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Vol. 51, No. 1
MICHIGA
Tribute to Ed Voss
A Journal of Great Lakes Botany
THE MICHIGAN BOTANIST (ISSN 0026- 203X) is published four times per year by the Michigan
Botanical Club (www.michbotclub.org) and is available online at http://quod.lib.umich.edu/my
mbot/. The subscription rate is $25.00 per year. Periodicals postage paid at Ann Arbor, MI 48103.
The office of publication is Western Michigan University, Kalamazoo, MI 49008.
On all editorial matters, please contact Todd J. Barkman, 3437 Wood Hall, Department of Biological
Sciences, Western Michigan University, Kalamazoo, MI 49008; 269. 387. 2776 (Phone), 269.
387. 5609 (FAX); todd.barkman@wmich.edu. All articles dealing with botany in the Great Lakes
region may be sent to the Editor at the above address. In preparing manuscripts, authors are
requested to follow the “Instructions for Authors” on the inside back cover.
For all inquiries about back issues and institutional subscriptions please contact Caroline Barkman,
The Michigan Botanist Business Office, 919 Dobbin Dr., Kalamazoo, MI 49006; 269. 544. 0034;
cbarkman9 1 47 @ mail.kvcc.edu
Todd J. Barkman, Editor
Editorial Board
Caroline Barkman, Business Manager
L. Alan Prather
Anton A. Reznicek
J. Dan Skean, Jr.
Anna K. Monfils
Timothy M. Evans
Catherine H. Yansa
THE MICHIGAN BOTANICAL CLUB
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the public to appreciate and preserve plant life; sponsorship of research and publication on the
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riverview.net
2012
THE MICHIGAN BOTANIST
1
EDWARD G. VOSS (1929-2012)
Anton A. Reznicek and Richard K. Rabeler
University of Michigan Herbarium
3600 Varsity Drive
Ann Arbor, MI 48108-2228
Edward Groesbeck Voss (Fig. 1), Professor and Curator Emeritus at the Uni¬
versity of Michigan Department of Ecology and Evolutionary Biology and Uni¬
versity of Michigan Herbarium, died on February 13, 2012 at his home in Ann
Arbor, after a brief illness. He was bom on Febmary 22, 1929 in Delaware,
Ohio. Ed’s abilities were clear early in his life. He attended Woodward High
School in Toledo, Ohio. He received the Achievement Award as the outstanding
senior in his 1946 high school class, and was later elected to the Woodward High
Hall of Fame. He then graduated with a bachelor’s degree with honors from
Denison University (1950); his undergraduate honors thesis, On the classifica¬
tion of the Hesperiidae, became a significant paper on Lepidoptera (Voss 1952).
He completed his education with a master’s in Biology (1951) and a doctoral de¬
gree in Botany (1954), both from the University of Michigan working with
Rogers McVaugh. He was awarded an honorary Doctor of Science from Denison
University in 2003.
In 1956, he began his long association with the University of Michigan and
Michigan’s flora when he was appointed as a research associate at the Herbar¬
ium, beginning what was expected to be a five-year project to document the flora
of Michigan. He joined the Botany faculty in 1960 as assistant professor and was
promoted to associate professor (1963), and later, professor (1969). He contin¬
ued his painstakingly precise study of the flora of Michigan, culminating 40
years later with the publication of the third volume of the Michigan Flora in
1996; the same year that he retired and was granted Professor Emeritus status.
He also served as Curator of Vascular Plants at the Herbarium from 1961 until
his retirement in 1996 and continued to work at the herbarium until late in 201 1.
Ed’s research focused on the vascular plants of the Great Lakes region: their
taxonomy, phytogeography, and status in natural environments; with a particular
interest in boreal plants and aquatic plants. He maintained a small herbarium at
his cottage in Mackinaw City, but most of his collections went to the University
of Michigan Herbarium, with modest numbers also helping to build up the Bio¬
logical Station herbarium. Ed founded and edited the first 15 volumes of The
Michigan Botanist and served on the Editorial Board until his death.
Ed also continued his early interest in Lepidoptera throughout his life. He col¬
lected and studied them extensively, publishing a number of articles on the Lep¬
idoptera of the Douglas Lake region (Emmet and Cheboygan counties). He was
honored in 201 1 as a founding member of the Lepidopterists’ Society.
Coincident with his floristic research, Ed was a long time instructor at The
University of Michigan Biological Station (UMBS). Ed began at the station in
2
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 1. Ed Voss beloved friend, mentor, teacher, and colleague. Photo by John Russell.
1949 as a teaching assistant for Rogers McVaugh; during the next several sum¬
mers he assisted in courses taught by Carroll Wood, Lloyd Shinners,1 and Elzada
Clover. From 1963 to 2003 (missing only 1975 and 1999-2002), Ed taught
Aquatic Flowering Plants (Fig. 2), Taxonomy of Flowering Plants, Boreal Flora,
and Field Botany of Northern Michigan at the Station. He continued teaching
“minicourses” through 2009. He was honored at the Station by a special “Dr. Ed¬
ward Voss Celebration” Weekend in 2009. Ed had received an early introduction
to the station when his parents took Ed, then 9 years old, to a “Visitor’s Day” at
UMBS. On the Ann Arbor campus, Ed taught Aquatic Flowering Plants, from
1961-93, Writing for Biologists in 1987, 1988, 1990, and 1992, and as one of the
instructors in the Plant Biology course from 1974-79. Post-retirement, Ed taught
popular courses and led field trips through the Adult Education Program of the
Matthaei Botanical Gardens and workshops in the Upper Peninsula on aquatic
plants, general plant identification, and boreal flora for Forest Service personnel
and other people working with stewardship.
Ed gave long service to plant nomenclature via membership on the General
and Editorial Committees responsible for the revision of the International Code
of Botanical Nomenclature. He was a member of the Editorial Committee from
1969 to 1993 (i.e. from the Seattle to the Berlin Codes), served as the Secretary
•See Rabeler & Reznicek, 2012. J. Bot. Res. Inst. Texas 6: 311-312.
2012
THE MICHIGAN BOTANIST
3
FIGURE 2. Ed Voss with his Aquatic Flowering Plants class in 1965. Photo by John Russell.
of the Committee for the Seattle and Leningrad Codes, and Chair for the Sydney
Code.2
The first volume of Michigan Flora was honored by a Resolution of the
Michigan Senate in 1972, while the second volume received the H.A. Gleason
Award from the New York Botanical Garden in 1986. The Michigan Botanical
Club also presented a lifetime achievement award to Ed in 1998, following pub¬
lication of the final volume of Michigan Flora in 1996.
He was devoted to the conservation of natural areas, and served two terms as
a member of the Board of Trustees of the Nature Conservancy of Michigan and
was a long time Trustee of the Little Traverse Conservancy. He served on a num¬
ber of citizens advisory committees for environmental stewardship and also
served on the Technical Advisory Committee for Plants for the Michigan Threat¬
ened & Endangered Species Program and was a member of the equivalent com¬
mittee for Ohio. Throughout all this service, he worked to educate others around
him, always concerned that the information about the flora being used in deci¬
sion making was as accurate as possible.
He was well known to students, professionals, and amateurs alike, sharing his
passion for botany equally with all. His precision, wit (often as clever puns),
command of the English language, and propensity for being thrifty all added up
to make him “Ed” — a unique man who left his mark on the study of the Michi¬
gan flora and nomenclature alike.
2 See: Rabeler & Reznicek, 2012. Taxon 61: 697.
4
THE MICHIGAN BOTANIST
Vol. 51
As a tribute to Ed and his work on the plants of Michigan, we present a series
of invited papers that we hope will give more insight into the various “sides” of
Ed, as well as basic biographical data including a hopefully complete bibliogra¬
phy, eponymy, list of new combinations and names resulting from Ed’s research,
plus a list of graduate students whose committee Ed chaired or co-chaired.
BIBLIOGRAPHY
This bibliography is split into three parts: Ed’s formal publications, his Michigan
Plants in Print series, and his numerous book reviews. Items in the formal bibliography
are listed by the actual date published, even if the date printed on the publication differs.
For the Michigan Plants in Print and book reviews, the date printed in the journal is ac¬
cepted. Numerous ephemeral items, such as his many pungent “Letters to the Editor” to
newspapers and journals, as well as editorial notices and other such items in The Michi¬
gan Botanist, are not included.
Note: Articles by Ed cited in the papers following are to be found here and are not re¬
peated in the literature cited (if any) in the individual articles.
1948
Voss, E. G. Observations on the Michigan flora: A survey of the St. Ignace Causeway
(Mackinac Co.). Jour. Sci. Labs. Denison Univ. 41: 39^f5.
1949
Voss, E. G. Observations on the Michigan flora, II: Some plants new or rare in Emmet,
Cheboygan, and Mackinac counties, Michigan. Jour. Sci. Labs. Denison Univ. 41:
77-81.
1950
Voss, E. G. Catocala feigning death. Lepidopterists’ News 4: 46.
Voss, E. G. Lepidoptera strays in Northern Michigan. Lepidopterists’ News 4: 46.
Voss, E. G. Making venation visible. Lepidopterists’ News 4: 70.
Voss, E. G. Observations on the Michigan flora, III: The flora of Green Island (Mackinac
County). Ohio Jour. Sci. 50: 182-190.
1952
Voss, E. G. Floristic study in the Douglas Lake region of Michigan, 1951. Asa Gray Bull,
N. S. 1: 51-52.
Voss, E. G. “Official Views” from the GPO. Science 115: 445.
Voss, E. G. On the classification of the Hesperiidae. Ann. Entomol. Soc. Amer. 45:
246-253.
Voss, E. G. The history of keys and phylogenetic trees in systematic biology. Jour. Sci.
Labs. Denison Univ. 43: 1-25.
1953
Voss, E. G. Observations on the Michigan flora, IV: A botanical survey in Huron County.
Asa Gray Bull., N. S. 2: 17-24.
Voss, E. G. Butterflies and crab spiders. Lepidopterists’ News 7: 54.
Voss, E. G. Observations on the Michigan flora, V: Plants of the St. Ignace Causeway
(Mackinac County) — The second four years. Jour. Sci. Labs. Denison Univ. 43: 53-60.
1954
Voss, E. G. The butterflies of Emmet and Cheboygan counties, Michigan, with other
notes on Northern Michigan butterflies. Amer. Midi. Naturalist 51: 37-104.
2012
THE MICHIGAN BOTANIST
5
Voss, E. G. The Vascular Plants of Emmet and Cheboygan counties, Michigan: Patterns
and Problems in Geographic Affinities. Ph.D. dissertation. 294 pp. [Abstract in Disser¬
tation Abstracts 15: 1020-1021. 1954.]
1955
Voss, E. G. Death of Frank C. Gates. Asa Gray Bull., N. S. 3: 32.
Voss, E. G. Charles W. Fallass (1854-1942), a pioneer Michigan Botanist. Asa Gray
Bull., N. S. 3: 77-96.
1956
Voss, E. G. A history of floristics in the Douglas Lake region (Emmet and Cheboygan
counties), Michigan, with an account of rejected records. Jour. Sci. Labs. Denison Univ.
44: 16-75.
Voss, E. G., & W. H. Wagner, Jr. Notes on Pieris virginiensis and Erora laeta — Two but¬
terflies hitherto unreported from Michigan. Lepidopterists’ News 10: 18-24.
Voss, E. G., & J. S. Wilson. Clarence R. Hanes. Rhodora 58: 272-273.
1957
Voss, E. G. New records of vascular plants from the Douglas Lake region (Emmet and
Cheboygan counties), Michigan. Pap. Michigan Acad. Sci. 42: 3-34.
Voss, E. G. Observations on the Michigan flora, VI: Distribution records of some an-
giosperms new, rare, or misinterpreted in the State. Brittonia 9: 83-101.
Voss, E. G. The Michigan flora project. Taxon 6: 156-157.
1958
Voss, E. G. Confusion in Alisma. Taxon 7: 130-133.
1959
Voss, E. G. The neglected sedges. Cranbrook Inst. Sci. Newsl. 23: 86-92.
Voss, E. G. Memorandum on the proposed diversion of additional water from Lake
Michigan at Chicago. Northern Michigan Skipper 7(4): 3 & 12.
1961
Voss, E. G. Which side is up? A look at the leaves of Oryzopsis. Rhodora 63: 285-287.
Voss, E. G. Harley Harris Bartlett. Bull. Torrey Bot. Club 88: 47-56. [Reprinted in
Michigan Pap. South & Southeast Asia 5: xv-xxiv (1973) and Bartlettia 2(3): 37-47
(1982)]
Voss, E. G. Burt Lake named after Michigan Baptist pioneer. Michigan Baptist 44(2): 7.
Voss, E. G. Harley H. Bartlett 1886-1960. Records Genetics Soc. Amer. 30: 13-15.
1962
Voss, E. G. Ailanthus promotion a fraud? Michigan Bot. 1: 22.
Rittenhouse, J. L., & E. G. Voss. Douglass Houghton’s botanical collections in Michigan,
Wisconsin, and Minnesota on the Schoolcraft Expedition of 1832. Michigan Bot. 1:
61-70.
Voss, E. G. (editor), The Michigan Bot. Vol. 1, No. 1 & 2.
1963
Guire, K., & E. G. Voss. Distributions of distinctive shoreline plants in the Great Lakes
region, Michigan Bot. 2: 99-114.
Voss, E. G. Arthur Ward Lindsey (1394-1963). Jour. Lepidopterists’ Soc. 17: 181-190.
Voss, E. G. American Association of Commons Clubs. In Baird’s Manual of American
College Fraternities, 17th ed., John Robson (ed.). George Banta Co., Menasha, Wis.,
pp. 362-364.
Voss, E. G. (editor), The Michigan Bot. Vol. 2, No. 1, 2, 3, & 4.
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THE MICHIGAN BOTANIST
Vol. 51
1964
Soper, J. H., & E. G. Voss. Black Crowberry in the Lake Superior region, Michigan Bot.
3: 35-38.
Voss, E. G. The Michigan Lily. Michigan Bot. 3: 44-47.
Voss, E. G. Skunk-cabbage in Michigan. Michigan Bot. 3: 97-101.
Voss, E. G. Entomologists and plants. Newsl. Michigan Entomol. Soc. 9. (2): 2-3.
Voss, E. G. Founding of First University Hospital. Jour. Med. Educ. 39: 797-798.
[reprinted in Ann Arbor News, Reader’s Viewpoint, Feb. 22, 1964 and Cincinnati
Alumnus, April 1964]
Voss, E. G. (editor), The Michigan Bot. Vol. 3, No. 1, 2, 3, & 4.
1965
Voss, E. G. Some rare and interesting aquatic vascular plants of Northern Michigan, with
special reference to Cusino Lake (Schoolcraft Co.). Michigan Bot. 4: 11-25.
Voss, E. G. Our lone native primrose. Michigan Audubon Newsl. 13(2): 4.
Soper, J. H., E. G. Voss, & K. E. Guire. Distribution of Primula mistassinica in the Great
Lakes region. Michigan Bot. 4: 83-86. [reprinted in Quart. Jour. Amer. Primrose Soc.
30: 108-112. 1972.]
Voss, E. G. A review on Crataegus. Michigan Bot. 4: 93-96.
Voss, E. G. Comparative revegetation of denuded areas in Northern Michigan. Pap.
Michigan Acad. Sci. 50: 139-160.
Voss, E. G. On citing the names of publishing authors. Taxon 14: 154-160.
Voss, E. G. A weedy orchid? Michigan Audubon Newsl. 13 (4): 4.
Voss, E. G. (editor), The Michigan Bot. Vol. 4, No. 1, 2, 3, & 4.
1966
Voss, E. G. The little bellwort. Michigan Audubon Newsl. 13(5): 3-4.
Voss, E. G. Nomenclatural notes on monocots. Rhodora 68: 435^463.
Voss, E. G. Two kinds of beauties. Michigan Audubon Newsl. 14(1): 7-8.
Voss, E. G. Carnivora of the plant world. Michigan Audubon Newsl. 14(3): 4-5.
Voss, E. G. (editor), The Michigan Bot. Vol. 5, No. 1, 2, 3, & 4.
1967
Voss, E. G. The status of some reports of vascular plants from Michigan. Michigan Bot.
6: 13-24.
Voss, E. G. A vegetative key to the genera of submersed and floating aquatic vascular
plants of Michigan. Michigan Bot. 6: 35-50.
Voss, E. G. Michigan mistletoe. Michigan Audubon Newsl. 14 (6): 5.
Voss, E. G. (editor), The Michigan Bot. Vol. 6, No. 1, 2, 3, & 4.
1968
Voss, E. G. The spring beauties ( Claytonia ) in Michigan. Michigan Bot. 7: 77-93.
Voss, E. G. By-ways of the North Shore. Michigan Audubon Newsl. 16(2): 4-5; 16(3): 4.
Voss, E. G. A preliminary report on the distribution of gymnosperms in Michigan. Michi¬
gan Bot. 7: 121-128.
Voss, E. G. (editor). The Michigan Bot. Vol. 7, No. 1, 2, 3, & 4.
1969
Voss, E. G. Appendix: Nomenclatural changes. In Our Northern Shrubs, by H. L. Keeler
(reprinted by Dover Publications), pp. 513-528.
Voss, E. G. Nomenclature. Encyclopaedia Britannica 21: 730A-731.
Stafleu, F. A., & E. G. Voss. Synopsis of proposals on botanical nomenclature Seattle
1969. Regnum Veg. 60. 124 pp.
Voss, E. G. (editor), The Michigan Bot. Vol. 8, No. 1, 2, 3, & 4.
2012
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7
1970
Voss, E. G. Charles H. Swift’s botanical work in Michigan. Michigan Bot. 9: 38 — 40.
Moore, H. E., F. A. Stafleu, & E. G. Voss. XI International Botanical Congress: Final mail
vote and congress action on nomenclature proposals. Taxon 19: 43-51.
Voss, E. G. Moths of the Douglas Lake region (Emmet and Cheboygan counties), Michi¬
gan: I. Sphingidae — Ctenuchidae (Lepidoptera). Michigan Entomol. 2: 48-54.
Voss, E. G. (editor), The Michigan Bot. Vol. 9, No. 1, 2, 3, & 4.
1971
Voss, E. G. (editor), The Michigan Bot. Vol. 10, No. 1, 2, 3, & 4.
1972
Voss, E. G. Additional nomenclatural and other notes on Michigan monocots and gym-
nosperms. Michigan Bot. 11: 26-37.
Voss, E. G. Michigan Flora. A Guide to the Identification and Occurrence of the Native
and Naturalized Seed-plants of the State. Part I Gymnosperms and Monocots. Cran-
brook Inst. Sci. Bull. 55 & Univ. Michigan Herbarium, xviii + 488 pp.
Stafleu, F. A., & E. G. Voss. Report on Botanical Nomenclature Seattle 1969. Regnum
Veg. 81. 133 pp.
Stafleu, F. A., C. E. B. Bonner, R. McVaugh, R. D. Meikle, R. C. Rollins, R. Ross, J. M.
Schopf, G. M. Schultz, R. De Vilmorin, & E. G. Voss. International Code of Botanical
Nomenclature adopted by the Eleventh International Botanical Congress Seattle, Au¬
gust 1969. Regnum Veg. 82. 426 pp.
Voss, E. G. The State of Things: Contributions of the plant taxonomist in Michigan.
Michigan Academician 5: 1-7.
Voss, E. G. (editor), The Michigan Bot. Vol. 11, No. 1, 2, 3, & 4.
1973
Voss, E. G. Wild Plants in Flower II: The Boreal Forest and Borders. By Torkel Korling,
with notes on the species and their distributions by Edward G. Voss. Dundee, IL. 72 pp.
Voss, E. G. General Committee Report, 1970-1971. Taxon 22: 153.
Voss, E. G. General Committee Report, 1972. Taxon 22: 313.
Voss, E. G. Errata in Seattle Code. Taxon 22: 503.
Voss, E. G. (editor), The Michigan Bot. Vol. 12, No. 1, 2, 3, & 4.
1974
Voss, E. G. Corrections in lists of Nomina Conservanda et Rejicienda. Taxon 23:
647-648.
Voss, E. G. (editor), The Michigan Bot. Vol. 13, No. 1, 2, 3, & 4.
1975
Stafleu, F. A., & E. G. Voss. Synopsis of proposals on botanical nomenclature Leningrad
1975. Taxon 24: 201-254.
Voss, E. G. (editor), The Michigan Bot. Vol. 14, No. 1, 2, 3, & 4.
1976
Voss, E. G., & G. E. Crow. Across Michigan by covered wagon: A botanical expedition in
1888. Michigan Bot. 15: 3-70.
Voss, E. G. XII International Botanical Congress: Mail vote and final Congress action on
nomenclature proposals. Taxon 25: 169-174.
Voss, E. G. General Committee Report, 1975-1976. Taxon 25: 511-512.
Voss, E. G. (editor), The Michigan Bot. Vol. 15, No. 1, 2, 3, & 4.
1977
Voss, E. G. Advice on proposals to amend the Code. Taxon 26: 309-310.
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THE MICHIGAN BOTANIST
Vol. 51
Wagner, W. H., E. G. Voss, J. H. Beaman, E. A. Bourdo, F. W. Case, J. A. Churchill, & P.
W. Thompson. Endangered, threatened and rare vascular plants in Michigan. Michigan
Bot. 16: 99-110.
Voss, E. G. Additions and corrections to the list of vascular plants from the Douglas Lake
Region, Michigan. Michigan Bot. 16: 126-141.
1978
Voss, E. G., & M. W. Bohlke. The status of certain hawkweeds ( Hieracium subgenus Pi-
losella) in Michigan. Michigan Bot. 17: 35—47.
Voss, E. G. Botanical Beachcombers and Explorers: Pioneers of the 19th Century in the
Upper Great Lakes. Contr. Univ. Michigan Herb. 13. viii + 100 pp.
Stafleu, F. A., V. Demoulin, W. Greuter, P. Hiepko, A. Linczevski, R. McVaugh, R. D.
Meikle, R. C. Rollins, R. Ross, J. M. Schopf, & E. G. Voss. International Code of
Botanical Nomenclature Adopted by the Twelfth International Botanical Congress,
Leningrad, July 1975. Regnum Veg. 97. xiv + 457 pp.
1980
Voss, E. G. Twelfth International Botanical Congress Leningrad 1975. Nomenclature
Section Report. Proc. XII Internatl. Bot. Congr., pp. 129-186.
Voss, E. G. General Committee Report 1977-1980. Taxon 29: 689.
1981
Voss, E. G., & W. Greuter. Synopsis of proposals on botanical nomenclature Sydney
1981. Taxon 30: 95-293.
Voss, E. G., & A. A. Reznicek. News of herbaria and collections: Hermann herbarium.
Taxon 30: 866.
Marquis, R. J., & E. G.Voss. Distributions of some western North American plants dis¬
junct in the Great Lakes region. Michigan Bot. 20: 53-82.
Voss, E. G. Moths of the Douglas Lake region (Emmet and Cheboygan counties), Michi¬
gan: II. Noctuidae (Lepidoptera). Great Lakes Entomol. 14: 87-101.
1982
Voss, E. G. Nomenclature at Sydney. Taxon 31: 151-154.
Voss, E. G. Nomenclature Committee Reports. Taxon 31: 310-318.
Greuter, W., & E. G. Voss. Report on Botanical Nomenclature — Sydney 1981. Englera
2. 124 pp.
Voss, E. G. Announcement: Nomenclature Committees. Taxon 31: 718.
1983
Voss, E. G., & R. E. Riefner, Jr. A Pyralid Moth (Lepidoptera) as pollinator of Blunt-Leaf
Orchid. Great Lakes Entomol. 16: 57-60.
Voss, E. G., H. M. Burdet, W. G. Chaloner, V. Demoulin, P. Hiepko, J. McNeill, R. D.
Meikle, D. H. Nicholson, R. C. Rollins, P. C. Silva, & W. Greuter. International Code
of Botanical Nomenclature Adopted by the Thirteenth International Botanical Con¬
gress, Sydney, August 1981. Regnum Veg. 111. xv + 472 pp.
1984
Voss, E. G. Moths of the Douglas Lake region (Emmet and Cheboygan counties), Michi¬
gan: III. Thyatiridae, Drepanidae, Lasiocampidae, Notodontidae, Lymantriidae (Lepi¬
doptera). Great Lakes Entomol. 16: 131-137.
1985
Voss, E. G. Nomenclatural notes on some Michigan dicots. Michigan Bot. 24: 117-124.
Beaman, J. H., E. A. Bourdo, F. W. Case, S. R. Crispin, D. Henson, R. W. Pippen, A. A.
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9
Reznicek, E. G. Voss, & P. W. Thompson. Endangered and threatened vascular plants in
Michigan. II. Third Biennial Review Proposed List. Michigan Bot. 24: 99-116.
Voss, E. G. Michigan Flora. Part II Dicots (Saururaceae — Cornaceae). Cranbrook Inst.
Sci. Bull. 59 & Univ. Michigan Herbarium, xx + 724 pp.
1986
Voss, E. G. Commission on Nomenclature of Plants [report]. Rep. XXIInd General As¬
sembly Inti. Union Biol. Sci., p. 130.
Voss, E. G. Major errata in Michigan Flora, Part II. Michigan Bot. 25: 55.
Voss, E. G. General Committee report 1982-1985. Taxon, 35: 551-552.
Voss, E. G. (23 1)— (242) Proposals to improve the Code. Taxon 35: 827-830.
1987
Voss, E. G. General Committee Report 1986. Taxon 36: 429
1988
Greuter, W., H. M. Burdet, W. G. Chaloner, V. Demoulin, R. Grolle, D. L. Hawksworth,
D. H. Nicholson, P. C. Silva, F. A. Stafleu, E. G. Voss, & J. McNeill. International Code
of Botanical Nomenclature Adopted by the Fourteenth International Botanical Con¬
gress, Berlin, July-August 1987. Regnum Veg. 118. 328 pp.
Voss, E. G., & A. A. Reznicek. Frederick J. Hermann (1906-1987): The evolution of a
botanical career. Michigan Bot. 27: 59-73.
Voss, E. G., & A. A. Reznicek. Frederick Joseph Hermann (1906-1987). Taxon 37:
509-510.
Voss, E. G., & A. A. Reznicek. Frederick Joseph Hermann. Bryologist 90: 466-467.
1990
Voss, E. G. The Practicality of Latin. Taxon 39: 239.
1991
Voss, E. G. Moths of the Douglas Lake region (Emmet and Cheboygan counties), Michi¬
gan: IV. Geometridae (Lepidoptera). Great Lakes Entomol. 24: 187-201.
1996
Voss, E. G. New combinations in the Michigan flora. Michigan Bot. 34: 139-140.
Orchard, A. E., W. R. Anderson, M. G. Gilbert, D. Sebsebe, W. T. Stearn, & E. G. Voss.
Harmonized bionomenclature — a recipe for disharmony. Taxon 45: 287-290.
Voss, E. G. Michigan Flora. Part III Dicots (Pyrolaceae — Compositae). Cranbrook Inst.
Sci. Bull. 61 & Univ. Michigan Herbarium, xii + 622 pp.
2000
Voss, E. G. Additions and corrections to Michigan Flora, Part II. Michigan Bot. 37:
12-13.
2001
Voss, E. G. A purple color form of Pitcher’s Thistle. Contr. Univ. Michigan Herb. 23:
349-350.
Voss, E. G. Flora of St. Helena Island (Straits of Mackinac), Michigan. Michigan Bot. 40:
27-47
2002
Voss, E. G. Labeling of Herbarium Specimens. Michigan Bot. 38: 57-63.
Voss, E. G. Moths of the Douglas Lake region (Emmet and Cheboygan counties), Michi¬
gan: VI. Miscellaneous small families (Lepidoptera). Great Lakes Entomol. 35: 53-61.
10
THE MICHIGAN BOTANIST
Vol. 51
2004
Voss, E. G. (087-088) Proposals to amend Arts. 37 & 46. Taxon 53: 222-223.
Voss, E. G. Additions and corrections to Michigan Flora, Part III. Michigan Bot. 43:
12-13.
A. A. Reznicek, E. G. Voss, & R. A. Simpson. Online Atlas of Michigan Plants. Edition
1. http://herbarium.lsa.umich.edu/website/michflora/onlinemaps.html
2005
Voss, E. G. Gazetteer of some possibly puzzling collecting localities for Michigan plants.
Contr. Univ.Michigan Herb. 24: 189-225. [also available online in a searchable format
at: http://www. Isa. umich. edu/herb/michvoss/index. html ]
2008
Voss, E. G. Seventy seasons (more or less) at or near UMBS. Pp. 246 in M. C. Scholtens,
and K. G. Williams, eds. Grapevine to Pine Point. University of Michigan Biological
Station, Pellston, MI.
2009
Voss, E. G. Demystifying nomenclature: The game of the name. Michigan Bot. 48:
38-44.
Voss, E. G. Chapter 5. Terrestrial Vascular Plants. Pp. 61-68 in K. J. Nadelhoffer, A. J.
Hogg, & B. A. Hazlett, eds., The Changing Environment of Northern Michigan. A Cen¬
tury of Science and nature at the University of Michigan Biological Station. University
of Michigan Press.
2011
Reznicek, A. A., E. G. Voss, & B. S. Walters. Michigan Flora Online, http://
michiganflora.net/
2012
Voss, E. G. & A. A. Reznicek. Field Manual of Michigan Flora. University of Michigan
Press, Ann Arbor, xiv + 990 pp.
THE 116 BOOK REVIEWS WRITTEN BY E. G. VOSS
Where more than one was published in a given year, we have arranged them alpha¬
betically by journal, not by the absolute chronology of when the publications would have
been distributed.
Voss, E. G. 1953. [Review of] H. A. Gleason. The New Britton and Brown Illustrated
Flora of the Northeastern United States and Adjacent Canada. Asa Gray Bull, N. S. 2:
25-28.
Voss, E. G. 1953. [Review of] H. A. Gleason. The New Britton and Brown Illustrated
Flora of the Northeastern United States and Adjacent Canada. Michigan Alumnus
Quarterly Rev. 60: 87-88.
Voss, E. G. 1962. [Review of] H. V. Smith. Michigan Wildflowers. Amer. Midi. Natural¬
ist 68: 509-510.
Voss, E. G. 1963. [Review of] H. A. Gleason & A. Cronquist. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada. Michigan Bot. 2: 61-62.
Voss, E. G. 1963. [Review of] H. A. Gleason & A. Cronquist. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada. Science 140: 637.
Voss, E. G. 1964. [Review of] Webster’s Third New International Dictionary of the Eng¬
lish Language, Unabridged. Sida 1: 389-392.
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THE MICHIGAN BOTANIST
11
Voss, E. G. 1964. [Review of] T. Korling. Wild Plants in Flower. Michigan Bot. 3: 121.
Voss, E. G. 1965. [Review of] H. T. Darlington. The Mosses of Michigan. Michigan Bot.
4: 25.
Voss, E. G. 1965. [Review of] V. S. Eifert. Tall Trees and Far Horizons. Michigan Bot. 4:
82. [reprinted in part in Audubon Magazine. 67: 334.]
Voss, E. G. 1965. [Review of] E. P. Kruschke. Contributions to the Taxonomy of Cratae¬
gus. Michigan Bot. 4: 93-96.
Voss, E. G. 1965. [Review of] T. G. Tutin, V. H. Heywood, N. A. Burges, D. H. Valentine,
S. M. Walters, & D. A. Webb. Flora Europaea. Volume 1. Lycopodiaceae to Pla-
tanaceae. Bull. Torrey Bot. Club 92: 4 1 0 — 4 1 3 .
Voss, E. G. 1965. [Review of] V. H. Heywood & R. E. G. Pichi-Sermolli. Proceedings of
the Second Flora Europaea Symposium. Bull. Torrey Bot. Club 92: 4 1 3^4 14.
Voss, E. G. 1965. [Review of] J. B. Moyle. Northern Non- Woody Plants. A Field Key to
the More Common Ferns and Flowering Plants of Minnesota and Adjacent Regions.
Quarterly Rev. Biol. 40: 295.
Voss, E. G. 1966. [Review of] E. P. Kruschke. Contributions to the Taxonomy of Cratae¬
gus. Amer. Midi. Naturalist 75: 252-253.
Voss, E. G. 1966. [Review of] O. Lakela. A Flora of Northeastern Minnesota. Michigan
Bot. 5: 119.
Voss, E. G. 1966. [Review of] K. L. Jones. Botanical Essays for Humanists. Michigan
Bot. 5: 119.
Voss, E. G. 1967. [Review of] H. W. Rickett. Wildflowers of the United States. Vol. I The
Northeastern States. Michigan Bot. 6: 30.
Voss, E. G. 1967. [Review of] H. V. Smith. Michigan Wildflowers, rev. ed. Michigan Bot.
6: 53.
Voss, E. G. 1967. [Review of] P. W. Thompson. Vegetation and Common Plants of Sleep¬
ing Bear. Michigan Bot. 6: 175.
Voss, E. G. 1967. [Review of] J. L. Taylor & R. Turner, eds. The Nut Jar. A Cookbook.
Michigan Bot. 6: 178.
Voss, E. G. 1967. [Review of] E. L. Braun. The Monocotyledonae Cat-tails — Orchids.
Michigan Bot. 6: 184.
Voss, E. G. 1967. [Review of] E. L. Braun. The Monocotyledonae Cat-tails — Orchids.
Ohio J. Sci. 67: 385.
Voss, E. G. 1968. [Review of] R. T. Peterson & M. McKenny. A Field Guide to Wild¬
flowers of Northeastern and North-central North America. Michigan Bot. 7: 190.
Voss, E. G. 1969. [Review of] J. Vosberg. Living With Your Land. A Guide to Conserva¬
tion for the City’s Fringe. Michigan Bot. 8: 10.
Voss, E. G. 1969. [Review of] A. A. Lindsey, D. V. Schmelz, & S. A. Nichols. Natural
Areas in Indiana and Their Preservation. Michigan Bot. 8: 150.
Voss, E. G. 1969. [Review of] C. D. Sculthorpe. The Biology of Aquatic Vascular Plants.
Econ. Bot. 23: 84-85.
Voss, E. G. 1969. [Review of] J. Stodola. Encyclopedia of Water Plants. Econ. Bot. 23:
195-196.
Voss, E. G. 1969. [Review of] T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore,
D. H. Valentine, S. M. Walters, & D. A. Webb. Flora Europaea. Volume 2. Rosaceae to
Umbelliferae. Bull. Torrey Bot. Club. 96: 499-500.
Voss, E. G. 1970. [Review of] G. W. Prescott. How to Know the Aquatic Plants. Michi¬
gan Bot. 9: 142.
Voss, E. G. 1970. [Review of] F. Swink. Plants of the Chicago Region. Michigan Bot. 9:
165.
12
THE MICHIGAN BOTANIST
Vol. 51
Voss, E. G. 1970. [Review of] J. A. Dorr & D. F. Eschman. Geology of Michigan. Michi¬
gan Bot. 9: 206.
Voss, E. G. 1971. [Review of] A. H. Smith & H. D. Thiers. The Boletes of Michigan.
Michigan Bot. 10: 93.
Voss, E. G. 1971. [Review of] J. E. Fitting. The Archeology of Michigan. Michigan Bot.
10: 62.
Voss, E. G. 1971. [Review of] H. R. Schoolcraft. Narrative Journal of Travels through the
Northwestern Regions of the United States Extending from Detroit through the Great
Chain of American Lakes to the Sources of the Mississippi River, Performed as a Mem¬
ber of the Expedition under Governor Cass (Facsimile Edition). Michigan Bot. 10: 205.
Voss, E. G. 1971. [Review of] L. Agassiz. Lake Superior: Its Physical Character, Vegeta¬
tion, and Animals, Compared with Those of Other and Similar Regions. With a Narra¬
tive of the Tour by J. Elliot Cabot. (Facsimile Reprint). Michigan Bot. 10: 206.
Voss, E. G. 1971. [Review of] A. Karlen, ed. Photographs by C. Steinhacker. Superior.
Michigan Bot. 10: 206.
Voss, E. G. 1971. [Review of] E. L. Little, Jr. Atlas of United States Trees Volume 1.
Conifers and Important Hardwoods. Michigan Bot. 10: 207.
Voss, E. G. 1972. [Review of] G. B. Ownbey. Common Wild Flowers of Minnesota.
Michigan Bot. 11: 12.
Voss, E. G. 1972. [Review of] I. H. Klein. Wild Flowers of Ohio and Adjacent States.
Michigan Bot. 11: 37.
Voss, E. G. 1972. [Review of] S. W. Jackman & J. F. Freeman, eds. American Voyager
The Journal of David Bates Douglass. Michigan Bot. 11: 121-122.
Voss, E. G. 1972. [Review of] N. Hotchkiss. Common Marsh Plants of the United States
and Canada. Econ. Bot. 26: 95-96.
Voss, E. G. 1972. [Review of] G. B. Ownbey. Common Wild Flowers of Minnesota.
Econ. Bot. 26: 198.
Voss, E. G. 1973. [Review of] P. Knauth. The North Woods/The American Wilderness.
Michigan Bot. 12: 128.
Voss, E. G. 1973. [Review of] A. H. Smith. The American Species of Psathyrella. Michi¬
gan Bot. 12: 137.
Voss, E. G. 1973. [Review of] T. Korling. Wild Plants in Flower: The Prairie — Swell and
Swale. Michigan Bot. 12: 128.
Voss, E. G. 1974 [“1973”]. [Review of] T. G. Tutin, V. H. Heywood, N. A. Burges, D. M.
Moore, D. H. Valentine, S. M. Walters, & D. A. Webb. Flora Europaea. Vol. 3. Diapen-
siaceae to Myoporaceae. Bull. Torrey Bot. Club 100: 241.
Voss, E. G. 1974. [Review of] D. Isely. Leguminosae of the United States: I. Subfamily
Mimosoideae. Michigan Bot. 13: 166.
Voss, E. G. 1974. [Review of] P. Dansereau. Inscape and Landscape. Michigan Bot. 13:
180.
Voss, E. G. 1974. [Review of] C. E. Herendorf, S. M. Hartley, & L. J. Charlesworth. Lake
Erie Bibliography in Environmental Sciences. Michigan Bot. 13: 180.
Voss, E. G. 1974. [Review of] C. Jeffrey. Biological Nomenclature. Taxon 23: 394-395.
Voss, E. G. 1975. [Review of] A. Love & D. Love. Cytotaxonomical Atlas of the Sloven¬
ian Flora. Michigan Bot. 14: 56.
Voss, E. G. 1975. [Review of] F. Swink. Plants of the Chicago Region, 2nd ed. Michigan
Bot. 14: 56.
Voss, E. G. 1975. [Review of] H. A. White & M Williams. The Alaska — Yukon Wild
Flower Guide. Michigan Bot. 14: 90.
Voss, E. G. 1975. [Review of] P. A. Herbert. Great Lakes Nature Guide. Michigan Bot.
14: 189.
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THE MICHIGAN BOTANIST
13
Voss, E. G. 1975. [Review of] T. Korling. Wild Plants in Flower III. Deciduous Forest.
Michigan Bot. 14: 202.
Voss, E. G. 1975. [Review of] F. Steams & N Kobriger. Environmental Status of the Lake
Michigan Region Vol. 10. Vegetation of the Lake Michigan Drainage Basin. Michigan
Bot. 14: 218.
Voss, E. G. 1975. [Review of] C. Jeffrey. Biological Nomenclature. Plant Sci. Bull. 21:
15.-16.
Voss, E. G. 1975. [Review of] J. C. Willis. A Dictionary of the Flowering Plants and
Ferns. Econ. Bot. 29: 1-2.
Voss, E. G. 1975. [Review of] D. S. Mitchell. Aquatic Vegetation and its Use and Control
a Contribution to the International Hydrological Decade. Quarterly Rev. Biol. 50: 475.
Voss, E. G. 1975. [Review of] E. C. Ogden. Anatomical Patterns of Some Aquatic Vascu¬
lar Plants of New York. Quarterly Rev. Biol. 50: 475^476.
Voss, E. G. 1975. [Review of] C. D. K. Cook ,B. J. Gut, E. M. Rix, J. Schneller, & M.
Seitz. Water Plants of the World. A Manual for the Identification of the Genera of
Freshwater Macrophytes. Econ. Bot. 29: 244+253.
Voss, E. G. 1976. [Review of] M. K. Wali, ed. Prairie: A multiple View. Michigan Bot.
15: 214.
Voss, E. G. 1976. [Review of] R. H. Premble. Native Grassland Ecosystems East of the
Rocky Mountains in North America: A Preliminary Bibliography. Michigan Bot. 15:
214.
Voss, E. G. 1976. [Review of] D. S. Correll & H. B. Cornell. Aquatic and Wetland Plants
of Southwestern United States, Vols. I and II. Quarterly Rev. Biol. 51: 134.
Voss, E. G. 1977. [Review of] T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore,
D. H. Valentine, S. M. Walters, & D. A. Webb. Flora Europaea. Vol. 4. Plantaginaceae
to Compositae (and Rubiaceae). Bull. Torrey Bot. Club 104: 73-75.
Voss, E. G. 1977. [Review of] N. C. Fassett. Spring Flora of Wisconsin, 4th ed., revised
and enlarged by Olive S. Thomson. Michigan Bot. 16: 90.
Voss, E. G. 1977. [Review of] D. Isely. Leguminosae of the United States: II. Subfamly
Caesalpinoidae. Michigan Bot. 16: 91.
Voss, E. G. 1977. [Review of] J. R. Wells & P. W. Thompson. Vegetation and Flora of the
Huron Mountains. Michigan Bot. 16: 91.
Voss, E. G. 1977. [Review of] R. H. Mohlenbrock. Sedges: Cyperus to Scleria. The Illus¬
trated Flora of Illinois. Michigan Bot. 16: 91-92.
Voss, E. G. 1977. [Review of] N. C. Fassett. Spring Flora of Wisconsin, 4th ed. Quarterly
Rev. Biol. 52: 90.
Voss, E. G. 1977. [Review of] W. L. Phillips & R. L. Stuckey. Index to Plant Distribution
Maps in North American Periodicals Through 1972. Quarterly Rev. Biol. 52: 204.
Voss, E. G. 1977. [Review of] T. Van Bruggen. The Vascular Plants of South Dakota.
Econ. Bot. 31: 7-8.
Voss, E. G. 1978. [Review of] J. B. Moyle. Northland Wild Flowers. Michigan Bot. 17:
24.
Voss, E. G. 1978. [Review of] N. F. Smith. Michigan Trees Worth Knowing, revised 5th
ed. Michigan Bot. 18: 24.
Voss, E. G. 1978. [Review of] J. K. Morton. The Flora of Manitoulin Island and the Ad¬
jacent Islands of Lake Huron, Georgian Bay, and the North Channel. Michigan Bot. 17:
48.
Voss, E. G. 1978. [Review of] N. Shelton. The Life of Isle Royale. Michigan Bot. 17: 48.
Voss, E. G. 1979. [Review of] R. L. Simonds & H. H. Tweedie. Wildflowers of Michi¬
gan. Michigan Bot. 18: 14.
14
THE MICHIGAN BOTANIST
Vol. 51
Voss, E. G. 1980. [Review of] J. T. Mickel. How to Know the Ferns and Fern Allies.
Michigan Bot. 19: 22.
Voss, E. G. 1980. [Review of] R. I. Ford, ed. The Nature of Ethnobotany. Michigan Bot.
19: 30.
Voss, E. G. 1980. [Review of] E. E. Weatherbee & J. G. Bruce. Edible Wild Plants of the
Great Lakes Region. Michigan Bot. 19: 30.
Voss, E. G. 1980. [Review of] F. Swink & G. Wilhelm. Plants of the Chicago Region, Re¬
vised and Expanded Edition. Michigan Bot. 19: 36.
Voss, E. G. 1980. [Review of] T. G. Tutin, V. H. Hey wood, N. A. Burges, D. M. Moore,
D. H. Valentine, S. M. Walters, & D. A. Webb. Flora Europaea. Volume 5. Alismataceae
to Orchidaceae (Monocotyledones). Bull. Torrey Bot. Club 107: 556-557.
Voss, E. G. 1981. [Review of] R. H. Mohlenbrock. Willows to Mustards. The Illustrated
Flora of Illinois. Michigan Bot. 20: 180.
Voss, E. G. 1981. [Review of] W. G. Dore & J. McNeill. Grasses of Ontario. Michigan
Bot. 20: 191.
Voss, E. G. 1981. [Review of] J. T. Kartesz & R. Kartesz. A Synoymized Checklist of the
Vascular Flora of the United States, Canada and Greenland. Syst. Bot. 6: 91-93.
Voss, E. G. 1982. [Review of] B. V. Barnes & W. H. Wagner, Jr. Michigan Trees, Revised
and Enlarged Edition. Michigan Bot. 21: 46.
Voss, E. G. 1982. [Review of] D. R. Given & J. H. Soper. The Arctic- Alpine Element of
the Vascular Flora at Lake Superior. Michigan Bot. 21: 65.
Voss, E. G. 1982. [Review of] J. B. Levenson. The Southem-mesic Forests of Southeast¬
ern Wisconsin: Species Composition and Community Structure. Michigan Bot. 21: 88.
Voss, E. G. 1982. [Review of] T. W. Hodler, R. Brewer, L. G. Brewer, & H. A. Raup. Pre¬
settlement Vegetation of Kalamazoo County, Michigan. Michigan Bot. 21: 88.
Voss, E. G. 1982. [Review of] R. H. Mohlenbrock. Flowering Plants: Magnolias to
Pitcher Plants. The Illustrated Flora of Illinois. Michigan Bot. 21: 116.
Voss, E. G. 1982. [Review of] J. A. Larsen. The Boreal Ecosystem. Michigan Bot. 21:
140.
Voss, E. G. 1982. [Review of] M. T. Stieber & A. L. Karg. Guide to the Botanical
Records and Papers in the Archives of the Hunt Institute. Syst. Bot. 7: 501.
Voss, E. G. 1983. [Review of] R. H. Mohlenbrock. Flowering Plants: Basswoods to
Spurges. The Illustrated Flora of Illinois. Michigan Bot. 22: 68.
Voss, E. G. 1983. [Review of] A. Cronquist. An Integrated System of Classification of
Flowering Plants. Econ. Bot. 37: 498.
Voss, E. G. 1984 [“1983”]. [Review of] G. Halliday & M. Beadle. Consolidated Index to
Flora Europaea. Bull. Torrey Bot. Club. 110: 544.
Voss, E. G. 1984. [Review of] R. L. Stuckey & K. J. Reese, eds. The Prairie Peninsula —
In the “Shadow of Transeau.” Michigan Bot. 23: 80.
Voss, E. G. 1984. [Review of] T. S. Cooperrider. Endangered Plants of Ohio. Michigan
Bot. 23: 88.
Voss, E. G. 1984. [Review of] R. Brewer. Proceedings of the Eighth North American
Prairie Conference. Michigan Bot. 23: 120.
Voss, E. G. 1985 [“1984”]. [Review of] J. K. Morton. The Flora of Manitoulin Island.
Ontario Field Biol. 38: 454-7.
Voss, E. G. 1986. [Review of] R. B. MacFarlane. Collecting and Preserving Plants for
Science and Pleasure. Michigan Academician 18: 468-469.
Voss, E. G. 1987. [Review of] R. H. Mohlenbrock. Flowering Plants: Smartweeds to
Hazelnuts. The Illustrated Flora of Illinois. Michigan Bot. 26: 186.
Voss, E. G. 1988. [Review of] M. L. Fernald. Gray’s Manual (Facsimile Reprint). Michi¬
gan Bot. 27:73.
2012
THE MICHIGAN BOTANIST
15
Voss, E. G. 1989. [Review of] D. A. Daubendiek & Edna S. Newman, eds. Michigan Na¬
ture Association in Retrospect. Michigan Bot. 28: 80.
Voss, E. G. 1989. [Review of] F. W. Case, Jr. Orchids of the Western Great Lakes Region.
Michigan Academician 21: 305-306.
Voss, E. G. 1990. [Review of] W. B. Crankshaw. Manual of the Seed Plants of Indiana.
Michigan Bot. 29: 35-36.
Voss, E. G. 1990. [Review of] R. I. Zimdahl. Weeds and Words. Michigan Bot. 29:
77-78.
Voss, E. G. 1990. [Review of] J. H. Soper, C. E. Garton, & D. R. Given. Flora of the
North Shore of Lake Superior (Vascular Plants of the Ontario Portion of the Lake Su¬
perior Drainage). Michigan Bot. 29: 78-79.
Voss, E. G. 1990. [Review of] R. H. Mohlenbrock. Flowering Plants: Nightshades to
Mistletoes. The Illustrated Flora of Illinois. Michigan Bot. 29: 140.
Voss, E. G. 1991. [Review of] J. S. Pringle. Botanical exploration of the Canadian water¬
shed of Lake Huron during the nineteenth century. Archives Nat. Hist. 18: 135.
Voss, E. G. 1991. [Review of] H. A. Gleason & A. Cronquist. Manual of Vascular Plants
of Northeastern United States and Adjacent Canada, 2nd ed. Michigan Bot. 30:
202-207.
Voss, E. G. 1992. [Review of] H. A. Gleason and Arthur Cronquist. Manual of Vascular
Plants of Northeastern United States and Adjacent Canada. Syst. Bot. 17: 508-510.
Voss, E. G. 1993. [Review of] J. C. Hickman. The Jepson Manual: Higher Plants of Cal¬
ifornia. Taxon 42: 736-738.
Voss, E. G. 1994. [Review of] R. W. Holzman, B. A. Daubenbiek, L. Rizor, & F. Sibley,
eds. Michigan Nature Association Nature Sanctuary Guidebook 7th Ed. Michigan Bot.
33: 94.
Voss, E. G. 1996. [Review of] T. S. Cooperrider. The Dicotyledonae of Ohio. Part 2.
Linaceae through Campanulacae. Michigan Bot. 35: 63.
Voss, E. G. 1996. [Review of] E. B. Pitcher. Of Woods and Other Things. Michigan Bot.
35: 63.
Voss, E. G. 1996. [Review of] R. A. Howard. An Almanac of Botanical Trivia. Rhodora.
98: 217.
MICHIGAN PLANTS IN PRINT
Between the inception of The Michigan Botanist , and the publication of Volume 3 of
Michigan Flora (Voss, 1996), Ed contributed this series to the Journal, referencing all
journal articles, books and other references of which he was aware that were about Michi¬
gan plants or had a bearing on the occurrence and distribution of plants in the State, in¬
cluding maps, biographies, and other useful materials. Before the easy digital availability
of literature and the ability to search large scholarly databases, this byproduct of Ed’s re¬
search on the Michigan Flora provided students and other researchers with an unparal¬
leled access to relevant literature on Michigan plants.
Voss, E. G. 1962. Michigan Plants in Print. Michigan Bot. 1: 43^46; 91-94.
Voss, E. G. 1963. Michigan Plants in Print. Michigan Bot. 2: 29-30; 55-61; 93-95.
Voss, E. G. 1964. Michigan Plants in Print. Michigan Bot. 3: 25-30; 102.
Voss, E. G. 1965. Michigan Plants in Print. Michigan Bot. 4: 33-38; 67-70; 100-102;
124-126.
Voss, E. G. 1966. Michigan Plants in Print. Michigan Bot. 5: 46; 91-94; 125-126.
Voss, E. G. 1967. Michigan Plants in Print. Michigan Bot. 6: 26-29; 68-70; 185-186.
Voss, E. G. 1968. Michigan Plants in Print. Michigan Bot. 7: 25-30; 94; 188-189.
16
THE MICHIGAN BOTANIST
Vol. 51
Voss, E. G. 1969. Michigan Plants in Print.
Voss, E. G. 1970. Michigan Plants in Print.
Voss, E. G. 1971. Michigan Plants in Print.
Voss, E. G. 1972. Michigan Plants in Print.
Voss, E. G. 1973. Michigan Plants in Print.
Voss, E. G. 1974. Michigan Plants in Print.
Voss, E. G. 1975. Michigan Plants in Print.
Voss, E. G. 1976. Michigan Plants in Print.
Voss, E. G. 1977. Michigan Plants in Print.
Voss, E. G. 1978. Michigan Plants in Print.
Voss, E. G. 1979. Michigan Plants in Print.
Voss, E. G. 1982. Michigan Plants in Print.
193-194; 204.
Voss, E. G. 1983. Michigan Plants in Print.
Voss, E. G. 1987. Michigan Plants in Print.
Voss, E. G. 1988. Michigan Plants in Print.
Voss, E. G. 1989. Michigan Plants in Print.
Voss, E. G. 1990. Michigan Plants in Print.
Voss, E. G. 1992. Michigan Plants in Print.
Voss, E. G. 1993. Michigan Plants in Print.
Voss, E. G. 1996. Michigan Plants in Print.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
Michigan Bot.
8; 44-46; 160-162.
9: 59-61; 205-206; 245-246.
10: 44^16.
11: 78.
12: 73-78; 139-144; 254.
13:44-46; 101-102; 186-189.
14: 61-62; 107-110.
15: 123-126.
16: 94-94; 160-163.
17: 163-166.
18: 30; 36; 42.
21: 57; 94-95; 128-130; 184;
22: 18; 25-26.
26: 174-175.
27: 26-30.
28: 39-40.
29: 107-111.
31: 67-68; 116-119.
32: 227-230.
35: 57-62.
NEW NAMES AND COMBINATIONS
The following list of new names and combinations which Ed Voss authored is
arranged alphabetically by genus. The dates given are as recorded in the International
Plant Name Index (IPNI).
Acer saccharum Marshall var. viride (Schmidt) E. G. Voss. Michigan Bot. 24: 122. 1985.
Agropyron dasystachyum (Hook.) Scribn. var. psammophilum (J. M. Gillett & H. Senn) E.
G. Voss. Rhodora 68: 437. 1966.
Agrostis hyemalis (Walter) Britton, Stems & Poggenb. f. setigera (Femald) E. G. Voss.
Rhodora 68: 438. 1966.
Bidens connata Willd. f. ambiversa (Fassett) E. G. Voss. Michigan Bot. 34: 139. [Dec
1996] “October, 1995”
Bidens connata Willd. f. anomala (Farw.) E. G. Voss. Michigan Bot. 34: 139. [Dec 1996]
“October, 1995”
Bromus pubescens Spreng. f. glabriflorus (Wiegand) E. G. Voss. Rhodora 68: 440. 1966.
Carex communis L.H. Bailey f. gynandra (Farw.) E. G. Voss. Rhodora 68: 451. 1966.
Carex rugosperma Mack. var. tonsa (Femald) E. G. Voss. Rhodora 68: 453. 1966.
Cirsium hillii (Canby) Femald f. albiflorum E. G. Voss. Michigan Bot. 34: 139. [Dec
1996] “October, 1995”
Cirsium pitcheri Torn & A. Gray f. magenteum E. G. Voss. Contr. Univ. Michigan Herb.
23: 350. 2001.
Liatris scariosa (L.) Willd. var. nieuwlandii (Lunell) E. G. Voss. Michigan Bot. 34: 139.
[Dec 1996] “October, 1995”
Nuphar variegata Durand f. lutescens (Farw.) E. G. Voss. Michigan Bot. 24: 120. 1985.
Orchis rotundifolia Pursh f. lineata (Mousley) E. G. Voss. Rhodora 68: 462. 1966.
Polygala polygama Walter f. ramulosa (Farw.) E. G. Voss. Michigan Bot. 24: 121. 1985.
Puccinellia fernaldii (Hitchc.) E. G. Voss. Rhodora 68: 445. 1966.
Salix exigua Nutt. f. wheeleri (Rowlee) E. G. Voss. Michigan Bot. 24: 1 17. 1985.
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Salix glaucophylloides Fernald var. brevifolia (C. F. Wheeler & E. F. Sm.) C. R. Ball ex
E. G. Voss. Ohio J. Sci. 50: 187. 1950.
Sparganium chlorocarpum Rydb. f. acaule (Macoun) E. G. Voss. Rhodora 68: 436. 1966.
EPONYMY OF ED VOSS
Asteraceae Solidago vossii J. S. Pringle & Laureto. Michigan Bot. 49(4): 108. 2010 [Oct.
2010]
Orchidaceae xBlepharopsis vossii (Case) Efimov. Novosti Sist. Vyssh. Rast. 40: 48. 2009
[2008 publ. 30 Mar. 2009]
Orchidaceae Platanthera xvossii Case. Michigan Bot. 22: 141. 1983
Orchidaceae xPlatanthopsis vossii (Case) P. M. Br. N. Amer. Native Orchid J. 8: 37. 2002
[30 Dec. 2002]
GRADUATE STUDENTS SUPERVISED BY ED VOSS
Ph.D. 1962. Alfred E. Schuyler, Curator Emeritus, Philadelphia Academy of Sciences
Ph.D. 1965. Ronald L. Stuckey, Professor Emeritus, Ohio State University
Ph.D. 1971. Richard W. Scott, Curator, Central Wyoming College Herbarium, Riverton.
(Co-chair with W. S. Benninghoff)
Ph.D. 1976. Thomas Duncan, Professor Emeritus, University of California, Berkeley (Co¬
chair with W. H. Wagner, Jr.)
M.Sc. 1983. Margaret Bliss, Private Sector
Ph.D. 1983. Kerry S. Walter, BG-BASE (UK) LTD. Royal Botanic Garden Edinburgh,
Scotland, UK (Co-chair with W. H. Wagner, Jr.)
Ph.D. 1983. Tom Rosatti, Scientific Editor, Jepson Herbarium, University of California,
Berkeley (Co-chair with W. H. Wagner, Jr.)
Ph.D. 1988. Brian Hazlett, Professor of Biology, Briar Cliff University
Ph.D. 1992. Rob Naczi, Arthur J. Cronquist Curator of North American Botany, The New
York Botanical Garden (Co-chair with A. A. Reznicek).
18
THE MICHIGAN BOTANIST
Vol. 51
“FRATERNALLY, ED” — A REMEMBRANCE OF ED VOSS
FROM UNDERGRADUATE DAYS
Tom S. Cooperrider
Department of Biological Sciences
Kent State University
Kent Ohio 44242
I met Ed Voss (Fig. 3) in 1947 (65 years ago!), when we were both freshmen
at Denison University in Granville, Ohio. Ed had enrolled in September 1946. I
was discharged from the Army in November and enrolled at Denison in January
1947. By the end of the spring semester, we were members of the same frater¬
nity. Over the years since then, he always closed his letters with the words, “Fra¬
ternally, Ed.”
Those who know us might not think of either Ed or me as a typical fraternity
man, and that would be correct. An understanding of how we came to be frater-
FIGURE 3. Ed Voss as an under¬
graduate at Denison University.
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THE MICHIGAN BOTANIST
19
nity brothers requires an understanding of the times and the environment. Acad¬
emically, Denison afforded its students a first-class education and a close work¬
ing relationship with the inspiring professors. Socially, college life at Denison in
the 1940s was in some ways like that depicted in early Hollywood movies, com¬
plete with everything from sweetheart serenades to pep rally bonfires. Among
other traditions, coats and ties were required for dinner, and following the meal
we remained at the table and sang college and fraternity songs. Of the male stu¬
dents, 95% joined a fraternity, and the other 5% were mostly students living at
home. For the men residing on campus, belonging to a social fraternity was al¬
most a necessity.
Fortunately for those of us inclined toward independence, there was the
American Commons Club (ACC). Despite the lack of Greek letters in the name,
ACC was a genuine fraternity. We had a residential house on campus with study
rooms, a sleeping porch, a dining room, and a large lounge for social events and
meetings of the brotherhood. Much of our college education came from life in
the ACC house, and it influenced us for the remainder of our years. For Ed Voss,
belonging to the American Commons Club was a special source of pride because
his father had been a member a generation earlier.
After graduating in 1950, 1 continued to do occasional ACC work for the next
few years, as did other members. Ed continued on for the next 60 years. His
work for the group ended in May 2009, with the final edition of “Commoner
News,” an alumni newsletter he edited. The American Commons Club at Deni¬
son had closed down some years earlier.
Ed and I both became botanists. We made our way to botany separately, by a
kind of convergent evolution. Our friendships started at Denison. So also did our
affection for the university and fraternity.
During our senior year, Ed served as secretary for the fraternity. The highlight
of Monday evening business meetings was Ed’s reading of the minutes of the
previous meeting, written and read with his customary logic and wit. He also
wrote and edited that year a weekly in-house newsletter for ACC members:
GRANVILLE GRAVEL, “We leave no stone unturned.”
20
THE MICHIGAN BOTANIST
Vol. 51
BEYOND THE BOTANY— LIFE WITH ED VOSS
John S. Russell
5260 Textile Rd.
Saline, MI 48176
I was honored to share a home with Ed Voss for three years, to assist him in
several courses at the Biological Station, and to be on his short list for camping
companions at the end of many summers. He was on my doctoral committee,
and was an advisor, advocate, and friend for many years.
Ed was the ultimate curator. His house was curated. His car was curated. His
medicine cabinet was curated, and his refrigerator was curated. At supper fol¬
lowing a night when I had left a half-empty bottle of wine in the refrigerator, I
was presented with the statement: “I trust that bottle of alcoholic beverage on the
milk products shelf is NOT for our mutual consumption?” Hard to answer that
correctly! His basement had a complete set of National Geographic from day
one, but no TV. Surprisingly, his yard held several alien species. I planted an
Abies concolor in the front yard by the drive only after a prolonged discussion
about what image that would convey.
I am sure that Ed was comfortable with me as a Teaching Assistant, not be¬
cause of my botanical ability, but because I could keep the boats running, tie
them on the trailer securely, and keep the cars running. I was happy with that
role, as my guesses at plant ID were usually wrong anyway. I learned to say
“Here’s an interesting find,” instead of “This looks like a gentian.” If it wasn’t a
gentian, even though it looked like one, I was in minor trouble. I increased my
value when I, somewhat guiltily, confessed to him that I had agreed to assist
Howard Crum in Bryophytes & Lichens. Ed snorted, and said “Good. Now I
won’t have to stoop to identify the little stinkers.”
The most relaxed and fun aspect of life with Ed was during the end-of-sum-
mer camping trips. Sometimes there would just be the two of us, and sometimes
three. We always sought out Lake Superior shorelines, for there we found the
greatest privacy and the best botany (Fig. 4). Each trip had a theme. We might
focus on exploring new areas, or on following a newly acquired map. One year
it was a search for a new record for Empetrum. Some themes recurred, such as
finding and consuming vast quantities of blueberries. Six species were available
on the shore in August, and here, mixing of species was permissible! Pies, pud¬
dings, or fresh — any way was fine. Ed’s appreciation of wild things edible ex¬
tended to the fungi. I honestly believe that he was saddened when fungi were de¬
graded and removed from the plant kingdom. That did not, however, affect their
edibility. Chanterelles were particularly relished. One summer I discovered a
Sulfur Shelf that fed us for several days — omelets, scrambled eggs, chili, and
mixed with vegetables. We never had the nerve to try boletes or amanitas, and
successfully lived on. We reminisced over Alex Smith’s contention that ALL
fungi were edible. Some, however, would kill you.
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THE MICHIGAN BOTANIST
21
FIGURE 4. Ed Voss with vasculum looking at Pictographs on Lake Supe¬
rior. Photo by John Russell.
A trip would begin with packing the car in a precisely calculated manner, so
the load would not shift, and so that the driver could just barely see out the back
window. Ed never got into the backpacking equipment thing, even though this
would have solved lots of problems. The nine by nine foot umbrella tent, te¬
diously coated with waterproofing, and its box (saved since purchase) of alu¬
minum poles, went in on the bottom. It was not pushed forward, as it had to
come out first so it could be erected and serve as a depot into which to unload
the rest of the gear. The two-burner Coleman stove in its original box, the Cole¬
man lantern, also in its box, the condiments, food, and utensils went in, and the
toiletries went on top, always at the ready. Once I returned the toilet tissue to the
condiment box. Ed’s subtle response was to hide it from me, as I had from him.
The cooler with frozen canned ice in ancient rusting tins went in on the left. A
custom-cut shelf of plywood was laid between the window ledges behind the
22
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 5. Ed Voss reading at 3 mile creek. Photo by John Russell.
rear seat to support small items, a plant press, small toolbox, and county maps.
Backpack gear would have solved space problems, but it never entered the pic¬
ture. One year, catastrophe nearly struck when Jeep changed models. Renault in¬
fluence caused the scrapping of the old large Cherokee design, with its lift gate
and tailgate, replacing it with a unit body design. The old light blue Cherokee
was badly rusted, tailgate nearly gone, and windows sticky. Ed got a new dark
blue Cherokee. It was calculated to be 3 1% smaller in load area, if you measured
all the way to the roof. I pointed out that the area above the shelf had to be left
free anyway, and the calculation was revised to under 20%. I designed and cut a
second piece of plywood to serve as a tailgate upon which to eat and press plants
(the new Cherokee had only a single liftgate). The old shelf was cut down to ac¬
commodate the narrower width. Now the only problem was to reduce the load by
about 15%. Packing tighter seemed the answer, not going to backpack gear! It
was done, and the trip was accomplished, with the only problem being the front
floor of the new Jeep being recessed so that it wasn’t possible to just sweep Lake
Superior sand out the door. Why would anyone design a four-wheel drive vehi¬
cle that way? Because they were French.
Eggs for breakfast — don’t cook the bottoms hard. Bring your own coffee or
tea, for Ed didn’t do stimulants. How did he ever get his degree? Oatmeal with
bilberries was better! We’d pack sandwiches for lunch and set out on a daily
hike, following maps from the DNR and always seeking blueberries and mush¬
rooms. A three-course dinner highlighted our evenings. Often we had brought
fresh corn from Clem Burr’s farm east of UMBS. We ate the perishables first, as
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THE MICHIGAN BOTANIST
23
FIGURE 6. Ed Voss having a lunch break amidst lichens. Photo by John Russell.
the canned ice could be expected to last 2.5 days if the cooler was placed appro¬
priately in the shade and the arc of the sun in the sky was properly calculated.
Evening was for bathing, and then reading (Fig. 5). Reading was continued in
the tent, warmed by the Coleman, until an hour or so after dark when it was time
to retire. This was also the time devoted to going over field notes, although
camping trips were probably the times that Ed did the least hardcore botany.
They were his “vacation” time, a time when perhaps he was most relaxed (Fig.
6) and under little pressure to perform his botanical duties. (I realize that a “re¬
laxed Ed” is somewhat oxymoronic.)
Often sadly, there was little room in the Jeep for souvenirs from the shore. I
wanted to bring a pail of beach sand back for my classes, but had to package it
in small containers that could be fitted into small interstices in the load. I ratio¬
nalized that we could bring back a volume equal to that of the food we had eaten.
A poor substitute for food, Ed opined. One thing I was able to smuggle back was
a very strangely shaped burl on a jack pine trunk, which, when skinned of its
bark, resembled the lower half of a human. When it was unloaded back in Mack¬
inaw, Ed professed surprise, but I think he knew it was contraband all along. We
did haul agates, the occasional pressed plant, and insects back to town. The
Mackinac Bridge seemed the gate back to reality.
I suspect that Ed enjoyed these weeks on the Superior shore more than any
other times of his busy life, and I take solace in the belief that he is spiritually
there today, in a place where essence is more important than the physical.
24
THE MICHIGAN BOTANIST
Vol. 51
ED VOSS— FIELD BOTANIST IN SUMMER,
CURATOR IN WINTER
A. A. Reznicek
University of Michigan Herbarium
3600 Varsity Drive
Ann Arbor, MI 48108-2228, USA
No, I’m not suggesting that Ed led a dual botanical life. He was devoted to the
vascular (especially the flowering) plants of Michigan at all seasons and
throughout his life. But his botanical seasons were definitely compartmentalized,
like so many other elements of his life.
In summers (from mid-May to the Labor Day weekend, slightly longer after
retirement) Ed was based “up north,” staying at the Biological Station on Dou¬
glas Lake while teaching (for 36 summers; 1963-1974, 1976-1998, 2003; plus
four years as a teaching assistant, 1949, 1951-1953) (Fig. 7). Ed attended the In¬
ternational Botanical Congress in Leningrad in 1975, John Thieret substituting
for him that summer. When not at the station, Ed stayed at his cottage on the
straits near the tip of the Lower Peninsula looking out over the Mackinac Bridge.
He likely went in the field more often than any other botanist in the region. I
always wished I could have as much field time as Ed. His field work, though,
was usually tightly focused and organized, like everything else in his life. When
he was teaching, he was scouting class trips and looking for new areas to take
people or show them special things, such as transient species of disturbed habi¬
tats, which were always a challenge. However, he was always keen to explore a
new bog or fen, and was especially interested in comprehensively documenting
the plants of favorite areas in the straits region, including Grass Bay, the Head¬
lands, or Saint Helena Island (Voss 2001).
He built a great body of knowledge about plants of the Biological Station
area, and his work, added to earlier collectors, made the Douglas Lake region
(Emmet and Cheboygan counties) one of the best known in the state. In fact, this
flora, including all the interesting Great Lakes endemics centered on the straits
region, was the subject of his doctoral thesis (Voss 1954) and most of his botan¬
ical publications prior to 1957. The Douglas Lake region was so well known that
it was a banner day when a new native species was added. So I was particularly
lucky on 9 August 1978, the very first time I went out with Ed in the region, to
spot right in the trail to Mud Lake Bog, Torreyochloa (Puccinellia) fernaldii — a
new, though inconspicuous, native grass to the Douglas Lake area (which ended
up as Voss No. 15,000). I felt then that I had passed muster as a field companion.
Ed also collected in southern Michigan in the fall, often in conjunction with
scouting for the well-known aquatic plants course which he taught for many
years in Ann Arbor. He was especially fond of the Lake Erie marshes, and was
interested in their dynamic changes with fluctuating lake levels. Whenever I was
with him, he always enjoyed seeing if this was a year that the very local Bolto-
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THE MICHIGAN BOTANIST
25
FIGURE 7. Ed Voss (second row, far
right) with the 1963 UMBS Aquatic Flow¬
ering Plants class. Photo UMBS archives.
nia asteroides or Sagittaria montevidensis would be in evidence. He was also in¬
terested in Great Lakes shoreline dynamics “up north,” with his Cecil Bay tran¬
sects, conducted over many years, and there is little doubt that the habitats of the
Great Lakes shorelines were a special interest to Ed his entire life.
Another important element for Ed was, of course, trying to make the maps in
the Michigan Flora (Voss 1972, 1985, 1996) represent the distributions of
species accurately. As time allowed in the summers, he ranged widely over the
state. He once noted that he had collected plants in every county in the State (and
on many of the islands). His collection numbers reached nearly to 17000, the
vast majority from Michigan, far exceeding the totals of any other Michigan col¬
lector. These collections will be one of Ed’s lasting legacies. Ed also made im¬
portant collections from the Ontario shore of Lake Superior, and occasionally
collected farther afield — usually in conjunction with International Botanical
Congress field trips. His greatest activity was in the early years of the Michigan
Flora project; in 1957, he collected 2088 numbers, on some days gathering
nearly 100 numbers from interesting sites. He made collections until 2008. Much
to some field companions dismay, Ed moved quickly through areas — but he
missed little. Ed’s collecting was very even handed. He liked sedges and never
avoided them, freely collected willows, and was particularly fond of aquatics,
though he did claim not to like ferns (despite collecting them assiduously). He
was fearless about getting wet feet (or more) and was very willing to wade in to
get a special wetland plant (Fig. 8); biting insects and other discomforts did not
seem to bother him when he was on a mission. When he was doing serious col¬
lecting, he always had his pick slung in a loop and clippers in their sheath on his
jeans — high botanical fashion — so was not afraid to tackle tough-rooted weeds
either.
His vehicles, for a long time Jeeps, then Ford Explorers, were selected with
the purpose of serving as a field vehicle. The ideal vehicle had a back tailgate
that lay down to produce a surface useful for pressing and other related duties.
When tailgates were converted to a lift up action it caused Ed consternation,
though he did realize it provided some rain shelter. He wanted a vehicle where
the floor did not have a raised lip at the doors, so that debris and sand could be
easily swept out, and he decried the loss of the front bench seat, which made for
26
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 8. Ed Voss sinking in Barclay
Lake in 1997. Photo by John Russell.
convenient placement of a box with collecting accoutrements. In fact, his next-
to-last vehicle was a special order to get that bench seat.
Ed was a careful collector, as one might expect from a student of Rogers Mc-
Vaugh, and his specimens were always of high quality; carefully prepared, com¬
plete, with good notes, and consistent, accurate labels. His scholarship was based
on specimens and he knew firsthand how difficult it was for a researcher to deal
with poor specimens. He carried a vasculum in the field where feasible, and
often carefully floated out aquatics to make the best collection possible. His
press was always handy in the vehicle (see Fig. 12). He recorded data as soon as
possible in the field and anybody who collected with him knew his pocket-sized
aluminum clipboard (see Fig. 20) with rubber bands disciplining the sheets
within. These roughly written, barely legible notes were neatly transferred into a
small three-ring binder (Ed freely admitted that his handwriting was not the
best), which served as his permanent field notes and were augmented with final
determinations, notations about herbaria to which duplicates were distributed,
and revisions by specialists. These will be deposited at MICH.
Besides collecting specimens, Ed always wanted to know and recognize
Michigan species as living plants in their habitats (Fig. 9). This was very impor¬
tant to him for many reasons. He described habitats in the Michigan Flora vol¬
umes from data compiled from herbarium specimens, but he always liked to note
specific information from his own experience, as information derived from spec¬
imens was often generalized and imprecise. He liked to include helpful field
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THE MICHIGAN BOTANIST
27
FIGURE 9. Ed Voss admiring Lithospermum caroliniense during the 2004 minicourse. Photo by
John Russell.
28
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 10. Ed Voss photographing at Crisp Point. Photo by John Russell.
recognition hints from his own experience. He also liked to test his keys as he
wrote the treatments. All this, of course, was essential for writing the original
keys that made his floras so accessible. He disdained floras that were uncritical
compilations from literature and specimens, and had choice words to say about
people who wrote about plants as if they knew them, but from their comments,
had obviously never seen them alive.
Ed deeply enjoyed being in the field, especially in bogs, fens, and on his
beloved Great Lake shores, dunes, and beaches (Fig. 10). It was far more than a
career, it was his, singular, life. A highlight every year was his annual week
camped on Lake Superior just before coming back to Ann Arbor, usually shared
with a botanically minded friend. This was the closest he ever came to taking a
vacation (see article by John Russell).
Ed’s life as a Curator, during the academic year was, of course, a natural ex¬
tension of his work as a field botanist collecting plants. But for Ed, curation was
also a critically important element of his research on the flora of Michigan. They
were inseparable, as specimens were his basic research tools, serving not only as
documentation for the occurrence of plants in Michigan, but also as the raw ma¬
terial for the development of his original keys, and his synthesis of habitats,
flowering times, etc., across the State, especially for plants with which he had lit¬
tle or no familiarity in the field. Ed was very concerned about specimen quality
even after collecting, and carefully watched the mounting of specimens, and the
herbarium staff heard about it if plants were mounted in such a way as to impede
study of diagnostic features.
Perhaps the most time consuming element of Ed’s curation was the enormous
2012
THE MICHIGAN BOTANIST
29
effort that he put into supplying accurate identifications for the 230,000 or so
specimens utilized for the Michigan Flora project. Ed was very strict about
records for the Michigan Flora; every dot on every map in his volumes was based
on specimens that he personally studied and determined. If he had not seen a
specimen, the record didn’t exist. Having to some degree assumed that mantle, I
know how much time it takes to do the painstaking determinations of plants that
Ed did for about 60 years, not only for specimens at the University of Michigan
Herbarium, but also numerous collectors and students all over the State and even
the Great Lakes region. It meant consistent late nights and weekends working at
the Herbarium. But his helping and encouraging others in this way was probably
as helpful to Great Lakes Botany over the years as his teaching and his flora.
Specimens labeled with obscure localities were a particular challenge to Ed,
and he worked assiduously to locate them at least to county. Very often, these
collections with obscure data were historical specimens of considerable potential
value for information about past distributions, and he would sometimes spend
many hours researching collectors habits, itineraries, and other details to produce
a more complete label or to resolve apparent contradictions. This was especially
important for Michigan because immediately upon achieving statehood in 1837,
the Michigan Legislature authorized a geological survey of Michigan’s natural
resources, headed by Douglass Houghton. One of the important duties of this
first survey was to collect plants to document the flora, and thus Michigan has an
unusually good series of collections before extensive settlement — but with many
hard to-place localities. This work led to a helpful catalog titled Gazetteer of
Some Possibly Puzzling Collecting Localities for Michigan Plants (Voss 2005;
and also at: http://www.lsa.umich.edu/herb/michvoss/index.html ) that summa¬
rized his notes about localities gained over a half century. Naturally, Ed’s own la¬
bels were models of consistency and precision. I recently learned that early con¬
tact with Lloyd Shinners in 1951 at the Biological Station may have provided Ed
with the format for his specimen labels. He would rail against vague and mean¬
ingless statements on labels, pointing out that locality descriptions like “south of
Ann Arbor” could mean Ohio, or worse (Voss 2002).
In the course of all this study, Ed became an expert on the handwriting, label
styles, and collecting and labeling habits of all of the major Michigan and Great
Lakes collectors, ranging from the highly productive and talented, but disorga¬
nized and eccentric, Oliver A. Farwell (whose initials Ed thought quite appropri¬
ate), to collectors he admired for their diligence and perspicacity in finding rare
plants and new records. He noted Charles K. Dodge’s obsession with having his
address on every label, sometimes even two or three times, regardless of where
the plant was collected, or certain collectors inability to correctly read maps.
With his focus on collections, and the tangible results — send in a collection and
a dot goes on a map — Ed stimulated many people to collect plants in their local
region to contribute to the Michigan Flora project. And, of course, contributors
always got back typewritten lists of corrections to their determinations.
With Ed’s attention to label accuracy and strong interest in historical collec¬
tions, it is no surprise that he also had a strong interest in Botanical history. A cu¬
rator working with historical specimens has to be part historian, part geographer,
part handwriting expert, and part detective. Ed was all of those in good measure.
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THE MICHIGAN BOTANIST
Vol. 51
His historical detective work culminated in his very readable 1978 book on
botanical exploration of the western Great Lakes region: Botanical Beach¬
combers and Explorers: Pioneers of the 19th Century in the Upper Great Lakes
(Voss 1978). This is a work notable for its accuracy, deep detail, and also for
having 343 sometimes lengthy footnotes crammed into its 100 pages! Ed de¬
lighted in having unearthed, from some specimen label or obscure reference, a
tidbit of information that shed new light on some collector and their method of
operation. All these he considered important items for a curator.
Ed recorded all label data, with exasperating literalness, for the collections he
examined for the Michigan Flora project for many years on record slips ( not
index cards, as Ed would quickly correct anyone who referred to them as such)
and, starting in 1991, into a computer database. Though never really warming to
computers and databases, Ed realized their value for his research immediately,
and the Michigan Flora database quickly grew to be the largest one at the
Herbarium, as Ed not only supervised entering all new collections he examined,
but also the conversion of his paper record slips into digital form. Ed developed
precise and detailed instructions — running to six single spaced pages — for enter¬
ing records into the database, and of course, he requested that a special proofing
routine be built so that he could carefully proof each individual record by com¬
paring the specimen label or the record slip to the database entry. Not being one
to change a well-developed routine, the printed output from the database had to
precisely match the format of the record slips — as Rich Rabeler and I discovered
when we created that database.
Ed decried carelessness, noting how commonly people misrepresented col¬
lection localities in the literature — mistaking the collectors address (often
printed on labels from the late 1800’s and early 1900’s), as the place the plant
was collected or coming to the wrong conclusion about which of the numerous
“Mud Lakes” a plant was from. Michigan specimens without County, or where
the locality was unclear (or obviously incorrect) were always annotated with the
correct data if Ed could discover them. It is a tribute to Ed that the distributions
of Michigan plants are so well presented in his flora, despite error-prone collec¬
tors like Farwell devoting their careers to the State. As of April, 2012, the task of
entering all the Michigan records into a database is now complete, finishing a
task Ed started at the inception of the Michigan Flora Project in 1956 (Fig. 11).
Ed’s approach writing his Michigan Flora work was very much an extension
of his curation, and was as methodical and deeply detailed. His command of
botanical literature was phenomenal, as reflected in his years of contributing
Michigan Plants in Print to The Michigan Botanist. He was a bibliophile, and
had an extensive (and, not surprisingly, well curated) personal reprint and book
collection centered on Michigan plants. He was very interested in the biology
and ecology of Michigan plants, and often had concise but perceptive comments
on those topics in the Michigan Flora. He also evaluated all literature that he
used for the flora with his deep and broad knowledge of plant nomenclature, de¬
rived from his years of service to the nomenclature committees of the Interna¬
tional Association of Plant Taxonomy, as well as with his knowledge of plants in
the field and the herbarium. Not surprisingly, the literature often was judged
sadly lacking. However, Ed did not himself delve into researching unresolved
2012
THE MICHIGAN BOTANIST
31
systematic questions in the flora, though he encouraged others to do so, and was
usually aware of the problems. Ed very much viewed his flora as a synthesis of
knowledge available to date. As a consequence, Ed made fewer nomenclatural
innovations for the flora than one might have expected, described no new
species, and only two new entities; rare color forms of two interesting Cirsium
species, the Great Lakes endemic Cirsium pitched, forma magenteum (Fig. 13),
and the prairie and jack pine barren Cirsium hillii forma albiflorum. This al¬
lowed others to name plants in honor of Ed, including the very rare hybrid Pla-
tanthera xvossii, and the Michigan endemic Solidago vossii.
FIGURE 11 (left). Ed Voss studying herbarium
specimens at the inception of the Michigan
Flora project. Photo: Asa Gray Bull. n.s. 3:
277-279. 1957.
FIGURE 12 (bottom left). Ed Voss pressing
plants. Photo by Bob Vande Kopple July 2007.
FIGURE 13 (bottom right). Holotype of Cirsium
pitcheri forma magenteum collected by Ed.
Photo by H. Huggins.
k
mmm
1 FI 5659
32
THE MICHIGAN BOTANIST
Vol. 51
THE BOTANICAL TEACHING LEGACY
OF EDWARD G. VOSS AT
THE UNIVERSITY OF MICHIGAN BIOLOGICAL STATION
Charles C. Davis
Harvard University Herbaria
22 Divinity Avenue
Cambridge, MA 02138
C. Eric Hellquist
Department of Biological Sciences
State University of New York Oswego
Oswego, NY 13126
Melanie Gunn
Point Reyes National Seashore
Point Reyes, CA 94956
The end of May marked Ed Voss’ annual trip to northern Michigan — his
botanical getaway since well before he became Professor of Biology and Cura¬
tor at the University of Michigan, Ann Arbor. This locale fortuitously provided
his inspiration for decades of pioneering research on the Michigan flora. His
grandfather had acquired their cabin on the Straits of Mackinac in 1930 to avoid
hay fever in their hometown of Dayton, Ohio. In September of that year, at the
age of one, Ed spent his first night along the Straits (Voss 2008). It was here in
northern Michigan that Ed established a lasting legacy from the many lives he
touched in the classroom, and field, at the University of Michigan Biological
Station (UMBS).
Ed became a faculty member at UMBS in 1963 and taught four courses during
his tenure: Systematic Botany (1967-1969, 1971-1974), Boreal Flora
(1967-1974, 1976-1994) (Fig. 14), Aquatic Flowering Plants (1963-1969; 1971;
1991), and Field Botany of Northern Michigan (1995-1998; 2003). From
1963-2003, he would miss only five summer sessions at UMBS. Ed treasured his
time at UMBS. As he once noted, “UMBS is a place you cannot stay away from!”
Ed taught his students with an enthusiasm that was legendary. It was peppered
with exclamations and tempered with precision to help students understand the
challenges, nuances, joys, and importance of botany. For example, on a class
field trip in 1995, Ed remarked with irritation about a motel named “The
Birches” that was surrounded by aspen. He curtly observed to the class that,
“There is a lot of botanical ignorance out there!” It was this dedication to the ap¬
preciation of the natural world, and of its accurate depiction, that were corner¬
stones of his teaching. The many students he inspired have become land man¬
agers, park rangers, policy makers, doctors, lawyers, professors, and of course,
well informed citizens of the botanical world (Fig. 14).
As we reflect on our experiences of Ed in the classroom and the field we are
struck by some of the traits that made Ed such a singular and inspiring botanical
educator.
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THE MICHIGAN BOTANIST
33
FIGURE 14. Ed Voss in 1974 with his Boreal Flora class. Photo UMBS archives.
Ed was undaunted by field conditions. Lightning, heat, rain, wind, and least
of all, mosquitoes, were no match for Ed. His sole focus was discovering and
teaching plants, with total disregard to environmental conditions that surrounded
him or his students. For example, when teaching the beautiful and diminutive or¬
chid Malaxis unifolia Michx. at Ryerse Lake in the Upper Peninsula (on years
when it could be found!), students would giggle as Ed was undeterred, and ut¬
terly enraptured, despite the mosquito sucking blood from his eyelid. Each new
discovery was a treasure for him, despite having seen these old friends many,
many times over the decades. His enthusiasm never dulled, and it was this in¬
fectious excitement that motivated us all (Fig. 15).
The only thing with the power to sway Ed from his botanical bliss was his
stomach. Despite his svelte physique, Ed was a voracious predator. Lunch or
dinner was never skipped. And he was notorious for taking heaping first and sec¬
ond helpings at the dinner line, and rarely made conversation while at his fa¬
vorite table. We suspect he felt that such discussions would get in the way of sat¬
isfying his enormous appetite.
Ed loved sharing the beauty of the Great Lakes region. Ed took great pride
in the beautiful places that he scheduled for the course. They included Grass Bay
(Cheboygan County) where students could traverse the upland mesic forest
down to the interdunal swales along the shore of Lake Huron; the mouth of the
Two Hearted River where students could stand atop a massive sand dune and
look to one side and see the vast open water of Lake Superior and the tannin rich
color of the river on the other; the old growth beech-maple-hemlock forests of
the Headlands near Mackinaw City; and of course the unforgettable bogs and
34
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 15. Ed Voss gleefully holding up Poison hemlock ( Conium maculatum) while teaching.
Photo by C. Mom.
fens of the Upper Peninsula (Fig. 16). Each of these habitats held a treasure trove
of plants that were unique to that site, and it was the combination of these plants
that elucidated the ecological context of these unforgettable landscapes.
When it came to spectacular locations, Boreal Flora students were fortunate
to experience a truly remarkable botanical excursion: the Canadian shore of
Lake Superior. For many “BoFlo” students, the Canada trip was the highlight of
the summer. Driving a procession of tightly packed vans, the class ventured
across the border synchronized to Ed’s precise itinerary. Although the logistics
of the trip could cause Ed anxiety, there was no doubt that this was the highlight
of his year. Ed’s love of northern floras, plant geography, geology, and history
were epitomized along to the panoramic backdrop of Lake Superior.
In addition to these many beautiful places, Ed also enjoyed waste ground
sites, or ‘stump dumps’ as he affectionately called them. These sites were not
easy to appreciate in part because, as Ed noted, waste grounds could be “as hot
as a pancake griddle”, but there was great wisdom in these choices. To Ed, these
sites served two purposes. First, they provided abundant material for teaching a
great variety of floral and vegetative forms, which was ideal for introductory stu¬
dents of plant diversity. Second, they helped to recreate the kind of destruction
and openness that resembled the post-glacial landscapes, so prominent in the
origination of Michigan’s flora. He was apt to remind students that the -2,800
species of Michigan’s land plants had arrived there only within the last -12,000
2012
THE MICHIGAN BOTANIST
35
FIGURE 16. Ed Voss teaching on the bog mat at
Ryerse Lake in the Upper Peninsula. Photo by C.
Davis.
years. It was in this context that he delivered lectures about how invasives, both
natives and non-natives, often found their way onto new landscapes. It was an
incredibly effective, and brilliant teaching tool. Not to mention that many of his
favorite plants-natives especially-could only be found in these environs (Fig.
17). The choice species of the waste ground of the Mackinaw City vicinity in¬
cluded Strawberry Blite ( Chenopodium capitatum (L.) Ambrosi), Leucophysalis
grandiflora (Hooker) Rydb., Pink Corydalis ( Corydalis sempervirens (L.) Pers.,
now known as Capnoides sempervirens (L.) Borkh. ), and Golden Corydalis
0 Corydalis aurea Willd.). He would spend hours scouting for these species long
after he retired from teaching. Each summer Ed would revel in rediscovering
these unpredictable species in his favorite vacant lots and gravel pits.
Ed was meticulously organized. Taxonomists like Ed are often guilty of
being overly tidy, and Ed was no different. For every class he was prepared with
a well-worn manila envelope containing relevant maps and other documents,
hand lens on sturdy upholster’s cord (Fig. 18), rock pick in a belt loop of his
trousers, and appropriate footwear (e.g., soccer cleats for wetlands) (Fig. 19).
Many will remember his carefully curated plant, insect, and book collections,
but a hallmark of Ed’s organization was exemplified by his field notes placed
neatly inside a small aluminum sided jacket. These notes were always typed on
the clean side of scrap paper that were cut to fit his notebook precisely (Fig. 20).
These cards served several purposes including expected, but yet undiscov¬
ered, county records for Michigan. Whenever there was an obvious gap in these
records, Ed was quite pleased to find the undocumented species in question no
matter how common or nondescript. These notes helped to produce glorious
maps of species’ range distributions of Michigan, which were the basis of his
classroom discussions of plant biogeography and the maps in the Michigan
Flora volumes (Voss 1972, 1985, 1996).
Ed was quite fond of the numerous distantly related species whose distribu¬
tions were found primarily in western North America, but were (strangely) also
36
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 17. Ed Voss at a waste ground site on the Darrow property near Mackinaw City teaching
Golden Corydalis, Corydcilis aurea. Photo by C. E. Hellquist.
H. s
... wM
FIGURE 18. Ed Voss in 1984
examining a specimen with
his hand lens. Photo: UMBS
archives
2012
THE MICHIGAN BOTANIST
37
FIGURE 19. EGV teaching
one of his last lectures in 2003
near Pictured Rocks National
Lakeshore. Photo by C. Davis.
FIGURE 20. EGV with his aluminum sided
notepad on the Bruce Peninsula, Ontario, Canada.
Photo by C. Davis.
38
THE MICHIGAN BOTANIST
Vol. 51
represented far from their center of diversity to include the Great Lakes region.
It was a fascinating topic on which he produced an excellent summary (Marquis
and Voss 1981), co-authored with one of his former students, Robert Marquis,
now a Professor of Ecology at the University of Missouri, St. Louis.
One of the most curious western disjuncts is Pterospora andromedea Nutt.,
an unusual mycoheterotroph found occasionally along the shores of the Great
Lakes. It was on a scouting trip for this plant near Pte. aux Chenes in the Upper
Peninsula that another important use of Ed’s note cards became apparent — di¬
rection finding. Ed typed the directions (to the tenth of a mile) to find
Pterospora. These directions included the obvious, but also the number of tele¬
phone poles to pass along the way, as well as the local botanical landmarks to
find the spot. It was truly remarkable, and led perfectly to the site that had not
been visited for at least a decade. Today, botany students visit many of the same
sites (and plants) that Ed vigilantly recorded in his notes season after season.
Ed was the “Program Director” of “Plant TV.” Although Ed was not the
originator of “Plant TV”, which traced back to a student of Professor Elzada
Clover in 1966 (Voss 2008), he became synonymous with its development and
use as a study aid. While in the field with students, Ed or his teaching assistant
(TA) would carefully collect a representative of each plant for display in glass
cases behind LaRue Library and later Gates Lecture Hall. Occasionally, he
would circulate the plant for closer inspection before carefully resting it in the
vasculum (“Into captivity!”, as he would gleefully exclaim). On one occasion,
while passing around the delicate Enchanter’s nightshade ( Circaea alpina L.), he
warned the students, “Don’t squeeze the pipes!” Suffice to say, Ed was very pro-
FIGURE 21. Plant specimens being prepared for “Plant TV”. Note Poison-ivy in beer bottle.
UMBS archives.
2012
THE MICHIGAN BOTANIST
39
FIGURE 22. Students study Plant TV at night.
Photo by M. Petru.
tective of these displays. At the end of each field trip, only Ed and the TA were
allowed to unpack the plants from the vasculum, place them carefully into water
bottles, and transfer them to plant TV with their proper typewritten name card
(Fig. 21). Once in plant TV, no student was allowed to touch any of the plants,
or even open the door to plant TV. Ever! Prior to quizzes and exams, students
would set up camp at plant TV, often staring into the glow of the fluorescent
lights late into the evening, creating mnemonics to remember each family and
scientific name (Fig. 22). Who among his students could ever forget that Poison-
ivy, Toxicodendron radicans (L.) Kuntze (now T. rydbergii (Rydb.) Greene), was
the only plant in plant TV stored in a brown beer bottle (Fig. 21), and always ap¬
peared as unlucky #13 on the first quiz? Fucky #13 still appears on quizzes every
year as a tribute to Ed.
Ed was a botanical monk. Ed did not suffer inexactitudes lightly. We re¬
member an unknowing student commenting on the value of common names, to
which Ed replied without hesitation, “Common names are for common people!”
We confess to having used this expression many, many times in our own teach¬
ing (always crediting Ed, of course). It would be easy to interpret this as mali¬
cious, but that was never Ed’s intent. And despite what many people thought, he
was more aware of his manner than we expected. When teaching Rubus parvi-
florus Nutt., for example, he offered about its fruits — “People say they are too
tart, but I have a tart tongue anyway.” We came to appreciate that Ed had a some¬
what monastic vision of learning and teaching plants. There was one way, and he
saw the path clearly. Any deflection from that path-be it rain, “dreadful” deer-
40
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 23: Ed Voss in a scene repeated virtually every summer, leading students from the 1994 Bo¬
real Flora class along the crest of the dunes near the “ghost forest,” Grand Sable Dunes, Pictured
Rocks National Lakeshore. Photo C. E. Hellquist
flies, delays while filing into the vans-was just that. His job was to get his stu¬
dents back on the path as quickly as possible to get on with the task at hand.
When in the field, it was time to learn, study, and appreciate the plants and
ecosystems of Michigan.
A final lasting lesson from Ed. Ed always finished the semester with one of
the most meticulously worded, detailed exams any student had experienced. The
exam always began with the instructions to follow the directions carefully (with
an T for emphasis). The second part of the exam was a “spot quiz” of 100 rep¬
resentative plants from the course. The installation of this quiz was so large it
could not be contained within Gates Lab, and spilled onto the dining hall tables.
The final exam marked a transformation of sorts for most students — a transfor¬
mation that is only characterized by the best biodiversity courses.
Ed’s passion for botany and Michigan resonated with his students. Few teach¬
ers, for example, receive hand-stitched embroidered field shirts from their
classes year after year (Figs. 15, 24). Of these cherished personal gifts Ed wrote
(2008), “. . . one of my most-appreciated mementos of my teaching days is a se¬
ries of shirts, personally embroidered for me with a depiction of some plant of
particular interest to the class.” The tradition began in 1977 with dwarf lake iris
(Iris lacustris Nutt.) and was followed by 13 additional shirts stitched with no¬
table species (Voss 2008) including Sand Dune Thistle ( Cirsium pitcheri (Torr.)
Torr. & A. Gray), Thimbleberry ( Rubus parviflorus Nutt.), Lake Huron Tansy
(Tanacetum bipinnatum (L.) Sch. Bip. subsp. huronense (Nutt.) Breitung),
2012
THE MICHIGAN BOTANIST
41
FIGURE 24. The back of one of Ed’s many embroidered gift shirts. Photo by C. Mom.
American White Waterlily ( Nymphaea odorata Aiton), and Chenopodium capi-
tatum (L.) Ambrosi.
Ed kept a record of every student he taught at UMBS (nearly 400), many with
whom he maintained correspondence throughout the years (Voss 2008). Under
his direction and mentorship, Ed helped these students make sense of the green¬
ery around them, providing the keys to unlock the natural world (see Fig. 7). For
many, including us, classes with Ed opened an entirely new dimension of won¬
der and discovery (Fig. 23). His instruction offered one of the most important
lessons that a beginning student of evolution and ecology can learn, which is
paraphrased so succinctly in the Chinese proverb: “Wisdom begins by calling
things by their right names.” Ed would often reflect this proverb in variations of
“You can’t understand the play unless you know the characters.” Finally, Ed’s
students always came to understand that sound conservation began with sound
taxonomy of the species of his beloved Great Lakes region. As Ed taught us, this
is also the gateway to becoming an informed steward of our planet and the ap¬
preciation that we are but one small leaf on this grand Tree of Life (Fig. 24).
ACKNOWLEDGEMENTS
We thank Irene Crum, Cindy Mom, Mary Scholtens, Karie Slavik, and Bob Vande Kopple for
their assistance locating information and pictures related to Ed’s teaching at UMBS. Thanks also to
Tony Reznicek and Rich Rabeler for their assistance and suggestions during preparation of the man¬
uscript.
42
THE MICHIGAN BOTANIST
Vol. 51
ED VOSS AND THE CONSERVATION OF
MICHIGAN’S RARE FLORA
Michael R. Penskar
Michigan Natural Features Inventory
Michigan State University Extension
Stevens T. Mason Building, PO Box 30444
Lansing, MI 48909-7944
Of Ed Voss’s many contributions and accomplishments as a botanist, professor,
and much lauded teacher, as amply recounted and celebrated in this volume, his
role as an indefatigable leader in plant conservation is one that should not be for¬
gotten. His dedicated work in protecting and conserving the rare flora of Michigan
and the Great Lakes region is an important legacy that will endure, his selfless de¬
votion in this regard serving as a model to all (see Fig. 25). Ed Voss contributed to,
and fostered, rare plant conservation throughout his career in numerous ways, as
will be described below. Moreover, he initiated his efforts well before the exis¬
tence, or significant expansion, of many national, state, and local conservation or¬
ganizations now known to lead in such pursuits. Voss’s work commenced prior to
the advent of Earth Day in 1970 and the subsequent use and proliferation of such
well-known terms or concepts as “biodiversity”, “biophilia” (Wilson 1984), or —
more topically — “sustainability.” His involvement in conservation also began long
before formal studies in the emerging fields of conservation biology and ecosys¬
tem management helped to articulate the scientific rationale for conserving rare
biota. It is therefore fitting and highly appropriate to honor Ed Voss’s important
work in rare plant and natural areas conservation by providing a brief overview
and description of his particular contributions in these areas.
As to how he may have regarded his purposeful work in this arena, it is easy to
surmise, to those who knew him well, that he considered his involvement in con¬
servation efforts to be much less an obligation than a natural and logical extension
of his scientific endeavors. Thus, this work was pursued with the same vigor and
thoroughness as applied to the Michigan Flora. One of Voss’s most important roles
in rare plant conservation was his long-time service as a member of the Technical
Advisory Committee for Plants, as appointed by the Michigan Department of Nat¬
ural Resources (MDNR) Threatened and Endangered Species Program. Initiated in
1974 following passage of the Michigan Endangered Species Act (Public Act 203
of 19741), the Technical Advisory Committee engaged Voss and his colleagues
with the important task of drawing up Michigan’s first Technical List of state en¬
dangered, threatened, and extirpated vascular plant species (encompassing pteri-
dophytes, gymnosperms, and flowering plant taxa). Preparation of a state rare
plant list was a ground-breaking task, yet the newly passed Michigan act provided
‘Now re-codified as Part 365, Endangered Species Protection, of the Natural Resources and En¬
vironmental Protection Act, Act 451 of the Public Acts of 1994 (Michigan Compiled Laws 1994).
2012
THE MICHIGAN BOTANIST
43
little specific guidance about how to do so. The act did note that in addition to the
Department’s own investigations and status determinations, it was given the
charge to seek “other available scientific and commercial data, which may include
consultation with scientists and others who may have specialized knowledge,
learning, or experience” to help promulgate a rule for the listing of such species. In
the absence of specific procedures and protocols, considerable credit is to be given
to Dr. Sylvia M. Taylor, Michigan’s first State Endangered Species Coordinator
(lodged within the MDNR Wildlife Division), for realizing the need to establish
both a procedural precedent as well as a methodology that would result in devel¬
oping a credible, objective, and scientifically defensible Technical List capable of
passing the periodic scrutiny of the Michigan Legislature. Enlisting the assistance
of Dr. Warren H. Wagner, Jr. to serve as its first Chair, the initial Technical Advi¬
sory Committee for Plants was formed, of which Ed Voss was a critically impor¬
tant member. Voss monitored the University of Michigan herbarium (MICH)
records for the committee, and also played the primary role in evaluating the sta¬
tus of aquatic flowering plants, in addition to providing guidance via his prodi¬
gious knowledge of the Michigan flora. His painstaking work to document the dis¬
tribution of rare species and other potential candidate taxa was only a portion of his
efforts, as he also carefully monitored the procedures and decisions of the com¬
mittee as well (S. Taylor, pers. comm.). I can vouch for this after having had the
distinct honor of serving with him for the last three of the six list reviews con¬
ducted by this committee. Voss’s voice of reason, his airtight logic, and his unim¬
peachable recommendations contributed enormously to a review system that
stands with high credibility to this day. It is also a testament to the breadth and
depth of his floristic knowledge that Ed Voss was also sought out to serve on the
state Rare Plant Advisory Committee for Ohio Department of Natural Resources,
on which he participated from 1992 until 2012.
In addition to his work on rare plant committees, Ed Voss contributed to plant
conservation through several other miscellaneous positions and activities. Notable
contributions included the Citizens Advisory Committee for the Cyrus H. Mc¬
Cormick Experimental Forest, U.S. Forest Service (1969), the Science Advisory
Committee of the Michigan Chapter of The Nature Conservancy (1992-2002), and
the Board of Technical Advisors, Tip of the Mitt Watershed Council (1993-2012).
Voss was an esteemed and long-time trustee for the Little Traverse Conservancy
(1983-2012), also serving on their Stewardship Committee and providing assis¬
tance by conducting a number of special botanical surveys (T. Bailey, pers.
comm.). Voss was also a highly dedicated and respected trustee for the Michigan
Chapter of The Nature Conservancy for two six-year terms of service (1986-1992
and 1993-1999). In the last known version of his curriculum vitae (2002), Ed
noted that he was a member of “25 or more local, regional, or national conserva¬
tion and natural history organizations, including The Nature Conservancy, Michi¬
gan Nature Association, Sierra Club, Wilderness Society, National Audubon Soci¬
ety, Environmental Defense Fund, Little Traverse Conservancy, Tip of the Mitt
Watershed Council, Michigan Natural Areas Council, Federation of Ontario Natu¬
ralists, Upper Peninsula Environmental coalition, etc. etc.”. These many activities
demonstrate his lifelong commitment to plant and natural area conservation, con¬
ducted during a vigorous professional career and a well occupied “retirement” that
44
THE MICHIGAN BOTANIST
Vol. 51
including ongoing Michigan Flora work and brief teaching stints at the University
of Michigan Biological Station (UMBS) in Pellston. Amidst these endeavors, how¬
ever, Voss somehow managed to find time to contribute yet more through various
kinds of outreach and education, such as delivering lectures to a wide diversity of
groups and organizations, leading numerous field trips (such as to his beloved
Grass Bay for The Nature Conservancy), and answering unending (and countless)
questions about plants (especially with regard to identifications), the inquiries ar¬
riving in all the standard ways, via phone, U.S. mail, personal visits, and the sub¬
sequent onslaught of e-mail (though he reserved a particular aversion to e-mail at¬
tachments).
Ed Voss understood the need to identify and protect natural areas as an essential
underpinning for the conservation of plants and all other organisms, as under¬
scored by the large number of organizations he served and supported . It is notable
that more than three decades ago he took a keen interest, while a member of the
editorial board of The Michigan Botanist , in the publication of an extended special
issue devoted entirely to an overview of all known, dedicated nature preserves in
Michigan (Crispin 1980). Susan Crispin, a botany graduate student advised by Dr.
John Beaman of Michigan State University (Beal-Darlington Herbarium, MSC),
prepared a master’s thesis devoted to such an overview, resulting in an unprece¬
dented compilation and description of the natural areas of the state. Although he
knew such a compilation would be highly dynamic through time, Voss felt it was
essential to present this work, both as a touchstone for accomplishment as well as
a measure of the considerable efforts that remain for us all. It was, in its day, a kind
of gap analysis and report card. Crispin (1980) acknowledges Voss’s “enormous
assistance in editing and arranging publication,” and once remarked to me that she
was deeply impressed by the level of his enthusiasm for this paper and his passion
and sense of urgency for bringing it to the readership of the journal and the con¬
servation community. As eloquently summarized by Tom Bailey, the Director of
the Little Traverse Conservancy, Ed Voss “was an amazing botanist, a dedicated
conservationist, and a wonderful friend. When he disagreed about something, he
was the embodiment of the Loyal Opposition, and always worked hard in service
to the conservation cause.” With the passing of Ed Voss, plant conservation has
lost one of its staunchest champions, one who left the richest and most admirable
of legacies and quietly taught us by his example and deed.
ACKNOWLEDGEMENTS
I am indebted to Sylvia Taylor for providing critical background information about the early
work and organization of the state Technical Advisory Committee for Plants, and to Tom Bailey and
Doug Pearsall for kindly verifying terms of service. I would also like to thank Tony Reznicek and
Rich Rabeler for guidance and support.
LITERATURE CITED
Crispin, S. R. 1980. Nature preserves in Michigan, 1920-1979. Michigan Bot. 19: 99-242.
Michigan Compiled Laws. 1994. Public Act 451 of 1994. Natural Resources and Environmental Pro¬
tection Act, Part 365, Endangered Species Protection, Sections 324.36501-324.36507.
Wilson, E. O. 1984. Biophilia, the human bond with other species. Harvard University Press, Boston.
2012
THE MICHIGAN BOTANIST
45
THE ROAD TO THE ISLES: REMINISCENCES OF
ED VOSS AS DISSERTATION CHAIRMAN
Brian T. Hazlett
Biology Department
Briar Cliff University
Sioux City, IA, 51104
At this time twenty-five years ago, I was writing my dissertation. The energy,
youthful enthusiasm, and purpose with which I entered The University of Michi¬
gan had not abated. Ann Arbor’s cultural stimulation and academic environment
remained unrivaled. I had served as a teaching assistant under notable faculty.
Nonetheless, my utmost reason for pursuing a doctorate at U-M was to study
under Edward G. Voss. No doubt, he stands highest among those who shaped my
botanical career. As a teacher (Fig. 14), he grounded me in nomenclature, aquat¬
ics, and the flora of northern Michigan. As mentor and dissertation chair, he di¬
rected my research, shaped my writing, penned letters of recommendation, and
served as an early reference on my resume. As a friend, he encouraged me dur¬
ing my early years as a college professor.
My passion for field botany and desire to teach at the college level developed
while still an undergraduate in western New York State. Midway through that
experience, a biology professor recommended (to those considering graduate
school) picking a research organism which one would still love after five years
of study. I tell my students that finding a dissertation committee chair with
whom one can develop and maintain a cordial relationship is considerably more
important.
My rapport with Ed developed during a summer at the University of Michi¬
gan Biological Station (UMBS) while still pursuing my Master’s at Michigan
State University. His invitation to take his Boreal Flora (Fig. 25) course came
when he visited East Lansing to speak about upper Great Lakes islands at a
Michigan Botanical Club meeting.
My research on the Nordhouse Dunes, a then proposed wilderness area, had
intensified an interest in regional flora. Moreover, I had concluded that contin¬
ued graduate study centered on floristics (within Michigan, at least) would most
likely only happen under Ed’s direction. My motivation to spend the summer at
UMBS centered on learning more about him and demonstrating that I was wor¬
thy of graduate study under his direction. Ed probably saw it as a prime oppor¬
tunity to evaluate me.
The respective insights into each other’s personalities gleaned during that
summer provided the foundation for our eventual collaboration on my disserta¬
tion research. In addition to taking Boreal Flora, I conducted an independent
study of the flora of Round Island, another proposed wilderness area, in the
Straits of Mackinac.
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THE MICHIGAN BOTANIST
Vol. 51
FIGURE 25. Boreal Flora, UMBS 1980. Photo by Gary Williams
Before heading to the Station, I had been forewarned that Ed was often
brusque. My informant, however, recommended that I not let that rough exterior
prevent me from knowing him better. Taking a cue from the 1973 film, Paper
Chase, I read much of what Ed had authored soon after arriving at the Station.
Among Ed’s passions were environmental stewardship, biogeography, philology,
and nomenclature. When he interspersed those subjects throughout his lectures
and field trips, I noticed. To some taking Boreal Flora, Ed spoke in inscrutable
parables. If so, I had discovered the key to their interpretation. One merely had
to listen and anticipate what he would say next.
Ed subtly found ways to connect to his students. When discussing Clintonia,
he would typically ask a student from New York State (were one present) to re¬
mind the rest of the class who DeWitt Clinton was. Such a seemingly random
question about an Empire State governor makes greater sense in light of the al¬
ternate name of the Erie Canal; Clinton’s Ditch. While on a joint collecting trip
to Round Island, soon after hearing that I had a year of Greek as an undergradu¬
ate, he immediately proceeded to quote (in Greek!) the beginning of the Gospel
of John. Fortunately, Professor Stockin had covered that material, thus by the
time Ed had reached “npog toy Oeov I recognized the quotation.
Ultimately, my habit of listening closely to what Ed said (and sometimes en¬
during a pun or a rant on the literal interpretation of a phrase or road sign) would
pay off when he became my dissertation chair. By that summer’s end, mean¬
while, I was fully established in the flora of northern Michigan, increasingly
2012
THE MICHIGAN BOTANIST
47
confident in conducting botanical inventories, and committed to pursuing re¬
search on Lake Michigan islands.
Ed and I had concluded, were I eventually accepted at U-M, he would over¬
see an investigation of the floras of the Manitou and Fox islands. He considered
islands, with their readily defined boundaries, to be worthy of special considera¬
tion. Thus, within Michigan Flora, seven islands (or island groups) in the Great
Lakes are mapped separately from their mainland counties. Leelanau County has
two distinct groups: the Manitou Islands and the Fox Islands.
Although my entry into a PhD program in Ann Arbor would begin three years
after that conversation, in the interim UMBS sought a primary investigator for a
National Park Service-funded botanical survey of North and South Manitou.
Fortuitously, Ed recommended me for the job. Thus, when I entered my doctoral
program, I already had a season of field work and a working relationship with
Ed.
Ed believed that the happiest graduate students were often the most produc¬
tive. I thoroughly enjoyed my tenure in the U-M herbarium. I eagerly used page
proofs of Michigan Flora while keying unknowns from my islands as well as
from the mainland portion of Sleeping Bear Dunes National Lakeshore. Under
Ed’s direction I became more disciplined; more task-driven rather than time-dri¬
ven.
Ed Voss personified persistence and precision. Words mattered. Just the right
word or phrase mattered more. Creating clear prose was his personal obsession.
Subsequently, my writing became more accurate. For instance, further referred
to time; farther denoted distance. Hyphens were handy. Placing a hyphen be¬
tween two dates automatically signified a range; one need not be encumbered
using to and from. One could refer to wind- and water-dispersed species’, wind
and water-dispersed species might imply two-part dispersal. Similarly, whereas
Fox and Manitou islands would signify two separate groups, Fox and Manitou
Islands would refer to only one island group. Accuracy in plant identification
was essential, yet my time-driven tendencies would inevitably surface. Ed,
nonetheless, cared enough to convey lists of my misidentifications to keep my
records current and to avoid subsequent mistakes. He could be gentle or sharp
(often using ! or ?) depending on the egregiousness of the error. Among his terse
comments:
3167 is Linaria canadensis, not Lobelia kalmii (which differs in having an in¬
ferior ovary, no spur on the corolla, and a big split down the middle of the
upper lip).
3766 Agrimonia striata [not gryposepala — a nice record! It keys perfectly to
striata in Mich. Flora; what went wrong?]
47 1 8 Bidens cernuus [not c onnatus] . . . This is the only misidentified Bidens
to have come through in at least 5 years. What key did you use and what eyes
examined the specimen?
As far as I could tell, these and similar blunders did not interfere with our
friendship. Ed was firm, yet forgiving.
48
THE MICHIGAN BOTANIST
Vol. 51
Ed, unlike some committee chairs, did not insist dissertations be perfect. Ide¬
ally, opportunities to improve one’s work would arise later during one’s career.
On the other hand, “one’s work” assuredly encompasses more than a disserta¬
tion. Another recollection from Professor Stockin’s Greek class applies here, “A
student is not above his teacher, but everyone who is fully trained will be like his
teacher” (Luke 6:40 NIV). In many respects, I am still Ed’s student and I reflect
his influence more than I realize. Among the habits that I do acknowledge: an in¬
sistence on clear writing, the necessity of prepping a field trip, and keeping ac¬
curate collecting records.
I also maintain a culinary tradition. During a Boreal Flora field trip to the
Upper Peninsula, we fixed dinner over Coleman stoves at a roadside rest along
Lake Michigan. That night’s meal was a hamburger-based creation inspired at
the conclusion of a collecting trip which Ed made to North Manitou Island in the
1950’s. I pirated the recipe for “Manitou Mystery” while assisting in its prepara¬
tion. Today, I continue to serve a version of it on nearly all my overnight field
trips with students. Typically, the circumstances behind the meal’s genesis plus
my involvement with the Manitou Islands are recounted. This narrative also in¬
cludes recalling a few noteworthy field trips when I have prepared it. From now
on, I am sure that I will add a few words about the person who invented the
meal. And who also inspired me.
2012
THE MICHIGAN BOTANIST
49
ED VOSS— AS AN EDITOR
Richard K. Rabeler
University of Michigan Herbarium
3600 Varsity Drive
Ann Arbor MI 48108-2228
Thomas J. Rosatti
University and Jepson Herbaria
1001 Valley Life Sciences Building #2465
University of California, Berkeley
Berkeley, CA 94720-2465
Ed Voss founded The Michigan Botanist , with the first issue appearing 1962.
It was “his” from the start. Not only did he edit submitted articles, he provided
many book reviews, and started several regular features. Michigan Plants in
Print was an annotated list of “significant literature relating to Michigan botany
since the beginning of 1960”; installments of this feature appeared in 27 volumes
of the journal (see list in the introductory article). News of Botanists included
notes about faculty assuming new positions, graduate students completing de¬
grees and assuming new positions, etc. He oversaw the editing and production of
the first 15 volumes, handing over the editorial reigns to Howard Crum in 1977;
the Club honored Ed for his service as the first editor with an award. He contin¬
ued to serve as an advisor to the journal, remaining on the Editorial Board until
his passing.
As many know, Ed was an editor’s editor. He developed a command for the
English language, no doubt fostered by his father’s experiences in teaching Latin
and Greek. Typos, incorrect usage, and factual errors were not for him. The edi¬
torial staff at the Ann Arbor News knew him well from the many letters to the
editor that he penned over the years.
I (RKR) remember when I submitted my second paper to The Michigan
Botanist around 1980. Although not the editor at the time, Ed reviewed the
paper. His review was nearly as long as the paper — scary to a grad student, but
the thoroughness made it a much better paper.
When I (RKR) became one of the co-editors of The Michigan Botanist in
1988, I knew that he always inspected the latest issue when it arrived. I always
approached the release of another issue with trepidation — would the “O” in
“Botanist” be precisely aligned over the Straits of Mackinac on the cover?
Would Ed spot some needless typos? Would the fourth number of a volume be
mailed during the same calendar year — Ed did notice the one time when that did
not take place. Actually, having the founding editor watching likely made my
products better!
Anyone who knew Ed was well aware that Ed “told it like it is”; no skirting
an issue for him. Ed received a copy of a volume entitled Manual of the Seed
50
THE MICHIGAN BOTANIST
Vol. 51
Plants of Indiana to review in 1989. One also readily knew when Ed did not like
something. He started his review with “That whirring noise you hear in the back¬
ground is Charles C. Deam [author of the 1940 Flora of Indiana] turning in his
grave” — and it got worse. In fact, I had to add an editorial note at the end of his
two-page review when it appeared in 1990 — “This book has been withdrawn by
the publisher.” How many reviewers can claim that distinction?
I (TJR) always will remember, with great fondness and admiration for the
man, the time I asked Dr. Voss to serve on my dissertation committee, as co-
chairman. I had been having trouble working up the nerve to do so, because I had
gotten the feeling that he had not been writing to any relatives about my work, at
least not in a good way. So, I began very cautiously, but before I could get to the
actual question, he announced, in that abrupt, loud, deep, clear, certain way of
his, “No way!”. Luckily, I had experienced this sort of thing before (my dad was
comparably decisive), and so was not completely deterred, but I did proceed
from that point even more cautiously in establishing that I had not yet asked him
anything, and that if he would clarify just what he was refusing to do, it might be
helpful. He responded that because he knew nothing about “species problems”,
he was not qualified to review my dissertation proposal. Now I was seriously
concerned, because my intention had been to ask for much more than this. I re¬
sponded with a very subdued, “Oh . . . uh, um, er, actually, I was intending to ask
about something far more involved: that you not only review my proposal, but
serve on my dissertation committee, as well . . . and that you not only serve on
it, but serve on it as, um, er . . . co-chairman.” Somehow, that did not send Dr.
Voss heading for the hills, possibly because I had quickly followed those re¬
marks with an explanation that I didn’t need him to know anything about
“species problems” per se, but rather that I simply wanted my work to benefit
from what I had observed to be his tough, thorough, rigorous, honest, fair, and
nearly always correct scrutiny.
I (TJR) had this experience in mind as I handed Dr. Voss an early draft of my
dissertation, while standing in his office, marveling over the fact he had a little
stand for his phone so he could make use of the desk surface underneath it. We
chatted about various problems I had had along the way in addressing my dis¬
sertation topic, as Dr. Voss paged through what I had handed him. At one point
he interrupted something I was saying to point out that something I had written
near the end contradicted something I had written a couple of hundred pages ear¬
lier. He was correct about the contradiction, of course, and this, together with the
fact that he had noticed it so easily and so readily, made me realize I had done
the right thing several years earlier in standing in there until he reversed his ini¬
tial position and agreed to serve on my dissertation committee, as co-chairman.
A couple years later, I (TJR) handed over to Dr. Voss a reprint of the first pub¬
lication to result from my post-doctoral gig, which was to prepare treatments for
the Generic Flora of the Southeastern United States. For what was the first time
in my recollection, Dr. Voss seemed pleased to have something I had given him.
As he slowly went through it, handling each page in a way that reminded me of
the way he would hold up a plant about which he was teaching, with delicacy
and appreciation, he said that he loved that series of papers, because “every sin¬
gle mark on every single page means something,” which I took as the ultimate
2012
THE MICHIGAN BOTANIST
51
compliment for my editors, and initially as a comment that had very little to do
with me. However, as it turned out, it had a lot to do with me, in the sense that it
had a large, important impact on my development as an editor, as well as author:
it planted in my brain, just as firmly and significantly, relatively speaking, as that
flag was planted atop Mount Suribachi, that each mark does matter, both in its
own right and in terms of the effect it has on higher levels of organization: the
groupings of marks into words, phrases, keys, descriptions, treatments, and
floras.
This thought occurred to me (TJR) countless times over the past 25 years,
over and over again, as I did my part in transforming manuscripts that had been
submitted by hundreds of authors into pieces that would fit together into a much
larger whole: The Jepson Manual — Higher Plants of California, and just recently
its Second Edition (TJM2).
For the past couple of months, I (TJR) have been making corrections to
TJM2. That they number in the hundreds has not been the most satisfying news,
but it has gotten me to think yet again about the great concern that Dr. Voss
demonstrated so many years ago for all those marks that are supposed to mean
something. Although I am sure that there would have been many fewer correc¬
tions had Dr. Voss himself been more directly involved with our project (i.e.,
more directly involved than having inspired and trained its Scientific Editor), I
was heartened by something that we determined just a few days ago: there are
over 10 million marks in TJM2 (if spaces are counted), so the fact that a few
hundred of them are incorrect is . . . completely unacceptable! As (nearly) al¬
ways, yes, you are correct. Dr. Voss; we needed you, we thank you, and we will
miss your influence.
52
THE MICHIGAN BOTANIST
Vol. 51
EDWARD G. VOSS,
AN ACCOMPLISHED ENTOMOLOGIST
Brian G. Scholtens
Biology Department
College of Charleston
Charleston, SC 29424
Edward Voss’ interest in entomology developed early in life, certainly in
large part due to his consistent exposure to nature and interesting habitats each
summer at his family’s summer cottage in Mackinaw City, MI. Ed’s parents
encouraged his natural history pursuits, allowing the rearing of caterpillars and
collections of plants and insects to accumulate in the cottage and at home in
Toledo, OH. Despite a strict bedtime, Ed was allowed special dispensation to
set an alarm and get up late at night to go out and collect moths at lights. In ad¬
dition, he was regularly allowed to ride his bicycle several miles out of town
to favorite collecting locations, e.g. Cecil Bay in Emmet Co., MI and Stimson
Rd. in Cheboygan Co., MI, both mentioned in his listing of the butterflies of
the area (Voss 1954). He also took his entomological interest with him to
Camp Miniwanca in Oceana Co., MI where he spent a week or two for several
summers. His counselors there also allowed Lepidoptera collecting, resulting
in several interesting records from the Lake Michigan shoreline. I was grati¬
fied to see that the camp still existed when I passed by while conducting an ex¬
tensive survey of the Lake Huron Locust on Great Lakes dunes. This early fas¬
cination turned into an award winning project when, in 1946, Ed was chosen
by the Science Clubs of America as one of the 300 high school senior students
with the most outstanding promise and ability in science based on his perfor¬
mance on a 3-hour exam and an essay about a scientific project, in Ed’s case,
“A Biological Survey” of the Mackinaw City area, focusing primarily on but¬
terflies. This was obviously a very prescient choice for the award on the part
of the Science Clubs of America.
Ed took his interest in entomology to college at Denison University from
1946 to 1950. Here he met and worked with the nationally known skipper expert,
Arthur Ward Lindsey, who had published extensively on this group of Lepi¬
doptera (Lindsey 1921; Lindsey et al. 1931). This collaboration resulted in a se¬
nior thesis examining the phylogeny of the skippers (Hesperioidea) worldwide
(Voss 1952). He used specimens from Lindsey’s extensive collection and also
borrowed specimens from around the world to complete tedious dissections and
an analysis of morphology in the group. Several others have worked on this
group subsequently (e.g. Evans 1937, 1949, 1951, 1952, 1953, 1955; and Watson
1893 before him) and the morphological diversity of the group has always made
determining relationships difficult. When molecular techniques were finally ap¬
plied to the problem along with the morphological evidence (Warren et al.
2012
THE MICHIGAN BOTANIST
53
2009), Ed’s original phylogeny was found to be very close to the currently ac¬
cepted hypothesis.
Ultimately, Ed’s interests transitioned into a greater focus on botany as a
graduate student at the University of Michigan (graduating with a Ph.D. in
1954), but he always remained actively involved with entomology, particularly
studying the Lepidoptera of northern Michigan. Shortly after arriving at
Michigan and taking on the career project of the “Michigan Flora,” he was
again actively publishing on butterflies and moths. Over a period of many
years, he wrote a series of papers cataloging all the known species of butter¬
flies and other Macrolepidoptera, the larger moths, of the Emmet Co. and
Cheboygan Co. region (Voss 1954, 1970, 1981, 1983, 1991, 2002). This has
proven to be one of the most complete regional surveys in North America, with
Ed having recorded 747 species in 19 families (as currently defined). I have
since taken over the survey and started work on listing the Microlepidoptera of
the region, so far adding 187 species in the Pyralidae and Crambidae
(Scholtens 1996) as well as tallying 509 species in other families (Scholtens
unpublished data), for a current total list of 1443 species in the region. Perhaps
the most comparable regional survey at a similar latitude was one done by
Procter on Mt. Desert Island in Maine (Procter 1946) in which he recorded
1479 species, so the Voss-initiated survey effort compares very favorably. My
sampling over 25 years, during and since the publication of the Voss lists, has
added very few additional species in the families that he covered, confirming
the thoroughness of his original work.
Ed’s sampling for this effort concentrated on the area around Mackinaw City
and the property of the University of Michigan Biological Station, where he was
a student and teaching assistant from 1949-1953 and taught botany courses from
1963 through 1998 and again in 2003. An extensive knowledge of the botany
and habitats of the region helped him seek out other unique areas for sampling
such as fens, bogs, and dunes. Notes about the occurrence of species in these
habitats pepper the accounts in his papers. I regularly tapped into that expertise
as I went looking for particular species of butterflies and moths, and I have often
based field trip locations for my summer Biology of Insects and Ecology classes
on this wealth of information.
Voss’ collecting and listing of Lepidoptera had an impact on entomology in
several ways, particularly on our knowledge of the natural history of northern
Michigan. Ed accumulated an extensive collection of Michigan Lepidoptera kept
at his cottage in Mackinaw City (Fig. 26), and donated many specimens to the
University of Michigan Museum of Zoology, the Michigan State University Col¬
lection and the collection at the University of Michigan Biological Station. The
remainder of his collection will be deposited at Michigan State University. His
listings have provided an invaluable dataset recording the occurrence of uncom¬
mon or rare species (e.g. Bates’ crescent: Phyciodes batesi, dusted skipper:
Atrytonopsis hianna, and cobweb skipper: Hesperia metea ), unusual vagrants
(e.g. California tortoiseshell: Nymphalis californica and black witch: Ascalapha
odorata ) (Voss 1950), and local eruptions of populations (e.g. galium sphinx:
Hyles gallii). His papers include excellent documentation of habitat associations
and specific localities where species were found.
54
THE MICHIGAN BOTANIST
Vol. 51
As I have discovered subsequently, these data are also crucial as we start try¬
ing to understand changes in species ranges and abundances due to invasive
species and climate change. Comparisons between Ed’s collection records and
mine in recent years have documented the northward migration of the dion skip¬
per, Euphyes dion , a wetland skipper previously known from the middle lower
peninsula, but now found north to the Straits of Mackinac, and the retreat north¬
ward of at least two butterflies, the greenish blue, Plebejus saepiolus, (evidently
retreating throughout its eastern range) and the branded skipper, Hesperia
comma , still found in the Upper Peninsula. Both of us witnessed the invasion of
the gypsy moth, Lymantria dispar , during the 1990s. Even though northern
Michigan experienced only a single large, extended defoliation event caused by
this European introduction, the legacy of its introduced enemies has apparently
left a lasting effect. In the last decade and a half, other large moths, particularly
Saturniidae, that Ed recorded as common (and still common in the 1980s and
early 1990s), have become very rare or locally extirpated, probably due to para¬
sitism by the introduced tachinid fly Compsilura concinnata. These include the
imperial moth, Eacles imperialism the Io moth, Automeris io , and the Polyphemus
moth, Antheraea polyphemus. Some of these same species suffered the same fate
in the Northeast (Boettner et al. 2000; Goldstein 2010), but populations there
have since shown signs of recovery. We have yet to note any positive trend of re¬
covery in northern Michigan.
Ed was an excellent observer and had a trained taxonomic eye (both in botany
and entomology). Generally, he did not try to obtain long series of species, but
worked primarily on documenting the diversity of the Lepidoptera in the region.
His ability to recognize fine differences among difficult species groups made for
a discerning eye as he collected specimens. His nearly photographic memory
was also valuable as he chose what to collect. He kept copious notes on his ob¬
servations, and occasionally turned these observations into short notes about the
biological interactions of insects, such as a very interesting paper on the pollina¬
tion of blunt-leaf orchids by pyralid ( senso lato) moths (Voss & Riefner 1983).
He also regularly incorporated these observations into his teaching in botany
classes, helping students learn about pollination systems and the influence of
herbivores on plants.
During his career, Ed built collaborations with many different entomologists.
Lindsey was probably his first major entomological contact, but he soon met
many other well-known lepidopterists, many through the Lepidopterists’ Soci¬
ety, in which he was a charter member and just recently elected as an honorary
life member. At the University of Michigan, his colleagues included W. H. Wag¬
ner, Jr., another multidisciplinary biologist who was a professor in botany but
also collected and published on Lepidoptera, including a note about the occur¬
rence of two rare species in northern Michigan (Voss & Wagner 1956). Ed be¬
came an important entomologist in the state and regularly consulted with other
entomologists from around the state, particularly through the Michigan Entomo¬
logical Society. He was also a longtime member of the Ohio Lepidopterists and
the Kentucky Lepidopterists, and built a large library of Lepidoptera literature
based on his contacts across the continent.
Edward Voss’ career will, of course, be marked primarily by the completion
2012
THE MICHIGAN BOTANIST
55
FIGURE 26. Ed Voss’s laboratory space in the family cottage where he did
his Lepidoptera work, and also some plant work in the summers. Photo by
Ed Voss, 18 June 1990.
and publication of the Michigan Flora , a monumental achievement by itself. He
completed this effort shortly before his retirement in 1996. In typical Voss fash¬
ion, he also planned his publication of the Lepidoptera lists of northern Michi¬
gan, and finished the final planned installment shortly after his retirement (Voss
2002). Ed’s was a well-planned and well-lived life and career with a great influ¬
ence in two complementary spheres of biology, botany and entomology.
Appropriately, Ed’s first visit to the Biological Station, a place where he spent
most of the entomological part of his career, was in 1938, when, at age 9, his fa¬
ther took him to the Station on visitor’s day and bought him his first insect pins.
56
THE MICHIGAN BOTANIST
Vol. 51
Likewise, his life was nicely bookended by a trip to the Biological Station on
visitor’s day in 201 1. He spent the day speaking with colleagues, visiting friends,
and enjoying one of the most beloved places of his life and career.
LITERATURE CITED
Boettner, G. H., J. S. Elkinton, & C. J. Boettner. 2000. Effects of a biological control introduction on
three non-target native species of satumiid moths. Conserv. Biol. 14: 1798-1806.
Evans, W. H. 1937. A catalogue of the African Hesperiidae in the British Museum. British Museum,
London. 212 pp.
- . 1949. A catalogue of the Hesperiidae from Europe, Asia and Australia in the British Museum
(Natural History). British Museum, London. 502 pp.
- . 1951. A catalogue of the American Hesperiidae indicating the classification and nomenclature
adopted in the British Museum. Part I. Introduction and Pyrrhopyginae. British Museum, London.
92 pp.
- . 1952. A catalogue of the American Hesperiidae indicating the classification and nomenclature
adopted in the British Museum. Part II. Pyrginae. Sec. 1. British Museum, London. 178 pp.
- . 1953. A catalogue of the American Hesperiidae indicating the classification and nomenclature
adopted in the British Museum. Part III. Pyrginae. Sec. 2. British Museum, London. 246 pp.
- . 1955. A catalogue of the American Hesperiidae indicating the classification and nomenclature
adopted in the British Museum. Part IV. Hesperiinae and Megathyminae. British Museum, Lon¬
don. 449 pp.
Goldstein, P. Z. 2010 [“2003”]. Life history of the imperial moth Eacles imperialis (Drury) (Satumi-
idae: Ceratocampinae) in New England, U.S.A.: distribution, decline, and nutritional ecology of a
relictual islandic population. J. Res. Lepid. 42: 34^-9.
Lindsey, A. W. 1921. The Hesperioidea of America north of Mexico. Univ. Iowa Studies Nat. Hist.
9: 1-114.
Lindsey, A. W., E. L. Bell, & R. C. Williams. 1931. The Hesperioidea of North America. Denison
Univ. Bull., J. Sci. Lab. 26: 1-142.
Procter, W. 1946. Biological survey of the Mount Desert region. The insect fauna with references to
methods of capture, food plants, the flora and other biological features. The Wistar Institute of
Anatomy and Biology, Philadelphia. 566pp.
Scholtens, B. G. 1996. Moths of the Douglas Lake region (Emmet and Cheboygan counties), Michi¬
gan: V. Crambidae and Pyralidae (Lepidoptera). Great Lakes Entomol. 29: 141-160.
Warren, A. D., J. R. Ogawa, & A. V. Z. Brower. 2009. Revised classification of the family Hesperi¬
idae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Syst. En¬
tomol. 34: 467-523.
Watson, E. Y. 1893. A proposed classification of the Hesperiidae, with a revision of the genera. Proc.
Zool. Soc. London 1893: 3-132.
New York Botanical Garden Library
3
5185 00269 4287
INSTRUCTIONS TO AUTHORS
1. Create text in 12-point Times New Roman font and double space paragraphs throughout. Papers
should be organized as follows: Title, Author(s) and address(es), Abstract with up to 5 keywords,
Introduction, Materials and Methods, Results, Discussion, Acknowledgements, Literature Cited,
Tables, Figure Legends, and Figures. Sections may be omitted if not relevant. All pages should
be numbered. Please contact the editor regarding any questions related to formatting.
2. For noteworthy collections, manuscripts should be formatted as described in The Michigan
Botanist, volume 27(3) p. 90. A brief description of the formatting follows. The following title,
“Noteworthy collections”, should begin each submitted manuscript followed on the next line by
the State or Province for the species reported. The next line should list the taxon of interest using
the following format: Species Author(s) (Family). Common name. The rest of the manuscript
should include the following named sections: Previous knowledge, Significance of the report.
Diagnostic characters (if desired), Specimen citations, and Literature cited. Each of these sec¬
tions are largely self explanatory; however, “specimen citations” should include the relevant
label data from the voucher specimen(s) including location data, collector(s), collection number,
etc. Also please include which herbarium the specimen(s) is deposited in using the Index
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3. Letters to the Editor can be formatted as general text without the specific sections listed above.
However, literature cited and any tables or figures should be formatted as described below.
4. Please create tables using either a tab delimited format or a spreadsheet using Excel or other sim¬
ilar program. Each table is to be submitted as a separate file. Table captions should be placed at
the top of the table. Any footnotes should appear at the bottom of the table. Please do not insert
tables within the body of the text.
5. Send each figure as a separate file in a high-resolution format — eps, jpg, or tif. Figures like bar
graphs that gain their meaning with color won’t work — use coarse-grained cross-hatching, etc.
Create figure legends as a separate text file, and the typesetter will insert them as appropriate.
Please do not insert the figure in the body of the text file.
6. Citations: Please verify that all references cited in the text are present in the literature cited sec¬
tion and vice versa. Citations within the text should list the author’s last name and publication
year (e. g. Smith 1990). For works with more than 2 authors, use “et al.”, and separate multiple
citations with a semicolon.
7. Literature Cited: List citations alphabetically by author’s last name. Author names are to be list¬
ed with surname first, followed by initials (e. g. Smith, E. B.). Separate author’s initials with a
single space. The year of publication should appear in parentheses immediately before the title
of the citation. The entire journal name or book title should be spelled out. Please put a space
after the colon when citing volume number and page numbers.
8. Italicize all scientific names. Voucher specimens must be cited for floristic works or any other
relevant study. Papers citing plant records without documenting vouchers are generally not
acceptable.
9. Manuscripts may be submitted electronically to the email address of the editor. Printed versions
of manuscripts may also be submitted in which case three copies should be provided. All manu¬
scripts will be reviewed by at least two referees. A more complete set of instructions is available
at http://www.michbot.org/publications/Botanist/instruct_authors.htm.
CONTENTS
Edward G. Voss (1929-2012)
Anton A. Reznicek and Richard K. Rabeler 1
“Fraternally, Ed” - A Remembrance of Ed Voss from
Undergraduate Days
Tom S. Cooperrider 18
Beyond The Botany— Life with Ed Voss
John S. Russell 20
Ed Voss— Field Botanist in Summer, Curator in Winter
A. A. Reznicek 24
The Botanical Teaching Legacy of Edward G. Voss at the
University of Michigan Biological Station
Charles C. Davis, Melanie Gunn, and C. Eric Hellquist 32
Ed Voss and the Conservation of Michigan’s Rare Flora
Michael R. Penskar 42
The Road to the Isles: Reminiscences of
Ed Voss as Dissertation Chairman
Brian T. Hazlett 45
Ed Voss— as an Editor
Richard K. Rabeler and Thomas J. Rosatti 49
Edward G. Voss, an Accomplished Entomologist
Brian G. Scholtens 52
On the cover: Ed Voss camping on Cusino Lake, near Lake Superior
with his Jeep in Aug. 1966. Photo by John Russell.
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April-June, 2012
Vol. 51, No. 2
THE
MICHIGAN BOTANIST
A Journal of Great Lakes Botany
THE MICHIGAN BOTANIST (ISSN 0026-203X) is published four times per year by the Michigan
Botanical Club (www.michbotclub.org) and is available online at http://quod.lib.umich.edu/rn/
mbot/. The subscription rate is $25.00 per year. Periodicals postage paid at Ann Arbor, MI 48103.
The office of publication is Western Michigan University, Kalamazoo, MI 49008.
On all editorial matters, please contact Todd J. Barkman, 3437 Wood Hall, Department of Biological
Sciences, Western Michigan University, Kalamazoo, MI 49008; 269. 387. 2776 (Phone), 269.
387. 5609 (FAX); todd.barkman@wmich.edu. All articles dealing with botany in the Great Lakes
region may be sent to the Editor at the above address. In preparing manuscripts, authors are
requested to follow the “Instructions for Authors” on the inside back cover.
For all inquiries about back issues and institutional subscriptions please contact Caroline Barkman,
The Michigan Botanist Business Office, 919 Dobbin Dr., Kalamazoo, MI 49006; 269. 544. 0034;
cbarkman9147@mail.kvcc.edu
Todd J. Barkman, Editor
Editorial Board
Caroline Barkman, Business Manager
L. Alan Prather
Anton A. Reznicek
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Catherine H. Yansa
THE MICHIGAN BOTANICAL CLUB
Membership is open to anyone interested in its aims: conservation of all native plants; education of
the public to appreciate and preserve plant life; sponsorship of research and publication on the
plant life of the state and the Great Lakes area in general, both in the USA and in Canada; spon¬
sorship of legislation to promote the preservation of Michigan’s native flora; establishment of
suitable sanctuaries and natural areas, and cooperation in programs concerned with the wise use
and conservation of all natural resources and scenic features.
Dues are modest, but vary slightly among the chapters. To become a chapter member please contact
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riverview.net
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2012
THE MICHIGAN BOTANIST
57
ASSESSMENT OF A NOVEL RECLAMATION EFFORT
IN GRAND MERE STATE PARK
Megan D. Edwards, Ranessa L. Cooper1,
Manning Serafin, and Brooke E. Toppen
Hillsdale College
Biology Department
33 E. College St.
Hillsdale, MI, 49242
ABSTRACT
The Great Lakes sand dunes, the largest system of freshwater sand dunes in the world, are a nat¬
ural resource with both ecological and economic value. In 2000, the Michigan DNR proposed a
novel reclamation plan to restore one of these dunes after being mined. This dune area, the former
Manley-Peters Sand Mine (MPSM), is located in Grand Mere State Park in Berrien County, Michi¬
gan. Local genotypes of 115 native dune species were planted within seven microhabitats over a
three- to four-year timeframe. As of 201 1, the vascular plant flora at the MPSM reclamation site con¬
sists of 150 taxa, representing 50 families. A 56.5% post-planting survival was observed for the orig¬
inally planted species from the reclamation plan, and 58 native species have established on their own
at the site. Of these native volunteers, Carex umbellata was a new report for Berrien County’s flora.
The 2011 study has provided a baseline assessment of species richness for use in long-term monitor¬
ing of this reclaimed sand dune area.
KEYWORDS: Grand Mere State Park, Great Lakes, Michigan flora, reclamation, sand dunes
INTRODUCTION
Michigan benefits from many natural resources, some of which are economi¬
cally important, while others have ecological value. As part of the largest fresh¬
water dune system in the world, the Lake Michigan sand dunes provide suitable
habitat for many plants and wildlife, giving the dunes high ecological worth.
Lake Michigan’s dunes are particularly valuable in industry because they contain
nearly pure quartz mineral content (Wilson, 2001), and they have been mined for
automobile foundries since 1910 because of the sand’s exceptional purity, grain
size, and texture (Ayers et al., 2001).
The Manley-Peters Sand Mine (MPSM), located in Grand Mere State Park of
Berrien County, Michigan, has been intermittently active since 1965, when the
Manley Brothers began operations. The passing of the 1976 Sand Dune Protec¬
tion Act ended unmonitored mining of sand dunes and required companies to re¬
claim used dunes. After several years of inactivity at the Peters mine, the Man-
ley Brothers requested a permit from the Michigan Department of Natural
Resources (MDNR) in 1979 to continue mining. After a couple of legal battles,
the Manley Brothers and the MDNR agreed to a Progressive Cell-Unit Mining
and Reclamation Plan (Palmgren, 2004).
'Author for correspondence (rcooper@hillsdale.edu)
58
THE MICHIGAN BOTANIST
Vol. 51
TechniSand, Inc. bought the mine from the Manley Brothers in 1999. A 1999
letter of agreement outlined the financial responsibilities of both TechniSand,
Inc. and the MDNR State Park Stewardship Program in restoring the mine back
to a natural area. Under a new permit with the MDNR (1999), TechniSand (now
Fairmount Minerals) was allowed to harvest sand from the site until June 17,
2003. The company promised to cooperatively implement a plan with MDNR to
reclaim the dunes which included mining below the water table to add to habitat
diversity for the reclamation effort (Palmgren, 2004).
Previous dune stabilization projects in Michigan have used Ammophila bre-
viligulata (beach grass) monocultures in their reclamation protocols (Wood-
house, 1982), as beach grass prevents dune loss due to wind and water erosion.
However, a novel reclamation approach was proposed by MDNR at MPSM to
develop “functioning ecosystems” by establishing seven different planting
zones, or microhabitats (Palmgren, 2004). The seven microhabitats established
at the MPSM reclamation site were steep slope, gentle slope, prairie, wetpanne,
wetland shelf, shallow water, and deep water (Fig. 1). A detailed planting sched¬
ule comprised of native dune taxa was outlined by Palmgren (2000) and then re¬
vised in 2004 after the reclamation effort was underway. Over 550 plant taxa
have been documented in Grand Mere State Park, and local genotypes of 115 of
these species (of plugs, seedlings, seeds, shrubs, transplants and/or tubers; see
Appendix I) were planted throughout the 40-acre site, according to the restora¬
tion plan (Palmgren, 2004; 2000).
In 2004, we began visits to the site while the reclamation was in progress, and
a follow-up evaluation took place in 2008 after the plantings had time to estab¬
lish. With comparison of the revised 2004 planting list from MDNR, the primary
objective of our 2011 study was to generate an official checklist of vascular
plants at the MPSM reclamation site to assist in determining post-planting sur¬
vival of the original plantings. Second, we estimated species richness for the site
overall, as well as for five (steep slope, gentle slope, prairie, wetpanne, and wet¬
land shelf) of the seven microhabitats through detailed surveys and additional
quadrat analyses. Furthermore, the 2011 survey of the MPSM reclamation site
flora will provide a baseline assessment for use in long-term monitoring of this
novel reclamation effort in Grand Mere State Park.
MATERIALS AND METHODS
A Michigan DNR permit (#RD-SU-201 1-010) was granted for field work and the collection of
plants at the MPSM reclamation site. From May to September of 2011, ten collecting trips were
made to the site in Grand Mere State Park to document the existing flora. Each species was recorded
as part of the original reclamation plan, as a newly established native of Michigan’s flora, or as a
newly established adventive (i.e., non-native) species. Post-planting survival of the planted species
from the MDNR list was calculated. Most of the documented species were collected as voucher spec¬
imens and deposited in the Hillsdale College Herbarium (HLSD). The Michigan Flora website
(http://michiganflora.net) was used to assign native or adventive rankings to the taxa that had estab¬
lished at the site, and taxonomy for all species follows that of the website too. Species were also
recorded specifically for the steep slope, gentle slope, prairie, wetpanne, and wetland shelf micro¬
habitats across the entire reclamation area to estimate species richness for each.
2012
THE MICHIGAN BOTANIST
59
Grand Mere State Park: Peters Sand Mine Reclamation
Planting Zones
| | | 1 1 Steep slopes (S/W/E aspects)
j Steep slopes (N aspects)
I - 1 Gentle slopes
| j Prairie
0 100 200 400
V/A Wetpanne (water table @ surface)
HHH Wetland shelf (1 - 2 ft deep)
Shallow water (2 - 6 ft deep)
Deep water (> 6 ft deep)
600
800
1,000
□ Feet
Planting Zones: Digitized by GRP 2002
Aerial Photo: Bing Imagery, 2010
Prepared: May 20, 2011 (GRP)
FIGURE 1 . The Manley-Peters Sand Mine reclamation area post-mining showing the planting zones,
or microhabitats. (Map courtesy of Glenn Palmgren; Graphic assistance provided by Angela Lash-
away.)
60
THE MICHIGAN BOTANIST
Vol. 51
■
FIGURE 2. Photo of the reclamation in progress in 2004. Gentle slope habitat is in the foreground
adjacent to the wetpanne habitat, while steep slopes occupy the background. (Photo by Ranessa
Cooper.)
RESULTS
The MPSM reclamation site has changed considerably since our first visits in
2004 (Fig. 2), and it is evident that the vegetation is denser in the steep slope,
gentle slope, and wetpanne habitats (Figs. 3, 4). The 2011 survey revealed that
150 vascular plant species occupy the site, representing 50 families (Table 1).
The presence of Carex umbellata (sedge) at the MPSM reclamation site repre¬
sents a new report for the native flora of Berrien County. Another native, Juncus
articulatus (jointed rush) is new to the Michigan Flora Online for Berrien
County, although it was previously reported from there by the Plants of the
Chicago Region (Swink and Wilhelm, 1994).
In Palmgren’s (2004) revised planting list, 115 species were planted across
the site among seven zones, or microhabitats (see also Appendix I). In the five
microhabitats surveyed here, 65 planted species were documented (Table 1), re¬
sulting in a 56.5% post-planting survival from the original planting list. The
steep slopes, gentle slopes, prairie, and wetpanne maintained at least 41% of the
taxa originally planted in each microhabitat, while the wetland shelf had 35.7%
of its originally planted species still persisting (Table 2).
2012
THE MICHIGAN BOTANIST
61
FIGURE 3. Photo of the reclamation area in 2008. Gentle slope habitat is in the foreground adjacent
to the wetpanne habitat, while steep slopes occupy the background. Vegetation is denser compared to
that seen in 2004. (Photo by Ranessa Cooper.)
FIGURE 4. Photo of the reclamation area in 2011. Gentle slope habitat is in the foreground, while
steep slopes occupy the background. Vegetation is denser compared to previous years. (Photo by
Ranessa Cooper.)
62
THE MICHIGAN BOTANIST
Vol. 51
TABLE 1. A 2011 checklist of vascular plants (including microhabitat occurrence) at the former
Manley-Peters sand mine in Grand Mere State Park.
MAJOR TAXONOMIC GROUP
Family — Species* [MicrohabitaC presence — Status^]
Dennstaedtiaceae
Equisetaceae
SEEDLESS VASCULAR PLANTS
Pteridium aquilinum (L.) Kuhn (Bracken fern) [GS— Planted]
Equisetum arvense L. (Common horsetail)* [GS, W — Native]
Equisetum laevigatum A. Braun (Smooth scouring rush)* [GS, W — Native]
Cupressaceae
Pinaceae
GYMNOSPERMS
Juniperus communis L. (Common juniper; ground juniper)* [SS — Planted]
Juniperus virginicinci L. (Red-cedar)* [SS — Native]
Pinus strobus L. (White pine)* [SS — Planted]
Adoxaceae
Alismataceae
Anacardiaceae
Apocynaceae
Asteraceae
ANGIOSPERMS
Viburnum acerifolium L. (Maple-leaved viburnum)* [SS, P — Planted]
Alisma sp. (Water-plantain)* [W — Planted]
Sagittaria latifolia Willd. (Wapato, duck-potato, common arrowhead)*
[WS— Planted]
Rhus typhina L. (Staghorn sumac)* [SS— Native]
Toxicodendron radicans (L.) Kuntze (Poison-ivy) [SS — Native]
Apocynum cannabinum L. (Indian-hemp)* [SS — Native]
Asclepias incarnata L. (Swamp milkweed)* [W — Planted]
Asclepias syriaca L. (Common milkweed)* [SS, GS, P — Native]
Ambrosia artemesiifolia L. (Common ragweed)* [SS— Native]
Anaphalis margaritacea (L.) Benth. & Hook. (Pearly everlasting)*
[W— Native]
Arnoglossum atriplicifolium (L.) H. Rob. (Pale Indian plantain)*
[GS— Planted]
Artemisia campestris L. (Wild wormwood)* [SS, GS, P, W — Planted]
Artemisia vulgaris L. (Mugwort)* [SS, GS — Adventive]
Bidens cernua L. (Nodding beggar-ticks)* [W — Planted]
Centaurea stoebe L. (Spotted knapweed)* [GS — Adventive]
Cirsium arvense (L.) Scop. (Canada thistle, field thistle)* [W — Adventive]
Cirsium muticum Michx. (Swamp thistle)* [W — Planted]
Conyza canadensis (L.) Cronq. (Horseweed)* [W — Adventive]
Coreopsis tripteris L. (Tall tickseed, tall coreopsis)* [P, W — Planted]
Erigeron annuus (L.) Pers. (Daisy fleabane)* [SS, GS, W — Native]
Erigeron philadelphicus L. (Common fleabane, Philadelphia fleabane)*
[W— Native]
Eupatorium perfoliatum L. (Boneset)* [GS, W— Planted]
Euthamia graminifolia (L.) Nutt. (Grass-leaved goldenrod)* [SS — Planted]
Helianthus divciricatus L. (Woodland sunflower)* [GS — Native]
Hieracium caespitosum Dumort. (King devil, yellow hawkweed)* [P —
Adventive]
Hieracium flagellare Willd. (Whip-lash hawkweed)* [GS— Adventive]
Hypochaeris radicata L. (Cat’s ear)* [P — Adventive]
Krigia virginica (L.) Willd. (Dwarf dandelion)* [SS, GS, P — Native]
Rudbeckia hirta L. (Black-eyed susan)* [GS, W — Planted]
Scorzoneroides autumnalis (L.) Moench (Fall-dandelion)* [W — Adventive]
Solidago altissima L. (Tall goldenrod)* [SS — Native]
Solidago nemoralis Aiton (Gray goldenrod, old-field goldenrod) [SS —
Planted]
Solidago rugosa Mill. (Rough-leaved goldenrod)* [W — Native]
(Continued)
2012
THE MICHIGAN BOTANIST
63
TABLE 1. (Continued).
Asteraceae (cont.)
Boraginaceae
Brassicaceae
Cactaceae
Campanulaceae
Caryophyllaceae
Ceratophyllaceae
Cleomaceae
Commelinaceae
Cornaceae
Cyperaceae
Elaeagnaceae
Euphorbiaceae
Fabaceae
Fagaceae
Gentianaceae
Hamamelidaceae
MAJOR TAXONOMIC GROUP
Family — Species* [MicrohabitaC presence — Status*]
Symphyotrichum dumosum (L.) G. L. Nesom (Bushy aster)* [W — Native]
Symphyotrichum lateriflorum (L.) A. Love & D. Love (Calico aster)*
[GS — Native]
Symphyotrichum pilosum (Willd.) G. L. Nesom (Frost aster, hairy aster)*
[SS— Planted]
Tragopogon dubius Scop. (Goat’s beard)* [SS, GS, P — Adventive]
Vernonia missurica Raf. (Missouri ironweed)* [GS — Planted]
Lithospermum caroliniense (Walter) MacMill. (Hairy puccoon, yellow
puccoon, plains puccoon)* [P — Planted]
Arabidopsis lyrata (L.) O’ Kane & Al-Shehbaz (Sand cress)* [SS — Native]
Lepidium campestre (L.) R. Br. (Field cress)* [W — Adventive]
Lepidium virginicum L. (Common peppergrass)* [P — Native]
Opuntia humifusa (Raf.) Raf. (Prickly-pear)* [GS — Planted]
Campanula rotundifolia L. (Bluebell, harebell)* [P — Planted]
Lobelia cardinalis L. (Red lobelia, cardinal-flower)* [W — Planted]
Lobelia kalmii L. (Kalm’s lobelia, brook lobelia, bog lobelia)* [P, W — Native]
Lobelia siphilitica L. (Great blue lobelia)* [W — Planted]
Cerastium semidecandrum L. (Small mouse-ear chickweed)*
[GS — Adventive]
Saponaria officinalis L. (Bouncing bet, soapwort)* [SS — Adventive]
Ceratophyllum demersum L. (Coontail)* [WS- — Native]
Polanisia dodecandra (L.) DC. (Clammy-weed)* [GS — Native]
Tradescantia ohiensis Raf. (Common spiderwort)* [GS, P — Planted]
Cornus drummondii C. A. Mey. (Rough-leaved dogwood)* [SS — Native]
Cornus sericea L. (Red-osier)* [SS — Planted]
Carex comosa Boott. (Sedge)* [WS — Planted]
Carex crinita Lam. (Sedge)* [WS — Native]
Carex flava L. (Sedge)* [W — Planted]
Carex hystericina Willd. (Sedge)* [W — Planted]
Carex lurida Wahlenb. (Sedge)* [W — Planted]
Carex muehlenbergii Willd. (Sedge)* [GS, W- — Native]
Carex pensylvanica Lam. (Sedge)* [W— Planted]
Carex stricta Lam. (Sedge)* [W — Planted]
Carex suberecta (Olney) Britton (Sedge)* [W — Planted]
Carex umbellata Willd. (Sedge)* [GS — Native]
Cyperus schweinitzii Torr. (Rough sand sedge)* [SS, GS — Native]
Cyperus strigosus L. (Long scale nut sedge)* [W — Native]
Eleocharis erythropoda Steud. (Spike-rush)* [W — Native]
Eleocharis obtusa (Willd.) Schult. (Spike-rush)* [W — Native]
Schoenoplectus acutus (Bigelow) A. Love & D. Love (Hardstem bulrush)*
[W, WS— Planted]
Schoenoplectus pungens (Vahl) Palla (Threesquare)* [W — Planted]
Scirpus cyperinus (L.) Kunth (Wool-grass)* [W — Planted]
Elaeagnus umbellata Thunb. (Autumn-olive)* [SS — Adventive]
Croton glandulosus L. (Tooth-leaved croton)* [SS — Adventive]
Medicago lupulina L. (Black medick)* [GS — Adventive]
Melilotus albus Medik. (White sweet-clover)* [GS — Adventive]
Quercus alba L. (White oak)* [SS, GS — Planted]
Quercus rubra L. (Red oak)* [SS, GS — Planted]
Quercus velutina Lam. (Black oak)* [SS, GS, P — Planted]
Sabatia angularis (L.) Pursh (Rose-pink; rose gentian)* [W — Planted]
Hamamelis virginiana L. (Witch-hazel)* [SS — Planted]
(Continued)
64
THE MICHIGAN BOTANIST
Vol. 51
TABLE 1. (Continued).
Hydrocharitaceae
Hypericaceae
Juncaceae
Lamiaceae
Lauraceae
Lemnaceae
Lentibulariaceae
Magnoliaceae
Moraceae
Nymphaeaceae
Oleaceae
Onagraceae
Orobanchaceae
Poaceae
Polygonaceae
Potamogetonaceae
Ranunculaceae
MAJOR TAXONOMIC GROUP
Family — Species* [MicrohabitaC presence — Status*]
Elodea canadensis Michx. (Common waterweed)* [WS— Native]
Hypericum kalmianum L. (Kalm’s St. John’s-wort)* [W — Planted]
Juncus alpinoarticulatus Chaix (Rush)* [W— Planted]
Juncus articulatus L. (Jointed rush)* [W — Native]
Juncus balticus Willd. (Rush)* [W — Planted]
Juncus brachycephalus (Engelm.) Buchenau (Rush)* [W — Planted]
Juncus canadensis Laharpe (Canadian rush)* [W — Native]
Juncus effusus L. (Soft-stemmed rush) [W, WS — Planted]
Juncus torreyi Coville (Torrey’s rush)* [W — Planted]
Lycopus americanus Muhl. (Common water horehound)*
[SS, GS, P, W— Planted]
Monarda punctata L. (Dotted mint, horse mint)* [GS — Planted]
Sassafras albidum (Nutt.) Nees (Sassafras)* [SS — Native]
Lemna sp. (Duckweed) [W, WS — Native]
Utricularia vulgaris L. (Common bladderwort)* [WS — Native]
Liriodendron tulipifera L. (Tulip tree) [GS — Planted]
Morus alba L. (White mulberry, Russian mulberry)* [SS — Adventive]
Nymphaea odorata Aiton (Sweet-scented waterlily) [WS — Planted]
Fraxinus americana L. (White ash)* [SS — Native]
Oenothera oakesiana (A. Gray) S. Watson & J. M. Coult. (Evening-
primrose)* [W — Native]
Oenothera parviflora L. (Evening-primrose)* [GS — Native]
Agalinis purpurea (L.) Pennell (Purple false foxglove)* [W — Native]
Ammophila breviligulata Femald (Beach grass, marram grass)*
[SS, GS— Planted]
Agrostis gigantea Roth (Redtop)* [P — Adventive]
Bromus japonicus Murray (Japanese brome)* [GS — Adventive]
Bromus tectorum L. (Downy chess, cheat grass)* [SS, GS, P, W— Adventive]
Calamagrostis canadensis (Michx.) P. Beauv. (Blue-joint)* [WS — Planted]
Dichanthelium commutatum (Shult.) Gould (Panic grass)* [GS, P — Native]
Dichanthelium commonsianum (Ashe) Freckmann (Panic grass)* [P — Native]
Dichanthelium latifolium (L.) Harvill (Broad-leaved panic grass)*
[GS, P — Native]
Dichanthelium oligosanthes (Schult.) Gould (Panic grass)* [P — Native]
Elymus canadensis L. (Canada wild rye)* [SS, GS, P — Planted]
Elymus repens (L.) Gould (Quack grass)* [GS, P — Adventive]
Koeleria macrantha (Ledeb.) Schult. (June grass)* [P — Planted]
Panicum virgatum L. (Switch grass)* [GS, P — Planted]
Poa compressa L. (Canada bluegrass)* [GS — Adventive]
Poa pratensis L. (Kentucky bluegrass)* [GS — Adventive]
Schizachyrium scoparium Michx. (Little bluestem)* [P — Planted]
Vulpia octoflora (Walter) Rydb. (Six-weeks fescue)* [SS — Native]
Fallopia cilinodis (Michx.) Holub (Fringed false buckwheat)* [SS — Native]
Rumex acetosella L. (Red sorrel, sheep sorrel)* [SS, GS, P — Adventive]
Potamogeton illinoensis Morong (Illinois pondweed)* [WS — Native]
Potamogeton natans L. (Pondweed)* [WS — Native]
Potamogeton nodosus Poir. (Pondweed)* [WS — Planted]
Stuckenia pectinata (L.) Bomer (Sago pondweed)* [WS — Planted]
Ranunculus longirostris Godr. (White water crowfoot)* [WS — Native]
(Continued)
2012
THE MICHIGAN BOTANIST
65
TABLE 1. (Continued).
MAJOR TAXONOMIC GROUP
Family — Species* [Microhabitat* presence — Status*]
Rosaceae
Rutaceae
Salicaceae
Scrophulariaceae
Solanaceae
Typhaceae
Verbenaceae
Vitaceae
Fragaria virginiana Mill. (Wild strawberry)* [GS — Native]
Rubus allegheniensis Porter (Common blackberry)* [SS, GS — Native]
Rubus flagellaris Willd. (Northern dewberry)* [SS, GS, P — Native]
Ptelea trifoliata L. (Wafer-ash, hop-tree)* [SS, GS — Planted]
Populus deltoides Marshall (Cottonwood)* [GS— Planted]
Populus grandidentata Michx. (Large-tooth aspen, big-tooth aspen)
[SS— Planted]
Salix amygdaloides Andersson (Peach-leaved willow)* [W— Native]
Salix cordata Michx. (Sand-dune willow, furry willow)* [GS — Planted]
Salix exigua Nutt. (Sandbar willow)* [W, WS— Planted]
Salix myricoides Muhl. (Blueleaf willow)* [W — Native]
Salix x pendulina Wenderoth (Weeping willow)* [W — Adventive]
Verbascum thaspus L. (Mullein, flannel plant, common mullein)
[GS, W — Adventive]
Physalis heterophylla Nees (Clammy ground-cherry)* [SS, GS — Native]
Solanum carolinense L. (Horse-nettle)* [SS, GS — Adventive]
Typha latifolia L. (Common cat-tail, broad-leaved cat-tail)* [W, WS — Native]
Verbena hastata L. (Blue vervain)* [SS, W — Planted]
Vitis aestivalis Michx. (Summer grape)* [SS — Native]
Vitis labrusca L. (Fox grape)* [SS — Native]
Vitis riparia Michx. (River-bank grape)* [SS — Planted]
*Voucher specimen(s) deposited in HLSD.
♦Microhabitat abbreviations (SS = Steep slope; GS = Gentle slope; P = Prairie; W = Wetpanne; WS
= Wetland shelf).
♦Status descriptions — “Planted” refers to being part of the original reclamation planting list. “Na¬
tive” (native to Michigan) or “Adventive” (non-native) status refers to establishment at MPSM
reclamation site.
TABLE 2. Numbers of planted, native and/or adventive species present*, including species richness,
in five microhabitats at the former Manley-Peters sand mine in Grand Mere State Park.
Microhabitat
Initial
species
planted
Planted species
Non-planted species
Species
richness
Still
persisting
Additional
colonizers*
Native
Adventive
Steep slopes
36
15
4
21
9
49
Gentle slopes
25
11
9
18
15
53
Prairie
15
7
5
9
7
28
Wetpanne
41
25
4
20
7
56
Wetland shelf
14
5
4
9
0
18
♦“Planted” refers to being part of the Palmgren’s (2004) original reclamation planting list. “Native”
(native to Michigan) or “Adventive” (non-native) status refers to separate establishment from out¬
side the MPSM reclamation site.
♦Species from Palmgren’s (2004) restoration plan that are present in the microhabitat, although not
planted there initially. These taxa likely colonized from original plantings of adjacent microhabitats.
66
THE MICHIGAN BOTANIST
Vol. 51
Some of the species from the planting list occupied more than one habitat, al¬
though only planted in one or two microhabitats (Tables 1, 2; Appendix I). For
example, Lycopus americanus (common water horehound) was only planted in
the wetpanne, yet it was also present on steep and gentle slopes, as well as in the
prairie area (Table 1). The wetpanne had 25 out of 41, or 61%, of its original
species present (Table 2), but 29 species overall, because four additional species
from adjacent microhabitats have established there too (Tables 1, 2). The steep
slopes had 19 planted taxa overall and the gentle slope had 20 species overall
from the original list, while the prairie and wetland shelf had 12 and nine of the
planted species present, respectively (Table 2).
Species richness for the site overall was 150, and as mentioned previously,
some taxa were found in more than one microhabitat (Table 1). Of the micro¬
habitats, the wetpanne had the highest species richness count at 56 taxa, fol¬
lowed by the gentle slopes with 53 taxa (Table 2). Fifty-eight species of Michi¬
gan’s native flora have volunteered at the site, with native taxa being present in
each microhabitat (Tables 1, 2). Twenty-seven adventive species were present
across the MPSM reclamation site too, however, none were found among the
wetland shelf flora (Tables 1,2).
DISCUSSION
As of the 2011 growing season, almost 57% of the original reclamation plant¬
ing list has been documented as part of the existing flora at the MPSM site (Ta¬
bles 1, 2). The success of a reclamation effort can be difficult to quantify; never¬
theless, MDNR originally stated that success would be evaluated based on
continuing colonization of at least 60% of all planted species (Palmgren, 2004).
Although our survey did not reveal 60% post-planting survival of the original
plantings, we recognize the possibility that a few plants may have been over¬
looked, especially where vegetation was dense (as in the wetpanne) or sub¬
merged, and we did not sample the shallow water or deep water vegetation. One
taxon, Lobelia kalmii (Kalm’s lobelia), was part of the original planting list but
then removed as it was difficult to propagate. In the revised plan, Palmgren
(2004) reported that L. kalmii had established naturally by 2003, suggesting an
indication of early progress of the restoration. We consider this restoration a suc¬
cess given that 58 species of native Michigan plants have also established at the
site since the reclamation began (Table 1).
Emory and Rudgers (2010) performed an ecological assessment of 18 dune
restorations in the Great Lakes region, including the MPSM site, and geograph¬
ically paired, natural dune sites. We cannot directly compare our findings with
those of Emory and Rudgers (2010), as data collection methodology and analy¬
ses differed, but they did report that the MPSM site had a higher species richness
than that of its paired, natural site. It would be interesting to acquire the locality
information of MPSM's natural counterpart in Grand Mere State Park, as deter¬
mined by Emory and Rudgers (2010), so that species richness between the two
2012
THE MICHIGAN BOTANIST
67
sites could be effectively compared using our microhabitat-delineated sampling
method.
The presence of newly colonized flora and fauna suggest that the reclamation
site has integrated with the surrounding dune areas of Grand Mere State Park.
Several species of fauna, including bluegill, herons, sandhill cranes, and a red¬
tailed hawk, have been observed at the dune over the years (R. Cooper et al. , per¬
sonal observation). Turkeys were present during several collections, and tracks
and scat provided evidence of the regular presence of deer, as well. One of the
objectives of the reclamation was to provide suitable habitat for both animals
and plants native to the area, and our field observations support this. The wet-
panne area had a species richness of 56, and over a third of the documented taxa
were native volunteers (Table 2). Interdunal wetlands, or wetpannes, are ranked
as imperiled in Michigan and globally (Palmgren, 2004) and thus, having such a
rich flora in the MPSM wetpanne microhabitat is another measure of success for
this thorough restoration effort.
Of the 27 adventive taxa documented at the MPSM site, Bromus tectorum
was found on steep slopes, gentle slopes, the prairie area, and the wetpanne,
while Rumex acetosella occurred in both slope habitats and the prairie (Tables 1 ,
2). At present, it might be appropriate to monitor these invasive species more
closely to judge whether action is needed to remove them from the reclamation
site. Most of the other adventive taxa were found in one or two microhabitats,
but none were documented in the wetland shelf (Tables 1,2).
In July 2012, a site visit was made to confirm identification of some of the
MPSM flora for this publication. The water levels were quite low to the point of
having submerged aquatics exposed in a terrestrial setting. Iris virginica (south¬
ern blue flag), part of the planting list outlined in the reclamation plan, was seen
in 2004, but not again until the recent 2012 visit (R. Cooper and J. Van Zant, per¬
sonal observation). It was present in a few places where the water levels were
low, but not where documented previously in 2004. Hopefully, /. virginica will
be confirmed as one of the post-planting survivors in future assessments.
Three of the 2011 surveys also included 1-m2 quadrat sampling, according to
methods described in Bonham (1989), and these took place across the seasons in
May, June, and July. A “hybrid” version of quadrat and transect methodologies
was used to ensure that each microhabitat was sampled according to the area
each occupied at the site, and at least 38 quadrats were analyzed in each season.
We do not show the data here for frequency and cover abundance, as only a sub¬
set of the total flora fell within the quadrat sampling areas. As might be ex¬
pected, A. breviligulata had high frequency and cover-abundance values, as it
occupies much of the slope habitats and was present in several of the quadrat
samples (Saunders, 2012). We have the GPS coordinates recorded for each
quadrat sample for use in further monitoring if deemed desirable by MDNR.
Human disturbance, in the form of litter and illegal fishing, is evident at the
MPSM reclamation site (M. Edwards and R. Cooper, personal observation).
A“Reclamation in Progress” sign, seemingly more effective during the early
monitoring years, has since been removed. We propose that some signage be
added to the gated area to warn against trespassing. Some level of supervision
68
THE MICHIGAN BOTANIST
Vol. 51
may be warranted in the future, especially during busy holiday weekends in the
spring and summer, to reduce human impacts on the restored site.
The baseline species richness observed in this study will be used in future
monitoring every five years as recommended by MDNR (G. Palmgren, personal
communication). Since planting methods differed among species (see Appendix
I), an analysis of post-planting survival based on planting method would allow
MDNR to better evaluate the value of each approach and, perhaps, modify fur¬
ther reclamation efforts. Future studies on bird, fish, insect, and/or mammal pop¬
ulations would also give a broader picture of the ecological viability of the site.
In our opinion, MDNR has shown the “practicality of replanting a more diverse
and natural vegetation cover which will complement the surrounding undis¬
turbed vegetation” (Palmgren, 2004), and we appreciate being part of such sound
ecological and conservation practices demonstrated at this innovatively re¬
claimed site in southwest Michigan’s Grand Mere State Park.
ACKNOWLEDGEMENTS
The authors are particularly grateful to Mr. Glenn Palmgren (MDNR.) and Mr. Bob Ledyard
(Fairmount Minerals) for granting permission to do research at the former Manley-Peters mine in
Grand Mere State Park, as well as the LAUREATES Program (at Hillsdale College) and the Hills¬
dale College Biology Department for funding this project. Dr. Frank Steiner provided comments on
an earlier draft of this manuscript, and Drs. Tony Reznicek and George Argus provided invaluable
assistance with plant identification. We appreciate the efforts of several friends and colleagues who
have helped us with field work and plant collection since 2004, especially Dr. Jeff Van Zant for his
recent support.
LITERATURE CITED
Ayers, Lewis, Norris, and May, Inc., and M. J. Chapman. (2001) An economic study of coastal sand
dune mining in Michigan. Government of Michigan Department of Environmental Quality, Geo¬
logic Survey Division.
Bonham, C. D. (1989) Measurements for Terrestial Vegetation. New York: John Wiley & Sons.
Emory, S. M. and Rudgers, J. A. (2010) Ecological assessment of dune restorations in the Great
Lakes region. Restoration Ecology 1 8(S 1 ): 184-194.
Michigan, Government of. (1999) Letter of Agreement between TechniSand, Inc. and Michigan
DNR, Parks and Recreation Bureau State Park Stewardship Program.
Palmgren, G. (2000) Sand mine restoration plan for Grand Mere State Park. Project #GL985669-01-
1. Government of Michigan. Prepared for MDNR, Parks and Recreation Bureau and US Environ¬
mental Protection Agency.
Palmgren, G. (2004) Sand mine restoration plan for Grand Mere State Park. Project #GL985669-01-
1. Government of Michigan. Prepared for MDNR, Parks and Recreation Bureau and US Environ¬
mental Protection Agency.
Reznicek, A. A., Voss, E. G., and Walters, B. S. (201 1) Michigan Flora Online. University of Michi¬
gan. Web. 7-13-2012. http://www.michiganflora.net/home.aspx.
Saunders, M. D. (2012) Taxonomic investigation of a revegetated mining site in Grand Mere State
Park: Part III. Undergraduate Honors Thesis in Biology. Hillsdale College.
Swink, F. and Wilhelm, G. 1994. Plants of the Chicago Region 4th ed. Indianapolis: Indiana Acad¬
emy of Science.
Wilson, S. E. (2001) Michigan’s Sand Dunes. Government of Michigan. Department of Environ¬
mental Quality, Geological Survey Division.
Woodhouse, W. W. Jr. (1982) Coastal Sand Dunes of the U.S.: Creation and Restoration of Coastal
Plant Communities. Boca Raton, Florida: CRC Press, Inc.
2012
THE MICHIGAN BOTANIST
69
APPENDIX I. Restoration plan (adapted from Palmgren, 2004) for the former Manley-Peters sand
mine in Grand Mere State Park.
Microhabitat (estimated area)
Species (common name)
Planting method
Steep slopes (18.7 acres)
Ammophila breviligulata (Beach grass, marram grass)
Plugs
Anemone canadensis (Canada anemone)
Seeds
Andropogon gerardii (Big bluestem, turkey foot)
Seeds
Aquilegia canadensis (Wild columbine)
Seeds
Arctostaphylos uva-ursi (Bearberry, kinnikinik)
Plugs
Artemisia campestris (Wild wormwood)
Plugs; seeds
Asclepias verticillata (Whorled milkweed)
Seeds
Calamovilfa longifolia (Sand reed grass)
Seeds
Campanula rotundifolia (Bluebell, harebell)
Seeds
Carpinus caroliniana (Hornbeam, blue-beech)
Seeds
Cornus sericea (Red-osier)
Plugs
Elymus canadensis (Canada wild rye)
Seeds
Euphorbia corollata (Flowering spurge)
Plugs
Hamamelis virginiana (Witch-hazel)
Plugs
Maianthemum stellatum (Starry false soiomon-seal)
Plugs
Juniperus communis (Common juniper; ground juniper)
Plugs
Lathyrus japonicus (Beach pea)
Seeds
Monarda punctata (Dotted mint, horse mint)
Seeds
Panicum virgatum (Switch grass)
Seeds
Pinus strobus (White pine)
Bare-root trees
Populus grandidentata (Large-tooth aspen, big-tooth aspen)
Rooted cuttings
Prunus pumila (Sand cherry)
Plugs
Ptelea trifoliata (Wafer-ash, hop-tree)
Plugs
Quercus alba (White oak)
Bare-root trees
Quercus rubra (Red oak)
Bare-root trees/seedlings; acorns
Quercus velutina (Black oak)
Bare-root trees
Rudbeckia hirta (Black-eyed susan)
Plugs
Salix cordata (Sand-dune willow, furry willow)
Rooted cuttings
Schizachyrium scoparium (Little bluestem)
Seeds
Smilax rotundifolia (Common greenbrier)
Seeds
Solidago nemoralis (Gray goldenrod, old-field goldenrod)
Plugs
Sorghastrum nutans (Indian grass)
Seeds
Symphyotrichum oolentangiense (Sky-blue aster, prairie heart-leaved aster) Plugs
Tradescantia ohiensis (Common spiderwoit)
Plugs
Vernonia missurica (Missouri ironweed)
Seeds
Viburnum acerifolium (Maple-leaved viburnum)
Plugs
Vitis riparia (River-bank grape)
Plugs
Gentle slopes (4.3 acres)
Amelanchier arborea (Juneberry)
Plugs
Arnoglossum atriplicifolium (Pale Indian plantain)
Plugs; seeds
Artemisia campestris (Wild wormwood)
Plugs
Calamovilfa longifolia (Sand reed grass)
Seeds
Carex pensylvanica (Sedge)
Transplants
Cornus florida (Flowering dogwood)
Plugs
Elymus canadensis (Canada wild rye)
Seeds
Hamamelis virginiana (Witch-hazel)
Plugs
Lindera benzoin (Spicebush)
Plugs
Liriodendron tulipifera (Tulip tree)
Plugs
Lupinus perennis (Wild lupine)
Seeds
(Continued)
70
THE MICHIGAN BOTANIST
Vol. 51
APPENDIX I. Continued.
Microhabitat (estimated area)
Species (common name) Planting method
Gentle slopes (4.3 acres) (Continued)
Opuntia humifusa (Prickly-pear)
Panicum virgatum (Switch grass)
Populus deltoides (Cottonwood)
Primus pumila (Sand cherry)
Pteridium aquilinum (Bracken fern)
Quercus alba (White oak)
Quercus rubra (Red oak)
Schizachyrium scoparium (Little bluestem)
Smilax rotundifolia (Common greenbrier)
Symphyotrichum pilosum (Frost aster, hairy aster)
Tilia americana (Basswood, linden)
Tradescantia ohiensis (Common spiderwort)
Vaccinium angustifolium (Low sweet blueberry)
Viburnum acerifolium (Maple-leaved viburnum)
Plugs
Seeds
Rooted cuttings
Plugs
Transplants
Bare-root trees
Bare-root trees
Seeds
Plugs; seeds
Plugs
Plugs
Plugs; quart pots
Quart and gallon pots
Plugs
Prairie (1.3 acres)
Calamovilfa longifolia (Sand reed grass) Plugs
Coreopsis tripteris (Tall tickseed, tall coreopsis) Chaff/seeds
Elymus canadensis (Canada wild rye) Plugs
Koeleria macrantha (June grass) Plugs
Lithospermum caroliniense (Hairy puccoon, yellow puccoon, plains puccoon) Plugs
Lupinus perennis (Wild lupine) Seeds
Monarda fistulosa (Wild-bergamot) Seeds
Monarda punctata (Dotted mint, horse mint) Seeds
Panicum virgatum (Switch grass) Plugs
Rudbeckia hirta (Black-eyed susan) Chaff/seeds
Schizachyrium scoparium (Little bluestem) Plugs
Solidago nemoralis (Gray goldenrod, old-field goldenrod) Seeds
Symphyotrichum ericoides (Heath aster, wreath aster, white prairie aster) Seeds
Tradescantia ohiensis (Common spiderwort) Plugs
Vernonia missurica (Missouri ironweed) Plugs
Wetpanne (2 acres)
Asclepias incarnata (Swamp milkweed) Plugs
Bidens cernua (Nodding beggar-ticks) Seeds
Calamagrostis canadensis (Blue-joint) Plugs
Carex comosa (Sedge) Plugs
Carex cryptolepis (Sedge) Plugs
Carex flava (Sedge) Plugs
Carex hystericina (Sedge) Plugs
Carex lurida (Sedge) Plugs
Carex stricta (Sedge) Plugs
Carex suberecta (Sedge) Plugs
Cirsium muticum (Swamp thistle) Seeds
Cladium mariscoides (Twig-rush) Plugs
Coreopsis tripteris (Tall tickseed, tall coreopsis) Seeds
Eupatorium perfoliatum (Boneset) Plugs
Euthamia graminifolia (Grass-leaved goldenrod) Plugs
Hibiscus moscheutos (Swamp mallow, rose mallow) Plugs
Hypericum kalmianum (Kalm’s St. John’s-wort) Plugs
Iris virginica (Southern blue flag) Plugs
(Continued)
2012
THE MICHIGAN BOTANIST
71
APPENDIX I. Continued.
Microhabitat (estimated area)
Species (common name) Planting method
Wetpanne (2 acres) (Continued)
Juncus acuminatus (Sharp-fruited rush)
Juncus alpinoarticulatus (Rush)
Juncus balticus (Rush)
Juncus biflorus (Two-flowered rush)
Juncus brachycephalus (Rush)
Juncus effusus (Soft-stemmed rush)
Juncus marginatus (Grass-leaved rush)
Juncus torreyi (Torrey’s rush)
Lobelia cardinalis (Red lobelia, cardinal-flower)
Lobelia siphilitica (Great blue lobelia)
Lycopus americanus (Common water horehound)
Peltandra virginica (Arrow-arum, tuckahoe)
Pontederia cordata (Pickerel-weed)
Rudbeckia hirta (Black-eyed susan)
Sabatia angularis (Rose-pink; rose gentian)
Sagittaria latifolia (Wapato, duck-potato, common arrowhead)
Schoenoplectus acutus (Hardstem bulrush)
Schoenoplectus pun gens (Threesquare)
Schoenoplectus tabernaemontani (Softstem bulrush)
Scirpus cyperinus (Wool-grass)
Thelypteris palustris (Marsh fern)
Verbena hastata (Blue vervain)
Vemonia missurica (Missouri ironweed)
Wetland shelf (3.2 acres)
Alisma sp. (Water-plantain)
Aronia prunifolia (Chokeberry)
Calamagrostis canadensis (Blue-joint)
Carex hystericina (Sedge)
Carex lupulina (Sedge)
Carex lurida (Sedge)
Carex stricta (Sedge)
Cephalanthus occidentalis (Buttonbush)
Cladium mariscoides (Twig-rush)
Cornus amomum (Pale dogwood, silky dogwood)
Cornus sericea (Red-osier)
Decodon verticillatus (Whorled loosestrife, swamp loosestrife)
Ilex verticillata (Michigan holly, winterberry, black-alder)
Iris virginica (Southern blue flag)
Juncus balticus (Rush)
Juncus effusus (Soft-stemmed rush)
Lindera benzoin (Spicebush)
Lobelia cardinalis (Red lobelia, cardinal-flower)
Nuphar advena (Yellow pond-lily)
Nymphaea odorata (Sweet-scented waterlily)
Peltandra virginica (Arrow-arum, tuckahoe)
Pontederia cordata (Pickerel-weed)
Physocarpus opulifolius (Ninebark)
Rosa palustris (Swamp rose)
Sagittaria latifolia (Wapato, duck-potato, common arrowhead)
Salix exigua (Sandbar willow)
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs; seeds
Plugs
Seedlings
Plugs
Seeds
Plugs
Plugs
Plugs
Seeds
Plugs; seeds
Transplants
Plugs
Plugs
Plugs; tubers
Plugs; seeds
Plugs
Plugs
Seeds
Plugs
Plugs
Plugs; seeds
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Plugs
Tubers
Tubers
Plugs
Plugs
Seeds
Plugs
Plugs
Rooted cuttings
Continued
72
THE MICHIGAN BOTANIST
Vol. 51
APPENDIX I. Continued.
Microhabitat (estimated area)
Species (common name)
Planting method
Wetland shelf (3.2 acres) (Continued)
Schoenoplectus pungens (Threesquare)
Plugs
Schoenoplectus tabernciemontani (Softstem bulrush)
Plugs
Scirpus atrovirens (Bulrush)
Plugs
Sparganium americanum (American bur-reed)
Seeds
Spiraea alba (Meadowsweet)
Plugs
Shallow water (1.9 acres)
Brasenia schreberi (Water-shield)
Transplants
Nuphar advena (Yellow pond-lily)
Tubers
Nymphaea odorata (Sweet-scented waterlily)
Tubers
Potamogeton amplifolius (Large-leaved pondweed)
Transplants
Potamogeton gramineus (Pondweed)
Transplants
Potamogeton nodosus (Pondweed)
Transplants
Schoenoplectus tabernaemontani (Softstem bulrush)
Plugs
Scirpus cyperinus (Wool-grass)
Plugs
Deep water (3.9 acres)
Potamogeton amplifolius (Large-leaved pondweed)
Transplants
Potamogeton gramineus (Pondweed)
Transplants
Potamogeton zosteriformis (Flat-stemmed pondweed)
Transplants
Stuckenia pectinata (Sago pondweed)
Transplants
Vallisneria americana (Tape-grass, wild-celery, eel-grass)
Transplants
2012
THE MICHIGAN BOTANIST
73
THE SUGAR MAPLE SAPSTREAK FUNGUS
0 CERATOCYSTIS VIRESCENS (Davidson) MOREAU,
ASCOMYCOTA) IN THE HURON MOUNTAINS,
MARQUETTE COUNTY, MICHIGAN
Dana L. Richter
School of Forest Resources & Environmental Science
Michigan Technological University
1400 Townsend Drive
Houghton, Michigan 4993 1
Phone: 906-487-2149
Email: dlrichte@mtu.edu
ABSTRACT
Sugar maple sapstreak disease is caused by the native fungus Ceratocystis virescens (Davidson)
Moreau, but is only a serious threat in disturbed areas or where soil conditions favor the disease.
Three of 16 trees along roads, in logged areas or other disturbance that showed crown symptoms of
sugar maple sapstreak disease in the Huron Mountains were confirmed or probable for the disease
based on xylem condition and laboratory isolation of the fungus. Trees diagnosed with the disease
had greater than 50% crown dieback, while all trees free of the disease had lesser degrees of crown
dieback. Although the presence of sugar maple sapstreak disease was confirmed in the Huron Moun¬
tains the incidence was found to be low. Results suggest this pathogen exists at low levels in the
Huron Mountain forests and is not an imminent threat to sugar maple there. Soil conditions and other
factors contributing to a generally healthy forest may be responsible for low incidence and spread of
the disease.
KEYWORDS: sugar maple, sapstreak disease, Huron Mountains, Ceratocystis virescens, fungi
INTRODUCTION
Maple sapstreak is a disease of sugar maple trees {Acer saccharum Marsh.)
throughout the eastern United States and Canada caused by the fungus Cerato¬
cystis virescens (Davidson) Moreau (Houston and Fisher 1964, Houston 1993,
Houston 1994). Previous names used for this fungus are Endoconidophora
virescens and C. coerulescens\ however, the latter species is now considered re¬
stricted to conifers (Witthuhn et al. 1998), while the former is the anamorph
(asexual stage) of C. virescens. A current listing of species within the genus Cer¬
atocystis includes 214 named taxa (CABI 2011). The taxonomic placement of C.
virescens is as follows: Family: Ceratocystidaceae; Order: Microascales; Class:
Sordariomycetes; Phylum: Ascomycota; Kingdom: Fungi (EPPO/CABI 2003).
Ceratocystis virescens is a xylem inhabitant of sugar maple, although it has
been reported in several other woody species in living trees and on cut logs
(Roth, et al. 1959, Shigo 1962). The fungus is considered among the most com¬
mon fungi in northern hardwoods (Shigo 1962). Early reports of the occurrence
of the fungus, however, must be taken with caution due to confusion in taxon¬
omy and difficulty in identification. For example, a monograph of the genus
74
THE MICHIGAN BOTANIST
Vol. 51
(Upadhyay 1981) does not separate C. coerulescens from C. virescens. These
fungi do not decay wood per se, nor do they produce large fruiting bodies.
Ceratocystis virescens is a native fungus related to the pathogens that cause
oak wilt (C. fagacearum (Bretz) Hunt, a native species), Dutch elm disease
0 Ophiostoma ulrni Buisman (formerly C. ulmi), an introduced species), and blue
stain defect in conifer and hardwood lumber (several native species of Cerato¬
cystis and/or their anamorphs). Native pathogens are generally present at low
levels in all forest ecosystems. Although once considered a potential threat to
sugar maple forests (Kessler 1978), sapstreak disease is rare in healthy stands of
sugar maple but occurs mainly where trees are stressed typically by root com¬
paction, logging or some other sort of human disturbance (Houston 1993, Hous¬
ton 1994). Water-logged soils prone to rutting may also contribute to spread of
the disease (Houston 1994).
The fungus infects trees through wounds, especially logging wounds, or dam¬
age along roads and in maple syrup operations (Hepting 1944, Mielke and
Charette 1989, Houston 1993, 1994). It is suspected the disease is vectored by in¬
sects, however, little is known about its spread and the exact species of insects
have not been identified (Kile 1993). It is likely that generalists such as sap bee¬
tles and fruit flies are involved in the spread of the disease. Symptoms of the dis¬
ease include wilting, yellowing and dwarfing of leaves, and dieback of branches
in the crown, leading to decline and eventual death of the tree, often within one
to two years (Mielke and Charette 1989, Houston 1993, 1994). Further symp¬
toms occur in the wood, where dark reddish-brown to blue-green streaking oc¬
curs (Figure 1); staining of wood is seen in wood at the base of the trunk in the
root-flare region of the tree (Houston 1993). This symptom makes the disease
identifiable in the field when bark from root flares is removed with a hatchet or
chisel. In the laboratory, dark hyphae present in xylem vessel and parenchyma
cells can be observed microscopically. The fungus can also reside in downed
logs and stumps, and was isolated in 2008 in the Huron Mountains from a small,
dead sugar maple tree along the road to Ives Lake (Richter 2008). In August
20 1 0, a further examination of 1 6 suspect sugar maple trees and one stump bear¬
ing living sprouts was conducted to assess the incidence of sugar maple sap-
streak disease in the Huron Mountains.
METHODS
The forests of the Huron Mountains in northern Marquette Co., Michigan are owned principally
by the private Huron Mountain Club (approximately 20,000 acres (8,094 ha)), and are used for recre¬
ation, research and a small amount of forest management (Figure 2). The forests are a mixture of
hemlock-northern hardwood, pine (red, white, jack) and wetlands (Simpson, et al. 1990). Small
parcels within the Huron Mountains area are owned by private individuals for recreation homes. The
other major landowner in the area is a commercial logging and real estate firm. Considerable logging
operations are conducted on lands bordering the Huron Mountain Club lands.
Sixteen sugar maple trees with varying degrees of crown dieback or other indication of stress
along roads or other forms of disturbance in the Huron Mountains were sampled Aug 20-22, 2010;
fifteen were standing stressed and declining trees, and one was a recently fallen tree. Additionally, a
stump of a tree cut three years before containing live wood and sprouts was also sampled. Table 1
gives the location, diameter, amount of crown dieback and xylem condition of the trees sampled.
2012
THE MICHIGAN BOTANIST
75
FIGURE 1. Thirty cm diameter section of a freshly cut sugar maple stem in Houghton Co., Michi¬
gan showing dark stain caused by infection by Ceratocystis virescens, the pathogen of sapstreak of
maple.
FIGURE 2. Location of the Huron Mountains in northern Marquette Co. in the Upper Peninsula of
Michigan.
76
THE MICHIGAN BOTANIST
Vol. 51
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2012
THE MICHIGAN BOTANIST
77
Sampling consisted of removing 2-3 chips of xylem per tree with a hatchet directly under the bark
of a root-flare at the base of a tree; chips were approximately 8x5 cm, and 1-2 cm thick.
Samples from four trees with dark streaking in the xylem characteristic of sapstreak disease were
cultured in the laboratory. Isolations were made on 2% malt-agar in 60 mm Petri dishes; trunk sam¬
ples were first surface sterilized with 10% chlorine bleach; per tree, nine pieces (approx. 2 x 2 x 0.3
mm) of stained inner xylem were excised using a sterile scalpel and placed in three Petri dishes
(three pieces per plate). Plates were grown at 22-24°C for 7-10 days and examined for fungus
growth. Fungus structures of C. virescens were examined at 400x with a Nikon Optiphot® micro¬
scope. At least ten fruiting structures and spores were measured to determine size range and confirm
identity of C. virescens according to Davidson (1944).
RESULTS
The sugar maple trees sampled ranged in diameter from 20-90 in diameter;
crown dieback ranged from 10-80%. Only four of the 17 trees sampled exhib¬
ited dark staining in the xylem characteristic of sugar maple sapstreak disease
(Table 1). Of these four trees, two (#6 and #17) were determined to be probable
infections based on color of xylem alone since C. virescens was not isolated;
other fungi that were isolated likely out-competed C. virescens in culture, #6 by
Trichoderma sp. (a mold fungus), and #17 by two decay fungi (unidentified).
One tree (#7) was confirmed infected with the sapstreak fungus by laboratory
isolation of C. virescens ; and one tree (#12, stump with living wood and sprouts)
was unlikely infected with sapstreak, the coloration in this case due to a dark
decay fungus which was isolated instead (tentative identification, Xylaria sp.).
The culture of C. virescens obtained from sugar maple tree #7 (DR-492) was
confirmed by examining growth on agar, hyphal characters, fruiting structures
and spores, and compared to the description provided by Davidson (1944). Dark
black mycelium covered the agar surface in a 90 mm petri plate in 10 days; the
culture had a sweet, musty odor. Abundant fruiting bodies (perithecia) were pro¬
duced which were approximately 0.6-0. 8 mm upright in length, with bases
30-50 pm diameter and tips 12-15 pm diameter (Figure 3); ascospores collected
at tip of asci were hyaline, slightly curved, 6-7 x 2 pm; conidiospores formed di¬
rectly from hyphae were hyaline, cylindrical and highly variable in size approx¬
imately 6-25 x 3-6 pm.
The three trees probable and confirmed (#6, #7, #17) for sapstreak disease
had from 50-80% crown dieback (Figure 4); two other trees with 50% crown
dieback had clear xylem and no evidence of sapstreak; all other trees with less
than 50% crown dieback also showed no evidence of sapstreak in the xylem.
Two of the trees showing sapstreak disease (#6 probable, #7 confirmed) were on
the edge of an excavation for gravel; the other probable sapstreak tree (#17) was
along the road to Mountain Lake. Serious root damage caused by excavation or
road compaction is the likely factor in these cases of sapstreak disease.
DISCUSSION
Although the presence of sapstreak disease of sugar maple was confirmed in
the Huron Mountains, the incidence was found to be low, and relatively few trees
78
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 3. Ascocarps (perithecia) of the sugar maple sapstreak fungus Ceratocystis virescens
(Davidson) Moreau; globose bases approx. 50 pm diameter; perithecial necks 0.6-0. 8 mm long.
with symptoms of crown dieback were infected with C. virescens as indicated by
xylem staining or laboratory isolation of the pathogen. The trees sampled do not
represent the sugar maple forest as a whole in the Huron Mountains, but were se¬
lected based on having some form of disturbance and susceptibility to the dis¬
ease. The disease was confirmed only in trees which had undergone major dis¬
turbance to the roots along a road and the edge of a gravel pit; these trees had
50-80% crown dieback. Results of the incidence of sapstreak disease may be dif¬
ferent if wounded trees were exclusively sampled; however, the single tree sam¬
pled having a logging wound (#4) did not have stained xylem, while the stump
with living tissue and stained xylem (#12) did not yield C. virescens by labora¬
tory culture; staining in the latter was likely due to a dark-colored decay fungus.
Thirteen of the 16 stressed sugar maple trees in the Huron Mountains that
exhibited 10-50% crown dieback, and one stump with living sprouts were ap¬
parently free from sapstreak disease; this implies that something other than sap¬
streak disease was affecting tree crown condition resulting in dieback. The
cause of the dieback in these trees is likely physiological stress due to root com¬
paction caused by roads and excavation. That these trees were not subsequently
infected by the sapstreak fungus suggests that this native pathogen exists at low
levels in the Huron Mountains. Factors contributing to a generally healthy for¬
est may be responsible for the low incidence and spread of the disease. The low
2012
THE MICHIGAN BOTANIST
79
FIGURE 4. Stressed sugar maple trees #6 and #7 on the edge of a gravel pit in the Huron Mountains
with 80% crown dieback (center and right); these trees contained stained xylem, evidence of sap-
streak disease; Ceratocystis virescens was isolated from tree on right. (Photo taken 21 Aug 2010)
80
THE MICHIGAN BOTANIST
Vol. 51
incidence of the disease might also be due to the vast extent of old growth and
the limited amount of logging that has been conducted in the forests of the
Huron Mountains.
ACKNOWLEDGEMENTS
This study was made possible with funds provided by the Huron Mountain
Wildlife Foundation (www.hmwf.org), and the School of Forest Resources and
Environmental Science, Michigan Technological University, Houghton, MI
(www.mtu.edu/forest). Ms. Tara L. Bal, School of Forest Resources and Envi¬
ronmental Science, provided valuable comments on the ecology of the sapstreak
fungus in sugar maple forests in the Upper Peninsula. This paper is dedicated to
Dr. William R. Manierre (1923-2011), meticulous student of the mosses and
lichens, who worked tirelessly to protect and deepen the understanding of the
ecology of the forests of the Huron Mountains; friend to many and all of nature.
LITERATURE CITED
Davidson, R. W. ( 1944). Two American hardwood species of Endoconidiophora described as new.
Mycologia 36: 300-306.
European Plant Protection Organization / Commenwealth Agricultural Bureau International. (2003).
Ceratocystis virescens. In: Data sheets for quarantine pests for Europe. (Ed. by Smith, I.M.) CAB
INTERNATIONAL, Wallingford, UK.
Hepting, G. H. (1944). Sapstreak, a new killing disease of sugar maple. Phytopathology 34:
1069-1076.
Houston, D. R. (1992). Importance of buttress root and taphole wounds as infection courts for the
sugar maple (Acer saccharum) sapstreak pathogen, Ceratocystis coerulescens. Phytopathology 82:
244.
Houston, D. R. (1993). Recognizing and managing sapstreak disease of sugar maple. Research Paper
NE-675. Radnor, PA: USDA, Forest Service, Northeastern Forest Experiment Station, 11pp.
Houston, D. R. (1994). Sapstreak disease of sugar maple: development over time and space. Re¬
search Paper NE-687. Radnor, PA: USDA, Forest Service, Northeastern Forest Experiment Sta¬
tion, 19pp.
Houston, D. R. and K. D. Fisher. (1964). Sapstreak of sugar maple found in the northeast. Plant Dis¬
ease Reporter 48:788.
Commenwealth Agricultural Bureau International. (2011). Index Fungorum. http://www.
indexfungorum.org/Index.htm
Kessler, K. J., Jr. (1978). How to control sapstreak disease of sugar maple. North Central Forest Ex¬
periment Station, Forest Service, US Department of Agriculture, St. Paul, MN. (pamphlet).
Kile, G. A. (1993). Plant diseases caused by species of Ceratocystis sensu stricto and Chalara. pp
173-184. In: Ceratocystis and Ophiostoma: Taxonomy, ecology and pathogenicity. Ed. by Wing¬
field, M.J. Seifert, K.A. and Webber, J.F. APS Press, St. Paul, MN. 293pp.
Mielke, M. E. and D. A. Charette. (1989). The incidence of sapstreak disease of sugar maple in
Menominee County, Wisconsin, and its relationship to wounds and season of logging. Northern
Journal of Applied Forestry 6: 65-67.
Richter, D. L. (2008). Continuing studies of fungi of the Huron Mountains: Xylem-inhabiting mi¬
crofungi. 2008 Field Season Report to the Huron Mountain Wildlife Foundation. 10 December
2008. 3pp.
Roth, E., G. H. Hepting, and E. R. Toole. (1959). Sapstreak disease of sugar maple and yellow poplar
in North Carolina. Phytopathology 49: 549.
Shigo, A. L. (1962). Observations on the succession of fungi on hardwood pulpwood bolts. Plant
Disease Reporter 46: 379-380.
Simpson, T. B., P. E. Stuart, and B. V. Barnes. (1990). Landscape ecosystem and cover types of the
2012
THE MICHIGAN BOTANIST
81
Reserve Area and adjacent lands of the Huron Mountain Club. Occasional Papers of the Huron
Mountain Wildlife Foundation. No. 4, Big Bay, MI 128pp.
Upadhyay, H. P. (1981). A monograph of Ceratocystis and Ceratocystiopsis. Univ. Georgia Press,
Athens, GA. 176 pp.
Witthuhn, R. C„ B. D. Wingfield, M.J. Wingfield, M. Wolfaardt and T. C. Harrington. (1998). Mono-
phyly of the conifer species in the Ceratocystis coerulescens complex based on DNA sequence
data. Mycologia 90 (1):96— 101 .
82
THE MICHIGAN BOTANIST
Vol. 51
NOTEWORTHY COLLECTION
MICHIGAN
Asplenium rhizophyllum L. Walking Fern.
Previous Knowledge. Asplenium rhizophyllum L. (synonym: Camptosorus
rhizophyllus L.) is currently listed as a threatened species in Michigan (Penskar
& Higman 1997) and has a global ranking of G5 S2/S3 indicating it is secure
through most of its range but in some regions it is considered critically imperiled
(Natureserve 2011). Limited habitat may account for its rarity in Michigan. As¬
plenium rhizophyllum is generally found on mossy boulders or outcrops com¬
posed of calcareous rocks, particularly limestone and dolomite, or rarely, on
sandstone (Cobb, 2005). It is most abundant in the eastern Upper Peninsula on
the dolomite and limestone outcrops of the Niagara Escarpment (Penskar & Hig¬
man, 1997). Seven Michigan counties have collection records: Alpena, Berrien,
Chippewa, Delta, Dickinson, Mackinac and Leelanau (on Manitou Islands only,
where it is found on mossy cedar logs in wooded dunes), (Michigan Flora On¬
line 2011) but it has also been observed in Houghton Co. (Penskar & Higman,
1997). It is also found in the border county of Florence, Wisconsin, where it is
given no special status (USDA 2011).
Significance of the Report. The colony was first observed in 2006, although
no collection was made. This is the first collection for Alger County and the only
collection from the Early Ordovician Au Train Formation, a light brown to white
dolomitic sandstone (National Park Service, 2012). Though our meander survey
of the area detected only two isolated colonies (Fig 1), more potential habitat ex¬
ists. The localized occurrence of this species in Michigan creates unique chal¬
lenges to the conservation of these vulnerable populations. It is important to doc¬
ument these occurrences in order to understand the full extent of the species
range in Michigan as well as its ecological requirements.
Diagnostic Characters. Asplenium rhizophyllum roots from the tips of its
arching fronds, hence its common name “walking fern” (Fig. 2). Another rare
Asplenium in Michigan is A.scolopendrium, which is only known from
Chippewa and Mackinac Counties. Both species occur in similar habitats and
have entire fronds. Several distinctions can be made between these species. The
most obvious field distinction is the tip rooting that occurs in A.rhizphyllum, but
not A.scolopendrium. Also, in A. rhizophyllum the frond apex tapers into a grad¬
ual point whereas in A. scolopendrium the frond apex tapers abruptly to a point,
or is more or less rounded (Michigan Flora, 2012).
Specimen citation. Alger Co. Michigan; Hiawatha National Forest. Two
colonies of no more than one square meter in size were growing on sandstone
boulders approximately four meters apart. The specimen was collected in a hard¬
wood forest. Also growing on the boulders hosting A. rhizophyllum were Dicen¬
tra americana and Sambucus racemosa. Associated species in the immediate
area included Acer saccharum, Abies balsamea, Picea glauca, Tilia americana,
2012
THE MICHIGAN BOTANIST
83
FIGURE 1. Collection area and habitat of A. rhizophyllum. Photo taken by Lauren Tarr, 2012.
FIGURE 2. Asplenium rhizophyllum. Illustrated by Susan Fawcett.
84
THE MICHIGAN BOTANIST
Vol. 51
Ostrya virginiana, Sambucus racemosa, Carex communis, Carex leptonervia,
Oryzopsis aspirifolia, Allium tricoccum, Claytonia caroliniana, Dicentra sp.,
Erythronium americanum, Galium sp., Hepatica americana, Maianthemum
canadense, Osmorhiza sp., Taraxacum officinale, Trillium grandiflorum, Viola
sp., Cystopteris fragilis, Dryopteris carthusiana, Polystichum lonchitis and
Polypodium virginianum. Specimens were collected by Susan Fawcett and
Michael Rotter on April 12th 2012 and deposited at NM (Rotter 528). Duplicates
were deposited at UM, and the Hiawatha National Forest. The collection was
made under a Forest Products Free Use Permit from the US Department of Agri¬
culture-Forest Service.
ACKNOWLEDGMENTS
We would like to thank Steve Caird and Lauren Tarr for field support. Thanks to Deb Le Blanc
and Dr. Tony Reznicek for collection and permitting advice and Dr. John Anderton for information
on the geology of the area.
LITERATURE CITED
Cobb, B., Lowe, C., and Farnsworth, E. 2005. Peterson Field Guide to Ferns: Northeastern and Cen¬
tral North America 2nd edition, Houghton, Mifflin Harcourt. New York.
Michigan Flora Online. Reznicek, A. A., Voss, E. G., and Walters B.S. 2011. University of Michigan
< http://michiganflora.net/species.aspx?id=201 accessed 12.29.2011
National Park Service, US Department of Interior. Pictured Rocks National Lakeshore. 2012.
www.nps.gov/piro/naturescience/geologicformations.htm accessed 4. 19.20 1 2
NatureServe. 2011. Nature Serve Explorer, Nature Serve Conservation Status. July 2011.
http://www.natureserve.org/explorer/ranking.htm accessed 12.29.201 1
Penskar, M.R. and Higman, P.J. 1997 Special Plant Abstract for Asplenium rhizophyllum (walking
fern). Michigan Natural Features Inventory, Lansing MI.
USDA, NRCS. 2011. The PLANTS database .National Plant Data Team NRCS. < http://
plants.usda.gov/ > accessed 12.29.2011
— Michael Rotter, Susan Fawcett, and Danny McConnell
Northern Michigan University
Northern Michigan Herbarium
1401 Presque Isle Ave
Marquette MI 49855 USA
2012
THE MICHIGAN BOTANIST
85
BOOK REVIEWS
Pringle, Peter. 2008. The Murder of Nikolai Vavilov: The Story of Stalin ’s Persecution
of One of the Great Scientists of the Twentieth Century. New York. Simon & Schuster.
370 pp. ISBN-13: 978-0-7432-6498-3. $26.00. Hardbound.
In this captivating biography of the Russian plant geneticist/botanist Nikolai Vavilov,
Pringle tells the story of how one of the twentieth centuries greatest botanists who
dreamed of feeding the world died of starvation. Though he worked for numerous years
for the Russian state, Vavilov was persecuted, incarcerated, interrogated and very likely
tortured by Stalin’s communist regime for his unwavering confidence in Mendelian ge¬
netics.
Vavilov, as portrayed by Pringle, was a passionate man full of energy, insatiable cu¬
riosity and unrelenting drive. He was first and foremost a scientist, though Pringle also
devotes time to discussing Vavilov’s family life. And though many of the charges brought
against Vavilov by the KGB were of subterfuge and disloyalty to the motherland, Pringle
works throughout the biography to show Vavilov as the incomparable patriot he is re¬
vealed to be in numerous personal journals, letters, interviews and records. Vavilov even
tried to convince American contemporaries working at the time with T. H. Morgan to
come to Russia and join him in his botanical cross breeding experiments. Of those who
accepted Vavilov’s invitation, only a few lived through the closing of Stalin’s fist around
scientists he deemed to be too theoretical.
Fueled in part by the common suspicion of the time that academics, particularly those
well born, did not have the interests of the common people at heart, the U.S.S.R eventu¬
ally stalled biological advancement in the motherland by fully backing the work of the
uneducated would-be scientist, Forim Fysenko. Ironically, Vavilov played a key part in
the recognition of Fysenko’s work on vernalization. For many years, Fysenko worked on
one of Vavilov’s experimental breeding stations. Throughout Fysenko’s career, Vavilov
repeatedly extended the inexperienced Fysenko invitations to join him at international
conferences on genetics and worked to give him opportunities to expand his knowledge
base and improve as a scientist. However, Lysenko preferred a Lamarckian theory of ge¬
netics and would not be moved, a preference ultimately shared by Stalin himself. It is not
surprising, therefore, after the forth of multiple famines hit Russia throughout the early
1900’s, Stalin turned to Lysenko instead of Vavilov for a plan to rapidly improve Russian
agriculture. Interestingly, the famines were caused in part by environmental factors, but
greatly exacerbated by Stalin’s grain collectivization program.
Lysenko, a formidable opportunist, worked at every turn to amass his own power and
undermine Vavilov’s own position. Pringle discusses Vavilov’s reluctance to engage Ly¬
senko in open conflict and confront his erroneous theory of inheritance and bad science.
By the time Vavilov grasped the gravity of the situation, it was too late. It is interesting that
a man so perceptive with plants was so blind to the darker side of people. Even throughout
the times of persecution, Pringle paints a picture of a painfully optimistic Vavilov. He con¬
stantly wrote encouragement to his experimental institutes around Russia, maintained con¬
tact with his international colleagues and believed that the truth of science would triumph.
None-the-less, Pringle writes, a different truth eventually dawned on the horizon: Vavilov
would be arrested. In an address to his workers at the Leningrad Institute, he stated, “We
will go into the pyre, we shall bum, but we shall not retreat from our convictions.” Ar-
86
THE MICHIGAN BOTANIST
Vol. 51
rested August 6, 1940 on the fictitious charges of sabotage of Soviet Agriculture and spy¬
ing for foreign countries, particularly Britain, Vavilov was brutally interrogated. Initially
sentenced to be shot on July 9, 1941, his sentence was later commuted to twenty years in
labor camps. Neither sentence carried; Vavilov was found dead on January 26, 1943. The
jail doctors pronounced the cause of death to be dystrophy from prolonged malnutrition.
Despite his unnecessarily early death, Vavilov made phenomenal advances in Russian
agriculture. He worked tirelessly to keep himself and his workers abreast of and contribut¬
ing to international agrarian developments. On each of his multiple and varied excursions
to locate “hot spots” of plant biodiversity and species origins, He would bring back fruits,
vegetables, and hoards of seeds and grains common to other continents but not yet seen in
Russia. Each specimen was saved in his ever-expanding seed library and many were di¬
rectly used in agricultural experiments at field stations throughout Russia. His now famous
and nearly globally inclusive collection of seeds, arguably one the first gene seed banks in
the world, even managed to survive World War II. Pringle writes of workers in the Insti¬
tute of Leningrad starving to death in their desks as they maintained vigil over the collec¬
tion. Vavilov, as Pringle reveals, was indeed a man of contagious vision.
In constantly searching for new varieties of existing crops, he dreamed of “directing
the evolution of cultivated plants and domestic animals according to our will,” seeing his
mission as one for all humanity. Although the director of multiple institutes over the span
of his life, Vavilov never sought administrative promotion, preferring instead to be in the
laboratory or the field performing genetic experiments on his preciously obtained vari¬
eties. None-the-less, his formidable education from the Petrovskaya Agricultural Insti¬
tute, post-graduate studies and internships in Russia, Britain, France and Germany, and
publically recognizable genius at one point landed him eighteen administrative posts. He
served a decade as the director of the Institute of Genetics of the Academy of Sciences of
the USSR, during which time period he also functioned as the vice president of the 6th In¬
ternational Congress of Genetics in Ithaca, New York. In the time since Vavilov’s death,
Russia has recanted. Vavilov was rehabilitated in 1955. Thirty-three years later the coun¬
try celebrated the centenary of his birth. Russian citizens today proudly recognize Vavilov
as one of their foremost scientists. The Russian Government posthumously granted Vav¬
ilov an honorary doctorate and presented it to his second son, Yuri, on November 27,
2007 at a special symposium honoring the 120th anniversary of Vavilov’s birth at the
Russian State Agrarian University in Moscow.
As Pringle mentions, it is starkly ironic that a man who dreamt of feeding the world
would die of starvation. One cannot help but wonder how the world would be different
had Vavilov lived. He carried out specimen collecting tours of extensive breadth on five
continents to better understand and hopefully harness the genetic variability globally pre¬
sent in wild and cultivated plants. Over his lifetime, Vavilov created a botanical library of
over 250,000 specimens and published prolifically. Yet even today, his dream of feeding
the world remains unrealized.
- Brianna Payne, Graduate Student, Biology Department
Andrews University, Berrien Springs, MI 49104-0410
brianna@andrews.edu
Quammen, D. 2008. On the Origin of Species: The Illustrated Edition. Sterling Pub¬
lishing Company, 387 Park Avenue South, New York, NY 100016, 544 pages. ISBN
1402756399 $35.00.
Charles Darwin is arguably the most well known scientist who has ever lived. The views
he developed on his fateful trip around the world would change the face of biology for-
2012
THE MICHIGAN BOTANIST
87
ever. He spent years after his voyage collecting evidence until he was forced to publish
his findings more than 23 years after his return. His book, titled On the Origin of Species
by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for
Life was in instant success and was followed by many further editions.
The interest in Darwin and his work continues to this day. Many who are interested in
biology have read at least part of The Origin of Species. However, this edition is different
than many. In contrast to the many small paperback versions available, this edition is
large (9x10 inches) and hardbound with a nicely illustrated dust jacket. The resulting de¬
crease in portability is more than made up for by the beautiful illustrations on almost
every page. They portray the diversity Darwin saw, the species he observed and snapshots
of his life. Some of the illustrations were made on the voyage of the Beagle, while others
are from Darwin’s later publication The Zoology of the Voyage of H.M.S. Beagle. In ad¬
dition to the illustrations, many related snippets were included from Darwin’s voyage
diary as well as other publications on his life. Although the text was taken directly from
Darwin’s Origin of Species, the liberal sprinkling of additional text and pictures greatly
enhances the reading experience.
Anyone interested in biology will find this book fascinating, whether you have previ¬
ously read The Origin of Species or not. The illustrations and glimpses into Darwin’s life
add further depth and meaning to his exploration of natural selection and the processes of
change. This book would be an excellent addition to any biological library.
- Jonathan Cowles, Graduate Student
Biology Department, Andrews University
Berrien Springs, MI 49104-0410
E-mail: jonathan.cowles@gmail.com
Theodoropoulos, D. I. 2003. Invasion Biology: A Critique of a Pseudoscience. Avvar
Books, Blythe, CA 92225. 236 pp. ISBN 0-9708504-1-7, paperback $14.50.
Our increasing awareness of just how dramatically humans can alter ecological systems
has spawned new subdisciplines of biology. I doubt that any of us today would, if given
a second chance, exterminate the passenger pigeon ( Ectopistes migratorius ): the idea of
conservation biology is too prominent in our minds. We wish to preserve the ecosystems
we observe around us. Invasion biology has sprung from that same mindset: we’ve seen
the damage we inflict when we transport species from a distant geographic locality and
introduce them to our native flora and fauna. Examples in our area include the house spar¬
row ( Passer domesticus ), brought from Europe and now competing with our beloved
eastern bluebirds ( Sialia sialis ), reedgrass ( Phragmites australis subsp. australis ) origi¬
nally from Europe and now abundant in our marshes, and the zebra mussel ( Dreissena
polymorpha ), which has become extremely prevalent in the Great Lakes.
But David Theodoropoulos has a different idea. In his controversial book Invasion Bi¬
ology , he argues that “invasive species” are actually not a threat to native wildlife; instead
of being harmful, “alien” species typically integrate quickly into their new ecosystem, in¬
duce natives to evolve more quickly, and increase diversity in the areas into which they
spread.
The first section, which is roughly half of the book, deals with the science behind this
issue. In rapid-fire succession Theodoropoulos cites many examples of species which in¬
vasion biologists claim are demonstrably harmful to native wildlife, but which he claims
are completely innocuous. “Non-natives” aren’t the problem, he says; anthropogenic de¬
struction of ecosystems is — and in fact, non-natives typically show up in areas disturbed
88
THE MICHIGAN BOTANIST
Vol. 51
by humans. Dispersal of exotics is nothing to be afraid of, he reiterates— it happens natu¬
rally all the time.
Theodoropoulos devotes the second half of his book to a discussion of the psychology
of invasion biology. He does not stop at refuting the scientific claims of “anti-alien” sci¬
entists: invasion biology is a pseudoscience, he says, and appears to be bom of some basic
human xenophobia. In quick succession the author lists several analogous principles or
behaviors in both invasion biology and Nazism. Theodoropoulos attributes paranoia, pro¬
paganda, a money connection with pesticide companies, and a hateful attitude towards
anything “foreign” to invasion biologists.
In the last and smallest section of Invasion Biology the author brings his arguments to
a conclusion, and directs the reader to what he thinks is the real crisis: bias against “alien”
species will hamper our ability to protect the planet’s biodiversity. If a particular plant is
rare, feel free to cultivate it in your backyard or greenhouse! says Theodoropoulos. By
keeping as much diversity as possible spread throughout the planet, species are much less
likely to go completely extinct. He cites several examples where the only members of a
species left in existence are in a zoological park or cultivation — Gladiolus citrinus,
Franklinia alatahama, and Tecophilia cyanocrocus for example. There are no theoretical
limits to diversity, he argues, so we should try to pack as many species as possible in ap¬
propriate environments; it doesn’t matter whether they are “native” or “alien”. Further¬
more, the language we use to describe exotics — aggressive, invasive, alien, biological
pollution — should be changed in order to eliminate bias and reflect a new understanding
of the role of “non-natives”. Anthropogenic dispersal is a critical duty not to be neglected,
not a matter for unscientific fear or regulation by bureaucracy.
This reviewer feels that Theodoropoulos’ argument would have been much better
made and received had he simply omitted the second section of his book, and left that can
of worms to open at a later date if he so desired. I am not convinced that yanking the gar¬
lic mustard (Alliaria petiolata) from my woods is tantamount to Ku Klux Klan activities.
That said, I found this book to be highly intriguing. Theodoropoulos raises questions wor¬
thy of our attention: What exactly does “invasive” mean? How exactly does anthro¬
pogenic disturbance affect ecosystems? Are there some pros to having certain exotic
species around? Have we overreacted and invoked too much red tape? Are the recent
studies indicating that zebra mussels may actually be improving water quality of our
Great Lakes correct? Will the house sparrows disappear and leave the bluebirds to live
their lives, if the agricultural disturbance ceases? How exactly do we carry out our role as
stewards on this solitarily verdant planet?
- Libby Megna
Department of Biology
Andrews University
Berrien Springs, MI 49104
megna@andrews.edu
INSTRUCTIONS TO AUTHORS
1. Create text in 12-point Times New Roman font and double space paragraphs throughout. Papers
should be organized as follows: Title, Author(s) and address(es), Abstract with up to 5 keywords.
Introduction, Materials and Methods, Results, Discussion, Acknowledgements, Literature Cited,
Tables, Figure Legends, and Figures. Sections may be omitted if not relevant. All pages should
be numbered. Please contact the editor regarding any questions related to formatting.
2. For noteworthy collections, manuscripts should be formatted as described in The Michigan
Botanist, volume 27(3) p. 90. A brief description of the formatting follows. The following title,
“Noteworthy collections”, should begin each submitted manuscript followed on the next line by
the State or Province for the species reported. The next line should list the taxon of interest using
the following format: Species Author(s) (Family). Common name. The rest of the manuscript
should include the following named sections: Previous knowledge, Significance of the report,
Diagnostic characters (if desired), Specimen citations, and Literature cited. Each of these sec¬
tions are largely self explanatory; however, “specimen citations” should include the relevant
label data from the voucher specimen(s) including location data, collector(s), collection number,
etc. Also please include which herbarium the specimen(s) is deposited in using the Index
Herbariorum acronym. The manuscript should end with the name and address of the author(s).
3. Letters to the Editor can be formatted as general text without the specific sections listed above.
However, literature cited and any tables or figures should be formatted as described below.
4. Please create tables using either a tab delimited format or a spreadsheet using Excel or other sim¬
ilar program. Each table is to be submitted as a separate file. Table captions should be placed at
the top of the table. Any footnotes should appear at the bottom of the table. Please do not insert
tables within the body of the text.
5. Send each figure as a separate file in a high-resolution format — eps, jpg, or tif. Figures like bar
graphs that gain their meaning with color won’t work — use coarse-grained cross-hatching, etc.
Create figure legends as a separate text file, and the typesetter will insert them as appropriate.
Please do not insert the figure in the body of the text file.
6. Citations: Please verify that all references cited in the text are present in the literature cited sec¬
tion and vice versa. Citations within the text should list the author’s last name and publication
year (e. g. Smith 1990). For works with more than 2 authors, use “et al.”, and separate multiple
citations with a semicolon.
7. Literature Cited: List citations alphabetically by author’s last name. Author names are to be list¬
ed with surname first, followed by initials (e. g. Smith, E. B.). Separate author’s initials with a
single space. The year of publication should appear in parentheses immediately before the title
of the citation. The entire journal name or book title should be spelled out. Please put a space
after the colon when citing volume number and page numbers.
8. Italicize all scientific names. Voucher specimens must be cited for floristic works or any other
relevant study. Papers citing plant records without documenting vouchers are generally not
acceptable.
9. Manuscripts may be submitted electronically to the email address of the editor. Printed versions
of manuscripts may also be submitted in which case three copies should be provided. All manu¬
scripts will be reviewed by at least two referees. A more complete set of instructions is available
at http://www.michbot.org/publications/Botanist/instruct_authors.htm.
CONTENTS
Assessment of a Novel Reclamation Effort in
Grand Mere State Park
Megan D. Edwards, Ranessa L. Cooper,
Manning Serafin, and Brooke E. Toppen
The Sugar Maple Sapstreak Fungus (Ceratocystis virescens)
(Davidson) Moreau, Ascomycota) in the Huron Mountains,
Marquette County, Michigan
Dana L. Richter
Noteworthy Collection
Michael Rotter, Susan Fawcett, and Danny McConnell
Book Reviews
On the cover: Opuntia humifusa {Prickly -pear) from the gentle slope microhabitat
at the former Manley-Peters sand mine in Berrien County s Grand Mere State Park.
(Photograph by Ranessa L. Cooper on 24 June 2008.)
XM
.1S3
v/. 5'
n o . ^>
July-September, 2012
Vol. 51 , No. 3
A Journal of Great Lakes Botany
THE MICHIGAN BOTANIST (ISSN 0026-203X) is published four times per year by the Michigan
Botanical Club (www.michbotclub.org) and is available online at http://quod.lib.umich.edu/rn/
mbot/. The subscription rate is $25.00 per year. Periodicals postage paid at Ann Arbor, MI 48103.
The office of publication is Western Michigan University, Kalamazoo, MI 49008.
On all editorial matters, please contact Todd J. Barkman, 3437 Wood Hall, Department of Biological
Sciences, Western Michigan University, Kalamazoo, MI 49008; 269. 387. 2776 (Phone), 269.
387. 5609 (FAX); todd.barkman@wmich.edu. All articles dealing with botany in the Great Lakes
region may be sent to the Editor at the above address. In preparing manuscripts, authors are
requested to follow the “Instructions for Authors” on the inside back cover.
For all inquiries about back issues and institutional subscriptions please contact Caroline Barkman,
The Michigan Botanist Business Office, 919 Dobbin Dr., Kalamazoo, MI 49006; 269. 544. 0034;
cbarkman9147@mail.kvcc.edu
Todd J. Barkman, Editor
Editorial Board
Caroline Barkman, Business Manager
L. Alan Prather
Anton A. Reznicek
J. Dan Skean, Jr.
Anna K. Monfils
Timothy M. Evans
Catherine H. Yansa
THE MICHIGAN BOTANICAL CLUB
Membership is open to anyone interested in its aims: conservation of all native plants; education of
the public to appreciate and preserve plant life; sponsorship of research and publication on the
plant life of the state and the Great Lakes area in general, both in the USA and in Canada; spon¬
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2012
THE MICHIGAN BOTANIST
89
HYBRIDIZATION DYNAMICS OF
INVASIVE CATTAIL ( TYPHACEAE ) STANDS AT
PIERCE CEDAR CREEK INSTITUTE:
A MOLECULAR ANALYSIS
Kelsey Huisman, Alex Graeff, and Pamela J. Laureto
Department of Biological Sciences
Grand Rapids Community College
143 Bostwick NE
Grand Rapids, Michigan 49503
plaureto @ grcc . edu
ABSTRACT
Three cattail taxa are recognized in Michigan USA: native Typha latifolia (broad-leaf cattail), the
invasive Typha angustifolia (narrow-leaf cattail), and the hybrid of these two species Typha x glauca.
Typha angustifolia and T. x glauca are of special interest because of their ability to aggressively
spread and out-compete the native cattail T. latifolia. Typha x glauca has been shown to out-com-
pete both its parental taxa and produce monospecific stands. We surveyed the Pierce Cedar Creek In¬
stitute (PCCI) property for cattails and located 25 distinct cattail marshes. We determined the total
area of cattail marsh at PCCI to be roughly 10% of the 267 ha property. Cattail individuals were sam¬
pled from each of the 25 stands and RAPD markers were used to identify the individuals to species.
We found that 20 of the 25 stands were monospecific for the native cattail, T. latifolia. Five of the
stands were mixtures of the native T. latifolia and the introduced T. angustifolia, and T. x glauca was
found in two of the mixed stands. We recommend removal of the invasive T. angustifolia and T. x
glauca individuals and the establishment of a monitoring plan in order to maintain the long-term
health of the cattail marshes at PCCI.
KEYWORDS: Typha spp., RAPD markers, invasive species
INTRODUCTION
Species of Typha L. (Typhaceae), commonly known as cattails, are highly
productive emergent plants that grow in a variety of wetland habitats throughout
the world (McManus et al. 2002). In the northern USA and Canada three taxa of
cattail have been recognized: Typha latifolia L. (broad-leaf cattail), Typha an¬
gustifolia L. (narrow-leaf cattail), and Typha x glauca Godr. (white cattail).
Typha latifolia is a native plant species that is a keystone emergent in marsh
communities throughout North America (Smith 2000). In the United States,
broad-leaf cattail is native to all 50 states. In Canada, it occurs in all provinces
and territories except Nunavut (USDA, NRCS 2012).
Typha angustifolia is thought to have been introduced to the eastern seaboard
from Europe in the early 19th century (Stucky and Salamon 1987; Selbo and
Snow 2004), although Shih and Finkelstein (2008) suggest it may be present in
pollen cores dating back 1,000 years. Following its colonization of the Atlantic
coast, T. angustifolia began to move inland slowly — but by the early 20th century
it had begun a rapid westward expansion (Mills et al. 1993). Today, T. angusti-
90
THE MICHIGAN BOTANIST
Vol. 51
folia is found in 42 of the 50 United States; it is absent from Florida, Georgia,
Alabama, Mississippi, Texas, Utah, Arizona, Hawaii and Alaska. In Canada it is
found in all provinces except Labrador and Newfoundland and is absent in the
Yukon, Northwest, and Nunavut Territories (USDA, NRCS 2012). Because of its
aggressive spread, T. angustifolia is considered an invasive species. It often out-
competes native wetland species, including T. latifolia, to produce very dense
monospecific stands (Grace and Harrison 1986).
Typha latifolia and T. angustifolia are obligate wetland species, meaning that
they are always found in or near water. Both species generally grow in flooded
areas; however, T. latifolia is typically found in waters that do not exceed 0.8 m
while T. angustifolia prefers deeper water, usually greater than 0.75 m. Where
the two species are sympatric, they typically segregate by water depth (Travis et
al. 2010).
According to Grace and Harrison (1986), both T. latifolia and T. angustifolia
are self-compatible, wind pollinated, clonal species. The reproductive shoots of
both species are monoecious, with staminate flowers occurring above pistillate
flowers, and protogynous (pistillate flowers produced prior to staminate flow¬
ers). The pistillate flowers remain receptive to pollen for four weeks (Kuehn et
al. 1999). While the protogynous condition would seem to facilitate out crossing
(Smith 2000), Krattinger (1975) showed that cattails are largely self-fertilized
and that vegetative reproduction occurs more frequently than sexual reproduc¬
tion. In part, the two species can be distinguished by the presence or absence of
a spike gap between the staminate and pistillate flowers ( T. latifolia generally
has no spike gap, whereas T. angustifolia has a spike gap ranging from 0.5 to
4 cm). T. latifolia often flowers later than T. angustifolia but overlap in flower¬
ing times can lead to hybridization between the species (Selbo and Snow 2004).
In fact, T. latifolia and T. angustifolia appear to hybridize wherever the two
occur sympatrically (Galatowitsch 1999; Kuehn et al. 1999; Olsen et al. 2009).
The third taxon, Typha x glauca, is the hybrid of T. angustifolia (maternal)
and T. latifolia (paternal) (Grace and Harrison 1986; Kuehn et al. 1999); how¬
ever, its hybrid status has long been disputed. Typha x glauca has been identified
as a distinct species; a stabilized hybrid taxon; an introgressed taxon with
T. x glauca representing a series of intermediates in a hybrid swarm; and also as
a sterile Fj hybrid (Kuehn et al. 1999 and references therein). Taxonomic de¬
scriptions of T. x glauca frequently identify the plant as a sterile F\ hybrid; how¬
ever, recent studies have revealed the presence of backcrossed and later genera¬
tion hybrids indicating that T. x glauca has at least some degree of fertility
(Snow et al. 2010; Kirk et al. 2011).
The spread of invasive taxa are of particular interest to evolutionary biolo¬
gists and ecologists because of their ability to alter community structure and
ecosystem function (Horvitz et al. 1998). Typha x glauca is considered to be a
highly invasive species due to its aggressive range expansion and ability to dom¬
inate wetland habitats. According to Galatowitsch et al. (1999), several hypothe¬
ses have been advanced in an effort to explain how introduced plant species be¬
come invasive species. The Introgression/Hybrid Speciation hypothesis suggests
that interspecific hybridization between an introduced taxon and a native taxon
results in novel phenotypes with selective advantages. Therefore, hybridization
2012
THE MICHIGAN BOTANIST
91
between native and introduced species is considered to be one of the driving
forces behind the evolution of invasiveness (Ellstrand and Schierenbeck 2000;
Schierenbeck and Ellstrand 2009). Hybrids between native and introduced
species have frequently been shown to have increased fitness with respect to
their parental species as they possess greater genetic and phenotypic diversity
than their parents (Kuehn et al. 1999; Ellstrand and Schierenbeck 2006; Kirk et
al. 2011). This appears to be the case for T. x glauca which can colonize the en¬
tire range of water depths in which the parental species segregate (Travis et al.
2010). Several researchers (e.g., Zedler and Kercher 2004; Travis et al. 2010)
have documented the competitive superiority of f x glauca indicating its po¬
tential as a highly invasive taxon.
Fi hybrids are generally expected to be morphologically intermediate to their
parental taxa but this is not necessarily the case for the hybrid T. x glauca. The
leaf width of the hybrid T. x glauca is believed to range from 6 mm to 16 mm;
although leaf widths up to 21 mm have been reported. Kuehn et al. (1999) found
considerable phenotypic variation in leaf width for each of the parental species.
The leaf width of T. angustifolia ranged from 4.5 mm to 12 mm and the leaf
width of T. latifolia ranged from 7.5 mm to 22 mm. Therefore, the leaf width of
T. x glauca overlaps with the parental species making this trait unreliable for hy¬
brid identification. Flowering in cattails is also an unreliable trait for taxonomic
identification because cattails often exhibit poor flower production. Dickerman
(1982) found that over a three-year period only three of 1,779 marked shoots
flowered at Lawrence Lake in Barry County, Michigan. The degree of shading,
including self-shading in dense stands (Grace and Wetzel 1982), and the depth of
rhizome submergence (Grace 1989) affect flowering success. In addition to poor
flowering and the overlap in morphological traits, backcrossed and advanced
generation hybrids are phenotypically more similar to one of the parental taxa
further complicating the identification of hybrid individuals through morpholog¬
ical traits (Kuehn et al. 1999; Selbo and Snow 2004; Snow et al. 2010). Because
morphological traits can be highly variable, DNA markers are considered to be
more reliable for the identification of cattail species and their hybrids (Kuehn et
al. 1999; Selbo and Snow 2004).
This study examined 25 discrete marsh populations at Pierce Cedar Creek In¬
stitute in Hastings, Michigan, USA in order to gain an understanding of the dis¬
tribution and abundance of T. latifolia , T. angustifolia, and T. x glauca. We used
random amplified polymorphic DNA (RAPD) markers coupled with intensive
field sampling of the 25 populations to identify individual cattails to species.
Kuehn et al. (1999) developed species-specific RAPD markers for T. latifolia
and T. angustifolia. RAPD analysis of genomic DNA is expected to produce
species-specific RAPD banding patterns while the hybrid, T. x glauca, is ex¬
pected to display the banding patterns of each parent. Because of the potential
for T. angustifolia and T. x glauca to invade the marsh communities at PCCI and
out-compete the native T. latifolia, baseline data on the presence and distribution
of Typha spp. is imperative to the development of an effective wetland manage¬
ment plan for the institute.
92
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 1 . Cattail stands identified at Pierce Cedar Creek Institute in Hastings, Michigan. Numbers
correspond with the coding assigned to sampled individuals. (See Table 1 for the species composi¬
tion of each stand.)
METHODS
Study Site
This research took place at Pierce Cedar Creek Institute (PCCI) in Hastings, Michigan, USA.
PCCI is an environmental field station and nature center whose property consists of forest, field,
open water, and a variety of wetland habitats. Of the 267 ha site, a total of 103 ha is considered wet¬
land. Brewster Lake and approximately 1.9 km of Cedar Creek are contained on the property and
many of the wetlands are associated with these bodies of water. Most of the property was previously
farmed and many of the wetland habitats along Cedar Creek appear to be recovering from distur¬
bance. In addition, the creek has also been dammed by beavers. This has led to flooding and the cre¬
ation of shallow water habitat which is ideal for cattail growth. The cattail composition at PCCI has
not been previously studied, but a 2003 vascular plant inventory of the property indicated that only
T. latifolia was present (Slaughter and Skean 2003).
2012
THE MICHIGAN BOTANIST
93
Mapping of Cattail Stands
In April of 2012, we visually surveyed the property by traveling on foot along the trails, and also
by canoeing Cedar Creek. After visual determination of the locations of cattail stands, each stand was
mapped using a hand-held GPS unit (Garmin c-Trex 30). Mapping consisted of traveling the perime¬
ter of each cattail stand with the GPS unit creating an electronic track file. This data was entered into
a geographical information system (ArcGIS 10) to create a digital map depicting the location of each
cattail stand on PCCI property (Fig. 1). Each track on the GPS, with the exception of the Brewster
Lake track, was created by walking the perimeter of the cattail stand. The Brewster Lake track was
created by rowing around the perimeter of the lake in a boat.
Taxon Sampling
Typha samples were collected from each identified cattail stand. Within each stand, we began
collecting at the edge and traveled straight-line transects, by foot, through the stand, stopping at least
every 10 m to collect leaf tissue from the nearest ramet (an individual shoot of a clonal organism). In
larger stands, the distance between sampled ramets often exceeded 10 m. For stands that were too
small to establish transects, or in cases where traveling in a straight line was impractical, sampled
ramets were separated by a distance of no less than 10 m. Having a minimum distance of 10 m be¬
tween samples has been shown to decrease the likelihood of sampling clone mates (Kuehn et al.
1999; Olsen et al. 2009; Snow et al. 2010). No a priori species assignments were made at the time
of collection since T. latifolia, T. angustifolia and T. x glauca display significant overlap in leaf width
(Kuehn et al. 1999, Olsen et al. 2009) and all samples were collected prior to flowering; however we
did sample from a representative range of morphological variants within each stand. We sampled by
clipping approximately 7 cm of leaf tissue from the youngest leaf. The tissue was immediately
placed into a plastic bag containing silica gel desiccant. Each sampled ramet was flagged and num¬
bered along with the corresponding tissue sample.
Molecular Identification of Typha Species and Hybrids
Total genomic DNA was extracted from approximately 3 cm2 of dried leaf tissue following a
variation of the 2 x CTAB (cetyl trimethylammonium bromide) extraction method for high polysac¬
charide plants described by Nickrent (2006). DNA extractions were stored at -20°C for use in ge¬
netic identification of species and hybrids.
We used RAPD molecular markers to identify T. latifolia, T. angustifolia, and T. x glauca. Total
genomic DNA was amplified with PCR using RAPD primers A2 (TGCCGAGCTG) and A8 (GT-
GACGTAGG) (Eurofins mwg/Operon Huntsville, AL, USA). Each 25 pL reaction mixture contained
19.7 pL ultra-sterile H20, 2.5 pL 1 x PCR buffer (200 mM Tris-HCl [pH 8.4], 500 mM KC1), 1 pL
50 mM MgCl2, 1 pL of either RAPD - A2 or RAPD - A8 primer, 0.25 pL 10 mM PCR Nucleotide
Mix (USB Corp. Cleveland, OH, USA), 0.05 pL Platinum® Taq DNA Polymerase (Invitrogen,
Carlsbad, CA, USA), and 0.5 pL total genomic DNA.
The 25 pL reaction mixtures were incubated in an iCycler Thermal Cycler (Bio-Rad Laborato¬
ries, Inc., Hercules, CA, USA) programmed for 2 min at 94°C (initial denaturation); 25 cycles of 1
min at 94°C, 1 min 30 sec at 34.5°C, and 2 min 20 sec at 72°C followed by 20 cycles of 1 min at
94°C, 1 min 30 sec at 34.5°C, and 2 min 20 sec at 72°C with a 10 sec increase after each cycle.
RAPD products were separated by gel electrophoresis on 1.2% agarose gels for 1 h at 90 V. Gels
were stained in an ethidium bromide bath for 15 min and photographed under a UV light source.
Molecular weights of amplification products were estimated using a 1 kb plus DNA ladder (Invitro¬
gen, Carlsbad, CA, USA). Primer A2 yields 5 species-specific bands with bands for T. latifolia at 1.5
kb, 1.0 kb, and 0.6 kb and bands for T. angustifolia at 1.2 kb and 0.8 kb; primer A8 produces 3
species-specific bands with T. latifolia having a band at 1 .0 kb and T. angustifolia having bands at
2.0 kb and 1.8 kb (Kuehn 1999). Ft T. x glauca individuals are expected to show all parental bands.
RESULTS
Twenty-five wetland communities containing Typha spp. were identified
from PCCI property (267 ha). These ranged in individual area from 8 m2 to
97,613 m2 (Table 1; Fig. 1). The largest stands were associated with the perime-
94 THE MICHIGAN BOTANIST Vol. 5 1
TABLE 1. Area, number of Typha individuals (n) identified, and species composition of the 25 cat¬
tail stands identified at Pierce Cedar Creek Institute. Area was calculated in Garmin e-Trex 30. The
species composition of each stand (L - Typha latifolia, A - Typha angustifolia, and G - Typha x
glauca) is presented as a percentage of each species per the total number of identified individuals
from the stand.
Stand
Area (m2)
n
% Species Composition
100
12181.00
9
L (100%)
200
717.82
4
L (100%)
300
27583.00
12
L (100%)
400
20048.00
15
L (100%)
500
3507.20
4
L (100%)
600
10751.00
8
L (75%), A (25%)
700
752.56
2
L (50%), A (50%)
800
22149.00
16
L (100%)
900
4607.00
16
L (100%)
1000
882.16
4
L (100%)
1200
97613.00
43
L (100%)
1300
14698.00
2
L (100%)
1400
14028.00
13
L (100%)
1500
5260.00
5
L (100%)
1600
3215.8.
8
L (100%)
1700S
207.03
3
L (33%), A (33%), G (33%)
1700N
177.02
3
L (100%)
1800
586.29
3
L (100%)
1900
95.06
1
L (100%)
2000
7133.76
8
L (100%)
2100
7695.90
14
L (77%), A (23%)
2200
2102.80
5
L (40%), A (20%), G (40%)
2300
1781.10
5
L (100%)
2400
371.96
3
L (66%), A (33%)
2500
1725.80
5
L (100%)
2600
7.63
1
L (100%)
ter of Brewster Lake and the wetland complexes adjoining Cedar Creek. The
smallest stand was a drainage ditch adjacent to the main road into the PCCI
property. This road was constructed between 1999 and 2001 (Brown, Pers.
Comm.). The cumulative area of all cattail stands totaled 259,878 m2 (26.0 ha)
which is approximately 9.72% of the PCCI property.
We collected tissue samples from a total of 370 individuals from across the 25
cattail stands. Of these, DNA was extracted from 252 randomly chosen individ¬
uals, representing all 25 stands. Photos of two representative electrophoresis gels
of RAPD DNA fragments produced by PCR amplification of cattail individuals
using primer A8 are shown in Figure 2. The species-specific bands and species
identification are indicated on the photos. We identified most individuals to
species using primer A8. For approximately 1% of the individuals we confirmed
the primer A8 species identification using Primer A2. No difference in species
identification was observed between the two primers. RAPD analysis allowed us
to identify 212 of these individuals to species: 200 (94.34%) were T. latifolia , 9
(4.25%) T. angustifolia , and 3 (1.42%) were the hybrid T. x glauca.
Of the 25 cattail stands at PCCI, 20 were monospecific, consisting of only the
2012
THE MICHIGAN BOTANIST
95
A.
B.
Figure 2. Two exemplary gel photos showing ran¬
dom amplified polymorphic DNA (RAPD) pheno¬
types for cattail individuals identified using primer
A8. The left lane contains a 1-kb plus DNA ladder
size standard. A species-specific band for Typha lat-
ifolia (■) occurs at lkb and two species-specific
bands for T. angustifolia (•) occur at 2.0kb and
1.8kb. A. One T. x glauca (G) and two T. latifolia
(L) individuals identified by RAPD analysis. B.
Eight T. latifolia (L) and four T. angustifolia (A) in¬
dividuals identified by RAPD analysis.
native cattail, T. latifolia , four were mixed for T. latifolia and T. angustifolia, and
two stands consisted of a mixture of both parental species and their hybrid T. x
glauca (Table 1).
DISCUSSION
This study documents the extent to which the native cattail Typha latifolia,
the introduced and widespread cattail T. angustifolia, and the hybrid of these two
species, T. x glauca occur at Pierce Cedar Creek Institute. Approximately 40%
of the PCCI property is characterized as wetland habitat based on its soil type
and hydrology (Brown, pers. comm.). However, many of these wetlands are vis¬
ibly dry throughout much of the year. Because cattails prefer standing water, we
were surprised to find that nearly 10% of the property consisted of cattail
marshes. The density and species composition within each sampled wetland var¬
ied from what appeared to be a monospecific stand of cattails with one ramet im¬
mediately adjacent to another, to stands in which individual cattail ramets were
separated by distances of approximately 10 m. Diversity appeared to be much
higher in these stands with a variety of sedges, forbs, and shrubs occurring in be¬
tween the Typha ramets.
On our initial observation most stands appeared to be the native T. latifolia
because the individual ramets consisted of broad leaves; however, variation in
96
THE MICHIGAN BOTANIST
Vol. 51
leaf width led us to suspect that genotyping would reveal mixed stands of both
parental species and their hybrid. Within the large stand that surrounded Brew¬
ster Lake (stand 1200; Fig. 1) a 50 m x 3 m patch of narrow-leaved ramets grew
out into the lake to a water depth of about 1 meter. A second patch of narrow¬
leaved ramets occurred in stand 2100 in an area of deeper water. We suspected
that individuals from these two patches of narrow-leaved, deeper water ramets
would genotype to T. angustifolia.
Our results demonstrate that the native cattail, T. latifolia, is the dominant cat¬
tail species at PCCI; 20 of 25 cattail stands were monospecific for T. latifolia.
Based on variation in leaf widths we believed that many of the cattails stands at
PCCI would be mixed for both parental species and their hybrid. We found evi¬
dence of the widespread, introduced species, T. angustifolia, in only six of the 25
stands and in two of these mixed stands the hybrid T. x glauca was present. We
had suspected that a patch of narrow-leaved individuals from the Brewster Lake
stand (stand 1200; Fig. 1) was T. angustifolia but genotyping did not reveal any
T. angustfolia individuals from Brewster Lake. This is likely due to random sam¬
pling of our collected tissue samples for genotyping which may have excluded
individuals from this patch. Genotyping did however reveal the patch of narrow¬
leaved individuals from stand 2100 to be T. angustifolia. It should be noted that
for most individuals we only analyzed a single RAPD locus and with only a sin¬
gle analyzed locus it is possible that individuals identified as T. latifolia could in
fact be T. angustifolia if introgression has occurred. Therefore, there is a chance
that our species assignments could be more complex if introgression is rampant
throughout the cattail stands at PCCI. Additionally, our % species composition
(Table 1) was calculated as a percentage of each species per the total number of
RAPD identified individuals per stand. Since several stands had limited RAPD
genotyping the % species composition should be considered tentative with re¬
gard to the entire stand.
The homogenous T. latifolia stands were primarily associated with the wet¬
lands surrounding Brewster Lake and Cedar Creek. It is likely that these stands
experience lesser disturbance from changes in hydrology and water depth. Typha
latifolia is reported to have superior growth and competitive ability over T. an¬
gustifolia in shallow, undisturbed, water habitats (Grace and Wetzel (1989). By
contrast, T. angustifolia was shown to be competitively superior to T. latifolia in
shallow water habitats when they were highly eutrophic (Weisner 1993). We
found that mixed parental stands were located in areas that are likely experienc¬
ing greater disturbance because of their proximity to the PCCI trail system,
roads, and other anthropogenic influences such as culverts which may be acting
to alter hydrology and increase sedimentation. Stand 1700S was of particular in¬
terest because it is a small retention pond that was constructed on the PCCI prop¬
erty sometime between 1999 and 2001 (Brown, Pers. Comm.). The very nature
of a retention pond suggests fluctuating water levels and sedimentation. Since
the hybrid T. x glauca can exist throughout the range of water depths occupied
by its parental species it is not surprising that we identified a hybrid individual
from this site.
It is well documented that when the native T. latifolia occurs sympatrically
with the introduced T. angustifolia they can produce the hybrid T. x glauca
2012
THE MICHIGAN BOTANIST
97
(Galatowitsch 1999; Kuehn et al. 1999; Olsen et al. 2009). Typha angustifolia it¬
self can out-compete the native cattail, T. latifolia, and the hybrid T. x glauca
can out-compete both parental species (Grace and Harrison 1986). Because the
2003 survey of vascular plants at PCCI (Slaughter and Skean 2003) did not iden¬
tify the occurrence of T. angustifolia , we believe it has moved onto the property
within the last ten years. Since a large patch of T. angustifolia is directly adjacent
to the trail system (stand 2100) it is unlikely that the species was overlooked in
the 2003 survey. Therefore, the presence of T. angustifolia is cause for concern
regarding the long-term health of the native T. latifolia cattail marshes at PCCI.
While the identification of hybrid individuals based on morphological characters
alone is not reliable, the leaf width and spike-gap differences between the
parental species seem to allow for the accurate identification of T. angustifolia in
the field. We recommend the removal of all Typha angustifolia individuals and
identified hybrids from the PCCI property in order to ensure the long-term health
of the T. latifolia cattail marshes. The removal of T. angustifolia, the maternal
parent of the hybrid, should prevent the formation of additional hybrid individu¬
als on the PCCI property. At the very least, we recommend that PCCI undertake
a monitoring program to record the health of, and/or changes to, the cattail
marshes on their property.
A cattail marsh monitoring program should keep track of fluctuations in hy¬
drology and water levels, as well as, the numbers and locations of the invasive cat¬
tail species. Typha angustifolia has been shown to prefer deeper water than T. lat¬
ifolia (Travis et al. 2010). Therefore, if water levels increase around Brewster Lake
or Cedar Creek, such as might be caused by beaver dams, the disturbance could
create T. angustifolia appropriate habitat. Additionally, if a cattail marsh regularly
receives additional water and nutrients from runoff, such as through the existing
culvert systems, they are likely to experience increased growth of the invasive cat¬
tail species. An appropriate control technique would be to manipulate water levels
so that the preferred T. angustifolia habitat is not available for colonization.
Optimally, all non-native cattail species would be removed from the PCCI
property. According to the USD A NRCS Plant Guide (USD A, NRCS 2012) cat¬
tails can be removed by mowing the plants after the flowering heads are well-
formed but before they have reached sexual maturity. This mowing treatment
should be followed by a second mowing after 0.5 m-1 m (2-3 feet) of new
growth has occurred (typically 1 month). This treatment opens up habitat for
other emergent vegetation and has been shown to be about 75% effective in
eliminating undesirable cattail stands. Additionally, herbicides can be applied to
the rooted portion of the cut ramets which should kill the submerged rhizome
and prevent regrowth. We predict that without intervention, T. angustifolia and T.
x glauca will spread through the marsh complexes at PCCI altering their species
composition and the foraging behavior of many of the animal species dependent
on them.
ACKNOWLEDGEMENTS
The authors thank the Willard G. Pierce and Jesse M. Pierce Foundation and the Pierce Cedar
Creek Institute Undergraduate Research Grants for the Environment program for providing us with
98
THE MICHIGAN BOTANIST
Vol. 51
financial support and for the opportunity for the first two authors to do independent undergraduate
research. We also thank Grand Rapids Community College for use of their laboratory facilities, and
the Michigan Botanical Foundation for financial support.
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THE MICHIGAN BOTANIST
Vol. 51
RELIC OR RECRUIT? NEWLY DISCOVERED
ARONIA ARBUTIFOLIA (ELL.) PERS. (RED CHOKEBERRY)
IN KENT COUNTY RAISES QUESTIONS REGARDING
PAST AND FUTURE DISTRIBUTIONS
Michael P. Ryskamp and David P. Warners
Department of Biology
Calvin College
Grand Rapids, MI 49546
ABSTRACT
The nomenclatural history of the genus Aronia is replete with name changes, misidentifications,
and debates regarding classification. In Michigan, collections of Aronia arbutifolia (Ell.) Pers. (Red
chokeberry) from the early 20th century are thought to be misidentifications of Aronia prunifolia
(Marsh.) Rehder (Purple chokeberry). A recent discovery of A. arbutifolia in West Michigan could
represent a rare species that has naturally occurred in Michigan in the past or an adventive individ¬
ual that has only recently arrived. We provide documentation of the new state record and discuss its
discovery in the context of earlier ambiguous records of Red chokeberry in the Great Lakes region.
We also consider whether this species is exhibiting a coastal plain disjunct distribution, and we ex¬
amine the possibility that Red chokeberry is expanding its range in response to climate change. Fu¬
ture genetic research and molecular analysis could help answer some of the questions raised by this
interesting discovery in Michigan.
KEYWORDS: Red chokeberry; Aronia arbutifolia, coastal plain disjunct, climate change
INTRODUCTION
In 1933, at the annual meeting of the Torrey Botanical Society, renowned
botanist and landscape architect Henry Teuscher (1933) gave an address on the
question “ what is a species ?” In his short talk, he used the Aronia (chokeberry)
complex to illustrate some of the inherent problems of using morphological cri¬
teria to define species of plants. Four decades later, Professor James Hardin
(1973) revisited “the enigmatic chokeberries” in a paper that outlined the taxo¬
nomic and nomenclatural confusion surrounding the shrubs since the early 20th
century. Today, the taxonomy of the two (or three) polymorphic chokeberry
species in the Aronia genus (or sub-genus) is still debated, its history likely
clouded by interbreeding (Hardin 1973; Voss 1985; Voss and Reznicek 2012),
different reproductive strategies (Campbell et al. 1991; Hovmolm 2004), and
inter-population variability (Hardin 1973; Uttal 1984; Brand 2010). Over the
years, several different genus names have been assigned to this small group, in¬
cluding Mespillus L., Pyrus L. f., Crataegus Lam., Adenorachis Nieuwl., and
Photinia K.R. Robertson & J.B. Phipps (MacMillin 1892; Nieuwland 1915; Brit¬
ton and Brown 1970) [In this paper, we follow the nomenclature outlined by
Voss (1985)]. More recently, studies involving nucleotide sequencing have es¬
tablished molecular phytogenies of Pyrinae ( Rosaceae ) that distinguish Aronia
2012
THE MICHIGAN BOTANIST
101
from Photinia (Campbell et al. 2007; Potter et al. 2007; Guo 2011). Campbell et
al. (2007) recommend maintaining Photinia and Aronia as separate genera, as
their results yielded no molecular evidence for placing them together, “or even
near one another,” on a phylogenetic tree.
The state of Michigan is no stranger to the botanical confusion associated
with these shrubs. Early plant surveys in the state, circa 1900, reported Aronia
arbutifolia (Ell.) Pers. (Red chokeberry) in St Clair (Dodge 1900), Berrien (Beal
1904; Hebert 1934), Ionia (Beal 1904), Kent (Cole 1901), and other counties.
Botanical literature from that period, e.g., W. J. Beal’s Michigan Flora (1904),
lists two species of Aronia, A. arbutifolia and A. melanocarpa (Michx.) Elliot.
(Black chokeberry) [the latter actually listed as A. nigra in Beal (1904)]. How¬
ever, these early records of Red chokeberry in Michigan are universally inter¬
preted as either reports or misidentifications of A. prunifolia (Marsh.) Rehder
(Purple chokeberry) (which sometimes appeared in older literature as a variety
of A. arbutifolia). This taxon (A. prunifolia ) is typically understood to have orig¬
inated as a hybrid between A. melanocarpa and A. arbutifolia (Farwell 1918;
Hardin 1973; Voss 1985). Our recent finding of A. arbutifolia in Kent County
clarifies the occurrence of Aronia in Michigan by providing the first confirmed
report of this taxon in the state. In this paper, we describe the A. arbutifolia dis¬
covery and then investigate some possible scenarios to explain its presence in
West Michigan.
MATERIALS AND METHODS
On November 4, 2011, a group of eight potentially clonal Red chokeberries was discovered dur¬
ing a field trip to Townsend Park, Kent County, MI. The shrubs were found in a moist bottomland
east of Bear Creek, growing in a backwater corridor that ran along the bottom of a west-facing hill¬
side. A small grove of large Thuja occidentalis (Northern White cedar) (10-18" dbh) filled the
canopy around the chokeberries, affording a relatively open understory characterized by groundwa¬
ter upwelling and several waste deep pockets of muck. To the west, a shrub zone of Comus spp.
(Dogwood), Betula pumila (Bog birch), Zanthoxylum americanum (Prickly ash), and Physocarpus
opulifolius (Ninebark) transitioned into an open wet meadow dominated by Carex stricta (Tussock
sedge), Onoclea sensibilis (Sensitive fern), and Cirsium muticum (Swamp thistle). A shrubby ripar¬
ian zone marked the western edge of the meadow, the eastern banks of Bear Creek.
At the time of the discovery in early November, dozens of bright-red cymose infructescences
populated the distal ends of the shrubs’ branches. Tomentose beneath, the scarlet-colored leaves were
elliptical and had short acuminate tips (Figure 1). The tallest shrub reached six meters in height and
was supported by a cedar trunk. All the shrubs were elongate, their lower reaches mostly un¬
branched. Collections of flowering material were made on May 8 2012.
VOUCHER SPECIMENS:
Michigan, Kent Co., Cannon Twp. Townsend Park, c. 200 m SE of Ramsdell Dr. and Six Mile
Rd„ 43° 03'N, 85° 21' W, 4 Nov. 2011, M.P. Ryskamp 1 (CALVIN, MICH); M.P Ryskamp 2
(CALVIN, NY); 8 May 2012, M.P Ryskamp 5 (CALVIN, MICH).
RESULTS AND DISCUSSION
The presence of Red chokeberry in Kent County raises a variety of questions
with regard to the origin, establishment, and mechanism of dispersal to West
102
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 1. Photograph taken November 4, 2011
at Townsend Park in Kent County, Michigan
showing the tomentose leaves and hairy stems of
Aronia arbutifolia, with canescent berries still at¬
tached.
Michigan. Below, we explore several different possibilities that might account
for this surprising discovery, we discuss some of the implications of these possi¬
bilities, and we propose some future research initiatives.
Given the history of misidentifications of Red chokeberry in Michigan (and
in other states), one possible explanation is that what we have identified as Aro¬
nia arbutifolia is actually an introgressant of the hybrid ‘A. prunifolia complex’
(Uttal 1984). This possibility, however, can be eliminated due to several distinct
morphological features of the Townsend Park chokeberries. Firstly, the red-
colored fruits observed on the shrubs at the time of discovery are unique to A. ar¬
butifolia, the fruits of A. prunifolia having a much darker purple to purple-black
hue when ripe (Gleason and Cronquist 1963; Hardin 1973; Voss 1985). Uttal
(1984) calls fruit color “the only reliable basis for separating chokeberry
species.” The fact that both leaves and berries remained on the shrub in Novem¬
ber is also indicative of A. arbutifolia; both Purple and Black chokeberry pro¬
duce fruit that ripen and drop earlier in the season (Hardin 1973; Voss 1985;
Brand 2010). Additionally, the size of the pomes, which we measured to range
4-7 mm in diameter, is congruent with descriptions of A. arbutifolia — the pomes
of A. prunifolia, in contrast, range from 8 to 10 mm (Gleason 1958; Gleason and
Cronquist 1963). Lastly, Robinson and Fernald (1908) describe Red chokeberry
as having cymes that are “more numerously fruited (9-18)” than those of A.
prunifolia , which have 3-10 fruits. Even as late as November, the Townsend
Park shrubs had some cymes with 10 or more pomes still attached (Figure 2),
and during a field trip in May 2012, with the shrubs in full bloom, we counted
10-16 florets per cyme (Figure 3). In light of all these traits [and with confirma-
2012
THE MICHIGAN BOTANIST
103
FIGURE 2. Another photograph taken on November 4, 2011 showing the bright red fall foliage and
pomes of Amnia arbutifolia.
tion from A. A. Reznicek (personal observation)], we are confident that the
shrubs in question are indeed A. arbutifolia.
A second possible explanation for the presence of these shrubs is that they
represent an escaped horticultural form. Again, we think this possibility is un¬
likely in light of how closely the shrubs match morphological descriptions of
wildtype Aronia arbutifolia (as described in the above paragraphs). The most
commonly sold cultivar (and perhaps the only one), A. arbutifolia ‘Brilliantis-
sima,’ has a more compact form, more lustrous leaves, and has larger, glossier
fruits than the wildtype species (Flint 1997; Dirr 2011). This description signifi¬
cantly contrasts with the small canescent berries and the tall elongate growth
habit of the Townsend Park shrubs. Additionally, the habitat immediately sur¬
rounding the Townsend Park chokeberries was replete with native species that
indicate relatively pristine conditions, which differs from the typical disturbed
habitat in which most non-native escapees are found (Herman et al. 2001).
Though we are confident the Townsend Park shrubs are wildtype Aronia ar¬
butifolia, there still remain several questions regarding how, when, and from
where this species arrived. Distribution patterns of coastal plain species may pro¬
vide important insight to these questions. There are several communities of dis¬
junct flora around the Great Lakes region that occur outside their primary range
of the Atlantic and Gulf coastal plains (Reznicek 1994). These concentrations
104
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 3. Amnia arbutifolia in full bloom, Townsend Park, Kent County, Michigan, May 8, 2012.
occur in three main areas: 1) a post-glacial drainage occupying southwest Michi¬
gan and northwest Indiana; 2) the region in Ontario east of the Georgian Bay;
and 3) the sand deposits of central and northern Wisconsin. Smaller representa¬
tions are scattered around the Great Lakes region, in northern Michigan and in a
few different locations along the shores of Lake Erie and Lake Ontario. One of
the most widespread of the disjunct species, Rhexia virginica (Meadow Beauty)
occurs in all three main regions and in some of the minor areas from the western
shore of Lake Erie eastward toward the St. Lawrence River (Figure 4a). The
range of A. arbutifolia is similar to Rhexia (Figure 4b); it centers on the south¬
eastern coastal plain, extending west along the Gulf coast into eastern Texas
(Hardin 1973) and northward into the coastal areas of New England and into the
southern portion of Nova Scotia (Eaton et al. 2010; Haines 2011). With A. arbu¬
tifolia identified in Kent County, it seems reasonable to ask if the same (or some
of the same) edaphic and dispersal factors responsible for the presence of dis¬
junct species around the Great Lakes are also responsible for bringing A. arbuti¬
folia to Michigan — however recent or long ago that may have been.
Purported records of Aronia arbutifolia in Ontario (Accessed through GBIF
Data Portal, data.gbif.org, 2012-01-21) would seem to support a coastal plain
2012
THE MICHIGAN BOTANIST
105
30
[ 100
virginica
loot
FIGURE 4. Distributions of Rhexia virginica (4a) and Aronia arbutifolia (4b). The original distribu¬
tion map of Rhexia appears in Reznicek (1994); images used with permission from the author. The
distribution of A. arbutifolia (4b) was based on Hardin (1973), Gardner et al. (2005), Haines (2011),
and on selected records from the New York State Museum (available upon request).
disjunct distribution pattern, but these records have never been confirmed, and
neither Shrubs of Ontario (Soper and Hiemburger 1982) nor the Ontario Plant
List (Newmaster et al. 1998) recognizes A. arbutifolia as a component of On¬
tario’s flora. Voucher specimens (collected in late summer and early fall) refer¬
enced in The New York Flora Atlas do show populations of Red chokeberries in
the western region of the state, near Lake Ontario and Lake Erie, isolated from
the main population along the Atlantic Coast. Further from the Great Lakes, in
Nova Scotia, Red chokeberry is considered a member of the community of dis¬
junct coastal flora that occurs in the southern region of that province (Eaton et al.
106
THE MICHIGAN BOTANIST
Vol. 51
2010), where Rhexia virginica and several other disjunct species occur. Although
some records remain dubious, it does appear that A. arbutifolia has a tendency to
initiate disjunct populations outside its coastal plain range.
Reznicek (1994) hypothesizes that birds or possibly other dispersal agents
played a role in shaping and sustaining populations of disjunct species. He adds
that migrations of flora from the coastal plains into post-glacial drainages near
the Great Lakes could have provided a seed source for dispersal agents. This
may describe the causative events behind the establishment of Aronia arbutifo¬
lia in Kent County — dispersed by migrating birds that ingested Red chokeberry
fruits in New York, Pennsylvania, Ontario, or elsewhere. If this scenario does de¬
scribe the origin of A. arbutifolia in West Michigan, then it could provide sup¬
port for Reznicek’s hypothesis regarding long-range dispersal of coastal plain
flora.
A more significant task than describing how the shrub came to Michigan,
however, might be discerning when this species arrived. Past records of Aronia
arbutifolia in Michigan, which date back to over a century ago, appear to be er¬
roneous. Many of these identifications had been made before the hybrid was de¬
scribed, when floral keys relied on leaf pubescence to distinguish the tomentose-
leaved A. arbutifolia from the glabrous A. melanocarpa. When Emma Cole
included Red chokeberry in Grand Rapids Flora , for example, she was almost
certainly using degree of leaf pubescence as a determining trait during her
spring-time field surveys, when fruit formation was still months away (Cole
1901). But it is difficult — if not impossible — to say with certainty that Red
chokeberry was never native to Michigan. However, if Red chokeberry has been
a longstanding component of Michigan’s flora, given the clear morphological
differences between A. arbutifolia and both A. melanocarpa and A. prunifolia,
we would expect it to have been documented at some point in the past, unless it
was very rare and local.
Another possibility, and perhaps the most plausible one, is that the Townsend
park shrubs are recent arrivals, and that Aronia arbutifolia is simply expanding
its range, perhaps in response to climate change. If climate change is in fact in¬
fluencing the northern and northwestern range expansion of A. arbutifolia, then
we would expect to see increasing frequencies of the shrub in Kentucky and
other places where it is considered infrequent or absent. A 2003 discovery of Red
chokeberry in southwestern Ohio, a state record collected from Clermont County
(Gardner et al. 2005), might fit the pattern of northward range expansion. Inter¬
estingly, two other state records were found in association with the Clermont
County discovery, Bartonia paniculata (Twining screwstem) and Woodwardia
areolata (Netted chain fern) (Gardner et al. 2005), both of which are listed in
Reznicek’s (1994) paper as disjunct coastal plain species that occur in south¬
western Michigan. The presence of a disjunct community of flora at the site in
Clermont County may indicate a more complex and potentially integrated inter¬
action between climate change and the distribution of coastal plain species, es¬
pecially if the populations in southwestern Ohio have only recently arrived to the
area. If climate change is expanding the ranges of coastal plain flora, then we
would expect to see increasing frequencies of A. arbutifolia and other coastal
2012
THE MICHIGAN BOTANIST
107
plain species occurring in the ‘hotspots’ defined by Reznicek (1994), or, con¬
versely, in new areas outside the classically recognized regions of disjuncts.
Further insight into the origin of this shrub could be gained through genetic
and molecular analyses. If the molecular composition of our Kent County Am¬
nia arbutifolia is found to be similar to representative individuals from Ohio or
from other regions close to Michigan, a recent dispersal event from that particu¬
lar population would be the most likely explanation. But if the Kent County
shrubs are notably distinct, such a finding would support the long-term presence
of an indigenous Michigan population. A more thorough examination of herbar¬
ium specimens from the Great Lakes region and elsewhere could also be valu¬
able in determining whether or not A. arbutifolia ever occurred in the state prior
to its recent discovery in West Michigan.
Genetic comparisons between local Amnia prunifolia, A. melanocarpa, and
A. arbutifolia could also add clarity to the debate over the relative uniformity or
variability of the Aronia genus in this region. Both diploid and polyploid popu¬
lations of A. arbutifolia are known to exist along the Atlantic Coast (Holvmam
2004; Brand 2010), and knowing the ploidy level of the Kent County shrubs as
well as other molecular traits may provide information as to their potential in¬
volvement in local genetic introgression within this enigmatic Aronia complex.
CONCLUSION
Trying to understand the presence of a recently discovered stand of Aronia
arbutifolia in Michigan raises several possible explanations, each with its own
set of implications. Its presence in an area known to be a ‘hotspot’ for coastal
plain disjuncts may support the possibility that an indigenous population of A.
arbutifolia has existed in Michigan for some time. However, in the present con¬
text of climate change, and given that the discovery of the species in Ohio is also
recent, we think it is more likely that this species is a recent arrival from the
southern or eastern United States. If this is indeed the case, then A. arbutifolia
could either be expanding its range opportunistically or, more complexly, doing
so in a manner that follows the patterns of the disjunct coastal plain species. It
remains to be seen whether this shrub represents a vestige of an undisturbed
Michigan ecosystem or a harbinger of a warming climate — molecular analyses
could help illuminate this and some of the many other unknowns surrounding
Aronia. If history is any indication, future investigations into this enigmatic
genus will elicit more questions than answers.
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Campbell, C. S., R. C. Evans, D. R. Morgan, T. A. Dickinson and M. P. Arsenault. (2007).
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cultivation in the vicinity of the Grand Rapids, Michigan. A. Van Dort, Book and Job Printer.
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Dirr, M. A. (2011). Din ’s encyclopedia of woody trees and shrubs. Timber Press. Portland, Oregon.
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multiple species of Atlantic coastal plain flora of Nova Scotia. Species at Risk Act Recovery
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Flint, H.L. (1997). Landscape plants for Eastern North America. 2nd ed. John Wiley and Sons, New
York.
Gardner, R. L., J. K. Bissell, D. Boone, G. Haase, R. McCarty, J. S. McCormac, S. McKee, & D.
Minney, (2005). Noteworthy collections: Ohio. Castanea 70: 74-76.
Gleason, H. A. (1958). The New Britton and Brown Illustrated Flora of the Northeastern United
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adjacent Canada. The New York Botanical Garden.
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Haines, A. (2011). New England Wild Flower Society’s Flora Novae Angliae: A Manual for the
Identification of Native and Naturalized Higher Vascular Plants of New England. Yale University
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Hardin, J. W. (1973). The enigmatic Chokeberries. Bulletin of the Torrey Botanical Club 100:
178-184.
Hebert, P. E. (1934). Ferns and flowering plants of Berrien County, Michigan. American Midland
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Herman, K. D., L. A. Masters, M. R. Penskar, A. A. Reznicek, G. S. Wilhelm, W. W. Brodovich, and
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110
THE MICHIGAN BOTANIST
Vol. 51
EFFECT OF SELECTION LOGGING ON
CAREX ASSINIBOINENSIS W. BOOTT
IN A NORTHERN HARDWOOD STAND
Margaret M. Fox and Blair On-
School of Forest Resources and Environmental Sciences
Michigan Technological University
Houghton, MI 49931
Susan J. Trull
Ottawa National Forest
Ironwood, MI 49938
strull@fs.fed.us
ABSTRACT
A population of Care x assiniboinensis W. Boott (Assiniboia sedge) was monitored during the
season before and for six years following a winter selection harvest in a sugar maple-eastern hem-
lock-black cherry stand on the Ottawa National Forest in the Upper Peninsula of Michigan. A mean
increase in sedge presence of 136% was observed over the seven years of monitoring. General linear
regression of sedge presence data indicated a significantly positive trend over the natural log-trans-
formation of time. Results indicate that canopy opening from winter selection logging in northern
hardwood forests may benefit populations of this threatened species.
KEYWORDS: Carex assiniboinensis, Assiniboia sedge, selection logging, threatened species,
Michigan
INTRODUCTION
A population of Carex assiniboinensis W. Boott (Assiniboia sedge) was mon¬
itored during the season before and for six years following a winter selection
harvest in a closed-canopy sugar maple-eastern hemlock-black cherry stand on
the Ottawa National Forest in the Upper Peninsula of Michigan. The site is rela¬
tively flat, dissected by a ravine associated with an intermittent stream. Soil is a
sandy loam with cobbles on the surface; litter accumulation is thin, typically less
than 2-3 cm. The understory is sparse, about 5% cover, mainly young sugar
maple with occasional Lonicera canadensis (American fly honeysuckle).
Ground cover is also sparse, with Dryopteris intermedia (intermediate wood-
fern) as the most common associate. The sedge forms solid patches in many
parts of the site.
Considering Carex assiniboinensis ’ status as a state threatened species, typi¬
cally a 250-foot buffer would have been established around the area covered by
the species. Within the population area plus the buffer zone, logging and other
ground-disturbing activity would have been prohibited (e.g. USDA Forest Ser¬
vice 2004). The population, however, was not discovered until after the timber
sale contract was awarded. Since harvest was already contracted, the Ottawa
2012
THE MICHIGAN BOTANIST
111
mmmm
§?«
iS
- ' 'Mm Mi
A’/>« life ile
S'«ag
FIGURES 1A AND IB. C. assiniboinensis at the study site.
National Forest Botany Program designed a monitoring program to investigate
potential effects of overstory selection logging on C. assiniboinensis. In addi¬
tion, to potentially lessen impacts to the sedge population, harvest activity was
delayed until snow cover was at least six inches.
112
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 2. Location of the study site.
C. assiniboinensis is a perennial sedge in the Cyperaceae family (Figures la
and lb). It is easily identified by its long above-ground vegetative shoots
(stolons) that can extend up to 2 meters in length, at the end of which new plants
form (Bernard 1959; Tolstead 1946; Hipp 2008; USDA NRCS 2012; Voss and
Reznicek 2012). The stolons are observable in late-summer, while flowering
usually occurs in June and July (Penskar and Higman 1999).
C. assiniboinensis is found on moist sites, in mesic deciduous and mixed
forests, floodplains, and river banks. Its native range extends from northern Iowa
in the south, southeastern Saskatchewan in the west, and east to the Lake of the
Woods region of Ontario near the southern border with Manitoba, and the Upper
Peninsula of Michigan (Penskar and Higman 1999; Flora of North America Ed¬
itorial Committee 2002; USDA NRCS 2012). Michigan is the only state in
which the plant is listed as threatened; its conservation status is imperiled (S2) in
Michigan, Ontario, and Saskatchewan, and vulnerable (S3) in Manitoba and
Iowa (NatureServe 2012). According to the Michigan Department of Natural Re¬
sources, it has a coefficient of conservatism of nine (on a scale of ten); that is, it
is believed to have high fidelity to a natural community that persists in a state
similar to pre-European settlement (Herman et al. 2001).
Effects of overstory management on C. assiniboinensis populations are
poorly understood. Tolstead (1946) and Bernard (1959) hypothesized that stolon
growth is encouraged by light, but no research since that time has confirmed
whether an increase in light would contribute to population growth. Bernard
(1959) also speculated that soil moisture and other characteristics or weather pat¬
terns could affect the degree of stolon formation. He believed that the prevalence
of stolons in C. assiniboinensis populations in North Dakota was due to long¬
standing woodland management and perhaps grazing. The Michigan Natural
Features Inventory recommends that until the effects of timber harvest on C.
assiniboinensis populations can be better understood, logging should be avoided
2012
THE MICHIGAN BOTANIST
113
altogether or at least limited to selection cutting during winter months (Penskar
and Higman 1999). In this study, it was hypothesized that sedge growth could be
promoted as the result of increased light penetration following the creation of
canopy gaps. This hypothesis was formed based on presence of the sedge on old
skid trails in the Ottawa National Forest.
METHODS
The surveyed population of C. assiniboinensis covers a roughly U-shaped area of approximately
five to ten acres located in Beechwood Quadrangle in Iron County, MI (T44N R36W S31) (Figure
2). The first monitoring was conducted in September of 1999 and was carried out every September
through 2005. Harvesting was conducted in the winter of 1999-2000 on the western portion of the
U-shaped area and in 2000-2001 on the eastern portion, with a target residual basal area of 85 square
feet per acre in both portions.
Permanent, parallel southeast-northwest running transects were established at randomly spaced
distances along the east and west sides of the U-shaped area. The sampling regime used permanent
transects in order to (1) be able to monitor changes in the population over time, and (2) allow for the
use of a fewer number of sampling points than would be required to meet the same level of accuracy
in non-permanent transects (Elzinga et al. 1998). Ten transects were established on the west side and
six on the east side. From the fixed starting point, sampling along each transect started at a random
distance between one and nine meters on the west and one and five meters on the east side. The start¬
ing point and subsequent sampling points, which were spaced one meter apart, therefore changed
every year. Thirty points were sampled on each transect on the west side and 25 on each transect on
the east side. At each point, presence or absence of sedge ramets was recorded. Data are summarized
in Table 1 . A sum value for each transect was recorded and then the values from west-side transects
were normalized to 25 points.
Data were analyzed in SAS software
(SAS Institute Inc., Cary NC) using PROC
GLM for general linear regressions. Gen¬
eral linear regressions were used to iden¬
tify a trend in sedge presence following
harvest. The data were analyzed as one
data set, with year one as the season before
harvest for the west (1999-2000) and the
east (2000-2001) sides. A single analysis
was conducted on monitoring data from
the six years following harvest for the west
transects and from the five years following
harvest for the east transects. Year seven
only included data from the west transect.
Results were considered significant at p
<0.05.
TABLE 1. Mean and standard error values for sedge
presence (as ramets) on all transects. Year 1 is the
year before harvest. n=16 for years 1 through 6. n=10
for year 7.
Year
Mean
Standard Error
1
8.44
1.04
2
10.00
1.25
3
16.44
1.37
4
17.13
1.23
5
18.75
1.11
6
19.19
1.02
7
19.99
1.42
RESULTS & DISCUSSION
A mean increase in sedge presence of 136% was observed over the seven
years of monitoring (Figure 3). A linear regression equation was used to model
the effect of time on sedge presence using the natural log of year of survey as the
independent variable and sedge presence as the dependent variable for each in¬
dividual transect and for the mean value of all transects. In order to analyze the
response of the sedge following logging, only the data from the second (2000)
114
THE MICHIGAN BOTANIST
Vol. 51
Year of survey
FIGURE 3. Mean number of sedge ramets per transect over the seven years of monitoring. n=16 for
years 1 through 6. n=10 for year 7.
through seventh year (2005), for the west side, and the third (2001) through sev¬
enth (2005) year for the east side, were used. The regression analysis showed a
significant positive trend in mean sedge presence over time (R2 = 0.89; p =
0.005) (Figure 4).
Results indicate that winter selection logging in a northern hardwood forest
where C. assiniboinensis is present had the effect of increasing sedge cover, at
least over the short term (as the canopy continues to fill in and light decreases,
the sedge population may decrease). Although we did not record size or maturity
of plants observed at the sample points, we assume that the observed increase in
total cover is due to vegetative reproduction, particularly via the characteristic
stolons, rather than from seed. Repeated recruitment from seed is less common
Mean
Expected
Mean
FIGURE 4. Linear regression of sedge ramets as a function of time, x = year of study beginning with
2 = first season after harvest.
2012
THE MICHIGAN BOTANIST
115
in woodland clonal plants where vegetative reproduction typically dominates
(Eriksson 1989). An increase in dominance of C. assiniboinensis in relation to
other species was also observed in the field.
This result is consistent with research (Schultz 1988) on the Hiawatha Na¬
tional Forest, also in Michigan’s Upper Peninsula, which found that a selection
harvest in mesic hardwoods caused a flush of reproductive effort, with an in¬
creased number of fertile culms, in Carex novae-angliae, another state-threat¬
ened sedge with a coefficient of conservatism of 9. However, this treatment did
not result in an increase in total sedge cover.
Increased penetration of light to the forest floor is likely a factor in support¬
ing sedge growth. Studies of the effects of canopy gaps on understory vegetation
cite the increase in light penetration as a causal factor in increased ground cover
following selection cutting (McComb and Noble 1982; Reader and Bricker
1992). Tolstead (1946) and Bernard (1959) both noted increased stolon forma¬
tion in Carex assiniboinensis populations with increased light.
Limiting harvest activities to winter months also could have contributed to
the positive response of the sedge. Harvesting over exposed ground causes soil
disturbance and compaction, which can change microclimates and site suitabil¬
ity for understory plants (Brais 2001). Restricting logging to winter months over
snowpack or frozen ground results in less impact on understory abundance and
percent cover than when logging is conducted during summer months. Species
that are more vulnerable to disturbance (coefficient of conservatism >6) are also
less likely to be found on summer-logged sites (Wolf et al. 2008). The high co¬
efficient of conservatism (9) attributed to C. assiniboinensis (Herman et al.
2001) suggests that if logging was not restricted to the winter, different results
might have been observed in this study.
The results indicate that winter selection logging in hardwood forests can
have a positive impact on populations of C. assiniboinensis , at least in the short
term. Based on this study, it may not be necessary to exclude selection cutting
from an area where C. assiniboinensis is present, despite its threatened status.
Without more research, it is advisable to limit harvest activities to winter months
over several inches of snow.
ACKNOWLEDGEMENTS
We thank I. Shackleford, C. Matula, R. Evans, B. Bogaczyk, S. Zoars, and L. Miskovich for field
assistance and an anonymous reviewer for helpful suggestions. This research was supported by the
Loret Miller Ruppe Scholarship Fund.
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abstracts/botany/Carex_assiniboinensis.pdf
Reader, R.J. and Bricker, B.D. (1992). Value of selection cut deciduous forest for understory herb
conservation: an experimental assessment. Forest Ecol. Manag. 51:317-327.
Schultz, J. (1988). Carex novae-angliae quadrat remeasurements within hardwoods in the Shingle-
ton Fen study area, Schoolcraft County, Michigan. Unpublished document. Marquette, Michigan:
Hiawatha National Forest, 7 pp.
Tolstead, W. L. (1946). Stolons of Carex assiniboinensis Boott in Iowa. Am. Midi. Nat. 35:797 .
USDA Forest Service. (2004). Camp 7 vegetation management project environmental assessment.
Ironwood, MI: Ottawa National Forest, pp. 2-9.
USDA NRCS PLANTS Database, http://plants.usda.gov/java/profile?symbol=CAAS2 (accessed
May 14, 2012).
Voss, E.G. and Reznicek, A. A. (2012). Field manual of Michigan flora. Ann Arbor, MI: University
of Michigan Press, p. 104.
Wolf, A. T., Parker L., Fewless, G., Corio. K., Sundance, J., Howe, R., and Gentry, H. (2008). Impact
of summer versus winter logging on understory vegetation in the Chequamegon-Nicolet National
Forest. Forest Ecol. Manag. 254:35-45.
2012
THE MICHIGAN BOTANIST
117
NOTEWORTHY COLLECTION
Michigan
Lygodium palmatum (Bemh.) Swartz — climbing fern, Hartford fern
Previous Knowledge. Lygodium palmatum is perhaps best known as the sub¬
ject of Henry David Thoreau’s affection and titling as “our most beautiful fern”.
He secreted a colony’s location in Concord, Massachusetts, from all but a few
close friends. It was 120 years later that the legendary colony was found by
botanists (Angelo 1985). Considered local and rare throughout its core eastern
range from New England to Mississippi, Lygodium is reported locally abundant
only in parts of Kentucky and Tennessee where, like the collection reported here,
it grows in moist, acidic soils (Flora of North America 1993).
FIGURE 1. Habitat of
Lygodium palmatum
Van Buren County,
Michigan. Copyright
James G. Krause
118
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 2. Detail of Sterile Pinnae on Lygodium palmatum. Copyright James G. Krause
Previous to this collection, Michigan’s only known station of Lygodium was
in the Gourdneck State Game Area in Portage, Michigan. Located and collected
by Richard W. Pippen and Western Michigan University students in 1965, it rep¬
resented, in addition to the only Michigan occurrence, the northern-most station
of the species in the United States by 250 miles (Pippen 1966). This colony is
documented to have persisted as of 1999 (McKenna 2004) but recent accounts
state it has become increasingly difficult to locate and may be extirpated from
this location (Mike Penskar MNFI and Todd Barkman WMU, pers. comm. Sept.
13, 2012 and Oct. 23, 2012, respectively). In Michigan the plant is listed as En¬
dangered with a rarity ranking of SI (critically imperiled).
Significance. In addition to representing the second known station of Ly¬
godium in Michigan, this collection now documents, at 19 kilometers northwest
of the Portage station, the most northerly station in the United States. As already
noted it may also be the only station in Michigan currently.
The colony reported here is vigorous and larger, at 34 m2, than the colony size
of 4 m2 noted by Pippin. Growing as a dominant species of this relatively large
area, on undeveloped private land, suggests this colony is not introduced. This
new discovery, growing in sandy soil near a sphagnum-floored shrub swamp of
Toxicodendron vernix, Osmunda cinnamomea , and Osmunda regalis , shares
many habitat features known to the Portage Lygodium station (Fig. 1). It is tan¬
talizing to think that these habitat echoes could lead to more Lygodium discov¬
eries in Michigan.
2012
THE MICHIGAN BOTANIST
119
FIGURE 3. Detail of Fertile Pinnae on Lygodium palmatum. Copyright James G. Krause
Diagnostic Characters. Lygodium is a distinct and easily recognized plant;
clearly a fern but unlike any other native fern in appearance and habit. It has ster¬
ile evergreen pinnae, ovate, deeply and palmately lobed, that fork on 1-2 cm
stalks resembling two human hands about to clap (Fig. 2). The fertile pinnae are
smaller than sterile pinnae, borne at the tips of the “vines” from late summer to
late fall, irregularly forked or lobed and notably smaller and more finely palmate
than sterile pinnae (Fig. 3) (Flora of North America 1993)(Cobb, et. al. 2005).
Arguably most unique is it’s twining and climbing, vine-like habit (Fig. 4).
The collection reported here represents a colony of >1000 sprouts that primarily
twine around each other forming mats over stunted Sassafras albidum. The col¬
lected specimen was one of only two climbing plants noted at the time of col¬
lection.
Specimen Citation. Van Buren County, Almena Township. Low, moist, open
transition zone between shrub swamp and dry upland forest; Thetford loamy
sand, pH: 3.9. Twining 55.9 cm vertically on Acer rubrum sapling. Sprouting
profusely to form 34 m2 colony. Dominant understory species; minority under¬
story associates: Sassafras albidum , Gaultheria procumbens, and Diphasiastrum
digitatum. Overstory Acer rubrum. Surficial rhizomes dark brown with red bris¬
tles. Rare. September 30, 2012. James G. Krause 1 with Grant D. Krause. Col¬
lected Under Michigan DNR Permit #2015. (MICH).
120
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 4. Twining and climb¬
ing habit of Lygodium palmatum.
Copyright James G. Krause
Special thanks to Mike Penskar MNFI and Anton A. Reznicek MICH for their
help and guidance with regard to this rare find, and to Todd Barkman for his
comments on this manuscript.
LITERATURE CITED
Angelo, R. (1985) Thoreau’s Climbing Fern Rediscovered. Amoldia 45: 24-26.
Cobb, B., Farnsworth, E., and Lowe, C. (2005) A Field Guide to Ferns and their Related Families,
Northeastern and Central North America. Second Edition. Houghton Mifflin Company. New York,
NY. 417 pp.
Flora of North America Editorial Committee. (1993) Flora of North America, North of Mexico. Vol¬
ume 2: Pteridophytes and Gymnosperms. Oxford University Press, New York. 475 pp.
McKenna, D. D. (2004) Flora and Vegetation of Kalamazoo County, Michigan. Michigan Botanist
43: 137-359.
Pippen, R.W. (1966) Lygodium palmatum, the climbing fern, in SW Michigan. Michigan Botanist 5:
64-65
— James G. Krause
Paw Paw, Michigan
INSTRUCTIONS TO AUTHORS
1. Create text in 12-point Times New Roman font and double space paragraphs throughout. Papers
should be organized as follows: Title, Author(s) and address(es). Abstract with up to 5 keywords.
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be numbered. Please contact the editor regarding any questions related to formatting.
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“Noteworthy collections”, should begin each submitted manuscript followed on the next line by
the State or Province for the species reported. The next line should list the taxon of interest using
the following format: Species Author(s) (Family). Common name. The rest of the manuscript
should include the following named sections: Previous knowledge. Significance of the report.
Diagnostic characters (if desired). Specimen citations, and Literature cited. Each of these sec¬
tions are largely self explanatory; however, “specimen citations” should include the relevant
label data from the voucher specimen(s) including location data, collector(s), collection number,
etc. Also please include which herbarium the specimen(s) is deposited in using the Index
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citations with a semicolon.
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single space. The year of publication should appear in parentheses immediately before the title
of the citation. The entire journal name or book title should be spelled out. Please put a space
after the colon when citing volume number and page numbers.
8. Italicize all scientific names. Voucher specimens must be cited for floristic works or any other
relevant study. Papers citing plant records without documenting vouchers are generally not
acceptable.
9. Manuscripts may be submitted electronically to the email address of the editor. Printed versions
of manuscripts may also be submitted in which case three copies should be provided. All manu¬
scripts will be reviewed by at least two referees. A more complete set of instructions is available
at http://www.michbot.org/publications/Botanist/instruct_authors.htm.
CONTENTS
Hybridization Dynamics of Invasive Cattail (Typhaceae)
Stands at Pierce Cedar Creek Institute: A Molecular Analysis
Kelsey Huisman, Alex Graeff, and Pamela J. Laureto 89
Relic or Recruit? Newly Discovered Aronia arbutifolia (Ell.)
Pers. (Red Chokeberry) in Kent County Raises Questions
Regarding Past and Future Distributions
Michael P. Ryskamp and David P. Warners 100
Effect of Selection Logging on Carex assiniboinensis W. Boott
in a Northern Hardwood Stand
Margaret M. Fox, Blair Orr, and Susan J. Trull 110
Noteworthy Collection
James G. Krause 117
On the cover: Cattail marsh bordering Cedar Creek at Pierce Cedar Creek Institute ,
Hastings, MI, summer 2012. (Photograph by Alex Graeff).
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October-December, 2012
Vol. 51, No. 4
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MICHIGAN BOTANIST
A Journal of Great Lakes Botany
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2012
THE MICHIGAN BOTANIST
121
SUITABILITY OF CARD AMINE CONCATENATA (MICHX.) SW.
AS AN INDICATOR SPECIES ON THE OTTAWA NATIONAL
FOREST
Margaret M. Fox, Susan J. Trull*, Blair Orr
School of Forest Resources and Environmental Science
Michigan Technological University
Houghton, MI 4993 1
^Ottawa National Forest
E6248 US Hwy. 2
Iron wood, MI 49938
strull@fs.fed. us
ABSTRACT
The cutleaf tooth wort ( Cardamine concatenata) is one of four Management Indicator Species
monitored on the Ottawa National Forest. It is a perennial spring ephemeral used to indicate effects
of forest management, particularly timber harvest, on site conditions in northern hardwoods, and on
populations of forest understory spring wildflowers. Surveys were conducted in 70 treated and 61
non-treated stands. Stands were monitored for abundance of C. concatenata as well as abundance of
three other spring ephemerals ( Cardamine diphylla, Claytonia caroliniana, and Allium tricoccum).
Additional site characteristics that could affect C. concatenata abundance were monitored, including
micro-topography, ground flora, earthworm disturbance, and deer browse. Analysis assessed both the
impact of management on site conditions and the suitability of using C. concatenata as an indicator
species. Results showed no significant relationship between hardwood management and C. concate¬
nata abundance. While C. concatenata was representative of the other three spring ephemerals mon¬
itored, its relative scarcity across the Ottawa National Forest raises questions as to its suitability as a
Management Indicator Species.
KEYWORDS: Cardamine concatenata, Dentaria laciniata, Management Indicator Species, for¬
est management, spring ephemerals, northern hardwoods
INTRODUCTION
Indicator species are used for efficient monitoring of the status of flora, fauna
and environmental conditions in a particular area, but their use in ecosystem and
biodiversity management has been widely critiqued (Niemi et al. 1997; Caro and
O’Doherty 1999; Landres et al. 1988; Lindenmeyer et al. 2000; Carignan and
Villard 2002; Nylen 2011). The USDA Forest Service uses Management Indica¬
tor Species (MIS) in the monitoring of effects from management activities on
plant and animal populations. The cutleaf toothwort, currently referred to as
Cardamine concatenata (Michx.) Sw. but previously known as Dentaria lacini¬
ata Muhl. ex Willd., is used as an indicator of impacts of northern hardwood for¬
est management, particularly harvesting, on woodland wildflower populations
on the Ottawa National Forest (ONF). As a spring ephemeral species representa¬
tive of that guild, C. concatenata was selected as an indicator because of its habi¬
tat specificity to mesic hardwood stands, its sensitivity to disturbance, its slow
122
THE MICHIGAN BOTANIST
Vol. 51
dispersal rate, and its presumed non-palatability to deer. In this paper, results
from monitoring C. concatenata and three other spring ephemerals, in stands
with different harvest histories, are analyzed to assess both the impact of hard¬
wood management on site conditions and the suitability of C. concatenata as a
management indicator.
Herbaceous Species as Ecosystem Health Indicators of Forest Management
There are a number of different types of, and uses for, indicator species. One
such type, ecosystem health indicator species, is used to obtain insights about the
environmental conditions of the resources upon which species depend (Landres
et al. 1988; Caro and O’Doherty 1999). Given their known site or resource re¬
quirements, the presence or abundance of plant species on a site is informative
of the moisture, soil, temperature, light and other conditions at that site. There
are, however, various factors unrelated to habitat quality, such as browse, dis¬
persal rates, and history of disturbance (Kashian et al. 2003) that can affect the
populations of indicator species without necessarily affecting related species.
Given the diversity of factors that affect populations, it can be difficult to assess
the impact of each on an indicator species and to correlate observations of indi¬
cator species populations to environmental conditions (Landres et al. 1988).
Natural resource managers use ecosystem health indicators to monitor the im¬
pacts of active management on site conditions at the stand and landscape levels.
Disturbance resulting from timber harvesting can cause a number of changes to
soil productivity, moisture, light regimes, and temperature on the forest floor.
When harvesting activities are not carried out over snow or frozen ground, skid-
der traffic can cause soil compaction and removal of the litter layer, resulting in
lower infiltration rates and increased erosion (Grigal 2000; Brais 2001; Zenner
and Berger 2008). Opening the canopy allows for increased light penetration to
the forest floor, causing an increase in temperature (Brooks and Kyker-Snow-
man 2008) and cover of herbaceous species (Metzger and Schultz 1984; Scheller
and Mladenoff 2002). While changes in soil moisture vary depending on harvest
intensity and soil disturbance, soil moisture is generally believed to increase fol¬
lowing tree removal due to lower transpiration rates (Bormann and Likens 1979;
Stoffel et al. 2010).
The degree of disturbance and the rate of recovery of site conditions and
stand composition and structure in hardwoods are affected by the intensity and
frequency of harvest (Metzger and Schultz 1984; Fredericksen et al. 1998; Zen¬
ner et. al. 2006). Group selection and clear-cutting cause significant changes im¬
mediately following harvest, including a shift in species composition from more
sensitive herbs to ruderals (Meier et al. 1995; Zenner and Berger 2008). Inten¬
sively harvested northern hardwood sites are slow to recover to pre-harvest con¬
ditions. The impact of single-tree selection and thinning is less severe because
uneven-aged management regimes more closely mimic natural stand succession
(Barnes et al. 1998; Zenner et al. 2006). Under uneven-aged management, initial
recovery may begin within three to five years (McComb and Noble 1982; Met¬
zger and Schultz 1984; Reader and Bricker 1992). As succession progresses and
light penetration to the forest floor decreases over time, the site gradually re-
2012
THE MICHIGAN BOTANIST
123
sembles pre-harvest conditions (Barnes et al. 1998). As time post-harvest in¬
creases, species composition and abundance will be expected to change and re¬
flect the process of stand succession.
Stand-level disturbance occurs less frequently under natural succession than
under timber management. Cutting cycles are typically 10 to 20 years in north¬
ern hardwoods in the upper Great Lakes area (Martin and Lorimer 1996), al¬
though closer to 20 on the Ottawa NF. Even the longest cycles impose changes
at a faster rate than natural processes (Barnes et al. 1998). With increased fre¬
quency of disturbance, understory species, especially those characteristic of late-
successional forests, are less capable of recolonizing a site (Brewer 1980; Duffy
and Meier 1992; Tonteri 1993).
When using herbaceous plants as indicators of management impacts on site
conditions, it is recommended to use species that are sensitive to change, fairly
specialized in their habitat type and site conditions requirements, and easy to
monitor (Caro and O’ Doherty 1999). Within each habitat type (dry conifer,
mesic hardwood, swamps, etc.), site characteristics such as soil conditions, light
availability, and topography vary at the stand level. Plants that favor specific
conditions, such as shade-intolerant species that colonize canopy gaps, are good
indicators of those conditions and for inferring stage of stand succession
(McLachlan and Basely 2001). Species that are sensitive to change are used to
correlate abundance of the indicator species to the degree of disturbance at a site
(Graul and Miller 1984). Species that are so sensitive or specialized that they are
rare are not good indicators because their low populations are difficult to find
and monitor and are incapable of providing useful monitoring data (Landres et
al. 1988). Considering that efficiency is a principal motivation for relying on in¬
dicator species, species selection should take into account their ease of monitor¬
ing. Species should be easily identifiable, their populations should be accessible
for survey, and their population levels should be high enough that absence is not
a problem (Caro and O’ Doherty 1999).
The ability to directly correlate specific site factors to abundance of an indi¬
cator species is essential to the utility of indicator species (Pearson and Cassola
1992). Susceptibility to external factors, such as deer browse, distorts this ability
and weakens the utility of an indicator species (Landres et al. 1988). Species
with low seed dispersal distances, such as ballistic or ant dispersal, have been
proposed as good indicators of stand recovery because they are slow to recolo¬
nize a site after disturbance (Lambeck 1997). Their limited dispersal ability is an
external factor that can prove misleading, however, if the species’ ability to re¬
colonize is limited, not by recovery of site conditions, but by an absence of rem¬
nant populations on sites within the same geographic area (Dzwonko and
Gawronski 1994). On sites where soil, moisture, and light conditions have re¬
covered to a suitable level, species composition recovery still may be limited by
the slow dispersal rates of certain species (Racke 2010). Therefore, the use of
such herbaceous plants as health indicators can confound analysis of manage¬
ment impacts; even when site conditions have recovered, ground flora may
never fully resemble species composition prior to the previous harvest and ear¬
lier disturbances.
124
THE MICHIGAN BOTANIST
Vol. 51
The Forest Service MIS System
The USDA Forest Service uses MIS to monitor the effects of forest manage¬
ment on species populations and viability. In order to comply with the 1976 Na¬
tional Forest Management Act (NFMA) “biodiversity mandate,” as it is com¬
monly referred to, the 1982 NFMA Planning Rule (36 C.F.R. § 219.19 200)
created the MIS system. The “biodiversity mandate” requires that all National
Forests “provide for diversity of plant and animal communities based on the suit¬
ability and capability of the specific land area in order to meet overall multiple-
use objectives” (16 U.S.C. § 1600(g)(3)(B)). Challenged with managing for mul¬
tiple uses, including plant and animal diversity, indicator species are relied upon
where monitoring of all plant and animal species would be prohibitively costly.
The MIS system has been criticized for being ineffective in fulfilling the bio¬
diversity mandate. The use of population indicators has been widely criticized
and is no longer grounded in the scientific literature (Owen 2010; Nylen 2011;
36 C.F.R. § 219 2012). Criticism of the use of MIS has focused on the inability
of a select group of indicator species to account for the full diversity of plant and
animal life on each national forest. The complexity inherent in managing for the
diversity of species and their resource dependencies makes it challenging for the
Forest Service to provide standardized protocols that offer sufficient guidance,
but are also sufficiently flexible to be adapted to the conditions at each national
forest (Nylen 2011).
Cardamine concatenata as an indicator of northern hardwood forest man¬
agement impacts
On the ONF, C. concatenata is one of four species or species suites monitored
under the MIS system since 2006 when the Forest Plan was revised. C. concate¬
nata is a low-growing native spring ephemeral in the mustard family (Figure 1). It
is found in the understory of northern hardwood stands on nutrient-rich sites. Car¬
damine concatenata population trends are used to monitor impacts of timber har¬
vesting of northern hardwood stands on species populations and site conditions.
Spring ephemerals are regarded as good indicators of site conditions because
they require moist conditions, nutrient-rich soil, and an absence of competition
from other ground flora in early spring (Eickmeier and Shussler 1993; McLachlan
and Basely 2001). Since their life cycle begins before leaf emergence, their pres¬
ence is influenced, not so much by canopy cover, as by soil characteristics, mois¬
ture, and temperature on the forest floor (Bratton 1994). Disturbance can result in
their absence from a site either as the direct result of harvest or indirectly due to
changes in site conditions associated with hardwood management activities (Dz-
wonko 1993; Meier et al. 1995). Spring ephemerals typically have limited seed
dispersal ability, through ant or ballistic methods, meaning that once absent from a
site, they are slow to recolonize (Dzwonko and Loster 1992). Their absence from
or decline within a site following timber harvest is considered an indication that
management activities resulted in deleterious changes on the forest floor, and their
recolonization may be an indicator of site recovery if a seed source is available.
Of the spring ephemerals found in northern hardwood stands on the ONF, C.
concatenata was chosen for monitoring because of its sensitivity to disturbance,
2012
THE MICHIGAN BOTANIST
125
Figure 1 . Cardamine concatenata at a study site on the Ottawa National Forest.
ease of monitoring, and its association with the hardwood cover type. Since deer
are presumed to be less likely to browse on plants in the mustard family, this ex¬
ternal factor might not distort monitoring data. Cardamine concatenata was cho¬
sen over the two-leaf tooth wort ( Cardamine diphylla), another mustard, because
C. concatenata is easier to distinguish and it is also a host plant for the West Vir¬
ginia White butterfly ( Pieris virginiensis ), a species on the Regional Forester’s
Sensitive Species list for the ONF. The leaves of C. concatenata turn yellow dur¬
ing senescence in late May or early June, facilitating spotting plants during field
surveys. Vegetative reproduction and ballistic dispersal limit its range and abil¬
ity to recolonize sites after disturbance. Cardamine concatenata is usually found
in northern hardwood stands, focusing insights from monitoring to conditions in
the northern hardwood habitat type across the ONF.
METHODS
Based on the known habitat preferences of C. concatenata , the Ottawa National Forest Geo¬
graphic Information System (GIS) was used to identify northern hardwood stands on nutrient-rich
sites. Acer-Viola-Osmorhiza (AVO) and Acer-Osmorhiza-Caulophyllurn (AOC) ecological landtype
phase map units were chosen because they designate rich, moist soils (Coffman et al. 1984). Stands
identified as northern hardwoods had sugar and red maple, hemlock, yellow birch, and basswood as
dominant tree species. Sites fulfilling both of these characteristics were considered to be suitable
habitat for C. concatenata. Individual stand management history was identified based on ONF
records. Forest management activities included, generally, selection harvests, stand improvements,
126
THE MICHIGAN BOTANIST
Vol. 51
Figure 2. Location of monitored stands across the Ottawa National Forest in Michigan’s Upper
Peninsula (spatial data from USDA Ottawa National Forest).
and commercial thinning. Three-hundred and twenty-nine non-treated stands totaling 5,668 acres and
165 treated stands totaling 4,636 acres were identified as potential monitoring sites. Treated sites are
those that were harvested since the widespread logging that occurred around the end of the 19th cen¬
tury and beginning of the 20th century. For each year of monitoring, a set of treated and non-treated
stands that were larger than 35 acres and with easy road access were chosen. Due to soil and mois¬
ture conditions on the ONF, northern hardwoods are more prevalent on the south and west portions of
the Forest, hence fewer stands were selected for monitoring on the east and north portions (Figure 2).
Starting in 2006, sampling was conducted every year in late May or early June, due to the plant’s
phenology. The number of stands monitored each year varied depending on staff workloads within the
short window of opportunity for monitoring during plant senescence. In each stand, a meander survey
was conducted, lasting about one hour (Goff et al. 1982). Abundance, on a scale from 0 (none) to 6 (ex¬
tensive) and pattern of occurrence of C. concatenata were recorded based on observations in the whole
stand. On a scale of 0 (none) to 4 (abundant), abundance of C. diphylla, Claytonia caroliniana (spring
beauty), and Allium tricoccum (wild leek) were also recorded. On a scale of 1 (scarce) to 4 (abundant),
coarse woody debris, micro-topography, weedy ground flora, and sugar maple regeneration were noted.
Dominant weedy ground flora species were identified and recorded. Earthworm infestation level and
deer browse on woody species in the stand were each rated as high, medium, or low.
Data collected through 2011 were analyzed in SAS software (SAS Institute Inc., Cary N.C.)
using the PROC FREQ and PROC GLM procedures. Because values in some abundance categories
were too low to be analyzed using a chi-square test, a Fisher’s exact test was used to analyze abun¬
dance data of the spring ephemeral plants to other categorical data. In cases where values in each cat¬
egory analyzed were too low to even use a Fisher’s exact test, abundance was generalized to “pre¬
sent” and “absent”. Results of all tests were declared significant at P < 0.05.
A Fisher’s exact test was used in the following procedures. Differences in abundance in treated
versus non-treated stands were tested on all four spring ephemeral species. Difference in abundance
2012
THE MICHIGAN BOTANIST
127
of C. concatenate! by harvest type was tested for each activity type. Activity types included single¬
tree selection cutting, overstory removal cutting, improvement cutting, commercial thinning, and
pre-commercial thinning. Presence versus absence of C. concatenata was tested against presence
versus absence of C. diphylla, C. caroliniana, and A. tricoccum. C. concatenata presence and ab¬
sence values were tested against abundance of weedy ground flora, coarse woody debris and micro¬
topography. A Tukey’s Studentized Range test was used to determine if number of years since har¬
vest influenced presence of C. concatenata. Number of years since harvest ranged from 5 to 33
(mean = 20.7).
A spatial autocorrelation was used to assess the pattern of C. concatenata abundance in each
stand monitored across the ONF. ArcGIS 10.1 [Esri Corporation, Redlands CA] software was used.
Cardamine concatenata abundance was set as the input field, using inverse distance as the concep¬
tualization of spatial relationships parameter and Euclidean distance as the distance method. The data
were not standardized.
RESULTS
In the following analysis, C. concatenata abundance was correlated with har¬
vest, based on the assumption that C. concatenata is a suitable indicator. This as¬
sumption is then tested by analyzing the ability of C. concatenata to indicate site
conditions. Suitability as an ecosystem health indicator species is determined by
the ability, first, of C. concatenata to represent the other spring ephemeral
species and, second, to directly correlate abundance to harvest impacts, control¬
ling for external factors that may also affect population abundance. Sound inter¬
pretation of monitoring results requires consideration of factors independent of
recent harvest activity.
Effects of harvest on site conditions
The Fisher’s exact test showed no significant difference in C. concatenata
abundance between treated and non-treated stands (P =0.98). The same test was
used to determine if the other spring ephemerals that were monitored showed a
difference under treated or non-treated conditions. There was no significant dif¬
ference in C. diphylla (P = 0.70), C. caroliniana ( P - 0.72), or A. tricoccum
( P = 0.23) abundance between treated and non-treated stands. There was also no
difference in C. concatenata abundance in relation to time in years since harvest
( P = 0.66). The results appear to indicate that harvest was not a factor affecting
abundance of spring ephemeral species.
The harvest activities would have produced disturbances that varied in degree
and character. The Fisher’s exact test showed no significant difference in abun¬
dance of C. concatenata under five out of the six activity types; the exception is
improvement cutting (Table 1). Given that only one of the six harvest types, one
with fewer than ten sample points, was statistically significant, it is likely that
this is a spurious relationship.
Suitability of C. concatenata as an indicator
The Fisher’s exact test was used to assess if C. concatenata is a good repre¬
sentative of the other spring ephemeral species. The results showed that the rela¬
tionship between abundance of C. concatenata and C. diphylla (P < 0.0001),
1 28 THE MICHIGAN BOTANIST Vol. 5 1
TABLE 1. Relationship between C. concatenata and management activity type.
n=66, df=6.
Activity Type
Table Probability
P-value
Single tree selection cut
>0.0001
0.41
Improvement cut
>0.0001
0.04
Commercial thin
0.0012
0.64
Sanitation cut
0.1207
0.52
Overstory removal cut
0.5606
1.00
Initiate natural regeneration
0.1698
1.00
Pre-commercial thin
0.0092
0.38
Claytonia caroliniana (P = 0.0014), and A. tricoccum (P < 0.0001) was signifi¬
cant. These results indicate that C. concatenata is representative of the other
spring ephemeral species.
To assess the reliability of using C. concatenata as an indicator species, addi¬
tional factors affecting its abundance were analyzed. Cardamine concatenata
was found to be absent from 68 out of 131 sites (Figure 3). Cardamine concate¬
nata presence versus absence was not significant in relation to coarse woody de¬
bris (P = 0.73), micro-topography abundance (P = 0.09), or weedy ground flora
abundance (i.e. competition) (P = 0.26). Results from the spatial autocorrelation,
used to assess the pattern of C. concatenata abundance in each stand monitored,
showed a clustering pattern across the ONF (P = 0.0008, z = 3.368). The data
were not standardized because of the low variance of 0.0045. The analysis shows
that on a range from 0 (absent) to 6 (extensive), sites with the same abundance
values are located close together on the landscape.
DISCUSSION
If it is assumed that C. concatenata is a good indicator of site conditions, then
the statistical similarity of spring ephemeral abundance in the treated and non-
45
Abundance of C. concatenata
■ Non-treated
■ Treated
Figure 3. Abundance of C. concatenata in treated and non-treated stands. n=131.
2012
THE MICHIGAN BOTANIST
129
treated stands indicates that harvest activities are not negatively affecting site
conditions in northern hardwood stands on the ONF. The harvest techniques,
protective design criteria, and best management practices employed by the ONF
thus would not be expected to significantly alter soil conditions or herbaceous
plant composition. The expected increase in soil moisture following harvest
(Bormann and Likens 1979; Stoffel et al. 2010) would provide the moist condi¬
tions required by spring ephemerals. Restriction of harvesting to winter months
prevents the soil compaction and organic layer disturbance that are the most
detrimental to site conditions (Grigal 2000; Brais 2001; Zenner and Berger
2008). Single-tree selection and thinning, as opposed to group selection and
clear-cutting, are known to more closely mimic dynamics within non-treated
stands (Zenner et al. 2006).
Results showed that C. concatenata is representative of the other spring
ephemerals and that as a group there was little distinction between the four
spring ephemeral species as they related to other factors. None of the spring
ephemerals appeared to be affected by harvest activities. Cardamine concate¬
nata therefore appears to be representative and an adequate indicator of the other
three spring ephemerals.
The ability to correlate C. concatenata abundance to impacts of harvest on
site conditions is weakened by the low population levels of the plant across the
surveyed stands. Cardamine concatenata was identified at only 68 out of 131
sites, and on sites where it was present, abundance was low (Figure 3). Where
present, the low population levels make it difficult to determine the degree to
which forest management may be a factor affecting abundance. Its widespread
absence indicates that factors independent from recent treatments are affecting
C. concatenata abundance. Coarse woody debris, micro-topography, and weedy
ground flora cover did not have an impact on C. concatenata abundance and
were not useful in explaining C. concatenata population dynamics.
A plausible explanation for the widespread absence of C. concatenata is that
as a result of its limited dispersal ability, it may still be recovering from the in¬
tensive disturbance that occurred at the turn of the 20th century (USDA Forest
Service 2006). At that time, forests across much of the Upper Peninsula of
Michigan were clearcut for their valuable timber and then burned over. In the
same period, over multiple years, fire swept through many parts of the area cur¬
rently within the ONF, with at least 20% burning three to four times (Karaman-
ski 1984; Karamanski 1989). Following such intensive disturbance, only isolated
populations of C. concatenata likely survived. Stands can take over a hundred
years to recover from intensive disturbance (Brewer 1980; Duffy and Meier
1992; Tonteri 1993), and recovery of C. concatenata populations would be lim¬
ited by their dispersal ability. Ballistic dispersal of woodland herbaceous plants
is on average only 1.5 meters per year (Cain et al. 1998). With the source for re¬
colonizing C. concatenata plants limited to remnant populations, recovery of the
plant across the approximately one million acres encompassed in the ONF may
still be in progress.
The clustering of C. concatenata populations across the ONF indicates that
population abundance is correlated to geographic position. Without survey data
covering a greater portion of stands across the ONF or population data from be-
130
THE MICHIGAN BOTANIST
Vol. 51
fore the acquisition of the ONF, a more precise mapping of C. concatenata pop¬
ulations in relation to late-successional stands is not possible. The spatial auto¬
correlation results do illustrate that absence of C. concatenata from sites may be
a factor of geographic position on the forest, suggesting that populations could
still be in the process of recolonizing areas of the ONF.
While the limited dispersal of spring ephemerals is considered to enhance
their usefulness as indicator species (McLachlan and Basely 2001), it may prove
misleading in cases where absence from a site is better explained by their slow
rate of recolonization than site conditions. In western Kentucky, transplant of
spring ephemeral species to northern hardwood sites with a history of distur¬
bance resulted in survival and self-propagation, indicating that absence from the
site was due to dispersal limitation and not site conditions (Racke 2010). In
Poland, landscape analysis of spring ephemeral population recovery determined
that close proximity to undisturbed forests was a factor affecting spring
ephemeral presence in secondary forests (Dzwonko and Gawronski 1994).
While it is not conclusive that the legacy of logging and fire from the turn of the
20th century are still affecting C. concatenata populations, it is one reasonable
explanation for the widespread absence of the plant across the ONF.
CONCLUSION
Since 2000, the Forest Service has proposed a number of changes to the orig¬
inal 1982 NFMA Planning Rule. The drafting processes and subsequent rules
have taken criticisms of MIS into account. The newest planning rule, effective as
of May 2012 (36 C.F.R. § 219 2012), acknowledges that the use of population
indicator species is no longer supported in science. The new rule is consistent
with previously proposed planning rules that moved away from “management
indicator species” (population indicators) to what are termed “focal species,”
which are meant to serve as indicators of “ecological conditions” and to provide
information regarding the effectiveness of the Forest Plan in meeting diversity
and ecological sustainability requirements (36 C.F.R. § 219 2012). Under the
2012 Planning Rule the National Forests will have to re-evaluate their monitor¬
ing programs by 2016 and determine which species will serve as focal species,
some of which may be species that were previously MIS.
Results of this study indicate that C. concatenata is not suitable as an indica¬
tor species. It has some of the characteristics that are recommended in indicator
species, and it is representative of other spring ephemerals. It is selective in its
colonization of mesic hardwood sites, sensitive to disturbance, presumed to be
avoided by deer, and has limited dispersal ability. However, its limited abun¬
dance across the ONF restricts C. concatenata' s usefulness. Cardamine concate¬
nata is not recommended for continued use as a MIS or for use as a focal
species.
Analysis of field data showed no significant difference in C. concatenata
abundance between treated and non-treated stands. Despite the widespread ab¬
sence of C. concatenata from both types of hardwood stands, this sample ap-
2012
THE MICHIGAN BOTANIST
131
pears to indicate that the ONF is managing northern hardwoods stands for reten¬
tion of tooth wort and other spring ephemerals.
The intensive disturbance that occurred near the beginning of the 20th century
may be one significant factor affecting C. concatenata absence. Northern hard¬
wood stands may take many years to recover, and on intensively and repeatedly
disturbed sites, species composition may forever be altered. Results of this study
show that it is difficult to monitor site conditions using health indicators given
the complexity of factors that can affect individual species, especially when the
species is not widely distributed. This study will help inform future selection of
focal species and other indicators on the Ottawa National Forest.
ACKNOWLEDGEMENTS
Thanks to A. Amman, L. Palmer, S. Pearson and I. Shackleford of the Ottawa National Forest for
field assistance.
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2012
THE MICHIGAN BOTANIST
133
THE DISTRIBUTION, ECOLOGY, AND
CONSERVATION STATUS OF RUBUSACAULIS MICHX.
(DWARF OR ARCTIC RASPBERRY) IN MICHIGAN
Bradford S. Slaughter
Michigan Natural Features Inventory
Michigan State University Extension
P.O. Box 13036, Lansing, MI 48901-3036
slaugh 1 4 @ msu.edu
Jan Schultz
Eastern Region United States Forest Service
626 E. Wisconsin Ave., Milwaukee, WI 53202
jschultz@fs.fed.us
ABSTRACT
Rubus acaulis Michx. (dwarf or arctic raspberry) is a dwarf herbaceous perennial plant in the
rose family (Rosaceae) characteristic of mesic to hydric open and forested habitats across boreal
North America. The species ranges south locally to the conterminous United States, where it is des¬
ignated a sensitive species by the United States Forest Service in Region 2 in Colorado and parts of
Regions 6 and 9 in Washington and Michigan, respectively. Rubus acaulis was first discovered in
Michigan in 1976, and several additional populations have been documented over the past decade
from minerotrophic peatlands in eastern Upper Michigan, extending west to Marquette County. In
these habitats, which are classified as patterned fen, northern fen, and poor fen, R. acaulis occurs
with a consistent group of vascular plant associates, and is typically most frequent on the tops, sides,
and margins of Sphagnum- dominated hummocks. Despite the recent documentation of several very
large populations, the long-term prospects for the persistence of R. acaulis in Michigan remain un¬
certain due to the threat of climate change and associated disruption of ecosystem processes in its
peatland habitats.
KEYWORDS: Rubus acaulis, Rosaceae, boreal flora, peatland, fen, conservation, climate
change. Upper Michigan
INTRODUCTION
Rubus acaulis Michx. (Rosaceae) is a small herbaceous, rhizomatous rasp¬
berry (Figure 1), traditionally placed in the polyphyletic subgenus or section Cy-
lactis (Raf.) Focke (Alice and Campbell 1999). The species is characterized by
largely herbaceous, generally unarmed, short stems; temate leaves with leaflets
rounded at the apex; solitary, showy pink or rose flowers with glandless sepals
and pedicels that are borne below (and often hidden within) the leaves; and edi¬
ble, red, nearly globular fruits (Soper and Heimburger 1982; Gleason and Cron-
quist 1991; Ladyman 2006). The species is widespread across boreal North
America, where it occurs primarily between 50° and 60° latitude (Soper and He¬
imburger 1982; USDA Forest Service, Eastern Region 2002; Kartesz, BONAP
2011). Rubus acaulis is sometimes taxonomically included within R. arcticus L.
134
THE MICHIGAN BOTANIST
Vol. 51
FIGURE 1 . Rubus acaulis (dwarf raspberry) in a Michigan patterned fen. Note the large flowers with
rose-pink petals and rounded leaflets characteristic of the species. (Photograph by Bradford S.
Slaughter, 18 June 2009.)
(as R. arcticus L. ssp. acaulis [Michx.] Focke), a circumpolar taxon that occurs
at high latitudes primarily in Eurasia and Alaska (Hulten 1968; USD A Forest
Service, Eastern Region 2002; USDA, NRCS 2012).
In the conterminous United States, Rubus acaulis is considered a boreal relict
species with isolated populations in alpine regions of Washington, Montana,
Wyoming, and Colorado and in peatland habitats in the northern Great Lakes re¬
gion of Minnesota, Wisconsin, and Michigan (USDA Forest Service, Eastern
Region 2002; Ladyman 2006; Kartesz, BONAP 2011; Robert W. Freckmann
Herbarium, University of Wisconsin-Stevens Point 2012; MNFI 2013a). In the
core of its boreal North American range, R. acaulis exhibits broad ecological
amplitude, occurring in a variety of generally mesic to hydric habitats, including
moist forests, meadows, bogs, muskegs, alpine slopes, tundra, and streambanks
(Hulten 1968; Ladyman 2006). In the conterminous United States, it occurs in a
restricted subset of habitats, including montane meadows, willow thickets, fens,
riparian areas, and Picea engelmannii (Engelmann spruce) forests in western
states and open to wooded peatlands, especially poor to rich fens, in the northern
Great Lakes States (Wheeler and Glaser 1982; Glaser 1992c; USDA Forest Ser¬
vice, Eastern Region 2002; Washington Natural Heritage Program and U.S.D.I.
Bureau of Land Management 2005; Ladyman 2006; Smith 2008; MNFI 2013a).
Although R. acaulis is considered globally secure, the species is listed as imper-
2012
THE MICHIGAN BOTANIST
135
TABLE 1. Rubus acaulis distribution, conservation status rank (S-rank; NatureServe 2013), state list¬
ing status, and USFS sensitive species status in the United States.
State
s-
State rank1
Listing
Status2
USFS Status
Notes
Sources
AK
SNR
None
None
Widespread
Hulten 1968
CO
SI
None
Sensitive,
Region 2
Four extant sites and one
historical site in the Pike and
Arapaho National Forests
Spackman et al. 1997;
Ladyman 2006
MI
SI
E
Sensitive,
Region 9
See text
MNFI 2007; Voss and
Reznicek 2012
MN
SNR
None
None
Core of conterminous U.S.
range; documented from
18 counties
Glaser 1992c;
MNDNR 2003;
Ladyman 2006; Smith
2008
MT
SNR
None
None
Known from five observations
in alpine peatlands
Montana Natural
Heritage Program 20 1 2
WA
SI
T
Sensitive,
Region 6
One extant and one historic site
Washington Natural
Heritage Program and
U.S.D.I. Bureau of Land
Management 2005;
Ladyman 2006
WI
SNR
None
None
Newly documented in 2007
(Horky #s.n., UWSP)
Robert W. Freckmann
Herbarium, University
of Wisconsin-Stevens
Point 2012
WY
S2
None
Sensitive,
Region 2
Four extant sites and one
historical site in Bighorn
National Forest, Yellowstone
Fertig 2000a, 2000b;
Ladyman 2006;1
Heidel 2012
National Park, and possibly
Medicine Bow National Forest.
Rank recently downgraded
from S 1 .
*SNR= Not Ranked; Sl = Critically Imperiled; S2= Imperiled (NatureServe 2013)
2E= Endangered; T= Threatened
iled or critically imperiled in four of the eight states in which it occurs, includ¬
ing Michigan (Table 1). Reports of the species from Oregon and Maine have not
been confirmed with specimens, and the taxon is not treated in recent floristic
works encompassing those states (Ladyman 2006; Cook and Sundberg 2011;
Haines 2011; C. S. Campbell, pers. comm., 7 February 2012).
Until recently, the status of Rubus acaulis in Michigan was poorly known.
The species was first discovered in the state in 1976, and no new sites were doc¬
umented until the mid-2000s (Voss 1985; MNFI 2013a). Recent discoveries and
collections necessitate a review of our current understanding of the distribution,
ecology, and conservation status of R. acaulis in Michigan. This paper aims to
(1) provide updated collection data to inform the conservation and listing status
of R. acaulis in Michigan; (2) clarify the distribution and ecology of R. acaulis
in Michigan to aid survey and conservation efforts; (3) discuss the challenges as-
136
THE MICHIGAN BOTANIST
Vol. 51
sociated with the conservation of a boreal relict species, particularly in the con¬
text of climatic warming; and (4) suggest avenues for monitoring and research.
COLLECTION HISTORY AND DISTRIBUTION
Rubus acaulis was first discovered in Michigan in 1976 by Charlotte Taylor
while exploring a peatland in Schoolcraft County known colloquially as Shin-
gleton Bog (C. M. Taylor, pers. comm., 11 March 2013). The first collections
from this site date to May 1977 ( Henson # 680 , Henson # 682 , MICH). Following
its initial discovery, 26 years passed before a second location was documented,
this one a partially forested peatland in western Luce County, north of Danaher
Junction (. MacKinnon # 03-283 , MICH). Since that time, the first author and col¬
leagues have documented the species from six additional peatland sites in
Chippewa, Luce, Schoolcraft, and Marquette counties between 46.1304° N and
46.3538° N latitude and 87.2521° W and 85.0236° W longitude (MNFI 2013a;
Figure 2). Most Michigan occurrences are located in the Seney Sand Lake Plain
ecoregion (Seney-Tahquamenon Sand Plain in Omernik and Bryce 2010) that
Legend
• Rubus acaulis ° 20 40 80 Miles
I _ i _ i _ i _ I _ i i i I
FIGURE 2. Known occurrences of Rubus acaulis in Michigan. VIII. 2.1 = Seney Sand Lake Plain
Ecoregion; VIII. 3. 2 = Gwinn Ecoregion (Albert 1995).
2012
THE MICHIGAN BOTANIST
137
encompasses much of the central eastern Upper Peninsula, including portions of
Chippewa, Mackinac, Luce, Schoolcraft, Alger, and Delta counties (Albert 1995;
Figure 2). This ecoregion is characterized by expansive peatlands developed
over poorly drained outwash and lacustrine sands associated with former em-
bayments of Glacial Lake Algonquin (Albert 1995; Jerome 2006). Although
Glacial Lake Algonquin last covered this area between 1 1,000 and 10,000 Y.B.P.,
the peatlands apparently established more recently (beginning approximately
5,000 to 3,000 Y.B.P.) following a climatic shift to cooler, wetter conditions (Fu-
tyma 1982; Futyma and Miller 1986; Madsen 1987; Brugam and Johnson 1997).
At the time of the General Land Office (GLO) surveys in the mid- 1800s, the
Seney Sand Lake Plain supported approximately 190,000 ha (470,000 ac) of
swamp forest dominated by Thuja occidentalis (northern white-cedar) and other
conifers, and 83,000 ha (200,000 ac) of open peatland (Comer et al. 1995). Al¬
though logging and attempts at drainage have altered wetlands in the ecoregion,
the peat deposits currently support extensive mixed conifer swamps dominated
by T. occidentalis, Larix laricina (tamarack), and Picea mariana (black spruce)
(Albert 1995). Open peatlands, including groundwater-fed fens and weakly
minerotrophic to ombrotrophic bogs and muskegs, also remain common, partic¬
ularly on the broadest peat deposits southwest of Seney and northeast of New¬
berry (Comer et al. 1995). Rubus acaulis has been documented from seven such
open peatland sites in the Seney Sand Lake Plain, at elevations ranging from 190
m (620 ft) near the mouth of the Tahquamenon River in Chippewa County to 254
m (830 ft) in the north-central portion of the ecoregion (Figure 2).
In 2010, the first author collected Rubus acaulis from an open peatland
known as the Cyr Swamp in south-central Marquette County that occupies much
of the poorly drained outwash plain in the Gwinn Ecoregion (included within the
Menominee Drumlins and Ground Moraine Level IV ecoregion of Omernik and
Bryce 2010). This is the first documented site for the species outside of the
Seney Sand Lake Plain, and also the westernmost (87.2521° W), southernmost
(46.1304° N), and highest elevation (approximately 330 m, or 1080 ft) docu¬
mented site for R. acaulis in Michigan. West of the Gwinn Ecoregion, higher-re¬
lief landforms developed over ancient metamorphic and igneous bedrock restrict
the development and scale of minerotrophic peatlands, limiting potential habitat
for R. acaulis. However, R. acaulis has been documented from forested peat¬
lands in Douglas County, Wisconsin and St. Louis County, Minnesota, suggest¬
ing the possibility that the species may occur in similar habitats in western Upper
Michigan (Smith 2008; Robert W. Freckmann Herbarium, University of Wis-
consin-Stevens Point 2012).
ECOLOGY
In Michigan, Rubus acaulis occurs in minerotrophic shrub- and sedge-domi¬
nated peatlands, corresponding to several vegetative associations described and
classified in the United States National Vegetation Classification (USNVC) (Na-
tureServe 2013) (Table 2). The Michigan Natural Features Inventory (MNFI)
138 THE MICHIGAN BOTANIST Vol. 5 1
TABLE 2. United States National Vegetation Classification (USNVC) Ecological Associations asso¬
ciated with Rubus acaulis populations in Upper Michigan (NatureServe 2013).
USNVC Ecological Association
Unique Identifier
USNVC Ecological Association Common Name
CEGL002494
Bog Birch - Leatherleaf Rich Fen
CEGL002495
Bog Birch - Shrubby-cinquefoil Rich Boreal Fen
CEGL002500
Boreal Sedge Rich Fen
CEGL005226
Tamarack Scrub Poor Fen
CEGL005227
Bog Birch - Willow Shore Fen
classifies all eight R. acaulis sites as one of three fen types: patterned fen, north¬
ern fen, and poor fen (Kost et al. 2007; MNFI 2013a). Patterned fens appear to
be particularly important habitat for R. acaulis , supporting five of the eight doc¬
umented populations in Michigan. These peatlands are named for their so-called
“ribbed” appearance associated with the presence of alternating peat ridges
(strings) and hollows (flarks) oriented along the contours of the peatland slope,
perpendicular to the flow of groundwater (Kost et al. 2007; Slaughter and Cohen
2010) (Figure 3). Patterned fens develop in “water tracks,” which are concave-
or flat-surfaced zones of drainage that channel sheet flow across peatland sur¬
faces (Glaser 1992a; Price and Maloney 1994). Water tracks begin as narrow
channels in swamp forest or other wetland communities at the upslope margins
of the open peatlands and coalesce and widen downslope (Glaser 1992a). Strings
and flarks are thought to develop due to the repeated inundation of depressions
on the surface of the peatland, which causes their eventual expansion and coa¬
lescence across the contours of the peatland slope (Foster et al. 1983; Foster and
King 1984; Glaser 1992b; Price and Maloney 1994; Quinton and Roulet 1998).
Although patterned fen is known from only 20 sites in Michigan and the com¬
munity is considered imperiled in the state (Kost et al. 2007), many occurrences
are expansive, and the statewide acreage currently occupied by high-quality pat¬
terned fen is approximately 15,600 ha (38,500 ac, or 60 square miles) (MNFI
2013a).
Within patterned fens, Rubus acaulis occurs primarily in hummocky areas
that support Sphagnum, ericaceous shrubs, and wetland conifers. This habitat oc¬
curs in the ecotone between swamp forest and open fen meadow and in “tails” of
woody vegetation that occur either as extensions of the swamp forest within the
fen meadow or immediately downslope of isolated dune ridges, where woody
vegetation develops in response to impeded water flow and increased nutrient in¬
puts from the adjacent mineral soils (Heinselman 1963, 1965, 1970; Glaser et al.
1981; Crum 1988; Glaser 1992a, 1992b, 1992c; Cohen et al. 2009; MNFI
2013a). Within the patterned water tracks that support fen meadow, R. acaulis is
concentrated on the strings, where it is typically rooted in Sphagnum spp (Figure
3). These mosses create a thin, highly acidic (pH= 4. 5-5. 5), fibric substrate over
peats that are otherwise less acidic (pH= 6. 0-7.0), derived from sedges, and more
decomposed. R. acaulis also occurs occasionally in mucky depressions and in
flarks that lack Sphagnum mosses, but the species appears to be less widely dis¬
tributed and less abundant in these habitats (MNFI 2013a). The relative impor-
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THE MICHIGAN BOTANIST
139
FIGURE 3. Poor fen in Marquette County, Michigan. This extensive poor fen supports a very lim¬
ited area of shrub-dominated strings (foreground) and sedge-dominated flarks (middle) where sheet
flow is impeded by scattered dune ridges covered in pine forest (background). Rubus acaulis occurs
primarily on Sphagnum hummocks within this fen. (Photograph by Bradford S. Slaughter, 12 Sep¬
tember 2010.)
tance of Sphagnum hummocks and strings for R. acaulis may be associated with
the water-holding capacity of hummock-forming Sphagnum species (Hajek and
Beckett 2008), which may buffer this microhabitat from the impacts of drought.
The other three Rubus acaulis sites are classified as either northern fen (two
sites) or poor fen (one site). Northern fens and poor fens lack strings and flarks,
but are otherwise characterized by vegetation structure and plant species compo¬
sition similar to that expressed in patterned fen (MNFI 2013a). The northern fen
sites are dominated by fine-leaved sedges and minerotrophic shrubs with limited
importance of Sphagnum and ericaceous shrubs and occur on slightly acid to
mildly alkaline (pH 6. 5-7. 5) sedge- or shrub-derived peats (Kost et al. 2007;
Cohen and Kost 2008a; MNFI 2013a). The single R. acaulis site characterized as
a poor fen supports extensive areas of Sphagnum and ericaceous shrubs with
lower importance of fine-leaved sedges and minerotrophic shrubs and occurs on
medium acid to slightly acid (pH 5. 5-6. 5) sedge- and shrub-derived peats (Kost
et al. 2007; Cohen and Kost 2008b; MNFI 2013a). Although northern fen and
poor fen differ in vegetation dominance and soil properties, R. acaulis tends to
occupy the same Sphagnum hummock microhabitat in both communities (MNFI
2013a). The species occurs with lower frequency and at lower density in sedge-
140 THE MICHIGAN BOTANIST Vol. 5 1
TABLE 3. Vascular plant taxa that occur at seven or eight of eight documented Michigan Rubus
acaulis sites. Taxa that occur at all eight sites indicated in bold. Data sources: MNFI (unpublished
data); Schultz (1987); MacKinnon (2005).
Tree
Shrub
Herb
Larix laricina
Alnus incana
Anemone quinquefolia
Picea mariana
Andromeda glaucophylla
Calamagrostis canadensis
Pinus strobus
Aronia prunifolia
Carex exilis
Thuja occidentalis
Betula pumila
Carex lasiocarpa
Chamaedaphne calyculata
Carex livida
Dasiphora fruticosa
Cladium mariscoides
Lonicera villosa
Comarum palustre
Myrica gale
Coptis trifolia
Rhamnus alnifolia
Doellingeria umbellata
Rhododendron groenlandicum
Drosera rotundifolia
Vaccinium oxycoccos
Equisetum fluviatile
Iris versicolor
Linnaea borealis
Maianthemum trifolium
Muhlenbergia glomerata
Oclemena nemoralis
Osmunda regalis
Pogonia ophioglossoides
Pyrola americana
Rhynchospora alba
Sarracenia purpurea
Solidago rugosa
Solidago uliginosa
Trichophorum alpinum
Trichophorum cespitosum
dominated areas within these fen systems. Of note, R. acaulis has not been doc¬
umented from poor fens weakly influenced by groundwater (pH< 5.5), nor has it
been observed in the moderately to strongly alkaline (pH 8.0-8. 5), marly north¬
ern fens that are especially characteristic of the Niagara Escarpment region of
the eastern Upper Peninsula (Albert 1995; MNFI 2013a).
Vegetative associates of Rubus acaulis in Michigan are consistent across its
three peatland habitats. A total of 22 taxa occur at all R. acaulis sites, and 40 taxa
occur in at least seven of the eight documented sites (Table 3). Stunted individ¬
uals of Larix laricina, Picea mariana, Thuja occidentalis, and Pinus strobus
(white pine) are characteristic of all R. acaulis sites. Fen hummocks are typically
dominated by ericaceous shrubs such as Andromeda glaucophylla (bog rose¬
mary), Chamaedaphne calyculata (leatherleaf). Rhododendron groenlandicum
(Labrador- tea), and Vaccinium oxycoccos (small cranberry), as well as shrubs in¬
dicative of more minerotrophic conditions, such as Betula pumila (bog birch),
Dasiphora fruticosa (shrubby cinquefoil), Myrica gale (sweet gale), and Rham-
nus alnifolia (alder-leaved buckthorn). Characteristic ground layer associates on
hummocks include Anemone quinquefolia (wood anemone), Carex lasiocarpa
(wiregrass sedge), Doellingeria umbellata (tall flat-topped white aster), Equise-
tum fluviatile (water horsetail), Maianthemum trifolium (false mayflower), Muh-
lenbergia glomerata (marsh wild-timothy), Oclemena nemoralis (bog aster). Os-
2012
THE MICHIGAN BOTANIST
141
munda regalis (royal fern), Pyrola americana (round-leaved pyrola), Sarracenia
purpurea (pitcher-plant), Solidago uliginosa (bog goldenrod), Trichophorum
alpinum (Alpine bulrush), and T. cespitosum (tufted bulrush). In depressions and
flarks, R. acaulis occurs with several additional herbaceous associates, including
Carex livida (livid sedge), Cladium mariscoides (twig-rush), Iris versicolor
(wild blue flag), and Rhynchospora alba (white beak-rush). One other notable
associate is Rubus pubescens (dwarf raspberry), a widespread species that differs
from R. acaulis in its smaller, white flowers, pedicels that bear at least a few
glands, and sharply acute to acuminate leaflets (Voss and Reznicek 2012). This
species occasionally hybridizes with R. acaulis , producing R. xparacaulis ,
which has been collected from Michigan (Henson 682; MICH; determination E.
G. Voss). In total, approximately 220 vascular plant taxa have been documented
from Michigan peatlands that support populations of R. acaulis (MNFI 2013b).
The only quantitative data on vegetative associates of Rubus acaulis in
Michigan was collected by Schultz (1987) at Shingleton Bog, the first docu¬
mented site for dwarf raspberry in Michigan. In all three fen openings studied by
Schultz, Trichophorum cespitosum was the most or secondmost important vas¬
cular plant associate (Table 4). Other important species included Anemone quin-
quefolia, Andromeda glaucophylla, Solidago uliginosa, Vaccinium oxycoccos,
Muhlenbergia glomerata, Oclemena nemoralis. Rhododendron groenlandicum,
Dasiphora fruticosa, and Maianthemum trifolium (Table 4). Sphagnum spp. pro¬
vided at least 20% average cover in all three study sites; only Sphagnum mosses
and Trichophorum cespitosum averaged greater than 10% cover in all three fen
openings (Table 5). Although most of the aforementioned taxa are broadly dis¬
tributed in Upper Michigan wetland communities, their co-occurrence indicates
potential habitat for R. acaulis in the Seney Sand Lake Plain Ecoregion.
Specimen labels and MNFI field survey data do not identify bryophyte asso¬
ciates of Rubus acaulis to the species level. In general, strings (in patterned fens)
and hummocks within minerotrophic fen systems support several Sphagnum
species, including S. angustifolium, S. capillifolium, S. magellanicum, S. re-
TABLE 4. Ten most important vascular and non-vascular plant associates in Rubus acaulis quadrats,
Shingleton Bog, ranked by Importance Value (IV = relative frequency + relative dominance).
Adapted from Schultz (1987).
Big Bog
Species
IV
South Bog
Species
IV
Section 12 Bog
Species
IV
Trichophorum cespitosum
121
Sphagnum spp.
132
Trichophorum cespitosum 144
Rubus acaulis
102
Trichophorum cespitosum
103
Sphagnum spp.
125
Rhamnus alnifolia
88
Rubus acaulis
102
Solidago uliginosa
103
Picea mariana
72
Anemone quinquefolia
101
Rubus acaulis
102
Sphagnum spp.
71
Vaccinium oxycoccos
84
Andromeda glaucophylla
102
Dasiphora fruticosa
70
Alnus incana
71
Anemone quinquefolia
101
Andromeda glaucophylla
69
Rhododendron groenlandicum
69
Vaccinium oxycoccos
100
Maianthemum trifolium
69
Andromeda glaucophylla
67
Lonicera villosa
87
Oclemena nemoralis
67
Oclemena nemoralis
67
Muhlenbergia glomerata
84
Solidago uliginosa
67
Viola spp.
67
Drosera rotundifolia
83
Muhlenbergia glomerata
67
1 42 THE MICHIGAN BOTANIST Vol. 5 1
TABLE 5. Ten vascular and non-vascular plants with the highest percent cover in Rubus acaulis
quadrats, Shingleton Bog. Adapted from Schultz (1987).
Big Bog
South Bog
Section 12 Bog
Species
IV
Species
IV
Species
IV
Trichophorum cespitosum
38
Sphagnum spp.
32
Trichophorum cespitosum
44
Sphagnum spp.
21
Trichophorum cespitosum
19
Sphagnum spp.
25
Gaultheria hispidula
6
Dasiphora fruticosa
5
Picea mariana
5
Picea mariana
6
Alnus incana
5
Thuja occidentalis
5
Thuja occidentalis
6
Picea mariana
4
Solidago uliginosa
3
Rhamnus alnifolia
4
Ilex verticillata
5
Lonicera villosa
3
Dasiphora fruticosa
4
Viburnum cassinoides
3
Carex exilis
2
Trichophorum alpinum
3
Rubus acaulis
2
Rubus acaulis
2
Maianthemum trifolium
2
Carex exilis
2
Andromeda glaucophylla
2
Rubus acaulis
2
Prunella vulgaris
2
Larix laricina
2
Andromeda glaucophylla
2
Rhododendron groenlandicum
2
Menyanthes trifoliata
2
Carex stricta
2
Sarracenia purpurea
2
Thuja occidentalis
2
Carex stricta
2
Cirsium spp.
2
curvum, and occasionally S. warnstorfii (Slaughter and Cohen 2010). The upper
portions of strings and hummocks, which can be highly acidic due to isolation
from groundwater, may support S.fuscum and Polytrichum juniperinum (juniper
polytrichum moss) (Wheeler et al. 1983; Crum 1988). Bryophytes are typically
of low importance in flarks and depressions, but are represented by several
species, including Aulacomnium palustre, Calliergon trifarium, Campylium
polygamum , C. stellatum , Dicranum undulatum , Drepanocladus spp., Poly¬
trichum strictum, Scorpidium scorpioides , several sphagnum mosses (e.g.,
Sphagnum angustifolium, S. cuspidatum, S. majus, S. magellanicum, S. papillo¬
sum , S. recurvum, S. subsecundum, and S. teres), and several liverworts (includ¬
ing Calypogeia spp., Cephalozia spp., Kurzia setacea, and Mylia anomala)
(Wheeler et al. 1983; Madsen 1987; Crum 1988; Janssens 1992). Further surveys
are needed to document microhabitat-specific bryophyte associates of R. acaulis
in Michigan.
CONSERVATION STATUS
Following its discovery in Michigan in 1976, Rubus acaulis was assigned
threatened status in the 1980 listing of state endangered and threatened plant
species. Subsequent to its discovery, R. acaulis was listed as a sensitive species
in the Hiawatha National Forest within USFS Region 9 (USDA Forest Service,
Eastern Region 2002). Over 20 years passed without any new sites being dis¬
covered, and the species was upgraded to endangered status in 1999. Since that
time, several additional populations of R. acaulis have been documented in
Upper Michigan, and a total of eight occurrences are currently tracked by MNFI
(MNFI 2013a). All of these populations appear to have good to excellent viabil-
2012
THE MICHIGAN BOTANIST
143
ity based on area of occupancy, population density, occurrence in relatively
undisturbed habitats, and other factors (MNFI 2013a). The first documented
population in Schoolcraft County occurs within the Shingleton Bog Candidate
Research Natural Area (cRNA) within the Hiawatha National Forest. Establish¬
ment of this area as a RNA would provide formal protection for this R. acaulis
population and its fen habitat. A second population of R. acaulis occurs in the
eastern unit of the Hiawatha National Forest. A third population is partly pro¬
tected within Tahquamenon Falls State Park, and extends into the adjacent Lake
Superior State Forest (LSSF). Four other populations occur within LSSF and ad¬
jacent commercial forest lands, and one population has been documented from
the Escanaba River State Forest (ERSF) (MNFI 2013a). Although the largest and
presumably most viable R. acaulis populations occur within LSSF and ERSF,
only one of these sites (Creighton Marsh) has received a special conservation
designation as an Ecological Reference Area (Cohen et al. 2009), and efforts to
provide additional measures of protection to the other sites have proven contro¬
versial (Casperson 2013; Garmon 2013).
One of the potential challenges to long-term conservation of Rubus acaulis in
Michigan is poor fruit set and, presumably, low levels of outcrossing (Ladyman
2006). Over a three year span, the second author monitored R. acaulis popula¬
tions in Shingleton Bog out of concern for the potential of a newly constructed
logging road to disrupt water flow within the fen and negatively impact the habi¬
tat (Schultz 1987, 1988, 1990). Over that three year-span, the number of stems,
average number of leaves per stem, percent cover, and number of fruits or dried
flowers showed little variability in 18 permanent 1 m2 quadrats (Schultz 1990).
Few fruits were observed in any year, consistent with the documentation of
scarce fruit production at the other Michigan sites (MNFI 2013a). Flower and es¬
pecially fruit production also appears to be poor in the western United States,
suggesting that R. acaulis reproduces primarily through cloning at the edges of
its range (Spackman et al. 1997; Fertig 2000b; Ladyman 2006). Clonal repro¬
duction is a life history strategy that has been demonstrated for several other
edge-of-range plant species, and appears to be associated with a variety of ge¬
netic and environmental factors (Dorken and Eckert 2001; Beatty et al. 2008).
Inferences from studies on R. arcticus ssp. arcticus suggest the lack of fruiting
may be due to the presence of triploid populations, a preponderance of self-in¬
compatible clones, or a lack of pollination (presumably by honeybees and/or
bumblebees) (Tammisola and Ryynanen 1970; Ladyman 2006).
Rubus acaulis is a relict species that occupies peatland habitats that are ex¬
pected to be sensitive to climatic warming and associated changes in water bal¬
ance and nutrient cycling (Galatowitsch et al. 2009; Kost and Lee 2011). A
warmer climate is expected to cause an increase in vascular plant cover, particu¬
larly that of ericaceous shrubs, as a result of lowered water tables and altered nu¬
trient cycling (Weltzin et al. 2003; Galatowitsch et al. 2009; Essl et al. 2012; van
Dijk et al. 2012; Bragazza et al. 2013; Jassey et al. 2013). Sphagnum cover, on
the other hand, is expected to decline. Because Michigan populations of R.
acaulis occur in close association with Sphagnum, a reduction of cover and vigor
of these mosses will likely have detrimental impacts to the species. In addition,
warmer, drier conditions will likely lead to more frequent and severe peatland
144
THE MICHIGAN BOTANIST
Vol. 51
wildfires that threaten to reduce moss cover and organic soils and alter peatland
ecology and vegetation (Galatowitsch et al. 2009). Galatowitsch et al. (2009)
suggest several climate change adaptation strategies to reduce the potential neg¬
ative impacts of climatic warming on peatland habitats, including the prohibition
of drainage projects in the vicinity of peatlands, the prohibition of groundwater
withdrawals in recharge areas of fens, and the control of peat fires. Regardless of
actions taken to mitigate the effects of climate change on peatland ecology, the
long-term persistence of R. acaulis in Michigan appears to be at risk. Canadian
climate change models for R. acaulis put Upper Michigan out of the climatic
range of the taxon under 19 of 20 climate projection scenarios by 2040 and in all
20 scenarios by 2070 (Natural Resources Canada, Canadian Forest Service
2011).
Based on the range of the species, area of occupancy, number of occurrences,
population sizes, condition of habitat, and threats, including climate change (Na-
tureServe 2013), the Endangered Species Technical Committee for Plants should
consider downgrading the conservation status of R. acaulis in Michigan from S 1
(critically imperiled) to S2 (imperiled).
SUGGESTIONS FOR FUTURE WORK
Additional surveys for Rubus acaulis should be conducted throughout Upper
Michigan. Three particular areas are likely to yield additional populations: (1)
the vicinity of Seney National Wildlife Refuge, encompassing much of northern
and eastern Schoolcraft County; (2) the peatlands of northwestern Chippewa
County and northern and eastern Luce County, in the vicinity of Trout Lake,
Eckerman Corner, Hulbert, Newberry, McMillan, and Paradise; and (3) the
vicinity of Cyr Swamp in Marquette County, south of Gwinn and primarily east
of the Escanaba River, with a smaller area of shrub-dominated fen west of the
river. Potential habitat in all three areas is extensive. High resolution aerial im¬
agery of this entire region is widely available and can be used to target likely
fens within these areas. The preponderance of boreal relict species in the Ke¬
weenaw Peninsula and especially Isle Royale (Given and Soper 1981; Marr et al.
2009) and presence of fen habitats (Slavick and Janke 1987; Cohen 2009) sug¬
gests at least a fair probability that R. acaulis may occur in one or both of these
locales, as well. Although R. acaulis may occur elsewhere in western Upper
Michigan, its typical fen habitat is less widely distributed and much less exten¬
sive in this region. At each newly documented R. acaulis site, detailed lists of
vascular plant associates and especially non-vascular plant associates (horn-
worts, liverworts, and mosses) should be recorded to improve our understanding
of R. acaulis habitat in Michigan.
Monitoring of R. acaulis and its habitat should be implemented to track
changes in population size, plant vigor, and fecundity and to document changes
in organic soils, hydrology, and species composition and vegetative structure of
the peatlands it occupies. Because R. acaulis in Michigan is a relict, edge-of-
range species, it is likely to be especially sensitive to climate change and associ-
2012
THE MICHIGAN BOTANIST
145
ated changes in its peatland habitats. Therefore, its population dynamics may
serve as a good indicator of the health of several of the largest peatland ecosys¬
tems in Upper Michigan. Although R. acaulis is considered secure globally,
these peripheral populations may possess genetic distinctiveness that may prove
important to the overall conservation of the taxon, particularly with regard to
adaptive potential in the face of climatic warming (Beatty et al. 2008). With this
in mind, population and genetic studies that address the viability and genotypic
richness of R. acaulis populations in Michigan are suggested in order to better
determine the long-term viability of this species in the state.
ACKNOWLEDGMENTS
Funding for field surveys was provided by Michigan Department of Natural Resources (MDNR)
Wildlife Division, MDNR Forest Resources Division, and MDNR Parks and Recreation Division.
We thank Joshua Cohen and David Cuthrell for their assistance in identifying, documenting, and
mapping Rubus acaulis populations. Tony Reznicek and Beverly Walters confirmed specimen iden¬
tification and entered the records into the Michigan Flora database, respectively. Charlotte Taylor
and Don Henson provided information on the initial discovery and first collections of R. acaulis in
Michigan. Alan Fryday and Alan Prather provided collection data from MSC. Christopher Campbell
provided information on the reported occurrence of R. acaulis in New England. Sally Sanderson pro¬
vided materials on Shingleton Bog Candidate Research Natural Area. Tyler Bassett, Joshua Cohen,
Scott Namestnik, Michael Penskar, Todd Barkman, and an anonymous reviewer provided insightful
comments and suggestions for improvements to the manuscript.
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Office Surveys 1816-1856. Report to the U.S. E.RA. Water Division and the Wildlife Division,
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2012
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149
NOTEWORTHY COLLECTIONS
Michigan
Mikania scandens (L.) Willd.
Asteraceae
Climbing Hempweed
Previous knowledge. Mikania scandens is a vine found in wetlands of the
eastern United States (USDA). It ranges from Florida to Maine and west to
Texas and is considered a noxious weed in Hawaii. It was first reported from
Michigan by C.D. McLouth (1896). The plant was brought to McLouth’s atten¬
tion in 1894, and that September, he observed it growing in the wetlands near the
mouth of the Muskegon River in what is now Muskegon State Game Area
(MSGA), Muskegon County. Inquiries to locals in the area allowed McLouth to
conclude that M. scandens had been present since at least the mid-nineteenth
century. This has remained the only report of M. scandens in Michigan up to the
present (Reznicek et al. 2011). The nearest known populations are about 200
kilometers to the south in northern Indiana (Swink & Wilhelm 1994). Attempts
to relocate this species in the areas described by McLouth have been unsuccess¬
ful, leading to this species being designated as extirpated from Michigan
(Reznicek et al. 2011).
Significance of the Report. This is a significant find as this is the first report
of this species in Michigan since it was originally reported by McLouth in 1896.
I observed three populations, all roughly within 2 kilometers of each other. Only
one of the populations was surveyed and there appeared to be about 50 healthy
individuals setting seed at the coordinates provided on the collection label. The
other populations were in remote areas and occurred in dense Cephalanthus oc¬
cidentals thickets and only a few individuals were observed at each location.
More thorough surveys are needed in the areas between Cedar Creek and the
Muskegon River to accurately describe the condition of the metapopulation.
Diagnostic characters. Mikania scandens is a climbing herb of floodplains
(Fig. 1), often twining up Cephalanthus occidentalis to heights of 3.0 m and oc¬
casionally forming mats over Phalaris arundinacea and Leersia oryzoides. The
leaves are oppositely arranged at swollen nodes, deeply cordate, palmatly
veined, and toothed. The flowers are small, pale-purple to white, and form loose
corymbs bom on axillary peduncles (MNFI 2007). Collected specimens were
observed to be blooming throughout September. M. scandens may be mistaken
for Ageratina altissima as the flowers are somewhat similar. However, M. scan¬
dens is a vine whereas A. altissima is a stout upright herb.
Specimen Citation. Mikania scandens (L.) Willd. Muskegon Co., Michigan:
Muskegon State Game Area. 5 Miles SSW of Twin Lake. Lat. & Long.: 43° 17'
34.2 N, 86° 07' 41.1 " W. Between Cedar Creek and the Muskegon River along
the margins of openings in the Acer saccharinum dominated floodplain forest.
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Vol. 51
FIGURE 1 . Mikania scandens in Muskegon State Game Area, MI. Photo taken by Jesse Lincoln on
Oct 2, 2012.
Primarily climbing on Cephalanthus occidentalis and Fraxinus pennsylvanica.
There seems to be a strong association of M. scandens and C. occidentalis. Also
found occasionally forming mats atop Phalaris arundinacea. Highly localized
but somewhat abundant. The majority of individuals observed were fruiting and
scenesent at the time of collection, 3 October 2012. Jesse M. Lincoln; (MICH).
ACKNOWLEDGMENTS
I would like to thank Dr. Tony Reznicek for his assistance editing this paper and for confirming
the identity of Mikania scandens in the field. Todd Barkman also reviewed the article and provided
helpful comments for improvements. Valerie Campbell assisted in the collection of samples and her
efforts were appreciated.
LITERATURE CITED
McClouth, C.D. (1896). Mikania scandens Willd. Asa Gray Bulletin 4: 68
Michigan Natural Features Inventory. 2007. Rare Species Explorer (Web Application). Available on¬
line at http://mnfi.anr.msu.edu/explorer [Accessed Nov 2, 2012]
Reznicek, A. A., E.G. Voss, & B.S. Walters. (2011). University of Michigan. Web. Available at
http://michiganflora.net/species. aspx?id=408 (Accessed 23 October 2012).
Swink, F. & G. Wilhelm. 1994. Plants of the Chicago region, 4th ed. Indiana Academy of Science for
Morton Arboretum, Indianapolis, Indiana, xiv + 921 pp.
- Jesse M. Lincoln
1510 Colorado SE, Grand Rapids, MI 49507
2012
THE MICHIGAN BOTANIST
151
BOOK REVIEW
Simmons, Eric. 2009. Darwin Slept Here. The Overlook Press. New York,
NY. 258 pp. ISBN 978-1-59020-220-3 $24.00. (Hardbound)
If a person is given a pairing game and they have placed Charles Darwin’s name
in the mix, most people would pair Darwin’s name with evolution. I know I
would. Because of Darwin’s theory of evolution by natural selection, his name
has become synonymous with evolution. People who saw me reading this book
thought that I was reading about Darwin’s theory because the title had his name
on it. However, after reading the book, I have a glimpse of the real Darwin most
people do not see. Most photos depicts Darwin as an old man, with “Santa
Clause beard”, quiet, wrinkled and maybe boring. But who is Darwin, really?
Was he always this “old man” or was he once a vibrant young man ready to face
the unknown?
Eric Simmons, a young man from San Francisco decided to journey and ex¬
perience first-hand how a young explorer Darwin might feel. He travelled to
South America and he compared his experience with that of Darwin’s, based on
his diary notes: “The Voyage of the Beagle”. Instead of unravelling the escapade
of an “old wrinkled man”, Simmons discovered the escapades of a young, fun-
loving Darwin whose adventures were exhilarating also to Simmons himself.
Simmons went to every place that Darwin visited based on his records and
walked where he perceived Darwin walked. Simmons attempted to see what
Darwin would have seen if he was looking at the same glorious views. Like Dar¬
win, the author viewed many things; from rheas to a beach filled with fossils in
Port San Julian. Reading this book, Simmons’ eyes become our eyes as to what
Darwin saw, but instead of us seeing it physically, we experience it through his
words. Simmons described the side of Darwin that most people never see, the
adventurous young man who goes places to learn. A most interesting part is that
Simmons exhibited the same adventurous spirit that Darwin had.
The book was divided into three parts: exploration of the industrialized South
America where Simmons explored the now changed South America, revolution
of the people where the author focused on the people of South America of pre¬
sent compared to the people during Darwin’s era, and Darwin’s discoveries
where Simmons looked at the most important things that Darwin discovered
while exploring South America. Each of these parts gives an idea as to what dif¬
ferences Simmons experienced as compared to Darwin’s time. At the end of the
book, Simmons shares his moving experience of sleeping at the same house
where Darwin slept and how he imagined the difference between his time and
Darwin’s time. The contrast of time may have been very different but the expe¬
riences that Simmons and Darwin experienced were irreplaceable, regardless of
the time difference.
So who was the young Darwin? Read this book and it will give you an idea
of who young Darwin was with less facial hair and how passionate Darwin was
152
THE MICHIGAN BOTANIST
Vol. 51
about his travels in South America discovering new species and places. Sim¬
mons’ book will give you a detailed and exhilarating experience that will keep
you wanting to know what happens next. This book will show you the different
side of Darwin, the man usually depicted as “the man with the long-white
beard”.
Rozaleth Jeanne Role, Graduate Student
Biology Department,
Andrews University,
Berrien Springs, MI 49104-0410
role@andrews.edu
New York Botanical Garden Library
3 5185 00269 4394
INSTRUCTIONS TO AUTHORS
1. Create text in 12-point Times New Roman font and double space paragraphs throughout. Papers
should be organized as follows: Title, Author(s) and address(es), Abstract with up to 5 keywords.
Introduction, Materials and Methods, Results, Discussion, Acknowledgements, Literature Cited,
Tables, Figure Legends, and Figures. Sections may be omitted if not relevant. All pages should
be numbered. Please contact the editor regarding any questions related to formatting.
2. For noteworthy collections, manuscripts should be formatted as described in The Michigan
Botanist , volume 27(3) p. 90. A brief description of the formatting follows. The following title,
“Noteworthy collections”, should begin each submitted manuscript followed on the next line by
the State or Province for the species reported. The next line should list the taxon of interest using
the following format: Species Author(s) (Family). Common name. The rest of the manuscript
should include the following named sections: Previous knowledge. Significance of the report,
Diagnostic characters (if desired), Specimen citations, and Literature cited. Each of these sec¬
tions are largely self explanatory; however, “specimen citations” should include the relevant
label data from the voucher specimen(s) including location data, collector(s), collection number,
etc. Also please include which herbarium the specimen(s) is deposited in using the Index
Herbariorum acronym. The manuscript should end with the name and address of the author(s).
3. Letters to the Editor can be formatted as general text without the specific sections listed above.
However, literature cited and any tables or figures should be formatted as described below.
4. Please create tables using either a tab delimited format or a spreadsheet using Excel or other sim¬
ilar program. Each table is to be submitted as a separate file. Table captions should be placed at
the top of the table. Any footnotes should appear at the bottom of the table. Please do not insert
tables within the body of the text.
5. Send each figure as a separate file in a high-resolution format — eps, jpg, or tif. Figures like bar
graphs that gain their meaning with color won’t work — use coarse-grained cross-hatching, etc.
Create figure legends as a separate text file, and the typesetter will insert them as appropriate.
Please do not insert the figure in the body of the text file.
6. Citations: Please verify that all references cited in the text are present in the literature cited sec¬
tion and vice versa. Citations within the text should list the author’s last name and publication
year (e. g. Smith 1990). For works with more than 2 authors, use “et al.”, and separate multiple
citations with a semicolon.
7. Literature Cited: List citations alphabetically by author’s last name. Author names are to be list¬
ed with surname first, followed by initials (e. g. Smith, E. B.). Separate author’s initials with a
single space. The year of publication should appear in parentheses immediately before the title
of the citation. The entire journal name or book title should be spelled out. Please put a space
after the colon when citing volume number and page numbers.
8. Italicize all scientific names. Voucher specimens must be cited for floristic works or any other
relevant study. Papers citing plant records without documenting vouchers are generally not
acceptable.
9. Manuscripts may be submitted electronically to the email address of the editor. Printed versions
of manuscripts may also be submitted in which case three copies should be provided. All manu¬
scripts will be reviewed by at least two referees. A more complete set of instructions is available
at http://www.michbot.org/publications/Botanist/instruct_authors.htm.
CONTENTS
Suitability of Cardamine concatenata (Michx.) SW. as an Indicator
Species on the Ottawa National Forest
Margaret M. Fox, Susan J. Trull, and Blair Orr 121
The Distribution, Ecology, and Conservation Status of
Rubus acaulis Michx. (Dwarf or Arctic Raspberry) in Michigan
Bradford S. Slaughter and Jan Schultz 133
Noteworthy Collections
Jesse M. Lincoln 149
Book Review 151
On the cover: No longer extirpated! Mikania scandens in Muskegon State Game Area, MI
where it has been rediscovered more than 100 years after its original discovery in the state.
Photo taken by Jesse Lincoln on Oct 2, 2012.