Proceedings of the United States National Museum
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PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 106 Washington: 1956 No. 3364 CHIGGERS OF THE GENUS EUSCHONGASTIA ( ACARINA: TROMBICULIDAE) IN NORTH AMERICA^ By Charles E. Faurell- Introduction In this paper the systematics of the genus Euschongastm in North America are studied; 15 previously described species of Euschongastia are redescribed; four previously described species are synonymized ; nine new species of Euschongastia are de- scribed (one of these having been divided into two subspecies) ; the morphology, geography, and ecology are considered in de- scriptions; a key for North American Euschongastia is given; and the nomenclature of E. oregonensis (Ewing, 1929), E. californica (Ewing, 1925), and E. setosa (Ewing, 1937) are con- sidered. The author wishes especially to thank Dr. G. W. Wharton for his assistance throughout all phases of this work and Dr. H. S. Fuller for making available a manuscript copy of his paper on the Trombiculidae in the Oudemans collection that was published in ^ This paper is brsRd on a thesis submitted in partial fulfillment for the degree of Doctor of Philosophy, Duke University, Durham, N. C., in 1951. The investivratior'.s were sui)ported in part by a research grant to Dr. G. W. Wharton fi'om the Division of Research Grants and Fellowships of the National Institutes of Health, U. S. Public Health Service. =* Department of Biology, Vanderbilt University, Nashville. Tenn. 85 — 86 PROCEEDINGS OF THE NATIONAL MUSEUM ^^^- ^^^ 1952. To all other persons who contributed materials or infor- mation, the author is also very grateful. Chiggers are the larvae of trombiculid mites. They are the first active developmental stage in a life history which includes seven steps egg, deutovum, larva, nymphochrysalis, nymph, imagochrysalis, and adult. Although very small, they can be seen by the naked eye. They are important parasites of terrestrial vertebrate animals, and they are the only stage of the trombiculid life cycle which is known to be parasitic. So far as is known, the two other active stages, nymph and adult, are predacious on small arthropods and arthropod eggs. Chiggers are widely distributed throughout the world. Several species are irritating pests of man. Wherever these species occur they have been given common names by the people living there ; and they are known by reputa- tion in places beyond where they are found. Recently there has been a great awakening of interest in the study of chiggers. During World War II, under the medical urgency to find out everything possible concerning the vector of scrub typhus in the Pacific area, intensive investigations were begun. The interest aroused in this effort, which still continues, has spread to chiggers in general and has brought about rapid development in the systematics of the group. Few areas have been studied intensively. Experience indicates that whenever a new region is investigated new forms as well as known species will be found. From September 1947 until Decem- ber 1949 small mammals were trapped in Duke Forest, Durham County, and Orange County, N. C, for the purpose of determining the chigger populations. During the same time, chiggers removed from mammals collected by the Pennsylvania Mammal Survey were sent to Duke University for identification. A number of shipments were received from Ohio. The most numerous chigger specimens and species secured were of the genus Euschongastia. With these collections as a nucleus, a study of the systematics of the genus for North America was undertaken. Review of the literature Systematics : There has been some confusion in the historical development of the family Trombiculidae. For many years the life histories of the mites were unknown; and, when different developmental stages were collected, they were named independ- ently. Later, as life histories were determined, shifts of nomen- clature were required by application of the law of priority. CHIGGERS—FARRELL 87 In 1905 Berlese described Tromhicula minor, a mite generally accepted to be a nymph with a body shaped like the figure 8. The monotypic genus Tromhicula was erected for it in the family Trombidiidae. In 1912 Oudemans published a taxonomic sum- mary of chiggers with descriptions and notes on all known species. He included these larvae in the genus Microtrombidium Haller, 1882, type species M. purjnireum Haller, 1882, or in sev- eral genera he himself had created in 1910 and 1911. These genera also were in the Trombidiidae. In 1916 the first successful rearing of nymphs from engorged chiggers occurred. In that year Kneissl reported rearing nymphs of the European pest chigger Microtrombidium aidumTialis Microtrom- (Shaw, 1790) . He observed they were shaped not like bidium but had the same form as Tromhicula. He changed the name to Tromhicula autumnalis. In 1917 Miyajima and Okumura made similar observations and changes with Tromhicula akamushi (Brumpt, 1910) . Later, other chiggers which had not been reared to nymphs were moved from Microtrombidium to Tromhicula by various workers. Ewing (1929d) established the subfamily Trombiculinae for the vertebrate-infesting chiggers, separating them from the arthropod-infesting mites of the family. Willmann, in 1941, published the results of his examination of the type specimen of Tromhicula minor. He determined it to be an adult female, not a nymph. However, this discovery had no effect on the classification. By this time adults of other species had been reared. These adults from cultures were congeneric with T. minor. Ewing (1944a) proposed the new family Trombiculidae with two subfamilies, Trombiculinae and Hemitrombiculinae. Womers- ley (1944) added the subfamily Leeuwenhoekiinae and in 1945 raised the group to full family status. Ewing (1946a) retained Leeuwenhoekiinae as a subfamily and added the Walchiinae. Wharton (1947b) agreed with Ewing's determinations, added the subfamily Apoloniinae, and redefined the family on the basis of larval characteristics. In his new definition Wharton excluded the subfamily Hemitrombiculinae, which Ewing (1949) has said could well be made a separate family. At the same time Wharton gave a new diagnosis to the subfamilies based chiefly on the num- ber of segments in the larval legs. With this development the classification of chiggers took on its present form at these inter- mediate levels. Wharton, Jenkins, et al. (1951) published a key 88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. lOG to the subfamilies and the genera. Euschdngaiitia is in the sub- family Trombiculinae. The historical development of the genus Euschongastia Ewing, 1938, has been confused only by differences of opinion or inter- pretation which have occurred during the recent growth in the systematics of chiggers. As groups of species were shifted into new genera in an effort to get more natural combinations, acarologists did not always view the problem in the same way and did not shift their nomenclature simultaneously and iden- tically. The oldest species included in Euscho^igastia, as it is defined at present, is E. trouessarti, described by Oudemans in 1910 and placed in his new genus Schongastia, type species Thromhidium vandersandei Oudemans, 1905. Only three species were included. Later, many species were added by several v/orkers. Ewing (1929d) separated several species from Schongastia and placed them in a new genus, Neoschongastia, type species Schongastia americana Hirst, 1921. E. trouessarti was included in the new genus in 1932. In 1938 Ewing erected the monotypic genus Euschongastia for his new species E. americana. His diagnosis in effect split Neoschongastia. In 1939 Womersley proposed a new genus, Paraschongastia (type species Neoschongastia yeomansi Gunther, 1939, by sub- sequent designation in Womersley and Heaslip, 1943) , for a group of four species within Neoschongastia. Vitzthum (1942) considered Euschongastia as a subgenus of Schongastia, but subsequent authors have retained it as a full genus. Ewing (1946b) showed that the four species placed in Para- schongastia by Womersley were congeneric with the type species of Neoschongastia. Therefore, Paraschongastia Womersley, 1939, was a synonym of Neoschongastia Ewing, 1929. Ewing then restricted Neoschongastia to this newly formed group. This action excluded many species formerly in Neoschongastia. To accommodate these excluded species, he erected a new genus, Ascoschongastia Ewing, 1946, type species Neoschongastia nialayensis Gater, 1932. Workers outside the United States have followed Ewing's diagnoses of these genera. However, Wharton (1948) has pointed out that the type species of Ascoschongastia had an unusual morphological feature—the posterolateral setae lying off the scutum—which made it almost . : CraGGERS—^FARRELL 89 unique. He held that Ascoschongastia should be restricted to species with this character, and he expanded Euschongastia to include all other species formerly placed in Ascoschongastia. Fuller (1948) followed Wharton's concept and placed in Euschon- gastia a number of old species, including E. trouessarti. In 1950 Ewing split off the monotypic genus Boshellia (type species Neoschongastia hirsuta Boshell and Kerr, 1942). Recently, Fuller (1952), along with a list of the known species, gave a more com- plete diagnosis of the genus, v/hich is accepted for this paper. Distribution : The genus Euschongastia is widely distributed over the world. It has been found wherever collections have been made. In Ewing's original description only the genotype, E. americana,