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The Phylogeny and Systematics of the Family Thaumastocoridae (Hemiptera: Heteroptera)1

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The Phylogeny and Systematics of the Family Thaumastocoridae (Hemiptera: Heteroptera)1 1957 Ward: Relationships of Mallophaga on Tinamiformes 353 Avulsos Dept. Zool., Sao Paulo, 9(21): 321-40, 1 Mayr, Ernst, and Dean Amadon. 1951. A classification fig., 9 tab., 3 graphs. of recent birds. Amer. Mus. Nat. Hist. Novitates 1953. S6bre alguns gfineros e espdcies de Heptapso- No. 1496: 1-42. gastridae (Mallophaga)-—IV. Arq. Mus. Nac, Rio Miranda-Ribeiro, Alipio de. 1938. Notas ornithologicas de Janeiro, 42: 233-54, 5 pis. (XIII). Tinamidae. Rev. Mus. Paulista 23: 667- Guimaraes, L. R., and Frederico Lane. 1937. Contri- 788, 18 pis. buicoes para o conhecimento das Mallophagas das Peters, J. L. 1931. Check-list of birds of the world. aves do Brasil. VI. Rev. Mus. Paulista 23: 1-21, Cambridge: Harvard Univ. Press, xviii+345 pp. 5 pis. Romer, A. S. 1945. Vertebrate Paleontology, 2d Edi- Hellmayr, C. E., and Boardman Conover. 1942. Cata- tion. Chicago: Univ. Chicago Press. viii+687pp. logue of birds of the Americas and the adjacent Rothschild, Miriam, and Theresa Clay. 1952. Fleas, islands. Zool. Ser., Field Mus. Nat. Hist., 13 (Pt. I, No. 1). vi+636 pp. flukes and cuckoos. London: Collins Son and Co., Hopkins, G. H. E., and Theresa Clay. 1952. A check Ltd. xiv+485 pp. list of the genera and species of Mallophaga. London: Todd, W. E. C. 1942. List of tinamous in the collection Downloaded from https://academic.oup.com/aesa/article/50/4/353/68897 by guest on 30 September 2021 British Museum (Natural History). 362 pp. of the Carnegie Museum. Ann. Carnegie Mus. 29: Keler, Stefan von. 1938. tlber brasilianische Mal- 1-29. lophagen. 1. Beitrag. Arb. Morph. Taxon. Ent., Ward, R. A. 1957. A study of the host distribution and Berlin, 5(4): 305-26, 11 figs. some relationships of Mallophaga (Insecta) parasitic 1939a. Uber brasilianische Mallopagen. 2. Beitrag. on the Tinamiformes (Aves). Part 2. Ann. Ent. Arb. Morph. Taxon. Ent., Berlin, 6(3): 222-53, 23 Soc. America, in press. figs. Wetmore, Alexander. 1951. A revised classification for 1939b. Baustoffe zu einer Monographic der Mai- the birds of the world. Smithsonian Misc. Coll. 117: lophagen. II. Teil: Uberfamilie der Nirmoidea (I). 1-22. Nova Acta Acad. Leop.-Carol. (n.F.) 8(51): 1-254, Wetmore, Alexander, and W. H. Phelps, Jr. 1951. 4 pis., 114 figs. Observations on the geographic races of the tinamou Lambrecht, Kalman. 1933. Handbuch der Palaeorni- Crypturellus noctivagus in Venezuela and Colombia. thologie. Berlin: Gebriider Borntraeger. xix + 1024 Bol. Soc. Venezolana Cienc. Nat. 13: 115-19, 1 map. pp., 209 figs., 4 pis. Wilson, F. H. 1936. The segmentation of the abdomen McDowell, Samuel. 1948. The bony palate of birds. of Lipeurus heterographus Nitzsch (Mallophaga). I. The Palaeognathae. Auk 65: 520-49, 6 figs. Jour. Morph. 60(1): 211-19, 5 figs. THE PHYLOGENY AND SYSTEMATICS OF THE FAMILY THAUMASTOCORIDAE (HEMIPTERA: HETEROPTERA)1 CARL J. DRAKE Smithsonian Institution, Washington, D. C. AND JAMES A. SLATER Department of Zoology and Entomology, University of Connecticut, Storrs, Connecticut The recent acquisition of considerable unworked terized the first genus and species from the material from several museums together with our western hemisphere, which he made the basis personal collections in the little known family for the creation of a new subfamily. In this Thaumastoeoridae has led us to review the latter paper, the higher groups within the family taxonomic relationships within the family, to were redefined and their phylogenetic characters characterize two new genera and seven new were discussed. These papers constituted all of species, and to investigate the systematic and the original contributions of a systematic nature phylogenetic position of the family. to the family until Kormilev (1955) recently The Thaumastoeoridae were first described by founded a new subfamily and new genus for a Kirkaldy (1908) as a subfamily of Lygaeidae, new species from Argentina. The affinities of based upon a new genus founded for the reception this form are discussed below. of a new species from Australia. Bergroth (1909) In this paper the literature on the family is erected a second genus for a second species from reviewed. The absence of trichobothria and the Tasmania, amplified the original diagnosis of the asymmetry of the male genitalia are noted as group, and commented upon the presumed indicating that the Thaumastoeoridae belong to antiquity of these singular insects. He followed the Cimicomorpha of Leston, Pendergrast, and Kirkaldy's classification and retained the group South wood (1954) rather than to the Pentato- within the Lygaeidae. Reuter (1912) elevated momorpha where they have previously been the Thaumastoeoridae to the status of a family assigned. The cimicomorph relationship of the and based on them a distinct new phalanx within family is further evidenced by the hind wing his series Onyehiophora. Barber (1920) charac- having radius and media fused beyond the basal cell (Plate 1: 17), by the lack of an ectodermal 'Accepted for publication Oct. 30, 1956. spermatheca, and by the structure of the eggs 354 Annals Entomological Society of America [Vol. 50 which lack micropylar processes but have a true MORPHOLOGICAL FEATURES operculum bearing follicular pits (Plate 1: 9). The morphological features of the Thau- The systematic position of the Thaumastocoridae mastocoridae that are of major importance in within the cimicomorph complex is less evident. understanding the systematic and phylogenetic Both Kirkaldy (1908) and Kormilev (1955) noted relationships of the family are as follows. their general anthocorid-like appearance. Male genitalia. Throughout the family the The opinion of Bergroth (1909) that the thau- ninth abdominal segment is highly asymmetrical mastocorids represent very ancient insects obvi- (Plate 2: 1, 6, 8, 11). This asymmetry is so ously influenced Reuter (1912) in his systematic strongly developed that it affects the preceding placement of the group. It is interesting to note segment or segments; and in Xylastodoris, at in passing that Reuter, who was blind at that least, it has resulted in modification of segments time, never actually saw a specimen of the five, six, and seven to accomodate the ninth seg- Downloaded from https://academic.oup.com/aesa/article/50/4/353/68897 by guest on 30 September 2021 family. Before we attempt to analyze the rela- ment (Plate 2: 6). Members of the subfamily tionships and phylogeny of these peculiar insects, Thaumastocorinae are much less specialized in it seems advisable to comment briefly upon their this regard. In these species the genital capsule previously ascribed primitive nature. itself is always extended into a prong-like pro- Of the characters mentioned as primitive by jection (Plate 2: 8, 11) which perhaps functions, Reuter (1910, p. 37), the Thaumastocoridae pos- together with the single paramere characteristic sess the following: 1) presence of ocelli; 2) of this subfamily, in spreading apart the orifice Four-segmented antennae; 3) Four-segmented of the female genital chamber during copulation. labium (Plate 1: 7); 4) Front wing divisible into Curiously, some species are modified so that the three primary units, the clavus, corium and opening of the genital capsule is to the right side membrane (Text figs. 1 and 2); 5) Metasternum (Plate 2: 8), while in others the modification turns with scent gland openings (Plate 1: 14); 6) Homo- the opening to the left side (Plate 2: 11). Rela- morphous pairs of legs; 7) Lack of arolia, in one tively few specimens of most species in this sub- subfamily; 8) ventral spiracles. On surveying family have been available for examination, but these characters it is evident that, with the pos- modification seems to be consistently dextral or sible exception of the arolia, their occurrence is sinistral for any given species. The phallus is so widespread through many heteropterous fami- difficult to interpret without fresh material, but lies that they are of little value, if any, in ascer- it appears to be divisible into phallolheca2 and taining whether one is dealing with a group of endosoma. The endosoma, however, is not further generally specialized species or a more primitive subdivided into conjunctiva and vesica. Exam- type of insect. ination of fresh material would be highly desirable The characteristics which Reuter regarded as here. specialized conditions are represented in the In Xylastodoris further specialization has Thaumastocoridae as follows: 1) Lack of occurred. Both the parameres are lacking, and venation in the membrane of the front wing; 2) 2Terminology of genital structures follows that of Separate mesosternum and metasternum (Plate Dupuis (1955). 1: 14); 3) Reduction of the metasternal scent gland opening; 4) Trochalopodous hind coxae; 5) Reduction of the tarsal segments from three to two (Plate 1: 8, 13); 6) Occurrence of arolia; EXPLANATION OF PLATE I 7) Absence of an ovipositor (Plate 2: 10); 8) 1. Baclozygum depressum Bergroth, dorsal view of Dissimilar male genital parameres; 9) reduction head. 2. Onymocoris hackeri n. sp., dorsal view of head. of the nymphal scent glands from three to two 3. Thaumastocoris hackerin. sp., dorsal view of head. or one. In addition to the above characters, the 4. Onymocoris izzardi n. sp., dorsal view of head. features that we consider significant specializa- 5. Thaumastocoris petilus n. sp., dorsal view of head. tions are the marked asymmetry of the ninth 6. Tliaumastocoris australicus Kirkaldy, dorsal view of head. abdominal segment of the male, loss of parameres 7. Onymocoris hackeri n. sp., ventral view of labium. (either one or both), modifications of the basal 8. Thaumastocoris hackeri n. sp., tarsus and apex of labial segments, jugal enlargements, and the tibia, showing lobate sensory appendage. absence of an ectodermal spermatheca. 9. Thaumastocoris hackeri n. sp., egg. 10. Onymocoris barberi n. sp., cephalic view of head. While the evidence presented above is con- 11. Onymocoris hackeri n. sp., cephalic view of head. flicting to some extent, it seems evident that the 12.
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