Diptera:Ephydridae) Dick L
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Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1966 A revision of the Nearctic species of Hydrellia (Diptera:Ephydridae) Dick L. Deonier Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Entomology Commons Recommended Citation Deonier, Dick L., "A revision of the Nearctic species of Hydrellia (Diptera:Ephydridae) " (1966). Retrospective Theses and Dissertations. 5360. https://lib.dr.iastate.edu/rtd/5360 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. I This dissertation has been microfihned exactly as received 6 7—5582 DBONIER, Dick Lee, 1936- A REVISION OF THE NEARCTIC SPECIES OF HYDRELLIA (DIPTERA; EPHYDRIDAE). Iowa State University of Science and Technology, Ph,D„ 1966 Entomology University Microfilms, Inc., Ann Arbor, Michigan A REVISION OP THE NEARCTIC SPECIES OF HYDBELLIA (DIPTEBA: EPHYDRIDAE) by Dick h\ Deonier A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of The Requirements for the Degree of DOCTOR OF PHILOSOPHY Major Subject: Entomology Approved: Signature was redacted for privacy. ka^e of M^ ork Signature was redacted for privacy. Head of Major Department Signature was redacted for privacy. te C llege Iowa State University Of Science and Technology Ames, Iowa 1966 il TABLE OF CONTENTS Page INTRODUCTION AND LITERATURE REVIEW 1 METHODS AND MATERIALS 9 MORPHOLOGY 21 ECOLOGY 62 SYSTEMATIC TREATMENT 102 LITERATURE CITED 305 ACKNOWLEDGMENTS 320 APPENDIX A 321 APPENDIX B 365 1 INTRODUCTION AND LITERATURE REVIEW Introduction The Ephydridae constitute a medium-sized family of acalyp- trate Diptera. They lead several different modes of life, mostly in aquatic habitats. Hydrellia and Lemnaphila are the only known genera of leaf-mining ephydrids. At least two species of Hydrellia, H, griseola and H. ischiaca, are economically important. In 1953j H. griseola destroyed 10 to 20 per cent of California's rice crop with an estimated loss of $l6 million. The same species caused heavy losses of rice in California, in 1922. It has damaged rice in Japan, in the last two decades. Lilljeborg (l86l) first re corded the pest status of H. griseola. Lilljeborg reported the fairly widespread damage of barley, oats, and timothy grass by H. griseola in southern and southeastern Sweden, during the summer of i860. After this, several authors report ed outbreaks of H. griseola in various places in the Palaeartic Region including Egypt. Balachowsky and Mesnil (1935) reported 50 per cent infestation of barley by H. griseola in northern Europe. Hydrellia ischiaca attacks wild rice, which is now a minor crop in Minnesota. Most of the known larvae of other Hydrellia species feed mainly in plants of Potamogetonaceae, Alismataceae, and Hydrocharitaceae. There are 182 available specific names in Hydrellia other than my designations. Of these, perhaps 130 have existing 2 holotypes and 132 have Palaearctic type localities. Possibly 110 of the available names are valid. These 110 species are distributed as follows: 52 Palaearctic, 35 Nearctic, 1 Holarctic, 2 Oriental, 12 Australian, 2 Ethiopian, and 7 Neotropical. This essentially cosmopolitan generic distribu tion has caused some interest in the dispersal center. Some data possibly indicate this was in the northern temperate zone. I started this research in the summer of I961, with the following objectives: 1) to describe, redescribe, and con struct keys to the adults and as many immature stages as possi ble of Nearctic species; 2) to describe the life cycles and behavior of as many as possible of the Nearctic species; 3) to present a brief morphology of the genus. Literature Review The taxonomic literature on Hydrellia dates from 1813» when Fallen described Notiphila griseola and several congeneric species from southern Sweden. In I83O, Robineau-Desvoidy erected the new genus Hydrellia. Macquart (1835), Zetterstedt (1846), Walker (I856), Loew (i860), Schiner (1864), Brischke (1883)J Gobert (1887), Kowarz (1894), and Becker (I896, I903) made many of the Initial contributions to the taxonomy of Palaearctic species. Strobl (1904), Grunberg (I9IO), Becker (1919), Collin (1928), Prey (1933), de Meijere (1939), Goetghebuer (1942), Grensted (1944), Kloet and Hincks (1945), 3 and Tsacas (1959, I960) added some descriptions of new species, "but primarily they presented reviews and new distribution records. Becker's (I896, 1926) synoptic keys to and descrip tions of Palaearctic adult Hydrellia and Hennig's (1943) specific key to larval Hydrellia including Palaearctic species contributed most to the taxonomy of Palaearctic species of Hydrellia. The recent revision by Dahl (1964) of the Stenhammar and Zetterstedt collections eliminated much confu sion and synonymy in Palaearctic Hydrellia. Loew (1861, 1862) described the first new species of Mearctic Hydrellia. Loew (1872), Osten Sacken (I878), Becker (1896), Aldrich (I905), Jones (I906), and Coquillett (1910a), presented short sections on Nearctic Hydrellia. Very little additional taxonomic study of the genus was done until 1915» . when Cresson started describing new species. Johannsen (1935) reviewed and presented a key to the known immature instars of Nearctic species. Hennig (19^3) summarized the literature on immature Hydrellia and presented a specific key to all known immature instars including the few in the Nearctic. Cresson's studies culminated in a key (1944b) to the adults of the 35 Nearctic species then known. Subsequent to this, Berg (1949, 1950) contributed to the biology and taxonomy, including a key, of the immature instars of six species; Hennig (1952) presented morphological interpretations of immature Hydrellia; Wirth and Stone (I956) included a key to California species and 4 a Nearctic generic key; Grigarick •( 1959) redescrlbed the life- cycle stages of H. griseola; Deonier (1964) wrote a key to the adults of species of Iowa and adjacent states; and Wirth (I965) cataloged many Nearctic species. Cresson (I918, 1947a) described and presented keys to the six species of Hydrellia known from the Neotropical Region. Little study has been made of Hydrellia in the Oriental Region. Cresson (1948) listed only two species, H. latipalpis Cresson and H. luteipes Cresson from the Oriental Region in his Indo-Australian synopsis. The latter species is known only from Formosa, which is more or less transitional between the Oriental and Palaearctic Regions. In the Australian Region, Coquillett (1903), Tonnoir and Malloch (1926), Cresson (1948) and Harrison (1959) published on some or all of the 12 known indigenous species. The rela tively small amount of attention given to Oriental and Australian Hydrellia is matched only for the Ethiopian Hydrellia. Cresson (1932, 1947b) listed two species, both from the Cape of Good Hope. The works of the following authors provide some framework for a more searching morphological study of adult Hydrellia. Becker (I896, 1926), GrUnberg (I910), Cresson (I9I8), and Wilke (1924) dealt mainly with chaetotaxy, but Cresson did briefly discuss facial contour and the supposed absence of vibrissae in Ephydridae. Frey (I92I) discussed the mouthparts of H. obscura and the cibaria of other ephydrlds. Seguy (193^) illustrated the head and wing of H. griseola, and Scotland (19^0) illustrated by photographs the proboscis, antenna, and wing of the closely related genus Lemnaphila. Bering (1950) briefly described and illustrated the excised male terminalia of H. xenophaga and H. nigricans. Kato (1955) and Kuwayana (1955) described and illustrated several aspects of the external morphology of adult H. griseola. Hennig (1958) discussed the external morphology of the head and the thoracic chaetotaxy of several ephydrid representatives including H. griseola. With phylogenetic interpretations as a primary objective, Hennig made an important contribution here, especially in proposing the presence of vibrissae in Ephydridae. Dahl (1959) illustrated the adult mouthparts, female abdomen, and hind tarsus of H. griseola. Grigarick (1959) illustrated the male and female abdomina, the wing, and the mesonotum of H. griseola. Harrison (1959) illustrated the head, wing, and portions of the male terminalia of five New Zealand species of Hydrellia. Dahl (1964) illustrated the male terminalia of 17 species of Hydrellia from the Stenhammar and Zetterstedt collections. Deonier (1964) illustrated the head of H. harti and the chaetotaxy and sclerite nomenclature of the head and thorax exemplified in Ephydra riparia. Sturtevant (I925, 1926) made the only contributions to the internal morphology of Hydrellia in his survey of 6 spermathecae in Acalyptratae. Bolwig (19^0, 19^1) described and illustrated the internal genitalia and mouthparts of Scatophila unicornis. These studies helped in understanding the morphology of these structures in Hydrellia. Several authors contributed morphological data on the immatures of Hydrellia; von Frauenfeld (1866) described the gross aspects of the metamorphosis of H. albilabris; Stein (1867) summarized briefly the life cycle of H. griseola; Gercke (1879> 1882, I889) briefly described and illustrated the puparia and feeding apparatus of H. mutata and H. fulviceps; and