Research Article |
David C. Houghton‡, R. Edward DeWalt§, Todd Hubbard|, Kurt L. Schmude¶, Jeffrey J. Dimick#, Ralph W. Holzenthal¤, Roger J. Blahnik¤, James L. Snitgen«
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Corresponding author: David C. Houghton ( david.houghton@hillsdale.edu ) Academic editor: Steffen Pauls
© 2022 David C. Houghton, R. Edward DeWalt, Todd Hubbard, Kurt L. Schmude, Jeffrey J. Dimick, Ralph W. Holzenthal, Roger J. Blahnik, James L. Snitgen.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Houghton DC, DeWalt RE, Hubbard T, Schmude KL, Dimick JJ, Holzenthal RW, Blahnik RJ, Snitgen JL (2022) Checklist of the caddisflies (Insecta, Trichoptera) of the Upper Midwest region of the United States. In: Pauls SU, Thomson R, Rázuri-Gonzales E (Eds) Special Issue in Honor of Ralph W. Holzenthal for a Lifelong Contribution to Trichoptera Systematics. ZooKeys 1111: 287-300. https://doi.org/10.3897/zookeys.1111.72345
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Abstract
Five hundred and fifty-two caddisfly species are reported from the Upper Midwest region of the United States, an area that includes 13 states and ~ 2 million km2. Of these, 62 species are reported for the first time from the state of Iowa, 25 from Wisconsin, 18 from South Dakota, 12 from Illinois, five from Indiana, four from North Dakota, four from Minnesota, and one from Nebraska. The Upper Midwest fauna contains nearly 40% of all species known from the United States and Canada, as well as 22 species endemic to the region. Overall species richness was highest in Michigan (319 species), Kentucky (296), Minnesota (292), and Wisconsin (284). Differences in state species assemblages within the region largely followed a geographic pattern, with species richness declining in the western prairie states. There are almost certainly further species remaining to be found in this large region.
Keywords
Caddisfly, checklist, diversity, Midwest, Trichoptera, USA
Introduction
The Upper Midwest region of the United States (Fig.
Materials and methods
We have been investigating the caddisflies of the Upper Midwest for ~ 20 years (Fig.
The 13 states and primary ecoregions of the Upper Midwest region, showing collecting localities within the last ~20 years by the authors or their colleagues. Citations are the most comprehensive taxonomic works for states where our collecting effort was low. State abbreviations, IA: Iowa, IL: Illinois. IN: Indiana, KS: Kansas, KY: Kentucky, MI: Michigan, MN: Minnesota, MO: Missouri, NE: Nebraska, ND: North Dakota, OH: Ohio, SD: South Dakota, WI: Wisconsin.
Adult specimens were identified using
We also utilized the distributional checklist of
Differences in caddisfly assemblages relative to geography were examined with a non-metric multidimensional scaling (NMDS) ordination using the program PC-ORD v. 7 for Windows (
Results
Based on examination of ~ 750,000 larval and adult specimens from nearly 4,000 collecting localities (Fig.
The 131 new state species records reported herein. Species organized by family and genus. More detailed collecting data are available in Suppl. material
| Taxon | IA | IL | IN | MN | ND | NE | SD | WI |
|---|---|---|---|---|---|---|---|---|
| BRACHYCENTRIDAE | ||||||||
| Brachycentrus fuliginosus Walker, 1852 | – | – | – | – | – | – | – | X |
| B. lateralis (Say, 1823) | X | – | – | – | – | – | – | – |
| B. numerosus (Say, 1823) | X | – | – | – | – | – | – | – |
| GLOSSOSOMATIDAE | ||||||||
| Agapetus tomus Ross, 1941 | – | – | – | – | – | – | – | X |
| Glossosoma parvulum Banks, 1904 | – | – | – | – | – | – | X | – |
| Protoptila erotica Ross, 1938 | X | – | – | – | – | – | – | – |
| Helicopsychidae | ||||||||
| Helicopsyche borealis (Hagen, 1861) | X | – | – | – | – | – | – | – |
| Hydropsychidae | ||||||||
| Cheumatopsyche aphanta Ross, 1938 | – | – | – | – | – | – | X | – |
| C. campyla Ross, 1938 | – | – | – | – | – | – | X | – |
| C. halima Denning, 1948 | X | – | – | – | – | – | – | – |
| C. lasia Ross, 1938 | – | – | – | – | – | – | X | – |
| C. minuscula (Banks, 1907) | – | X | – | – | – | – | – | – |
| C. oxa Ross, 1938 | X | – | – | – | – | – | – | – |
| C. pasella Ross, 1941 | X | – | – | – | – | – | – | – |
| Diplectrona modesta Banks, 1908 | X | – | – | – | – | – | – | – |
| Homoplectra doringa (Milne, 1936) | – | X | – | – | – | – | – | – |
| Hydropsyche aerata Ross, 1938 | X | – | – | – | – | – | – | – |
| H. alternans (Walker, 1852) | X | – | – | – | – | – | – | – |
| H. arinale Ross, 1938 | X | – | – | – | – | – | – | – |
| H. betteni Ross, 1938 | – | – | – | – | – | – | X | – |
| H. dicantha Ross, 1938 | X | X | – | – | – | – | – | – |
| H. hageni Banks, 1905 | X | – | – | – | – | – | – | – |
| H. morosa Hagen, 1861 | – | – | – | – | – | – | X | – |
| H. phalerata Hagen, 1861 | – | – | – | – | X | – | – | – |
| H. scalaris Hagen, 1861 | X | – | – | – | – | – | – | – |
| H. slossonae Banks, 1905 | X | – | – | – | – | – | – | – |
| H. sparna Ross, 1938 | X | – | – | – | – | – | – | – |
| Macrostemum carolina (Banks, 1909) | X | – | – | – | – | – | – | – |
| Parapsyche apicalis (Banks, 1908) | X | – | – | – | – | – | – | – |
| Hydroptilidae | ||||||||
| Agraylea multipunctata Curtis, 1834 | X | – | – | – | – | – | – | – |
| Hydroptila ajax Ross, 1938 | – | – | – | – | – | – | X | – |
| H. albicornis Hagen, 1861 | X | – | – | – | – | – | – | – |
| H. ampoda Ross, 1941 | – | – | – | – | – | – | – | X |
| H. angusta Ross, 1938 | – | – | – | – | X | – | X | X |
| H. arctia Ross, 1938 | – | – | – | – | – | – | X | – |
| H. consimilis Morton, 1905 | – | – | – | – | – | – | X | – |
| H. delineata Morton, 1905 | – | – | – | – | – | – | – | X |
| H. grandiosa Ross, 1938 | X | – | – | – | – | – | – | – |
| H. gunda Milne, 1936 | – | X | – | – | – | – | – | – |
| H. metoeca Blickle & Morse, 1954 | – | – | – | – | – | – | – | X |
| H. perdita Morton, 1905 | X | – | – | – | – | – | – | – |
| H. quinola Ross, 1947 | – | – | – | – | – | – | – | X |
| H. scolops Ross, 1938 | – | – | X | – | – | – | – | – |
| H. tusculum Ross, 1947 | – | – | – | – | – | – | – | X |
| H. xera Ross, 1938 | – | – | – | – | – | – | – | X |
| Neotrichia minutisimella (Chambers, 1873) | X | – | – | – | – | – | – | – |
| N. vibrans Ross, 1938 | X | – | – | – | – | – | – | – |
| Ochrotrichia alsea Denning & Blickle, 1972 | – | – | – | – | – | – | X | – |
| O. arva (Ross, 1941) | – | – | – | – | – | – | – | X |
| O. riesi Ross, 1944 | – | – | – | – | – | – | – | X |
| Orthotrichia cristata Morton, 1905 | X | – | – | – | – | – | – | – |
| O. curta Kingsolver & Ross, 1961 | – | – | – | – | – | – | – | X |
| Oxyethira forcipata Mosely, 1934 | X | – | – | – | – | – | – | – |
| O. novasota Ross, 1944 | – | X | – | – | – | – | – | – |
| LEPIDOSTOMATIDAE | ||||||||
| Lepidostoma griseum (Banks, 1911) | – | X | – | – | – | – | – | – |
| L. liba Ross, 1941 | X | – | – | – | – | – | – | – |
| L. sommermanae Ross, 1946 | – | X | – | – | – | – | – | – |
| L. togatum (Hagen, 1861) | X | – | X | – | – | – | – | – |
| LEPTOCERIDAE | ||||||||
| Ceraclea alagma (Ross, 1938) | X | – | – | – | – | – | – | – |
| C. alces (Ross, 1941) | X | – | – | – | – | – | – | – |
| C. ancylus (Vorhies, 1909) | X | – | – | – | – | – | X | – |
| C. cancellata (Betten, 1934) | X | – | – | – | – | – | X | – |
| C. enodis Whitlock & Morse, 1994 | X | – | – | – | – | – | – | – |
| C. erratica (Milne, 1936) | – | – | – | X | – | – | – | – |
| C. maculata (Banks, 1899) | – | – | – | – | – | – | X | – |
| C. neffi (Resh, 1974) | X | – | – | – | – | – | – | – |
| C. nepha (Ross, 1944) | X | – | – | – | – | – | – | – |
| C. ophioderus (Ross, 1938) | – | – | – | – | – | – | – | X |
| C. resurgens (Walker, 1852) | X | – | – | – | – | – | – | – |
| C. spongillovorax (Resh, 1974) | X | – | – | – | – | – | – | – |
| C. transversa (Hagen, 1861) | X | – | – | – | – | – | – | – |
| Leptocerus americanus (Banks, 1899) | – | – | – | – | – | X | X | – |
| Mystacides interjectus (Banks, 1914) | X | – | – | – | – | – | – | – |
| Nectopsyche diarina (Ross, 1944) | X | – | – | – | – | – | – | – |
| N. exquisita (Walker, 1852) | X | – | – | – | – | – | – | – |
| N. pavida (Hagen, 1861) | X | – | – | – | – | – | – | – |
| Oecetis avara (Banks, 1905) | – | – | – | – | – | – | X | – |
| O. ditissa Ross, 1966 | – | – | – | – | – | – | – | X |
| O. immobilis (Hagen, 1861) | X | – | – | – | – | – | – | – |
| O. nocturna Ross, 1966 | – | – | – | – | X | – | – | X |
| O. ochracea Curtis, 1825 | X | – | – | – | – | – | – | – |
| Triaenodes aba Milne, 1935 | X | – | – | – | – | – | – | – |
| T. baris Ross, 1938 | X | – | – | – | – | – | – | – |
| T. cumberlandensis Etnier & Way, 1973 | – | X | – | – | – | – | – | – |
| T. ignitus (Walker, 1852) | X | – | – | – | – | – | – | – |
| T. marginatus Sibley, 1926 | X | – | – | – | – | – | – | – |
| T. melaca Ross, 1947 | X | – | – | – | – | – | – | X |
| LIMNEPHILIDAE | ||||||||
| Asynarchus mutatus (Hagen, 1861) | – | – | – | – | – | – | – | X |
| Chilostigmodes aeroelatus (Walker, 1852) | – | – | – | X | – | – | – | – |
| Hydatophylax argus (Harris, 1869) | X | – | – | – | – | – | – | – |
| Ironoquia punctatissima (Walker, 1852) | X | – | – | – | – | – | – | – |
| Limnephilus castor Ross & Merkley, 1952 | – | – | – | – | – | – | X | – |
| L. femoralis Kirby, 1837 | – | – | – | – | – | – | – | X |
| Platycentropus amicus (Hagen, 1861) | X | – | – | – | – | – | – | – |
| Pseudostenophylax uniformis (Betten, 1934) | X | – | – | – | – | – | – | – |
| Psychoglypha subborealis (Banks, 1924) | – | – | – | X | – | – | – | – |
| Pycnopsyche guttifera (Walker, 1852) | X | – | – | – | – | – | – | – |
| PHILOPOTAMIDAE | ||||||||
| Chimarra aterrima Hagen, 1861 | X | – | – | – | – | – | – | – |
| C. obscura (Walker, 1852) | X | – | – | – | – | – | – | – |
| Dolophilodes distincta (Walker, 1852) | – | X | – | – | – | – | – | – |
| Wormaldia moesta (Banks, 1914) | X | – | – | – | – | – | – | – |
| W. shawnee (Ross, 1938) | – | – | X | – | – | – | – | X |
| PHRYGANEIDAE | ||||||||
| Agrypnia straminea Hagen, 1873 | – | – | X | – | – | – | – | – |
| A. vestita (Walker, 1852) | X | – | – | – | – | – | – | – |
| Oligostomis pardalis (Walker, 1852) | – | – | – | – | – | – | – | X |
| Ptilostomis angustipennis (Hagen, 1873) | – | X | – | – | – | – | – | – |
| Polycentropodidae | ||||||||
| Cernotina spicata Ross, 1938 | – | – | – | X | – | – | – | X |
| Holocentopus melanae Ross, 1938 | – | – | – | – | – | – | – | X |
| H. picicornis (Stephens, 1836) | – | – | – | – | X | – | – | – |
| Neureclipsis piersoni Frazer & Harris, 1991 | – | X | X | – | – | – | – | – |
| Nyctiophylax moestus Banks, 1911 | – | – | – | – | – | – | X | – |
| Plectrocnemia albipuncta Banks, 1930 | – | – | – | – | – | – | – | X |
| P. clinei Milne, 1936 | – | – | – | – | – | – | – | X |
| P. icula (Ross, 1941) | – | – | – | – | – | – | – | X |
| Polycentropus centralis Banks, 1914 | X | – | – | – | – | – | – | – |
| P. confusus Hagen, 1861 | X | – | – | – | – | – | – | – |
| Psychomyiidae | ||||||||
| Psychomyia flavida Hagen, 1861 | X | – | – | – | – | – | – | – |
| Rhyacophilidae | ||||||||
| Rhyacophila vibox Milne, 1936 | X | – | – | – | – | – | – | – |
| Thremmatidae | ||||||||
| Neophylax ayanus Ross, 1938 | – | X | – | – | – | – | – | – |
| Total | 62 | 12 | 5 | 4 | 4 | 1 | 18 | 25 |
Michigan (319) had the greatest species richness, followed by Kentucky (296), Minnesota (292), and Wisconsin (284) (Fig.
The 22 species that are global endemics to the Upper Midwestern region, organized by family and genus, and with known number of collection localities and recent collection year. Superscript references are after the table.
| Taxon | IL | KY | MI | MN | MO | ND | NE | OH | No. localities | Collected |
|---|---|---|---|---|---|---|---|---|---|---|
| GLOSSOSOMATIDAE | ||||||||||
| Agapetus artesus Ross, 1938 | – | – | – | – | X | – | – | – | 3 | 2017a |
| Protoptila talola Denning, 1948 | – | – | – | X | – | – | – | – | 1 | 1941b |
| Hydroptilidae | ||||||||||
| Hydroptila danieli Harris & Armitage, 2011 | – | – | – | – | – | – | – | X | 6 | 1998c |
| H. howelli Houp, Houp & Harris, 1998 | – | X | – | – | – | – | – | – | 3 | 1998d |
| H. kuehnei Houp, Houp, & Harris, 1998 | – | X | – | – | – | – | – | – | 5 | 1998d |
| H. paraxella Harris & Armitage, 2011 | – | X | – | – | – | – | – | X | 3 | 2008c |
| Neotrichia paraokopa Keth, 2015 | – | – | – | – | X | – | – | – | 1 | 2013d |
| N. staufferi Keth, 2015 | X | – | – | – | – | – | – | – | 1 | 2013d |
| Oxyethira itascae Monson & Holzenthal, 1993 | – | – | X | X | – | – | – | – | ~20 | 2014e |
| LEPTOCERIDAE | ||||||||||
| Ceraclea brevis (Etnier, 1968) | – | – | – | X | – | – | – | – | 1 | 1965b |
| C. erulla (Ross, 1938) | – | – | – | – | – | – | – | X | 1 | 1930sb |
| C. maccalmonti Moulton & Stewart, 1992 | – | – | – | – | X | – | – | – | 2 | 2002f |
|
Setodes truncatus |
– | – | X | – | – | – | – | – | 2 | 2019e |
| Triaenodes phalacris Ross, 1938 | – | – | – | – | – | – | – | X | 1 | 1930sb |
| LIMNEPHILIDAE | ||||||||||
| Chilostigma itascae Wiggins 1975 | – | – | – | X | – | – | – | – | 4 | 2020e |
| Glyphopsyche missouri Ross, 1944 | – | – | – | – | X | – | – | – | 2 | 2017a |
| Ironoquia plattensis Alexander & Whiles, 2000 | – | – | – | – | – | – | X | – | ~25 | 2013g |
| Polycentropodidae | ||||||||||
| Cernotina ohio Ross, 1939 | – | – | – | – | – | – | – | X | 1 | 1930sb |
| Holocentropus chellus (Denning, 1964) | – | – | – | – | – | X | – | – | 1 | 1960sb |
| H. milaca (Etnier, 1968) | – | – | X | X | – | – | – | – | 6 | 2021e |
| Plectrocnemia sabulosa (Leonard & Leonard, 1949) | – | – | X | – | – | – | – | – | 5 | 2019e |
| Polycentropus neiswanderi Ross, 1947 | X | X | – | – | – | – | – | X | 4 | 1990sc,d |
The NMDS ordination of species presence or absence per state produced a two-dimensional solution (Fig.
Hydroptilidae (117 species) was the most species rich family, followed by Limnephilidae (82), and Leptoceridae (76) (Fig.
Discussion
The majority of our reported new state records are species found in at least one other Upper Midwest state. Many of these species, such as Ceraclea maculata (Banks) (Leptoceridae) in South Dakota or Psychomyia flavida Hagen (Psychomyiidae) in Iowa, are common and widespread throughout the region. Thus, their recent discovery almost certainly reflects a lack of collecting in particular states.
Conversely, a few of our reported species represent some interesting range extensions. Chilostigmodes aeroelatus (Walker) (Limnephilidae) is known throughout Alaska and Canada (
The 22 documented endemic species represent 4% of the total caddisfly fauna of the Upper Midwest. Not surprisingly, most of these species are rare and have been found at < 10 total localities throughout their ranges (Table
The congruence of state species assemblages with geographic location was noteworthy and probably due to a combination of factors. Both latitude and longitude have been previously shown to affect caddisfly assemblages (
The total determined caddisfly species richness of the Upper Midwest region currently represents 37% of all described species from the United States and Canada, as well as 63% of genera and 81% of families (
Acknowledgements
Primary funding for this research came from a U.S. Environmental Protection Agency Science to Achieve Results Fellowship and Minnesota Natural Heritage and Nongame Research Program grant to DCH, and from U.S. Department of Interior (INT RD X-1-R-1), National Science Foundation (DEB 09-18805 ARRA), Illinois State Wildlife (IDNR FWS T-121-R-1), and Indiana Department of Natural Resources (E16-21-40777, 0017556043) grants to RED. Further support came from the Hillsdale College (HC) biology department, grants from the Huron Mountain Wildlife Foundation, several HC LAUREATES grants to DCH and affiliated students, and a HC Faculty Summer Leave grant. Funding to support KLS for surveys of state-owned properties in Wisconsin came from the Wisconsin Department of Natural Resources, Natural Heritage Inventory.
We appreciate the efforts of all who have collected, sorted, and identified caddisflies from this large region over the last 20 years, including Benjamin Albers, Geoffrey Archibald, Doug Bidlack, Dean Blinn, Chris Bowyer, Kelsey Brakel, Kiralyn Brakel, Henrey Deese, Mikayla Dove, Lily Erickson, David Etnier, Christine Fenendael, Erin Flaherty, Mike Floyd, Erin Furmaga, Mark Galatowitsch, James Glover, Hannah Goble, Kim Ha, Lily Hart, Robert Kintz, Kyler Kuzio, Ryan Lardner, Grace Lewis, Travis Ling, Faith Linton, Brooklyn Little, Caitlin Lowry, Connie Loruss, Emily Malcolm, Bilyana McLeod, Evan Newman, Bridget O’Leary, Joel Parker, Sally Petrella, Megan Phelps, David Ruiter, Sarah Salow, Karen Schultz, Guenter Schuster, Logan Shoup, Mary Clare Smith, Eric South, Erich Steger, Peter Thistleton, Shelby Tone, Sydney Tone, Eleanor Valle, Jeff Van Zant, Lydia Wassink, Daniel Wright, Mia Young, Jennifer Zaspel, and Jessica Zeglin. Special thanks are due to Johanna Birchem, Nick Connell, Jared Engresser, Kyle Johnson, Gretchen Mehmel, and Charlie Tucker for collecting specimens of Chilostigmodes aeroelatus. We also appreciate access to the vast larval specimen databases maintained by the Iowa and Wisconsin Departments of Natural Resources.
Permits to collect in the state parks of Michigan and Minnesota were provided by Alicia Selden (Michigan Department of Natural Resources) and Ed Quinn (Minnesota Department of Natural Resources), respectively. The sampling of Sleeping Bear Dunes National Lakeshore was conducted under permit SLBE-2014-SCI-0002, facilitated by Kevin Skerl. The Saint Croix National Scenic Waterway was sampled under permits SACN-2013-SCI-0003 and SACN-2016-SCI-0001, facilitated by Jill Midland. The staff at Pictured Rocks National Lakeshore granted access to streams under permits PIRO-2010-0008 and PIRO-2013-0002. Permission to sample in the Huron Mountains of Michigan was provided by the Huron Mountain Wildlife Foundation. Permission to sample at the Black River Ranch of Michigan was provided by the Black River Ranch Board of Directors. Permission to sample at Sarah Jane’s Natural Area of Michigan was provided by John Bagley and Andrew Bacon (Michigan Nature Association).
Google Earth base maps were used following permission guidelines (https://www.google.com/permissions/geoguidelines/attr-guide/). The valuable comments of Desiree Robertson and Paul Frandsen improved earlier version of the manuscript. This is paper #30 of the G.H. Gordon BioStation Research Series.
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Supplementary materials
Current checklist of 552 caddisfly species known from the Upper Midwest region
Data type: species data
Collection data for the 131 new state species records
Data type: species data