A GENERIC REVISION--AND PHYLOGENETIC STUDY OF THE FAMILY KALOTER- MITIDAE (ISOPTERA)
KUMAR KRISHNA
BUL-LETIN OF THE, AMERICAN MUSEUM OF NATURAL HISTORY VOLUM'E 122 ARTICLE 4 NEWV. YORK: 1961
A GENERIC REVISION AND PHYLOGENETIC STUDY OF THE FAMILY KALOTERMITIDAE (ISOPTERA)
A GENERIC REVISION AND PHYLOGE- NETIC STUDY OF THE FAMILY KALOTERMITIDAE (ISOPTERA)
KUMAR KRISHNA Department of Zoology The University of Chicago
THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY AT THE UNIVERSITY OF CHICAGO
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 122 : ARTICLE 4 NEW YORK :1961 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 122, article 4, pages 303-408, figures 1-81, tables 1-6
Issued September 15, 1961
Price: $1.50 a copy CONTENTS INTRODUCTION. 309
Acknowledgments...... 310 . Material and Technique ...... 310 . Terminology 311
SYSTEMATIC REViSIONS.IN...... 312 .
Family Kalotermitidae Banks, 1919...... 312 . Key to the Genera of the Family Kalotermitidae . 315
Genus Proelectrotermes von Rosen, 1913 ...... 317 .
Genus Electrotermes von Rosen, 1913...... 318 .
Postelectrotermes, New Genus...... 319 .
Genus Neotermes Holmgren, 1911 ...... * ...... 321 Genus Rugitermes Holmgren, 1911. 325
Genus Eucryptotermes Holmgren, 1911...... 328
? Genus Prokalotermes Emerson, 1933 ...... 331 . Genus Kalotermes Hagen, 1853 ...... 331 .
Genus Paraneotermes Light, 1934 ...... 336 .
Ceratokalotermes, New Genus...... 338.
Comatermes, New Genus...... 341 .
Genus Glyptotermes Froggatt, 1896 ...... 343 .
Genus Calcaritermes Snyder, 1925...... 348 .
Genus Pterotermes Holmgren, 1911...... 349 .
Incisitermes, New Genus...... 353 .
Genus Allotermes Wasmann, 1910...... 358 .
Marginitermes, New Genus...... 358 .
...... Tauritermes, New Genus ...... 361 .
Genus Proneotermes Holmgren, 1911...... 363 .
Bifiditermes, New Genus...... 365 .
Bicornitermes, New Genus ...... 370 .
Genus Epicalotermes Silvestri, 1918 ...... 374 .
Genus Procryptotermes Holmgren, 1910...... 376 .
Genus Cryptotermes Banks, 1906 ...... 379 .
PHYLOGENY...... 383 .
DISCUSSION...... 389
SUMMARY...... 400 .
BIBLIOGRAPHY...... 400 . 307
INTRODUCTION THE FAMILY Kalotermitidae consists of 353 the traditional classical practice of weighting living and fossil species, grouped under 24 characters. They argue that the procedure of genera in the present revision. The family is assigning significance to either adaptive or primitive in its morphology, nesting behavior, non-adaptive characters brings in subjectiv- and social organization. The species found ity in classification. They have proposed what and maintain their colonies strictly in wood, they consider to be a more objective, quanti- without necessary soil connections, often in tative approach in grouping 97 species of sound damp or dry wood. They do not con- megachilid bees. Their basic procedure con- struct definite nests, as do the higher ter- sisted of measuring correlations in 122 char- mites, but instead excavate an irregular acters weighted equally. They thus arrived at system of galleries with constructed parti- a classification by grouping species according tions. Because of such an ecology, they have to the extent to which the characters are escaped competition with higher termites. correlated. They claim that the resulting The Kalotermitidae have intestinal flagellates classification is closely similar to the one pre- but do not have any staphylinid beetles as viously established by classical methods. guests. Inger (1958) attacked the quantitative The family Kalotermitidae consists of a methodology of Michener and Sokal and number of living genera, which have existed strongly objected to their practice of giving presumably since Cretaceous times or earlier, equal weight to all characters. He states that even though they have given rise to phylo- their method has no advantage over the more genetic series of advanced genera. Also, the traditional taxonomic approaches, as their transitional forms in the phylogenetic se- technique also rests on a subjective base. quence have persisted to a remarkable degree Emerson (in Schmidt and Emerson, 1960) to the present time, so that we have "living states: "The tendency to give equal impor- fossils" or connecting links between many tance to all comparative characters of organ- genera. Because of the presence of such isms often leads to bias and to errors of judg- transitional forms, it is possible to theorize ment. Because the order of taxonomic classi- with considerable assurance about the deriva- fication is based upon homology through tion of a particular living genus from another descent, principles of evolution are both living genus. For these reasons, the family is substantiated by taxonomic data, and in exceptionally satisfactory for the analysis of turn, may be used for taxonomic interpreta- evolutionary history. tion. For example, it is possible to distinguish The objectives of this study are: (1) the homology from analogy and both from fortui- regrouping of the formerly classified species tous resemblance, strongly adaptive from into several genera consistent with the mod- weakly adaptive or nonadaptive characters, ern concept of the genus unfolded by the primitive from derivative characters, rapidly studies of the imago mandibles and their asso- evolving from slowly evolving characters, ciated structural patterns in all castes and persistent conservative characters from labile redescription and illustration of the various changing characters, progressively functional new and old genera; (2) the formulation of a characters from regressed vestiges, relatively hypothetical phylogeny; and (3) a synthesis complex from relatively simple characters, of taxonomic, phylogenetic, and biogeograph- and individually acquired characters from ical data, in an attempt to elucidate the hereditary characters." All characters are not evolutionary history of the group. equivalent in their genetic control and in their The basic method of taxonomy is to com- development and ecological effects, and there- pare and evaluate the characteristics of the fore cannot be treated equally (Inger, 1958). structure by morphological comparisons and A competent taxonomist after long training interpret them in the light of comparative and experience can determine which charac- genetics, biochemistry, embryology, behav- ters are constant, which are variable, and ior, ecology, and geography. which are more important for generic classifi- Michener and Sokal (1957) have criticized cation. That character weighting has been 309 310 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 used injudiciously by poor taxonomists is not different castes with the same basic genetic sufficient reason for rejecting the method. system corrects the errors due to the confu- Stroud (1953) used the technique of multi- sion of convergence with homology and pro- ple factor analysis in the evaluation of char- vides excellent data for generic classification acters, the elucidation of evolutionary proc- and phylogenetic analysis. In the present esses by detection of adaptive trends, the study, I have conformed to the traditional detection of phylogenies, and the definition of approach of character weighting in defining categories. By this quantitative technique he categories. I have studied a number of analyzed correlations of 14 characters for morphological characters-wing venation, soldiers of 48 species and imagoes of 43 species imago-nymph mandible, arolium, eye size, of the genus Kalotermes. The application of pigmentation, pilosity, tibial spines, and multiple factor analysis as used by Stroud in other characters in the imago, and head establishing categories has limitations, be- shape, size, and pronotum in the soldier- cause in his study of the genus Kalotermes he and have defined genera based on a constella- selected characters in such a way that homol- tion of conservative, adaptive, and regressed ogous dimensions of soldier and imago characters. anatomy could be measured. He limited him- The imago mandible is a conservative self to only 14 characters found in both character which has been found valuable in imagoes and soldiers and thus omitted impor- generic classification. In a number of super- tant characters such as wing venation, imago- generic groups within the family Kalotermiti- nymph mandible, arolium, anterior margin of dae, the genera show no differences in the soldier pronotum, and tibial spines. The imago mandible. Yet it has been possible to statistical approach used by Stroud failed to recognize genera by other sets of characters, correct the errors resulting from earlier in- both in the imago and the soldier caste. An vestigations. The phylogenetic analysis used animal can retain primitive characters and at in the present paper conforms more closely to the same time have obvious derivative modi- classical methods than did that of Stroud fications. Different portions of the same (1953). The method of multiple factor analy- individual may show conservative characters sis used by Stroud when applied to the with little evolutionary change and also have categories distinguished by the method used more rapidly evolving adaptive specializa- in this paper may indicate more subtle rela- tions. tionships and correlations of characters here ACKNOWLEDGMENTS treated as independent units. (Also see under I am greatly indebted to Dr. Alfred E. the genus Kalotermes.) Emerson for suggesting the problem and Inger (1958) suggests a biological approach taking keen interest in the progress of the to the delimitation of genera. He outlines a work. My wife, Valerie Krishna, has pro- method for establishing genera on the basis of vided editorial assistance. This work was done adaptive characters. He thinks that the use of during tenures of a University Fellowship, adaptive characters has more advantages John M. Prather Fellowship, and Chicago than the use of non-adaptive conservative Natural History Museum Fellowship of the characters. Many errors have been made University of Chicago. because of convergence, when adaptive trends The author is also grateful to Mr. Rupert were used as the basis for classification. Also, Wenzel of the Chicago Natural History many characters of importance in the classifi- Museum and to Mr. Victor Harris of the cation of genera and higher taxonomic cate- British Museum (Natural History), London, gories do not have known functions. for the loan of specimens. In termites, we have the soldier caste, which is adaptive for defense, and the imago, MATERIAL AND TECHNIQUE a more conservative reproductive caste. The This study is based primarily on the ter- ability to contrast slowly evolving, relatively mites in the collection of the American conservative, weakly adaptive characters Museum of Natural History, now in the with strongly adaptive characters in the custody of Alfred E. Emerson, the University 1961 KRISHNA: KALOTERMITIDAE 311 of Chicago. The majority of species were TERMINOLOGY represented by primary types. A total of 344 species was studied in alcohol The morphological terms and measure- under a binocular dissecting microscope. The ments used in this paper are explained in imago-nymph mandibles were dissected. The Emerson (1945, 1952b) and Ahmad (1950). In drawings were made by two methods. Most this paper, the first plus second left marginal were drawn with the help of a camera lucida. tooth and third left marginal tooth are equal The drawings of wings were made with a to the first left marginal tooth and second left microprojector. All drawings are of the type marginal tooth, respectively, as referred to by species of the different genera. Ahmad (1950). SYSTEMATIC REVISION FAMILY KALOTERMITIDAE BANKS, 1919 >Genus Calotermes HOLMGREN, 1911d, p. 209. > sent (fig. 21) .3 ...... Paraneotermes 2. Left imago-nymph mandible with anterior Left imago-nymph mandible with no dent in margin of third marginal tooth longer than posterior.... margin of first plus second mar- one and one-half times the length of the ginal tooth, evenly rounded; base of first posterior margin of first plus second mar- plus second marginal tooth equal to base of ginal tooth (fig. 70) . . . . Epicalotermes third marginal tooth (fig. 28) ...... Left imago-nymph mandible with anterior ...... Ceratokalotermes margin of third marginal tooth clearly 12. Radial sector without branches; media run- longer but usually not more than one and ning close to radial sector to tip of wing one-half times longer than the posterior (fig. 36) . . . Glyptotermes, Calcaritermes margin of first plus second marginal tooth Radial sector with branches ...... 13 (figs. 63, 67) . Bifiditermes, Bicornitermes 13. Media very small, coalescing with radial sec- 3. Middle tibia with three apical spurs and addi- tor very close to suture (fig. 13) . tional spines (figs. 2, 4, 6) ...... 4 ...... Rugitermes Middle tibia with three apical spurs only, Media running close and parallel to radial additional spines absent ...... 6 sector to tip of wing (figs. 9, 17) . . . 14 4. Middle tibia with two outer spines and one 14. Anterior margin of forewing scale convex (fig. inner spine; fossil only (fig. 2). 9); eye diameter, 0.46-0.78 mm. ...... Proelectrotermes ...... Neotermes Middle tibia with one or two outer spines, Anterior margin of forewing scale almost inner spine absent ...... 5 straight (fig. 17); eye diameter, 0.43 mm. 5. Middle tibia with two outer spines; fossil only ...... Eucryptotermes. (fig. 4) ...... Electrotermes 15. Media slightly sclerotized and running closer Middle tibia with one outer spine; living (fig. to radial sector than to cubitus (fig. 60) 6) ...... Postkectrotermes ...... Proneotermes 6. Left imago-nymph mandible with anterior Media.... not sclerotized and running midway margin of third marginal tooth equal to the between radial sector and cubitus (figs. 47, posterior margin of first plus second mar- 54, 75) ...... 16 ginal tooth (fig. 20) ...... 7 16. Media bending up and joining radial sector in Left imago-nymph mandible with anterior middle or beyond middle of wing (figs. 75, margin of third marginal tooth clearly 79) . . . . Procryptotermes, Cryptotermes longer than posterior margin of first plus Media running clear to tip of wing (figs. 47, second marginal tooth (fig. 46) . . . . 15 54) ...... 17 7. Forewing with media unsclerotized, weak, and 17. Arolium always absent ...... running midway between radial sector and ...... Marginitermes, Aliotermes cubitus to tip of wing (fig. 21) . . . . .8 Arolium usually present; occasionally absent Forewing with media sclerotized (figs. 5, 9, 13, ...... Incisitermes, Tauritermes 25) ...... 9 8. Wings dark, with pimple-like nodules (fig. 21); BASED ON THE CHARACTERS OF THE SOLDIER arolium present...... Kalotrmes 1. Middle tibia with one outer spine . Wings hyaline with no nodules (fig. 43); aroli- ...... Postekectrotermes um absent ...... Pterotermes Middle tibia with no spine ...... 2 9. Forewing with media sclerotized, but not so 2. Front tibia with a thick conspicuous spur near strongly as radial sector (figs. 25, 29, 32) apex of outer side, much larger than two ...... 10 apical spurs (fig. 40) . . . Calcaritermes Forewing with media as strongly sclerotized Front tibia with an apical spur not enlarged as radial sector (figs. 9, 13, 17, 36) . . 12 ...... 3. 10. Head and pronotum densely covered with 3. Head short and highly phragmotic (figs. 18, very long and wavy hairs (fig. 31) . 69, 76, 80) ...... 4 ...... Comatermes Head long or short, not phragmotic, sloping in Head and pronotum covered with short bris- front at an approximate angle of 20 to 45 tles; long hairs absent ...... 11 degrees; with or without anterolateral 11. Left imago-nymph mandible with distinct prominences (figs. 10, 14, 22, 26, 30, 33, 37, dent in posterior margin of first plus second 48, 55, 61, 65) ...... 8 marginal tooth; base of first plus second 4. Anterior margin of pronotum serrated (figs. 18, marginal tooth slightly larger than base of 69, 80) ...... 5 1961 KRISHNA: KALOTERMITIDAE 317 Anterior margin of pronotum even and not ser- five to seven segments together . . . . 15 rated (fig. 76) ...... 7 15. Head dorsoventrally flat (figs. 48, 72) . . 16 5. Anterior margin of pronotum sharply serrated; Head thick (figs. 30, 51, 56, 61) . . . . 17 head bilobed, with region between lobes 16. Head sloping anteriorly from near middle; depressed; ridge between vertex and frons mandibles long in proportion to rest of head absent (fig. 69) ...... Bicornitermes capsule and strongly curved (fig. 72); femur Anterior margin of pronotum wavy, weakly not swollen ...... Epicalotermes or finely serrated; ridge between frons and Head not sloping anteriorly from near middle; vertex present (figs. 18, 80) ...... 6 mandibles short in proportion to rest of 6. Head in front strongly scooped out; antennal head capsule (figs. 48, 49); femur strongly socket at one-third to half of the length of swollen ...... Incisitermes head from front (fig. 18) . Eucryptotermes 17. Head with a horn-like projection below and in Head in front vertical and not so strongly front of antennal socket (fig. 51) .... scooped out; antennal socket close to side ...... Allotermes base of mandibles (fig. 80) . Cryptotermes Head without horn-like projection . . . 18 7. Ridge between vertex and frons distinct; head 18. Head with distinct anterolateral prominences in front not vertical; third joint of antenna (figs. 30, 56) ...... 19 large and club-shaped (fig. 76). Head with indistinct anterolateral promi- ...... Procryptotermes ....nences...... 20 Ridge between vertex and frons absent; head 19. Third segment of antenna modified, longer in front vertical; third joint of antenna not and darker than fourth; anterior margin of large ...... Glyptoter- pronotum distinctly wavy (fig. 56) . . . mes (some spp.), Kalotermes (some spp.) ...... Tauritermes 8. Anterior margin of pronotum not deeply con- Third segment of antenna not modified, equal cave or incised ...... 9 to fourth; anterior margin of pronotum Anterior margin of pronotum deeply concave faintly wavy (fig. 30) . . Ceratokalotermes or incised ...... 13 20. Femur swollen; head in front dark; faint ridge 9. Head with a prominent ridge in front formed between vertex and frons (fig. 61). by a continuation of the dorsal antennal ...... Proneotermes margin medially (fig. 14) . Rugitermes Femur not swollen; ridge between vertex and Head without the prominent ridge . . 10 frons absent (fig. 65) . . . . Bifiditermes 10. Head usually with distinctanterolateral promi- GENUS PROELECTROTERMES VON ROSEN, 1913 nences, sometimes phragmotic (fig. 37) ...... Glyptotermes ...... Kalotermes, Neotermes p. 331. 12. Postmentum very broad in front (fig. 26); = Genus Proelectrotermes EMERSON, 1942, pp. nymph mandible as in figure 24 9, 10. ...... Paraneotermes = Genus Proelectrotermes SNYDER, 1949, p. 356. Postmentum not very broad in front (fig. 33); = Genus Proelectrotermes GRASSE, 1949, p. 530. head slope in front concave (fig. 33) . = Genus Proelectrotermes WEIDNER, 1955a, pp. ...... Comatermes 46, 68. 13. Sides of head distinctly and clearly rounded = Genus Proelectrotermes WEIDNER, 1955b, pp. (fig. 44); sclerotized swelling at base of post- 64, 65, 70. mentum ...... Pterotermes =Genus Proelectrotermes EMERSON, 1955, p. Sides of head almost straight; no swelling at 507. base of postmentum ...... 14 14. Third segment of antenna very large, as long as TYPE SPECIES: Proelectrotermes berendtii next five to seven segments together; head (Pictet). with a ridge between vertex and frons (fig. The type species of this genus, Proelectro- 55) .Marginitermes termes berendtii, was described by Pictet in Third segment of antenna not so long as next 1854, and the holotype specimen was depos- 318 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 1. Forewing of Proelectrotermes berendtii (Pictet). Baltic amber. ited in the collection of Berendt in Danzig. communication). Recently, I have discovered Weidner (1955b) states that at present its a specimen of Proelectrotermes berendtii Pictet depository is unknown, and perhaps the type in the Kohlman collection of arthropods in is lost. In addition to the type specimen, Baltic amber acquired by the Chicago Nat- Hagen had two additional specimens at that ural History Museum in 1953 (Wenzel, time which were probably deposited in the 1953). If the type specimens and the other Munich Museum. Emerson in 1926 studied specimens are lost, as suspected, then this is two specimens in the Munich Museum, one the only existing specimen of Proelectrotermes labeled "Termes berendtii Pict. (? belongs to berebdtii (Pictet), and I am tentatively select- Konigsberg Museum) No. 14681, No. 61," ing it as a neotype. the other, "K7808, collected by Dr. Klebs." IMAGO: Color generally dark, probably Weidner states that these specimens were lost owing to preservation. Eyes large; diameter, during the Second World War (personal 0.52 mm. Ocellus about 0.03 mm. from eye. Antenna with 19 to 20 articles. Pronotum wider than head; anterior margin concave. Forewing with all major veins arising inde- pendently at wing suture; radius (R1) simple; media weak, running slightly closer to radial sector than to cubitus; cubitus weak (fig. 1). Tibial spurs 3:3:3. Fore tibia with no addi- tional spines; middle tibia with two spines on outer edge and one spine on inner edge near the apical spur (fig. 2); hind tibia with one 3 inner spine. Arolium present. F; 3 L SPECIES INCLUDED Proelectrotermes berendtii (Pictet), 1854 GEOGRAPHICAL DISTRIBUTION: Baltic am- ber of East Prussia. GEOLOGICAL DISTRIBUTION: Upper Eo- cene. GENUS ELECTROTERMES VON ROSEN, 1913 FIG. 3. Forewing of Electrotermes aftinis (Hagen). Baltic amber. < Genus Calotermes HAGEN, 1858b, p. 1. FIG. 5. Forewing and hind wing of Postelectrotermes praecox (Hagen). Homotype colony from Camancha, Madeira Island. 1961 KRISHNA: KALOTERMITIDAE 321 GEOGRAPHICAL DISTRIBUTION: Indo-Ma- layan: Ceylon, Pakistan. Ethiopian: Africa. Malagasy: Madagascar, Reunion. Palearctic: Madere Island. GENUS NEOTERMES HOLMGREN, 1911 2 0 10 FIG. 7. Imago of Neotermes castaneus (Burmeister). A. Head and pronotum from above. B. Head from side. Punta de San Juan, Santa Clara Province, Cuba. 1961 KRISHNA: KALOTERMITIDAE 323 FIG. 8. Mandibles of imago of Neotermes castaneus (Burmeister). Pine Crest, Monroe County, Florida. placed in the genus Rugitermes, by Snyder bles as in the genus Kalotermes, i.e., left (1949). In the present paper I am also exclud- mandible with posterior margin of first plus ing the following species from the genus second marginal tooth equal to anterior Neotermes that were formerly included by margin of third marginal tooth (fig. 8). Right Snyder (1949): N. amplus, N. gracilignathus, mandible with posterior margin of second N. perfectus, N. howa, N. militaris, N. pishi- marginal tooth subequal to molar plate (fig. nensis, N. praecox, and N. semilunaris. 8). Pronotum slightly narrower, as broad as, IMAGO (FIG. 7): Color ranging from light or broader than head. Forewing with all castaneous brown to dark brown. Eyes large; major veins arising independently at wing diameter, 0.46-0.78 mm. Ocellus touching eye suture; radial sector with variable number of or approximately 0.01 mm.-0.10 mm. from branches; media as strongly sclerotized as eye. Antenna with 12 to 21 articles. Mandi- radial sector and running close and parallel to FIG. 9. Forewing and hind wing of Neotermes castaneus (Burmeister). Punta de San Juan, Santa Clara Province, Cuba. 324 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 radial sector; cubitus weak and unsclerotized third article either shorter or longer than (fig. 9). Tibial spurs 3:3:3. Arolium usually second or fourth. Mandibles long or short, present, but absent in a few species. strong, and with variable dentition. Prono- SOLDIER (FIG. 10): Larger than that of the tum as broad as head; anterior margin usually genus Kalotermes. Head usually long, sides shallowly concave or widely emarginate. parallel; frons faintly depressed in middle, Femur in some species swollen. Tibial spurs sloping less than 45 degrees; in some spe- 3:3:3. cies antennal carinae large and projecting. COMPARISONS: The imago closely resembles Eyes usually unpigmented; pigmented in a that of Postelectrotermes in wing venation, but few species. Antenna with 12 to 19 articles; differs from it in that the forewing has the 2 B 0 .)U--C FIG. 10. Soldier of Neotermes castaneus (Burmeister). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Miami, Florida. 1961 KRISHNA: KALOTERMITIDAE 325 median vein much more strongly sclerotized; N. medius Oshima, 1923 spine in the middle tibia is absent. The soldier N. meruensis (Sjostedt), 1907b of Kalotermes is smaller and usually has the N. microphthalmus Light, 1930 frons sloping more steeply. N. modestus (Silvestri), 1901 N. mona (Banks), new combination=Kalotermes mona Banks, 1919 SPECIES INCLUDED N. nigeriensis (Sjostedt), 1911 N. aburiensis Sjostedt, 1926 N. ovatus Kemner, 1931 N. agilis (Sj6stedt), 1902a N. pallidicollis (Sjostedt), 1902b N. andamanensis Snyder, 1933a N. papua (Desneux), 1905 N. araguaensis Snyder, 1959 N. paraensis da Costa Lima, 1942 N. arthur-mulleri von Rosen, 1912 N. parviscutatus Light, 1930 N. artocarpi (Haviland), 1898 N. rainbowi Hill, 1926a N. assmuthi Holmgren, 1913a N. rouxi N. and K. Holmgren, 1915 N. brevinotus Snyder, 1932 N. saleierensis Kemner, 1932b N. camerunensis (Sjostedt), 1897 N. samoanus (Holmgren), 1912 N. castaneus (Burmeister), 1839 N. sanctae-crucis Snyder, 1925c N. chilensis (Blanchard), 1851 N. sarasini N. and K. Holmgren, 1915 N. chilensis subsp. cayutuensis, Goetsch, 1933 N. schultzei Holmgren, 1911b N. chilensis subsp. zaallarensis Goetsch, 1933 N. sepulvilus Emerson, 1928 N. collarti Coaton, 1955 N. setifer Snyder, 1956 N. connexus Snyder, 1922 N. sinensis (Light), 1924 N. cryptops (Sjostedt), 1900a N. sjostedti (Desneux), 1908 N. cubanus (Snyder), new combination = Kalo- N. sonneratiae Kemner, 1932b termes cubanus Snyder, 1922 N. superans Silvestri, 1927 N. dalbergiae (Kalshoven), 1930 N. tectonae (Dammermann), 1915 N. desneuxi (Sjbstedt), 1904 N. voeltzkowi (Wasmann), 1897 N. erythraeus Silvestri, 1918 N. wagneri (Desneux), 1904c N. europae (Wasmann), 1910 N. zuluensis Holmgren, 1913b N. ferruginus Holmgren, 191lb N.firmus (Sjostedt), 1911 GEOGRAPHICAL DISTRIBUTION: Australian: N. fletcheri K. and N. Holmgren, 1917 Australia and New Zealand. Papuan: Am- N. fulvescens (Silvestri), 1901 boina, Caroline Islands, Hawaii, New Guinea N. gardneri Snyder, 1933a and adjoining islands, New Caledonia, New N. gestri Silvestri, 1912 Hebrides, Samoa, Society Islands. Indo- N. gracilidens Sj6stedt, 1925 Malayan: Andaman Islands, Celebes, Ceylon, N. grandis Light, 1930 N. grasseil Piton, 1940 south China, India, Java, Philippines, Suma- N. greeni (Desneux), 1908 tra. Ethiopian: Africa. Malagasy: Madagas- N. hirtellus (Silvestri), 1901 car, Seychelles. Nearctic: Southern North N. holmgreni Banks, 1918 America. Neotropical: Central and South N. insularis (White), 1846 America, Cocos Island, Mona Island, and the N. jouteli (Banks), new combination = Kalotermes West Indies. jouteli Banks, 1920 GEOLOGICAL DISTRIBUTION: Eocene, Re- N. kanehirae Oshima, 1917a cent. N. kartaboensis Emerson, 1925 N. ketelensis Kemner, 1932b GENUS RUGITERMES HOLMGREN, 1911 N. koshunensis Shiraki, 1909 BE E~~~~~Cdg~~~~~0 ii0 FIG. 11. Imago of Rugitermes nodulosus (Hagen). A. Head and pronotum from above. B. Head from side. Brazil. 1961 KRISHNA: KALOTERMITIDAE 327 FIG. 12. Mandibles of imago of Rugitermes nodulosus (Hagen). Brazil. simple; radial sector with seven to eight nal ridge medially, forming a prominent ridge branches; media as heavily sclerotized as in front. Eyes unpigmented. Antenna with 12 radial sector, coalescing with radial sector to 18 articles; third article in all cases longer about one-fifth to one-tenth of length of wing than second, clavate. Mandibles short and from suture, close to radial sector, with an stout, outer margin evenly rounded, with or upward bend soon after suture; cubitus weak without basal hump. Pronotum either slightly and unsclerotized; cross branches between narrower than, as broad as, or broader than radial sector and cubitus in distal end of wing head; anterior margin shallowly concave or (fig. 13). Hind wing with media absent, angular. Legs not swollen. Tibial spurs 3:3:3. present in all other genera of the family COMPARISONS: The imago can be distin- Kalotermitidae. Tibial spurs 3:3:3. Arolium guished from all other genera in having a present in all species. short, sclerotized median vein in the forewing SOLDIER (FIG. 14): Head long and narrow, which joins the radial sector very close to the sides parallel; medially depressed, only wing suture. The median vein in the hind faintly bilobed; continuation of dorsal anten- wing is absent. The soldier closely resembles FIG. 13. Forewing and hind wing of Rugitermes nodulosus (Hagen). Brazil. 328 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 C If 1I'i 0 I. FIG. 14. Soldier of Rugitermes nodulosus (Hagen). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Brazil. that of the genus Neotermes, but differs from = Subgenus Eucryptotermes EMERSON, 1928, p. it in having the third joint of the antenna 405. clavate and in every case longer than the = Genus Eucryptotermes SNYDER, 1949, P. 44. second; a prominent ridge is present in front = Genus Eucryptotermes GRASSE, 1949, p. 531. of and medial to the antennal socket. =Genus Eucryptotermes AHMAD, 1950, pp. 44, 50. SPECIES INCLUDED = Genus Eucryptotermes WEIDNER, 1955a, pp. R. athertoni (Light), 1932 48, 68. R. bicolor Emerson, 1925 =Genus Eucryptotermes EMERSON, 1955, pp. R. costaricensis Snyder, 1926a 469, 492, 508. R. flavicinctus Emerson, 1925 R. kirbyi Snyder, 1926b TYPE SPECIES: Eucryptotermes wheelerz R. magninotus Emerson, 1925 Snyder and Emerson. R. nodulosus (Hagen), 1858a R. occidentalis (Silvestri), 1901 IMAGO (FIG. 15): Uniformly brownish R. panamae (Snyder), 1925a yellow. Diameter of eye, 0.43 mm. Ocellus R. rugosus (Hagen), 1858a touching eye. Antenna with 14 to 16 articles. R. unicolor Snyder, 1952 Mandibles as in the genus Glyptotermes, i.e., left mandible with first plus second marginal GEOGRAPHICAL DISTRIBUTION: Papuan: tooth equal to third marginal tooth (fig. 16). Marquesas Islands. Neotropical: Central and Right mandible with posterior margin of South America. second marginal tooth slightly longer than GENUS EUCRYPTOTERMES HOLMGREN, 1911 molar plate (fig. 16). Pronotum narrower = Subgenus Eucryptotermes HOLMGREN, 1911a, than head. Anterior margin of forewing scale pp. 53, 55, 59-61. almost straight or very slightly convex as in 1961 KRISHNA: KALOTERMITIDAE 329 FIG. 15. Imago of Eucryptotermes wheeleri Snyder and Emerson. A. Head and pronotum from above. B. Head from side. Cotype from Blumenau, Brazil. Drawn by A. E. Emerson. the genera Glyptotermes and Calcaritermes (fig. 17). Forewing with all major veins aris- ing independently at wing suture; radius (R1) simple; radial sector with five to six branches, first branch arising at one-half of length of the wing from suture; media as strongly sclero- tized as radial sector and running very close and parallel to radial sector; cross veins between media and radial sector; cubitus FIG. 16. Mandibles of imago of Eucryptotermes weak and unsclerotized (fig. 17). Tibial spurs wheeleri Snyder and Emerson. Cotype from 3:3:3. Arolium present. Blumenau, Brazil. Drawn by A. E. Emerson. SOLDIER (FIG. 18): Head blackish and phragmotic, short and thick; sides fairly vertex and front and sides; frontal area ex- straight and parallel, somewhat constricted in tending out in front of base of mandibles; region of antenna; frontal area smooth, hol- profile of front steep; antennal socket at lowed out, with no sculpturing; a sharp about one-third or one-half of length of head circular ridge extending between frons and from front sides. Eyes present, but indistinct 330 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 17. Forewing and hind wing of Eucryptotermes wheeleri Emerson and Snyder. Cotype from Blumenau, Brazil. A FIG. 18. Soldier of Eucryptotermes wheeleri Snyder and Emerson. A. Head and pronotum from above. B. Head and pronotum from side. C. Head from below. Cotype from Blumenau, Brazil. Drawn by A. E. Emerson. 1961 KRISHNA: KALOTERMITIDAE 331 and narrow. Antenna with 11 articles; third branches at the suture, and the radial sector article shorter than second. Labrum not has seven to eight branches. The media runs visible from above, black concave front of midway between the radial sector and the head extending beyond its normal position. cubitus. There are no other distinguishing Mandibles short, broad at base, and with a characters for this genus, with the exception distinct basal hump. Postmentum short, of the three to four branches of the radius oval. Pronotum with anterior margin rising (R1). However, this character may be vari- abruptly from flatly arched posterior portion able. From the known characters of the in profile; from above, anterior margin angu- species, this genus could be synonymous with larly emarginate and strikingly serrated. Kalotermes. Until more specimens can be ex- Femur not swollen. Tibial spurs 3:3:3. amined, it is thought best to retain the pres- COMPARISONS: The wing venation and ent name with a question mark. imago mandible are similar to those of Neo- GEOGRAPHICAL DISTRIBUTION: Florissant termes. Neotermes differs, however, in having beds of Colorado. the anterior margin of the forewing scale GEOLOGICAL DISTRIBUTION: Oligocene. more convex. Otherwise, it is difficult to distinguish the imago in the two genera. The GENUS KALOTERMES HAGEN, 1853 soldier resembles that of the genus Crypto- =Genus Kalotermes HAGEN, 1853, p. 479 (no termes because of convergent evolution of the type mentioned). phragmotic head. = Genus Prokalotermes EMERSON, 1933, p. 189. B FIG. 19. Imago of Kalotermesflavicollis (Fabricius). A. Head and pronotum from above. B. Head from side. France. 334 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 similar; the soldiers are alike. Proglyptotermes should therefore be treated as a synonym of Kalotermes, as K. flavicollis is the generitype of the genus, and also bears an older name. The method of multiple factor analysis used by Stroud (1953) in defining taxonomic categories thus failed to show that Kalotermes flavicollis was related to some of the species FIG. 20. Mandibles of imago of Kalotermes included tentatively in the genus Proglypto- flavicollis (Fabricius). France. termes. The genus Proglyptotermes as treated by Stroud (1953, p. 88) is a heterogeneous group; the statistical technique also failed to as handled by Snyder (1949) and others and show this. In the present work I have reas- therefore was dealing with a large hetero- signed some of its species to other genera. geneous group. In the present study I am IMAGO (FIG. 19): Size small. Darkly pig- restricting the concept of the genus Kalo- mented. Eyes small; diameter, 0.25-0.35 mm. termes to K. flavicollis and its relatives only Ocellus in contact with eye or approximately (see list), and have reassigned most of the 0.02 mm. from eye. Antenna with 12 to 17 species to other genera, many of them new. articles. Left mandible with posterior mar- Stroud was thus not evaluating the genus gin of first plus second marginal tooth equal Kalotermes as restricted in this paper (K. to anterior margin of third marginal tooth flavicollis and its relatives) but was instead (fig. 20). Right mandible with posterior mar- examining a much larger group. gin of second marginal tooth subequal to The present study has shown that the molar plate (fig. 20). Pronotum broader than generitype species of Proglyptotermes is con- head; anterior margin concave. Anterior generic with K. flavicollis and its relatives. margin of forewing scale convex. Wing mem- The imagoes of the genus Proglyptotermes, as brane densely covered with small, pimple- are those of K. flavicollis and its relatives, are like, pigmented nodules. Region between small, darkly pigmented, and in all cases have costal margin and radial sector broad. Fore- an arolium; the imago-nymph mandibles are wing with all major veins arising independ- FIG. 21. Forewing and hind wing of Kalotermesflavicollis (Fabricius). Megiddo, Israel. 1961 KRISHNA: KALOTERMITIDAE 335 Ei 0 i4 FIG. 22. Soldier of Kalotermes flavicollis (Fabricius). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Homotype from Gennazano, Italy. ently at wing suture; short subcosta present; degrees. Kalotermes atratus (Hill) has a short, radius (R1) strong and simple; radial sector thick, truncated head, as in the genus Crypto- strongly sclerotized, with approximately termes. Eyes small, faceted, non-pigmented. eight to 12 branches; media weak, unsclero- Antenna with 11 to 18 articles; third article tized, running almost midway between radial variable, longer or shorter than second; in K. sector and cubitus, in some species joining flavicollis (Fabricius) third slightly longer distal part of radial sector and emerging than second. Mandibles strong, dentition again to join wing tip; cubitus weak and un- variable. Left mandible with two to four sclerotized, with variable number of branches marginal teeth. Right mandible with two to inner margin of wing (fig. 21). Tibial spurs marginal teeth. Pronotum as wide as or wider 3:3:3. Arolium present in all species. than head, anterior margin broadly emargi- SOLDIER (FIG. 22): Head shape variable; nate. Femur in some species slightly swollen. usually long, narrow, sides parallel; some- Tibial spurs 3:3:3. times dorsoventrally flat; frontal area de- SPECIES INCLUDED pressed medially; sloping downward to K. approximatus Snyder, 1920a mandibles at angle of approximately 40 to 45 K. atratus (Hill), 1933 336 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 6jEli FIG. 23. Imago of Paraneotermes simplicicornis (Banks). A. Head and pronotum from above. B. Head from side. Thermal, California. K. banksiae Hill, 1942 K. piacentinii' (Piton and Theobald), 1937 K. bosniaskii' Handlirsch, 1908 K. rhenanus' Hagen, 1863 K. brouni Froggatt, 1896 K. rufinotum Hill, 1925a K. capicola Coaton, 1949 K. sinaicus Kemner, 1932a K. convexus (Walker), 1853 K. tillyardi Hill, 1932b K. dispar Grasse, 1938 K. umtatae (Coaton), new combination=? Glyp- K. flavicollis (Fabricius), 1793 totrmes umtatae Coaton, 1949 K. gracilignathus (Emerson), new combination= Neotermes gracilignathus Emerson 1923 GEOGRAPHICAL DISTRIBUTION: Australian: K. hllui (Emerson), 1949 Australia and New Zealand. Papuan: Juan ? K. inamurae Oshima, 1912 Fernandez Island. Indo-Malayan: Ceylon, K. isaloensis (Cachan), new combination = Neo- Formosa. Ethiopian: Africa. Malagasy: Mad- termes isaloensis Cachan, 1949 agascar. Palearctic: Europe, Canary Islands. K. jepsoni Kemner, 1932b Nearctic: North America. K. oeningensis' von Rosen, 1913 K. pallidinotum Hill, 1942 GENUS PARANEOTERMES LIGHT, 1934 FIG. 25. Forewing and hind wing of Paraneotermes simplicicornis (Banks). Thermal, California. 338 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 C 3 3 FIG. 26. Soldier of Paraneotermes simplicicornis (Banks). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Sabino Canyon, Santa Catalina Mountains, Arizona. 15 articles; third article equal to second. GEOGRAPHICAL DISTRIBUTION: Nearctic: Mandibles short in proportion to head length; Southwestern United States and western strong, with a hump in basal region. Post- Mexico. mentum very broad in front. Pronotum as broad as head; anterior margin broadly CERATOKALOTERMES, NEW GENUS emarginate. Femur noticeably swollen. Tibial FIG. 27. Imago of Ceratokalotermes spoliator (Hill). A. Head and pronotum from above. B. Head from side. Morphotype colony from Federal Capital Terri- tory, Australia. I am erecting this genus for the single arising independently at wing suture; radius species Ceratokalotermes spoliator. Hill (1942) (R1) usually branched; radial sector with placed it in the genus Kalotermes, sensu lato, seven to eight branches, the first branch without assigning it to a subgenus. Emerson arising about one-sixth of length of wing from (1949) and Stroud (1953) included it in the suture; media as in the genera Paraneotermes genus Proglyptotermes (see discussion under and Comatermes, less heavily sclerotized than the genus Kalotermes). IMAGO (FIG. 27): Dark brown. Eyes small; diameter, 0.25 mm. Ocellus touching eye. Antenna with 13 to 14 articles. Mandibles as in the genus Kalotermes, i.e., left mandible with posterior margin of first plus second marginal tooth equal to anterior margin of third marginal tooth (fig. 28). Right mandi- ble with molar plate subequal to posterior margin of second marginal tooth (fig. 28). FIG. 28. Mandibles of imago of Ceratokalotermes Pronotum wider than head; anterior margin spoliator (Hill). Morphotype colony from Federal concave. Forewing with all major veins Capital Territory, Australia. FIG. 29. Forewing and hind wing of Ceratokalotermes spoliator (Hill). Autotype colony, Australia. C A-- E FIG. 30. Soldier of Ceratokalotermes spoliator (Hill). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Morpho- type colony from Federal Capital Territory, Aus- tralia. 340 1961 KRISHNA: KALOTERMITIDAE 341 radial sector, running closer to radial sector species in the genus Neotermes. Snyder than to cubitus; a few cross branches between (1926d) described Neotermes manni, based on radial sector and media in distal third of the soldier caste alone, from Mapiri, Bolivia. wing; cubitus weak (fig. 29). Tibial spurs In the collection of A. E. Emerson there is a 3:3:3. Arolium present. homotype series with both imagoes and SOLDIER (FIG. 30): Head somewhat phrag- soldiers, collected by A. M. Adamson, from motic, narrow, sides parallel; antennal cari- El Tucuche, Trinidad, the West Indies, nae large and markedly projecting; frons determined and compared by Emerson as concave and sloping down sharply to ante- Neotermes manni. Emerson in 1957 studied clypeus; anterodorsal surface with two large the unique type of Comatermes perfectus at the prominences or lobes, with an even curve British Museum (Natural History), London, between. Eyes unpigmented. Antenna with and made a detailed description. A recent 11 to 14 articles; third article unmodified and comparison by Emerson and myself of the subequal to second. Mandibles long and associated homotype series of Neotermes slender. Pronotum as wide as head, arched manni with Comatermes perfectus shows that transversely; anterior margin raised, deeply Neotermes manni is conspecific with Coma- and angularly emarginate, with a notch in the termes perfectus and therefore should be middle. Legs short and stout. Tibial spurs treated as a synonym. 3:3:3. IMAGO (FIG. 31): Dark brown. Head and COMPARISONS: The imago most closely pronotum with remarkably long hairs, which resembles that of Paraneotermes and that of are not stiff, but thin and somewhat wavy. Comatermes. Ceratokalotermes can be distin- Eyes small; diameter, 0.29-0.31 mm. Ocellus guished from Comatermes by the absence of in contact with eye. Antenna with 14 articles. long wavy hairs on the head and pronotum. Mandibles as in the genus Kalotermes, i.e., Paraneotermes has an imago mandible with a left mandible with posterior margin of first distinct angle in the posterior margin of the plus second marginal tooth equal to anterior first plus second marginal tooth, and the margin of third marginal tooth. Pronotum arolium is absent. The soldiers of Paraneo- broader than head. Wing venation closely termes and Comatermes differ in having short resembling that of Paraneotermes and that of and strong mandibles; the anterodorsal Ceratokalotermes. Forewing with all major prominences in the head are very faint or veins arising independently at wing suture; absent; the anterior margin of the pronotum radial sector with nine to 10 branches; media is not deeply emarginate. more weakly sclerotized than radial sector, running closer to radial sector than to cubi- SPECIES INCLUDED tus; many cross branches between media and Ceratokalotermes spoliator (Hill), new combina- radial sector; cubitus weak (fig. 32). Tibial tion = Calotermes (?) spoliator Hill, 1932b spurs 3:3:3. Arolium present. SOLDIER (FIG. 33): Head usually long; GEOGRAPHICAL DISTRIBUTION: Australian: sometimes two types of soldiers are found (a Australia. long-headed and a short-headed); head dorso- COMATERMES, NEW GENUS ventrally flattened; frons depressed medially, concave in with two faint anterodorsal FIG. 32. Forewing and hind wing of Comatermes perfectus (Hagen). El Tucuche, Trinidad, the West Indies. 342 1961 KRISHNA: KALOTERMITIDAE 343 A C E 0 FIG. 33. Soldier of Comatermes perfectus (Hagen). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. El Tucuche, Trinidad, the West Indies. wavy hairs in the head and pronotum. The GENUS GLYPTOTERMES FROGGATT, 1896 imago mandible of Paraneotermes is different B I FIG. 34. Imago of Glyptotermes tuberculatus Froggatt. A. Head and pronotum from above. B. Head from side. Homotype colony from Galston, New South Wales, Australia. = Subgenus Glyptotermes HOLMGREN, 1910b, p. >Subgenus Glyptotermes SNYDER, 1925b, pp. 140. 152, 157. > FIG. 35. Mandibles of imago of Glyptotermes tuberculatus Froggatt. Homotype colony from Galston, New South Wales, Australia. = Genus Glyptotermes LIGHT, 1930, pp. 15, 16, = Genus Glyptotermes SNYDER, 1949, pp. 45, 18. 359. > .,.. FIG. 36. Forewing and hind wing of Glyptotermes canellae (Fr. Muller). Cotype from Blumenau, Brazil. Drawn by A. E. Emerson. 346 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 C 0 FIG. 37. Soldier of Glyptotermes tuberculatus Froggatt. A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Homotype colony from Galston, New South Wales, Australia. TYPE SPECIES: Glyptotermes tuberculatus brown. Diameter of eye, 0.29-0.48 mm. Froggatt. Ocellus in most species touching the eye. Holmgren (191la) designated the species Antenna with 11 to 17 articles. Left mandible Glyptotermes borneensis (Haviland) as the with posterior margin of first plus second generitype of the genus Glyptotermes. This marginal tooth equal to anterior margin of species was not mentioned by Froggatt (1896) third marginal tooth; in some species no when he named the genus Glyptotermes. The distinct notch between first plus second species he mentioned were G. tuberculatus, G. marginal tooth and third marginal tooth (fig. iridipennis, G. brevicornis, and G. eucalypti. 35). Right mandible with posterior margin of According to the Rules of Zoological Nomen- second marginal tooth slightly longer than clature (Article 30e), G. borneensis cannot be molar plate (fig. 35). Pronotum as wide as or the generitype. Snyder (1949) selected G. slightly narrower than head. Anterior margin tuberculatus as the generitype. of forewing scale almost straight or slightly IMAGO (FIG. 34): Reddish brown to dark convex. Wing membrane smoky brown in 1961 KRISHNA: KALOTERMITIDAE 347 FIG. 38. Mandibles of nymph of Calcaritermes imminens Snyder. Cotype from type colony from Cincinnati, Colombia. color, with small, pimple-like, pigmented with an anterior rim between the frons and nodules, as in the genera Kalotermes and the vertex and a horn-like projection in front, Rugitermes. Forewing with all major veins either above or below the antennal socket. arising independently at wing suture; radius The anterior margin of the pronotum is (R1) short; radial sector heavily sclerotized, serrated. running very close and parallel to costal border, without branches; media as heavily SPECIES INCLUDED sclerotized as radial sector, running very close G. almorensis Gardner, 1944 to and parallel to it; faint cross branches G. angustus Snyder, 1925a between radial sector and media, and between G. asperatus (Snyder), 1926a G. bilobatus Snyder, 1934a media and cubitus, in distal end of wing; G. borneensis (Haviland), 1898 cubitus weak (fig. 36). Tibial spurs 3:3:3. G. brevicaudatus (Haviland), 1898 Arolium always present. G. brevicornis Froggatt, 1896 SOLDIER (FIG. 37): Head more or less elon- G. buttel-reepeni Holmgren, 1914 gated; some species (G. asperatus) having G. canellae (Muller), 1873 extremely phragmotic head, with frontal area G. caudomunitus Kemner, 1932b sloping quite steeply to vertical, and in some G. ceylonicus Holmgren, 191 id species concave; either distinctly or faintly G. chapmani Light, 1930 bilobed, with V- or U-shaped depression be- G. contracticornis (Snyder), 1925b tween lobes. Eyes unpigmented. Antenna G. coorgensis K. and N. Holmgren, 1917 with 10 to 15 articles; third article shorter G. dentatus (Haviland), 1898 G. dilatatus (Bugnion and Popoff), 1910 than second and not modified. Mandibles G. eucalypti Froggatt, 1896 short and broad, with or without basal hump. G. franciae Snyder, 1958 Pronotum slightly narrower, as broad as, or G. fuscus Oshima, 1912 wider than, head; anterior margin either shal- G. guianensis Emerson, 1925 lowly or deeply emarginate or concave. G. hospitalis Emerson, 1925 Femur not swollen. Tibial spurs 3:3:3. G. ignotus Wilkinson, 1959 COMPARISONS: The imago can be distin- G. insulanus Silvestri, 1912 guished from that of all other genera except G. iridipennis Froggatt, 1896 Calcaritermes in having the radial sector with G. kawandae Wilkinson, 1954 no branches and the media heavily sclero- G. liberatus (Snyder), 1929 G. luteus Kemner, 1931 tized and running close to and parallel to the G. magsaysayi Snyder, 1958 radial sector. The soldier resembles that of G. marlatti, Snyder, 1926a the genus Cryptotermes because of convergent G. minutus Kemner, 1932b evolution of the phragmotic head. Crypto- G. montanus Kemner, 1934 termes has an extremely phragmotic head, G. neotuberculatus Hill, 1933 with the frontal area scooped or vertical and G. nevermani Snyder, 1926a 348 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 C E 0. FIG. 39. Soldier of Calcaritermes imminens Snyder. A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Cotype from type colony from Cincinnati, Colombia. G. niger Kemner, 1934 Malayan: Borneo, Ceylon, Formosa, India, G. parvulus (Sjostedt), 1907a Japan, Java, Malacca, Philippines, Sumatra. G. pellucidus Emerson, 1925 Ethiopian: Africa. Malagasy: Seychelles. G. perparvus Emerson, 1925 Neotropical: Central and South G. pinangae (Haviland), 1898 America, the G. planus Snyder, 1925b West Indies, Puerto Rico. G. posticus (Hagen), 1858a GEOLOGICAL DISTRIBUTION: Oligocene, Re- G. pubescens Snyder, 1924 cent. G. pusilus' (Heer), 1849 GENUS CALCARITERMES SNYDER, 1925 G. reticulatus Wilkinson, 1954 G. satsumensis (Matsumura), 1907 = Genus Calcaritermes SNYDER, 1949, p. 52. = Genus Calcaritermes GRASSE, 1949, p. 531. = Genus Calcaritermes AHMAD, 1950, pp. 44, 50. = Genus Calcaritermes WEIDNER, 1955a, pp. 48, 68. = Genus Calcaritermes EMERSON, 1955, pp. 472, 490, 492, 498, 508. TYPE SPECIES: Calcaritermes imminens Snyder. IMAGO: Dark brown. Eyes small; diameter, 0.29-0.34 mm. Ocellus touching eye in most species. Antenna with 13 to 14 articles. Left mandible with posterior margin of first plus second marginal tooth equal to anterior mar- gin of third marginal tooth; in most species, notch between first plus second and third marginal tooth absent; posterior margin of first plus second merging with anterior mar- gin of third, forming a concave cutting edge (fig. 38). Right mandible with posterior margin of second marginal tooth slightly longer than molar plate (fig. 38). Pronotum FIG. 40. Left foreleg of Calcaritermes imminens slightly narrower than head. Wing venation Snyder. Cotype from type colony from Cincin- similar to that of the genus Glyptotermes. nati, Colombia. Tibial spurs 3:3:3. Arolium present in all species. SPECIES INCLUDED SOLDIER (FIG. 39): Head dark, thick, and C. brevicollis (Banks), 1918 semicylindrical; distinctly bilobed, with V- or C. emarginicollis (Snyder), 1925a U-shaped depression between lobes; front C. fairchildi Snyder, 1926a profile steep, varying from greater than verti- C. guatemalae Snyder, 1926a cal to less than vertical, in some species with C. imminens Snyder, 1925b C. nearcticus Snyder, 1933b dorsal rim overhanging, or concave between C. nigriceps (Emerson), 1925 upper rim and clypeus; in some species a C. parvinotus Light, 1933 horn-like projection of ventral genae be- C. recessifrons Snyder, 1925b tween mandible base and below antennal C. temnocephalus (Silvestri), 1901 socket. Eyes either distinct or indistinct and unpigmented. Antenna with 10 to 12 articles; GEOGRAPHICAL DISTRIBUTION: Nearctic: third article short and unmodified. Mandibles North America (Florida). Neotropical: Cen- usually short and broad, either with or with- tral and South America. out basal hump. Pronotum either narrower, as broad as, or broader than, head. Anterior GENUS PTEROTERMES HOLMGREN, 1911 margin shallowly angular or conspicuously B FIG. 41. Imago of Pterotermes occidentis (Walker). A. Head and pronotum from above. B. Head from side. St. Xavier Mission, Tucson, Arizona. FIG. 42. Mandibles of imago of Pterotermes occidentis (Walker). St. Xavier Mission, Tucson, Arizona. 1961 KRISHNA: KALOTERMITIDAE 351 FIG. 43. Forewing and hind wing of Pterotermes occidentis (Walker). St. Xavier Mission, Tucson, Arizona. 352 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 44. Soldier of Pterotermes occidentis (Walker). A. Head and pronotum from above. B. Head and pronotum from side. C. Postmentum from below. San Jose Island, Baja California. tooth plainly longer than posterior margin of on each side of postclypeus, as in Rugitermes. first plus second marginal tooth. Right mandi- A horn-like projection present, in front of and ble with posterior margin of second marginal below antennal socket, near side base of tooth equal to molar plate (fig. 42). Pronotum mandibles. Eyes pigmented. Antenna with 17 as broad as head. Forewing with all major articles; third article longer than second or veins arising independently at wing suture; fourth. Mandibles long and stout, toothed, subcosta long and unbranched; radial sector with a distinct basal hump. Postmentum with five to six branches, first branch arising with a sclerotized swelling in basal region. at one-fourth of length of wing from suture; Pronotum broader than head, somewhat media weak, unsclerotized, running midway saddle shaped; anterior margin covering between radial sector and cubitus, in some posterior region of head, deeply concave; specimens branching in distal end of wing; anterolateral corners raised; mesonotum and cubitus weak and unsclerotized (fig. 43). metanotum with distinct wing pads. Femur Tibial spurs 3:3:3. Arolium always absent. swollen. Tibial spurs 3:3:3. SOLDIER (FIG. 44): Largest in the family COMPARISONS: The imago closely resembles Kalotermitidae. Head very large; sides that of Incisitermes and that of Margini- rounded; frons sloping in front gradually from termes in wing venation and other characters. almost middle of head; mesial ridge present It can be distinguished from the imagoes of 1961 KRISHNA: KALOTERMITIDAE 353 these genera by the large eye size, and the FIG. 45. Imago of Incisitermes schwarzi (Banks). A. Head and pronotum from above. B. Head from side. Cotype from Paradise Key, Florida. 354 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 I am proposing this genus for species that were previously placed by Snyder (1949) and others in the genus Kalotermes. This group is distinct from the genus Kalotermes and be- longs to the Procryptotermes-Cryptotermes branch, as is indicated by the imago mandi- ble. The genus Incisitermes differs from the genus Kalotermes in the following ways: (1) the left imago mandible has the anterior FIG. 46. Mandibles of imago of Incisitermes margin of the third marginal tooth longer schwarzi (Banks). Homotype colony from Coral than the posterior margin of the first plus Gables, Florida. second marginal tooth; (2) the imago is larger, not so darkly pigmented, and has FIG. 47. Forewing and hind wing of Incisitermes schwarzi (Banks). Homotype colony from Coral Gables, Florida. 1961 KRISHNA: KALOTERMITIDAE 355 C FIG. 48. Large soldier of Incisitermes schwarzi (Banks). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Cotype from Paradise Key, Florida. plus second marginal tooth (fig. 46). Right (fig. 47). Tibial spurs 3:3:3. Arolium present mandible with molar plate subequal to pos- in all species except I. minor, I. galapagoensis, terior margin of second marginal tooth (fig. and L marginipennis. 46). Pronotum as broad as or broader than SOLDIER (FIGS. 48 AND 49): Head long, head. Wing transparent and lacking pimple- dorsoventrally flat, sides parallel, head slop- like nodules present in the genus Kalotermes. ing very slightly in front, at angle of approxi- Forewing with all major veins arising inde- mately 20 degrees; some species having, in pendently at wing suture; costal area much addition, a short-headed soldier. Eyes mostly narrower than Kalotermes; radius with three unpigmented; slightly pigmented in I. to eight branches; media weak, unsclerotized, schwarzi, I. minor, I. marginipennis, I. running midway between radial sector and snyderi, and I. taylori. Mandibles massive and cubitus; cross branches between radial sector short, compared to rest of head capsule; with and media; cubitus weak and unsclerotized or without basal hump. Antenna with 10 to 17 356 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 C /s El ,' i FIG. 49. Small soldier of Incisitermes schwarzi (Banks). A. Head and pro- notum from above. B. Head from side. C. Postmentum from below. Homo- type from Coral Gables, Florida. articles; third article longest, sclerotized, ? I. marianus (Holmgren), new combination darker, and often club-shaped, attaining its = Calotermes marianus Holmgren, 1912 maximum size in I. marginipennis. Pronotum I. marjoriae (Snyder), new combination = Kalo- as broad as or broader than head; anterior termes marjoriae Snyder, 1924 margin deeply angular, sides evenly rounded. I. mcgregori (Light), new combination = Kalo- Femur usually swollen. Tibial spurs 3:3:3. termes mcgregori Light, 1921 I. milleri (Emerson), new combination = Kaho- SPECIES INCLUDED termes milleri Emerson, 1943 I. minor (Hagen), new combination= Calotermes I. arizonensis (Snyder), new combination = Kalo- minor Hagen, 1858a termes arizonensis Snyder, 1926c I. nigritus (Snyder), new combination =Kalo- I. banksi (Snyder), new combination= Kalotermes termes nigritus Snyder, 1946 banksi Snyder, 1920b I. (Banks), new I. bequaerti (Snyder), new combination = Kalo- pacificus combination= Calo- termes bequaerti Snyder, 1929 termes pacificus Banks, 1901 I. emersoni (Light), new combination = Kalotermes I. platycephalus (Light), new combination = Kalo- emersoni Light, 1933 termes platycephalus Light, 1933 I. galapagoensis (Banks), new combination= I. repandus (Hill), new combination= Calotermes Kalotermes galapagoensis Banks, 1901 repandus Hill, 1926d I. immigrans (Snyder), new combination = Kalo- I. schwarzi (Banks), new combination = Kalo- termes immigrans Snyder, 1922 termes schwarzi Banks, 1920 I. incisus (Silvestri), new combination= Calo- I. seeversi (Snyder and Emerson), new combina- termes incisus Silvestri, 1901 tion=Kalotermes seeversi Snyder and Emerson, I. marginipennis (Latreille), new combination= 1949 Termes marginipenne Latreille, 1811-1832 I. semilunaris (N. and K. Holmgren), new com- 1961 KRISHNA: KALOTERMITIDAE 357 bination = Neotermes semilunaris N. and K. Holmgren, 1915 I. snyderi (Light), new combination=Kalotermes snyderi Light, 1933 I. tabogae (Snyder), new combination= Kalotermes tabogae Snyder, 1924 I. taylori (Light), new combination=Kalotermes taylori Light, 1930 FIG. 50. Mandibles of nymph of Allotermes I., new species, from Gibson Island paradoxus Wasmann. Homotype colony from 6 I., new species, from Nissan Island kilometers southeast of Tulear, Madagascar. GEOGRAPHICAL DISTRIBUTION: Papuan: B ) \ ~C FIG. 51. Soldier of Allotermes paradoxus Wasmann. A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Homotype from 6 kilometers southeast of Tulear, Mada- gascar. 358 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 Fiji, Guam, Hawaii, Loyalty Islands, Samoa, SOLDIER (FIG. 51): Head long, narrow; sides Nissan Island. Indo-Malayan: Philippine parallel, sloping in front at angle of approxi- Islands. Nearctic: North America. Neotropi- mately 45 degrees; ridge absent between frons cal: Central and South America, Galapagos and vertex; faint horn-like projection formed Islands, the West Indies, western Mexico. by prolongation of ventral genae, below and in front of antennal socket. Eyes distinct, in GENUS ALLOTERMES WASMANN, 1910 some species pigmented. Antenna with 10 to -Genus Allotermes WASMANN, 1910, p. 121. 12 articles, third article darker and heavily 0 3 FIG. 52. Imago of Marginitermes hubbardi (Banks). A. Head and pronotum from above. B. Head from side. Homotype from Sabino Canyon, Santa Catalina Mountains, Arizona. 360 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 53. Mandibles of imago of Marginitermes hubbardi (Banks). Homotype from Sabino Can- yon, Santa Catalina Mountains, Arizona. than molar plate (fig. 53). Pronotum slightly base. Eyes distinct, slightly pigmented. narrower than head. Wings hyaline. Forewing Antenna with 10 to 11 articles; first and with all major veins arising independently at second segments dark brownish and heavily wing suture; radial sector with eight to nine sclerotized; third segment greatly elongated, branches, first branch arising at approxi- clavate, dark-colored, and as long as next five mately one-third of length of wing from su- to seven segments together. Mandibles long, ture; media weak and unsclerotized, running slender, toothed, with a prominent basal midway between radial sector and cubitus, to hump. Pronotum as broad as head; anterior tip of wing; cubitus weak and unsclerotized margin deeply concave, with anterolateral (fig. 54). Tibial spurs 3:3:3. Arolium always corners irregularly serrated; sides evenly absent. rounded, with no median incision. Femur SOLDIER (FIG. 55): Head long, sloping in slightly swollen. Tibial spurs 3:3:3. front at angle of approximately 40 degrees; COMPARISONS: The imago so closely resem- raised ridge between frons and vertex which is bles that of the genus Incisitermes that it is thicker and more raised just above antennal difficult to separate them. The soldier differs FIG. 54. Forewing and hind wing of Marginitermes hubbardi (Banks). Homotype from Sabino Canyon, Santa Catalina Mountains, Arizona. 1961 KRISHNA: KALOTERMITIDAE 361 C FIG. 55. Soldier of Marginitermes hubbardi (Banks). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Homotype from Sabino Canyon, Santa Cata- lina Mountains, Arizona. from that of Incisitermes in having a ridge GEOGRAPHICAL DISTRIBUTION: Nearctic: between the frons and the vertex and the Western United States and western Mexico. third antennal segment greatly enlarged. Also, the anterolateral corners of the prono- TAURITERMES, NEW GENUS tum are serrated. B~~~ E 0 FIG. 56. Soldier of Tauritermes -4aurocephalus (Silvestri). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Cotype from Corumba, Mato Grasso, Brazil. 1961 KRISHNA: KALOTERMITIDAE 363 nences; horn-like projections absent in front of and above or below antennal socket; anten- nal carinae slightly prominent and rounded; a very faint ridge between frons and vertex, not so sharp as in the genus Procryptotermes. Eyes distinct and unpigmented. Antenna with 10 to 11 articles; third article wider and longer FIG. 57. Mandibles of nymph of Tauritermes than second. Mandibles slender, with marked taurocephalus (Silvestri). Cotype from Corumba, basal hump. Pronotum as broad as head; Mato Grasso, Brazil. anterior margin raised and covering posterior region of head, deeply emarginate, sinuate, B N~~~~ ~ ~~ FIG. 58. Imago of Proneotermes perezi Holmgren. A. Head and pronotum from above. B. Head from side. Homotype from near La Gloria, 7 miles south of San Jose, Costa Rica. radius (R1) simple, extending about half of above and below antennal socket absent. Eye length of wing from suture; radial sector with spots distinct, unpigmented. Antenna with 13 about four to six branches, first branch arising to 14 articles; third article longer than either at about half of length of wing from suture; fourth or second, club-shaped. Mandibles media more weakly sclerotized than radial strong, with a distinct basal hump. Pronotum sector, running parallel and closer to radial as broad as head; anterior margin slightly sector than to cubitus; a few cross veins be- raised, deeply emarginate; sides and hind tween radial sector and media. Cubitus weak and unsclerotized (fig. 60). Tibial spurs 3:3:3. Arolium present. SOLDIER (FIG. 61): Color nearly black in front of head, grading to yellowish in back. Head with parallel sides; front somewhat truncate, sloping at angle of approximately 45 degrees to clypeus; dorsolateral corners of frons slightly raised; a very faint ridge be- FIG. 59. Mandibles of imago of Proneotermes tween vertex and frons; sides above antennal perezi Holmgren. Homotype from near La Gloria, socket roughened; horn-like projections 7 miles south of San Jos6, Costa Rica. 1961 KRISHNA: KALOTERMITIDAE 365 FIG. 60. Forewing and hind wing of Proneotermes perezi Holmgren. Homotype from near La Gloria, 7 miles south of San Jos6, Costa Rica. margin fairly rounded. Tibial spurs 3:3:3. front, at an angle of approximately 20 de- Femur thick and short. grees; the faint ridge between the frons and COMPARISONS: The imago differs from that the vertex is absent. of Incisitermes and that of Marginitermes in having the forewing with the median vein SPECIES INCLUDED weakly sclerotized and running closer to the P. perezi Holmgren, 1911a radial sector than to the cubitus. This wing P. latifrons (Silvestri), new combination = C. lati- venation resembles that of Paraneotermes and frons Silvestri, 1901 that of Comatermes because of convergent GEOGRAPHICAL DISTRIBUTION: Neotropi- evolution. The imago of Procryptotermes has cal: Central and South America. the forewing with a weak and unsclerotized median vein which runs midway between BIFIDITERMES, NEW GENUS the radial sector and the cubitus in the prox- C FIG. 61. Soldier of Proneotermes perezi Holmgren. A. Head and pronotum from above. B. Head and pronotum from side. C. Postmentum from below. D. Mandibles. Homotype from near La Gloria, 7 miles south of San Jos6, Costa Rica. Drawn by A. E. Emerson. 1 1 -.4IC FIG. 62. Imago of Bifiditermes madagascariensis (Wasmann). A. Head and pro- notum from above. B. Head from side. Homotype from Marohogo, 20 kilometers east of Majunga, Madagascar. present in Kalotermes, absent in Bifiditermes. longer) than posterior margin of first plus In Ialotermes the forewing has all the major second marginal tooth (fig. 63). Right mandi- veins arising independently at the wing ble with molar plate clearly smaller than suture; in Bifiditermes the media and cubitus posterior margin of second marginal tooth arise from a common stem beyond the wing suture. The soldier of Kalotermes is much smaller, and the mandibles are not so long and robust, as in the genus Bifiditermes. The genus Procryptotermes differs in wing venation and several soldier characters. IMAGO (FIG. 62): Light brown to dark brown. Diameter of eye, 0.37-0.64 mm. Ocellus usually touching eye, but in a few species 0.01-0.03 mm. from eye. Antenna with 15 to 20 articles. Left mandible with FIG. 63. Mandibles of imago of Bifiditermes anterior margin of third marginal tooth madagascariensis (Wasmann). Type from Nossi- clearly longer (but not more than 1.5 times bU, Madagascar. 368 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 64. Forewing and hind wing of Bifiditermes madagascariensis (Wasmann). Type from Nossi-be, Madagascar. (fig. 63). Pronotum as broad as or slightly sides almost straight, curved near apex; basal broader than head. Wings hyaline, slightly hump present; dentition variable. Pronotum iridescent, usually covered with deposits of arched transversely; anterior margin deeply unpigmented papillae, in some species cov- or broadly concave; sides evenly rounded. ered with pigmented papillae, as in B. sibay- Femur not swollen. Tibial spurs 3:3:3. ensis. Forewing with costal border, subcosta, COMPARISONS: This genus is related to the radius, and radial sector strongly and heavily genera Epicalotermes and Bicornitermes. Bifid- sclerotized, arising independently at wing itermes differs in having the anterior margin suture; radial sector with five to seven of the third marginal tooth shorter than that branches; media and cubitus weak and un- of the genus Epicalotermes; otherwise the sclerotized, forking from a common stem imago is similar in wing venation and other beyond wing suture; media running midway respects. The imago of the genus Bifiditermes between radial sector and cubitus; cross is indistinguishable from that of the genus branches between radial sector and media Bicornitermes. The soldier of Bifiditermes (fig. 64). Tibial spurs 3:3:3. Arolium absent. differs from that of Epicalotermes in having a SOLDIER (FIG. 65): Head long and thick; thicker head; also, the mandibles are not so sides straight and parallel; frontal area rough, curved as in the genus Epicalotermes. Bicorni- depressed medially, moderately steep, sloping termes has a short and much more truncated downward towards anteclypeus at angle of head, the mandibles are short, and the ante- approximately 45 degrees; in some species rior margin of the pronotum is distinctly lateral-dorsal corners of frons above antennal serrated. base slightly prominent and elevated. Eyes large, either pigmented or unpigmented. SPECIES INCLUDED Antenna with 10 to 18 articles; third article B. angulatus (Wilkinson), new combination longest and clavate. Mandibles long, outer = Kalotermes angulatus Wilkinson, 1959 1961 KRISHNA: KALOTERMITIDAE 369 E 0 FIG. 65. Soldier of Bifiditermes madagascariensis (Wasmann). A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Homotype from 20 kilometers east Majunga, Madagascar Marohogo, of B. beesoni (Gardner), new combination = Kalo- B. madagascariensis (Wasmann), new combination termes beesoni Gardner, 1944 = Calotermes madagascariensis 1897 B. condonensis = Wasmann, (Hill), new combination Calo- B. mutabae (Harris), new combination= Calo- termes (C.) condonensis Hill, 1922 termes mutabae 1948 B. durbanensis Harris, (Haviland), new combination B. pintoi (Kemner), new combination = Kalo- = Calotermes durbanensis Haviland, 1898 termes B. pintoi Kemner, 1932b indicus (Holmgren), new combination= Calo- B. sibayensis (Coaton), new combination = Kalo- termes (C.) indicus Holmgren, 1913a termes sibayensis Coaton, 1949 ? B. improbus new combination = (Hagen), Calo- B. sylvaticus (Wilkinson), new combination = Kal- termes improbus Hagen, 1858a * otermes B. sylvaticus Wilkinson, 1959 jeannelanus (Sj6stedt), new combination= Cal- B., new species, from Transvaal, South Africa otermesjeannelanus Sjostedt, 1915 B., new species, from India 370 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 66. Imago of Bicornitermes bicornis, new species. A. Head and pro- notum from above. B. Head from side. Paratype from type colony. B, new species, from 10 kilometers south of than head. Wing membrane hyaline, with Besangoa, Madagascar unpigmented, indistinct papillae. Wing vena- GEOGRAPHICAL DISTRIBUTION: Australian: tion as in the genera Epicalotermes and Australia, New Zealand. Indo-Malayan: Cey- Bifiditermes. Forewing with costal border, lon, India, Thailand. Ethiopian: Africa. Mal- subcosta, radius, and radial sector heavily agasy: Madagascar. sclerotized and arising independently at wing suture; radial sector with six or seven BICORNITERMES, NEW GENUS branches; media and cubitus weak and fork- TYPE SPECIES: Bicornitermes bicornis, new ing from a common stem beyond wing suture; species. media running midway between radial sector IMAGO (FIG. 66): Light yellowish brown. Eyes large; diameter, 0.41-0.46 mm. Ocellus touching eye. Antenna with 12 to 15 articles. Mandibles as in the genus Bifiditermes, i.e., left mandible with anterior margin of third marginal tooth clearly longer (but not more than 1.5 times longer) than posterior margin of first plus second marginal tooth (fig. 67). Right mandible with molar plate distinctly shorter than posterior margin of second FIG. 67. Mandibles of imago of Bicornitermes bi- marginal tooth (fig. 67). Pronotum narrower cornis, new species. Paratype from type colony. 1961 KRISHNA: KALOTERMITIDAE 371 and cubitus (fig. 68). Tibial spurs 3:3:3. tum. The ridge between the vertex and frons Arolium absent. is absent. SOLDIER (FIG. 69): Head dark, short, and truncate, sloping steeply in front; latero- SPECIES INCLUDED frontal area raised into prominent and B. bicornis, new species rounded lobes; region between lobes de- B. exsertifrons (Wilkinson), new combination pressed; lateral corners of frons near postclyp- = Kalotermes exsertifrons Wilkinson, 1958 eus B. spinicollis (Wilkinson), new combination drawn forward into a horizontal projec- =Kalotermes spinicollis Wilkinson, 1958 tion or ridge, linked to dorsal margin of B., new species, from Leopoldville, the antennal socket; another prominent, horn- Congo like projection in front of and below antennal GEOGRAPHICAL DISTRIBUTION: Ethiopian: socket. Eyes unpigmented. Antenna with 11 Africa. to 15 articles; third article subequal to second or fourth. Mandibles short, with a prominent Bicornitermes bicornis, new species basal hump. Pronotum narrower than, or as IMAGO (FIG. 66): Head yellowish brown, broad as, head; anterior margin raised, ser- lighter behind eyes; ocellus white; eyes black; rated, and deeply emarginate. postclypeus smoky brown and darker than COMPARISONS: The imago closely resembles head; anteclypeus whitish; labrum yellow; that of the genus Bifiditermes. It can be dis- pronotum yellowish, almost same color as tinguished from that of Epicalotermes in labrum or slightly darker; tibia pale yellow; having a shorter anterior margin in the third antenna and femur brownish yellow; wing left marginal tooth. The soldier head is highly scale yellowish brown; wing membrane clear phragmotic and analogously resembles that of and transparent, with a slight brownish tinge; the genus Cryptotermes in the strong, sharp costal region yellowish brown, five or six serration of the anterior margin of the prono- branches of cubitus near wing scale with a FIG. 68. Forewing and hind wing of Bicornitermes bicornis, new species. Paratype from type colony. 372 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 C 2 0 Co FIG. 69. Soldier of Bicornitermes bicornis, new species. A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Paratype from type colony. brownish tinge; sternites brownish yellow; terior margin slightly concave in middle. tergites pale yellow. Head with a few scat- Wing membrane reticulated with indistinct tered, short bristles; pronotum with either papillae. Forewing with subcosta simple; long or short stiff bristles on the margin and radius long and unbranched; radial sector lobe; costal margin of forewing and hind wing with six to seven branches; cubitus with 13 to with short hairs. Frons sloping slightly 14 branches (fig. 68). In hind wing radial towards postclypeus. Anteclypeus with sides sector with five to six branches. Tibial spurs straight, converging anteriorly; anterior mar- 3:3:3. Arolium absent. gin straight or slightly emarginate. Eyes SOLDIER (FIG. 69): Head brown posteriorly, large, moderately convex; lower margin not grading forward to reddish brown, with far removed from lower margin of head; anterior part almost black; mandible dark anterior margin straight. Ocellus suboval, reddish brown; labrum reddish brown; prono- touching eye. Antenna with 14 to 15 articles; tum yellow, anterior raised portion brown; second article longer than third or fourth; legs and antenna light brownish yellow. third subequal to fourth. Pronotum trans- Head sparsely covered with minute bristles. versely arched, narrower than head; anterior Pronotum moderately clothed with various- margin concave, with an incision in middle; sized bristles. Head short and truncate; sides sides rounded, converging posteriorly; pos- straight up to eyes, converging anteriorly up terolateral corners broadly rounded; pos- to side base of mandibles; frontal region 1961 KRISHNA: KALOTERM ITIDAE 373 rough and wrinkled; laterofrontal area raised TABLE 2 into prominent and rounded lobes, region MEASUREMENTS (IN MILLIMETERS) OF EIGHT between lobes depressed and sloping down to SOLDIERS OF Bicornitermes bicornis, postclypeus; lateral corners of frons drawn NEW SPECIES forward into a horizontal projection, linked to dorsal margins of antennal socket; another prominent horn-like process in front and Range Mean type below antennal socket. Eyes large and unpig- mented. Antenna with 12 to 13 articles; third Length of head to article subequal to second or fourth. Mandi- side base of man- bles short, broad at base, curved towards tip, dible 1.62-1.71 1.67 1.71 with a prominent basal hump. Left mandible Maximum width of with three not very prominent marginal head 1.37-1.43 1.40 1.40 teeth, posterior margin of first marginal tooth Maximum height of head 1.22 1.22 1.22 merging with anterior margin of second mar- Length of left man- ginal tooth, forming a concave cutting edge; dible 0.70-0.73 0.73 0.73 concave cutting edge between second and Maximum width of third marginal teeth not so broad as between postmentum 0.46-0.49 0.49 0.49 first and second marginal teeth. Right mandi- Minimum width of ble with two marginal teeth; posterior margin postmentum 0.40-0.46 0.43 0.43 of first marginal tooth longer than anterior Length of postmen- margin. Labrum with sides converging anteri- tum 0.79-0.85 0.81 0.79 orly into a blunt point. Postmentum as in Median length of figure 69C. Pronotum narrower than pronotum 0.67-0.85 0.78 0.85 head, Maximum length of transversely arched; anterior margin raised pronotum 0.85-1.00 0.96 0.98 and distinctly serrated; posterolateral cor- Maximum width of ners broadly rounded, in some specimens flat- pronotum 1.28-1.37 1.33 1.28 tened. Femur not swollen. Length of hind tibia 0.98-1.10 1.05 1.10 TABLE 1 MEASUREMENTS (IN MILLIMETERS) OF EIGHT COMPARISONS: The imago of Bicornitermes IMAGOES OF Bicornitermes bicornis, bicornis, new species, is larger in all respects NEW SPECIES than that of B. spinicollis (Wilkinson) and that of B. exsertifrons (Wilkinson). The Range Mean soldier of B. spinicollis is smaller; the mandi- bles are shorter; the laterofrontal lobes are Length of head to tip of lab- not so prominent; the frontal area is not so rum 1.52-1.68 1.58 depressed medially; the eyes are smaller; the Length of head to side base postmentum shape is different; and the pro- of mandibles 1.10-1.22 1.16 notum has the corners more Width of head 1.25-1.28 1.27 posteriolateral Diameter of eye 0.47-0.59 0.53 broadly rounded. The soldier of B. exserti- Eye from lower margin 0.06-0.12 0.09 frons is smaller; the head anteriorly is reddish Length of ocellus 0.10-0.13 0.12 brown; the medial depression between the Median length of pronotum 0.61-0.70 0.65 frontal lobes is deeper and narrower; and the Maximum length of prono- anterior margin of the pronotum is more tum 0.70-0.79 0.73 deeply emarginate and has fewer serrations, Width of pronotum 1.04-1.10 1.08 which are not so prominent. Length of hind tibia 0.98-1.10 1.05 TYPE LOCALITY AND DISTRIBUTION: Camp Length of forewing from su- Putnam (latitude 10 24' N., longitude 280 36' ture 10.15-10.72 10.37 Width of forewing 2.82- 2.91 2.84 E.) on the Epulu River, the Congo (type Length of forewing scale 0.98- 1.07 1.05 locality), soldiers (holotype and paratype) and imagoes (morphotype and paratype), 374 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 collected by A. E. Emerson, April 14, 1948, in hard dry wood of standing dead tree on the edge of Epulu River. GENUS EPICALOTERMES SILVESTRI, 1918 -Genus Epicalotermes SILVESTRI, 1918, p. 347. =Genus Epicalotermes SJ6STEDT, 1926, pp. 23, 44. =Subgenus Epicalotermes EMERSON, 1928, p. 405 (no description). FIG. 70. Mandibles of nymph of Epicalotermes FIG. 71. Forewing and hind wing of Epicalotermes kempae (Wilkinson). Lake Chala, near Taveta, Kenya. 1961 KRISHNA: KALOTERMITIDAE 375 posterior margin of first plus second marginal sclerotized, arising from a common stem tooth; first plus second marginal tooth shorter beyond wing suture; media may be simple or at base than third marginal tooth (fig. 70). forked, running midway between radial sector Right mandible with molar plate shorter than and cubitus (fig. 71). Tibial spurs 3:3:3. posterior margin of second marginal tooth Arolium absent. (fig. 70). Pronotum slightly narrower than or SOLDIER (FIG. 72): Head long and broad; nearly as broad as head; anterior margin dorsoventrally flat; sides converging anteri- concave. Wing membrane clear, covered with orly; genal region below the antennal fossa hyaline papillae. Forewing with costal mar- depressed; frons sloping in front, the slope gin, subcosta, radius, and radial sector gradual and beginning at about half of length strongly developed and sclerotized and of head; frons depressed medially, concave; arising independently at wing suture; radial dorsal rim of antennal foveolae very promi- sector with five to nine branches to costal nent and dorsally appearing in form of small margin; media and cubitus weak and un- tube. Eyes large, pigmented. Labrum sub- 0\B .MV C FIG. 72. Soldier of Epicalotermes aethiopicus Silvestri. A. Head and pronotum from above. B. Head from side. C. Postmentum from below. Cotype from Mayabal, Eritrea. 376 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 rectangular, broader than long, anterior Genus Procryptotermes GRASSE, 1949, p. 530. margin convex, with a small, median, anterior I0 0 C0 FIG. 73. Imago of Procryptotermes fryeri Holmgren. A. Head and pronotum from above. B. Head from side. C. Left eye. Cotype from type colony from Takamaka, Aldabra Islands. brown. Eyes small; diameter, 0.34-0.39 mm. wing, bending up near middle or beyond Ocellus touching eye. Antenna with 16 to 18 middle of wing to meet radial sector. Cubitus articles. Left mandible with first plus second weak and unsclerotized (fig. 75). Tibial spurs marginal tooth much smaller at base than 3:3:3. Arolium present. third marginal tooth; anterior margin of third marginal tooth about one and one-half times longer than posterior margin of first plus second marginal tooth (fig. 74). Right mandi- ble with molar plate distinctly shorter than posterior margin of second marginal tooth (fig. 74). Pronotum as broad as or broader than head. Forewing with all major veins arising independently at wing suture; radius (R1) simple; radial sector with four to five branches, first branch arising at one-third of length of wing from suture; media weak and FIG. 74. Mandibles of imago of Procryptotermes unsclerotized, running midway between ra- fryeri Holmgren. Cotype from type colony from dial sector and cubitus in proximal half of Takamaka, Aldabra Islands. 378 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 FIG. 75. Forewing and hind wing of Procryptotermes fryeri Holmgren. Cotype from Takamaka, Aldabra Islands. SOLDIER (FIG. 76): Head long, sides almost COMPARISONS: The imago is similar in all parallel; front sloping at angle of approxi- respects to that of the genus Cryptotermes. mately 45 degrees to clypeus; prominent ridge The soldier differs from that of Cryptotermes (frontal flange of Hill, 1942) present between in having longer mandibles and a less phrag- frons and vertex, with a shallow, median, motic head; also the anterior margin of the U-shaped incision in P. fryeri and a few other pronotum is even, rather than serrated. species; in some species ridge less prominent or absent; anterolateral region of head with SPECIES INCLUDED two pairs of horn-like projections: one situ- P. fryeri Holmgren, 1910b ated near lateral margin of postclypeus, in P. corniceps (Snyder), 1923 front of antennal socket; the other formed by P. dioscurae Harris, 1954 a prolongation of ventral genae in front and P. krishnai Emerson (new name for the soldier below antennal socket; in some the upper or described by Hill, 1942, p. 73, under the name lower projection absent or not so prominent. Procryptotermes canalensis) 1936 Eyes unpigmented. Antenna with 11 to 16 P. rapae Light and Zimmermann, third article usually long and club- P. speiseri N. and K. Holmgren, 1915 articles; P, new species, from west of Plumb Point Light- shaped, longer than second; in P. rapae, third house, Jamaica article shorter than second. Mandible length P., new species, from Port de Galles, R6union ranging from 1.28 to 1.58 mm.; basal region Island thick and with a hump. Teeth present in most species; in some, as P. fryeri, absent. GEOGRAPHICAL DISTRIBUTION: Papuan: Pronotum as broad as head; markedly arched New Caledonia, New Hebrides, Rapa Island. transversely; anterior margin shallowly or Ethiopian: Socotra Island. Malagasy: Alda- deeply emarginate. Femur not swollen. Tibial bara Island, Reunion Island. Neotropical: spurs 3:3:3. Jamaica, Mona Island, Puerto Rico. 1961 KRISHNA: KALOTERMITIDAE 379 FIG. 76. Soldier of Procryptotermes fryeri Holmgren. A. Head and pro- notum from above. B. Head and pronotum from side. Cotype from type colony, Takamaka, Aldabra Islands. Drawn by A. E. Emerson. GENUS CRYPTOTERMES BANKS, 1906 2E 61 FIG. 77. Imago of Cryptotermes cavifrons Banks. A. Head and pronotum from above. B. Head from side. Near Strickland, Hammock, Florida. >Subgenus Planocryptotermes EMERSON, 1928, = Genus Cryptotermes Wu, 1935, p. 217. p. 405. Subgenus Cryptotermes HARE, 1937, pp. 461, =Genus Cryptotermes LIGHT, 1930, pp. 15, 16, 474, 475. 18. >Subgenus Planocryptotermes LIGHT, 1930, pp. 15, 16, 18. Genus Cryptotermes LIGHT, 1931, p. 586. FIG. 79. Forewing and hind wing of Cryptotermes cavifrons Banks. Near Strickland, Hammock, Florida. 382 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 El 'a'- FIG. 80. Soldier of Cryptotermes cavifrons Banks. A. Head and pro- notum from above. B. Head from side. C. Postmentum from below. Near Strickland, Hammock, Florida. projections lacking in some species. Eyes C. gearyi Hill, 1942 distinct, unpigmented. Mandibles short, C. havilandi (Sj6stedt), 1900b 0.38-0.95 mm. in length, humped basally, C. kirbyi Moszkowski, 1955 bent sharply near middle, weakly toothed or C. longicollis Banks, 1918 not toothed. Antenna with 11 C. merwei Fuller, 1921 to 15 articles; C. pallidus (Rambur), 1842 third article not especially long. Labrum C. perforans Kemner, 1932b pointed at tip. Pronotum either slightly C. primus Hill, 1921 narrower than or as broad as head; anterior C. queenslandis Hill, 1933 margin strongly concave, emarginate, usually C. senegalensis Silvestri, 1914 irregularly wavy or serrated; anterolateral C. solidus (Hagen), 1858a corners sharp; sides and hind margin evenly C. sumatrensis Kemner, 1930 rounded. C. verruculosus Emerson, 1925 SPECIES INCLUDED GEOGRAPHICAL DISTRIBUTION: Australian: C. albipes N. and K. Holmgren, 1915 Australia. Papuan: Hawaii, Loyalty Islands, C. brevis (Walker), 1853 Marquesas Islands, New Guinea, New Brit- C. canalensis (N. and K. Holmgren), 1915 C. cavifrons Banks, 1906 ain, Society Islands, Solomon Islands. Indo- C. cubicoceps Emerson, 1925 Malayan: Ceylon, Formosa, India, Java, C. cynocephalus Light, 1921 Philippines, Thailand, Singapore, Sumatra. C. dolei Light, 1932 Ethiopian: Africa. Malagasy: Madagascar, C. domesticus (Haviland), 1898 Mauritius. Nearctic: North America. Neo- C. dudleyi Banks, 1918 tropical: Central and South America, Gala- C. fatulus Light, 1933 pagos Islands, the West Indies. PHYLOGENY THERE ARE VARIOUS APPROACHES to the ble is equal to the anterior margin of the third study of phylogeny, namely, comparative marginal tooth. The second line is represented morphology, ecology, embryology, physiol- by the Incisitermes-Cryptotermes complex and ogy, genetics, zoogeography, and paleontol- is characterized by the elongation of the ogy. In the following pages I present the anterior margin of the third marginal tooth phylogeny of the genera of the family Kalo- and the shortening of the posterior margin of termitidae, based mainly on the comparative the first plus second tooth of the left mandi- morphology of living forms, with some sup- ble. plemental knowledge from Tertiary fossils. In the first line, Proelectrotermes, a fossil From a study of present-day geographical genus from Baltic amber of East Prussia, is distribution and ecological limitations, and a the most primitive, as is indicated by the knowledge of ancient land masses, land presence of extra spines in the middle tibia- bridges, climate, and the geological possibili- two spines on the outer edge and one spine on ties of dispersal and isolation, correlated with the inner edge. The wing venation is more clear phylogenetic stages and direction of evo- advanced; the median vein in the forewing, lution, one may find some indications of the though weak and unsclerotized, is closer to place of origin, the time of origin, and the the radial sector than to the cubitus. The direction of ancient dispersion. Emerson arolium is present. The antenna has 19 to 20 (1952a, 1955) studied the biogeography of articles. termites, using such an analysis. In the pres- The fossil genus Electrotermes from Baltic ent study I also include the biogeography of amber of East Prussia probably arose directly the genera as used by Emerson, and some of from the genus Proelectrotermes, having lost the conclusions are also taken from his study. the outer spine from the middle tibia. The Because the conclusions presented here are wing venation is similar to that of Proelectro- based on circumstantial evidence, with many termes. The arolium is present. The antenna gaps, they are therefore tentative (fig. 81 and has 15 to 18 articles. table 3). Postelectrotermes was probably derived The family Kalotermitidae probably arose directly from Electrotermes. In the middle in the remote past from a Mastotermes-like tibia only one inner spine remains, one having ancestor (Ahmad, 1950). The ancestral been lost. The wing venation is more ad- kalotermitid may be presumed to have had vanced than that of Proeiectrotermes and the following characteristics: (1) the imago Electrotermes: the media in the forewing is mandible Mastotermes-like, i.e., the marginal weakly sclerotized and has advanced closer to teeth in the left mandible reduced from three the radial sector than in the other two genera. to two by means of the reduction and fusion The genus lives in damp wood and survives of the second marginal tooth with the first; only in islands or on continental edges. The the posterior margin of the first plus second distribution of the species is: Indo-Malayan marginal tooth equal to the anterior margin (two), Ethiopian (one), Malagasy (seven, of the third marginal tooth; (2) the arolium including two new species), and Palearctic present; (3) the median vein in the forewing (two). Because of the presence of the ances- weak, unsclerotized, and running midway tral genus Electrotermes in the Baltic amber of between the radial sector and the cubitus to East Prussia and its present-day distribution, the tip of the wing; and (4) the middle tibia it is suspected that the genus originated in the with extra spines in addition to the apical Old World in Mesozoic times. The presence of spurs. a large number of species in Madagascar is From such an ancestor two major lines probably due to reduced competition. diverged. The first line is represented by Neotermes probably descended directly the Proelectrotermes-Calcaritermes complex in from Postelectrotermes, as is indicated by the which the imago mandible is like that of the morphology and ecology. Neotermes, as is ancestor, i.e., the posterior margin of the first Postelectrotermes, is a damp-wood dweller. It plus second marginal tooth of the left mandi- shows an advance over Postelectrotermes in 383 384 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 having-lost all the spines in the middle tibia, radial sector close to the wing suture. The and in having the median vein in the forewing distribution of species is: Papuan (two, in- more strongly sclerotized and closer to the cluding one new species) and Neotropical (19, radial sector. The distribution of species is: including eight new species). The distribution Australian (one), Papuan (20, including one of species indicates that the genus originated new species), Indo-Malayan (21, including in the Neotropical region in early Tertiary one new species), Nearctic (one), and Neo- times. Emerson (1955) states that the pres- tropical (24, including seven new species). ence of two endemic species in the eastern The genus is essentially tropical, except for oceanic islands (Marquesas, Society, and one species, Neotermes castaneus, which is Cook) of the Papuan Region indicates a found in subtropical regions of Florida. Its western dispersal from America, and he wide distribution indicates an origin in either thinks that they could probably have been the Neotropical or Indo-Malayan region in carried in branches of floating trees. Mesozoic times. Eucryptotermes is a highly specialized Rugitermes probably arose from a Neo- monotypic genus endemic to South America. termes-like stock and is more advanced than It probably arose from Neotermes-like stock in Neotermes. The sclerotized median vein in the South America. The wing venation is Neo- forewing, instead of running parallel to the termes-like; the soldier, however, is derivative radial sector to the tip of the wing, joins the and has developed a highly phragmotic head, TABLE 3 THE NUMBER OF LIVING SPECIES IN EACH GENUS ARRANGED BY ZOOGEOGRAPHICAL REGIONS - 0 2 c~~co~~~ c.b cl) >-.~~~~~~~~~~~0 c ~ ~,%1 V m Cd ~~ ~~~~~~~Cd~co 4-~0Q0 o ,q_.) CO Cd C00 Postelectrotermes - 2 1 7 2 - - - 12 10 2 Neotermes 1 20 21 17 6 1 24 90 73 17 Rugitermes 2 - - - 19 - 21 12 9 Eucryptotermes - - - - - 1 - 1 1 0 Kalotermes 7 1 1 2 1 3 1 - 16 16 0 Paraneotermes 1 - - 1 1 0 Ceratokalotermes 1 - - - 1 1 0 Comatermes - - -- 1 1 1 0 Glyptotermes 5 8 21 12 2 - - 25 1 74 51 23 Calcaritermes - - - - 1 13 - 14 10 4 Pterotermes - - - - 1 - - 1 1 0 Incisitermes - 8 2 1 7 16 - 34 23 11 Allotermes 3 - - - 3 1 2 Marginitermes - - - 1 - - 1 1 0 Tauritermes ------2 2 2 0 Proneotermes - - - - 2 - 2 2 0 Bifiditermes 2 - 4 7 2 - - - - 15 12 3 Bicornitermes - 4 - - - - 4 3 1 Epicalotermes - 4 1 - - - 5 4 1 Procryptotermes - 3 1 2 - - 3 - 9 5 4 Cryptotermes 6 8 5 4 2 - 1 9 1 36 24 12 Total 22 50 56 52 27 5 14 115 2 343 254 89 Y CD .0 a bo U. X. S as 5 0 2zID .X 0. c f ¢ij ON'DSN S3R3W1IHlV:Y1V *o*M3&JQI.'"fd'TN S3MIM3±OdA19 * U~~~~~~~~~~0C . u0? 00ON S3NV31VK0O ; *lV S3WU310VIOIVH30 a .0 *DON S3WH3LO3NVIVd 0~~~ 'DOl0cDWD73'IOtDCflV S3WI&l3101V 3I!JO1-410 S3WH3i0lV>I0d Vo co a - Z.E.d .00~~~Cd *OONDNd S3W3ilIn w - 09@N S3M3i1doi On3 0.~~~~~ o9OS`4'OWfWJ3'VODd'1nv S3I H3103N - C0 10DcDWI20 S3WH31081O3J31SOd - j*qwo 0Ilog S31rn31OI1.313- *qwD 0!11D8 S3WH31081O0133Od - 'DN S3WI31083Ld zD~~~~A co Co 'ON' D,Nb':'O'Dd S3WU3J.tSIONI 'DON S3WH3±1NIOl9VW 'DIW S3WPH31011V z ~~~~~~)CdC °ON 83w1H3±Lv±, >.C~ ~~~C Wic 0OON S3WI3103NOUd c~~~~0. ~ ~ ~ 0bW 00. "DW33@0fV S3MH3iatutii 00 13 S3WV311NNODI9- *Dwl'J3 S3WH3101VOd3 - ~~~*0~~~~~ ac.) 0 *~~.0O.s5i CA.0 O°SONAj3"Dd S3UH310.LdM1AO0d 0~~~~~~W0 oe0N`9N`DW13!oD, nV S3VU3LO.LdALIO 30~ ~~~ U- U 385 386 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 which is convergent with that of Crypto- that of Paraneotermes and Comatermes. The termes, Glyptotermes, Calcaritermes, and Bi- soldier head is specialized, has two antero- cornitermes. lateral prominences on the head, and closely The validity of the fossil genus Prokalo- resembles that of Glyptotermes because of termes is questionable, and its relationship is convergent evolution. obscure. From its known characters it could Glyptotermes shows an advance over Coma- be synonymous with Kalotermes. Until more termes, Paraneotermes, and Ceratokalotermes. It specimens can be examined, it is thought best probably arose from a Kalotermes-like ances- to retain the present name. tor near the base of the Comatermes complex. The genus Kalotermes probably arose The costal area is more consolidated. The directly from the ancestral kalotermitid stock median vein in the forewing is as heavily after losing all the additional spines. It has sclerotized as the radial sector and runs very retained the primitive wing venation, imago close and parallel to the radial sector; the mandible, and arolium. The distribution of radial sector has lost its branches to the the species is: Australian (seven), Papuan costal margin. The soldier has a phragmotic (one), Indo-Malayan (one), Ethiopian (two), head and is somewhat convergent with the Malagasy (one), Palearctic (seven, with four specialized head of Cryptotermes soldiers. The fossil species), and Nearctic (one). The genus distribution of the species is: Australian is found mainly on islands and continental (five), Papuan (eight, including five new edges in tropical regions of Australia and species), Indo-Malayan (21, including three Africa, and in the central continental area in new species), Ethiopian (12, including six temperate regions of Europe. The place of new species), Malagasy (two, including one origin is obscure; the world-wide distribution new species), and Neotropical (25, including of species probably indicates an early or mid- eight new species). The genus is strictly tropi- Mesozoic origin. The large number of species cal. Because of its abundance in the Neotropi- in Australia may be due to a somewhat re- cal Region, and the decline in numbers in a duced competition within its hardwood regular order in the Indo-Malayan, Ethio- ecological niche. pian, Papuan, anxd Australian regions along From the Kalotermes-like stock probably the lines of dispersion, Emerson (1955) arose three endemic genera, Paraneotermes, postulates an origin probably in the Neotropi- Comatermes, and Ceratokalotermes, in the cal Region, with late Jurassic or early Cre- Nearctic, Neotropical, and Australian regions. taceous dispersal. Paraneotermes is a monotypic endemic Calcaritermes probably was derived directly genus of temperate Mexico and southwestern from Glyptotermes. It is closely related in United States. The median vein in the fore- morphology and ecology to the genus Glypto- wing is weakly sclerotized and has shifted termes. The imago is similar to that of the slightly closer to the radial sector, as com- genus Glyptotermes. The soldier has a promi- pared to the position midway between the nent spur on the fore tibia. The head is phrag- radial sector and the cubitus in Kalotermes. motic, as in the genus Glyptotermes. The The arolium is absent. distribution of species is: Nearctic (one) and Comatermes is a specialized, monotypic, Neotropical (13, including four new species). endemic genus of the Neotropical Region. Because of its abundance and the limited The wing venation is like that of Paraneo- distribution of the species, it may be postu- termes, i.e., the media is weakly sclerotized lated that the genus originated in the Neo- and is closer to the radial sector than to the tropical Region. Emerson (1955) states that cubitus. The imago head and pronotum have it originated in Central America during post- unique, very long, and wavy hairs. There are Eocene times, but the presence of a species in no living species of Kalotermes in the Neo- Ilha Grande, near Rio de Janeiro, Brazil, tropical Region, but the Kalotermes-like casts some doubt on this hypothesis. stock, from which Comatermes was derived, Pterotermes probably also arose directly was probably present in ancient times. from the kalotermitid ancestor. It has re- Ceratokalotermes is an endemic genus of tained the ancestral imago mandible and Australia. The wing venation is similar to wing venation. The posterior margin of the 1961 KRISHNA: KALOTERMITIDAE 387 first plus second marginal tooth of the left cies), Indo-Malayan (two), Malagasy (one mandible is equal to the anterior margin of new species), Nearctic (seven, including one the third marginal tooth. The median vein in new species), and Neotropical (16, including the forewing is weak and unsclerotized and six new species). The distribution of species runs midway between the radial sector and suggests a New World origin. the cubitus. The antenna has 20 to 21 articles. Marginitermes is an endemic genus of The arolium is absent. The soldier is large and western United States and western Mexico. highly specialized in some respects. The eyes It probably arose from an Incisitermes-like are pigmented. The third article of the an- stock. The imago mandible and wing vena- tenna is dark, large, and club-shaped. The tion are similar to those of Incisitermes. The posterior margin of the postmentum has a arolium is always absent. The soldier is distinct, sclerotized swelling. Its large size specialized and has a somewhat phragmotic and multi-segmented antenna suggest a head, with a raised margin between the ver- primitive structure, in spite of the loss of the tex and the frons. The third antennal segment arolium. is enlarged and as long as the next five to The second line is represented by the seven segments together. Incisitermes-Cryptotermes complex. From the Allotermes is an endemic genus of Madagas- second line there developed three branches: car which probably arose from an Incisi- the first branch developing into Incisitermes termes-like stock. The wing venation resem- and its relatives, i.e., Allotermes, Margini- bles that of the genus Incisitermes. The termes, Tauritermes, and Proneotermes; the imago mandible is slightly more advanced, second branch developing into Bifiditermes, i.e., the length of the anterior margin of the Bicornitermes, and Epicalotermes; and the third marginal tooth of the left mandible is third branch developing into Procryptotermes more than one and one-half times the length and Cryptotermes. The trend of the dentition of the posterior margin of the first plus second of the left mandible of the imago in this line marginal tooth. has been towards further proportional elonga- Tauritermes also was probably derived tion of the anterior margin of the third margi- from an Incisitermes-like stock. It is endemic nal tooth compared to the posterior margin of to South America. The imago is unknown, the fused first plus second marginal tooth. but the nymph mandible is Allotermes-like. The maximum length of the anterior margin The soldier is specialized and has a phrag- of the third marginal tooth is more than one motic head. and one-half times as long as the posterior Proneotermes probably arose from an margin of the first plus second marginal Incisitermes-like base. The imago mandible is tooth. This extreme type of dentition is found like that of Incisitermes. The wing venation is in Tauritermes, Allotermes, Epicalotermes, more advanced than that of Incisitermes. The Procryptotermes, and Cryptotermes, and has median vein in the forewing is weakly sclero- probably evolved independently in several of tized, running closer to the radial sector than these genera.of the three branches. to the cubitus. The wing venation resembles Incisitermes has retained the ancestral wing that of the genera Paraneotermes, Ceratokalo- venation. The imago mandible, however, has termes, and Comatermes of the first main line, advanced, i.e., the anterior margin of the because of convergent evolution. The genus is third marginal tooth in the left mandible is endemic to Central and South America. plainly longer, but not more than one and The second branch is represented by Bifidi- one-half times longer, than the posterior termes, Bicornitermes, and Epicalotermes, and margin of the first plus second marginal arose from a common ancestor. The wing tooth. The arolium may be present or absent, because of convergent reduction. The soldier venation is similar in the three genera, i.e., head is dorsoventrally flat; the anterior mar- the media and cubitus in the forewing branch gin of the pronotum is deeply incised; the from a common stem outsidce the wing scale. third article of the antenna is dark, long, and An arolium is lacking. club-shaped. The distribution of the species Bifiditermes is the most primitive genus of is: Papuan (eight, including three new spe- this branch. The imago mandible is similar to 388 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 that of the genus Incisitermes, i.e., the ante- marginal tooth of the left mandible is more rior margin of the third marginal tooth in the than one and one-half times longer than the left mandible is plainly longer, but not more posterior margin of the first plus second than one and one-half times longer, than the marginal tooth. The median vein in the fore- posterior margin of the first plus second wing is weak and unsclerotized, running marginal tooth. The distribution of the midway between the radial sector and the species is: Australian (two), Indo-Malayan cubitus in the proximal half of the wing, (four, including one new species), Ethiopian bending up near the middle or beyond the (seven, including one new species), and middle to meet the radial sector. Malagasy (two, including one new species). Procryptotermes is more 'primitive than The genus is confined to the tropics of the Old Cryptotermes, as is indicated by the soldier, World. The distribution pattern suggests a which does not have such a highly developed, probable origin in the Ethiopian Region in phragmotic head, or other adaptations that Cretaceous times. are associated with the phragmotic head. The Bicornitermes probably descended directly distribution of species is: Papuan (three, from Bifiditermes. The imagoes of the two including one new species), Ethiopian (one), genera are similar. The soldier is specialized Malagasy (two, including one new species), and has developed a highly phragmotic head. and Neotropical (three, including two new The distribution of species is Ethiopian (four, species). It is strictly a tropical genus and is including one new species). The genus prob- found only on islands, an indication that it is ably originated in Tertiary times in the an ecological relict. The place of origin is Ethiopian Region. obscure. Epicalotermes is more advanced than Cryptotermes probably arose directly from Bifiditermes and Bicornitermes, as is indicated Procryptotermes, which is indicated by a by the imago mandible. The anterior margin transition in the development of the phrag- of the third marginal tooth of the left mandi- motic head from Procryptotermes to Crypto- ble is more than one and one-half times longer termes. The distribution of species is: Austra- than the posterior margin of the first plus lian (six, including two new species), Papuan second marginal tooth. The soldier has a flat (eight, including five new species), Indo- head, and the mandibles are strongly curved. Malayan (five), Ethiopian (four), Malagasy The distribution of species is: Ethiopian (two, including one new species), Nearctic (four) and Malagasy (one new species). The (one), and Neotropical (nine, including five genus probably originated in the Ethiopian new species). The genus is primarily tropical, Region and dispersed to Madagascar in the with the exception of a single species in the late Cretaceous or early Eocene before it warm temperate region of Florida, and capa- became isolated from the mainland. ble of living under dry conditions; therefore it The third branch is represented by Pro- has often been introduced into other conti- cryptotermes and Cryptotermes. The imago nents. The world-wide distribution indicates mandible and wing venation are similar in the a very ancient origin. The place of origin is two genera. The anterior margin of the third obscure. DISCUSSION CONSERVATIVE CHARACTERS On the basis of this adaptive character, ALTHOUGH IT IS DIFFICULT to prove that a some species were erroneously placed in given structure has no adaptive value, the Cryptotermes or Procryptotermes. The imago evidence sometimes points strongly in that mandible, wing venation, and other associ- direction. In termites, the imago-worker ated characters show that this adaptation has mandibles are adaptive in their biting and evolved independently many times. chewing functions, but the morphological The third segment of the soldier antenna is differences in the dentition of various genera greatly enlarged, dark, and highly sclero- and higher categories are very conservative, tized. The adaptive function of this enlarged non-adaptive, or weakly adaptive, slowly segment is unknown; it is probably sensory. evolving characters, compared to the more Such an adaptation has evolved independ- rapidly evolving and highly adaptive char- ently in almost all the genera of the Incisi- acters of the soldier, which are presumably termes-Cryptotermes complex (except Crypto- under high selection pressure. The dentition termes), and in Pterotermes, Rugitermes, and a of the imago mandible does not change with few species of Neotermes. change of food habits. Termites that eat the In Proneotermes, Paraneotermes, Cerato- same food have entirely different dentitions kalotermes, and Comatermes, the median vein according to their systematic relationships; in the forewing is slightly sclerotized, running conversely, closely related termites that midway between the radial sector and the occupy different niches and utilize different cubitus. This wing venation has evolved food have similar mandibles. Emerson (in independently in the two major branches of Allee et al., 1949) and Ahmad (1950) postu- the phylogenetic tree. late that multiple genes that influence the The arolium has convergently regressed in development of the imago-worker mandible some or all of the species of the following have pleiotropic effects, some of them affect- genera: Pterotermes, Paraneotermes, Neo- ing the development of vital adaptations in termes, Incisitermes, Marginitermes, Allo- other bodily activities. Selection for these termes, Bifiditermes, Epicalotermes, Bicorni- multiple effects gradually produces evolu- termes, and Cryptotermes. tionary changes in the imago-worker mandi- ble. Circumstantial evidence indicates that REGRESSION these gene complexes have been stable for long geological periods. Therefore this highly The modern theory of regressive evolution conservative pattern gives us a clue to basic formulated by Emerson (in Allee et al., 1949) phylogenetic relationships. is based on the following factors: most genes (pleiotropic) affect more than one character; CONVERGENCE most characters (polygenic) are affected by The present study reveals many cases of more than one gene; each gene mutates at a convergence in the phylogeny of the kalo- specific statistical rate; most mutations are termitid genera. Serious mistakes have been deleterious; and entire organisms or unitary made in the use of such analogous characters populations are selected as whole entities as for phylogenetic interpretations. well as the parts of each. The phragmotic or truncate head is a A character would be expected to remain defensive adaptation for a plugging of the functional as long as it is maintained by a passageways in the wood. The evolution of positive selection pressure. Decrease or loss of the extremely phragmotic head has taken this selection pressure will not prevent the place seven times in both major branches of gradual regression of the structure. Regres- the phylogenetic tree of the family Kalo- sion may also take place in the absence of termitidae. The genera that show this adap- negative selection pressure. If one character tation are: Kalotermes, Eucryptotermes, Glyp- decreases in importance relative to others, totermes, Calcaritermes, Tauritermes, Bicorni- there will be a shift in the allelic frequency in termes, and Cryptotermes. many gene systems, with consequent degen- 389 390 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 TABLE 4 CHARACTERS THAT SHOW PHYLOGENETIc ADVANCE IN THE FAMILY KALOTERMITIDAE Primitive Condition Derivative Condition IMAGO Left imago-nymph mandible with anterior margin Left imago-nymph mandible with anterior margin of third marginal tooth equal to posterior mar- of third marginal tooth longer than posterior gin of first plus second marginal tooth margin of first plus second marginal tooth 01 Right mandible with posterior margin of second Right mandible with posterior margin of second marginal tooth equal to molar plate marginal tooth longer than molar plate Median vein weak and unsclerotized, running mid- Media strongly sclerotized, running closer to ra- way between radial sector and cubitus, to tip of dial sector than to cubitus, to tip of wing or wing joining radial sector Damp-wood dwellers Dry-wood dwellers SOLDIER Head long, thick, or flat Head short, truncate, or phragmotic Third segment of antennae short, and not strongly Third segment of antennae long, club-shaped, and sclerotized highly sclerotized Anterior margin of pronotum slightly emarginate Anterior margin of pronotum deeply incised Anterior margin of pronotum smooth Anterior margin of pronotum serrated eration of this less important character and HOST-PROTOZOA RELATIONSHIP an increased development of the others. Thus The Protozoa have accompanied the a positive selection pressure for other char- termites during their phylogenetic develop- acters which are genetically and epigeneti- ment, so are particularly related to the cally interconnected with the regressed limited group of hosts in which they occur. characters would secondarily produce a more For these reasons, the Protozoa have been rapid regression of the neutral or less valuable used as indicators of relationship of the hosts character. (Kirby, 1937a, 1944, 1947, 1949b). Concomi- TABLE 5 CHARACTERS THAT SHOW PHYLOGENETIC REGRESSION IN THE FAMILY KALOTERMITIDAE Primitive Condition Derivative Condition IMAGO Median vein in hind wing present Median vein in hind wing absent Anal vein present Anal vein absent Subcosta present Subcosta absent Tibial spines in middle tibia present Tibial spines in middle tibia absent Arolium present Arolium absent Antennal segments numerous (21 in Pterotermes) Antennal segments fewer (11 in Glyptotermes) Branches of radial sector present Branches of radial sector absent SOLDIER Mandibles long Mandibles reduced Left mandible with three prominent marginal Left mandible with reduced number of or no mar- teeth ginal teeth Right mandible with two marginal teeth Right mandible with reduced numberof or no mar- ginal teeth Eyes pigmented Eyes unpigmented Wing pads present Wing pads absent Antenna with numerous segments (maximum num- Antenna with fewer segments (minimum number ber 19) 10) Spines in middle tibia present Spines in middle tibia absent 1961 KRISHNA: KALOTERMITIDAE 391 tant studies of Protozoa and their hosts have not occur in conjunction with the evolution of been made a basis for conclusions about their the kalotermitid genera. Because certain phylogeny and biogeographical and evolu- genera of Protozoa are characteristic of the tionary history. I compiled a Protozoa and family Kalotermitidae rather than of the host list (table 6) to find corroborative evi- genus of the host, it seems that the generic dence for my present generic classification. differentiation of the Protozoa took place It is evident from the Protozoa-host data before the differentiation of the kalotermitid that evolution of the genera of Protozoa did genera. TABLE 6 SYSTEMATIC LIST OF THE PROTOZOA SYMBIOTIC OF THE FAMILY KALOTERMITIDAE Protozoa Hosts POLYMASTIGIDA FAMILY CALONYMPHIDAE Calonympha grassii Foa Cryptotermes brevis (Walker) Coronympha clevelandi Kirby Incisitermes immigrans (Snyder) Coronympha octonaria Kirby Incisitermes emersoni (Light) Incisitermes pacificus (Banks) Incisitermes tabogae (Banks) Incisitermes platycephalus (Light) Incisitermes lighti (Snyder) Diplonympha foae Grassi Glyptotermes parvulus (Sj6stedt) Metacoronympha senta Kirby Incisitermes tabogae (Banks) Incisitermes lighti (Snyder) Incisitermes emersoni (Light) Incisitermes platycephalus (Light) Snyderella bandeirantium de Mello Cryptosermes brevis (Walker) Snyderella tabogae Kirby Cryptotermes longicollis Banks Snyderella ypiranga de Mello Rugitermes rugosus (Hagen) Stephanonympha campinae de Mello Neotermes hirtellus (Silvestri) Stephanonympha erythrei Grassi Neotermes erythraeus Silvestri Stephanonympha havilandi Grassi Cryptotermes havilandi (Sjostedt) Stephanonympha nelumbium Kirby Cryptotermes domesticus (Haviland) Stephanonympha reenstierna de Mello Cryptotermes sp. Stephanonympha silvestrii Janicki Neotermes connexus Snyder FAMILY DEVESCOVINIDAE Achemon platycaryon Grass6 and Hollande Cryptotermes havilandi (Sjostedt) Bullanympha silvestrii Kirby Neotermes erythraeus Silvestri Caduceia bugnioni Kirby Neotermes greeni (Desneux) Caduceia kalshoveni Kirby Neotermes dalbergiae (Kalshoven) Caduceia kofoidi Kirby Glyptotermes guianensis Emerson Glyptotermes perparvus Emerson Caduceia monile Kirby Neotermes tectonae (Dammermann) Caduceia nova (Grassi) Neotermes, new species, from Tanganyika Neotermes erythraeus Silvestri Caduceia pruvoti (Duboscq and Grass6) ?Kalotermes sp. from Lifou, Loyalty Islands Caduceia theobromae Franca Neotermes gestri Silvestri Neotermes aburiensis Sjostedt Bifiditermes mutabae (Harris) Bifiditermes, new species, from Transvaal Devescovina arta Kirby Glyptotermes angustus Snyder Glyptotermes minutus Kemner Neotermes castaneus (Burmeister) 392 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 TABLE 6-(Continued) Protozoa Hosts Devescovina coghilli Kirby Cryptotermes merwei Fuller Devescovina cometoides de Mello and de Britto ?Kalotermes sp. from Damann, India Cryptotermes dudleyi Banks Glyptotermes dilatatus (Bugnion and Popoff) Glyptotermes, new species, from Java Incisitermes repandus (Hill) Devescovina cuneata Kirby Neotermes, new species, from El Salvador Devescovina exilis Kirby Neotermes connexus Snyder Devescovina fissa Kirby Procryptotermes, new species, from Reunion Devescovina glabra Grassi Cryptotermes havilandi (Sj6stedt) Cryptotermes kirbyi Moszkowski Cryptotermes dudleyi Banks Glyptotermes caudomunitus Kemner Glyptotermes, new species, from Sumatra Bifiditermes madagascariensis (Wasmann) Neotermes zuluensis Holmgren Neotermes meruensis (Sj6stedt) Neotermes europae (Wasmann) Devescovina hawaiensis Janicki Neotermes connexus Snyder Devescovina insolita Kirby Neotermes, new species, from Tanganyika Bifiditermes, new species, from Transvaal Devescovina lemniscata Kirby Cryptotermes domesticus (Haviland) Cryptotermes dudleyi Banks Cryptotermes fatulus Light Cryptotermes havilandi (Sj6stedt) Cryptotermes cynocephalus Light Cryptotermes queenslandis Hill ?Kalotermes sp. from Damann, India Glyptotermes tuberculatus Froggatt ?Glyptotermes caudomunitus Kemner Neotermes insularis (White) Neotermes larseni Light Postelectrotermes castaneiceps (Sj6stedt) Postelectrotermes, new species, from near Ihosy, Madagascar Postelectrotermes longiceps (Cachan) Postelectrotermes longus (Holmgren) Devescovina lepida Kirby Calcaritermes brevicollis (Banks) Calcaritermes emarginicollis (Snyder) Calcaritermes nearcticus Snyder Calcaritermes parvinotus Light Calcaritermes nigriceps Emerson Calcaritermes, new species, from El Salvador Cryptotermes longicollis Banks Neotermes castaneus (Burmeister) Neotermes holmgreni Banks Devescovina minor Kirby Glyptotermes perparvus Emerson Glyptotermes guianensis Emerson Devescovina parasoma Kirby Neotermes tectonae (Dammermann) Neotermes dalbergiae (Kalshoven) Neotermes sonneratiae Kemner Cryptotermes cynocephalus Light Cryptotermes havilandi (Sj6stedt) 1961 KRISHNA: KALOTERMITIDAE 393 TABLE 6-(Continued) Protozoa Hosts Devescovina robusta Kirby Neotermes erythraeus Silvestri Devescovina similis Kirby Cryptotermes havilandi (Sj6stedt) Devescovina striata Foa Cryptotermes brevis (Walker) Devescovina tendicula Kirby Bifiditermes mutabae (Harris) Devescovina transita Kirby Cryptotermes cynocephalus Light Glyptotermes, new species, from Java Glyptotermes dilatatus (Bugnion and Popoff) Incisitermes repandus (Hill) Devescovina uniflexa Kirby Proneotermes perezi Holmgren Devescovina vestita Kirby Glyptotermes niger Kemner Devescovina vittata Kirby Bifiditermes jeannelanus (Sj6stedt) Evemonia punctata Grass6 Neotermes aburiensis Sjostedt Evemonia ramosa Kirby Neotermes aburiensis Sjostedt Foaina ? acontophora Kirby Glyptotermes guianensis Emerson Foaina acontophora Kirby Glyptotermes perparvus Emerson Cryptotermes havilandi (Sjostedt) Postelectrotermes castaneiceps (Sj6stedt) Postelectrotermes longiceps (Cachan) Postelectrotermes longus (Holmgren) Foaina appendicula Kirby Glyptotermes minutus Kemner Foaina calmoni de Mello Cryptotermes havilandi (Sjostedt) Foaina costata Kirby Neotermes insularis (White) Foaina decipiens (Grass6) Neotermes aburiensis Sj6stedt Foaina delicata Kirby Glyptotermes iridipennis Froggatt Glyptotermes brevicornis Froggatt Glyptotermes neotuberculatus Hill Foaina dogieli (Duboscq and Grass6) Kalotermes flavicollis (Fabricius) Foaina duo Kirby Glyptotermes niger Kemner Foaina exempta Kirby Neotermes insularis (White) Foaina falcifera Kirby Glyptotermes contracticornis (Snyder) Glyptotermes montanus Kemner Foaina fontesi de Mello Cryptotermes havilandi (Sjostedt) Foaina funifera Kirby Cryptotermes queenslandis Hill Glyptotermes tuberculatus Froggatt Foaina gracilis Janicki Neotermes connexus Snyder Foaina grassii (Duboscq and Grass6) Neotermes, new species, from El Salvador Kalotermes flavicollis (Fabricius) Kalotermes dispar Grasse Foaina hamata Kirby Glyptotermes hospitalis Emerson Glyptotermes ueleensis Coaton Glyptotermes parvulus (Sjostedt) Glyptotermes caudomunitus Kemner Glyptotermes sp. from Java Cryptotermes kirbyi Moszkowski Neotermes, new species, from El Salvador Foaina hilli (Duboscq and Grasse) Glyptotermes iridipennis Froggatt Glyptotermes brevicornis Froggatt Foaina humilis Kirby Cryptotermes brevis (Walker) Foaina inflata Kirby Neotermes, new species, from El Salvador Foaina minuscula Kirby Allotermes, new species, from near Mahabo, Mada- gascar Foaina nana (Kirby) Calcaritermes, new species, from El Salvador Calcaritermes brevicollis (Banks) 394 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 TABLE 6-(Continued) Protozoa Hosts Foaina nana (Kirby) Calcaritermes emarginicollis (Banks) Calcaritermes nearcticus Snyder Calcaritermes nigriceps (Emerson) Cryptotermes cubicoceps Emerson Cryptotermes domesticus (Haviland) Cryptotermes cynocephalus Light Cryptotermes brevis (Walker) Cryptotermes dudleyi Banks Cryptotermes fatulus Light Cryptotermes havilandi (Sj6stedt) Glyptotermes dilatatus (Bugnion and Popoff) Glyptotermes taveuniensis Hill Glyptotermes, new species, from Java Glyptotermes, new species, from Sumatra ?Kalotermes sp. from Damann, India Bifiditermes condonensis (Hill) Bifiditermes madagascariensis (Wasmann) Incisitermes repandus (Hill) Neotermes connexus Snyder Neotermes dalbergiae (Kalshoven) Neotermes greeni (Desneux) Neotermes larseni Light Neotermes sonneratiae Kemner Neotermes tectonae (Dammermann) Allotermes, new species, from near Mahabo,'Mada- gascar Rugitermes magninotus Emerson Rugitermes kirbyi Snyder Rugitermes panamae (Snyder) Foaina ovata Kirby Postelectrotermes howa (Wasmann) Neotermes desneuxi (Sj6stedt) Neotermes gracilidens Sj6stedt Foaina parvula Kirby Glyptotermes contracticornis (Snyder) Glyptotermes montanus Kemner Glyptotermes neotuberculatus Hill Glyptotermes taveuniensis Hill Bifiditermes jeannelanus (Sjostedt) Bifiditermes, new species, from Transvaal Neotermes erythraeus Silvestri Postelectrotermes castaneiceps (Sjostedt) Postelectrotermes longus (Holmgren) Postelectrotermes longiceps (Cachan) Neotermes, new species, from Tanganyika Foaina ramulosa Kirby Cryptotermes kirbyi Moszkowski Foaina reflexa Kirby Calcaritermes nigriceps (Emerson) Cakcaritermes brevicollis (Banks) Calcaritermes emarginicollis (Banks) Calcaritermes nearcticus Snyder Calcaritermes parvinotus Light Proneotermes perezi Holmgren Cryptotermes brevis (Walker) Cryptotermes cavifrons Banks Cryptotermes longicollis Banks 1961 KRISHNA: KALOTERMITIDAE 395 TABLE 6-(Continued) Protozoa Hosts Foaina reflexa Kirby Cryptotermes havilandi (Sjostedt) Bifiditermes jeannelanus (Sj6stedt) Bifiditermes mutabae (Harris) Epicalotermes mkuzii (Coaton) Neotermes castaneus (Burmeister) Neotermes erythraeus Silvestri Neotermes holmgreni Banks Neotermes meruensis (Sj6stedt) Neotermes zuluensis Holmgren Neotermes europae (Wasmann) Neotermes, new species, from Tanganyika Epicalotermes kempae (Wilkinson) Bifiditermes, new species, from Transvaal Foaina serrata Kirby Neotermes, new species, from El Salvador Foaina signata Kirby Bifiditermes jeannelanus (Sjostedt) Neotermes meruensis (Sjostedt) Neotermes zuluensis Holmgren Neotermes europae (Wasmann) Foaina solita Kirby Cryptotermes dudleyi Banks Cryptotermes longicollis Banks Cryptotermes havilandi (Sjostedt) Glyptotermes angustus Snyder Glyptotermes dilatatus (Bugnion and Popoff) Glyptotermes parvulus (Sj6stedt) Glyptotermes tuberculatus Froggatt Glyptotermes ueleensis Coaton Glyptotermes, new species, from near Rutchuru, the Congo Glyptotermes, new species, from Ruanda-Urundi, the Congo Bifiditermes mutabae (Harris) ?Kalotermes sp. from Damann, India Bifiditermes madagascariensis (Wasmann) Incisitermes repandus (Hill) Cryptotermes queenslandis Hill Neotermes castaneus (Burmeister) Neotermes connexus Snyder Neotermes greeni Holmgren Neotermes holmgreni Banks Neotermes sonneratiae Kemner Neotermes tectonae (Dammermann) Procryptotermes, new species, from R6union Rugitermes kirbyi Snyder Rugitermes panamae (Snyder) Rugitermes magninotus Emerson Foaina taenicola Kirby Paraneotermes simplicicornis (Banks) Hyperdevescovina balteata Kirby Ceratokalotermes spoliator (Hill) Hyperdevescovina caudata Kirby Kalotermes umtatae (Coaton) Hyperdevescovina calotermitis (Nurse) Kalotermes brouni Froggatt Hyperdevescovina falcifera Kirby Kalotermes rufinotum Hill Kalotermes banksiae Hill Hyperdevescovina insignita Kirby Postelectrotermes militaris (Desneux) Hyperdevescovina mitrata Kirby Kalotermes pallidinotum Hill Kalotermes tillyardi Hill i. 396 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 TABLE 6-(Continued) Protozoa Hosts Hyperdevescovina riciniata Kirby Kalotermes atratus (Hill) Hyperdevescovina torquata Kirby Kalotermes hilli (Emerson) Macrotrichomonas directa Kirby Incisitermes, new species, from El Salvador Macrotrichomonas emersoni Kirby Glyptotermes hospitalis Emerson Macrotrichomonas hirsuta Grass6 and Hollande Postelectrotermes barretoi (Grass6) Postelectrotermes praecox (Grasse) Macrotrichomonas lighti (Connell) Paraneotermes simplicicornis (Banks) Macrotrichomonas procera Kirby Calcaritermes brevicollis (Banks) Calcaritermes emarginicollis (Banks) Calcaritermes nearcticus Snyder Calcaritermes parvinotus Light Calcaritermes, new species, from El Salvador Macrotrichownoas pulchra Grassi Glyptotermes parvulus (Sjostedt) Glyptotermes ceylonicus Holmgren Glyptotermes contracticornis (Snyder) Glyptotermes brevicornis Froggatt Glyptotermes iridipennis Froggatt Glyptotermes montanus Kemner Glyptotermes neotuberculatus Hill Glyptotermes taveuniensis Hill Glyptotermes ueleensis Coaton Glyptotermes, new species, from the Congo Glyptotermes, new species, from Ruanda-Urundi, the Congo Glyptotermes magsaysayi Snyder Macrotrichomonas ramosa Kirby Glyptotermes brevicaudatus (Haviland) Macrotrichomonas restis Kirby Neotermes jouteli (Banks) Macrotrichomonas unguis Kirby Glyptotermes caudomunitus Kemner Glyptotermes sp. from Java Glyptotermes minutus Kemner Macrotrichomonas virgosa Kirby Allotermes, new species, from Mahabo, Madagascar Metadevescovina carina Kirby Bifiditermes madagascariensis (Wasmann) Metadevescovina cristata Kirby Bifiditermes angulatus (Wilkinson) Metadevescovina cuspidata Kirby Incisitermes minor (Hagen) Metadevescovina debilis Light Marginitermes hubbardi (Banks) Metadevescovina fulleri Kirby Bifiditermes madagascariensis (Wasmann) Bifiditermes durbanensis (Haviland) Epicalotermes munroi (Coaton) Epicalotermes mkuzii (Coaton) Epicalotermes, new species, from Tulear, Madagascar Metadevescovina magna Kirby Incisitermes marginipennis (Latreille) Metadevescovina mediocris Kirby Glyptotermes, new species, from near Rutchuru, the Congo Glyptotermes, new species, from Ruanda-Urundi, the Congo Metadevescovina modesta Kirby Glyptotermes angustus Snyder Metadevescovina modica Kirby Incisitermes marginipennis (Latreille) Metadevescovina nitida Kirby Allotermes paradoxus Wasmann Allotermes, new species, from Tsihombe, Madagascar Metadevescovina nudula Kirby Neotermes jouteli (Banks) Metadevescovina patula Kirby Bifiditermes condonensis (Hill) Metadevescovina polyspira (Lewis) Pterotermes occidentis (Walker) Metadevescovina stereociliata (Grassi) Glyptotermes parvulus (Sj6stedt) Glyptotermes ueleensis Coaton 1961 KRISHNA: KALOTERMITIDAE 397 TABLE 6-(Continued) Protozoa Hosts Metadevescovina turbula Kirby Neotermes jouteli (Banks) Parajoenia grassii Janicki Neotermes connexus Snyder Pseudodevescovina brevirostris Grass6 Neotermes aburiensis Sj6stedt Pseudodevescovina uniflagellata Sutherland Neotermes insularis (White) Stellaria nucleoflexa (Kirby) Cryptotermes merwei Fuller Cryptotermes havilandi (Sjostedt) Cryptotermes kirbyi Moszkowski FAMILY MONOCERCOMONADIDAE Hexamastix claviger Kirby Incisitermes marginipennis (Latreille) Cryptotermes dudleyi Banks Calcaritermes brevicollis (Banks) Hexamastix conigaviger Kirby Cryptotermes brevis (Walker) Hexamastix disclaviger Kirby Cryptotermes brevis (Walker) Cryptotermes longicollis Banks Tricercomitus cunhai de Mello Cryptotermes havilandi (Sjostedt) Tricercomitus damasmorai de Mello Cryptotermes sp. Tricercomitus divergens Kirby Marginitermes hubbardi (Banks) Incisitermes minor (Hagen) Kalotermes flavicollis (Fabricius) Incisitermes immigrans (Snyder) Incisitermes lighti (Snyder) Incisitermes marginipennis (Latreille) Incisitermes tabogae (Snyder) Incisitermes sp. from Galapagos Island Incisitermes sp. from Costa Rica Calcaritermes brevicollis (Banks) Calcaritermes emarginicollis (Snyder) Glyptotermes angustus Snyder Glyptotermes contracticornis (Snyder) Cryptotermes longicollis Banks Neotermes connexus Snyder Neotermes holmgreni Banks Rugitermes kirbyi Snyder Rugitermes panamae (Snyder) Cryptotermes domesticus Haviland Cryptotermes brevis (Walker) Cryptotermes dudleyi Banks Cryptotermes domesticus (Haviland) Cryptotermes sp. from Galapagos Island FAMILY OXYMONADIDAE Barroella coronaria Cross Postelectrotermes howa (Wasmann) Barroella zeteki Zeliff Calcaritermes brevicollis (Banks) Microrhopalodina hofmanni (de Mello and de Mello) Cryptotermes sp. Microrhopalodina inflata Grassi and Foa Kalotermes flavicollis (Fabricius) Microrhopalodina multinucleata (Kofoid et al.) Cryptotermes dudleyi Banks Microrhopalodina occidentis (Lewis) Pterotermes occidentis (Walker) Oxymonas barbouri Zeliff Glyptotermes angustus Snyder Oxymonas brevis Zeliff Cryptotermes brevis (Walker) Oxymonas caudata Cross Proneotermes perezi Holmgren Oxymonas clevelandi Zeliff Incisitermes immigrans (Snyder) Oxymonas dimorpha Connell Paraneotermes simplicicornis (Banks) Oxymonas di-undulata Nurse Kalotermes brouni Froggatt 398 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 TABLE 6-(Continued) Protozoa Hosts Oxymonas gracilis Kofoid and Swezy Rugitermes magninotus Emerson Oxymonas grandis Cleveland Neotermes dalbergiae (Kalshoven) Neotermes tectonae (Dammermann) Oxymonas granulosa Janicki Neotermes connexus Snyder Oxymonas hubbardi Zeliff Marginitermes hubbardi (Banks) Oxymonasjouteli Zeliff Neotermes jouteli (Banks) Oxymonas kirbyi Zeliff Rugitermes kirbyi Snyder Oxymonas lutzi de Mello Cryptotermes havilandi (Sjostedt) Oxymonas megakaryosoma Cross Glyptotermes, new species, from Uganda Oxymonas minor Zeliff Incisitermes minor (Hagen) Oxymonas ovata Zeliff Calcaritermes brevicollis (Banks) Oxymonas parvula Kirby Cryptotermes domesticus (Haviland) Oxymonas pediculosa Kofoid and Swezy Calcaritermes nigriceps (Emerson) Oxymonas projector Kofoid and Swezy Glyptotermes perparvus Emerson Oxymonas rotunda Cross Calcaritermes emarginicollis (Banks) Oxymonas snyderi Zeliff Cryptotermes domesticus Haviland FAMILY TRICHOMONADIDAE Pentatrichomonoides scroa Kirby Cryptotermes dudleyi Banks Cryptotermes longicollis Banks Trichomonas barbouri Kirby Glyptotermes angustus Snyder Trichomonas brevicollis Kirby Calcaritermes brevicollis (Banks) Trichomonas cartagoensis Kirby Glyptotermes contracticornis (Snyder) Trichomonas holmgreni Kirby Neotermes holmgreni Banks Trichomonas sp. (Duboscq and Grasse) ?Kalotermes sp. from Lifou, Loyalty Islands HYPERMASTIGIDA FAMILY TRICHONYMPHIDAE Eulophomonas calotermitis Grassi Kalotertnes flavicollis (Fabricius) Hoplonympha natator Light Paraneotermes simplicicornis (Banks) Joenia annectens Grassi Kalotermes flavicollis (Fabricius) Kofoidia loriculata Light Paraneotermes simplicicornis (Banks) Mesojoenia decipiens Grassi Kalotermes flavicollis (Fabricius) Pseudotrichonympha sertaneja de Mello Neotermes, new species, from Sao Paulo, Brazil Spirotrichonympha polygyra Cupp Paraneotermes simplicornis (Banks) Staurojoenina assimilis Kirby Incisitermes minor (Hagen) Staurojoenina sp. Marginitermes hubbardi (Banks) Staurojoenina mirabilis Grassi Epicalotermes aethiopicus Silvestri Trichonympha ampula Kirby Pterotermes occidentis (Walker) Trichonympha chattoni Duboscq and Grass6 Glyptotermes iridipennis (Froggatt) Glyp4otermes brevicaudatus (Haviland) Glyptotermes brevicornis Froggatt Glyptotermes ceylonicus Holmgren Glyptotermes contracticornis (Snyder) Glyptotermes montanus Kemner Glyptotermes neotuberculatus Hill Glyptotermes parvulus (Sj6stedt) Glyptotermes taveuniensis Hill Glyptotermes ueleensis Coaton Glyptotermes, new species, from Ruanda-Urundi, the Congo Glyptotermes, new species, from the Philippines Incisitermes milleri (Emerson) Incisitermes schwarzi (Banks) 1961 KRISHNA: KALOTERMITIDAE 399 TABLE 6-(Continued) Protozoa Hosts Trichonympha corbula Kirby Allotermes, new species, from near Mahabo, Mada- gascar Postelectrotermes castaneiceps (Sjostedt) Postelectrotermes longus (Holmgren) Postelectrotermes longiceps (Cachan) Trichonympha divexa Kirby Kalotermes umtatae (Coaton) Trichonympha lighti Kirby Incisitermes emersoni (Light) Trichonympha peplophora Kirby Postelectrotermes howa (Wasmann) Neotermes desneuxi (Sjostedt) Neotermes gracilidens Sj6stedt Postelectrotermes amplus (Sjostedt) Trichonympha quasilla Kirby Proneotermes perezi Holmgren Trichonympha saepicula Kirby Rugitermes kirbyi Snyder Rugitermes panamae (Snyder) Trichonympha subquasilla Kirby Incisitermes immigrans (Snyder) Trichonympha tabogae Kirby Incisitermes tabogae (Snyder) Trichonympha teres Kirby Neotermes meruensis (Sj6stedt) Trichonympha zeylanica (Dobell) Postelectrotermes militaris (Desneux) Kalotermes hilli (Emerson) SUMMARY 1. THE FAMILY Kalotermitidae is redescribed. two main evolutionary lines. The first line is The subfamily names "Electrotermitinae" represented by the Proelectrotermes-Calcari- and "Kalotermitinae" are placed in synon- termes complex, and the second line by the ymy. The fossil genus Eotermes is removed Incisitermes-Cryptotermes complex. from the family Kalotermitidae and placed in 5. Several cases of convergence are illus- the family Hodotermitidae. trated. In both the main lines of the family 2. Three hundred and fifty-three species, Kalotermitidae, the phragmotic head, the fossil and living, are classified into 24 genera. enlarged third antennal segment, and the Of these 24 genera, the following eight are slightly sclerotized median vein have all new: Postelectrotermes, Ceratokalotermes, Co- evolved independently many times. Also, the matermes, Incisitermes, Marginitermes, Tauri- arolium has been convergently lost in many termes, Bifiditermes, and Bicornitermes. The genera. genera Pterotermes, Proneotermes, Allotermes, 6. A discussion on conservative and re- and Epicalotermes are resurrected. The genus gressed characters is included. Characters name "Proglyptotermes" is relegated to synon- that show phylogenetic advancement or ymy. All the genera are described, and the regression are also listed. generitype species are illustrated. 7. It is evident from the data on the hosts 3. The generic classification is based on a and Protozoa that the evolution of the genera constellation of conservative, adaptive, and of the Protozoa did not occur in conjunction regressed characters of both the imago and with the evolution of the host genera and that the soldier castes. the differentiation of the Protozoa genera 4. The phylogeny of the genera is dis- took place before the differentiation of the cussed. The imago-nymph mandible indicates host genera. BIBLIOGRAPHY AHMAD, M. 1920. (See Banks and Snyder, 1920.) 1950. The phylogeny of termite genera based BANKS, N., AND T. E. SNYDER on imago-worker mandibles. Bull. Amer. 1920. A revision of the Nearctic termites with Mus. Nat. Hist., vol. 95, art. 2, pp. 37- notes on biology and geographic dis- 86. tribution. Bull. U. S. Natl. Mus., no. 1955. Termites of West Pakistan. Biologia, 108, viii+228 pp. vol. 1, no. 2, pp. 202-264. BATHELLIER, J. 1958. Key to the Indomalayan termites.. Ibid., 1927. Contribution a l'6tude systematique et vol. 4, nos. 1-2, pp. i-xii, 33-198. biologique des termites de l'Indochine. ALLEE, W. D., A. E. EMERSON, 0. PARK, T. PARK, In Gruvel, A., Faune des Colonies AND K. P. SCHMIDT Frangaises. Paris, vol. 1, no. 4, pp. 125- 1949. Principles of animal ecology. Philadel- 365. phia and London, W. B. Saunders Co., E. 837 pp. BLANCHARD, BANKS, N. 1851. Insectes. In Gay, Claudio, Historia fisica 1901. Thysanura and Termitidae. Papers from y politica de Chile. Santiago, Chile, vol. the Hopkins Stanford Galapagos Ex- 6, pp. 87-91 (Termianos). pedition 1898-99. Proc. Washington BRUES, C. T., AND A. L. MELANDER Acad. Sci., vol. 3, pp. 541-546. 1932. Classification of insects. A key to the 1906. Two new termites. Ent. News, vol. 17, known families of insects and other ter- no. 9, pp. 336-337. restrial arthropods. Bull. Mus. Comp. 1918. The termites of Panama and British Zool., Harvard College, vol. 73, pp. 1- Guiana. Bull. Amer. Mus. Nat. Hist., 672. vol. 38, art. 17, pp. 659-667. BUGNION, E. 1919. Antillean Isoptera. Bull. Mus. Comp. 1914. La biologie des termites de Ceylan. Bull. Zool., Harvard College, vol. 62, no. 10, Mus. Natl. d'Hist. Nat., Paris, vol. 20, pp. 473-489. no. 4, pp. 170-204. 400 1961 KRISHNA: KALOTERMITIDAE 401 BUGNION, E., AND N. POPOFF "Devescovina" du termite africain 1910. Les Calotermes de Ceylan. M6m. Soc. "Cryptotermes havilandi" (Sj6stedt), fix6 Zool. France, vol. 23, no. 9, pp. 124-144. dans le sol bresilien. Rev. Brasileira BURMEISTER, H. Biol., vol. 12, no. 4, pp. 433-445. 1839. Handbuch der Entomologie. Berlin, vol. 1953. Sur une oxymonade de l'intestin du 2, pt. 1, pp. 757-768. termite africain " Cryptotermfes havilandi" CACHAN, P. Sj6stedt, recolt6 A Santos (Bresil). Ibid., 1949. Les termites de Madagascar. Mom. vol. 13, no. 1, pp. 65-72. Inst. Sci. Madagascar, ser. A, vol. 3, no. 1954a. Contribution a l'etude des micropara- 2, pp. 177-275. sites des termites bresiliens. II. Un 1951. Les termites de Madagascar. Premier nouveau calonymphide, "Snyderella supplement. Ibid., ser. A, vol. 5, no. 1, ypiranga" sp. n., de "Rugitermes ru- pp. 1-18. gosus" (Hagen, 1858). Ibid., vol. 14, COATON, W. G. H. no. 1, pp. 71-78. 1949. Notes on some South African species of 1954b. Pseudotrichonympha sertaneja n. sp. the families Hodotermitidae and Kalo- (Protozoa, Mastigophora), from the in- termitidae. Jour. Ent. Soc. South Africa, testine of a new termite (Rugitermes sp.) vol. 12, pp. 13-77. collected in Brazil. Parasitology, vol. 44, 1950. Cryptotermes of the Union of South Af- nos. 1-2, pp. 24-29. rica. Ibid., vol. 13, pp. 3-32. 1954c. On a new species of Stephenonympha 1955. New Isoptera from the Belgian Congo (Protozoa, Mastigophora) from the in- (with description of some named spe- testine of the Brasilian termite, Neo- cies). Ibid., vol. 18, no. 2, pp. 109-136. termes hirtellus. Ibid., vol. 44, nos. 1-2, CONNELL, F. H. pp. 30-33. 1930. The morphology and life cycle of Oxy- DESNEUX, J. monas dimorpha sp. nov., from Neo- 1904a. Notes termitologiques. Ann. Soc. Ent. termes simplicicornis (Banks). Univ. Belgique, vol. 48, no. 3, pp. 146-151. California Publ. Zool., vol. 36, no. 2, pp. 1904b. A propos de la phylogenie des termitides. 51-66. Ibid., vol. 48, no. 8, pp. 278-286. CROSS, J. B. 1904c. Trois termites nouveaux. Ibid., vol. 48, 1941. A study of Oxymonas minor Zeliff from no. 8, pp. 286-289. the termite Kalotermes minor Hagen. 1904d. Isoptera, family Termitidae. In Wyts- Univ. California Publ. Zool., vol. 43, no. man, P., Genera insectorum. Brussels, 15, pp. 379-404. fasc. 25, pp. 1-52. 1946. The flagellate subfamily Oxymonadinae. of New Guinea vol. no. pp. 67-162. 1905. Isoptera collected by Ibid., 53, 3, L. Bir6. Ann. Hist. Nat. Mus. Natl. Cupp, E. E. Hungarici, vol. 3, pp. 367-377. 1930. Spirotrichonympha polygyra sp. nov. from Neotermes simplicicornis Banks. 1908. Vari6t6s termitologiques. II. Ann. Soc. Univ. California Publ. Zool., vol. 33, no. Ent. Belgique, vol. 51, no. 12, pp. 388- 17, pp. 351-378. 400. DAMMERMANN, K. W. EHRLICH, P. R. 1915. On a new species of Calotermes (Calo- 1958. Problems of higher classification. Syst. termes tectonae) which attacks living teak Zool., vol. 7, no. 4, pp. 180-184. trees. Tijdschr. Ent., vol. 58 (suppl.), pp. EMERSON, A. E. 98-100. 1923. A new termite from the Juan Fernandez DE MELLO, I. F. Islands (Swedish Expedition 1916-17). 1946. Estudos ulteriores sbbre os protozoirios In Natural history of Juan Fernandez intestinais de uma termite indiana do and Easter Island. Uppsala, vol. 3, pt. genero Cryptotermes. An. Inst. Med. 3, pp. 392-394. Trop., vol. 3, pp. 29-52. 1925. The termites of Kartabo, Bartica Dis- 1952a. S6bre um flagelado do g6nero Tricer- trict, British Guiana. Zoologica, vol. 6, comitus, parasita do termita africano no. 4, pp. 291-459. Cryptotermes havilandi Sjostedt, cole- 1928. Termites of the Belgian Congo and the tado em terra brasileira (Santos). Arq. Cameroon. Bull. Amer. Mus. Nat. Biol., vol. 36, no. 310, pp. 1-4. Hist., vol. 57, art. 7, pp. 401-574. 1952b. Contribution & l'6tude des micropara- 1933. A revision of the genera of fossil and re- sites des termites br6siliens. I. Sur une cent Termopsinae (Isoptera). California 402 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 Univ. Publ. Ent., vol. 6, no. 6, pp. 165- FULLER, C. 196. 1921. The termites of South Africa, being a 1942. The relations of a relict South African preliminary notice. South African Jour. termite (Isoptera: Hodotermitidae, Nat. Hist., vol. 3, no. 1, pp. 14-52. Stolotermes). Amer. Mus. Novitates, no. GARDNER, J. C. M. 1187, pp. 1-12. 1944. New Termitidae from India and Burma 1943. Kalotermes milleri, a new species of ter- (Isoptera). Indian Jour. Ent., vol. 6, pp. mite from the Florida Keys and Ja- 103-110. maica (Isoptera, Kalotermitidae). GIEBEL, C. G. Psyche, vol. 50, pp. 18-22. 1856. Fauna der Vorwelt mit steter Beriuck- 1945. The neotropical genus Syntermes (Isop- sichtigung der lebenden Thiere mono- tera: Termitidae). Bull. Amer. Mus. graphisch dargestellt, 2. Gliedertheire 1. Nat. Hist., vol. 83, art. 7, pp. 427-472. Abt. Insecten und Spinnen. Leipzig, 1949. In Kirby, H., Devescovinid flagellates 18+411 pp. of termites. V. The genus Hyperdeves- GIRARD, M. covina and the genus Bullanympha, and 1879. Les insectes. In Trait6 6l6mentaire d'en- undescribed or unrecorded species. tomologie. Paris, vol. 2, pp. 261-294. Univ. California Publ. Zool., vol. 45, no. GOETSCH, W. 5, pp. 319-422. 1933. Die chilenischen Termiten. Zool. Jahrb. 1951. Termite studies in the Belgian Congo. (Syst.), vol. 64, no. 2, pp. 225-243. Deuxieme Rapp. Ann. (1949) Inst. GRASSi, P. P. Rech. Sci. en Afrique Centrale, pp. 149- 1938. Calotermes dispar n. sp., termite nouveau 160. des Iles Canaries. Le polymorphisme des 1952a. The biogeography of termites. Bull. soldats chez les calotermitides. Bull. Amer. Mus. Nat. Hist., vol. 99, art. 3, Soc. Ent. France, vol. 42 (1937), no. 20, pp. 217-225. pp. 291-295. 1952b. The neotropical genera Procornitermes 1939. Les termites de l'ile de Madere. Ibid., and Cornitermes (Isoptera, Termitidae). vol. 44, nos. 13, 14, pp. 179-185. Ibid., vol. 99, art. 8, pp. 479-539. 1949. Trait6 de zoologie. Paris, Insectes, vol. 1955. Geographical origins and dispersions of 9, pp. 1-1117. termite genera. Fieldiana: Zool., vol. 37, GRASS{, P. P., AND A. HOLLANDE pp. 465-521. 1950. Recherches sur les flagelles termiticoles. ENDERLEIN, G. Les sous-familles des Devescovininae 1909. Die Klassifikation der Embiidinen, nebst Kirby et des Macrotrichomonadinae morphologischen und physiologischen nov. Ann. Sci. Nat. Zool., ser. 11, vol. Bemerkungen, besonders uber das 12, pp. 25-64. Spinnen derselben. Zool. Anz., vol. 35, HAGEN, H. A. no. 6, pp. 166-191. 1853. Hr. Peters Berichtete uiber die von ihm FABRICIUS, J. C. Gesammelten und von Hrn. Dr. Her- 1793. Entomologia systematica. Copenhagen, mann Hagen bearbeiteten Neuropteren vol. 2, p. 87, no. 1 (sine syn. cit.), Termes aus Mosambique. Ber. K. Preussischen fatale = T. bellicosus, Afrika; p. 88, no. 2, Akad. Wiss. Berlin, pp. 479-481. destructor; p. 90, no. 3, morio; no. 4, T. 1854. tYber die Neuropteren der Bernstein arda=Eutermes atrox, Sierra Leone; Fauna. Verhandl. Zool. Bot. Verein p. 91, no. 5, T. mordax, Sierra Leone; no. Wien, vol. 4, pp. 221-232. 6, T. flavicolle. 1856. (See Pictet and Hagen, 1856.) FROGGATT, W. W. 1858a. Monographie der Termiten. Part 2. 1896. Australian Termitidae. Part II. Proc. Linnaean Ent., vol. 12, pp. 1-342. Linnean Soc. New South Wales, vol. 1858b. Catalogue of the specimens of neurop- 21, pt. 2, pp. 510-552. terous insects in the collection of the 1905. White ants (Termitidae>, with sugges- British Museum. London, pt. 1, Ter. tions for dealing with them in houses mitina, pp. 1-34. and orchards. Agr. Gaz. New South 1862. In Peters, W. C. H., Naturwissenschaft- Wales, vol. 16, pp. 632-656, 752-774. liche Reise nach Mossambique ... in Reprinted in Dept. Agr. New South Jahren 1842 bis 1848. Berlin, Zoologie, Wales Misc. Publ., no. 874, pp. 1-47. no. 5, Neuroptera, Termitina, pp. 57- 1915. White ants. Farmers Bull. Dept. Agr. 89. New South Wales, no. 60, pp. 1-46. 1863. Neuropteren aus der Braunkohle von 1961 KRISHNA: KALOTERMITIDAE 403 Rott im Siebengebirge. Palaeontograph- 1926d. Termites (Isoptera) from South Sea and ica, vol. 10, pp. 247-269. Torres Strait islands. Ibid., new ser., HANDLIRSCH, ANTON vol. 39, pt. 1, art. 4, pp. 20-24. 1908. Die fossilen Insekten und die Phylo- 1926e. New termites from Samoan Islands. genie der rezenten Formen. Leipzig, Entomologist, vol. 59, no. 762, pp. 296- Wilhelm Engelmann, 1430 pp. 300. 1930. Zwolfte Ordnund der Pterygogenea: 1932a. Termites (white ants) in southeastern Isoptera oder Termiten. In Kukenthal, Australia. A simple method of identifica- W., and T. Krumbach, Handbuch der tion and a discussion of their damage in Zoologie. Berlin and Leipzig, Walter de timber and forest trees. Pamphlet Gruyter und Co., vol. 4, no. 8, pp. 840- Counc. Sci. Indus. Res., no. 25, pp. 1-28. 858. 1932b. Australian termites (Isoptera). Biologi- HARE, L. cal notes and descriptions of new spe- 1937. Termite phylogeny as evidenced by sol- cies. Proc. Roy. Soc. Victoria, new ser., dier mandible development. Ann. Ent. vol. 44, pts. 1-2, pp. 134-154. Soc. Amer., vol. 37, no. 3, pp. 459-486. 1933. Notes on Porotermes and Calotermes HARRIS, W. V. (Isoptera) from the Australian region, 1948. Termites of the Uganda Protectorate. with descriptions of new species. Ibid., Proc. Roy. Ent. Soc. London, ser. B, new ser., vol. 46, pt. 1, art. 4, pp. 36-53. vol. 17, nos. 5-6, pp. 73-83. 1942. Termites (Isoptera) from the Austra- 1954. Termites from Socotra (Isoptera). Ann. lian region. Melbourne, Australian Mag. Nat. Hist., ser. 12, vol. 7, pp. 493- Council for Scientific and Industrial Re- 496. search, pp. 1-479. 1957. British Museum expedition to South- HOLMGREN, N. West Arabia (Isoptera). London, pp. 1909. Termitenstudien. 1. Anatomische Un- 421-433. tersuchungen. K. Svenska Vetensk. HAVILAND, G. D. Akad. Handl., vol. 44, no. 3, pp. 1-215. 1898. Observations on termites: with descrip- 1910a. Das System der Termiten. Zool. Anz., tions of new species. Jour. Linnean Soc. vol. 35, pp. 284-286. London, vol. 26, pp. 358-442. 1910b. The Percy Sladen Trust Expedition to HEER, 0. the Indian Ocean in 1905. No. VIII- 1849. Die Insektenfauna der Tertiiargebilde Isoptera. Trans. Linnean Soc. London, von Oeningen und von Radoboj in Cro- Zool., ser. 2, vol. 14, no. 8, pt. 1, pp. atien. Leipzig, vol. 2, pp. 22-36 (Ter- 135-148. miten). 191 la. Termitenstudien. 2. Systematik der Ter- HILL, G. F. miten. Die Familien Mastotermitidae, 1921. New and rare Australian termites with Protermitidae and Mesotermitidae. K. notes on their biology. Proc. Linnean Svenska Vetensk. Akad. Handl., vol. Soc. New South Wales, vol. 46, pt. 4, 46, no. 6, pp. 1-88. pp. 433-456. 1911b. Neu-Guinea-Termiten. Mitt. Berlin 1922. A new Australian termite. Ibid., vol. 47, Zool. Mus., vol. 5, no. 3, pp. 451-465. pt. 3, pp. 275-277. 1911c. Bemerkungen uber einige Termiten- 1925a. Termites from the Australian region, Arten. Zool. Anz., vol. 37, no. 26, pp. descriptions of new species and hitherto 545-553. undescribed castes. Proc. Roy. Soc. Vic- 1911d. Ceylon-Termiten. In Escherich, K., toria, new ser., vol. 37, pt. 2, pp. 206- Termitenleben auf Ceylon. Jena, pp. 229. 185-212. 1925b. The Victorian termites. Victorian Nat., 1912. Neue Termiten aus dem Deutschen En- vol. 42, no. 4, pp. 85-91. tomologischen Museum. Ent. Mitt., vol. 1926a. Termites from the Ellice group. Proc. 1, no. 9, pp. 280-282. Roy. Soc. Victoria, new ser., vol. 38, 1913a. Termitenstudien. 4. Versuch einer sys- art. 10, pp. 95-99. tematischen Monographie der Termiten 1926b. The genus Porotermes (Isoptera). Ibid., der orientalischen Region. K. Svenska new ser., vol. 38, pp. 143-149. Vetensk. Akad. Handl., vol. 50, no. 2, 1926c. Australian termites (Isoptera). Notes on pp. 1-276. Stolotermes, Calotermes, and Copto- 1913b. Termiten aus Natal und dem Zululande. termes, with descriptions of new species. Gesammelt von Dr. Ivar Triigiardh. Ibid., new ser., vol. 38, pp. 192-214. Ent. Tidskr., yr. 34, pp. 321-366. 404 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 1914. Wissenschaftliche Ergebnisse einer 1932b. Neue Termiten aus der orientalischen Forschungsreise nach Ostindien, ausge- Region I-IL. Ibid., vol. 53, nos. 2-3, pp. ftihrt im Auftrage der Kgl. Preuss. 133-155. Akademie der Wissenschaften zu Berlin 1934. Systematische und biologische Studien von H. v. Buttel-Reepen. III. Termiten uiber die Termiten Javas und Celebes. aus Sumatra, Java, Malacca und Cey- K. Svenska Vetensk. Akad. Handl., ser. lon. Gesammelt von Herrn. Prof. Dr. 3, vol. 13, no. 4, pp. 1-241. v. Buttel-Reepen in den Jahren 1911- KIRBY, H. 1912. Zool. Jahrb., Abt. Syst., vol. 36, 1926a. On Staurojoenina assimilis sp. nov., an nos. 2-3, pp. 229-290. intestinal flagellate from the termite HOLMGREN, K. AND N. Kalotermes minor Hagen. Univ. Cali- 1917. Report on a collection of termites from fornia Publ. Zool., vol. 29, no. 3, pp. India. Mem. Dep. Agr. India, vol. 5, 25-102. no. 3, pp. 138-171. 1926b. The intestinal flagellates of the termite HOLMGREN, N. AND K. Cryptotermes hermsi Kirby. Ibid., vol. 1915. Termiten aus Neu-Caledonien und den 29, no. 4, pp. 103-120. Benachbarten Inselgruppen. Nova Cale- 1928. A species of Proboscidiella from Kalo- donia, A, Zoologie, vol. 2, pt. 2, no. 6, termes (Cryptotermes) dudleyi Banks, a pp. 85-93. termite of Central America, with re- IMMS, A. D. marks on the oxymonad flagellates. 1925. A general textbook of entomology. Lon- Quart. Jour. Micros. Sci., vol. 72, pp. don, Methuen and Co., Order 7, Isoptera, 355-386. pp. 249-276. 1929. Snyderella and Coronympha, two new INGER, R. F. genera of multinucleate flagellates from 1958. Comments on the definition of genera. termites. Univ. California Publ. Zool., Evolution, vol. 13, no. 3, pp. 370-384. vol. 31, no. 18, pp. 417-432. JOHN, 0. 1930. Trichomonad flagellates from termites. 1925. Termiten von Ceylon, der Malayischen I. Tricercomitus gen. nov. and Hex- Halbinsel, Sumatra, Java und der Aru amastix Alexeieff. Ibid., vol. 33, no. 19, Inslen. Treubia, vol. 6, nos. 3-4, pp. pp. 393-444. 360-419. 1931. Trichomonad flagellates from termites. KALSHOVEN, L. G. E. II. Eutrichomastix and the subfamily 1930. De biologie van de djatitermiet (Kalo- Trichomonadinae. Ibid., vol. 36, no. 10, termes tectonae Damm.) in verband met pp. 171-262. zijn bestrijding. Thesis, Wageningen, 1932. Flagellates of the genus Trichonympha Holland; simultaneously published as in termites. Ibid., vol. 37, no. 15, pp. Meded. Inst. v. Plantenziekten, no. 76, 349-476. Buitenzorg, Java; with 8 pages sum- 1937a. Host-parasite relations in the distribu- mary in English: "Bionomics of Kalo- tion of protozoan termites. Ibid., vol. termes tectonae Damm. as a base for its 41, no. 15, pp. 189-212. control," pp. 1-154. 1937b. The devescovinid flagellate Parajoenia KELSEY, J. M. grassii from a Hawaiian termite. Ibid., 1944. The identification of termites in New vol. 41, no. 16, pp. 213-224. Zealand. New Zealand Jour. Sci. and 1938a. The devescovinid flagellates Caduceia Technol., sect. B, vol. 25, no. 6, pp. theobromae Franca, Pseudodevescovina 231-260. ramosa new species and Macrotricho- KEMNER, H. A. monas pulchra Grassi. Ibid., vol. 43, no. 1930. Fauna Sumatrensis (Bijdrage no. 66). 1, pp. 1-40. Termitidae. Tijdschr. Ent., vol. 73, nos. 1938b. Polymastigote flagellates of the genus 3-4, pp. 298-324. Foaina Janicki and two new genera, 1931. Die Termitenfauna von Amboina. Lunds Crucinympha and Bullanympha. Quart. Univ. Arsskr., new ser., Avd. 2, vol. 27, Jour. Micros. Sci., vol. 81, pt. 1, pp. 1- no. 13; Fysiodr. Sallsk. Handl., new 25. ser., vol. 42, no. 13, pp. 1-53. 1939. Two new flagellates from termites in the 1932a. Zwei Termiten aus Sinai mit beschrei- genera Coronympha Kirby and Meta- bung der neuen Art Kalotermes sinaicus. coronympha Kirby, new genus. Proc. Ent. Tidskr., vol. 53, nos. 2-3, pp. 87- California Acad. Sci., ser. 4, vol. 20, 92. no. 10, pp. 207-220. 1961 KRISHNA: KALOTERMITIDAE 405 1941. Devescovinid flagellates of termites. I. 1930. Notes on Philippine termites, IV. Philip- The genus Devescovina. Univ. California pine Jour, Sci., vol. 42, no. 1, pp. 13-58. Publ. Zool., vol. 45, no. 1, pp. 1-92. 1931. Present status of our knowledge of the 1942a. Devescovinid flagellates of termites. II. termites of China. Lingnan Sci. Jour. The genera Caduceia and Macrotricho- (1929), vol. 7, pp. 581-600. monas. Ibid., vol. 45, no. 2, pp. 93-166. 1932. Termites of the Marquesas Islands. 1942b. Devescovinid flagellates of termites. Bernice P. Bishop Mus. Bull., no. 98, III. The genera Foaina and Para- pp. 73-86. joenia. Ibid., vol. 45, no. 3, pp. 167- 1933. Termites of western Mexico. California 246. Univ. Publ., Ent., vol. 6, no. 5, pp. 79- 1944. The structural characteristics and nu- 164. clear parasites of some species of Tricho- 1934. In Kofoid, C. A., et al., Termites and nympha in termites. Ibid., vol. 49, no. 8, termite control. Second edition. Berke- pp. 185-282. ley, University of California Press, pp. 1945. Devescovinid flagellates of termites. IV. 117-135, 206-216, 311-313. The genera Metadevescovina and Pseudo- 1935. The Templeton Crocker Expedition of devescovina. Ibid., vol. 45, no. 4, pp. the California Academy of Sciences, 247-318. 1932. No. 20. The termites. Proc. Cali- 1947. Flagellate and host relationship of tricho- fornia Acad. Sci., ser. 4, vol. 21, no. 20, monad flagellates. Jour. Parasit., vol. pp. 233-258. 33, pp. 214-228. 1937. Contributions to the biology and taxon- 1949a. Devescovinid flagellates of termites. V. omy of Kalotermes (Paraneotermes) sim- The genus Hyperdevescovina, the genus plicicornis Banks (Isoptera). California Bullanympha and undescribed or unre- Univ. Publ., Ent., vol. 6, no. 16, pp. corded species. Univ. California Publ. 423-464. Zool., vol. 45, no. 5, pp. 319-422. LIGHT, S. F., AND E. C. ZIMMERMANN 1949b. Systematic differentiation and evolu- 1936. Termites of southeastern Polynesia. tion of flagellates in termites. Rev. Soc. Occas. Papers Bernice P. Bishop Mus., Mexicana Hist. Nat., vol. 10, pp. 57-79. vol. 12, no. 12, pp. 1-12. LATREILLE, P. A. 1811-1832. Insectes de l'Am6rique Equinoxi- LIMA, A. DE COSTA ale. In von Humboldt, F. H. A., Re- 1942. S6bre Kalotermes (Neotermes) wagneri cueil d'observations de zoologie. Paris, e especies afins (Isoptera: Kalotermi- vol. 1 (1811), p. 39 (Termes margini- tidae). Bol. Soc. Brasileira Agron., vol. penne); vol. 2 (1832), p. 111. 5, no. 1, pp. 1-14. LIGHT, S. F. MATSUMURA, S. 1921. Notes on Philippine termites, II. Philip- 1907. [Systematic entomology.] Tokyo, * pine Jour. Sci., vol. 19, no. 1, pp. 23-63. Konchv Bunruigaku, vol. 1. 1924. The termites (white ants) of China, MARTYNOV, A. V. with descriptions of six new species. 1937. [Wings of termites and phylogeny of China Jour. Sci. and Arts, vol. 2, nos. Isoptera and of allied groups of insects.] 1-4, pp. 50-60, 140-142, 242-254, 354- Moscow, Acad6mie des Sciences de 358. l'URSS (N. Y. Nassonov memorial vol- 1926a. On Hoplonympha natator gen. nov., sp. ume), pp. 83-150 (summary in English). nov. A non-xylophagous hypermastigote MICHENER, C. D. from the termite Kalotermes simplici- 1957. Some bases for higher categories in cornis Banks, etc. Univ. California Publ. classification. Syst. Zool., vol. 6, pp. Zool., vol. 29, no. 5, pp. 121-140. 160-173. 1926b. On Metadevescovina debilis gen. nov., MICHENER, C. D., AND R. R. SOKAL sp. nov. A xylophagous polymastigote 1957. A quantitative approach to a problem from the termite Kalotermes hubbardi in classification. Evolution, vol. 11, pp. Banks. Ibid., vol. 29, no. 6, pp. 141-157. 130-162. 1927. Kofoidia, a new flagellate from a Cali- MOSZKOWSKI, L. I. fornia termite. Ibid., vol. 29, no. 18, pp. 1955. Cryptotermes kirbyi, new species from 467-492. Madagascar and C. havitandi (Sj6stedt) 1929. New termite records for Lower Cali- from Africa and introduced into Mada- fornia. Pan-Pacific Ent., vol. 6, no. 2, gascar, India and South America (Isop- pp. 67-72. tera, Kalotermitidae). M6m. Inst. Roy. 406 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 Sci. Madagascar, ser. E, vol. 6, pp. 15- Staatssammlung in Mulnchen sowie 41. einigen anderen Sammlungen. Zool. M#LLER, F. Anz., vol. 39, pp. 221-232. 1873. Beitriage zur Kenntniss der Termiten. 1913. Die fossilen Termiten. Eine kurze I-III. Jenaische Zeitschr. Med. Natur- Zusammenfassung der bis jetzt be- wiss., (I-II), vol. 7, no. 3, pp. 333-358; kannten Funde. Trans. 2d Internatl. (III), vol. 7, no. 4, pp. 451-463. Congr. Ent., Oxford, vol. 2, pp. 318- OSHIMA, M. 335. 1912. [The third official report on termites.] SCHMIDT, K. P., AND A. E. EMERSON Taihoku (Pub. Govt. Formosa). 1960. Taxonomy. In Encyclopaedia Britan- 1917a. Three new species of termites from Caro- nica. Chicago, vol. 21, pp. 854-856. line Islands. Annot. Zool. Japonenses, SCUDDER, S. H. vol. 9, pt. 3, pp. 195-198. 1890. The Tertiary insects of North America. 1917b. Notes on a collection of termites from Bull. U. S. Geol. Surv. Territories, vol. Luzon obtained by R. C. McGregor. 13, pp. 102-116. Philippine Jour. Sci., vol. 12, no. 4, pp. SHIRAKI, T. 221-225. 1909. On the Japanese termites. Trans. Ent. 1920. Philippine termites collected by R. C. Soc. Japan, vol. 2, pp. 229-242. McGregor, with descriptions of one new SILVESTRI, F. genus and nine new species. Ibid., vol. 1901. Nota preliminare sui Termitidi sud- 17, no. 5, pp. 489-512. americani. Bol. Mus. Zool. Anat. Comp. 1923. Fauna Simalurensis-Termitidae. Cap. Torina, vol. 16, no. 389, pp. 1-8. Zool., vol. 2, no. 3, pp. 1-22. 1903. Contribuzione alla conoscenza dei Ter- PICTET, F. J. mite e Termitofili dell'America merid- 1854. In Hagen, H., tVber die Neuropteren der ionale. Redia, vol. 1, pp. 1-234. Bernstein Fauna. Verhandl. Zool.-Bot. 1912. Termite raccote da L. Fea alla Guinea Ver. Wien, vol. 4, pp. 221-232. Portoghese e alla isole S. Thom6, PICTET, F. J., AND H. HAGEN Annobon, Principe e Fernando Poo. 1856. Die im Bernstein befindlichen Neurop- Ann. Mus. Civ. Stor. Nat. Genova, ser. teren der Vorwelt. In Berendt, G. C., 3, vol. 5, no. 45, pp. 211-255. Die im Bernstein befindlichen organ- 1914. Contribuzione alla conoscenza dei Ter- ischen Reste der Vorwelt. Berlin, vol. 2, mitide e Termitofili dell'Africa occiden- pp. 41-125. tale. I. Termitidi. Bol. Lab. Zool. Gen. PITON, Louis-E. Agr. Portici, vol. 9, pp. 1-146. 1940. Paleontologie du gisement Eocene de 1918. Un genere e due nuove specie di Calo- Menat (Puy-de-D6me) (Flore et Faune). termitidi (Insecta, Isoptera) dell'Eritrea Paris, Lechevalier (Isoptera, pp. 144- (Africa Or). Ibid., vol. 12, pp. 347-351. 145). 1927. Descrizioni di dur nuove specie di PONGRAcZ, A. Isoptera dell'Africa. Ibid., vol. 21, pp. 1917. ttj harmadidoszaki termeszfaj Radoboj- 91-95. rol. A. Magyar kir Foldtani Intezet ev 1934. Compendio di entomologia applicata Konyve, vol. 25, no. 2, pp. 25-35. (agraria - forestale - medica - veterinaria), 1928. Die fossilen Insekten von Ungarn, mit 1-28). besonderer Berucksichtigung der Ent- parte speciale, vol. 1 (fogli wicklung der Europaischen Insekten- Portici, pp. 1-448 (Isoptera, pp. 27- Fauna. Ann. Hist. Nat. Mus. Natl. 41). Hungarici, vol. 25, pp. 91-194. SJOSTEDT, Y. RAMBUR, J. P. 1897. Neue Termiten aus Sierra Leone und 1842. Histoire naturelle des insectes. N6vrop- Guinea. Ent. Tidskr., vol. 18, no. 4, teres. Paris, 534 pp. p. 212. RATCLIFFE, F. N., F. J. GAY, AND T. GREAVES 1900a. Vorliiufige Diagnosen einiger afrikan- 1952. Australian termites; the biology, recog- ischen Termiten. Ibid., vol. 20, no. 4, nition and economic importance of the p. 278. common species. Melbourne, Common- 1900b. Monographie der Termiten Afrikas. wealth Science and Industrial Research K. Svenska Vetensk. Akad. Handl., Organization, pp. 1-124. vol. 34, no. 4, pp. 1-236. ROSEN, K. VON 1902a. Eine neue Termite aus Kamerun. Ent. 1912. Neue Termiten aus der zoologischen Tidskr., vol. 23, no. 4, p. 252. 1961 KRISHNA: KALOTERMITIDAE 407 1902b. Neue afrikanische Termiten. Ibid., vol. Sci., vol. 15, no. 7, pp. 152-162. 23, no. 4, pp. 302-304. 1925c. New termites from the Solomon Islands 1904. Monographie der Termiten Afrikas. and Santa Cruz Archipelago. I. Ibid., Nachtrage. K. Svenska Vetensk. Akad. vol. 15, no. 17, pp. 395-407. Handl., vol. 38, no. 4, pp. 1-20. 1925d. Notes on fossil termites with particular 1907a. ttber eine Termitensammlung aus reference to Florissant, Colorado. Proc. Kongo. Ent. Tidskr., vol. 28, no. 4, pp. Biol. Soc. Washington, vol. 38, pp. 149- 233-250. 166. 1907b. Wissenschaftliche Ergebnisse der 1925e. A new Rugitermes from Panama. Jour. Schwedischen Zoologischen Expedition Washington Acad. Sci., vol. 15, no. 9, nach dem Kilimandjaro, dem Meru pp. 197-200. und den umgebenden Massaisteppen 1926a. New termites from Guatemala, Costa Deutsch-Ostafrica, 1905-1906 unter Rica and Colombia. Ibid., vol. 16, no. Leitung von Prof. Y. Sj6stedt. Uppsala, 1, pp. 18-28. Schwedisch Akademie der Wissenschaf- 1926b. Five new termites from Panama and ten, vol. 3, no. 15, pp. 1-28 (Corroden- Costa Rica. Proc. Ent. Soc. Washing- tia, no. 1, Termitidae). ton, vol. 28, no. 1, pp. 7-16. 1911. Neue Ost- und Westafrikanische Ter- 1926c. Notes on termites from Arizona with miten. Ent. Tidskr., vol. 32, nos. 3-4, description of two new species. Cali- pp. 173-188. fornia Univ. Publ. Zool., vol. 28, no. 21, 1914. Termiten aus Madagascar eingesam- pp. 389-397. melt von Herrn Dr. W. Kaudern 1911- 1926d. Termites collected on the Mulford bio- 1912. Arkiv Zool., vol. 8, no. 27, pp. logical exploration to the Amazon 1-19. Basin, 192 1-1922. Proc. U. S. Natl. 1915. Termitidae, Pseudonevropteres I. In Mus., vol. 68, art. 14, pp. 1-76. Alluaud, Charles A., and R. Jeannel, 1929. New termites from the Antilles and Voyage de Ch. Alluaud et R. Jeannel Middle America. Proc. Ent. Soc. Wash- en Afrique Orientale 1911-1912. Paris, ington, vol. 31, no. 4, pp. 79-87. vol. 3, Insectes, pp. 1-18. 1932. Two new termites from Costa Rica. 1925. Neue Termiten aus Afrika und Mada- Ibid., vol. 34, no. 6, pp. 98-100. gaskar. Konowia, vol. 4, nos. 1-2, pp. 1933a. New termites from India. Proc. U. S. 53-55. Natl. Mus., vol. 82, art. 16, pp. 1-15. 1926. Revision der Termiten Afrikas. 3. 1933b. Calcaritermes in the United States. Monographie. K. Svenska Vetensk. Proc. Ent. Soc. Washington, vol. 35, Akad. Handl., ser. 3, vol. 3, no. 1, pp. no. 5, pp. 67-69. 1-419. 1934a. Two new termites from Costa Rica. SNYDER, T. E. Proc. Biol. Soc. Washington, vol. 47, 1920a. (See Banks and Snyder, 1920.) pp. 95-98. 1920b. Two new termites from Arizona. Proc. from Ent. Soc. Washington, vol. 22, no. 2, 1934b. New termites India. Indian Forest p. 38. Rec., ent. ser., vol. 20, pt. 2, pp. 1-28. 1922. New termites fiom Hawaii, Central and 1946. A small dark-colored new Kalotermes South America, and the Antilles. Proc. from Guatemala. Proc. Ent. Soc. Wash- U. S. Natl. Mus., vol. 61, art. 20, pp. ington, vol. 48, no. 6, pp. 158-160. 1-32. 1948. Our enemy the termite. Revised edition. 1923. A new Glyptotermes from Porto Rico. Ithaca, New York, Comstock Publish- Proc. Ent. Soc. Washington, vol. 25, ing Company. no. 4, pp. 89-94. 1949. Catalog of the termites (Isoptera) of the 1924. Descriptions of new species and hitherto world. Smithsonian Misc. Coll., vol. 112, unknown castes of termites from Amer- pp. 1-490. ica and Hawaii. Proc. U. S. Natl. Mus., 1952. A new Rugitermes from Guatemala. vol. 64, art. 6, pp. 1-40. Proc. Ent. Soc. Washington, vol. 54, 1925a. New termites and hitherto unknown no. 6, pp. 303-305. castes from the Canal Zone, Panama. 1956. A new Neotermes from Panama (Isop- Jour. Agr. Res., vol. 29, no. 4, pp. 179- tera: Kalotermitidae). Ibid., vol. 58, 193. no. 6, p. 352. 1925b. New American termites, including a 1958. Two new Glyptotermes from the Philip- new subgenus. Jour. Washington Acad. pines (Isoptera: Kalotermitidae). Ibid., 408 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 122 vol. 60, pp. 229-231. Comoren und Inseln Ostafrikas. In 1959. New termites from Venezuela with Voeltzkow, A., Reise in Ostafrika in keys and a list of the described Vene- den Jahren 1903-1905. Wissenschaft- zuelan species. Amer. Midland Nat., liche Ergebnisse. Stuttgart, vol. 3, no. 2, vol. 61, no. 2, pp. 313-321. pp. 115-127. SNYDER, T. E., AND A. E. EMERSON WEIDNER, H. 1949. (See Snyder, 1949.) 1955a. Korperbau, Systematik und Verbreitung SOKAL, R. R., AND C. D. MICHENER der Termiten. In Schmidt, H., Die Ter- 1958. A statistical method for evaluating miten. Leipzig, pp. 5-81. systematic relationship. Univ. Kansas 1955b. Die Bernstein-Termiten der Sammlung Sci. Bull., no. 38, pp. 1409-1438. des Geologischen Staatsinstituts, Ham- STATZ, G. burg. Mitt. Geol. Staatsinst., vol. 24, 1939. Geradfliigler und Wasserkafer der oli- pp. 55-74. gocWnen Ablagerungen von Rott. De- WENZEL, R. L. cheniana, no. 99A, pp. 1-102. 1953. Research aided by amber collector. Bull. STROUD, C. P. Chicago Nat. Hist. Mus., vol. 24, no. 8, Factor in pp. 6-7. 1953. analysis termite systematics, WHITE, A. Kalotermes. Syst. Zool., vol. 2, no. 2, 1846. Insects. In Richardson, John, and J. E. pp. 76-92. Gray, The zoology of the voyage of TILLYARD, R. J. H.M.S. Erebus and Terror, under the 1926. The insects of Australia and New Zea- command of ... J. C. Ross ... during land. Sydney, pp. 100-106 (chapter 11, ..... 1839-43. London, pl. 2 (pl. 35, fig. 4). Isoptera). WILKINSON, W. WALKER, F. 1954. Three new species of Kalotermitidae 1853. List of the specimens of neuropterous in- (Isoptera) from East Africa. Proc. Roy. sects in the collection of the British Mu- Ent. Soc. London, ser. B, vol. 23, pts. seum. London, pt. 3 (Termitides), pp. 5-6, pp. 75-82. 501-529. 1958. Two new species of Kalotermes Hagen WASMANN, E. from West Africa (Isoptera: Kalo- 1897. Termiten von Madagaskar und Ost- termitidae). Ibid., ser. B, vol. 27, pts. afrika. Abhandl. Senckenbergische 7-8, pp. 109-115. Naturf. Gesell., vol. 21, no. 1, pp. 137- 1959. Four new species of Kalotermitidae 182. from East Africa (Isoptera). Ibid., ser. 1909. Kurze tVbersicht uiber die Systematik B, vol. 28, pts. 5-6, pp. 61-72. der Termiten. In Escherich, K., Die Wu, C. F. Termiten oder weissen Ameisen. Leip- 1935. Catalogus insectorum Sinensium (Cata- zig, Werner Klinkhardt, pp. 169-179. log of Chinese insects). Peiping (Order 1910. Termiten von Madagaskar, den V, Isoptera), vol. 1, pp. i-iv, 217-222. It