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The Termite Family Serritermitidae (Isoptera) ~~~~~S.U.,Wsjx-9Uw R¶Ua-48U .Wwiy, Ypm R

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The Termite Family Serritermitidae (Isoptera) ~~~~~S.U.,Wsjx-9Uw R¶Ua-48U .Wwiy, Ypm R 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 <Family Mesotermitid6s: Bathellier, 1927, pp. Serritermes of the 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 <Genus Calotermes: Hagen, 1858a, pp. 32, 33; with numerous irregularly scattered medium- 1858b, p. 5. Wasmann, 1897, p. 150. length thin hairs (fig. 1); forewing scale with = Subgenus Serritermes Wasmann, 1897, p. 150. = Genus Serritermes: Silvestri, 1901, p. 3; 1903, hairs on costal margin; wings with no hairs p. 40. Desneux, 1904, p. 47. Holmgren, 1909, beyond humeral suture on margins, veins, or pp. 91, 109-111,169,170; 1911 a, pp. 62,63, membranes; sternites and tergites each with short 82, 83. Bathellier, 1927, p. 130. Emerson, hairs and a marginal row of bristles hardly longer 1928, p. 406; 1952, p. 508; 1955, p. 515. than hairs, hairs near tip of abdomen longer; Armbruster, 1941, p. 20. Snyder, 1949, p. pleural membrane covered with short hairs. 100. Ahmad, 1950, p. 74; 1958, p. 35. Harris, Head egg-shaped from above (fig. 1). Fonta- 1961, p. 64. Weesner, 1969a, pp. 27, 28. nelle in small depression, white, moderate-sized, Krishna, 1970, p. 137. Wilson, 1971, p. 105. forked, with somewhat thickened arms. Y-suture Araujo, 1972, pp. 67-70. vestigial, with short arms from fontanelle visible, Type Species. Galotermes (Serritermes) ser- very faint stem connecting with more distinct rifer (Bates) (Wasmann; 1897, p. 150), a syno- short stem at rear. Frontal gland beneath fonta- nym of Serritermes serrifer (Hagen and Bates), nelle possibly present but small, not seen through only included species in the genus. head chitin. Two muscle insertions in front of Diagnostic Characters. The same as those of ocelli, each smaller than ocellus. Eye propor- the family Serritermitidae. tionally moderately large, round, about a fourth of its diameter from lower margin of head, Serritermes serrifer (Hagen and Bates) prominent from above with convex margin. Ocellus moderately large, subtriangular, facing Termes serrifer Bates In Hagen, 1858a, p. 73 (field notes). toward side and front, much less than width Calotermes serrifer Hagen, 1858a, p. 72 (soldier, from eye. Antenna with 13 or 14 articles, third nymph), pl. 1, fig. 6 (soldier); 1858b, p. 11 either smallest or equal to second or fourth. (soldier, nymph). Antennal fossa larger than ocellus, less than Calotermes (Serritermes) serrifer: Wasmann, width from eye. Mandibles probably close to 1 897, p. 1 50 (systematics). those of nymph or worker (fig. 2). Postclypeus Serritermes serrifer: Silvestri, 1901, p. 3; 1903, large, not quite so long as half its width, with p. 41 (imago, soldier, worker), p. 112 (biol- median longitudinal line, profile convex, with ogy), pi. 1, figs. 39, 40, 43 (imago), figs. 41, small but distinct angle at junction with front of 42 (soldier).
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