, Die Fauna Sudwest- 1, pp. 125-160. Australiens. Jena, G. Fischer, vol. 2, Weidner, H. pt. 17, pp. 279-314. 1955. Korperbau, Systematik und Ver- Snyder, T. E. breitung der Termiten. In Schmidt. H., 1949. Catalog of the termites (Isoptera) of Die Termiten. Leipzig, Akad. Verlagsge- the world. Smithsonian Misc. Coll., vol. sellschaft, Geest und Portig, pp. 5-81. 112, pp. 1490. Wilson, E. 0. Springhetti, A. 1971. The insect societies. Cambridge, Mass., 1963. Sulla struttura della vescicole seminali Belknap Press of Harvard Univ. Press, delle Termiti. Atti Accad. Naz. Italiana x+548 pp. Ent. Rend., vol. 11, pp. 212-219. Tu, T. 1955. The termites of China. Jour. Formosan Sci., vol. 9, no. 1, pp. 30-39.

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