(1758–2021) (Acari: Trombiculoidea), with Notes on Nomenclature, Taxonomy, and Distribution (Zootaxa 4967) 243 Pp.; 30 Cm
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Zootaxa 4967 (1): 001–243 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Monograph ZOOTAXA Copyright © 2021 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4967.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:96B0632C-788A-493A-AA42-A237973EBF48 ZOOTAXA 4967 Annotated world checklist of the Trombiculidae and Leeuwenhoekiidae (1758– 2021) (Acari: Trombiculoidea), with notes on nomenclature, taxonomy, and distribution DAVID H. NIELSEN1, RICHARD G. ROBBINS2 & LEOPOLDO M. RUEDA3 1Nebraska Department of Agriculture, 301 Centennial Mall South, Lincoln, Nebraska 68509 USA. �[email protected] https://orcid.org/0000-0001-6651-4424 2 Walter Reed Biosystematics Unit, Department of Entomology, Smithsonian Institution, MSC, MRC 534, 4210 Silver Hill Road, Suitland, Maryland 20746 USA, and Department of Entomology, Walter Reed Army Institute of Research, 503 Robert Grant Avenue, Silver Spring, Maryland 20910, USA. �[email protected] https://orcid.org/0000-0003-2443-5271 3Walter Reed Biosystematics Unit, Department of Entomology, Smithsonian Institution, MSC, MRC 534, 4210 Silver Hill Road, Suitland, Maryland 20746 USA, and Department of Entomology, Walter Reed Army Institute of Research, 503 Robert Grant Avenue, Silver Spring, Maryland 20910, USA. Mailing address: 204 Draymore Way, Cary, North Carolina 27519, USA �[email protected] https://orcid.org/0000-0003-0038-2857 Magnolia Press Auckland, New Zealand Accepted by O. Seeman: 24 Feb. 2021; published: 7 May 2021 Licensed under Creative Commons Attribution-N.C. 4.0 International https://creativecommons.org/licenses/by-nc/4.0/ DAVID H. NIELSEN, RICHARD G. ROBBINS & LEOPOLDO M. RUEDA Annotated world checklist of the Trombiculidae and Leeuwenhoekiidae (1758–2021) (Acari: Trombiculoidea), with notes on nomenclature, taxonomy, and distribution (Zootaxa 4967) 243 pp.; 30 cm. 7 May 2021 ISBN 978-1-77688-250-2 (paperback) ISBN 978-1-77688-251-9 (Online edition) FIRST PUBLISHED IN 2021 BY Magnolia Press P.O. Box 41-383 Auckland 1041 New Zealand e-mail: [email protected] https://www.mapress.com/j/zt © 2021 Magnolia Press ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 4967 (1) © 2021 Magnolia Press NIELSEN ET AL. Table of Contents Abstract . 3 Introduction . 3 Approach . 4 Scope . 5 Future prospects. 7 Acknowledgements. 194 References. 195 Abstract The superfamily Trombiculoidea is a large and diverse group of acarines that comprises six families; of these, the families Trombiculidae and Leeuwenhoekiidae are characterized by larvae commonly known as chiggers that are parasites of terrestrial vertebrates, including humans, and some species are of medical importance as vectors of chigger-borne rickettsiosis (scrub typhus), caused by the rickettsia Orientia tsutsugamushi. This paper presents an annotated checklist of 3,013 generally accepted chigger species, together with their distribution by zoogeographic region, and a non-comprehensive list of synonyms. A total of 58 new combinations are proposed by transferring species to different genera, treating some subgenera as genera, or updating current generic names. The checklist updates earlier catalogs of the world chigger fauna, but because the literature on chigger taxonomy is voluminous and scattered, similar, regional monographs were utilized in constructing the classificatory foundation. This list may not contain every species, synonym or taxonomic rearrangement that has been published, but it reflects the most recent arrangement of chigger taxa. Distribution records are primarily drawn from collection sites listed in the original species descriptions, together with subsequently published geographical records. Key words: checklist, chigger mites, Trombiculoidea, taxonomy, biogeography Introduction The superfamily Trombiculoidea is a large and diverse group of mites. Its fundamental classification has occasionally been changed by several acarologists, particulary the arrangement of recognized superfamilies, families, subfamilies, tribes, genera and subgenera. To cite a few examples, Krantz (1978) noted that in addition to the family Chyzeridae, Vercammen-Grandjean (1973) recognized 8 families within what he referred to as the Trombidioidea, namely: Trombellidae, Johnstonianidae, Neotrombidiidae, Stygothrombiidae, Podothrombiidae, Trombidiidae, Trombiculidae and Leeuwenhoekiidae. Zhang et al. (2011), however, revised the classification of what is now called the Trombiculoidea and listed seven families: Trombellidae, Johnstonianidae, Neotrombidiidae, Audyanidae, Trombiculidae, Walchiidae, and Leeuwenhoekiidae. Larvae of the families Trombiculidae and Leeuwenhoekiidae, commonly known as chiggers, have been associated with human discomfort through their bite or through transmission of disease organisms (Audy 1968), but little was known about them until the early 20th century. The first chigger to receive a formal scientific name was Acarus batatas Linnaeus, now known as Eutrombicula batatas (Linnaeus) (Michener 1946). In 1912, Oudemans and Berlese separately published monographs containing a total of 33 known species, but until the outbreak of World War II, most studies on trombiculid taxonomy, ecology and disease relationships were conducted by Walch (1922-1927) in Europe and the Netherlands East Indies, Kawamura and Ikeda (1936) in Japan, and Ewing (1913-1937) in North America. During his long career, Ewing (active 1913-1950) described 64 new chigger species, 16 new genera, and two subfamilies, the Trombiculinae and Walchiinae. Ewing’s work in the 1920s and 1930s established a classificatory framework for chigger genera that became the foundation of modern chigger taxonomy. Propelled by studies of chigger-borne rickettsiosis (scrub typhus, tsutsugamushi disease) in Southeast Asia and the western Pacific during World War II, the post-war era saw an explosion of trombiculid mite research, resulting in great numbers of new genera and species. In 1952, Wharton and Fuller published “A Manual of the Chiggers,” a worldwide review that included four subfamilies, 31 genera, 17 subgenera, and 440 species. This work also summarized valuable data on species distribution and host preferences. Concurrently, Womersley (1952) published his voluminous monograph, “The Scrub-Typhus and Scrub Itch Mites CHECKLIST OF THE TROMBICULIDAE AND LEEUWENHOEKIIDAE Zootaxa 4967 (1) © 2021 Magnolia Press · 3 (Trombiculidae, Acarina) of the Asiatic-Pacific Region,” which described some 260 species, 75 of them new. This work was soon revised by Audy (1957), who also published a checklist of trombiculid mites of the Oriental and Australasian Regions. In 1968, Audy published a series of his lectures on chigger-borne rickettsiosis, the only major human disease associated with chiggers, in a book entitled “Red Mites and Typhus.” From the 1960s through the 1980s, Domrow conducted research on Australasian chigger taxonomy, while Vercammen-Grandjean, starting in the 1950s, began studying the trombiculid mites of Africa. During the 1960s and 1970s, Vercammen-Grandjean emerged as one of the most prolific chigger taxonomists of all time, describing more than 50 genera and 370 species with colleagues around the world. He also proposed a number of fundamental changes in chigger classification, including the creation of tribes within subfamilies, the tribes being further divided into complexes where genera were grouped based on combinations of characters. In 1971, Vercammen-Grandjean and Langston launched a monumental program to catalog the world chigger fauna. In a projected series of volumes entitled “The Chigger Mites of the World,” they endeavored to illustrate all accepted chigger species, together with known locality and host data. Unfortunately, only two volumes, the Leptotrombidium complex (volume III), published in 1976, and the Guntherana complex (volume VIII), published in 1971, were ultimately completed. While significant portions of Vercammen-Grandjean’s oeuvre are now disputed, his contributions to the study of trombiculoid mites continue to influence researchers. The 1960s and 1970s saw a number of other major publications addressing chigger taxonomy, particularly at the regional level. During the 1960s Brennan and collaborators published numerous papers on the chiggers of South America, bringing attention to a part of the world that had gone largely unstudied. Brennan’s work (1946-1982) still stands as the foundation for chigger taxonomy in the Neotropical region. A 1974 study by Nadchatram and Dohany provided diagnoses of all the chigger genera then known from Southeast Asia, as well as a classificatory arrangement that is still widely accepted. The late 1960s witnessed the first of many publications by the Russian acarologist Kudryashova, followed a decade later by the first publications by Goff; between them, they described almost 300 new species and multiple genera. Both authors also focused on refining the genus- level classification of chiggers. Goff, Loomis and Wrenn (1986) published a chigger bibliography, and Goff et al. (1983-1989) comprehensively defined the terms used in chigger taxonomy, setting a universal standard for chigger morphology. Kudryashova’s research (1965-2011) remains central to studies of chiggers in Russia and the Palearctic Region. More recent research by Stekolnikov (1992-2021) and a multitude of collaborators has expanded upon this base, yielding an understanding of chigger diversity in that part of the world, as well as Africa and Cuba. In 1984, Wen published “The