RHODORA The Journal of the New England Botanical Club CONTENTS Atlas of the flora of New England: pteridophytes and gymnosperms. Ray elo and David E. Boufford First ae of Uncinia macrolepis and U. tenuis (Cyperaceae) in Colombia ald A. Whee w Colombian sites for U. paludosa. Gerald A. CLC e nesee 80 The Paralychnophora group of Eremanthus (Vernonieae: Asteraceae). Harold Robinson 85 NEW ENGLAND NOTES A flora of the vascular and non-vascular plants of Nantucket, Tucker- nuck, and Muskeget Islands. Peter W. Dunwiddie and Bruce A. Sorrie 94 Seleria reticularis (Cyperaceae) new to New Hampshire. Daniel D. rduto 99 BOOK REVIEWS The Vascular and Non-Vascular Flora of Nantucket, Tuckernuck and Muskeget Islands 103 Native Trees, Shrubs and Woody Vines of Cape Cod and the Islands ...... 105 NEBC MEETING NEWS 107 eae EMENTS C Graduate Student Research Award 110 1996 Jesse M. Greenman Award 111 Ronald L. Stuckey Endowment Fund 112 Information for Contributors 114 NEBC Membership Form 115 NEBC Officers and Council Members inside back cover Vol. 98 Winter, 1996 No. 893 Issued: April 23, 1997 The New England Botanical Club, Inc. 22 Divinity Avenue, Cambridge, Massachusetts 02138 RHODORA JANET R. SULLIVAN, Editor-in-Chief Department of Plant Biology, University of New Hampshire, Durham, NH 03824 MARGARET P. BOGLE, Managing Editor Department of Plant Biology, University of New Hampshire, Durham, NH 03824 Associate Editors HAROLD G. BROTZMAN THOMAS D. LEE CHRISTOPHER S. CAMPBELL LESLIE J. MEHRHOFF DAVID S. CONANT THOMAS MIONE GARRETT E. CROW LISA A. STANDLEY NANCY M. EYSTER-SMITH K. N. GANDHI—Latin diagnoses and nomenclature RHODORA (ISSN 0035-4902). Published four times a year (January, April, July, and October) by The New England Botanical Club, 810 East 10th St., Lawrence, KS 66044 and printed by Allen Press, Inc., 1041 New Hampshire St., Lawrence, KS 66044-0368. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send _ address changes to RHODORA, PO. Box 1897, Lawrence, KS 66044-8897. RHODORA is a journal of botany devoted primarily to the flora of North America. Monographs or scientific papers concerned with systemat- ics, floristics, ecology, paleobotany, or conservation biology of the flora of North America or floristically related areas will be considered. SUBSCRIPTIONS: $75 per calendar year, net, postpaid, in funds pay- e at par in United States currency. Remittances payable to RHO- DORA. Send to RHODORA, P.O. Box 1897, Lawrence, KS 66044- 8897 MEMBERSHIPS: Regular $35; Family $45; Student $25. Application form printed herein NEBC WEB SITE: Information about The New England Botanical Club, its history, officers and councillors, herbarium, monthly meet- ings and special events, annual graduate student award, and the jour- nal RHODORA is available at http://www.herbaria.harvard.edu/nebc/ BACK ISSUES: Information on be soueeel of back issues should be addressed to Dr. Cathy A. Paris, Department of Botany, University of Vermont, Burlington, VI 05405-0086. E-mail: cparis@ moose.uvm.edu ADDRESS CHANGES: In order to receive the next number of RHO- ORA, changes must be received by the business office prior to the first day of January, April, July, or October. INFORMATION FOR CONTRIBUTORS: See contents. Submit manu- scripts to the Editor-in-Chief. © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). RHODORA, Vol. 98, No. 893, pp. 1-79, 1996 ATLAS OF THE FLORA OF NEW ENGLAND PTERIDOPHYTES AND GYMNOSPERMS RAY ANGELO New England Botanical Club, 22 Divinity aa Cambridge, Massachusetts 02138-202 aes ie BOUFFORD Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Ma pena 02138-2020 ABSTRACT. Dot maps are provided to depict the distribution at the county level os a pteridophytes (Lycopodiophyta, erent dis Seeercetlacs and gymnosperms (Coniferophyta) growing outside of cultivation in six New E gland states of the northeastern United States. The 171 taxa pee subspecies, varieties and hybrids, but not forms) are mapped at the county level based on —o in the a herbaria of Maine, New ai aes Vermont, Massachu- for names used in recent manuals and floras for the area, habitat and chromo- some information and common names also are provided Key Words: Flora, New England, atlas, distribution, pteridophytes, gymnosperms This article is the first in a series that will present the distributions of the vascular flora of New England in the form of dot distribution maps at the county level. New England com- prises the six northeastern states of the United States: Maine, New Hampshire, Vermont, Massachusetts, Rhode Island and Connecticut. These states range in latitude from 47°28'N and in longitude from 67°W to 73°45'W. The elevation of the region extends from sea level to 1,898 m (6,228 ft.) at the summit of Mt. Washington, New Hampshire, the highest moun- tain in eastern North America outside of North Carolina. The total area of the region is 172,515 km* (66,608 mi.”) including bodies of water; the total land area is 163,157 km? (62,995 mi.”). The effects of continental glaciers, which covered the entire region several times during the Pleistocene, and the last of which did not retreat until about 15,000 years ago, are still clearly evident throughout the area. The state by state statistics on size and elevation are shown in Table 1 (Merriam-Webster, Inc. 1984 Table 1. Total England states. Rhodora | Vol. 98 area, land area, and range in elevation for the six New Total Area Land Area Elevation Range Maine New Hampshire Vermont Massachusetts Rhode Island Connecticut 86,028 km? 12,973 km? 80,117 km? 23,395 km? 24,020 km? 20,287 km° 2,725 km°* 12.613 km? O—1,606 m O—1,898 m 29—1,339 m O—1,064 m 0-248 m 0-725 m Ye The more southerly states of Massachusetts, Rhode Island, and Connecticut are densely populated (the least densely populated of these three states, Connecticut, had an average of 262 persons per km’ in 1990; Hoffman 1992). The northern states of Maine, New Hampshire, and Vermont are mostly rural (the most densely populated of these three states, New Hampshire, had an average of 48 persons per km* in 1990; Hoffman 1992). This work encompasses all vascular plants (pteridophytes and spermatophytes) at the rank of species, subspecies, and variety growing outside of cultivation in the New England states. Hy- brids also will be included, but forms and other ranks below the level of variety will not. The dots are based primarily on voucher specimens in the herbaria of New England representing reproduc- ing populations, or plants persisting after cultivation when it is uncertain that they are actually naturalized. This first installment includes the pteridophytes (Lycopodiophyta, Equisetophyta, Poly- podiophyta) and gymnosperms (Coniferophyta). Future accounts will treat the distribution of angiosperms. We intend to gather this series of articles, together with addi- tional background material, into a separate volume upon comple- tion of all the maps, and to make it available on the World Wide Web as various parts are finished. It is our hope that, in the mean- time, these articles will stimulate additional field work to supple- ment the distributions portrayed in the maps. The New England Botanical Club herbarium, which has proven to be the most 1m- portant resource for this project, is especially eager to receive specimens documenting range extensions. We also would like to be informed of such specimens in other herbaria. Similarly, be- cause the atlas of the New England flora will be continuously updated as new information becomes available, we are eager to receive notification of published corrections of cytological infor- 1996] Angelo and Boufford—Atlas of New England Flora 3 mation and new, documented chromosome counts for taxa in the New England flora. MATERIALS AND METHODS This atlas grew out of an inventory of the New England Bo- tanical Club vascular plant herbarium. The purpose of the inven- tory, conducted at the county level, was to serve as a filter for the processing of a large number of unmounted specimens being considered for accession. Consequently, the core of the database for this atlas consists of specimens in the herbarium of vascular plants of the New England Botanical Club (NEBC). The NEBC herbarium, which limits its geographical scope to the New En- gland states, presently comprises more than 251,000 specimens and is the largest collection of plants for the New England re- gion. In the mid 1980s the herbarium inventory was expanded into an atlas project, since data from the NEBC herbarium essentially depict the distributions of the vascular plants of New England. Additional data from specimens in other major herbaria with large holdings of New England plants have allowed us to portray the distributions more completely. The data represented currently by specimens in New England herbaria best depict vascular plant distributions when displayed at the county level. Attempting to display such data by the exact geographical coordinates of each collection locality, or even at the township level, distorts the actual distributions. By such de- tailed mapping, patterns tend to reflect concentrated collecting along major highways and clustering near popular resort areas or botanical hot spots. This was evident from some of the detailed distribution maps produced about 15 years ago by the plant dis- tribution committee (now defunct) of the New England Botanical Club. The distortions portrayed by such detailed mapping are re- solved by using data only at the county level. The counties of New England range in size from Bristol County, Rhode Island (65 km*, or 25 mi.”), to Aroostook County, Maine (17,666 km’, or 6821 mi.°—land only). The great majority of counties, however, are between 1300 km? (S500 mi.”) and 3900 km (1500 mi.”). To reduce the size disparity at the large end, and to depict distributions more evenly, the largest Maine counties are subdivided. Penobscot, Piscataquis, Somerset, and Oxford Coun- ties are divided arbitrarily into roughly equal-sized, north and 4 Rhodora [ Vol. 98 south sectors along township boundaries. Aroostook County, the argest county, is divided into three sectors. Cods County, New Hampshire, is divided into north and south sectors. The division of Coés County isolates the White Mountains, which harbor an alpine flora associated mostly with New England’s highest peak, Mount Washington. Figure | shows the location of states and counties of New England and the subdivisions of the largest coun- ties. Data were collected primarily from herbarium specimens of major herbaria in New England. The herbaria used to compile data for this work were those of the New England Botanical Club (NEBC), Harvard University (A, AMES, GH), University of Maine (MAINE), University of Massachusetts (MASS), University of New Hampshire (NHA), University of Rhode Island (KIRI), University of Vermont (vT), and Brown University (BRU). Records from the herbaria at the University of Connecticut (CONN), Connecticut Bo- tanical Society (NCBS), and Yale University (YU) were accessed from data compiled by Leslie Mehrhoff. Lastly, the notebooks of Harry Ahles at the University of Massachusetts were consulted. Ahles visited these same herbaria, plus one or two others, in the 1960s and 1970s to gather locality information at the county level. We obtained some records from systematic treatments where the information on voucher specimens was specific enough to pro- — vide county data and the name of the herbarium where the speci- mens were deposited. In recording the collection data no attempt was made to verify the accuracy of the identification of every specimen, but obvious misidentifications were annotated with corrections. Even though the basis for a particular dot might be a misidentified specimen, for plants common within New England, our view is that the taxon most likely occurs within the county. Specimens documenting dis- junct or marginal occurrences (or otherwise rare, endangered or threatened taxa) usually have received greater scrutiny, because of a heightened interest in rare plants nationwide, and are less likely to be misidentified. The data were entered into a simple computer database using one of several letters in the county field as a code to indicate the herbarium in which the voucher specimen ts deposited. The letter code indicates the first herbarium where a voucher specimen was seen. No letter codes were added for subsequent voucher speci- mens at other herbaria, because our primary objective has been to plot the distribution of the taxon, not to inventory the holdings 1996] Angelo and Boufford—Atlas of New England Flora 5 of any particular herbarium. While such information might be of value for some purposes, we felt that our time at other herbaria was best devoted to searching for data not found in the New En- gland Botanical Club herbarium, or in each subsequently searched herbarium, to fill in gaps in distribution. The data in the database were converted to dot distribution maps using a method devised by Angelo (1994). A more detailed and technical description of the mapping methodology will appear as an appendix in the final volume. Interested parties may obtain a draft of the appendix upon request to the first author. TAXONOMY AND FORMAT The taxonomy and nomenclature adopted for this work essen- tially follow the Flora of North America (Flora of North America Editorial Committee 1993). With the exception of the arrangement for the major divisions within the pteridophytes (Lycopodiophyta, Equisetophyta, and Polypodiophyta), which follows the Flora of North America (Flora of North America Editorial Committee 1993) sequence, the families are ordered alphabetically. The gen- era are alphabetical within families, as are species within genera. Named and unnamed hybrid taxa are placed alphabetically at the end of the genus. Unnamed hybrids combine the names of the progenitors alphabetically by epithet. Species deemed to be introduced, that is, not native in New England at the time of European contact with North America, are indicated by the use of all upper case type for the scientific name. No single source of information was used in determining intro- duced species. For most taxa there is little dispute as to nativity. Where differences of opinion exist in the literature, we have used our own judgment based on the evidence available. A common name is supplied when it appears to be a name in general use. Names used in New England have been given pref- erence. Cited chromosome numbers are taken from indices prepared by Cave (1958a, 1958b, 1959a, 1959b, 1960, 1961, 1962, 1963, 1964, 1965), Goldblatt (1981, 1984, 1985, 1988), Goldblatt and Johnson (1990, 1991, 1994), Moore (1973, 1974, 1977), Ornduff (1967, 1968, 1969), and reports by Takamiya and Kurita (1983), Taylor (1970), Tryon (1978), Tryon and Tryon (1982), and Wag- ner (1971). Very few of the counts are based on material from 6 Rhodora [Vol. 98 New England, but instead reflect counts made from throughout the range of the taxon. The habitat data are distillations from a variety of sources aug- mented by our own field observations. An attempt was made to indicate habitat information as it applies to a particular taxon in New England rather than to the entire range of the taxon. Synonymy is provided primarily with respect to names used in the standard manuals covering New England published from 1950 onward, including Fernald (1950), Gleason (1952), Gleason and Cronquist (1963, 1991), and Seymour (1969a, 1982). A selection of references, including many not cited above, is provided. This list consists of the standard manuals published from 1950 to date, regional and county floras and checklists pro- duced after 1950, and what we believe to be the key articles and books on the pteridophytes and gymnosperms pertinent to New England. We would appreciate being notified of papers we may have overlooked. LYCOPODIOPHYTA ISOETACEAE Isoétes acadiensis Kott—Acadian Quillwort (Figure 2). 2n = 44; the report of 2n = 44 (Kott and Britton 1980) for /. macro- spora Durieu forma hieroglyphica (A. A. Eaton) N. E. Pfeiffer was later attributed (Kott 1981) to /. acadiensis. Shallow wa- ter of ponds and slow moving streams. Isoétes echinospora Durieu—Braun’s Quillwort (Figure 2). 27 = 22, 33. Shallow water and shores of slightly acidic lakes, ponds, and streams. [/. echinospora var. braunii (Durieu) En- gelmann; /. echinospora var. muricata (Durieu) Engelmann; I. muricata Durieu] Isoétes engelmannii A. Braun—Engelmann’s Quillwort (Figure 2).n = 11; 2n = 22. Shallow water of ponds, lakes, streams, or in muddy ditches. Isoétes lacustris Linnaeus—Lake Quillwort (Figure 2). 27 = [10. Shores and shallow water of slightly acidic ponds, lakes, and streams. [/. macrospora Durieu; /. hieroglyphica A. A. Eaton] 1996] Angelo and Boufford—Atlas of New England Flora a [soétes prototypus D. M. Britton (Figure 3). 2n = 22. Deep wa- ter of cold, acidic lakes. Isoétes riparia Engelmann ex A. Braun—Riverbank Quillwort (Figure 3). 2n = 44. Muddy or sandy shores (including tidal shores and estuaries), usually of rivers. [/. riparia var. cana- densis Engelmann; /. saccharata Engelmann var. amesii A. A. Eaton] Isoétes tuckermanii A. Braun ex Engelmann—Tuckerman’s Quill- wort (Figure 3). 2” = 44. Pond and lake margins, shores (fre- quently tidal), usually submersed in quiet water. —Isoétes Hybrids— Isoétes X eatonii Dodge (Figure 3). [= /. echinospora Durieu X I. engelmannii A. Braun; /. * gravesii A. A. Eaton; /. eatonii Dodge var. gravesii (A. A. Eaton) Clute] Isoétes X harveyi A. A. Eaton. This hybrid was recognized only recently by Britton (1991). Herbarium specimens annotated as this hybrid are from southern Penobscot Co., Maine (us); Somerset Co., Maine (MAINE); and Mt. Desert Island, Maine (MO). Since the holdings of NEBC have not been examined criti- cally for this hybrid, it is not mapped. [= /. lacustris Lin- naeus J. tuckermanii A. Braun] LYCOPODIACEAE Diphasiastrum complanatum (Linnaeus) Holub—Northern Run- ning Pine (Figure 4). n = 22-24, 23; 2n = 40, 44, 46, 48. Dry, open coniferous or mixed forests. [Lycopodium com- Planatum Linnaeus | Diphasiastrum digitatum (Dillenius ex A. Braun) Holub—South- ern Running Pine (Figure 4). 7 = 23; 2n = 46. Usually dry woods, thickets, or open fields, in acid soils. [Lycopodium complanatum Linnaeus var. flabelliforme Fernald] Diphasiastrum sitchense (Ruprecht) Holub—Sitka Clubmoss isu 4). n = 23; 2n = 46. Alpine meadows, open rocky 8 Rhodora [Vol. 98 barrens, conifer woods (rarely). [Lycopodium sitchense Ru- precht] Diphasiastrum tristachyum (Pursh) Holub—Ground Cedar (Fig- ure 4). n = 23; 2n = 46. Acid soils in open conifer or oak woods, thickets. [Lycopodium tristachyum Pursh]| —Diphasiastrum Hybrids— Diphasiastrum alpinum (Linnaeus) Holub < Diphasiastrum com- planatum (Linnaeus) Holub (Figure 5). Diphasiastrum complanatum (Linnaeus) Holub * Diphasiastrum digitatum (Dillenius ex A. Braun) Holub (Figure 5). Diphasiastrum digitatum (Dillenius ex A. Braun) Holub < Dipha- siastrum X sabinifolium (Willdenow) Holub (Figure 5). Diphasiastrum X* habereri (House) Holub (Figure 5). [= D. digi- tatum (Dillenius ex A. Braun) Holub X D. tristachyum (Pursh) Holub; Lycopodium X habereri House] Diphasiastrum X issleri (Rouy) Holub (Figure 6). [= D. alpi- num (Linnaeus) Holub X* D. tristachyum (Pursh) Holub; Ly- copodium X issleri (Rouy) Lawalree | Diphasiastrum X_ sabinifolium (Willdenow) Holub—Savin- ee Clubmoss (Figure 6). 2” = 46. Woods (often subal- pine spruce), thickets, clearings. [= D. sitchense (Ruprecht) Holub X D. tristachyum (Pursh) Holub; Lycopodium sabini- folium Willdenow } Diphasiastrum X zeilleri (Rouy) Holub (Figure 6). [= D. complanatum (Linnaeus) Holub xX D. ¢tristachyum (Pursh) Holub] Huperzia appalachiana Beitel & Mickel (Figure 6). Cliffs, talus slopes, damp, acidic rocks in alpine zones. [Specimens mapped are from the herbaria of the New England Botanical Club and Harvard University. | 1996] Angelo and Boufford—Atlas of New England Flora 9 Huperzia lucidula (Michaux) Trevisan—Shining Clubmoss (Fig- ure 7). 2 = 67; 2n = 134. Cool, moist woods in rich and acid soils. [Lycopodium lucidulum Michaux ] Huperzia selago (Linnaeus) Bernhardi ex Schrank & Martius— Mountain Clubmoss (Figure 7). 7 = 1361]; 21 = 268. Damp rocks, barrens, cold woods, in mountainous regions. [H. se- lago var. densa Trevisan; Lycopodium selago Linnaeus, Ly- copodium selago Linnaeus var. appressum Desvaux] —Huperzia Hybrids— Huperzia appalachiana Beitel & Mickel * Huperzia lucidula (Michaux) Trevisan (Figure 7). Huperzia appalachiana Beitel & Mickel * Huperzia selago (Lin- naeus) Bernhardi ex Schrank & Martius (Figure 7). Huperzia X buttersii (Abbe) Kartesz & Gandhi (Figure 8). [= H. lucidula (Michaux) Trevisan X H. selago (Linnaeus) Bernhardi ex Schrank & Martius] Lycopodiella alopecuroides (Linnaeus) Cranfill—Foxtail Club- moss (Figure 8). 2n = 156. Bogs, marshes, ditches, sandy or peaty shores, wet barrens, in acid soils. [Lycopodium alope- curoides Linnaeus | Lycopodiella| appressa (Chapman) Cranfill—Appressed Bog Clubmoss (Figure 8). 27 = 156. Damp, acid or peaty soils in the open. [Lycopodium appressum (Chapman) Lloyd & Un- derwood; Lycopodium inundatum Linnaeus var. bigelovii Tuckerman; Lycopodium inundatum Linnaeus var. robustum R. J. Eaton] Lycopodiella inundata (Linnaeus) Holub—Bog Clubmoss (Figure 8). nm = 78; 2n = 156. Damp, acid, sandy or peaty soils in the open. [Lycopodium inundatum Linnaeus] Lycopodiella margueritae J. G. Bruce, W. H. Wagner & Beitel (Figure 9), 2” = 312. 10 Rhodora [ Vol. 98 Lycopodium annotinum Linnaeus—Stiff Clubmoss (Figure 9). n = 34, 6811; 2n = ca. 50, 66, 68. Woods, cool shaded thick- ets, exposed rocky or peaty habitats at higher elevations. [L. annotinum var. acrifolium Fernald; L. annotinum var. alpestre C. Hartman; L. annotinum var. pungens (Bachelot de la Py- laie) Desvaux ] ca clavatum Linnaeus—Staghorn Clubmoss (Figure 9). 3411, 341, 341V, 68 (69, 70); 2n = 34, 44, 68, io, 136. Open, usually dry, woods and thickets, clearings, exposed rocky situations, in acid soils. [L. clavatum var. sub- cs remotum Victorin] en ycopodium dendroideum Michaux—Tree Clubmoss (Figure 9). 2n = 68. Dry, open woods and clearings. [L. obscurum Lin- naeus var. dendroideum (Michaux) D. C. Eaton] ycopodium hickeyi W. H. Wagner, Beitel & R. C. Moran—Hick- ey’s Clubmoss (Figure 10). 2” = 68. Mainly deciduous woods and thickets, usually in acid soils. [L. obscurum Linnaeus var. isophyllum Hickey] Oy = ycopodium lagopus (Laestadius ex C. Hartman) G. Zinserling ex Kuzeneva-Prochorova (Figure 10). 27 = 68. Grassy fields and openings in second-growth woods. [L. clavatum Linnaeus var. megastachyon Fernald & Bissell; L. clavatum Linnaeus var. monostachyon Hooker & Greville] Distribution data are primarily from the herbaria of the New England Botanical Club and Harvard University. ycopodium obscurum Linnaeus—Tree Clubmoss (Figure 10). n = 34, 3411; 2n = 68. Rich woods, clearings, edges of boggy forests, usually in acid soils. i —Lycopodium Hybrids— Lycopodium dendroideum Michaux & Lycopodium obscurum Linnaeus (Figure 10). Pseudolycopodiella caroliniana (Linnaeus) Holub—Carolina Clubmoss (Figure 11). 2 = 35, 68, 70, 115; 27 = 70, 78, 140. 1996] Angelo and Boufford—Atlas of New England Flora 11 Damp, open sands and peat in acid soils. [Lycopodium caro- linianum Linnaeus | SELAGINELLACEAE Selaginella apoda (Linnaeus) Spring—Meadow Spikemoss (Fig- ure 11). 2n = 18. Meadows, stream banks, moist lawns, wet rocks, often in circumneutral soils. [S. eclipes W. R. Buck] Selaginella rupestris (Linnaeus) Spring—Rock Spikemoss (Fig- ure 11). 27 = 18. Exposed, dry rocks or packed sands. Selaginella selaginoides (Linnaeus) Palisot de Beauvois ex Martius & Schrank—Northern Spikemoss (Figure 11). 2n = 18. Damp shores and banks, bogs, springs, often calcar- cous. EQUISETOPHYTA EQUISETACEAE Equisetum arvense Linnaeus—Field Horsetail (Figure 12). nm = 108; 2n = 200+, 216. Open, low ground, roadsides, damp, open woods and thickets. [F. arvense var. boreale (Bongard) Ruprecht] Equisetum fluviatile Linnaeus—Water Horsetail (Figure 12). = 108; 2n = 216. Shores, shallow water of pond and river mar- gins, swales. Equisetum hyemale Linnaeus subsp. affine (Engelmann) Calder & Roy L. Taylor—Common Scouring Rush (Figure 12). n = 108. Sandy banks and shores, alluvium, moist wooded slopes, roadsides. [F. hyemale var. affine (Engelmann) A. A. Eaton; FE. hyemale var. pseudohyemale (Farwell) C. V. Mor- ton] Equisetum palustre Linnaeus—Marsh Horsetail (Figure 12). 1 = 108. Shores, flooded meadows, swamps, and marshes. 2 Rhodora [Vol. 98 al pratense Ehrhart—Meadow Horsetail (Figure 13). n = 108; 2n = 216. Rich slopes in humus-rich neutral soils, meadows, alluvial thickets. Equisetum scirpoides Michaux—Dwarf Scouring Rush (Figure 13). n = 108. Well-drained slopes, in cool woods, frequently under 7Tsuga, often growing in moss. Equisetum sylvaticum Linnaeus—Wood Horsetail (Figure 13). 2n = 216. Cool moist woods, swamps, around springs. [F. sylvaticum var. pauciramosum Milde] Equisetum variegatum Schleicher ex F. Weber & D. Mohr—Var- iegated Horsetail (Figure 13). 27 = 216. Riverbanks, shores, bog margins, moist gravel, and damp, calcareous sands. —Equisetum Hybrids— Equisetum X ferrissti Clute (Figure 14). Sandy shores, embank- ments, roadsides. |= FE. hyemale Linnaeus < EF. laevigatum A. Braun; FE. hyemale Linnaeus var. intermedium A. A. Eaton: E. hyemale Linnaeus var, e/atum (Engelmann) C. V. Morton] Equisetum X litorale Kuhlewein ex Ruprecht—Shore Horsetail (Figure 14). Shores, ditches, meadows. [= FEF. arvense Linnaeus X EF. fluviatile Linnaeus | Equisetum X mackaii (Newman) Brichan (Figure 14). 2n = 432. Riverbanks, shores, bog margins, gravelly areas, damp, cal- careous sands. [= EF. hyemale Linnaeus < E. variegatum Schleicher ex F. Weber & D. Mohr: FE. X trachyodon (A. Braun) Koch; -. variegatum Schleicher ex F. Weber & D. Mohr var. jesupii A. A. Eaton| POLY PODIOPHYTA ASPLENIACEAE Asplenium montanum Willdenow—Mountain Spleenwort (Figure I = 72. Moist, sheltered crevices of chiefly non- calcareous cliffs and ledges. 1996] Angelo and Boufford—Atlas of New England Flora 13 Asplenium platyneuron (Linnaeus) Britton, Sterns & Poggen- burg—Ebony Spleenwort (Figure 15). n = 36, 3611. Well- drained, rocky, circumneutral soils on slopes and ledges in de- ciduous forests (occasionally on crumbling mortar). [A. platyneuron var. incisum (E. C. Howe) B. L. Robinson] Asplenium rhizophyllum Linnaeus—Walking Fern (Figure 15). n = 36. Shaded, basic or circumneutral rocks, rarely on earth or at tree bases. [Camptosorus rhizophyllus (Linnaeus) Link] Asplenium ruta-muraria Linnaeus—Wall-rue (Figure 15). 7 = 36, 72; 2n = 144. Shaded crevices of calcareous cliffs and ledges. [A. cryptolepis Fernald; A. ruta-muraria var. cryptolepis (Fer- nald) Wherry ] Asplenium trichomanes Linnaeus—Maidenhair Spleenwort (Fig- ure 15). n = 36, 36II, 361] + 361, 72, 81, 108, 144; 2n = 72, 144, ca. 216. Shaded crevices of mainly calcareous rocks (oc- casionally on crumbling mortar). [A. tichomanes subsp. quadrivalens D. E. Meyer] Asplenium trichomanes-ramosum Linnaeus—Green Spleenwort (Figure 16). m = 36; 2n = 72. Shaded, moist crevices and talus of calcareous and serpentine rocks and outcrops. [A. viride Hudson | —Asplenium Hybrids— Asplenium X clermontae Syme (Figure 16). 3x = 108. [= ruta-muraria Linnaeus < A. trichomanes Linnaeus | Asplenium X ebenoides R. R. Scott—Scott’s Spleenwort (Figure 16). 2n = 72, 144. [= A. platyneuron (Linnaeus) Britton, Sterns & Poggenburg x A. rhizophyllum Linnaeus; X Asplenosorus ebenoides (R. R. Scott) Wherry] AZOLLACEAE AZOLLA CAROLINIANA Willdenow—Mosquito Fern (Figure 16). n = 48; 2n = 44, 66. Floating in quiet waters. From south of our range. 14 Rhodora | Vol. 98 BLECHNACEAE Woodwardia areolata (Linnaeus) T. Moore—Netted Chain Fern (Figure 17). 2 = 35. Swamps and boggy woods in acid solls. Woodwardia virginica (Linnaeus) Smith—Virginia Chain Fern (Figure 17). n = 35, ca. 36. Acid swamps, bogs, and wet, wooded bottoms. DENNSTAEDTIACEAE Dennstaedtia punctilobula (Michaux) T. Moore—Hay-scented Fern (Figure 17). nm = ca. 33, 34. Open woods, roadsides, stream banks, pastures and rocky slopes in sterile soils. Preridium aquilinum (Linnaeus) Kuhn subsp. /atiusculum (Des- vaux) Hultén—Bracken (Figure 17). 7 = 52; 2n = 104. Clear- ings, pastures, burns, thickets and woods, usually in dry, ster- ile soils. [P. aquilinum var. latiusculum (Desvaux) L. Underwood ex A. Heller] Preridium aquilinum (Linnaeus) Kuhn subsp. pseudocaudatum (Clute) Hultén—Bracken (Figure 18). Same habitats as in subsp. /atiusculum. [P. aquilinum var. pseudocaudatum (Clute) A. Heller] DRYOPTERIDACEAE Athyrium filix-femina (Linnaeus) Roth ex Mertens subsp. angus- tum (Willdenow) R. T. Clausen—Lady Fern (Figure 18). 1 = 40; 2n = 80. Moist woods, thickets, ravines, fields, meadows and swamps. [A. filix-femina var. angustum (Willdenow) G. Lawson; A. filix-femina var. michauxii (Sprengel) Farwell] Athyrium filix-femina (Linnaeus) Roth ex Mertens subsp. asple- nioides (Michaux) Hultén—Southern Lady Fern (Figure 18). 2n = 80. Moist woods, thickets, swamps, meadows. [A. filix- femina var. asplenioides (Michaux) Farwell] 1996] Angelo and Boufford—Atlas of New England Flora 15 Cystopteris bulbifera (Linnaeus) Bernhardi—Bulblet Fern (Fig- ure 18). n = 42; 2n = 42. Calcareous, usually moist, ledges, steep banks, rocky slopes, and ravines. Cystopteris fragilis (Linnaeus) Bernhardi—Fragile Fern (Figure 19). n = 42, 84, 126; 2n = 168. Moist rocks, rocky (mostly wooded) slopes. Cystopteris laurentiana (Weatherby) Blasdell. n = 126; 2n = 252. Cliffs, often calcareous. [C. fragilis (Linnaeus) Bernhardi var. laurentiana Weatherby] Distribution data were not collected for this taxon, since it was generally not distinguished from C. fragilis (Linnaeus) Bernhardi in New England herbaria. Cystopteris protrusa (Weatherby) Blasdell—Southern Bladder Fern (Figure 19). 7 = 42, 63. Moist, often calcareous, wooded slopes, rocky banks, and alluvium. [C. fragilis (Linnaeus) Bernhardi var. protrusa Weatherby | Cystopteris tenuis (Michaux) Desvaux—Mackay’s Fragile Fern (Figure 19). n = 84; 2n = 168. Moist, shaded rocks, rocky (mostly wooded) slopes, alluvium, open woods. [C. fragilis (Linnaeus) Bernhardi var. mackayi G. Lawson] —Cystopteris Hybrids— Cystopteris fragilis (Linnaeus) Bernhardi X Cystopteris tenuis (Michaux) Desvaux (Figure 19). n = 5311 + 601 + 2II, 531 + 561 + 2 I, 521 + 641. Dry to moist cliffs and talus, moist ravines. Information for this hybrid is from Paler and Barrington (1995). Deparia acrostichoides (Swartz) M. Kato—Silvery Spleenwort (Figure 20). n = 40; 2n = 80. Rich moist woods, bottom- lands, shaded slopes. [Athyrium thelypterioides (Michaux) Desvaux; Diplazium acrostichoides (Swartz) Butters] Diplazium pycnocarpon (Sprengel) M. Broun—Narrow-leaved Spleenwort (Figure 20). » = 40. Moist, mostly calcareous, wooded slopes, ravines, and bottoms. [Athyrium pycnocarpon (Sprengel) Tidestrom; Hc us pycnocarpos (Sprengel) Small ex Pichi-Sermolli] 16 Rhodora [ Vol. 98 Dryopteris campyloptera (Kunze) Clarkson—Mountain Wood Fern (Figure 20). 7 = 82; 2n = 164. Moist, cool woods, thick- ets, and banks. [D. austriaca (Jacquin) Woynar ex Schinz & Thellung var. austriaca,; D. spinulosa (O. F. Mueller) Watt var. americana (Fischer ex Kunze) Fernald; D. spinulosa (O. F. Mueller) Watt var. concordiana (Davenport) Eastman; D. aus- triaca (Jacquin) Woynar ex Schinz & Thellung var. concordi- ana (Davenport) Morton] Dryopteris carthusiana (Villars) H. P. Fuchs—Spinulose Wood Fern (Figure 20). n = 82; 2n = ca. 160, 164. Low, moist (oc- casionally dry) woods and thickets, stream banks, swamps. [D. austriaca (Jacquin) Woynar ex Schinz & Thellung var. spinu- losa (O. F. Mueller) Fiori; D. spinulosa (O. F. Mueller) Watt] Dryopterts cristata (Linnaeus) A. Gray—Crested Wood Fern (Fig- ure 21). n = 41, 82, 123; 2n = 82, 164. Swamps, wet woods, marshes, boggy thickets. Dr yoprers filix-mas (Linnaeus) Schott—Male Fern (Figure 21). n = 82, 83; 2n = 41, 164. Rocky wooded slopes, cold ra- vines, rich woods, upland pastures in chiefly calcareous soil, trap, or slate. Dryopteris fragrans (Linnaeus) Schott—Fragrant Cliff Fern (Fig- ure 21). n = 41, 42. Dry, often north-facing and calcareous, cliffs, ledges, talus, and rocky banks. [D. fragrans var. remo- fluscula Komarovy | Dryopteris goldiana (Hooker ex Goldie) A. Gray—Goldie’s Fern (Figure 21). = 41; 2n = 41. Rich, moist woods, rocky slopes (mostly calcareous), ravines. Dryopteris intermedia (Muhlenberg ex Willdenow) A. Gray— Glandular Wood Fern (Figure 22). n = 41; 2” = 82. Moist, rocky woods and moist thickets. [D. austriaca (Jacquin) Woy- nar ex Schinz & Thellung var. intermedia (Muhlenberg ex Willdenow) C. V. Morton; D. spinulosa (O. F. Mueller) Watt var. intermedia (Muhlenberg ex Willdenow) L. Underwood] Dryopteris pest sai s (Linnaeus) A. Gray—Marginal Shield Fern (Figure 22). n = 41. Rocky, usually rich, wooded slopes, ra- vines. 1996] Angelo and Boufford—Atlas of New England Flora 17 —Dryopteris Hybrids— Dryopteris * benedictii Wherry (Figure 22). [= D. carthusiana (Villars) H. P. Fuchs * D. X clintoniana (D.C. Eaton) Dowell] Dryopteris * boottii (Tuckerman) Underwood—Boott’s Wood Fern (Figure 22). n = 82, 123; 2n = 123. Moist woods and thickets in mildly acid soils. [= D. cristata (Linnaeus) A. Gray < D. intermedia (Willdenow) A. Gray]. This common and widespread hybrid is not treated in Flora of North America (Flora of North America Editorial Committee 1993). Dryopteris * burgessii Boivin (Figure 23). [= D. X* clintoniana (D. C. Eaton) Dowell * D. marginalis (Linnaeus) A. Gray] Dryopteris campyloptera (Kunze) Clarkson X Dryopteris inter- media (Muhlenberg ex Willdenow) A. Gray (Figure 23). Dryopteris campyloptera (Kunze) Clarkson < Dryopteris margi- nalis (Linnaeus) A. Gray (Figure 23). Dryopteris carthusiana (Villars) H. P. Fuchs * Dryopteris goldi- ana (Hooker ex Goldie) A. Gray (Figure 23). There is one known collection for this taxon. Other collections so identi- fied are variations of D. goldiana (report in preparation, W. H. Wagner, Jr., pers. comm.). Dryopteris X clintoniana (D. C. Eaton) Dowell—Clinton’s Wood Fern (Figure 24). n = 123; 2n = 246. Wet (rarely only moist), often circumneutral woods and swamps. [=D. cristata (Lin- naeus) A. Gray < D. goldiana (Hooker ex Goldie) A. Gray; D. cristata (Linnaeus) A. Gray var. clintoniana (D. C. Eaton) L. Underwood] Dryopteris X clintoniana (D. C. Eaton) Dowell * Dryopteris cristata (Linnaeus) A. Gray (Figure 24). Dryopteris X clintoniana (D. C. Eaton) Dowell * Dryopteris filix-mas (Linnaeus) Schott (Figure 24). Dryopteris X clintoniana (D. C. Eaton) Dowell * Dryopteris goldiana (Hooker ex Goldie) A. Gray (Figure 24). 18 Rhodora | Vol. 98 Dryopteris X dowellii (Farwell) Wherry (Figure 25). [= D. X clintoniana (D. C. Eaton) Dowell X D. intermedia (Muhlen- berg ex Willdenow) A. Gray] Dryopteris filix-mas (Linnaeus) Schott * Dryopteris marginalis (Linnaeus) A. Gray (Figure 25). Dryopteris goldiana (Hooker ex Goldie) A. Gray * Dryopteris intermedia (Muhlenberg ex Willdenow) A. Gray (Figure 25). Dryopteris intermedia (Muhlenberg ex Willdenow) A. Gray X Dryopteris marginalis (Linnaeus) A. Gray (Figure 25). Dryopteris X neowherryi W. H. Wagner (Figure 26). [= D. goldi- ana (Hooker ex Goldie) A. Gray * D. marginalis (Linnaeus) A. Gray] Dryopteris X pittsfordensis Slosson (Figure 26). [= D. carthusi- ana (Villars) H. P. Fuchs * D. marginalis (Linnaeus) A. Gray | Dryopteris X slossoniae Wherry ex Lellinger (Figure 26). [= D. cristata (Linnaeus) A. Gray < D. marginalis (Linnaeus) A. Gray | Dryopteris X triploidea Wherry (Figure 26). 37 = 123. Rocky or swampy (often rich) woods. [= D. carthusiana (Villars) H. P. Fuchs * D. intermedia (Muhlenberg ex Willdenow) A. Gray; D. austriaca (Jacquin) Woynar ex Schinz & Thellung var. fructuosa (Gilbert) C. V. Morton; D. spinulosa (O. F. Mueller) Watt var. fructuosa (Gilbert) Trudell]. This common and widespread hybrid is not treated in Flora of North America (Flora of North America Editorial Committee 1993). Dryopteris * uliginosa (A. Braun ex Dowell) Druce (Figure 27). [= D. carthusiana (Villars) H. P. Fuchs * D. cristata (Lin- naeus) A. Gray | Gymnocarpium dryopteris (Linnaeus) Newman—Oak Fern (Fig- ure 27). n = 40, ca. 73, 80; 2n = 80, 160. Cool, moist, often rocky, woods and banks. [Dryopteris disjuncta (Ruprecht) C. V. Morton (misapplied) |] 1996] Angelo and Boufford—Atlas of New England Flora 19 Gymnocarpium jessoense (Koidzumi) Koidzumi subsp. parvu- lum Sarvela—Nahanni Oak Fern (Figure 27). 2n = 80, 160. Acid or neutral substrates of cool talus slopes, cliffs, and outcrops. This taxon also is reported to occur in Connecticut in Flora of North America (Flora of North America Editorial Committee 1993), but neither specimens nor specific citations of specimens in the literature have been seen. Kathleen Pryer (pers. comm.) indicates that the Connecticut report may be based on a mislabeled speci- men. Matteuccia struthiopteris (Linnaeus) Todaro—Ostrich Fern (Fig- ure 27). n = 39, 40; 2n = 78. Alluvium, bottomland thickets, swamps, and moist woods, often in circumneutral soils. [M. pensylvanica (Willdenow) Raymond; M.. struthiopteris var. pensylvanica (Willdenow) C. V. Morton; Preretis pensylvanica (Willdenow) Fernald] Onoclea sensibilis Linnaeus—Sensitive Fern (Figure 28). n = 37; n = 74. Swamps, low woods, alluvial thickets, meadows, in neutral to slightly acid soils. [O. sensibilis var. obtusilobata (Schkuhr) Torrey] Polystichum acrostichoides (Michaux) Schott—Christmas Fern (Figure 28). n = 41; 27 = 82. Rocky woods and slopes, oc- casionally swamps. Polystichum braunii (Spenner) Féee—Braun’s Holly Fern (Figure 28). nm = 82; 2n = 164. Cool, damp, rocky woods, ravines, and banks in circumneutral soils. [P. braunii var. purshii Fer- nald; P. braunti subsp. purshii (Fernald) Calder & Roy L. Tay- lor] —Polystichum Hybrids— Polystichum X potteri Barrington (Figure 28). n = 711 + 26H, 74.-F 231) 4 TT. 7/1 a 23h, 79). 2 221k Bll 21, 85 = 191], 871 + I8IL. [= P. acrostichoides (Michaux) Schott X P. brauni (Spenner) Fée] 20 Rhodora [ Vol. 98 Woodsia alpina (Bolton) S. F. Gray—Northern Woodsia (Figure 29). n = 82: 2n = 156, 164. Cool, calcareous (sometimes slaty) rock crevices. [W. a/pina var. bellit Lawson] Woodsia glabella R. Brown ex Richardson—Smooth Woodsia (Figure 29). n = 39, ca. 40, ca. 82; 2n = 78, ca. 80. Moist, shaded crevices of calcareous rocks. Woodsia ilvensis (Linnaeus) R. Brown—Rusty Woodsia (Figure 29). n = 39, 41; 2n = 78, 82. Dry, often exposed, cliffs and talus. Woodsia obtusa (Sprengel) Torrey—Blunt-lobed Woodsia (Figure 29). n = 76. Shaded ledges and rocky woods, mostly in cal- careous soils. —Woodsia Hybrids— Woodsia X* gracilis (Lawson) Butters (Figure 30). [= W. alpina (Bolton) S. F. Gray * W. i/vensis (Linnaeus) R. Brown] HYMENOPHYLLACEAE Trichomanes intricatum Farrar—Appalachian Bristle Fern (Fig- ure 30). Sheltered, moist, noncalcareous rock crevices and other protected places in deep ravines and gorges. Known only as gametophytes. LYGODIACEAE Lygodium palmatum (Bernhardi) Swartz—Climbing Fern (Figure 30). 2 = 30. Damp thickets and woods, stream banks, in acid soils. MARSILEACEAE MARSILEA QUADRIFOLIA Linnaeus—Water Clover (Figure 30). n = 20; 2n = 40. Lakes, ponds, and quiet streams. From Europe. 1996] Angelo and Boufford—Atlas of New England Flora 21 OPHIOGLOSSACEAE Botrychium dissectum Sprengel—Cut-leaved Grape Fern (Figure 31). n = 45. Woods, open thickets, clearings, sandy barrens. |B. dissectum Sprengel var. obliquum (Muhlenberg ex Will- denow) Clute] Botrychium lanceolatum (S. G. Gmelin) Angstrém subsp. angus- tisegmentum (Pease & A. H. Moore) R. T. Clausen —Triangle Grape Fern (Figure 31). 1 = 45; 2n = 90. Moist, cool, rich woods, swamp margins, meadows, peaty slopes, clearings. [B. lanceolatum var. angustisegmentum Pease & A. H. Moore] Botrychium lunaria (Linnaeus) Swartz—Common Moonwort (Figure 31). n = 45; 2n = 90. Open, gravelly or rocky slopes, pastures, meadows, shores, chiefly in calcareous soils. Botrychium matricartifolium (D6) A. Braun ex W. D. J. Koch— Daisyleaf Grape Fern (Figure 31). 2 = 90. Rich, usually moist, deciduous woods and thickets. Botrychium minganense Victorin—Mingan Moonwort (Figure 32). n = 90; 2n = 180. Habitats similar to those of B. luna- ria. [B. lunaria (Linnaeus) Swartz forma minganense (Vic- torin) Clute; B. /unaria var. minganense (Victorin) Dole] Botrychium multifidum (S. G. Gmelin) Ruprecht—Leathery Grape Fern (Figure 32). 1 = 45; 2n = 90, 180. Peaty, sandy, or grav- elly slopes, sparse woods, thickets, clearings, usually in acid soils. [B. multifidum var. intermedium (D. C. Eaton) Farwell] Botrychium oneidense (Gilbert) House—Oneida Grape Fern. 1 = 45; 2n = 90. Moist, acidic woods and swamps. [B. dissectum Sprengel var. oneidense (Gilbert) Farwell; B. multifidum (S. G. Gmelin) Ruprecht var. oneidense (Gilbert) Farwell]. Distribution data were not collected for this taxon as it was inadequately distinguished from B. dissectum Sprengel and B. multifidum (S. G. Gmelin) Ruprecht in New England her- baria. Botrychium pallidum W. H. Wagner. 2n = 90. A species reported in Flora of North America (Flora of North America Editorial Rhodora [ Vol. 98 i) N Committee 1993) as occurring in Maine; no voucher seen, no specific voucher for New England cited. Botrychium rugulosum W. H. Wagner (Figure 32). 1 = 45. Open fields and secondary forests. We found no annotated vouch- ers for this relatively recently recognized taxon in New En- gland herbaria. The distribution data are taken from W. H. Wagener, Jr., and F. S. Wagner (1982). Botrychium simplex E. Hitchcock—Least Grape Fern (Figure 32). n = 45. Meadows, moist woods, edges of swamps and pas- tures, open shores, usually in poor soils. [B. simplex var. tene- brosum (A. A. Eaton) R. T. Clausen] Botrychium virginianum (Linnaeus) Swartz—Rattlesnake Fern (Figure 33). n = 92; 2n = 184. Rich woods and thickets in neutral to slightly acid soils. [B. virginianum var. europaeum Angstrom] Ophioglossum pusillum Rafinesque—Northern Adder’s Tongue (Figure 33). n = 480; 27 = 960. Peaty or grassy swales, mead- ows, damp thickets, damp sands. [O. vu/gatum Linnaeus var. pseudopodum (S. F. Blake) Farwell] OSMUNDACEAE Osmunda cinnamomea Linnaeus—Cinnamon Fern (Figure 33). = 22. Swamps, damp thickets, stream banks, and other moist, shaded places in acid soils. [O. cinnamomea var. glan- dulosa Waters] Osmunda claytoniana Linnaeus—lInterrupted Fern (Figure 33). n = 22; 2n = 44. Moist woods, margins of swamps, moist thickets, occasionally in the open, in acid or neutral soils. Osmunda_ regalis Linnaeus’ var. spectabilis (Willdenow) A. Gray—Royal Fern (Figure 34). n = 22; 2n = 44. Swamps, peaty thickets, shores, meadows, bogs, mostly in acid soils. 1996] Angelo and Boufford—Atlas of New England Flora = 23 —Osmunda Hybrids— Osmunda cinnamomea Linnaeus * Osmunda claytoniana Lin- naeus (Figure 34). Osmunda X ruggti R. M. Tryon (Figure 34). [= O. claytoniana Linnaeus X O. regalis Linnaeus var. spectabilis (Willdenow) A. Gray] POLYPODIACEAE Polypodium appalachianum Haufler & Windham—Appalachian Polypody (Figure 34). 2 = 37; 2n = 74. Cliffs and rocky slopes. [Data are from the herbaria of the New England Bo- tanical Club and Harvard University. | Polypodium virginianum Linnaeus—Common Polypody (Figure 35). n = 74; 2n = 148. On rocks, cliffs, tree bases, rocky (occasionally sandy) slopes, usually in acid soils. [P. vulgare Linnaeus var. virginianum (Linnaeus) D. C. Eaton] PTERIDACEAE Adiantum aleuticum (Ruprecht) Paris—Aleutian Maidenhair Fern (Figure 35). n = 29. Serpentine barrens. [A. pedatum Linnaeus var. aleuticum Ruprecht; A. pedatum subsp. calderi Cody | ADIANTUM HISPIDULUM Swartz. n = 171. A species from Asia reported in Flora of North America (Flora of North America Editorial Committee 1993) as escaped in Connecti- cut; no voucher seen, no specific voucher for New England cited. Adiantum pedatum Linnaeus—Common Maidenhair Fern (Figure 35). n = 29, 581, 5711 + 21, 56 + R4, 561] + chlV, 541 + 211; 2n = 58, 60. Rich, deciduous woods, in circumneutral or basic soil. 24 Rhodora [ Vol. 98 —Adiantum Hybrids— Adiantum X viridimontanum Paris (Figure 35). n = 58. [= A. aleuticum (Ruprecht) Paris * A. pedatum Linnaeus] Cheilanthes lanosa (Michaux) D. C. Eaton—Hairy Lip Fern (Fig- ure 36). n = 30. Rocks and cliffs, calcareous or siliceous. [C. vestita (Sprengel) Swartz] Cryptogramma. stelleri (S. G. Gmelin) Prantl—Slender Cliff- brake (Figure 36). n = 30. Shaded, moist, calcareous cliffs and rock ledges, typically in coniferous woods. Pellaea atropurpurea (Linnaeus) Link—Purple Cliff-brake (Fig- ure 36). 2 = 87. Dry, exposed, calcareous rocks (occasionally on crumbling mortar), trap rock. Pellaea glabella Mettenius ex Kuhn—Smooth Cliff-brake (Fig- ure 36). n = 116. Dry, calcareous rocks and cliffs. SALVINIACEAE SALVINIA NATANS (Linnaeus) Allioni—Floating Fern (Figure 37). n = 18; 2n = 18, 36. On still waters of ponds or slow streams. From Europe and Asia. [Records from New England are likely to be misidentified as S. rotundifolia Willdenow (Nauman 1993)]. This species is mentioned, but not treated in Flora of North America (Flora of North America Editorial Committee 1993). THELYPTERIDACEAE Phegopteris connectilis (Michaux) Watt—Long Beech Fern (Figure 37). n = 60, 90; 27 = 90. Rich, damp woods, cool, shaded, rocky banks, moist cliffs. [Dryopteris phegopteris (Linnaeus) C. Christensen; Thelypteris phegopteris (Linnaeus) Slosson]} 1996] Angelo and Boufford—Atlas of New England Flora = 25 Phegopteris hexagonoptera (Michaux) Fée—Broad Beech Fern (Figure 37). 27 = 60. Rich woods and thickets. [Dryopteris hexagonoptera’ (Michaux) C. Christensen; Thelypteris hexagonoptera (Michaux) Nieuwland] Thelypteris noveboracensis (Linnaeus) Nieuwland—New York Fern (Figure 37). n = 27. Moist woods and_ thickets. [Dryopteris noveboracensis (Linnaeus) A. Gray] Thelypteris palustris Schott var. pubescens (Lawson) Fernald— Marsh Fern (Figure 38). 2 = 35. Swamps, low damp thick- ets, meadows, marshes, bogs, along ditches and streams. [Dryopteris thelypteris (Linnaeus) A. Gray var. pubescens (Lawson) Weatherby | Thelypteris simulata (Davenport) Nieuwland—Massachusetts Fern (Figure 38). n = 64. Swampy or boggy woods and thick- ets, on knolls in bogs, in acid soils. [Dryopteris simulata Dav- enport] CONIFEROPHYTA CUPRESSACEAE Chamaecyparis thyoides (Linnaeus) Britton, Sterns & Poggen- burg—Atlantic White Cedar (Figure 38). Swamps, wet thick- ets. Juniperus communis Linnaeus var. depressa Pursh—Ground Ju- niper (Figure 38). 27 = 22. Poor soils in open areas, espe- cially old pastures and ledges, persisting in recent woodlands. [Larger individuals have been misidentified as var. commu- nis. | Juniperus horizontalis Moench—Creeping Juniper (Figure 39). n = 22. Ledges, sandy or rocky banks. Juniperus virginiana Linnaeus—Eastern Red Cedar (Figure 39). Mostly dry, open sites (typically old fields), rocky slopes, bor- 26 Rhodora [ Vol. 98 ders of salt marshes, persisting in recent woodlands. [J/. vir- giniana var. crebra Fernald & Griscom] —Juniperus Hybrids— Juniperus communis Linnaeus var. depressa Pursh X Juniperus virginiana Linnaeus (Figure 39). Juniperus horizontalis Moench X Juniperus virginiana Linnaeus (Figure 39). Thuja occidentalis Linnaeus—Arbor-vitae (Figure 40). 2n = 22. Swamps, rocky banks of ponds and rivers, old pastures, in cal- careous soils. PINACEAE Abies balsamea (Linnaeus) Miller—Balsam Fir (Figure 40). 2n = 24. Damp, rich, cool woods, swamps, mountain slopes. [A. balsamea var. phanerolepis Fernald] LARIX DECIDUA Miller—European Larch (Figure 40). 2n = 24. Near habitations in moist woods and clearings. From Europe. Larix laricina (Du Roi) K. Koch—Tamarack (Figure 40). n = 12; 2n = 24. Northward in poorly drained lowlands, swamps, bogs, shaded hillsides, sometimes to tree line on mountain slopes; southward in cold swamps and bogs. PICEA ABIES (Linnaeus) H. Karsten—Norway Spruce (Fig- ure 41). 2 = 12; 2n = 24. Woods and hedgerows. From Eu- rope. Picea glauca (Moench) Voss—White Spruce (Figure 41). 2 = 12; 2n = 24. Moist woods, dry sandy soils, old pastures, rocky slopes, exposed hilltops, ocean cliffs. 1996] Angelo and Boufford—Atlas of New England Flora = 27 Picea mariana (Miller) Britton, Sterns & Poggenburg—Black Spruce (Figure 41). 2 = 12; 217 = 24. Northward on moun- tain slopes, rocky hillsides, high banks of streams, bogs; southward in sphagnum bogs and swamps. Picea rubens Sargent—Red Spruce (Figure 41). 2 = 12; 2n = 24. Rocky, well-drained, upland woods, mountain slopes, oc- casionally in swamps (southward). Pinus banksiana Lambert—Jack Pine (Figure 42). 2n = 18, 24. Sterile sand or rocky soils, rocky slopes and ledges, es- caped from plantations at roadsides and bog margins (south- ward). PINUS MUGO Turra—Swiss Mountain Pine (Figure 42). 2n = 24. Old pastures. From Europe. PINUS MUGO is not treated in Flora of North America (Flora of North America Editorial Committee 1993). PINUS NIGRA Arnott—Austrian Pine (Figure 42). 2n = 18, 24. From Europe. PINUS N/GRA is mentioned, but not treated in Flora of North America (Flora of North America Editorial Committee 1993). Pinus resinosa Aiton—Red Pine (Figure 42). 2n = 24. Sandy or gravelly soil in dry woods and barrens, mountain slopes, ledges, occasionally in peat bogs. Pinus rigida Miller—Pitch Pine (Figure 43). 2n = 24. Dry, ster- ile, sandy or gravelly soils, rocky slopes, occasionally in cold swamps and bogs. Pinus strobus Linnaeus—Eastern White Pine (Figure 43). 1 = 12; 2n = 24. Sandy or fertile, well-drained soils, but tolerating wet ground (rarely in bogs). PINUS SYLVESTRIS Linnaeus—Scotch Pine (Figure 43). 1 = 12; 2n = 24. Roadsides, fallow fields, dry woods. From Eurasia. Our plants are apparently all var. SYLVESTRIS. 28 Rhodora [ Vol. 98 PINUS THUNBERGI Parlatore—Japanese Black Pine (Figure 43). n = 12; 2n = 24. Dry sandy soils. From Asia. PINUS THUNBERGI/ its mentioned, but not treated in Flora of North America (Flora of North America Editorial Committee 1993). Tsuga canadensis (Linnaeus) Carri¢re—Eastern Hemlock (Figure 44). n = 12; 2n = 24. Cool moist slopes, ravines, moist woods, swamps, mucky soils near ponds and streams. TAXACEAE Taxus canadensis Marshall—American Yew (Figure 44). 2 = 24. Damp, rich, or swampy woods, mostly in the shade of coni- fers, ravines and near streams, occasionally on moist, rocky slopes at higher elevations. TAXUS CUSPIDATA Siebold & Zuccarini—Japanese Yew (Fig- ure 44). From Asia. TAXUS CUSPIDATA is mentioned but not treated in Flora of North America (Flora of North America Editorial Committee 1993). ACKNOWLEDGMENTS. We thank the curators and directors of the herbaria of Brown University, Harvard University, the University of Maine, University of Massachusetts, the University of New Hampshire, the University of Rhode Island, and the University of Vermont for allowing us access to their collections. We particu- larly appreciate the kindness of David Barrington, Chris Campbell and Karen Searcy for allowing use of the collections in their care outside of normal hours of operation. We are grateful also to Karen Searcy for allowing access to the notebooks of Harry E. Ahles at the University of Massachusetts. Leslie J. Mehrhoff was very generous in sharing data on pteridophytes and gymnosperms that he has collected from the herbaria at the University of Con- necticut and Yale University. James Hinds assisted by checking the University of Maine herbarium for some discrepancies in the Harry Ahles data. Thomas Vining, at the University of Maine, checked additional specimen data for us at that institution. David Barrington, David S. Conant, and James Hickey made many help- ful comments on various versions of the manuscript and Warren H. “Herb” Wagner was particularly helpful and encouraging. Lastly, we thank Bruce A. Sorrie for collecting data from the Harvard University herbaria in the early stages of this project. 1996] Angelo and Boufford—Atlas of New England Flora = 29 NEW HAMPSHIRE VERMONT Lis oem MASSACHUSETTS i a =e Bee BLOCK Si CONNECTICUT RHODE ISLAND Figure 1. Key map for counties of the New England states (and Mt. Desert Island, Maine and Block Island, Rhode Island and arbitrary divisions of larger Maine counties and Coés County, New Hampshire). 30 Rhodora [ Vol. 98 1996] Angelo and Boufford—Atlas of New England Flora = 31 Isoétes tuckermanii Isoétes X eatonii Figure 3. Distribution maps for /soétes prototypus, I. riparia, 1. tuackerma- ni, and I. X eatonit. ao Rhodora [ Vol. 98 Diphasiastrum complanatum Diphasiastrum digitatum Diphasiastrum sitchense Diphasiastrum tristachyum Figure 4. Distribution maps for Diphasiastrum complanatum, D. digita- m, D. sitcher tu ise, and D. tristachyum. 1996] Angelo and Boufford—Atlas of New England Flora = 33 Diphasiastrum alpinum Diphasiastrum complanatum X D. complanatum X D. digitatum Diphasiastrum digitatum Diphasiastrum X habereri X D. X sabinifolium Figure 5. Distribution maps for Diphasiastrum alpinum * D. complana- tum, D. complanatum X D. digitatum, D. digitatum * D. & sabinifolium, and D. X habereri. 34 Rhodora [ Vol. 98 Diphasiastrum X issleri Diphasiastrum X sabinifolium Diphasiastrum X zeilleri Figure 6. Distribution maps for Diphasiastrum & issleri, D. X sabinifo- lium, zeillert, and Huperzia appalachiana D. X 1996] Angelo and Boufford—Atlas of New England Flora = 35 Huperzia appalachiana Huperzia appalachiana X lucidula selago Figure 7. Distribution maps for Huperzia lucidula, H. selago, H. appala- chiana . ducidula, and H. appalachiana * H. selago. 36 Rhodora [| Vol. 98 Lycopodiella appressa Lycopodiella inundata Figure 8. Distribution maps for Huperzia X buttersti, Lycopodiella alo- pecuroides, L. appressa, and L. tnundata. 1996] Angelo and Boufford—Atlas of New England Flora = 37 Lycopodium clavatum Lycopodium dendroideum Figure 9. Distribution maps for Lycopodiella margueritae, Lycopodium unnotinum, L. clavatum, and L. dendroideum. 38 Rhodora | Vol. 98 1996] Angelo and Boufford—Atlas of New England Flora 39 Selaginella rupestris Selaginella selaginoides Figure 11. Distribution maps for Pseudolycopodiella caroliniana, Selag- inella apoda, S. rupestris, and S. selaginoides. 40 Rhodora [ Vol. 98 Equisetum hyemale subsp. affine Figure 12. Distribution maps for Eguisetum arvense, FE. fluviatile, E. hyemale subsp. affine, and E. palustre. 1996] Angelo and Boufford—Atlas of New England Flora 41 42 Rhodora [ Vol. 98 1996] Angelo and Boufford—Atlas of New England Flora 43 Asplenium platyne uron Asplenium rhizophyllum Asplenium ruta-muraria Asplenium trichomanes ure 15. Distribution maps for Asplenium platyneuron, A. rhizophyllum, A. ruta-muraria, and A. trichomanes. 44 Rhodora [ Vol. 98 Asplenium trichomanes-ramosum Asplenium X clermontae AZOLLA CAROLINIA NA Figure 16. Distribution maps for Asplenium trichomanes-ramo clermontae, A. * ebenoides, and AZOLLA CAROLINIANA. sum, A, X 1996] Angelo and Boufford—Atlas of New England Flora = 45 Dennstaedtia punctilobula Pteridium aquilinum subsp. latiusculum Figure 17. Distribution maps for Woodwardia areolata, W. virgin Dennstaedtia punctilobula, and Pteridium aquilinum subsp. latiusculum. 46 Rhodora [ Vol. 98 Pteridium aquilinum Athyrium filix-femina subsp. pseudocaudatum subsp. angustum Athyrium filix-femina Cystopteris bulbifera subsp. asplenioides Figure 18. Distribution maps for Preridium aquilinum subsp. pseudocau- datum, Athyrium filix-femina subsp. angustum, A, filix-femina subsp. asple- nioides, and Cystopteris bulbifera. 1996] Angelo and Boufford—Atlas of New England Flora = 47 Cystopteris fragilis Cystopteris protrusa Cystopteris fragilis X C. tenuis Figure 19. Distribution maps for Cystopteris fragilis, C. protrusa, C. tenuis, and C. fragilis * C. tenuis. 48 Rhodora [ Vol. 98 opteris campyloptera Figure 20. Distribution maps for Deparia acrostichoides, Diplazium pyc- nocarpon, Dryopteris campyloptera, and D. carthusiana 1996] Angelo and Boufford—Atlas of New England Flora 49 Dryopteris fragrans Dryopteris goldiana Figure 21. Distribution maps for Dryopterts cristata, D. filix-mas, D. fra- grans, and D. goldiana. 50 Rhodora [ Vol. 98 1996] Angelo and Boufford—Atlas of New England Flora = 51 Dryopteris X burgessii Dryopteris campyloptera X D. intermedia Dryopteris campyloptera Dryopteris carthusiana x D. marginali xX D. goldiana Figure 23. Distribution maps for Dryopteris X< burgessii, D , loptera X D. intermedia, D. campyloptera * D. marginalis, and D. carthu- siana X D. goldiana. 2 Rhodora [| Vol. 98 Dryopteris X clintoniana Dryopteris X clintoniana x D. filix-mas xX D. goldiana Figure 24. Distribution maps for Dryopteris * clintoniana, D. X clinto- niana X D. cristata, D. X clintoniana X D. filix-mas, and D. X clintoniana X D. goldiana. 1996] Angelo and Boufford—Atlas of New England Flora 53 Dryopteris goldiana X D. intermedia Dryopteris intermedia X D. marginalis Figure 25. Distribution maps for Dryopteris * dowellii, D. filix-mas x D. marginalis, D. goldiana X D. intermedia, and D. intermedia * D. mar- ginalis. 54 Rhodora [| Vol. 98 Dryopteris X neowherryi Dryopteris X pittsfordensis Dryopteris X triploidea Figure 26. Distribution maps for Dryopteris X neowherryi, D. X pittsfor- pnsis, D. & slossoniae, and D. X triploidea. 1996] Angelo and Boufford—Atlas of New England Flora = 55 Gymnocarpium jessoense Matteuccia struthiopteris subsp. parvulum Figure 27. Distribution maps for Dryopteris * uliginosa, Gymnoc ‘arpium dryopteris, G. jessoense subsp. parvulum, and Matteuccia struthiopteris. 56 Rhodora [ Vol. 98 Polystichum braunii Polystichum X potteri Figure 28. Distribution maps for Onoclea sensibilis, Polystichum acrosti- choides, P. braunti, and P. X pottert. 1996] Angelo and Boufford—Atlas of New England Flora = 57 Woodsia ilvensis Woodsia obtusa Figure 29. Distribution maps for Woodsia alpina, W. glabella, W. ilvensis, and W. obtusa. 58 Rhodora [Vol. 98 Lygodium palmatum MARSILEA QUADRIFOLIA Figure 30. Distribution maps fi ips for Woodsia gracilis, Trichomanes intri- catum, Lygodium palmatum, and MARSILEA QUADRIFOLIA. 1996] Angelo and Boufford—Atlas of New England Flora) 59 Botrychium dissectum Botrychium lanceolatum subsp. angustisegmentum Botrychium lunaria Botrychium matricariifolium Figure 31. Distribution maps for Botrychium dissectum, B. lanceolatum subsp. angustisegmentum, B. lunaria, and B. matricariifolium. 60 Rhodora [ Vol. 98 Botrychium rugulosum Botrychium simplex 32. Distribution maps for Botrychium minganense, B. multifidum, d B. simplex. and B. simpl 1996] Angelo and Boufford—Atlas of New England Flora 61 Botrychium virginianum Ophioglossum pusillum Osmunda cinnamomea Osmunda claytoniana Figure 33. Distribution maps for Botrychium virginianum, Ophioglossum pusillum, Osmunda cinnamomea, and O. claytoniana. 62 Rhodora [ Vol. 98 var. spectabilis Osmunda X ruggii Polypodium appalachianum Figure 34. Distribution maps for Osmunda regalis var. spectabilis, O. cin- namomea X O. claytoniana, O. X ruggti, and Polypodium appalachianum. 1996] Angelo and Boufford—Atlas of New England Flora 63 Adiantum pedatum Adiantum X viridimontanum Figure 35. Distribution maps for Po/lypodium virginianum, Adiantum aleu- ticum, A. pedatum, and A. X viridimontanum. 64 Rhodora [ Vol. 98 Pellaea atropurpurea Pellaea glabella Figure 36. Distribution maps for Cheilanthes lanosa, Cryptogramma stel- leri, Pellaea atropurpurea, and P. glabella. 1996] Angelo and Boufford—Atlas of New England Flora 65 Phegopteris hexagonoptera Thelypteris noveboracensis Figure 37. Distribution maps for SALV/NIA NATANS, Phegopteris connec- tilis, P. hexagonoptera, and Thelypteris noveboracensis. 66 Rhodora | Vol. 98 Chamaecyparis thyoides Figure 38. Distribution maps for Thelypreris palustris var. pubescens simulata, Chamaecyparis thyoides, and Juniperus communis var. depressa 1996] Angelo and Boufford—Atlas of New England Flora 67 Juniperus communis var. depressa Juniperus horizontalis X J. virginiana X J. virginiana Figure 39. Distribution maps for Juniperus horizontalis, J. virginiana, J. communis var. depressa X J. virginiana, and J. horizontalis * J. virginiana. 68 Rhodora [| Vol. 98 LARIX DECIDUA Figure 40. Distribution maps for Thuja occidentalis, Abies balsamea LARIX DECIDUA, and L. laricina. 1996] Angelo and Boufford—Atlas of New England Flora 69 Figure 41. Distribution maps for PICEA ABIES, P. glauca, P. mariana, d P. rubens. 70 Rhodora [Vol. 98 PINUS NIGRA Pinus resinosa Figure 42. Distribution maps for Pinus banksiana, P. MUGO, P. NIGRA, and P. resinosa. 1996] Angelo and Boufford—Atlas of New England Flora 71 PINUS SYLVESTRIS PINUS THUNBERGII ire 43. Distribution maps for Pinus rigida, P. strobus, P. SYLVESTRIS, and P. THUNBERGII. de Rhodora [ Vol. 98 TAXUS CUSPIDATA Figure 44. Distribution maps for Tsuga canadensis, Taxus canadensis, and T. CUSPIDATA. 1996] Angelo and Boufford—Atlas of New England Flora — 73 REFERENCES ANGELO, R. 1994. A computer method for producing dot distribution maps. Rhodora 96: 190-1 BLASDELL, R. F. 1963. A monograph study of the fern genus Cystopteris. Mem. Torrey Bot. Club 21(4): 1-102. Britton, D. M. 1991. A Fa Isoétes, 1. X harveyi in northeastern North America. sae J. Bot. 69: 634-640. Brown, D. F 1964. . monographic study of the fern genus Woodsia. Beih. Nova Sane 16: | Bruce, J. G. 1975. ee and Morphology of Subgenus Lepidotus of the Genus Lycopodium (Lycopodiaceae). Ph. D. thesis. University of Michigan, Ann Arbor BRUNTON, D. E AND D. M. BRITTON. — oe prototypus (ISOETACEAE) in the United ae Rhodora 95: 12 CAMPBELL, C. S. AND L. M. EASTMAN. ee Flora of Oxford County, Maine. Life ae and Arca Experiment Station, University of Maine, Technical Bull. 99: 1-244. Orono, ME. . ADAMS, P. cane A. C. Dipste, L. M. EASTMAN, S. C. GAWLER, bs a Grecory, B. A. GRUNDEN, A. D. HAINES, K. JONSON, S. C. ROONEY, T. F. VINING, J. E. WEBER, AND W. A. WRIGHT. 1995. Checklist of the Vascular Plants of Maine. 3rd Revision, Maine Agricultural and Forest Experiment Sta- tion, ee eee of Maine, Bull. 844. Orono, M CARLSON, T. J. AND W. H. WAGNER, JR. 1982. The North American distribution of the genus o yopteris. Contr. Univ. Michigan Herb. 15: 141-162. Cave, M. S., ed. 1958a. ae to Plant Chromosome Numbers for 1956. [No. 1] Calif. Bot. Soc., Berkeley, CA. , ed. 1958b. nk © Plant Chromosome Numbers for 1957. [No. 2] Calif. Bot. Soc., Berkeley, , ed. 1959a. se to > Plant Chromosome Numbers, Supplement (previous to 1956). Calif. Bot. Soc., Berkele —., ed. 1959b. Index to Plant on ne ape for 1958. No. 3. The University of North Carolina Press, Chapel Hill, 1960. Index to Plant Chromosome Simi ee 1959. No. 4. The Uni- iersity of North Carolina Press, Chapel Hill, ,ed. 1961. Index to Plant Chromosome —— for 1960. Vol. II, No. 5. The University of North Carolina Press, Chapel Hill, NC. ed. 1962. Index to Plant Chromosome Numbers for 1961. Vol. II, No. 6. The University of North Carolina Press, Chapel Hill, NC. ed. 1963. Index to Plant Chromosome Numbers for 1962. Vol. II, No. 7. The ner of North Carolina Press, Chapel Hill, NC. ,ed. 1964. Index to Plant Chromosome ee for 1963. Vol. II, No. 8. The eas of North Carolina Press, Chapel Hill, NC. 1965. Index to Plant Chromosome ee for 1964. Vol. Il, No. 9. The University is ee Carolina Press, Chapel Hill, NC CHADWICK, L. C. A R. A. KEEN. 1976. A study of the genus Taxus. Ohio Agric. Exp. Sta. Ball, 1086. Wooster, OH. Cuurcu, G. L. AND R. L. CHAMPLIN. 1978. Rare and Endangered Vascular Plant a in Rhode Island. The New England Botanical Club in cooperation with e U.S. Fish and Wildlife Service [Region 5, Newton Corner, MA]. 74 Rhodora [ Vol. 98 CLAUSEN, * : 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club 19: 1-177. Coss, B. en A Field Guide to the Ferns. Houghton Mifflin Company, Boston, MA. CODDINGTON, J. AND K. G. FIELD. 1978. Rare and Endangered Vascular Plant Spe- cies in Massachusetts. The New England Botanical Club in cooperation with The U. - ot and Wildlife Service [Region 5, Newton Corner, MA]. Copy, W. J. . M. BRITTON. 1989. Ferns and Fern Allies of Canada. Agric. Can. a ee Ottawa. COUNTRYMAN, W. D. 1978. Rare and Endangered Vascular Plant Species in Ver- mont. The New England Botanical Club in cooperation with The U.S. Fish and Wildlife Service [Region 5, Newton Corner, MA] CRANFILL, R. 1983. The distribution of Woodwardia areolata. Amer. Fern J. 73: 46-52 .M. BRITTON. 1983. Typification vee the Polypodium virginia- num ear pe aes Hi 32: 557-56 CRITCHFIELD, W. B. AND E. L. L . IR. 1966. ie Distribution of the Pines of the World. U. S. D. A Mise. Pub. 991, Washington, D.C Crow, G. E. 1982. New England’s Rare, Threatened, and Endangered Plants. U.S. Department of the Interior, Fish and Wildlife Service, in cooperation with the New Hampshire Agricultural Experiment Station, University of New Hamp- shire. Washington, D.C. DowHan, J. J. 1979. Preliminary Checklist of the Vascular Flora of Connecticut. State Geological and Natural History Survey of Connecticut, Report of Inves- tigations No. 8, Hartfor EASTMAN, L. M. 1978. Rare and Endangered Vascular Plant Species in Maine. The New England Botanical Club in cooperation with The U.S. Fish and Wildlife Service [Region 5, Newton Corner, MA]. FARJON, A. 1990. A Bibliography of Conifers. Konigstein. Regnum Veg. 122: 1-129. FARRAR, D. R. 1992. Trichomanes intricatum: The se Trichomanes ga- metophyte in the eastern United States. Amer. Fern J. 82: 68—74. FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th a American Book Com- any, New York. FLORA OF NorTH AMERICA EpITORIAL COMMITTEE, ed. 1993. Flora of North America. Volume 2. Pteridophytes and Gymnosperms. Oxtord University Press, New FRANCO, J. D. A. 1962. Taxonomy of the common juniper. Bol. Soc. Brot., ser. 2a, 36: 107-120. Gastony, G. J. ao The Pellaea glabella complex: Electrophoretic evidence for the derivations of the agamosporous taxa and a revised synonymy. Amer. Fern J. 78: 44-67. GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of the North- eastern United States and Adjacent Canada. The New York Botanical Garden. Bronx, NY. AND A. CRONQUIST. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. D. Van Nostrand Company, New 199], Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx, NY. 1996] Angelo and Boufford—Atlas of New England Flora a GoLpBLATT, P., ed. 1981. Index to Plant Chromosome Numbers 1975— 1978. Monographs in Systematic Botany. Vol. 5. Missouri Bot. Gard., St. Louis, ,ed. 1984. Index to sar Chromosome Numbers 1979-1981. Monographs in Systematic Botany. Vol. 8. Missouri Bot. Gard., St. Louis, MO. ed. 1985. Index to ee Chromosome uinbers 1982— 1983. Monographs in Systematic Botany. Vol. 13. Missouri Bot. Gard., St. Louis, MO. ed. 1988. Index to Plant Chromosome Numbers 1984-1985. Monographs in Systematic Botany. Vol. 23. Missouri Bot. Gard., St. Louis, MO. _E. Jounson, eds. 1990. Index to Plant Chromosome Numbers 1986-1987. Monographs in Systematic Botany. Vol. 30. Missouri Bot. Gard., St. Louis, MO. 991. Index to Plant Chromosome Numbers 1988— 99 - Monogh in Systematic Botany. Vol. 40. Missouri Bot. Gard., St. Loui ae ———., eds. 1994. Index to Plant Chromosome Numbers 1990— cor Ronee in Systematic Botany. Vol. 51. Missouri Bot. Gard., St. Louis, M 996. Index to Plant Chromosome hae 1992-1993. Monographs in ane Botany. Vol. 58. Missouri Bot. Gard., St. Louis, MO. Gorpon, A. G. 1976. The taxonomy and genetics of Picea see and its rela- re to Picea mariana. Canad. J. Bot. 54: 781-81- HALL, M. 1952. Variation and hybridization in Siniser us. Ann. Missouri Bot. Gard. 1-04. Harris, S. K. Oe The Flora of Essex County, Massachusetts. Peabody Museum, Salem, MA. HAUuELER, C. H. AND Z. R. WANG. 1991. Chromosomal analyses and the origin of allopolyploid Polypodium virginianum. Amer. J. Bot. 78: 624-629. M. D. WINDI 991. A new species of North American Cystop- teris and Polypodium, with comments on ‘heir reticulate relationships. Amer. Fern J. 81: HAUKE, R. L. 1963. A taxonomic monograph of Equisetum subgenus Hippoc haete. Beih. Nova Hedwigia 8: 1-123. 966. A systematic study of Equisetum arvense. Nova Hedwigia 13: 81- ! ——.. 1978. A oe monograph of Equisetum subgenus Equisetum. Nova Hedwigia 30: 385 HEWITSON, W. 1962. Compara morphology of the Osmundaceae. Ann. Mis- souri Bot. Gard. 49: 57-93. Hickey, R. J. 1977. The as aie obscurum complex in North America. Amer. 5-49. 1980. New pane allies, Ophioglossaceae, Hymenophyllaceae, and Marsileaceae. Rhodora 82: 607. 1986. /soétes megaspore Geen morphology: Nomenclature, variation, and systematic ae ance. Amer. Fern J. 76: 1-16. W. C. TA p N. T. LUEBKE. 1989. The cee concept in pterido- phyta with sil ee to /soétes. Amer. Fern J. 8-89, Hopacpon, A. R. AND F. L. STEELE. 1958. The Woody nee of New Hampshire. Agricultural el Station, Bulletin 447. University of New Hampshire. Durham, NH 76 Rhodora [ Vol. 98 HOFFMAN, M. S., ed. 1992. The World Almanac and Book of Facts, 1993, p. 389. Pharos Books, New York. ian 4 1975. Diphasiastrum, a new genus in Lycopodiaceae. Preslia 14; 97— i B. F., C. R. WERTH, AND S. I. GUTTMAN. 1984. Genetic relationships in Abies (fir) of eastern United States: An electrophoretic study. Canad. J. Bot 62: 609-616. Kato, M. 1980. A new combination in Athyrium ( (Athyriaceae) of North America. Ann. prices Mus. 49: | ) D. DARNAEDI. 1988. Taxonomic and phytogeographic relationships of Dine ium ae D. pycnocarpon, and Diplaziopsis. Amer. Fern J. 78: —_ 77_8 Kort, L. . 1981. a acadiensis, a new species from eastern North America. anes J. Bot. 59: 2592-2594. > D. M. BRITTON, 1980. eaey Nae Numbers for /soéfes in north- eastern ue America. Canad. J. Bot. 58: 980-6 aS AND, 1985, Sore mor ie .. one of /soétes in north- eastern North a Canad. J. . 61: 3140-3163. LELLINGER, D. B. 1985. Field ne of i i & Fern-allies of the United States & Canada. aetna Institution Press, Washington, D.C. LESTER, D. T. 1968. Variation in cone morphology of balsam fir, Abies balsamea. Rhodora 70: 83-94. Liew, F.S. 1972. Numerical taxonomic studies on North American lady ferns and their allies. Taiwania 17: 190-221. Liu, T. 1971. A ono of the Genus Abies. National Taiwan University, Tai- pel, Taiwar LLoypb, R. M. AND F. A. Lana. a The Polypodium vulgare complex in North America. Brit. Fern eer 9: —177. Love, A., D. LOVE, AND R. E. ICHT SERMOLLI. 1977. Cytotaxonomical Atlas of the oO ae Liechtenstei MACKEEVER, F. C. 98. Plants of Sean The University of Massachusetts Press, cee we MARSHALL, M. P. 1977. A Vascular Flora of pate Harta Vermont. Mas- ter’s Thesis, poe, of North Carolina. Chapel Hill, NC. MCMAHON, J. . L. JAcoBson, Jk. AND F. HYLAND. 1990. An Atlas of the Native Woo Gody Plants of Maine: A Revision of the Hyland Maps. Maine Agri- Sama Experiment Station Bulletin 830. U niversity of Maine. Orono, ME. MCNEILL, J. AND K. M. Pryer. 1985. The status and typification of Phegopteris and saa ‘arpium. Taxon 34: 136-143, ME pore L. J. 1978. Rare and Endangered Vascular Plant Species in Connecti- cut. The New England Botanical Club in cooperation with the U.S. Fish and aa Service [Region 5, Newton Corner, MA]. MERRIAM-WEBSTER, INC. 1984. Webster’s New Geographical Dictionary. Spring- field, MA MICKEL, J. T. 1979, ee fern genus Cheilanthes in the continental United States. ules 41: 431-437. Mirov, N. T. 1967. os Genus Pinus. Ronald Press ae New Yor ae BOTANICAL GARDEN. As of 27 April Search the plant eee numbers. gopher://mobot. ue org:70/77/. Chiemeli nee chrom 1996] Angelo and Boufford—Atlas of New England Flora = 77 MONTGOMERY, J. D. 1982. i ale in North America. Part II. The hybrids. Fiddlehead Forum 9(4): 23-3 M. PAULTON, ae Dryopteris in North America. Fiddlehead Fo- rum. 8(4): 25-31 Moore, R. J., ed. 1973. eg Plant Chromosome Numbers for 1967-1971. Reg- num aon 90: 39. 1974, eee to aa Chromosome Numbers for 1972. Regnum Veg- dae 91: 1-108. ,ed. 1977. Index to Plant Chromosome Numbers for 1973-1974. Regnum Vegetabile 96: 1-257. Moran, R. C. 1982. The — wrichomanes complex in the United States ata ee. Canada. r. Fern J. 72: 5-11. ene tenuis (Michx.) Desv.: A poorly understood species. Cas- tanea 48: 218-223. MORGENSTERN, E. K. AND J. L. FARRAR. 1964. Introgressive sg a in Red Spruce and Black Spruce. Univ. Toronto, Fac. Forest., Techn. Rep. 4. Toronto. NAUMAN, C. E. 1993. Salviniaceae Reichenbach, pp. 336-337. /n: Flora of North America Editorial Committee, ed., Flora of North America. Volume 2. Pterido- phytes and Gymnosperms. Oxford University Press, New OGDEN, E. B. 1948. The te of Maine. University of Maine Studies. Second a ries, No. 62. Orono OGDEN, E. C. 1981. Field Gui to Northeastern Ferns. New York State Museum pai ts 444, Alban (LLGAARD, B. 1987. A aa classification of the Lycopodiaceae s. lat. Opera Bot. 92: 153-178. ———. 1989. Index of the Lycopodiaceae. Biol. Skr. 34: 1-135. OrNDuFF, R., ed. 1967. Index to Plant Chromosome Numbers for 1965. Regnum bes 50: 1-128. , ed. 1968. Index to Plant Chromosome Numbers for 1966. Regnum Veg- etabile ue 1-12 1969. Index to Plant Chromosome Numbers for 1967. Regnum Veg- 129. PAGE, C. N. 1976. The taxonomy and phytogeography of bracken-A review. Bot. > 1-34. PALER, M. H. AND D. S. BARRINGTON. 1995. The hybrid Cystopteris fragilis X C. tenuis (Dryopteridaceae) and the relationships between its tetraploid progeni- tors. ai Bot. 20: 528-545 PALMA-OTAL, M., W. S. Moore, R. P. ADAMS, AND G. R. JOSWIAK. 1983. Genetic and biogeographical analyses of natural hybridization between ] Juniperus virginiana L. and J. horizontalis Moench. Canad. J. Bot. 61: 2733-2746 Paris, C. A. 1991. Adiantum Be a new maidenhair fern in eastern North America. Rhodora 93: 105-122. AND M. D. WINDHAM. 1988. A biosystematic investigation en Adiantum cae complex in eastern North America. Syst. Bot. 13: 240-2 PEAS .S. 1964. A Flora of Northern New Hampshire. The New er Bo- sane Club, Inc. Cambridge, MA. RICHARDS, C. D., F. HYLAND, AND L. M. EASTMAN. 1983. en Check-list of the Vascular Plants of Maine. Bull. Josselyn Bot. Soc. 11: 78 Rhodora [Vol. 98 SARVELA, J. 1978. A synopsis of the fern genus Gymnocarpium. Ann. Bot. Fenn. 15: 101-106. SCAMMAN, E. 1947. Ferns and Fern Allies . New gta The New Hamp- shire Academy of Science, Bulletin No. 2. Durham, N SEYMOUR, F. C. me The Flora of New England. The Charles E. Tuttle Com- sis Rutland, V 969b. aa Plots of Vermont. Sse Experiment Station, Bulletin 660. a University of Vermont. Burlington, 1982. The Flora of New England, 2nd a easel Memoirs V. Plain- field, NJ. Storks, I. M. AND G. E, Crow. 1978. Rare and Endangered Vascular Plants of New Hampshire. The New England Botanical Club in cooperation with The U.S. Fish and Wildlife Service [Region 5, Newton Corner, MA]. SVENSON, H. K. AND R. W. PYLE. 1979. The Flora of Cape Cod. The Cape Cod Museum of ine — Brewster, MA. TAKAMIYA, M. AND S. KuRITA. 1983. Cytotaxonomic studies on Japanese ee of the genus eee sensu lato. Acta Phytotax. Geobot. 34: 79, aes T. M. C. 1970. Pacific Northwest Ferns and their Allies. University of Toronto Press, aa TaAYLorR, W. C. AND R. J. Hickey. 1992. Habitat, evolution, and speciation of Tsoétes. Ann. Missouri Gard. 79: 613-62 T. LUEBKE. 1988. /soétes X hickeyi a naturally occurring hybrid we ey ec oe ~~ macrospora. Amer. Fern J. 78: 6-13. THC 89. H THORNE, F. AND L. THORNE, enry Potter’s Field Guide to the Hybrid Ferns - i Northeast. esate Institute of Natural Science, Wood- stock, VT. Tryon, A. FE. 1957. A revision of the fern genus Pellaea section Pellaea. Ann. Missouri Bot. Gard. 44: 125-193. 1968. Comparisons of sexual and apogamous races in the fern genus Pe/l- ea. ene 70: 1-24. 78. New England ferns. Rhodora 80: 558-569. . M. TRYON, AND F. BADRE. 1980. Classification, spores, and nomencla- ture of the marsh fern. Rhodora 82: 461-474. sas R. M. 1955. Selaginella rupestris and its allies. Ann. Missouri Bot. Gard. : 1-99, plates 1-6. ree A review of the genus Dennstaedtia in America. Contr. Gray Herb. 187: at . TRYON. 1982. Ferns and Allied Plants, with Special Reference to ee America. Springer-Verlag, New York. VERMONT BOTANICAL AND BIRD CLuB. 1973, Check List of Vermont Plants. Bur- lington, VT VICKERY, B. 1981. The Rare Flora Me Aroostook County, Maine. Unpublished manuscript. Bates College. Lewiston, ME. WAGNER, W. H., JR. 1971. Evolution - Dr yopteris in relation to the be ate 47-192. In: Holt, P. C., ed., The Distributional History of the Bio the ene Appalachians. Part 2. Flora. Virginia Polytechnic Inst. and ee a 2s. Div. Monograph 2. Blacksburg, VA BEITEL. 1992. Gack cisectieanion of modern North Ameri- can Lycopodiaceae. Ann. Missouri Bot. Gard. 79: 676-686 1996] Angelo and Boufford—Atlas of New England Flora = 79 AND F. S. WAGNER. 1976. Asplenium * clermontae Sim from Clifton Gorge, Greene County, Ohio—a second North American record. Ohio J. Sci. 76: 99-102. WEATHERBEE, P. B. 1990. Flora of Berkshire County, Massachusetts. Master’s The- sis. University of New Hampshire. Durham 1996. Flora of Berkshire County, Massachusetts. The Berkshire Museum, Pittsfield, MA. RHODORA, Vol. 98, No. 893, pp. 80-84, 1996 FIRST REPORT OF UNCINIA MACROLEPIS AND U. TENUIS (CYPERACEAE) IN COLOMBIA AND NEW COLOMBIAN SITES FOR U. PALUDOSA ALD A. peas LER Department of — Biology. University of Minnesota Herbarium, . Paul, Minnesota 55 108- 1095 ABSTRACT. Uncinia macrolepis and U. tenuis are reported for the first time from Colombia. Additionally, four new Colombian sites are recorded for U. palu- dosa, previously known from only a single site in the country. A key to the five Uncinia species known to occur in Colombia is provided. Key Words: Colombia, Cyperaceae, Uncinia macrolepis, U. paludosa, U. tenuis The genus Uncinia Pers. (Cyperaceae), with some 60 to 70 spe- cies worldwide, is well represented in South America, with be- tween 20 and 30 growing in the mountains and cooler regions of the continent (Wheeler and Goetghebeur 1995). Of these, Uncinia hamata (Sw.) Urban and U. phleoides (Cav.) Pers. have long been known trom Colombia (Kiikenthal 1909; Mora-Osejo 1966). More recently, U. paludosa G. A. Wheeler and Goetghebeur, well known from Ecuador, has been reported from a single Colom- bian site (Wheeler and Goetghebeur 1995). While working on a revision of the Uncinia of Ecuador (Wheeler and Goetghebeur, in manuscript), many specimens from Colombia also were examined, some of which form the basis of this note. In this paper, two species of Uncinia, U. macrolepis Decne. and U. tenuis Poeppig ex Kunth, are reported for the first time from Colombia and four new Colombian sites are recorded for U. paludosa. A key to the five Uncinia species known to oc- cur in Colombia is provided at the end of this report. Uncinia macrolepis Decaisne in Dumont D’Urville, Voy. Péle Sud 213, [53 This species has a wide and rather remarkable distribution, oc- curring in northern South America, in Patagonia and Tierra del Fuego, and on the south-Atlantic islands of South Georgia and Tristan da Cunha (Wheeler 1994, 1995). In northern South America, specimens have been seen from Ecuador, Peru, and Ven- ezuela; also, single specimens have been seen from Bolivia (Beck SO 1996] Wheeler—Uncinia $1 21503 [MIN]) and Colombia (specimen cited below). The north- ern plants previously had been called U. meridensis Steyerm., whose type comes from Venezuela (Steyermark 1951), and Hooper (1968) considered similar-appearing plants from South Georgia (originally called U. smithii Philcox) and Tristan da Cunha to be conspecific with U. meridensis. In a separate paper (Wheeler 1995), however, it was demonstrated that the populations from northern South America, South Georgia, and Tristan da Cunha are morphologically indistinct from the Patagonian-Fuegian populations of U. macrolepis, whose name has priority. In northern South America Uncinia macrolepis occurs near the upper limit of paramo, at elevations between 4000 and 4600 m (Wheeler and Goetghebeur, in manuscript). In Colombia, the single known locality for this species is at Alto Ritacuva in the Sierra Nevada del Cocuy, in the Department of Boyaca, where it grows at about 4550 m. At this site the species is common “‘be- tween rocks on high slopes” and frequently is embedded in cush- ion plants. Because some paramo uncinias tend to look very simi- lar, some Colombian specimens originally identified as U. macrolepis (as U. meridensis) actually are referable to U. palu- dosa (for more details and citation of specimens see under that species further below). Although similar in aspect, U. macrolepis and U. paludosa differ in several morphological features, and, in fact, are in different sections of the genus, with the former in sec- tion Uncinia and the latter in section Platyandrae C. B. Clarke (sensu Clarke 1883 and Kiikenthal 1909). In addition to the dif- ferences given in the key, the style base of U. macrolepis is con- spicuously enlarged, whereas that of U. paludosa is little thick- ened Colombia. Dpto. BoYACA: Cordillera Oriental, Sierra Nevada del Cocuy; Alto Ritacuva; alt. 4550 m, 11-29 Apr 1959, Barclay & Juajibioy 7449 (US) Uncinia paludosa G. A. Wheeler & Goetghebeur, Aliso 14: 142. 1995. This species occurs in northern South America, with many sites known from Ecuador (Wheeler and Goetghebeur 1995, in manu- script), five from Colombia (specimens cited below), and a single one from Peru (Leé6n and Young 1663 [NyY]). In Colombia, it oc- curs in paramo, at elevations from about 3500 to 4000 m, grow- ing primarily in bogs and swamps dominated by low shrubs and 82 Rhodora [| Vol. 98 herbs, such as Chusquea sp., Espeletia lopezii, and Oritrophium limnophilium. Uncinia paludosa (sect. Platyandrae) appears to be closely re- lated to U. lacustris G. A. Wheeler, thus far known only from Ecuador, but differs by having wider leaves, longer rachillae, and slightly larger perigynia and achenes (Wheeler and Goetghebeur 1995: see Figures 2 and 3 for a comparison of the two species). It also is related to U. macloviana Gaudich., which grows in Pat- agonia and on the Falkland Islands (Wheeler 1994), but differs by having smaller perigynia and achenes and pistillate scales that are about equal to or slightly exceed the perigynia. As mentioned earlier, all of the specimens cited below origi- nally were identified as Uncinia meridensis (here called U. mac- rolepis), but they actually are referable to U. paludosa. For dit- ferences between these two species, see the key and the discussion under U. macrolepis. Colombia. Dpto. ARAUCA. Sierra Nevada del sk Quebrada El Play6n: Hoya S. Luis, | km al N de la finca El Play6n, alt. 3500 m, 13 Mar 1973, Cleef 9/53 (NY, U); Plan de S. José, alt. 3625 m, 9 Jun 1973, C a 10057 (GB, NY, U).—Dpto. BOYACA: paramos al NW de Belén, cabaceras Quebrada Minas, Hoya Clla. Larga, alt. 3775 m, 2 Mar 1972, Cleef 2092 (Ny, U); Sierra Nevada del Cocuy, Alto Valle Lagunillas, NNE de la Laguna Pintada, alt. 3980 m, 3 Oct 1972, Cleef 5782 (Ny, U); Paéramo de la Rusia, NW-N de Duitamo, Pena Blanca, 2 km al NE de Buenos Aires, alt. 3720 m, 16 Dec 1972, Cleef 7318 (Ny, U). Uncinia tenuis Poeppig ex Kunth, Enum. Pl. 2: 525. 1837. This species ranges from Tierra del Fuego (Wheeler 1994) northward to west-central Argentina (Barros 1969) and central Chile (Marticorena and Quezada 1985; Munoz-Schick 1980) and is disjunct on the Juan Fernandez Islands (Skottsberg 1922); fur- ther north it occurs in Ecuador (Wheeler and Goetghebeur, in manuscript) and is also reported from Costa Rica in Central America (Chater 1994). In Colombia, a single locality for this spe- cies is known from the summit of Quindio Pass, near the Caldas- Tolima boundary, where it grows on a “wind-beaten, bushy, wet crest” at about 3500 m. Uncinia tenuis occurs primarily in forests and thickets, where it grows in loose mats in moist soil and also epiphytically on de- caying, fallen trees. It is separated readily from the other Colom- bian uncinias by elas glabrous perigynia and deciduous scales. Also, the inflorescence axes of this species, even when greatly 1996] Wheeler—Uncinia 83 (or entirely) divested of perigynia, are identified easily by the presence of whitish or pale brown saccate appendages (about | mm long), which are the persistent portions of otherwise decidu- ous scales (Wheeler 1994). Notably, there are only two South American uncinias with deciduous scales and persistent saccate appendages, U. tenuis and U. subsacculata G. A. Wheeler and Goetghebeur, but the latter has shorter appendages (less than 0.5 mm long) and, thus far, is known only from the type locality in Ecuador (Wheeler and Goetghebeur 1995). Colombia. Dpto. CALDAS—TOLIMA boundary: summit of Quindio Pass, 04° 28’ N. lat., 75° 37’ W. long., alt. 3500 m, 29 Nov 1944, Fosberg 22389b (us). A KEY TO THE UNCINIA SPECIES OF COLOMBIA 1. Perigynia glabrous; pistillate scales deciduous (however, saccate ap- pendages about | mm long persist); filaments narrow (ca. 0.1 Mint: wide); plants ThizOMalous 23.2 45e25bc eos U. tenuis 1. Perigynia slightly to densely pubescent, the margins ciliate (at least distally); pistillate scales persistent; filaments wide (ca. 0.2—0.3 mm) or narrow; plants cespitose or rhizomatous .......... (2) 2. Spikes less than 2.5 cm long, with fewer than 20 perigynia per spike; filaments narrower than the anthers; culms frequently scabrous beneath the inflorescence .......... U. macrolepis 2. Spikes 2.5 cm long or longer, with more than 20 perigynia per spike; filaments as wide as anthers; culms usually smooth be- NAH INE WNLIORESCEHCE aducdkochcg hoes daa eR eee debaes (3) 3. Longest perigynia more than 6.5 mm; achenes narrowly oblong; spikes thickish, clavate (or sometimes subclavate); robust plans ap tol me tall oaks ereuaeee U. phleoides 3. Perigynia less than 6.5 mm long; achenes not narrowly oblong; spikes linear or subclavate; plants short or tall ...... (4) 4. Achenes more than 3 mm long; rachilla often bent just be- neath the hook; spikes subclavate; leaves flaccid, some CAUNING: - cone ciat eae nod eurese esse nesses U. hamata 4. Achenes less than 3 mm long; rachilla straight or nearly so; spikes linear; leaves rigid and mostly ascending, usually Ol Dasa) -~ cirereracatees ton gesoveuate ws U. paludosa ACKNOWLEDGMENTS. I thank Paul Goetghebeur (Gent Univer- sity, Belgium) for correspondence regarding Uncinia and for in- formation on the Colombian specimens of U. paludosa at the 84 Rhodora [Vol. 98 Utrecht Herbarium (U), The Netherlands. Thanks also to the cu- rators and directors of the following herbaria for the loan (or in- house study) of specimens from northern South America: AAU, F G, GB, K, LD, MICH, MIN, MO, NY, P, QCA, US, and WIS. LITERATURE CITED BARROS, M. 1969, Cyperaceae, pp. 38-92. /n: M. N. Correa, ed., Flora Patagénica, Pt. Hl, Typhaceae a Orchidaceae (exceptio Gramineae). Colec. Cient. Instituto Nacional de Tecnologia Agropecuaria. Vol. 8. CHATER, . o 94. Uncinia. In: Davidse, G. et al., eds., Flora Mesoamericana CLARKE, a. - 1883. On Hemicarex Benth., and its allies. J. Linn. Soc., Bot. 20: eee - “be era Uncinia. Results Norweg. Sci. Exped. Tristan da Cunha 1937-1938 8. KUKENTHAL, G. 1909. eae Caricoideae, pp. 1-824. /n: A. Engler, ed., Das Pflanzenreich, IV. 20, Heft 38. Wilhelm ie Leipzig. MARTICORENA, C. AND M. QUEZADA. 1985. Catalogo de la flora vascular de Chile. ce a Bot. 42(1—2): 1-157. MoraA-OSEJO, L. E. 1966. Cyperaceae, pp. 95-128. /n: P. Pinto-Escobar y L. E. Mora- oe Catalogo ilustrado de las plantas de eee Vol. I. Insti- tuto de Ciencias Natu , Universidad Nacional, Bog ombia. MUNOZ-SCHICK, M. 198( . F fore del parque nacional ae Mis Nac. Hist. San- tiago, Chile. SKOTTSBERG, C. 1922. The oo of the Juan Fernandez Islands. Nat. Hist. Juan Fernandez and Easter Is. —240, STEYERMARK, J. A. 19SL. Bownical exploration in Venezuela. I. Fieldiana, Bot. 28: 42 a Y 1-242. WHEELER, G. A. 1994. The Uncinia (Cyperaceae) of Tierra del Fuego, the Falk- land Islands, a South Georgia. Ans. Inst. Pat., Punta Arenas, Chile 22 (1993- 94): 21-31 1995. The status of Uncinia macrolepis, U. meridensis, and U. smithii Keypeee) in the New World. Hickenia 2: —164. ——— AND GOETGHEBEUR, P. 1995. Four new ee of Uncinia (Cyperaceae) from northern South America. Aliso 14: 141— nN RHODORA, Vol. 98, No. 893, pp. 85-93, 1996 THE PARALYCHNOPHORA GROUP OF EREMANTHUS (VERNONIEAE: ASTERACEAE) HAROLD ROBINSON Department of Botany, National Museum of Natural History, Smithsonian Institution, adie ok. D.C., 20560 ABSTRACT. The species placed in Paralychnophora by MacLeish are returned to the genus Eremanthus, and two related species, E. harleyi and E. santosii, are described as new, Key Words: Asteraceae, Compositae, Vernonieae, Eremanthus, Paralychnophora Extensive work has been done on Eremanthus Less. and its relatives during the last 20 years. Some genera such as Chresta Vell. ex DC. (Robinson 1980) and Paralychnophora MacLeish (MacLeish 1984) have been removed from Eremanthus. One traditional genus, Vanillosmopsis Sch. Bip., has been placed in the synonymy of Eremanthus (MacLeish 1987). Recent authors still do not agree completely on all the generic limits. In two genera, Eremanthus and Chresta, the concepts of the present author remain more inclusive than those of MacLeish (1984, 1985a, b, 1987). The group treated in the present paper, Paralychnophora, is regarded by the present author as part of Eremanthus. MacLeish (1987) states that Paralychnophora is like Lychnophora in having a biseriate pappus and glomerules that are derived from the internode reduction of corymbose subglomerules. She distinguishes Paralychnophora on the basis of axillary, solitary glomerules and angled achenes. Of these features, only the rather solitary nature of the glomerules of heads, a feature noted by Baker (1873), can be used to distinguish the group from the broad concept of Eremanthus. The present study of the Paralychnophora group has consid- ered the generic problem, but it concerns problems mostly at the species level. As a result of the study, an unvalidated Glaziou name, Eremanthus schwackei, is placed in synonymy and two new species, FE. harleyi and E. santosii, are described. The records of the names and synonyms are needed for a listing being prepared of all known Vernonieae of the Western Hemisphere. Paralychnophora was a nom. nov. by MacLeish (1984) for Sphaerophora Sch. Bip., Jahresber. Pollichia 20—21: 402. 1863, non Sphaerophora Blume, 1850 (Rubiaceae). The type of Para- 85 86 Rhodora | Vol. 98 lychnophora, P. bicolor (DC.) MacLeish, and two other species were included in Paralychnophora by MacLeish. All had been treated previously as Eremanthus. An initial problem was noticed in this study in Eremanthus (Paralychnophora) schwackei, a Gla- ziou species, that was recognized as distinct by MacLeish (1987), but which never has been validated. The proposed combination by MacLeish did not validate the species. In spite of the initial concern, the present study has shown that there now is no need to validate the Glaziou name. Instead, E. bicolor needs to be re- defined, and two new species need to be distinguished and de- scribed. A listing also can be provided of specimens of Ereman- thus reflexo-auriculatus Barroso that’ previously have been identified as E. bicolor. The four species of the Paralychnophora group of Eremanthus can be distinguished by the following key. |. Leaf blades with depressed areoles on upper surface; fertile branches with subopposite or opposite median bracts; corolla lobes with few hairs not extending beyond tips of lobes... 1. E. bicolor . Leaf blades with weakly bullate or rugulose upper surfaces; fertile branches with strictly alternate bracts; corolla lobes with dense hairs distally, some extending beyond tips of lobes ....... (2) 2. Individual heads with 2 or 3 florets; leaf blades oblanceolate with NAMOWY ClNedle DASES i stces dace cseunesnxa ey ty SOMOS 2. Individual heads with 5—9 florets; leaf blades oblong to obovate with shortly acute to auriculate bases ..............0... (3) 3. Bases of leaf blades auriculate, often clasping the stem ..... ee ee ee ee re ene 4. E. reflexo-auriculatus 3. Bases of leaf blades obtuse to shortly acute, distinctly petio- MOUS, heated ucap din dius eneheendaseenteiacien ee dae — EKremanthus bicolor (DC.) Baker in Mart., Fl. brasil. 6(2): 1873. (Figure 1) — Albertinia bicolor DC., Prodr. 5: 81. 1836. Sphaerophora bicolor (DC.) Sch. Bip., Jahresber. Pollichia 20-21: 402. 1863. Eremanthus schwackei Glaziou, Bull. Soc. Bot. France 3: 380. 1909, nom. nud. Paralychnophora bicolor (DC.) MacLeish, Taxon 33: 106, 1984 Paralychnophora schwackei (Glaziou) MacLeish, Taxon 33: 106. 1984, nom. nud. Shrubs or trees 1.5—5.0 m tall, moderately branched; stems terete, usually grayish velutinous, distal internodes 0.3—1.0 cm long. Leaves alternate, petioles 0.5—1.8 cm long; blades oblong- 1996] Robinson—Eremanthus 87 elliptical, 5-19 cm long, 3-6 cm wide, base rounded to attenuate, margins entire, apex rounded, upper surface minutely alveolate when dry, 9-15 secondary veins on each side. Inflorescences on long, ascending branches from upper leaf axils, branches with re- duced, easily deciduous bracts, median bracts usually suboppo- subequal series, linear, to 5 mm nied. with reflexed, white- tomentose tips. Florets 2—4 in a head; corollas violet, ca. 7 mm long, tube 3-4 mm long, throat nearly lacking, lobes ca. 3 mm long, erect, with few hairs distally not extending beyond lobe tip, with numerous glands. Achenes ca. 3 mm long, glabrous, with idioblasts on surface; inner pappus bristles ca. 4.5 mm long, with slightly but distinctly broadened tips, outer pappus elements nar- rowly linear, ca. 1.5 mm long. Pollen ca. 45 jum in diameter. Brazil. MINAS GERAIS: on sandstone outcrop ca. 3—5 km east of Serra, road to Diamantina, 9 Aug 1960, Maguire, Magalhaes & Maguire 49119 (NY, US); slopes and summit of Serra Grao Mogul, 900-1100 m, frequent, on sandstone, 17 Aug 1960, Maguire, Magalhaes & Maguire 49247 (Ny, US); Serra do Espinhago, ca. 18 km E of Diamantina, elev. 1100 m, eee summits with soil-filled crevices and small areas of white sand, 19 1970, Irwin, Fonséca, Souza, Reis dos pie & Ramos 27887 — UB, US); ca. oe km SW of Gouveia, km 243 on M.G. , LO80 m, very rocky sandstone ra with occasional wet hee 2 Feb 1972, pee Stieber & Kirkbride 35116 (NY, UB, US); Ribeirao (mun. Grao Mogol), dos pared6es rochosos na beira do corrego, 23 Apr 1978, aes 41427 (MBM, US); between Mendanha and Diamantina, 10 km NE of Diamantina on Estrada 367, campo rupestre, 4 Oct 1980, MacLeish, Ferreira, Smith & Stutts 701 (GA, us); Grao Mogol, subida para Pasto, campo rupestre, encosta de morro junto a — afloramentos rochosos, 17 May 1988, Hatschbach & Ribas 52102 (MBM, US): Penha da Franga, ca. de 100 km ao nordeste de aman, 18°S’S 43°5'O, ca. 1000 m, 11 Mar 1995, Splett 86S (BONN, UB, US). Glaziou (1909) named the plant the same way he named many others, indicating locality, “Diamantina,” habit, “‘arbusto,” and flower color, lilac. Such names have been regarded as nom. nud. by recent authors, and such species are accepted only after valida- tion by later authors. No such validation has occurred for the Gla- ziou name, and now it seems that such validation is not necessary. Material annotated as Eremanthus schwackei was distinct from extensive material from Bahia recently determined as E. bicolor. However, the types of both FE. schwackei and E. bicolor are from Minas Gerais, and the original description (Candolle 1836) and subsequent critical description (Baker 1873) of EF. bicolor state 88 Rhodora [| Vol. 98 that there are 2 or 3 florets in a head, indicating that EF. bicolor and FE. schwackei are the same species. It is the extensive mate- rial determined as F. bicolor from Bahia that needs to be placed in other species, including two that are new. The material deter- mined as EF. schwackei seems to differ from more typical F. bi- color only by the somewhat shorter, oblong leaves that are 5—12 11-19 cm long. One of the most distinctive features of the combined concept of E. bicolor and E. schwackei is the elongate, ascending inflorescence branches. The branches are particularly noticeable after the bracts have fallen. The median bracts of the branches are few, and they seem to be opposite or subopposite. Other less obvious, but apparently useful differences are the regu- larly alveolate upper surface of the dried leaf and the limited pu- bescence of the corolla lobes. All other members of the Paraly- chnophora group have minutely bullate or rugulose upper leaf surfaces and longer and denser pubescence on the corolla lobes. The Bahia material previously named as EF. bicolor or E. schwackei proves to belong to either FE. reflevo-auriculatus Bar- roso or one of the two following new species. iw) Eremanthus santosii H. Robinson, sp. nov. TYPE: BRAZIL. Bahia: Santa Maria Eterna, | a 2 km de estrada a Canavieires, campo arenoso coberta de érvore e arbusto, 18 May 1970, 7.S. dos Santos S20 (HOLOTYPE: CEPEC; ISOTYPE: US). (Figure 2) Plantae frutescentes 2.5—4.0 m altae leniter ramosae; internodes distales 0.5—2.0 cm longis. Folia in laminis oblanceolata ad 18 cm longa base cuneata margine saepe recurvata apice rotundata in nervis secundariis utrinque ca. 15. Inflorescentiae in ramis in- terdum elongatis ascendentibus dispositae, bracteis alternis; flores in capitulo 2 aut 3; corollae in lobis distaliter dense longe pilo- sae. Shrubs 2.5—4.0 m high, sparsely branched; stems terete below, sulcate when dried young, usually yellowish-brown velutinous, distal internodes 0.5—2.0 cm long. Leaves alternate, petioles mostly 1.0—2.5 cm long; blades oblanceolate, 13-18 cm long, 4.0— 4.5 cm wide, base narrowly cuneate, margins entire, often broadly recurved, with sides nearly parallel in distal half, apex rounded, upper surface minutely bullate or rugulose when dry, ca. 15 close- set, spreading secondary veins on each side. Inflorescences on long, ascending branches from upper leaf axils or with spreading branches from lower axils, branches with reduced, easily decidu- 1996] Robinson—Fremanthus 89 ous bracts, all bracts alternate. Inflorescences becoming spherical when mature, 2.5—3.0 cm in diameter. Heads sessile; involucral bracts, in ca. 3 subequal series, linear, to 7 mm long, with re- flexed, white-tomentose tips. Florets 2 or 3 in a head; corollas violet, ca. 7 mm long, tube ca. 3.5 mm long, throat nearly lack- ing, lobes ca. 3.5 mm long, erect, with many hairs distally, some extending beyond lobe tip, with numerous glands. Achenes ca. 3 mm long, glabrous, idioblasts not seen; inner pappus bristles ca. 5 mm long, shghtly broadened from bases to tips, outer pappus elements linear to setiform 0.5—1.0 mm long. Pollen ca. 50 wm in diameter. PARATYPES: Brazil. BAHIA: Mt. Peludo, H. M. Curran 387 (Us); Santa Maria Eterna km 1, a Canavieires, campo arenoso, 13 Aug 1971, 7. S. dos Santos 1512 (CEPEC, US); Serra do Sincora, 3—13 km W of Barra da Estiva on road to Jussiape, ee 300 m, 13°40'S 4 1°25’W, low grassland with scattered woodland — small streams on white quartzite soils, growing on slope of sandstone ridge 1980, Harley, Bromley, Carvalho & Martinelli 20809 (CEPEC, K, US); aes de Abaira, Campos de Ouro Fino, pr6ximo a Serra dos Bicanos, 1650 m, 13°16'S 41°54'W, campo rupestre, entre rochas, 16 July 1992, Wilson Ganev 671 (K, SPF, US). Specimens of Eremanthus santosii have been identified as Er- emanthus bicolor or as the synonymous Paralychnophora schwackei (Glaz.) MacLeish, which is identified here as EF. bi- color. The new species includes specimens that have elongate, as- cending inflorescence branches as in E. bicolor, and the two spe- cies share the characteristic of few-flowered heads. The first two specimens examined by this author initially seemed different from most specimens of E. bicolor because of the longer cuneate leaf bases. However, cuneate bases apparently occur in some speci- mens of £. bicolor (Baker 1873, pl. 45). The more important dis- tinctions are the often less congested distal leaves with longer in- ternodes, the slight difference in the bulging rather than depressed areoles on the upper surface of the leaves, the strictly alternate bracts on the inflorescence branches, and the presence of the larger, more numerous hairs on the corolla lobes. At present, F. santosti is known only from Bahia, while &. bicolor is known only from Minas Gerais. The leaves of the two dos Santos col- lections are pressed flat, but the leaves of the other three speci- mens show a strong tendency for the margins to recurve, a ten- dency noticed only in E. reflexo-auriculatus among other species of the group. 90 Rhodora [ Vol. 98 The new species is named for the collector of the type speci- men, T. S. dos Santos of the Herbario Centro de Pesquisas do Cacau (CEPEC) in Ilhéus, Bahia. 3. Eremanthus harleyi H. Robinson, sp. nov. TYPE: BRAZIL. Bahia: Mun. Rio de Contas, Pico das Almas, Vertente leste, alto do vale acima da Faz, Silvina, 13°31'S 41°58'W, 1550 m, sobre rochas, 16 Dec 1988, Harley 27255 (HOLOTYPE: CEPEC; ISOTYPES: K, SPF, US). (Figure 3) Plantae frutescentes vel arborescentes 1—5 m altae; internodes distales 0.3—-1.0 cm longis. Folia in laminis oblongo-elliptica plerumque 8—17 cm longa 3.5—8.0 cm lata base obtusa vel breviter acuta apice rotundata supra minute bullata vel rugulosa in nervis secundaris utrinque 6—12. Inflorescentiae saepe distincte latera- les plerumque mediocriter vel late patentes, ramis brevibus vel mediocriter elongatis, bracteis alternis; flores in capitulo 5—9; corollae in lobis distaliter dense longe pilosae. Shrubs to small trees 1.5—5.0 m high, moderately branched; trunks to 8 cm in diameter, terete below, young parts sulcate when dried, yellowish or brownish velutinous, internodes 0.5—2.5 cm long. Leaves alternate, petioles 1.0—3.5 cm long; blades oblong- ovate to elliptical, 8-17 cm long, 3.5—8.0 cm wide, base obtuse to shortly acute, margins entire, flat, apex rounded to obtuse, up- per surface minutely bullate or rugulose, lower surface whitish to sordid tomentose, with prominent veins, with 6—12 secondary veins on each side of midrib, strongly arching distally. Inflores- cence on short terminal peduncle or on spreading lateral branch mostly 8-15 cm long, bracteoles of branch alternate; inflorescence becoming spherical or subspherical when mature, 3.0-4.5 cm in diameter, with many lanceolate to elliptical foliose bracts at base, these I—5 cm long. Heads sessile; involucral bracts in ca. 3 sub- equal series, linear, 5-7 mm long, densely white-tomentose out- side. Florets 5—9 in a head: corollas rosaceous to violet, ca. 8.5 mm long, tube ca. 5 mm long, throat ca. 0.5 mm long, lobes ca. 3 mm long, with many hairs at tip, some extending beyond lobe tip, with numerous glands. Achenes ca. 3 mm long, glabrous, id- ioblasts not seen; pappus bristles ca. 7 mm long, slightly broad- ened from bases to tips, outer pappus elements setiform, 1-2 mm long, not broadened distally. Pollen ca. 48 zm in diameter. PARATYPES: Brazil. BAHIA: Serra do Rio de Contas, 12-14 km N of town of Rio de Contas on road to Mato Grosso, ca. 1200 m, approx. 13°28'S 41°S50'W, dry quartzite hillside with disturbed woodland on lower slopes and scrub above, 1996] Robinson—Eremanthus 9] 17 Jan 1974, Harley, Renvoize, Erskine, Brighton & Pinheiro 15187 (CEPEC, K, us); Mun. de Rio de Contas, a 4 km ao de Rio de Contas, campo rupestre, ca. 1000 m, 21 July 1979, King, Mori, T. S. dos Santos & Hage S069 any Us); base de Pico das Almas, a 18 km ao NW de Rio de Contas, ca. 1300 m, 24 July 1979, ec Mori, T. S. dos Santos & Hage 8132 (CEPEC, US); Mur Mucugé, a 3 km ao S de Mucugé, na estrada que vai par Jussiape, ca. 1000 m, 26 July 1979, King, Mori, T. S. dos Santos & Hage 8152 (CEPEC, US); Estrada e liga ee com Andarai a Il km de scenes ca. 1150 m, 27 July 1979, a. Mori, T. S. dos Santos & Hage 8170 (CEI us); Mun. Lengois, Serras dos Lengois, Serra do Brejao ca. 14 km NW of ae western face of sandstone serra with mre bedded rocks, extensive grassland & marsh on low slopes & small streams with low woodland, rock outcrops at eee altitudes, oe humid forest in rocky declivity below summit, 700- 1000 m, 12°27'S 41°27'W, E6, 22 May 1980, Harley, Bromley, Carvalho, Soares Nunes, . & bE. B. dos Santos 22335 (CEPEC, K, US); Vicinity of Pico das Almas, ca. 20 km NW of town of Rio de Contas, 3500-4600 ft., 25 Jan 1981, King Bishop 8668 (UB, US); km along road S of Mucugé, 2600 ft., 31 Jan 1981, King & Bishop 8712 coe, us); 52 km E of Seabra, along road toward Itaberaba, 1900 ft 2 ed 1981, King & Bishop 8774 (uB, US); Mun. of Lengois, along BR242, ca. 15 km NW of Lencois at km 225, ca. 900 m, campo rupestre, 10 June ere ee & Boom 14271 (NY, oe Estrada de Lengois BR242, 5 km ao N de Lengois, campo natural, 19 Dec 1981, Carvalho, Lewis & Hage 100] (CEPEC, US); Rio de Contas, arredores, campo rupestre, 17 May 1983, Hatschbach 46552 (MBM, US); Mucugé, 5 km S, campo peed solo arenoso, 22 Jan 1984, Hatschbach 47486 (MBM, us); Mun. Piataé: Quebrada da Serra do one 13°13'S 41°50’W, 1600-1700 m, 26 Dec 1991, Hind, a Souza, Sakaragui e Ganev SO385 (CEPEC, HUEFS, K, SPF, US); Mun. de Abaira, Belo Horizonte, acima do Jambreiro, pr6ximo a Serra do Sumbaré, 13°18’S 41°52’W, 1350 m, campo rupestre com solo arenoso e grandes matag6es de rochas, 27 Oct 1992, ree ne (HUEFS, K, SPF, US); . Abaira, Campo de Ouro Fino (alto), 13°1S’S 4 4'W, 1700-1800 m, em capoeira e sobre e entre rochas na encosta do morro, 17 Jan 1992, Hind & Queiroz SOO71 (CEPEC, HUEFS, K, SPF, US). Eremanthus harleyi includes the bulk of the material that has been identified in recent years as E. bicolor from Bahia. The present study has shown that true E. bicolor is mostly, if not to- tally, restricted to Minas Gerais and differs in having fewer flo- rets in the heads, a different upper leaf surface, more closely set secondary veins of the leaf, opposite or subopposite median bracts on the inflorescence branches, and fewer, shorter hairs on the lobes of the corollas. Eremanthus harleyi is more closely related to E. reflexo-auriculatus, which also occurs in Bahia, but the lat- ter differs in having the strong basal auricles on the leaf blade, the often strongly recurved leaf margins, and the foliose bracts at the base of the inflorescences that are more broadly ovate. The species is named for Ray M. Harley of Kew, the collector of the type specimen and a specialist in the Lamiaceae, who has organized a number of trips to collect Bahian specimens. Figures 1-4. & Boom 14271, Leaves of Eremanthus spp. 1. E. bicolor ). US). 4. E. reflexo-auriculatus (Harley 22766, US). Scale bar = 2 cm. ( YY a Hatschbach 52102, Us). 2. E. santosii (Ganev 671, US). 3. E. harleyi (Mori c ‘6 vIOPOYY 86 1A] 1996] Robinson—Eremanthus 93 4. Eremanthus reflexo-auriculatus Barroso, Rodriguésia 35-36: 6. 1960-61 [1962]. (Figure 4) Paralychnophora reflexo-auriculata (Barroso) MacLeish, Taxon 33(1): 106. 1984. The species originally was described from Pernambuco (Bar- roso, 1960-1) to accommodate a specimen differing from Eremanthus bicolor in having strongly auriculate bases of the leaf blades. The leaf blades also commonly have broadly recurved margins. The species seems to be closely related to FE. harleyi, which shares the shorter, usually more spreading branches and the 6—9-flowered heads, but the series of specimens seen from Bahia shows a consistent difference in the leaf base. The following specimens of Eremanthus reflexo-auriculatus have been extracted from material in the U.S. National Herbarium previously identified as E. bicolor. Brazil. BAHIA: Serra do Tombador, 20 Feb 1971, /rwin, Harley & Smith 30689 (NY, US); Munic. Morro do 30-31 May 1980, Harley, Bromley, de Car- valho, Soares Nunes, Hage & 3. dos Santos 22766, 22867 (CEPEC, K, US); Morro do Chapeu, 16 June 1981, Mori - Boom 14446 (NY, US): ae Jacobina, Serra do Tombador, ca. 25 km na estrada Jacobina/Morro do Chapéu, 20 Feb 1993, Carvalho, Amorin & Jardim 4177 (CEPEC, US). ACKNOWLEDGMENTS. John Pruski, of the Smithsonian Depart- ment of Botany, is thanked for many helpful suggestions. LITERATURE CITED BAKER, J. B. 1873. Compositae, I. Vernoniaceae, I]. Eupatoriaceae. /n: Martius, C , Flora ache Vol. 6(2): 1-398, pl. 1-102. CANDOLL LE, A. Pi 1836. Prodromus systematis naturalis regni vegetabilis. Vol. 5. Paris. Giaziou, A. F M. 1909. Plantae Brasiliae centralis a Glaziou lectae. Liste des ies du Brésil Central recueillies en 1861—1895. Mém. Soc. Bot. France 1(3): 297-392. MACLEISH, F. F. 1984. Argyrovernonia and Par ee new names in the tribe are (Asteraceae/Compositae). Tax 105-106. 985a. Revision of Glaziovianthus Sonera Vernonieae). Syst. Bot. 10: Pee 1985b. Revision of Chresta and Pycnocephalum (Compositae: Vernon- ieae). ae Bot. 10: 459-4 987, Revision of Eremanthus (Compositae: Vernonieae). Ann. Missouri cee 265-290 ee H. 1980. Notes on the Phy eae genera Chresta and Eremanthus See Asteraceae). Phytologia 45: 89— RHODORA, Vol. 98, No. 893, pp. 94-98, 1996 NEW ENGLAND NOTE A FLORA OF THE VASCULAR AND NON-VASCULAR PLANTS OF NANTUCKET, TUCKERNUCK, AND MUSKEGET ISLANDS R W. DUNWIDDIE Massa aie a Society, Lost Far Hummock Pond Rd., Nantucke a ASSé chhusetts 02554 Current Address: The Nature Conservanc 217 Pine St., Suite 1100, cite nea: O8 101 BRUCE A. SORRI Massachusetts Natural ea age and Endangered Species Program, stboro, MA O1581 Current Address: 160 West Rhode Island Ave., Southern Pines, orth Carolina 28387 Nantucket has been recognized widely as a botanical “hotspot” with one of the greatest concentrations of rare species in Massa- chusetts. A host of well-known 19th century botanists forayed on the island in search of unusual material. Many of their collec- tions still are intriguing. The first recorded plant specimen from Nantucket dates from 1827, when Thomas A. Greene of New Bed- ford made a collection of slender marsh-pink (Sabatia campanu- lata) trom the island. Two years later, James W. Robbins (of Po- tentilla robbinsiana fame) made a second collection of the same species from Greene’s site “in moist hollows in Squam.” Interest in this species continues to be keen as it is known in Massachu- setts only from two sites on Cape Cod and Nantucket. It was to be the first in a long line of notable records to come from Nan- tucket. Another of the early collections made on the island was seabeach amaranth (Amaranthus pumilus), collected by 5, 12 Ot- ney in 1849. It has not been seen again in the state. In 1888, Maria Owen published a flora of the island that in- cluded over 600 species. Her list was by no means definitive, but it was one of the first floras published for any county in Massa- chusetts. Eugene P. Bicknell produced the most definitive work on the Nantucket flora with his “Ferns and Flowering Plants of Nantucket,” published in a series of installments between 1908 and 1919 in the Bulletin of the Torrey Botanical Club. Now, nearly eight decades later, we have compiled a comprehensive flora that builds upon the work of these earlier botanists (Sorrie and Dun- widdie 1996; Dunwiddie and Sorrie 1996). 94 1996] New England Note oD Our flora is somewhat broader both geographically and taxo- nomically than earlier works. In addition to the island of Nan- tucket, we have included records from the other two vegetated islands in Nantucket County, Muskeget and Tuckernuck. Our work also includes lichens, mosses, and other bryophyte records. The published flora includes common and scientific names and synonymy for each species, as well as habitat and abundance in- formation, state ranks for rare species, data on origin (native ver- sus introduced), and notations for potentially invasive species. The flora also is available in a computerized relational database. This database contains all of the above information, in addition to fields noting life form (tree, shrub, graminoid, etc.), duration (annual, biennial, or perennial), records of our own collections and sightings, and notations indicating which previous authors also reported a Nantucket occurrence for a species. Our flora includes a total of 1265 vascular taxa, including spe- cies, varieties, subspecies, and hybrids. Thirty-nine percent (489) were introduced to the island. This compares to the total of 1136 reported by Bicknell, 33 percent of which were non-native. Due to taxonomic revisions and re-annotations of specimens, we cur- rently accept 982 of Bicknell’s records in our flora. The number of rare species is high, with 23 endangered, 13 threatened, 15 spe- cial concern, 46 watch list, and 7 historical, according to the Mas- sachusetts Natural Heritage and Endangered Species Program 992). The non-vascular flora comprises 79 mosses, 19 liver- worts, | hornwort, and 99 lichens. Nantucket has received some of the greatest scrutiny by bota- nists of any area of similar size (ca. 14,000 ha) in the state. As an example of one of the most completely inventoried regional floras, therefore, it is valuable to examine what deficiencies re- main in our knowledge of Nantucket species’ occurrence and dis- tribution. From such an analysis, inferences may be drawn regard- ing the thoroughness of our understanding of the many less intensively studied areas in New England. Currency of records. Three hundred and twenty-six species, over one-quarter of the flora, have not been recorded in over 60 years. This partially reflects the predilections of many recent bota- nists, ourselves included, to scour natural habitats for unusual records, and to ignore the back yards, waste dumps, and weed lots of the island. Yet nearly half of this total (153) includes na- tive species, many of which are not weedy. Their absence from 96 Rhodora [Vol. 98 recent lists may not be due to inadequate searching in appropri- ate habitats. Number of sites. Many species have been found only rarely on Nantucket. Bicknell noted that almost one-third of the flora (352 species: 182 native, 170 introduced) was recorded from only 1-3 sites. A comparable statistic is not available from our database, and will be the focus of future work. However, our impression is that this number has not declined greatly. About two-thirds of the 326 species for which we have no recent record (>60 years) fall into this group, and many other currently rare species are known only from the same sites where earlier botanists reported them. Lone occurrences of non-native taxa may reflect chance introduc- tions of plants that failed to become permanently established and may no longer be present. The abundance of native species with few records suggests that many may have extremely limited dis- tributions on Nantucket, and may be particularly susceptible to extirpation. Examples include Gaultheria hispidula, Schwalbea americana, Botrychium simplex, and Cornus amomum. Completeness of inventory. It is impossible to determine whether every species occurring on an island the size of Nan- tucket (ca. 30,000 acres) has been found. Furthermore, the total is always changing as new species are introduced and become naturalized, and others disappear from the island. One approach to assessing the thoroughness of our inventory is to examine the number of new species observations in recent decades. Based on his work from 1958 to 1966 on Nantucket, MacKeever (1968) reported about 50 taxa new to the island, 9 of which were native. Since beginning our work on the island in 1980, about 60 names have been added to the Nantucket vascular plant flora that previ- ously were not recorded; half of these were native. While some of these native species represented taxonomic changes that had not been recognized when earlier compilations of the Nantucket flora were prepared, others were undoubtedly species that simply had been overlooked by previous botanists on the island. Examples include Scleria pauciflora var. caroliniana, Amphi- carpum purshit, Aster acuminatus, Polystichum acrostichoides, and Amelanchier arborea. Our results suggest that even within a relatively small and heavily botanized area such as Nantucket, many questions remain unanswered regarding the nature of the local flora. Several points are worth considering as they may apply elsewhere. 1996] New England Note o7 1) Despite over a century and a half of extensive study, additional species continue to be found. These include species new to the island that are common elsewhere, as well as species that are rare regionally. A large number of species are known only from a few sites. Ef- forts should be made to identify which of these are truly limited in distribution and not merely undercollected. Measures then can be taken to protect sites of locally rare taxa to prevent their ex- tirpation. Our knowledge of much of the current flora may be woefully dated. The preparation and maintenance of large, private herbaria by serious amateurs as well as professionals was relatively wide- spread at the turn of the century. This practice virtually has disap- peared. As a result, much of our information on the presence and distributions of many species is out of date by many decades. Recent work often is conducted by professionals employed by conservation organizations and agencies who focus primarily on rare native species. The tendency to overlook other components of the flora, particularly non-native taxa, leaves conspicuous gaps in our knowledge of local and regional plant geography. NO — we) ww K ~~” ACKNOWLEDGMENTS AND ADDITIONAL INFORMATION. The produc- tion and publication of the Nantucket flora represent a coopera- tive effort by the Massachusetts Audubon Society, Massachusetts Natural Heritage and Endangered Species Program, Nantucket Maria Mitchell Association, and The Nature Conservancy. Ken- dra Buresch and Carol Moffitt assisted with much of the docu- ment research, data input, and editorial portions of this project. In addition to the support of our own organizations and the co- sponsors who helped bring this work together, we are especially grateful for the generosity of the Cox Foundation, Inc. for fund- ing this project. Copies of the flora are available to libraries and serious researchers free of charge while supplies last. The com- puter database also is available on two 3.5 inch diskettes. LITERATURE CITED BICKNELL, E. P. 1908-1919. Ferns a ee plants of Nantucket. ne ees Bot. Club 35: 49-62, 181-201, 498; 36: 1-29, 441-456; 37: 51-72; 69-80, 103-133, 447-460; 39: hae 605-624; 41: 27-47, 2 ee 411-427; 42: 265— A 549-570; 44: 365-369, 383-387; 46: 423-440. aie PW. A . A. SorriE. 1996. A computerized database of vascular non- oe a from Nantucket, Tuckernuck, and Muskeget Islands. anne Audubon Society, Massachusetts Natural Heritage and Endan- 98 Rhodora [ Vol. 98 gered Species Program, Nantucket Maria Mitchell Association, and The Nature Conservancy. MACKEEVER, F.C. 1968. Native te Naturalized Plants of Nantucket. University of Massachusetts Press, Amherst, MA. MASSACHUSETTS NATURAL HERITAGE AND ENDANGERED SPECIES PROGRAM. 1992. Massachusetts list of endangered, threatened, and special concern species. Bos- ton, MA. Owen, M. L. 1888. A Catalogue of Plants Growing Without Cultivation in the County of Nantucket, Massachusetts. Gazette Printing Co., Northampton, MA. SoRRIE, B. A. AND P. W. DuNWibpbIE. 1996. The Vascular and Non-Vascular Flora of Nantucket, Tuckernuck, and Muskeget Islands. Massachusetts Audubon So- ety, Massachusetts Natural Heritage and Endangered Species Program, Nan- aa Maria Mitchell Association, and The Nature Conservancy. Nantucket, RHODORA, Vol. 98, No. 893, pp. 99-102, 1996 NEW ENGLAND NOTE SCLERIA RETICULARIS (CYPERACEAE) NEW TO NEW HAMPSHIRE DANIEL D. SPERDUTO New Hampshire Natural Heritage Program/The Nature Conservancy, Department of Resources and Economic Development, P.O. Box 1856, Concord, New Hampshire 03302 Scleria reticularis Michx., reticulated nut-rush, is a plant of wet, sandy soil and southern pine barrens ranging from_ tropical America and southern U.S., north mainly along the coast to New England, with disjunct extensions into the midwest and south cen- tral states of Arkansas, Oklahoma, Missouri, [llinois, Indiana, Michigan, and Wisconsin (Reznicek 1994). Throughout much of its range S. reticularis is found on damp, sandy shores of coastal plain ponds that have broadly fluctuating water levels. Sorrie (1994) reports on the distribution of these ponds in New England, noting their primary concentration on the glacio-fluvial deposits of southeastern Massachusetts, adjacent Rhode Island and discon- tinuous localities in Connecticut, inland Massachusetts and the Ossipee Lake region of New Hampshire and Maine. Records com- piled by the Massachusetts Endangered Species and Natural Heri- tage Program indicate that S. reticularis is extant and locally abun- dant on pond borders in 7 towns in Plymouth, Barnstable and Nantucket Counties, with 61 stations recorded as of 1989. At this time, documentation of new occurrences was discontinued. An ex- tant station in the Connecticut River valley in the south-central Massachusetts town of Ludlow, Hampden County, is disjunct from the main distribution. A single historic station at Winter Pond, Winchester, Middlesex County, has not been observed since 1933 and possibly is extirpated. There are 3 extant populations in Washington County in southern Rhode Island and an historic col- lection from Providence County to the north (Rhode Island Natu- ral Heritage Program). Thus the New Hampshire population dis- cussed here is nearly 75 miles (123 km) disjunct from the nearest extant population in Massachusetts. A small population of Scleria reticularis (Sperduto 2344 NHA, NEBC, MICH; Sperduto 3/62 NHA) was discovered on the sandy, shallow-peat upper shore of Rocky Hill Pond in Litch- field, Hillsborough County, New Hampshire on September 16, 1992 by the author, Lesley Sneddon and Mark Anderson. 99 100 Rhodora [Vol. 98 The majority of several hundred plants observed was found along a short section of the pond margin. Numerous other species characteristic of coastal plain pondshores in New En- gland were associated with the Sc/eria including Rhexia_ vir- ginica, Eleocharis tenuis, Agrostis hyemalis, Glyceria borealis, G. acutiflora, Juncus pelocarpus, J. canadensis, Drosera inter- media, Gratiola aurea, Hypericum boreale, H. gentianoides, Dulichium arundinaceum, Panicum spretum, P.. tuckermanit, Cyperus dentatus, Viola lanceolata, Rhyncospora_ capitellata, Bulbostylis capillaris, and Fimbristylis autumnalis. Other spe- cies elsewhere in the basin include Euthamia tenuifolia, Xyris difformis var. difformis, Vaccinium macrocarpon, Lindernia ana- gallidea, Scirpus smithit, Eleocharis smallit, E. olivacea, E. obtusa, Potamogeton bicupulatus, P. oakesianus, Nymphoides cordata, Utricularia radiata and Myriophyllum humile. A total of 80 species was recorded in the wetland habitats of the pond basin. Rocky Hill Pond is found in a 2.7 acre basin blanketed by one to several meters of sand deposits overlying deeper, fine-textured lake-bed sediments of the postglacial Lake Merrimack (Gold- thwait et al. 1969; J. E. Cotton, USGS, pers. comm.). This por- tion of the southern Merrimack River valley contains an exten- sive groundwater aquifer associated with the reworked glacio- fluvial and = glacio-lacustrine deposits of the former Lake Merrimack. The pond basin has a moderately shallow profile with a maximum depth of 1.2 meters below the upland margin. Inter- mittent observations over several years from 1991—1994 revealed widely fluctuating seasonal and annual water levels. The pond level rises every spring, occupying essentially the entire 2.7 acre basin, and rises and falls with seasonal variation in precipitation and groundwater levels. At its lowest observed point in Septem- ber 1993, the water depth was only 10—20 cm and occupied less than a quarter acre. In general, the upper borders of the basin (e.g., where Scleria was observed) had several centimeters of peat over a sandy muck layer to a depth of 0.2—0.3 m, over fine to coarse sands to at least 1.2 m (extent of auger). Some middle areas of the basin had a more rapid transition to sand, while the lowest part of the basin had a full meter of surface muck over sand. Or- ganic matter content determined by ignition of soil samples in a muffle furnace indicated a 63% organic matter content in the vi- cinity of greatest Sc/eria abundance. Several meters away in wet- 1996] New England Note 101 ter habitat where Sc/eria was sparse, organic matter content was 20%. A series of small wetland basins similar to Rocky Hill Pond can be found elsewhere in Litchfield and adjacent towns of the lower Merrimack River valley sandplain, ranging from single ba- sins less than | acre in size to complexes of basins totaling as much as 11 acres. Although detailed surveys have not revealed any other populations of Scleria reticularis, many of these wet- lands do harbor various combinations of the pondshore associ- ates mentioned above (Sperduto 1994). Rocky Hill Pond is unique among these basins with its broad gradient of wetland habitats ranging from semipermanently to temporarily flooded zones. Each zone occupies significant area due to the moderate slope of the basin profile. This diversity of conditions seems to contribute to the higher species richness as compared to other, often larger, basin complexes of the sandplain system. The discovery of Sc/e- ria reticularis and associated plants in New Hampshire’s lower Merrimack River valley expands the known range of coastal plain pondshore flora from distributions described by Sorrie (1994) for New England. During the 1980s several houses were constructed immediately around the pond. A road has been built approximately 10 meters from the pond’s west shore with a culvert directed into the basin. Direct water pumping from the pond and other local residential and commercial aquifer pumping have the potential to modify natural fluctuations. As with other coastal plain pondshores in the region, development and its associated secondary impacts may di- minish the potential long-term viability of the Scleria reticularis and other constituents of the coastal plain pondshore flora unless there are concerted local conservation efforts, including commu- nity cooperation (Keddy and Reznicek 1982; McCarthy 1987; Keddy and Wisheu 1989). ACKNOWLEDGMENTS. I thank T. J. Rawinski for his earlier field work in the lower Merrimack River valley which led to and in- spired my interest in NH’s coastal plain affinities. I also thank A. A. Reznicek for identification confirmation of Scleria collec- tions, and T. D. Lee for critical review of this note. The Nature Conservancy and the Environmental Protection Agency are grate- fully acknowledged for their support for coastal plain pondshore research in NH 102 Rhodora [ Vol. 98 LITERATURE CITED GOLDTHWAIT, J. W., L. GOLDTHWAIT, AND R. P. GOLDTHWAIT. 1969. The Geology of New Hampshire. Part I—Surficial Geology. New Hampshire Department of Resources and Economic Development. Concord, NH. Kebby, P. A. AND A. A. REZNICEK. 1982. The role - seed ea in the persistence of ones. s coastal op flora. Amer. J. Bot. 69(1): Ecology, Seine and conservation of coastal slain plants some ae principles from the study of Nova Scotia wetlands. Rhodora 91: .K. MCCARTHY A. 1987. bake ‘ani temporal distributions of species in two in- termittent ponds in Atlantic County, New Jersey. M.S. thesis, Rutgers Univer- ark, NJ. REZNICEK, A. A. 1994. The disjunct coastal plain flora in the Great Lakes region. Biological Conservation 68: 203-215 Sorrik, B. A. 1994. Coastal plain ponds in New England. Biological Conserva- tion 68: 225-253. SPERDUTO, D. S. 1994. Coastal plain pondshores and basin marshes in New Hamp- shire. New Hampshire Natural Heritage Program, Department of Resources and Economic Development, Concord, NH. RHODORA, Vol. 98, No. 893, pp. 103-104, 1996 BOOK REVIEW Sorrie, Bruce A. and Peter W. Dunwiddie. 1996. The Vascu- lar and Non-Vascular Flora of Nantucket, Tuckernuck and Muskeget Islands. 145 pp. Published jointly by the Massa- chusetts Audubon Society, the Massachusetts Natural Heri- tage and Endangered Species Program, Nantucket Maria Mitchell Association, and The Nature Conservancy, Nan- tucket, MA. ISBN 0-932691-18-8. The authors have compiled an extraordinary list of the recorded flora of Nantucket and its adjacent islands, to provide informa- tion for further study on floristic changes. In addition to the ba- sic flora and descriptions of community types, the book provides an excellent account of the history of botany on Nantucket (sur- prisingly, Fernald never visited the island). The authors of this book, through extensive research in 9 regional herbaria and 16 years of field work, have produced a list of 2165 taxa (including interspecific hybrids) documented from Nantucket and nearby is- lands, with 79 species of mosses, 19 liverworts, | hornwort, and 99 lichens. Muskeget and Tuckernuck, although small and with few plant community types, still support considerable plant spe- cies diversity (160 and 353 species, respectively). The Nantucket flora contains 51 state-listed endangered, threat- ened, or special concern species. A significant proportion (39%) of the flora consists of introduced species. This may in fact be an underestimate, as there are several introduced species that are not so designated in the text (Mentha spicata, M. piperata, Origa- num). The list also includes a number of species known only from cultivation, such as silver maple and dill. The authors note that 153 of the native species have not been collected during the past 60 years, with the question of whether this represents extirpation or lack of collection left for future research. For each listed taxon, the book lists available synonyms, com- mon names, habitat types, state list category, status as introduced or introduced from the mainland, whether the plant is invasive, and notes on frequency. Habitat types are not provided for many species of mosses, and there are no moss records from Muskeget or Tuckernuck. The Flora is largely free of typographic and printing errors. Separate indices are provided for vernacular and scientific names. Unfortunately, the vegetation maps provided use a different clas- 103 104 Rhodora [ Vol. 98 sification system than was developed by the authors. A comput- erized database, on Microsoft Access 2.0, is available on diskette from The Nature Conservancy. This allows searches by family, genus, species, variety/subspecies, author, common name, growth form, state rank, synonymy, habitat, location, and collected speci- mens. The nomenclature used is eclectic. The authors largely follow Gleason and Cronquist (1991), occasionally substituting names from Kartesz (1994) or Flora North America, Vol. 2. However, the authors have chosen to list taxa (particularly infraspecific taxa) that they consider valid but that are not recognized by Cronquist, Kartesz, or the FNA authors. This may cause prob- lems for many botanists attempting to correlate field observa- tions to the Flora. —LISA A. STANDLEY, VHB, Inc., 101 Walnut Street, Watertown, MA 02272. RHODORA, Vol. 98, No. 893, pp. 105-106, 1996 BOOK REVIEW Buckley, Ann and Theodore O. Hendrickson. 1996. Native Trees, Shrubs and Woody Vines of Cape Cod and the Islands. vi + 70 pp. The Natural History of Southeastern New Eng- land, University of Massachusetts Dartmouth Print Shop. Available from UMass Dartmouth Bookstore, Dartmouth, MA 02714. $12.95. Any book which attempts to explain any portion of the nat- ural world to the general public is desirable and important. This book(let), which is part of a series devoted to the natural history of southeastern New England, is particularly important because the large numbers of summer visitors who visit the area now have a source of information about, at least, the common native and naturalized woody plants. The authors apparently set a very limited goal for themselves, viz., to provide keys to, small line drawings of the twigs and leaves of, and very brief descriptive comments about, 125 com- mon woody species. What they have done is successful. It is un- fortunate that they did not try to do more. If they wish public support (e.g., money), scientists have an obligation to try to show the general public what it is that they find exciting about their field of study, what interesting information they uncover. The pub- lic is interested in information. It would have been nice if the readers could learn: — How many kinds of higher plants are native to Cape Cod and the Islands? How many of them are trees, shrubs and vines? (Svenson and Pyle’s Flora of Cape Cod, 1979 is apparently long out of print and not readily available to the public.) 2. How were the particular species selected? One wonders, for ex- ample, at the absence of Al/lanthus. It would be worth noting, in these days when species extinction is a topic of conversation, how quickly some exotics have be- come naturalized (and what did they replace?). ies) Ailanthus altissima, introduced to this country in 1784, was be- ginning to naturalize by 1870, and was noted as a roadside es- cape around New Bedford and Buzzards Bay in 1911. Berberis thunbergti, introduced into cultivation in’ North America about 1864, was still noted as a garden plant around New 105 106 Rhodora [ Vol. 98 Bedford and Buzzards Bay in 1911; it is not mentioned in the 7th edition of Gray’s Manual (1907). Berberis vulgaris (introduced in colonial times), which only re- ceived passing mention in the booklet, was widely distributed in 1907. Has it become more uncommon in the intervening 90 years? Lonicera japonica was introduced in 1806, naturalized from Connecticut to Florida by 1907, but was still only a garden plant around New Bedford and Buzzards Bay in I9I1. Robinia pseudoacacia was esteemed for ship building (in par- ticular for the production of trunnels [tree nails]) and for rot-resistant posts. A southern species, it was widely planted in New England by farmers. It seems, in many areas, not to spread by seed, so the clumps we find are memorials to some former farmer’s diligence. Rosa rugosa was introduced into cultivation in this country about 1845. It was noted, with Rosa mu/tiflora (introduced some- time before 1868), as a garden plant around New Bedford and Buzzards Bay in 1911. It was first noted as an escape in 1912. And this doesn’t mention the deforestation and reforestation of Cape Cod. There is so much that the visitor would find interest- ing. —GORDON P. DEWOLF, RD |, Box 9, 125 Long Hill Road, West Brookfield, MA O1585. RHODORA, Vol. 98, No. 893, pp. 107-109, 1996 NEBC MEETING NEWS February 1996. Judith Warnement, Head Librarian of the Botany Libraries at Harvard, spoke on “Harvard Botanists and Their Li- braries.” These five botanical libraries are closely tied to Harvard’s history and to the personalities and relationships among its botanists, since most libraries were started as the personal col- lections of Harvard botanists. The first botanical institution, the Botanical Garden, was started by Peck in 1807, and continued by Nuttall. Asa Gray succeeded Nuttall in 1842, having already demonstrated a passion for books. Before coming to Harvard, Gray had traveled to Europe and pur- chased 3700 books in Europe to start a botanical library at the University of Michigan. The Bussey Institute was started in Ja- maica Plain in 1871, and focused on farming, agriculture, and hor- ticulture. The Arnold estate, deeded to Harvard for the promotion of agriculture and horticulture, was used to establish an arbore- tum at the Bussey Institute. Gray assembled a group of botanists at Harvard, each of whom was the center of a different institution and library. These included Sereno Watson at the Herbarium, Farlow in cryptogamic botany, Goodall in economic botany and physiology, and Sargent at the Botanic Garden. Goodall developed the Botanical Museum, with its collections in economic botany, paleobotany, and the Glass Flowers. Sargent, who apparently did not get along with the oth- ers, moved to Jamaica Plain to develop the Arboretum. During his 55-year tenure, he built the Arnold Arboretum’s library, and met its budget shortfalls from his own fortune. Farlow’s large per- sonal library was given to Harvard on the condition that a fire- proof building would be constructed. In 1864, Gray gave his li- brary of 2200 volumes to the Botanical Garden. The modern Gray Herbarium library was shaped by its librar- ian, Mary Day, who worked under B. L. Robinson for 30 years. She created the library’s unique classification system, replacing an even more eccentric system of filing by size. She also devel- oped the Gray Index, and personally prepared 170,000 cards. Oakes Ames saw the libraries and botanical institutions through WWI and the Depression, created an economic botany library, and donated his orchid library and collection to Harvard. By the 1950s, a century of “rampant individualism” had cre- ated a dispersed collection of botanical institutions, including 11 organizations funded by 9 separate endowments. Irving W. Bailey 107 108 Rhodora [ Vol. 98 recommended their consolidation into a single institution and building. The Friends of the Arnold resisted consolidation, and initiated a lawsuit that was settled after 13 years of litigation. De- spite the ongoing legal action, the libraries of the Gray Herbarium, Arnold Arboretum, and Economic Botany, the Ames Orchid Li- brary, and the Farlow Library were merged in the mid-1950s. These libraries are now managed by a single staff, but are main- tained in accordance with various bequests and a court order. To- gether, there are 230,000 volumes in Cambridge, and an additional 35,000 volumes at Jamaica Plain. The five different classification systems are being converted to the Library of Congress system, and are being catalogued onto the electronic database at a rate of 10,000 volumes a week. Judy ended her talk with a quote from George Bentham, cor- recting our plant-centered viewpoint and reminding us that “A bo- tanical library is useful without a herbarium, but not a herbarium without a library.” March 1996. Outgoing President Barre Hellquist spoke on “Non- native and Weedy Aquatic Plants of Northeastern United States.” Over the years, he has experimented with growing native and in- troduced species, and explored an interest in the aggressive qualli- ties of aquatic species. The USDA has listed several noxious weeds which it is illegal to possess, including Azolla pinnata, Eichhornia azurea, Lagaro- siphon major, and Salvinia molesta. Despite these prohibitions, several of these are already present in parts of the U.S., and can be obtained from suppliers or institutions. Botanical gardens ap- pear to have been the source of many problem introductions. This group includes Hydrocharis, Marsilea quadrifolia, Myriophyllum spicatum, Najas minor and Potamogeton crispus. Trapa natans, the only plant legally outlawed in Massachusetts, was first recorded at the Cambridge Botanical Garden in 1877. Emergent aquatics such as Butomus, Iris pseudacorus, and Lythrum salicaria are more attractive nuisances, and may not pose as much of a threat to aquatic systems. Phragmites is the most aggressive species in this group, and is almost impossible to eradi- cate. The debate over whether this species is introduced or native has not been resolved—the first New England record is from 1879 near Hartford, but specimens were collected in Texas in 1828. South American aquatics contain some of North America’s worst weeds, but are not found in New England—yet. Several of 1996] NEBC Meeting News 109 these (Alternanthera, Eichhornia, Pistia) cannot survive winter conditions. Others, including Egeria densa and Myriophyllum aquaticum, occur in New England but are sparse and localized. Barre identified several non-native species adapted to cold wa- ter that may be of concern as possible weeds. Aponogeton dis- tachyus, Nelumbo nucifera, Typha laxmaniti and Typha minima have aggressive growth forms and are tolerant of New England climates. Several North American natives (outside of New En- gland) also have weedy tendencies and could be a problem here. Cabomba caroliniana, first recorded from Rhode Island in 1892, has become a widespread problem in acidic ponds of southern New England and in the Albany area. Myriophyllum heterophyl- lum is a robust native first reported in 1932 in Connecticut. Re- cently outcompeted there by Eurasian Milfoil, it is now spread- ing in New Hampshire. Threats from the aggressive aquatics lurk in every pond. Even the attractive water lilies can become a nuisance unless grazed by moose, and Typha latifolia requires control by muskrats to keep it from dominating shallow aquatic sites. Potamogeton amplifo- lius, Coleogeton pectinatus, Lemna spp., Wolfia spp., and Utricu- laria (both inflata and purpurea) may all form dense stands in New England waters, reducing the value of ponds or lakes for swimming and boating. Barre closed with the question of how conservation groups should regard those native aquatics that have recently expanded their distributions into New England states: as rare—or as weeds’? —LISA A. STANDLEY, Recording Secretary. RHODORA, Vol. 98, No. 893, pp. 110, 1996 NEW ENGLAND BOTANICAL CLUB GRADUATE STUDENT RESEARCH AWARD The New England Botanical Club will offer an award of $1,000 in support of botanical research to be conducted by a graduate student in 1997. This award its made annually to stimulate and encourage botanical research on the New England flora, and to support visits to the New England region by those who would not otherwise be able to do so. The award will be given to the graduate student submitting the best research proposal dealing with systematic botany, biosystem- atics, plant ecology, or plant conservation biology, although re- search in other areas will be considered. Applicants must submit a proposal of no more than three double-spaced pages, a budget (the budget will not affect the amount of the award), a curriculum vitae, and two letters in sup- port of the proposed research, one from the student’s thesis advi- sor. Three copies of the proposal must be submitted. Proposals and supporting letters must be received no later than March 1, 1997. The recipient will be notified by April 30, 1997. Send proposals to: Awards Committee, The New England Botani- cal Club, 22 Divinity Avenue, Cambridge, MA 02138. The 1996 Graduate Student Research Award was presented to Bruce Henning Lindwall, University of Massachusetts (Amherst), for his study titled “The genetic consequences of long-term habi- tat fragmentation.” RHODORA, Vol. 98, No. 893, pp. I11, 1996 THE 1996 JESSE M. GREENMAN AWARD The 1996 Jesse M. Greenman Award has been won by Paul Kores for his publication “A systematic study of the genus Acianthus (Orchidaceae: Diurideae),” published in Allertonia 7: 87—220 (1995). This study is based on a Ph.D. dissertation from Tulane University under the direction of Dr. Steven P. Darwin. The Greenman Award, a certificate and a cash prize of $1,000, is presented each year by the Missouri Botanical Garden. It rec- ognizes the paper judged best in vascular plant or bryophyte sys- tematics based on a doctoral dissertation published during the pre- vious year. Papers published during 1996 are now being accepted for the 29th annual award, which will be presented in the sum- mer of 1997. Reprints of such papers should be sent to Dr. P. Mick Richardson, Greenman Award Committee, Missouri Botani- cal Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. In order to be considered for the 1997 award, reprints must be received by 1 June 1997. 111 RHODORA, Vol. 98, No. 893, pp. 112-113, 1996 STUCKEY PRESENTS ENDOWMENT GIFT TO OSU STONE LABORATORY Ronald L. Stuckey, Professor Emeritus of Botany at The Ohio State University, announced his donation of a $15,000 contribu- tion to the Franz Theodore Stone Laboratory, on the occasion of the Laboratory’s 100th Anniversary Celebration on Gibraltar Is- land, Put-in-Bay, Ohio, 24 August 1996, Dr. Stuckey’s contribution was made in November 1995 and approved by the Board of Trustees of The Ohio State University, 2 February 1996. The donation establishes an endowment whose income will provide annual financial support to employ a faculty- level individual to teach a course in vascular aquatic plants. The aquatic plants course was Professor Stuckey’s chief instructional responsibility at the Laboratory when he taught there from 1966 to 1991. He offered the course 18 of the 25 summers he worked at the Laboratory. Stuckey chose to announce the gift in the form of a poem he wrote, titled, “Of Life, Friends, Death, and Giv- ing.” To show their appreciation for the gift, officials presented Pro- fessor Stuckey with two awards. The Director of the Laboratory, Jeffrey M. Reutter, honored Stuckey with a Partnership Award, a plaque with the inscription, “/a Recognition of Your Numerous Contributions to Stone Laboratory and Your Efforts to Perma- nently Endow a Faculty Position For Your Course—Higher Aquatic Plants, 24 August 1996.” The second award was given by Michael Ahern, 1995—1996, President of the Friends organi- zation. This group, who simultaneously were holding their annual meeting, consists of former students, faculty, staff, and others who volunteer their time, make financial gifts, or offer other kinds of donations to the Laboratory. Dr. Stuckey’s second plaque was in- scribed, “For your outstanding support and dedication to Stone Laboratory.” The award also carried the title “Centennial, 1896— 1996, 100 Years of Education and Research at Franz Theodore Stone Laboratory.” Stuckey’s endowment, the eighth one estab- lished by the Friends organization for the Laboratory, is the first one for a specific course of instruction, and is believed to be the first of its kind in the country for a course in aquatic plants. Dr. Stuckey’s major research at the Laboratory focuses on the changes in abundance and distribution of aquatic and wetland plants in western Lake Erie. In 1971, Stuckey published a paper 112 1996] Announcement 113 in the Ohio Journal of Science documenting that 50% of the aquatic plant species in Put-in-Bay Harbor had disappeared since 1900. With the invasion of the zebra mussels, the water is much clearer and cleaner now, and several of the species have returned following this environmental change. This recent study, published in the same journal (1995) is co-authored by Stuckey and his former student, David L. Moore, who now teaches the aquatic plants course at the Laboratory. Their paper represents the first known study of its kind documenting the return of aquatic plants to a site where they once had disappeared. Their work has been reported recently in the Cleveland Plain Dealer, Chicago Tribune, Detroit News, and in the magazines Business Week and Discover. Stuckey has participated in the Laboratory during five decades, first as a student while he attended Heidelberg College (1959), as a faculty member (1966—1991), and as the Associate Director for the Educational Program (1977-1985). He is currently writing the 100-year history of the Laboratory. INFORMATION FOR CONTRIBUTORS TO RHODORA Submission of a manuscript implies it is not being considered for publication simul- taneously elsewhere, either in whole or in par GENERA anuscripts should be submited in triplicate. The text must be double- spaced throug Haut, inc ke tables, figure legends, and literature citations. Use a non- ha font throughout and do not wustify the right margin, Do not indicate the style Eee | thr yugh the use of capitals, underscoring, or bold, except for names of genera and species which should be in italics or underscored throughout. Do not underline punc- tuation, All pages should Be uals in the upper right-hand corner. For sues in matters not addressed here, sult the editorial office by phone at (603) 862- , FA (603) 862-4756, or e-mail: pete aae unh.edu. Brevity is urged for 2 submissions TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital let . Omit au- thors of scientific names. Below title, include author(s) name(s), affil a and ad- dress(es). If “present address” is different, it should follow immediately below, not as a STRACT: An abstract and a list of key words should be included with each paper, for shorter papers submitted as Notes. An abstract must be one paragraph, and should not include literature citations or taxonomic authorities. Please be concise, while including information about the paper’s intent, materials and methods, results, and sig- cae of findings. ain headings are all capital letters and centered on one line. Examples are: MATERIALS ene METHODS, RESULTS, and DISCUSSION. Do not title the Intro- duction. Do not combine sections of the paper (such as Results and Discussion), or use Conclusions or pes Second level headings should be in bold, indented, upper and lower case, and end with a period. Taxonomic authorities should sited for all species names at their first usage in the text, or in a referenced table. Cite ea figure and table in the text in numerical order. Each reference cited in the text must be in the Literature Cited. Cross-check spelling of author(s) os and dates of publication, ne rature Cl- tations in the text should be as follows: Hill (1982) or (Hill 1982). For two or more authors, cite Angelo and Boufford (1996) or (Angelo and Boufford 1996). Within paren- the use a semicolon to ny le different types of citations (Hill 1982; Angelo and Houtiowl 1996) (Figure 4; Table FLORAS AND TAXC One TREATMENTS: reece citation should be selected critically, especially for common species of broad distributio eys and synonymy for systematic revisions should be eae in the style of “A Nese of the Genus Mal- vastrum,” S. R. Hill, RHODORA 84: 159-264, 1982. Designation of a new taxon should carry a Latin diagnosis (rather than a im Latin description), which sets forth succinctly how the new ety di ee rom its congeners. ATTERATURE CITED: All bibliographic entries must be cited in the paper, unless a special ae ion has ant made by the Editor (such papers w will be allowe d a REFER- ENCES section). Verify all entries hae original sources, paying special attention to spelling and details of publication. Cite references in strict siphaEeEGaL order by firs author’s surname. Do not write authors’ names in all capital letters. References a cae author precede multi-authored works of same senior author, regardless of date. Use a long dash when the author(s) are the same as in the entry immediately eee (see recent issues). Refer to Botanico-Periodicum-Huntianum (B-P-H, 1968) and B-P-H/ ss ie) for standardized abbreviations for journals. TABLES: Tables must be double-spaced. Tables may be continued on an extra page, if eo As much as possible, the ttle should be self-explanatory. Do not use foot- notes; ins tead, add notes after the end of the table title. Broadside ane should be ided, if possible. Each table should be cited in the text in numerical order. “Fl UR ES: Illustrations must be either black and white half-tones (photograph), draw- ings, or graphs. Illustrations must be camera-ready; flaws cannot be corrected by the Edi- tor or the printer. Add symbols or shading with press-on sheets. The printed plate will be 4 * 6 inches; be sure that illustrations are proportioned to reduce correctly. Allow space orac rine if possible. Magnification/reduction values should be se to reflect the actual printed size. Maps must indicate scale and compass direction. The double- spaced list i. legends for figures should be —— on a separate page. Each figure should be cited in the text in numerical order on = 114 THE NEW ENGLAND BOTANICAL CLUB 22 Divinity Avenue Cambridge, MA 02138 The New England Botanical Club is a nonprofit organization that promotes the study of plants of North America, especially the flora of New England and adjacent areas. The Club holds regu- lar meetings, and has a large herbarium of New England plants and a library. It publishes a quarterly journal, RHODORA, which is now in its 99th year and contains about 400 pages per volume. Visit our web site at http://www.herbaria.harvard.edu/nebc/ Membership is open to all persons interested in systematics and field botany. Annual dues are $35.00, including a subscription to RHODORA. Members living within about 200 miles of Boston receive notices of Club meetings. To join, please fill out this membership application and send with enclosed dues to the above address. Regular Member $35.00 Family Rate $45.00 Student Rate $25.00 For this calendar year ee For the next calendar year ee Name Address City & State Zip Special interests (optional): i Es, THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 1996—1997: President: W. Donald Hudson, Jr., Chewonki Foundation, RR 2, Box 1200, Wiscasset, ME 04578 Vice-President (and Program Chair): David S. Conant, Depart- ment of Natural Sciences, Lyndon State College, Lyndon- ville, VT 05851 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Bi- ology, North Adams State College, North Adams, MA 01247-4100 Recording Secretary: Lisa A. Standley Curator of Vascular Plants: Raymond L. Angelo Assistant Curator of Vascular Plants: Pamela B. Weatherbee Curator of Non-Vascular Plants: Anna M. Rei Librarian: Paul Somers Councillors: C. Barre Hellquist (Past President) Garrett E. Crow | Matthew Hickler (Graduate Student Member) 1997 Edward Hehre, Jr. 1998 Michael J. Donoghue 1999 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora Volume 97, No. 892 including pages 291-394 was issued January 3, 1997 OO SS a a — 7 ee ; — 7 re a - a - : - 7 : " oe : ~~ OO ee : RHODORA The Journal of the New England Botanical Club CONTENTS A classification of coastal heathlands and sandplain grasslands in Massa- chusetts. Peter W. Dunwiddie, Robert E. Zaremba, and Karen A. Harper 1 The vascular flora of Hopewell Culture National Historical Park, Ross County, Ohio. James P. Bennett and Jennifer E. J. Course ........... 146 Notes on the genus Scirpus sensu lato in Missouri. S. Galen Smith and George Yatskievych 168 The flora of Latimer Point and vicinity, New London County, Connecticut. Steven R. Hi 180 BOOK REVIEWS An Almanac of Botanical Trivia 217 A Revision of Heterotheca sect. ae eet (Nutt.) Harms (Compo itae: Astereae): The Prairie and Montane Goldenasters of ean America 218 NEBC MEETING NEWS 220 Information for Contributors 223 NEBC Membership Form 225 Bylaws of The New England Botanical Club, Inc. 226 NEBC Officers and Council Members ............. .......0e0 eee inside back cover ad oad Vol. 98 Spring, 1996 No. 894 Issued: May 28, 1997 The New England Botanical Club, Inc. 22 Divinity Avenue, Cambridge, Massachusetts 02138 RHODORA JANET R. SULLIVAN, Editor-in-Chief Department of Plant Biology, University of New Hampshire, Durham, NH 03824 MARGARET P. BOGLE, Managing Editor Department of Plant Biology, University of New Hampshire, Durham, NH 03824 Associate Editors HAROLD G. BROTZMAN THOMAS D. LEE CHRISTOPHER S. CAMPBELL LESLIE J. MEHRHOFF DAVID S. CONANT THOMAS MIONE GARRETT E. CROW LISA A. STANDLEY NANCY M. EYSTER-SMITH K. N. GANDHI—Latin diagnoses and nomenclature RHODORA (ISSN 0035-4902). Published four times a year (January, April, July, and October) by The New England Botanical Club, 810 East 10th St., Lawrence, KS 66044 and printed by Allen Press, Inc., 1041 New Hampshire St., Lawrence, KS 66044-0368. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send address changes to RHODORA, P.O. Box 1897, Lawrence, KS 66044-8897. RHODORA is a journal of botany devoted primarily to the flora of North America. Monographs or scientific papers concerned with systemat- ics, floristics, ecology, paleobotany, or conservation biology of the flora of North America or floristically related areas will be considered. SUBSCRIPTIONS: $75 per calendar year, net, postpaid, in funds pay- able at par in United States currency. Remittances payable to RHO- DORA. Send to RHODORA, PO. Box 1897, Lawrence, KS 66044- 8897. MEMBERSHIPS: Regular $35; Family $45; Student $25. Application form printed herein. NEBC WEB SITE: Information about The New England Botanical Club, its history, officers and councillors, herbarium, monthly meet- ings and special events, annual graduate student award, and the jour- nal RHODORA is available at http://www.herbaria.harvard.edu/nebc/ BACK ISSUES: Information on availability of back issues should be addressed to Dr. Cathy A. Paris, Department of Botany, University of Vermont, Burlington, VT 05405-0086. E-mail: cparis@ moose.uvm.edu. ADDRESS CHANGES: In order to receive the next number of RHO- DORA, changes must be received by the business office prior to the first day of January, April, July, or October. INFORMATION FOR CONTRIBUTORS: See contents. Submit manu- scripts to the Editor-in-Chief. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). RHODORA, Vol. 98, No. 894, pp. 117-145, 1996 A CLASSIFICATION OF COASTAL HEATHLANDS AND SANDPLAIN GRASSLANDS IN MASSACHUSETTS PETER W. DUNWIDDIE The Nature Conservancy, 217 Pine St., Suite 1100, Seattle, WA 98101 ROBERT E. ZAREMBA The Nature Conservancy, 91 Broadway, Albany, NY 12204 KAREN A. HARPER 9826 76th Avenue, Edmonton, Alberta, T6E 1K5 Canada ABSTRACT. A set of 372 ee relevés was collected from coastal sandplain communities on Nantucket, Cape Cod, and Martha’s Vin a Sites on Nantucket had the highest average number of rare species < e lowest number of non-native species. We used TWINSPAN to identify 1 major vegetation types from these data. The first division separated the samples into heathlands and grasslands. Heathlands subsequently were classified into two major types: 1) Tall Shrub, including Huckleberry-Scrub Oak (Gaylussacia baccata-Quercus ilicifolia) and Mixed Maritime Shrublands (G. baccata, Myrica pensylvanica, and others), and 2) Low Shrub, including Broom Crow- berry (Corema conradit) and Bearberry (Arctostaphylos uva-urst) Heathlands. o major grassland types were designated Hairgrass (Deschampsia flex- uosa) and Little Bluestem ets zachyrium scoparium), after the dominant taxa. Hairgrass Grasslands included a Beach subtype often dominated by coastal species a as es iligulata, Hudsonia tomentosa), and a Heathy Grassland that includes many ericads and graminoids. Little Bluestem Grass- lands were divided into several subtypes, including Weedy Grasslands with many introduced species, Pennsylvania Sedge Grasslands dominated by Car- ex pensylvanica, and High Diversity Native Sandplain Grasslands, which in- cluded many rare species, few exotics, and high species diversity. We discuss successional relationships among sandplain vegetation types. Key Words: oe grassland, coastal heathland, community classifica- tion, two-way indicator species analysis, Schizachyrium sco- parium, Arctostaphylos uva-ursi, Gaylussacia baccata, Des- champsia flexuosa, Quercus ilicifolia The easternmost grasslands in North America occur in the coastal sandplains of the northeastern United States from Long Island (New York) to Cape Cod (Massachusetts). Similar to mid- western prairies, they usually are dominated by Schizachyrium scoparium, but also include a distinctive association of primarily coastal taxa. Many regionally and globally rare plants occur in 117 118 Rhodora [Vol. 98 the sandplain grasslands, including such taxa as bushy rockrose (Helianthemum dumosum), yellow sandplain flax (Linum inter- cursum), sandplain blue-eyed grass (Sisyrinchium fuscatum), sandplain gerardia (Agalinis acuta), and eastern silvery aster (As- ter concolor). The grasslands also provide critical habitat for var- ious uncommon or declining fauna, including Short-eared Owls (Asio flammeus), Northern Harriers (Circus cyaneus), Upland Sandpipers (Bartramia longicauda), Grasshopper Sparrows (Am- modramus savannarum), and Regal Fritillary Butterflies (Spey- eria idalia). This community is referred to variously as maritime grassland (Reschke 1990), New England sandplain grassland (Maine Natural Heritage Program | ), or terrestrial Schiza- chyrium scoparium medium-tall graminoid vegetation (Sneddon and Metzler 1992). It is considered endangered throughout its range, where considerable efforts are being made to preserve and manage the remaining examples (Massachusetts Natural Heritage Program 1982; Rawinski 1984; Godfrey and Alpert 1985). The coastal sandplain grasslands frequently include a large component of woody species, many of which are ericads. Where heath taxa are dominant, the community has been termed mari- time or coastal heathlands (Reschke 1990), and can be found from New Jersey to the Canadian Maritimes. In Massachusetts, heath- land and grassland plants are often intermixed, and the commu- nities sometimes are not readily distinguished. They often share many species with other coastal communities, such as sand dunes and scrub oak/pitch pine (Quercus ilicifolia/Pinus rigida) barrens. he past extent of coastal grasslands and heathlands is uncer- tain. A few historical accounts describe the presence of maritime grasslands along the coast, and suggest that they may predate European settlement (Harper 1912). Furthermore, it is unlikely that the many rare native species found in the coastal sandplain communities arrived in the last 300 years, the period since set- tlement in this area by Europeans. However, fossil pollen studies on Cape Cod and Nantucket indicate that grasslands and heath- lands became extensive only following settlement (Winkler 1985; Tzedakis 1987; Dunwiddie 1989, 1990). The species found in these communities today most likely existed prior to European settlement in openings close to the coast where salt spray sup- pressed the growth of woody plants, and in openings created by windstorms, fires, and aboriginal agricultural activities. Forest clearance to create fields for crops and pastures quickly 1996] Dunwiddie et al.—Heathlands and Grasslands 119 followed settlement in the 1600s (Dunwiddie 1990). Cultivation of crops often was brief, as soils were exhausted quickly (Macy 1835), but grazing, fire, salt spray, and other disturbances prob- ably helped to maintain grasslands and heathlands in many areas for several hundred years. Introduced pasture grasses did poorly in the acidic, droughty soils, and native grassland species thrived in the pastures and abandoned fields. With the widespread aban- donment of sheep grazing in the late 1800s on Cape Cod, Nan- tucket, and Martha’s Vineyard many grasslands and heathlands have been overtaken by taller shrub and forest vegetation (Dun- widdie 1992, 1994; Dunwiddie and Adams 1994). Today, many conservation organizations are working to pre- serve rare grasslands and heathlands (Godfrey and Alpert 1985). A top priority has been to develop ecological management strat- egies for slowing or reversing the succession of shrubs and trees (Dunwiddie and Caljouw 1990). This management has been ham- pered by an incomplete understanding of the compositional, en- vironmental, successional, and historical relationships among these vegetation types. This study was undertaken to develop a classification of coastal grasslands and heathlands in Massachu- setts based on their vegetational composition. Using data sets from sites on Cape Cod, Nantucket, and Martha’s Vineyard, we differentiate major types of grasslands and heathlands in this re- gion, identify those types that are most likely to contain rare species, and propose likely successional relationships among them. METHODS The grasslands and heathlands on Nantucket, Martha’s Vine- yard, and Cape Cod (Figure |) all occur within a few kilometers of the coast on coarse-grained glacial moraine and outwash de- posits. Sites were chosen for sampling based on personal knowl- edge and on information from the files of the Massachusetts Nat- ural Heritage and Endangered Species Program. At each site we noted the area and extent of communities and made general de- scriptions of topography, land use, location, recent disturbance history (if known), and species composition. Locations for relevés were selected to characterize the range of vegetation types that occurred at each site. The initial data set for this study consisted of 233 relevés collected on Nantucket in 120 Rhodora [Vol. 98 Pad et? km SSS 0 10 20 = Cape Cod Cuttyhunk Noman’s Land Figure |. Locations in southeastern Massachusetts mentioned in the text. The larger grasslands and heathlands in which many relevés were located are indicated with shading. Small sites are not depicted. 1982. To maintain consistency among the data, we used the same methodologies to collect 119 additional relevés on Cape Cod and Martha’s Vineyard in 1992, and an additional 20 from Nantucket. This methodology consisted of listing all species encountered within a 10 m radius plot and ranking them on a dominance scale of | (low) to 4 (high). Dominance values (d.v.) of | corresponded to rare species in each plot with 50O percent. Values of 2 and 3 distinguished intermediate levels of abundance, with a cover of about 15 per- cent roughly separating these two classes. This methodology ad- equately distinguished dominant from minor species on sites and 1996] Dunwiddie et al.—Heathlands and Grasslands 121 yielded comparable results when applied by different investiga- tors. A total of 372 relevés, including 253 from Nantucket, 28 from 13 sites on Martha’s Vineyard, and 91 from 37 sites on Cape Cod, were ordinated and classified using two-way indicator spe- cies analysis (TWINSPAN; Hill 1979) from the Cornell Ecology Program. TWINSPAN distinguishes groupings of sites with sim- ilar species associations and is particularly helpful where the high number of species and samples may obscure elements in common between sites (Jongman et al. 1987). Community classifications make more intuitive sense when based on common species rather than on species that only occur infrequently, so we explored var- ious combinations of weightings of the dominance values. Several trials yielded similar results, and we present here results in which weights of 10 and 5 were attached to the more common species (d.v. = 4 and 3), and less common species (d.v. = 2 and 1) were left unweighted. We included all 274 taxa recorded from the relevés in the first trial TWINSPAN classification, and made several adjustments in subsequent trials. Species for which some taxonomic confusion in the field may have resulted in differential identification by the various data gatherers were combined into several composite “‘su- pertaxa,”’ such as species of Agrostis, mosses, and a few minor taxa. Lichens were omitted entirely as input to the classification since they were not identified to species. Ten relevés formed anomalous small groupings, and were discarded in the final clas- sification. Nomenclature follows Gleason and Cronquist (1991). For each of 20 Nantucket relevé sites, four replicate soil sam- ples were taken from the top 15 cm of the soil profile, excluding the litter layer. Soil pH was measured in 0.01 M CaCl,, and tex- ture (percent sand, silt, and clay) was determined using Bouy- oucos hydrometer methods. Exchangeable cations (magnesium, calcium, sodium, potassium) were measured by shaking samples in IN ammonium acetate for 30 minutes, and analyzing the su- pernatant using a Jobin-Yvon Inductively Coupled Plasma spec- trophotometer. Organic carbon:nitrogen ratios were measured us- ing a Perkin-Elmer 2400 CHN. RESULTS Most of the groups of relevés produced by the dichotomies in the TWINSPAN classification include recognizable associations LZ Rhodora [Vol. 98 Table 1. Percent frequency of species in grassland and heathland relevés. Vegetation types are derived from TWINSPAN classification (Figure 2): GL = Grassland, LB = Little Bluestem, W = Weedy, PS = Pennsylvania Sedge, = High Diversity Native Sandplain Grassland, Hg = Hairgrass, HG = Heathy Grassland, B = Beach Grassland, HL = Heathland, LS = Low Shrub, BB Bearberry, BC = Broom Crowberry, TS = Tall Shrub, HSO = Huckleberry/Scrub Oak, MMS = Mixed Maritime Shrubland. In general, spe- cies are arranged according to their decreasing frequency in several major TWINSPAN Group I A Al A2a A2b Vegetation Type GL LB Ww PS HDN Schizachyrium sc oo 92 96 95 92 99 Carex pensylvanica 384 9] 72 100 97 Arctostaphylos uva-ursi 76 62 15 50 9] Danthonia spicata 75 80 54 77 95 Lechea maritima 69 60 15 50 88 Myrica pensylvanica 68 72 77 50 77 Aster linaritfolius 68 60 33 54 76 Quercus ilicifolia 63 53 38 73 54 Aster paternus 61 67 26 54 93 Vaccinium angustifolium 61 71 56 81 74 Gaylussacia baccata 57 70 49 62 84 osa virginiana 52 64 4] 77 72 Deschampsia Jenene 50 25 21 54 18 Quercus prinoides 25 27 8 35 35 Hudsonia ericoides 46 35 5 27 54 ek iene one 19 22 18 35 20 Aster dun 47 68 69 58 72 Rubus Wie 43 55 74 50 47 Pinus rigida 4] 30 44 46 18 Epigaea repens 29 40, 23 23 55 Helianthemum dumosum 37 43 8 54 58 Viburnum dentatum 34 46 46 35 50 Solidago puberula 23 27 26 3] 26 Hieracitum venosum 38 38 8 19 61 Aronia arbutifolia 14 19 21 15 19 Helianthemum Abie ea 33 35 10 3] 50 Amelanchier nar citesilaals 13 17 8 23 20 Vaccinium palli a 10 11 18 19 4 Gaultheria oO enneee 8 12 13 15 9 Panicum spp. 40 37 31 38 4] Corema conradit 8 4 3 8 3 Corylus cornuta 3 3 0 4 4 Solidago nemoralis 51 57 4] 35 73 Chrysopsis a 51 42 10 38 61 Juncus greenei 43 55 62 54 53 OSTIS § 22 32 85 46 45 Pp. eins tenuifolia 31 45 4] 58 43 1996] Dunwiddie et al.—Heathlands and Grasslands 123 el. Extended. sandplain vegetation types identified in the column headings, including (top to bottom): Grasslands (1), Heathlands (II), HDN Sandplain Grasslands (A2b), Weedy Grasslands (A1), and Beach Grasslands (B2). The table includes only species that occurred in >25 percent of the relevés of at least one vegetation type, as well as a few taxa listed as endangered, threatened, or of special concern by the Massachusetts Natural Heritage and Endangered Species Pro- gram (1992). B Bl B2 II Cc Cl C2 D D1 D2 Hg HG B HL LS BB BC TS HSO MMS 84 93 70 69 76 79 71 64 74 45 74 9 42 81 82 88 71 81 91 62 98 100 94 84 100 100 100 73 94 34 66 91 24 54 75 79 65 42 56 17 83 79 9] 25 35 35 35 19 26 7 61 61 61 62 53 56 47 67 67 69 80 86 70 62 78 79 D2 72 14 79 84 70 90 98 97 100 86 98 62 52 80 3 a7 63 65 9 54 76 14 45 71 0 77 76 79 71 77 89 55 36 52 9 87 76 74 82 93 94 90 31 45 9 58 3] 26 4] 75 65 O35 88 80 100 58 86 9] 76 4] 52 21 20 30 3 63 63 50 88 63 80 31 64 13 48 57 75 74 76 46 61 17 13 18 6 52 55 47 71 51 a7 38 13 21 0 43 31 26 41 49 a2 45 25 2 12 43 25 24 29 53 4] 76 58 50 fe) 4] 80 79 82 17 22 7 10 16 0 36 27 15 53 41 50 24 Pe 39 6 33 37 32 47 30 39 14 16 25 0 31 20 29 0 37 37 38 18 29 0 31 39 35 47 25 39 0 37 48 18 29 29 35 18 29 41 7 8 13 0 26 20 15 29 30 28 34 30 46 3 25 3] 32 29 22 30 7 7 11 0 25 24 18 35 27 24 3] 9 14 0 25 27 24 35 24 31 10 2 4 0 22 22 15 35 23 35 0 45 39 55 21 39 35 47 10 13 E 15 14 15 20 51 26 100 | 2 0 2 4 0 10 8 6 12 12 7 21 42 59 12 16 24 26 18 1] 13 7 65 61 73 1] 14 2] 0 10 15 0 22 30 9 4 0 0 0 6 6 7 9 1] 6 3 0 0 0 5 2 10 8 i3 0 15 8 12 0 19 Ie ot 124 Rhodora [Vol. 98 Table 1. Continued TWINSPAN Group I A Al A2a A2b Vegetation Type GL LB Ww PS HDN Potentilla canadensis 23 32 23 15 43 Rubus hispic 28 40 49 19 42 Sisvrinchium seauin 15 22 3 8 36 Aster patens 17 21 3 23 30 Aster solidagineus 11 17 8 15 23 Linum intercursum 7 12 8 12 15 Aster concolor | 2 0 0 4 fatris: SCarlosa 5 6 0 4 11 Baptisia tinctoria 28 37 4] 42 32 Festuca ovina 37 44 72 38 31 Solidago rugosa 14 20 64 8 l Festuca rubra 16 19 56 12 3 Rumex acetosella 28 35 46 35 28 Toxicodendron radicans 23 27 44 15 22 Poa pratensis 11 15 44 12 | Anthoxanthum odoratum 11 15 4| 4 5 Juniperus virginiana 14 14 4] 8 3 Hypochoeris radicata 11 13 4] 8 0) Prunus serotina 29 26 38 23 20 Cirsium pumilum 17 24 36 IS 22 Potentilla simplex 14 21 36 19 14 olcus lanatus 7 10 33 0 l Hieracium piloselloides 6 9 28 4 | Panicum virgatum 6 10 28 12 0 ea odora 15 14 26 23 5 Hypericum perforatum 9 10 26 4 4 Panes lanceolata > 8 23 0 3 Asclepias syriaca 4 6 23 0 0 Asclepias tuberosa 3 5 13 8 0 trex umbellata 21 12 10 19 9 Ammophila breviligulata 2] 12 5 23 12 Prunus maritima 19 9 5 27 5 Asclepias amplexicaulis 6 4 5 8 I Solidago sempervirens 12 9 3 4 14 Hudsonia tomentosa 1S 2 3 4 | Polygonella articulata I] 4 0 4 5 Number of vascular species 26.5 29.6 32.8 26.4 29.0 Number of rare species 0.8 1.0 0.4 1.0 1.3 Number of non-native spp. 1.6 2.2 5.3 1.1 0.9 Number of ericaceous SEP: 4.1 4.2 3.0 4.0 4.9 Number of other 4.6 5.0 6.1 5.2 4.4 Number of cd spp. 6.4 6.9 8.9 6.8 5.9 Number of forb species 11.4 135 14.8 10.3 13.8 1996] Dunwiddie et al.—Heathlands and Grasslands 125 Table 1. Extended, Continued. B Bl B2 II Cc Cl C2 D D1 D2 Hg HG B HL LS BB BC TS HSO MMS 8 13 0) 10 6 9 0 13 17 7 9 13 3 19 4 3 6 28 30 24 4 7 10) 5) 4 6 0) 6 7 3 11 18 10) 15 18 2 12 13 15 10 l 2 0 | 2 O 6 l 2 0 0 0 O l O 0 0 l 2 O 10) 0) O 0) 0 0 l O 3 2 4 0) 8 6 3 12 10 13 3 15 20 6 18 24 12 47 14 19 7 26 32 15 Bi 4 6 0 2 2 3 3 2 6 2 0 O 0 4 6 0 11 > 21 ] 2 3 0) 0 0 0 18 14 24 ] O 0 0 1 2 0 17 14 21 2 3 0 I 2 0 3 5 0) ] 2 3 0 0 0 0 3 4 3 ] O 0 0 1 2 0 13 13 15 2 2 3 0 2 4 0 8 4 15 O 0 0 0 0 0 0 35 30 42 12 14 21 0 11 vf 17 4 D) 3 4 4 6 0 5 4 7 3 5 0 3 0 0 0 5 6 3 1 2: 0 0 O 0 0 0 0 0 1 2 0 0 O 0 0 0 0 0 O O 0 0 O 0 O O O 0 16 13 21 4 4 6 0 5 7 0 7 9 3 2 2 3 0 2 4 0 ] 2 0 | 0 0 O ] O 3 1 2 0 0 0 O 0 0 O 0 0 0 0 0 0 0 0 0 O 0 35 16 67 4 10 15 0 | 2 0 34 16 64 5 10 12 6 2 2 3 35 21 58 14 12 12 12 16 7 31 10 2 24 2. 6 9 O O O 0 17 4 39 | 2, 3 0) O O 0 35 7 82 l Z 3 O 0 O 0 21 4 52 | 4 6 0) 0 0 0 21.7 23.2 19.2 19.4 20.9 20.9 20.8 18.5 20.9 14.1 0.5 0.7 0.2 0.9 1.2 0.8 1.8 0.7 0.8 0.5 0.8 0.8 0.8 0.1 0.1 0.2 0.0 0.1 0.1 0.1 4.0 4.4 3.2 5.3 5.7 5.2 6.7 5.1 5.9 3.4 4.0 4.2 3.7 4.7 4.5 4.5 4.6 4.8 4.7 5.0 5.6 5.8 323: Bees, 4.] 4.3 3.8 2.8 3.2 2.0 8.2 8.8 7.0 6.1 6.5 7.0 5.6 5.8 re0) 3.7 126 Rhodora [Vol. 98 of species that are encountered frequently within the coastal sand- plain communities (Figure 2; Table 1). We have attached informal names that describe distinctive features of most groups, which also are distinguished by Roman numerals, letters, and numbers to differentiate hierarchical levels in the dichotomies. Distinguishing grasslands and heathlands. The first TWIN- SPAN division separates the samples into two groups that reflect graminoid-dominated sandplain grasslands (Group I, with 228 re- levés) and shrub-dominated heathlands (Group I], with 134 re- levés). Three of the five species that occur most frequently in the grasslands are graminoids, including Schizachyrium scoparium, Carex pensylvanica, and Danthonia spicata, whereas four of the top five species in heathlands are woody, including Quercus ili- cifolia, Gaylussacia baccata, Arctostaphylos uva-ursi, and Vac- cinium angustifolium, Nevertheless, there is significant overlap in species composition between these communities; seven of the ten most frequent species in each community are the same, and these species occur in over 60 percent of the relevés. These include A. uva-ursi, C. pensylvanica, S. scoparium, Q. ilicifolia, Aster lin- artifolius, Myrica pensylvanica, and V. angustifolium. Overall, the grasslands are considerably more species-rich than the heathlands; on average, 37 percent more species occur in the former (26.5 vs. 19.4 spp./relevé; Table 1). This richness is con- tributed largely by forbs, which are nearly twice as abundant in the grasslands than in the heathlands (11.4 vs. 6.1 spp./relevé; Table 1). Species considered to be endangered, threatened, or of special concern by the Massachusetts Natural Heritage and En- dangered Species Program (1992) have similar frequencies in both grassland and heathland relevés, occurring in 58% and 57% of the relevés, respectively. Thirteen of the 22 endangered, threat- ened, special concern, or watch list species found in grasslands or heathlands in Massachusetts were encountered in the relevés (Table 2). Grasslands also tend to have more introduced species than heathlands (1.6 vs. 0.1 spp./relevé; Table 1). Of the 45 spe- cies of non-native taxa found in the relevés, only 10 occurred in heathlands, and all of these were much more abundant in grass- lands. Relevés in Nantucket grasslands and heathlands tend to have more rare species (1.0 + 0.9) than those from either Cape Cod or Martha’s Vineyard (0.3 + 0.5; 0.7 + 0.7). They also tend 1996] Dunwiddie et al.—Heathlands and Grasslands 127 Table 2. Rare native plants occurring in coastal sandplain grasslands and heathlands. Species rank within Massachusetts (Mass. Nat. Her. & End. Spp Prog. 1990, 1992): E = Endangered, T = Threatened, SC = Special Concern, WL = Watch List. Species preceded by > were present in relevés in ie study. Occurrence data based on this study and personal records. G = Pre- dominantly sandplain grasslands, H = Predominantly heathlands, G-H = Found in both grasslands and heathlands, ! = Exclusively in one community, vé occurrences predominantly in HDN Sandplain Grassland (Group A2b). cur- Species Rank — rence Agalinis acuta (Sandplain Gerardia) E G! >Amelanchier nantucketensis Shay ans Shadbush) SC G-H Aristida purpurascens (Purple Needlegrass) T G Asclepias purpurascens (Purple Malkw eed) T G >Asclepias tuberosa (Butterflyweed) WL G >Aster concolor (Eastern Silvery Aster) E G-H* >Cirsium horridulum (Yellow Thistle) WL G-H* >Corema conradti (Broom Crowberry) SC H Panicum commonsianum (Commons’ Panic-grass) SC G Gnaphalium purpureum (Purple Cudwee E G! >Helianthemum dumosum (Bushy Rockrose) SC G-H Hypericum stragulum (St. Andrew’s Cross) E G-H Lactuca hirsuta var. sanguinea (Hairy wild- lettuce) T G Lechea minor (Thyme-leaved Pinweed) WL G-H fatris scariosa var. novae-angliae (New England Blaz- ing-Star) SC G-H* >Linum intercursum (Sandplain Flax) ts | G* >Lupinus perennis (Wild Lupine) WL G S| nuttallii (Nuttall’s Milkwort) WL G >Quercus stellata (Post Oak) WL G-H >Scleria pauciflora (Papillose Nut-sedge) E G >Sisyrinchium fuscatum (Sandplain Blue-eyed Grass) SC G* Spiranthes vernalis (Early Ladies’-tresses) Sc G to have fewer non-native species (0.4 + 0.7 vs. 2.4 + 3.7 on Cape Cod, and 2.4 + 2.9 on Martha’s Vineyard). The abundance of shrubs, particularly heath species, in the grasslands is noteworthy. There is little difference between grass- lands and heathlands in terms of the average number of ericads (4.1 vs. 5.3 spp./relevé; Table 1) or other shrubs (4.6 vs. 4.7). Comparisons of the percentage of relevés in which many of the smaller heath species occur illustrate this. Species such as Arc- tostaphylos uva-ursi (76 vs. 84%), Vaccinium angustifolium (61 vs. 77%), and Hudsonia ericoides (46 vs. 57%) are similarly com- 128 Rhodora [Vol. 98 Table 3. Characteristics of grassland and heathland soils on Nantucket. Only -vegetation types from which soil samples -were collected are included. Values + | SD. Sand Clay n pH (%) (%) GRASSLANDS All grasslands 44 3.52 +0.20 904+58 44+ 1.9 Pennsylvania Sedge GL (Group A2a) 24 3.55 £0.20 88.8 + 7.0 5.0 + 1.8 HDN Sandplain GL (Group A2b) 12 355+ 0.15 897+ 14 52 +08 Heathy Grassland (Group B1) S&S. 230202) 963204 1.7 2°10 HEATHLANDS All heathlands (Group II) 36 3.30 +£0.24 910+45 43+08 Low Shrub (Group C) 8 3.17 0.22 925+27 38 +03 Huckleberry-Scrub Oak (Group D1) 28 3.35 40.24 906+50 44+08 mon in both vegetation types. However, the dominance of taller shrub species in heathlands is pronounced, as demonstrated by the percent frequencies of several taxa in grasslands versus heath- lands: Quercus tlicifolia (63 vs. 90%), Gaylussacia baccata (57 vs. 87%), Quercus prinoides (25 vs. 63%), Comptonia peregrina (19 vs. 52%), and Aronia arbutifolia (14 vs. 26%). This pattern also is reflected in another measure, the average of dominance values, which is generally higher for these shrubs in the heath- lands (e.g., Gaylussacia baccata (d.v. = 2.2 vs. 3.2), Quercus ilicifolia (d.v. = 1.5 vs. 2.3). Samples collected from Nantucket suggest that heathland soils tend to be more acidic (pH 3.30 vs. 3.52, p < 0.005) and have higher organic C:N ratios (31.5 vs. 24.8, p < 0.005) than grass- lands (Table 3). Types of sandplain grasslands. Within the group of grass- land relevés (Figure 2; Table 1), the first dichotomy separates those in which Deschampsia flexuosa is a major component (Group B—Hairgrass Grasslands) from those in which Schiza- chyrium scoparium is the primary dominant grass (Group A— 1996] Dunwiddie et al.—Heathlands and Grasslands 129 Table 3. Extended. t Mg Ca Na (%) (ppm) (ppm) (ppm) (ppm) C:N 5224) 68240. 190-2140 Bet 16 6823.9 246 247 62255-62240 1692130 2e2 1s. Je=43 244254 S110 64244 1502108 222712 70234 232224 a) i) + So o) oO i) + oe) in vs) oS ue I+ ~ i) ion) I+ Na n be I+ w 28.2 + 3.0 Sotds 7024 2352167 2524) W237 3lo S46 3.7424 92+40 3634192 28+09 91434 3444 43 65 249 69243 225220) 2321) b4236 301244 Little Bluestem Grasslands). Although Schizachyrium scoparium is common in both groups (84% in Group B vs. 96% in Group A), Deschampsia flexuosa is not (88% in Group B vs. 25% in Group A). In general, the Hairgrass Grasslands tend to have fewer vascular species than the Little Bluestem Grasslands (21.7 vs. 29.6 spp./relevé), a pattern which is most noticeable among the forbs (8.2 vs. 13.5 spp./relevé). The frequencies of many rare species also are lower in the Hairgrass Grasslands: Sisyrinchium fuscatum (4 vs. 22%), Helianthemum dumosum (27 vs. 43%), Amelanchier nantucketensis (7 vs. 17%), Linum intercursum (O vs. 12%), Asclepias tuberosa (O vs. 5%), and Liatris scariosa vat. novae-angliae (2 vs. 6%). Hairgrass Grasslands are well repre- sented on Cape Cod and Nantucket, but appear to be scarce on Martha’s Vineyard, where two relevés were collected of this type (Table 4). Many Hairgrass Grasslands are dominated by a mixture of gra- minoids and heath species. The TWINSPAN dichotomy that sub- divided Group B distinguished a large subgroup of relevés, termed Heathy Grasslands (Group B1; Figure 2; Table 1), that had high frequencies of grasses (Schizacl ium scoparium—93%, Schizachyrium scoparium Festuca fa Lechea Ci sos jl Agro |_1- GRASSLAND (n=228) D - TALL SHRUB |Gaylussacia baccata |Quercus ilicifolia Il - HEATHLAND (n=134 Gaylussacia baccata Quercus ilicifolia C-LOWS ee HL (n=51) Corema conr Deschampsia Tan B - HAIRGRASS GL (n=89) Deschampsia flexuosa Arctostaphylos uva-ursi D2 - MIXED SHRUBLAND (n=29) —_ WwW Osa Spp. Prunus maritim oO Corylus cornuta Rubies teas D1- eee SCRUB OAK HEATHLAND (n=54) Arc Suva-ursi Aster paternus sin sa c HEATHLAND (n=17) Wal 4 Fre ea inne eae - BEARBERRY HEATHLAND (n=34 Arctostaphylos uva-ursi Hudsonia tomentosa Ammophila breviligulata a e) B1 - HEATHY GRASSLAND (n=56) 2. ster paternus Danthonia spicata = Vseciniinn sncucdiiol ta) v A2b - HDN SANDPLAIN GRASSLAND (n=74) Aster linariifolius - (n=100) Aster paternus | L (A - LITTLE BLUESTEM GL (n=139)_| |.A Schizachyrium scoparium ech ostis Aster dumosus Euthamia spp. Aster paternus Sisyrinchium fuse echea maritima eee uva-ursi rctostaphylos uva-ursi | A2a - PENNSYLVANIA SEDGE GRASSLAND (n=26) Carex pensylvanica Al - WEEDY a (n=39) Solidago rugos Poa sie i Holcus lanatus Anth th A, 86 1A] 1996] Dunwiddie et al.—Heathlands and Grasslands 131 Table 4. Summary of relevés by vegetation type from Nantucket, Cape Cod, and Martha’s Vineyard. % = Number of relevés in each vegetation type sampled in a particular region divided by the total number of relevés sampled in all regions for that vegetation type. Cape Martha’s Total Nantucket Cod Vv. n = 243) (n = 91) (n = 28) 362) No. % No. % No. % No. Heathland (Group I) 116 87 16 12 2 | 134 Tall Shrub (Group D) 81 98 0 O 2 2 83 Mixed Maritime (Group D2) 29 100 0 O GO: Ay 229 Huckleberry-Scrub Oak (Group D1) 52 96 0 O 2 4 54 Low Shrub (Group C) 35 69 16 31 Oo O 51 Broom Crowberry (Group C2) 14. 82 3 18 QO O 17 Bearberry (Group C1) 21 62 13 38 QO O 34 Grassland (Group I) 127 56 75 33 26 I1 = 228 Hairgrass (Group B) 41 46 46 52 2 2 89 Beach (Group B2) 0 O 31 94 2 6 33 Little Bluestem (Group A) 86 62 29 21 24 17 139 HDN Sandplain (Group A2b) 70 = 95 0 O 4 5 74 Pennsylvania Sedge (Group A2a) IS 58 7 27 4 15 26 Weedy (Group Al) | 3 22 5 16 41 39 Carex pensylvanica—93%; Danthonia spicata—91%; Deschampsia flexuosa—80%), ericoids (Arctostaphylos uva-ursi-100%, Hud- sonia ericoides—73%, Vaccinium angustifolium—71%), and other common heathland plants. Soil samples from sites within this subgroup (Group B1; Table 3) are significantly sandier, have a lower pH, and have lower exchangeable Mg:Ca and Na:K ratios than the other grasslands (p < 0.005). The other branch of the Hairgrass Grassland dichotomy iden- tified a smaller subgroup of relevés we termed the Beach Grass- lands (Group B2; Figure 2; Table 1). These grasslands often occur close to the coast in areas that receive some input of eolian sand Figure 2. TWINSPAN dendrogram of coastal sandplain relevé groupings. Informal names describe major distinguishing features of grassland (GL) and heathland (HL) vegetation types. Species identified by TWINSPAN to be ‘indicator species’’ are listed under each dichotomy. (n = number of relevés comprising group). [32 Rhodora [Vol. 98 and usually include such species as Ammophila breviligulata (64%), Hudsonia tomentosa (82%), Lechea maritima (91%), and several species of Cladonia and Cladina lichens. This group had the lowest diversity of any of the grassland types and generally did not contain any rare species. Little Bluestem Grasslands (Group A) include several types of coastal sandplain vegetation that were distinguished in the TWIN- SPAN ordination. Weedy Grasslands (Group Al) are character- ized by the greatest number of non-native species, with an av- erage of 5.3 spp./relevé (16% of the vascular plants), compared to 1.6 spp./relevé (6% of the vascular plants) for all grasslands (Group I). Common non-native species include grasses such as Festuca ovina (72%), Anthoxanthum odoratum (41%), Poa pra- tensis (44%), Holcus lanatus (33%), and Phleum pratense, and forbs such as Hypochoeris radicata (41%), Rumex acetosella (46%), Hieracium piloselloides (28%), and Hypericum perfora- tum (26%). Weedy Grasslands also tend to have fewer heath spe- cies than any other grasslands (3.0 vs. 4.1 ericads/relevé). Two native grasses, Schizachyrium scoparium and Agrostis species, are also common. These grasslands are frequent on Cape Cod and Martha’s Vineyard, but only one of the Nantucket relevés was of this type (Table 4). The dichotomy that distinguished subgroups within the Little Bluestem Grasslands contrasted the Weedy Grassland samples with a group of relevés (Group A2) that is noteworthy for the presence of Helianthemum dumosum, a species listed as “‘Special Concern” in Massachusetts (Mass. Natural Heritage and Endan- gered Species Program 1992). This species is found in both coast- al grasslands and heathlands, and occurred in over 50% of the relevés in this group. Group A2, which might be considered as the “‘non-weedy” Little Bluestem Grasslands, is best described by examining the two distinct subgroups of which it is comprised. Group A2a, Pennsylvania Sedge Grasslands, contains abundant Carex pensylvanica, occurring in 100 percent of the relevés with an average d.v. = 3.7. This type is more common on Nantucket and Cape Cod, with only a couple of locations sampled on Mar- tha’s Vineyard (Table 4). The second group (Group A2b) includes the largest number of native grassland species (28 spp./relevé), the most rare species (1.3 spp./relevé), and the fewest non-native species (0.9 spp./relevé) in any of the Little Bluestem Grasslands. We refer to it as High Diversity Native (HDN) Sandplain Grass- 1996] Dunwiddie et al.—Heathlands and Grasslands [33 land, because it provides the most diverse expression of the native grasslands, with a rich flora of rare species and few introduced taxa. In addition to the greater dominance of Carex in Group A2a than in Group A2b, these two grassland types also can be distin- guished from one another by the greater frequency of ericads (4.9 vs. 4.0 spp./relevé) and forbs (13.8 vs. 10.3 spp./relevé) in the HDN Sandplain Grasslands. This type was found most frequently on Nantucket, with only a few examples on Martha’s Vineyard, and none on Cape Cod (Table 4). Types of coastal heathlands. The first dichotomy within the heathland relevés (Figure 2; Table 1) separates those dominated by tall (>0.7 m) shrubs (Group D—Tall Shrub Heathlands) from others that are generally of low stature (Group C—Low Shrub Heathlands). Gaylussacia baccata and Quercus ilicifolia are com- mon in both groups, but both are more dominant in Group D (d.v. = 3.7 and 2.6) than in Group C (d.v. = 2.3 and 1.9). Most other tall shrubs found in these coastal habitats, including such species as Rosa virginiana and Rubus flagellaris, also occur with a great- er frequency in Tall Shrub Heathlands than in any other types. Tall Shrub Heathlands (Group D) are divided into Mixed Mar- itime Shrublands (Group D2) and Huckleberry-Scrub Oak Heath- lands (Group D1). The samples in our study come primarily from Nantucket, although both vegetation types occur frequently throughout the coastal sandplains of Cape Cod and Martha’s Vineyard. Many tall, non-ericaceous species are found most fre- quently in Group D2, including Aronia arbutifolia, Corylus cor- nuta, Prunus maritima, and Rubus flagellaris. This group tends to be the most species-poor of all heathlands (14.1 spp./relevé) and contains the lowest numbers of ericads (3.4 spp./relevé), gra- minoids (2.0 spp./relevé), and forbs (3.7 spp./relevé). Unlike other heathland types, where Quercus ilicifolia occurs in >86% of the relevés, oak is much less frequent in Mixed Maritime Shrublands (62%). As the name implies, Mixed Maritime Shrublands are ex- tremely variable in composition; quite different associations of shrub species may be lumped into this group. In many places, they probably would not be considered to be types of heathlands. However, the relevés that comprise Group D2 mostly include Gaylussacia baccata and Vaccinium angustifolium as dominants, and thus are arguably heathlands. The other Tall Shrub Heathland subtype, the Huckleberry-Scrub Oak Heathlands (Group D1), is 134 Rhodora [Vol. 98 noteworthy in having the greatest dominance of both Quercus ilicifolia (d.v. = 2.8) and Gaylussacia baccata (d.v. = 3.7) of any heathland type. In some cases, this type approaches the com- position, structure, and appearance of many scrub oak barrens. e other main group of heathlands, Low Shrub Heathlands (Group C), is dominated by Arctostaphylos uva-ursi (100% fre- quency, d.v. = 3.6), Hudsonia ericoides (75%, d.v. = 2.4), and a diversity of other low ericoid shrubs, such as Corema conradii and Vaccinium angustifolium. Compared to the Tall Shrub Heath- lands, both grasses (4.1 vs. 2.8 spp./relevé) and forbs (6.5 vs. 5.8 spp./relevé) are more frequent in this vegetation type. Low Shrub Heathlands are well developed on Nantucket and Cape Cod, but are infrequent on Martha’s Vineyard (Table 4). Low Shrub Heathlands were divided into two types. Arcto- staphylos uva-ursi is extremely dominant (d.v. = 3.9) in the Bear- berry Heathlands (Group C1). Although bearberry occurs in most heathlands and many grasslands, it reaches its greatest abundance in this type. Many examples exist within the Cape Cod National Seashore and on Nantucket. Broom Crowberry Heathlands (Group C2) have many of the same species as Bearberry Heath- lands, but are strongly dominated by the rare shrub Corema con- radii (100%, d.v. = 4.0). This vegetation type is most common on Cape Cod, with several sites on Nantucket (Table 4). DISCUSSION The results of a TWINSPAN classification of relevés from grasslands and heathlands on Cape Cod, Martha’s Vineyard, and Nantucket highlight the compositional variability of the shrub, grass, and forb-dominated vegetation types found in this region. The fact that seven of the ten most frequent taxa are shared by both communities emphasizes the degree of overlap in these veg- etation types. While some sites are relatively distinct, others in- corporate so many taxa from both communities that it may be arbitrary into which type they are classified. The inclusion of samples from other coastal grasslands and heathlands in the Northeast would further clarify the taxa that define the communities described in this classification. Data from such well-known sites as the Hempstead Plains and Montauk in New York (Taylor 1923; Conard 1935; Reschke 1990), as well as lesser-known examples from Rhode Island and Connecticut, 1996] Dunwiddie et al.—Heathlands and Grasslands 135 would likely expand the number of types beyond those we have described. Even within Massachusetts, some types were not rep- resented in our samples, including the Panicum virgatum grass- lands of Noman’s Land and Cuttyhunk (Elizabeth Islands) and the Smilax-Gaylussacia heathlands and Myrica_ pensylvanica shrublands elsewhere in the Elizabeth Islands (pers. obs.). The distribution and extent of grasslands and heathlands on Nantucket, Martha’s Vineyard, and Cape Cod can be evaluated from various vegetation maps of these areas (Harshberger 1914; Robertson 1973; MacConnell et al. 1984; Nature Conservancy 1995). However, the different vegetation types identified in these studies make it difficult to compare the grassland and heathland types we describe. On 1980 aerial photos, MacConnell et al. (1984) delineated “‘heath plant community” acreages that amounted to about 576 ha on Cape Cod, 280 ha on Martha’s Vineyard, and 3638 ha on Nantucket. Their definition appears to have lumped most of the grassland and heathland vegetation types we sampled. MacConnell et al.’s Nantucket figure compares fa- vorably with a total of 3239 ha reported in a GIS study by The Nature Conservancy (1995) based on 1978 aerial photographs. Although our selection of relevé sites was neither random nor exhaustive, we collected data from most major native grasslands and heathlands on Cape Cod, Martha’s Vineyard, and Nantucket (Dunwiddie et al. 1993). The number of relevés sampled in each of the three regions is similar to the relative acreages reported by MacConnell et al. (1984). Thus our relevé data from different vegetation types, summarized by region (Table 4), may provide a general indication of the occurrence and abundance of these communities in southeastern Massachusetts. These data suggest that most types of grassland and heathland are found throughout the areas sampled, but that significant regional variation exists. For example, the grassland type with the greatest concentration of rare species (HDN Sandplain Grassland, Group A2b) occurs primarily on Nantucket. Weedy Grasslands (Group A1), in con- trast, are more prevalent on Cape Cod and Martha’s Vineyard. Hairgrass Grasslands appear most frequently on Cape Cod, and are uncommon on Martha’s Vineyard. The great variability in the composition and distribution of these communities presents an intriguing and difficult problem in understanding the origin, maintenance, and dynamics of the coastal sandplain vegetation. Why grasslands develop in some 136 Rhodora [Vol. 98 areas while heathlands and maritime shrublands predominate in others, or why some sites appear to be relatively stable while others succeed to forests within a few decades, are questions that may have several answers in different places. A combination of factors probably is important in most areas, including differences in climate, substrate, disturbance and land use history, and initial floristic composition. Proximity to the ocean was considered as one factor that might affect the distribution of vegetation types. Wind and salt spray play an evident role in suppressing arboreal species near the coast (Boyce 1954), which may explain the prevalence of both grass- lands and heathlands within a few miles of the shoreline. How- ever, we could find no clear relationship between the occurrence of different vegetation types identified in our study and their dis- tance from the ocean. For example, Bearberry Heathlands are abundant on coastal bluffs on Cape Cod, while Little Bluestem Grasslands flourish close to the Nantucket shore. Only the Beach Grassland (Group B2) showed a tendency to occur primarily within a few hundred meters of the shore. We also examined whether grassland and heathland types occur preferentially on different substrates. However, a categorization of sites as located on either glacial moraine or outwash revealed no correlation; the different vegetation types occur on both types of deposits. Comparisons of soils collected from some Nantucket grasslands or heathlands showed no significant differences in available cations or particle size (Table 3), also suggesting that underlying parent material composition is not controlling the dis- tribution of these vegetation types. Additional soil samples from Cape Cod, Martha’s Vineyard, and other coastal grassland and heathland sites are needed to determine whether these patterns can be generalized throughout the region. Several differences in soil characteristics are worth noting. The significantly higher organic C:N ratios in heathland soils than in grasslands may largely result from biotic activity. Grassland spe- cies tend to have a higher percent N in tissues (lower C:N ratio), generally producing soils with relatively high exchangeable N; perennial woody species often produce litter with a lower percent N in tissues, giving rise to soils with a higher organic C:N ratio and lower N availability. Within the grassland samples, the data provide some evidence that the occurrence of at least one of the grassland types may 1996] Dunwiddie et al.—Heathlands and Grasslands 137 relate to soil parameters. The samples from the Heathy Grasslands subgroup (Group B1) are significantly sandier, more acid, and have lower Mg:Ca and Na:K ratios than the other grassland sam- ples. The other grassland type in the Hairgrass group (Beach Grasslands, Group B2) might also be expected to have a similar pattern of occurrence on droughty, low nutrient soils. Both of these grassland types have lower species diversity than the rela- tively species-rich Little Bluestem Grasslands. Our results suggest that substrate and geographic location play roles in the distribution of some of the coastal vegetation types. However, clear environmental correlates do not emerge to explain many of the patterns of grassland and heathland distribution we observed. Instead, we hypothesize that much of the variation in composition and distribution of coastal sandplain grasslands and heathlands reflects differences in the disturbance and successional histories of sites, rather than resulting from differences in the physical or environmental characteristics of sites. According to this hypothesis, the mosaic of current vegetation on the coastal sandplain documented in our relevés is largely attributable to dis- turbances of different types and intensities occurring at different times and frequencies. Each disturbance acts upon a unique as- semblage of species that is continually reshaped at each site. A similar situation has been described in midwestern prairies, where sites that have a recent history of fire and grazing support dif- ferent communities than comparable sites where these distur- bances are absent (Curtis 1959). This hypothesis needs to be examined using sites with a long and detailed land use history. Such an examination would be difficult, however, as most grasslands and heathlands in south- eastern Massachusetts are many decades removed from the period when grazing, fires, and agriculture were actively shaping the vegetation. Records of 18th and 19th century fires, land use, and vegetation that are tied to specific sites are scant, and were unob- tainable for most of our relevé sites. While it is likely that all of these coastal sites received some degree of livestock grazing dur- ing the last 300 years, plowing histories may differ considerably between locations. Unfortunately, we did not collect data on the presence of plow horizons and other evidence of past soil tillage. In the absence of detailed, site-specific land use histories for the relevé sites, we have drawn upon photographic studies on Nantucket (Dunwiddie 1992), Martha’s Vineyard (Dunwiddie Plowed Fields Weedy GL (1) Low (C Bearberry HL (C1) Broom Crowberry HL (C2) Pennsylvania Sedge GL (A2a) Other Little Bluestem GL (A2) | HDN Sandplain GL (A2b) | orest , Tall Shrub Heathlands (D) H uckleberry-Scrub Oak HL (D1) Mixed Maritime Shrubland (D2) [eo hk OaL/D; ace h Pine Barrens Black/White Oak Forest Oak/Pitch Pine Forest Figure 3. Proposed successional relationships of major coastal sandplain vegetation types. GL = Grassland; HL = Heathland. eIOpOyY 86 10A] 1996] Dunwiddie et al.—Heathlands and Grasslands 139 1994), and Cape Cod (Dunwiddie and Adams 1994) to provide anecdotal information on how different vegetation types have changed over time in this region. Combining this information with our observations of the distribution of sites and a regional knowledge of past land use, we developed a model of the suc- cessional relationships among coastal vegetation types (Figure 3). Agriculture was widespread throughout the coastal sandplain, and many sites no doubt were plowed, at least for a while. Thus our model depicts several successional pathways beginning with plowed fields. Factors that determine which pioneering commu- nities develop on bare ground may include the type and extent of disturbance creating the bare ground, the composition of both the prior and surrounding vegetation, the current land use, and differences in substrate. For example, many agricultural fields and improved pastures develop into some form of the Weedy Grass- land (Group Al), especially if they have received additions of lime and fertilizer and have been cultivated for sufficient time to eliminate much of the native seed bank. These areas may increas- ingly resemble the other Little Bluestem Grassland types (Group A2) if sources of native seed are available, and fires, salt spray, grazing, or other factors prevent extensive shrub encroachment. Otherwise, they may succeed to various Tall Shrub Heathlands (Group D) Weedy Grasslands are less likely to develop in agricultural sites that received little or no nutrient or lime amendments and in pastures that were created by clearing open woodland or shrub- land and were never plowed. Instead, these sites may develop > Figure 4. (a) The ee of sandplain grassland and heathland veg- etation on Nantucket is visible in the photograph, taken about 1890. Light areas are dominated by nee flexuosa and Schizachyrium scoparium. Small dark cushions in the foreground are Hudsonia ericoides; dark patches in the distance are primarily Gaylussacia baccata clones. Heavy grazing by sheep and other livestock had declined in this area about 40 years earlier, although peta animals may have continued to graze when this photo was taken (reprinted courtesy of the Sigkenee pape: Association). (b) Photograph taken from the same location a y later, showing an increase in taller woody plants, especially eae ae Many areas of Nantucket, Cape Cod, and Martha’s Vineyard have experienced much more rapid suc- cession of shrubs and trees into the me and heathlands than depicted in these photographs. [Vol. 98 Rhodora 140 14] Dunwiddie et al.—Heathlands and Grasslands 1996] 142 Rhodora [Vol. 98 other types of grassland or Low Shrub Heathland. Old photo- graphs and some contemporary examples of unimproved grazing land on Nantucket, Martha’s Vineyard, and the Elizabeth Islands suggest that they frequently develop into types of Little Bluestem Grassland (Group A; Dunwiddie 1992, 1994). The high diversity HDN Sandplain Grasslands (Group A2b) may develop from these sites if they are not overtaken by woody plants. On Cape Cod, Hairgrass Grasslands (Group B) appear to have developed more frequently in pastured land than in the other locales. It is not clear under what conditions the Pennsylvania Sedge dominated (Group A2a) type develops, although it may be related to particularly heavy grazing. The high organic C:N ratios observed in Low Shrub Heathlands suggest that nitrogen may be particularly lim- iting in these sites; Broom Crowberry (Group C2) and Bearberry Heathlands (Group Cl) often appear to develop in areas where the topsoil is removed, and the subsoil is widely exposed. Tall Shrub Heathlands (Group D) develop from both grasslands and Low Shrub Heathlands (Group C; Figure 4). The factors that determine which of the different tall shrub vegetation types may develop are unclear, although seed availability and land use his- tory probably play major roles. Huckleberry-Scrub Oak Heath- land (Group D1) often replaces Low Shrub Heathland, but where it succeeds grassland types, low ericaceous shrubs are generally already common. Mixed Maritime Shrublands (Group D2) often develop from grasslands and less often from Low Shrub Heath- lands. In some cases, tall shrub stages may be bypassed and succes- sion directly to forest can occur. Many of the pitch pine forests on Cape Cod are replacing Low Shrub Heathlands and Hairgrass Grasslands (Dunwiddie and Adams 1994), On Martha’s Vineyard, forests of black, white, and post oaks have developed in Little Bluestem Grasslands in many areas during the last 80—100 years, with no intervening shrub stage (Dunwiddie 1994). The loss of grassland and heathland to scrub oak, pitch pine, and other taller woody plant associations often is very rapid (Dunwiddie 1992). MacConnell et al. (1984) reported a 30 percent loss of heathland on Martha’s Vineyard and Nantucket between 1951 and 1971 as a result of succession of these species. Our model does not attempt to describe disturbances that arrest or reverse succession. Such events usually would correspond to a different set of arrows in the reverse direction in Figure 3. While 1996} Dunwiddie et al.—Heathlands and Grasslands 143 the model begins with plowed fields, succession following other disturbances, such as fire, livestock grazing, and forest clearance, is also likely to follow similar pathways. However, the starting point of these successional sequences would vary, depending on the initial species composition and the nature of the disturbance. Most of the maritime grasslands and heathlands of southeastern Massachusetts are dominated by native species, including a di- versity of rare taxa. In many places, these vegetation types may have been created by plowing or grazing. In other cases, agri- cultural practices modified the composition of existing natural grasslands or heathlands. Today, many organizations are focusing their efforts on preserving the remaining examples of these veg- etation types. The classification presented here provides a means of identifying more clearly how particular sites compare to the compositional range of grasslands and heathlands in the region. Furthermore, it can provide the basis for ranking sites to establish priorities for acquisition and management (Dunwiddie et al. 1993) ACKNOWLEDGMENTS. Partial funding for this project was pro- vided by a grant from the Massachusetts Natural Heritage and Endangered Species Program. P. Swain, T. Chase, R. Johnson, and T. Simmons helped in many ways to facilitate field work. We would also like to thank K. Lajtha for assisting with the soil analyses. T. Simmons and other participants at a Nature Conser- vancy-sponsored workshop provided useful ideas on the succes- sional mode LITERATURE CITED Boyce, S. G. 1954. es salt spray community. Ecol. Monogr. 24: 29-67. CoNnaARD, H. S. 1935. — aie of Central Long Island. Amer. Midl. Naturalist 16: oe Curtis, J. T. 1959. The eee of Wisconsin. University of Wisconsin Press, Madison, WI. Dunwippig, P. W. 1989. Forest and heath: The shaping ie the vegetation on Nantucket Seer Journal of Forest History 33: 126-133. 1990. Postglacial vegetation history of coastal ae in southeast- erm . New England. Natl. Geogr. Research 6: 178-195 1992. eae Landscapes: A pictorial field guide to a century of ofan nge on Nantucket. Nantucket Conservation Foundation, Nantucket Historical renin and Massachusetts Audubon Society, Nantucket, MA 144 Rhodora [Vol. 98 1994. Martha’s Vineyard Landscapes: The Nature of Change. The Vineyard Conservation Society and Peter W. Dunw / : s. 1994. Fire suppression and nde ape change on outer Cape Cod: 1600-1994. Unpublished See " the National Park — Cape Cod National ent Wellflee pC. CaALsjouw. 1990. Prescribed an i mowing of coastal ene and grasslands in een pp. 271-275. In: Mitchell, - Cc lak, an . J. Leopold, eds., Ecosystem Management: e Spacies and pistaneee Habitats, Pioceelings of the 15th Annual ne Areas Conference, New York State Museum Bull. 471 , K. A. HARPER AND R "E. ZAREMBA. 1993. Classification and ranking of ‘coastal heathlands and sandplain stages in Massachusetts. Un- published report to the Mass. Natural Heritage and Endangered Species — Nantucket, MA. GLEASON, H. A. AND A. Cronguist. 1991. Manual of Vascular Plants of eaueeea, United States and Adjacent Canada. 2nd ed. The New York Botanical pees Bronx, NY. GoprrRey, P. J. AND P. ALPERT. 1985. Racing to save the coastal heathlands. Nature oe, News 35: 10-13. — R. M. 1912. The Hempstead plains of Long Island. Bull. Torrey ot. Club 12: 277-287. ee es J. W. 1914. The vegetation of Nantucket. Bulletin of the pre Society of Philadelphia 7 0-79. HILL, M 979. TWINSPAN-—A FORTRAN program for ge os mul- tivariate ie in an ordered two-way ae by classification of the indi- ore and attributes. Ecology and Systematics, Cornell University, Ith- jena a. G H., C. J. EF TER BRAAK AND D. E R. VAN TONGEREN. 1987. Data oe in a and landscape ecology. Pudoc, Wageningen, Netherlan MACCONNELL, W, D. SWaRTOUT AND J. STONE. 1984. Land use update for aie Cod and the Islands with area statistics for 1951, 1971 and 1980. Res. Bull. 686, Mass. Agricultural Experiment Station, Univ. of Mass., pian MA. Macy, O. 1835. The History of Nantucket. Research Reprints (1970). New ork. MAINE NATURAL HERITAGE PROGRAM. 1991. Natural landscapes of Maine: A classification of ecosystems and natural communities. Department of Economic and Community Development, State House Station 130, Au- gusta, ME MASSACHUSETTS NATURAL HERITAGE PROGRAM. 1982. The ecological signif- icance of the Katama Plains. plang report. Massachusetts Division Fisheries and Wildlife. Boston Masseuse NATURAL ace: AND Seen SPECIES PROGRAM. 990). Watch list uncommon or rare Massachusetts plants. Massachusetts cae of Fisheries and Wildlife. Boston, MA. MASSACHUSETTS NATURAL HERITAGE AND ENDANGERED SPECIES PROGRAM. 1992. Massachusetts list of endangered, threatened, and special concern species. Massachusetts Division of Fisheries and Wildlife. Boston, MA 1996} Dunwiddie et al.—Heathlands and Grasslands 145 NATURE CONSERVANCY. 1995. aeeeneee map of Nantucket. Eastern Re- RESCHKE, C. 1990. ee communities of New York state. New York Natural Heritage Program, Latham, NY. Rosertson, D. 1973. The use of time lapse aerial ae analysis to evaluate land use and vegetational change in Nantucket and Dukes counties be- tween 1951 and 1971. M.S. thesis, Univ. of f Mass. ae SNEDDON, L. AND K. METZLER. 1992. Base: regional community classifi- cation, organizational hierarchy, and cross-reference to state heritage community classifications. Unpublished report. The Nature Conservancy, Eastern Regional Task Force, Boston, MA. Taytor, N. 1923. The vegetation of Montauk A study of grassland and forest. Brooklyn Bot. Gard. Mem |- TZEDAKIS, P. C. 1987. Holocene Goes ee in Cape Cod, Massa- chusetts. M.S. thesis, Brown University, Providence, WINKLER, M. G. 1985. A 12,000-year history of ee aes and climate for Cape Cod, Massachusetts. Quaternary Research 23: RHODORA, Vol. 98, No. 894, pp. 146-167, 1996 THE VASCULAR FLORA OF HOPEWELL CULTURE NATIONAL HISTORICAL PARK, ROSS COUNTY, OHIO JAMES P. BENNETT AND JENNIFER E. J. COURSE National Biological Service, University of Wisconsin, 610 Walnut St., Rm. 1007, Madison, WI 53705 ABSTRACT. Hopewell Culture National Historical Park, a unit of the Unit- ed States National Park Service located in Ross County in south central Ohio, was created to restore, protect, and eae t the legacy of the mound building Hopewell prehistoric peoples. The vascular flora of the park had been esti- mated to be only 20% known prior to the undertaking of this project. During the spring, summer, and fall of 1995, almost 700 plant specimens were col- lected by three investigators from five units of the park. Totals of 438 species, 281 genera, and 93 families of vascular plants were discovered, representing 40% of the flora of Ross County, and 17% of the flora of Ohio. Introduced species constituted 32% of the flora. Sixty-five species are new records for Ross County. Two species of spect concern, Spiranthes ovalis and Eleo- charts ovata, are on the state’s threatened and endangered species list. The Hopewell unit had the ee oc diversity of the five units Key Words: Hopewell, national park, flora, vascular plants Hopewell Culture National Historical Park is a unit of the Unit- ed States National Park Service located in Ross County in south central Ohio. It was created to restore, protect, and interpret the legacy of the mound building Hopewell, a group of prehistoric Native American peoples and cultures existing approximately 2,200 to 1,500 years ago. The park is nationally significant be- cause it contains some of the few remaining Hopewell earthworks in the world. However, it also contains areas of land with natural vegetation. Prior to this study, three partial plant checklists and some plant specimens in the park herbarium constituted the knowledge on plant species in the park. As of 1994, about 85 plant species had been recorded in the park and this was estimated to be about 20% of the total probably present (Allison Cusick, Ohio Department of Natural Resources, pers. comm.). This made the park’s flora the least known flora of any national park in the midwestern region (Bennett 1996). Recent floristic work in the region includes the flora of unglaciated southeastern Ohio, in- cluding Ross County (Cusick and Silberhorn 1977), and a flora of Fort Hill State Memorial in adjacent Highland County (Braun 146 1996] Bennett and Course—Hopewell Culture Flora 147 1969). The current study aimed to increase knowledge of the park’s flora with the following goals in mind: (1) to investigate the presence of federally and/or state listed threatened and en- dangered species, (2) to gain a better understanding of the park’s habitats and floristic diversity, (3) to document biodiversity in- formation to assist future park management, and (4) to contribute to the knowledge of Ohio’s flora and the floras of parks in the midwestern U.S. Hopewell Culture park and Ross County are situated on a ma- jor dividing line running NE/SW between two phytogeographic regions in Ohio: “... the Miami region, mainly a calcareous, glaciated till plain” to the northwest of the park, and the “ southern non-glaciated area of the Allegheny Plateau”’’ to the southeast (Braun 1989). This latter area contains an older and more relatively undisturbed flora than that of the glaciated area northwest of the park. Because of this phenomenon the flora of central Ross County is likely to be richer than the surrounding areas, assuming that the abiotic environment is fairly constant. However, human disturbances have had major influences on the park’s vegetation. e five units of Hopewell Culture park are High Banks, Hope- well, Hopeton, Mound City, and Seip (Figure 1). In 1995, the designated acreage was 1,134, of which 404 is federal land and the remainder non-federal. The separate units are all located in Ross County with the greatest distance between any two units being approximately 25 miles. Each of the five units is slightly different, and all are described in detail elsewhere (Bennett and Course: 1996). The area occupied by the park has a long history of modern human disturbance. The first park unit, Mound City Group, was originally a military base (the War Department’s Camp Sherman) before being transferred to the National Park Service in 1923 by presidential proclamation as a national monument. Barracks and training facilities were removed to restore the mounds. Most of the Hopewell Mound Group unit’s land has been farmed at one time and continues to be farmed today. As a historical park, Hopewell Culture’s priority has been to protect Hopewell archeo- logical resources. The park’s long term objectives include restor- ing park land to the historic landscape. However, lack of research and funds have impeded that work. In many areas, maintaining hayfields has been the least costly and the least damaging man- 148 Rhodora [Vol. 98 Hopewell Culture National Historical Park Ross County, Ohio Figure 1. Map of Hopewell Culture National Historical Park (1 cm = approximately 3.5 miles). agement of the earthworks. Consequently, a large percentage of the park is managed for this habitat. MATERIALS AND METHODS The taxonomic scope of this survey included all terrestrial and aquatic vascular plants. The field season ran from 15 March 1995 to 13 October 1995. The objective was to collect at least one specimen with reproductive parts of every naturally established plant species occurring within existing and/or proposed park boundaries. The following information was recorded with each specimen: locality, habitat, elevation, and collection date. Standards for writing floras suggested by Palmer et al. (1995) were adhered to as much as possible for this study. Relative abun- dance was determined for each species collected, using a three- level abundance scale modified from Palmer et al.’s five rank scale: 1996] Bennett and Course—Hopewell Culture Flora 149 COMMOM sicaxcece Dominant or co-dominant in one or more common habitats, or easily seen or found in one or more common habitats but not dom- inant in any common habitat. Intermediate ...... Widely scattered but not difficult to find. Fate wiciditnsssewedes Difficult to find with few individuals or col- onies but found in several locations, or very difficult to find and limited to one or very few locations or uncommon habitats. In addition, species presence at other units of the park was noted periodically in order to determine frequency throughout the park. Lastly, the origin (native or introduced) of each species was recorded as determined by Gleason and Cronquist (1991). Because it was uncertain how much land the park ultimately will purchase from the unacquired Hopewell, Seip, and High Banks units, some acreage was surveyed outside the currently proposed park unit boundaries. However, in some units, land was excluded from the survey because it was still under agriculture. The total acreage surveyed at each site was as follows: Hig Banks, 8; Hopewell, 247; Hopeton, 155; Mound City, 120; and Seip, 53. The grand total area surveyed was 583 acres. Acreages were estimated from park property maps. Out of the approximate seven month field season, there were 82 field collecting days, with the majority of them occurring June through September. Size and habitat complexity were primary factors in determining frequency and length of visits to the five different units. The total number of collecting visits to each park unit was the following: High Banks, 7; Hopewell, 34; Hopeton, 29; Mound City, 41; and Seip, I3. Specimens were identified using the following references: Braun (1967, 1989), Britton and Brown (1970), Catling (1983), Cobb (1963), Cooperrider (1995), Courtenay and Zimmerman (1978), Fisher (1988), Gleason and Cronquist (1991), Hitchcock (1935), Newcomb (1977), Petrides (1972), and Weishaupt (1971). The nomenclature followed for specimen and museum record la- bels is that of Gleason and Cronquist (1991). New species for Ross County were determined from the county dot maps in Braun (1967), Cooperrider (1995), Fisher (1988), unpublished maps of J. Furlow, and the county presence data in Cusick and Silberhorn (1977). 150 Rhodora [Vol. 98 RESULTS Six hundred and eighty-six specimens representing 438 species were collected for this study (Table 1). Collection numbers for each species are available in Bennett and Course (1996). The voucher specimens were deposited in the park’s museum collec- tion at park headquarters and duplicates were deposited at the Ohio State University Herbarium in Columbus (0s). Ten speci- mens were unidentified. A floristic summary for the park is shown in Table 2. Almost 80% of the species are dicots. The Hopewell unit had the greatest number of species (348). This was followed by the Mound City unit (268), Hopeton (238), Seip (219), and High Banks (175). These numbers roughly correspond with the acreages of the units. None of the species collected were new to the state of Ohio. Two species were collected that are on the 1994-95 Ohio Di- vision of Natural Areas and Preserves (1994) Rare Native Ohio Plant List: Eleocharis ovata (Roth) Roemer & Schultes is listed as a state endangered species. Two specimens (JCO147 and JCO385) were found at Hopewell unit along the old logging road in a recently logged area approximately 300 meters east of the northwestern entrance of the forest. They were found in sat- urated soil near other species of the Cyperaceae. It should be noted that we are using the broad definition of this species according to Gleason and Cronquist (1991), which includes Eleocharis obtusa (Willd.) Schultes. Allison Cusick (pers. comm.) considers the specimen to be the latter and quite common throughout the state. Eleocharis ovata sensu stricto is quite rare and is found only in the northern part of the state. Spiranthes ovalis Lindl. var. erostellata Catling is listed as a po- tentially threatened species in the state. It was listed formerly as a threatened species but recent findings indicate it may be more frequent in southern Ohio than previously thought (Cat- ling 1983). One specimen (JC0633) was found 20 feet east of the service road in the Mound City unit in an open area of a moist, early successional woods. Two species in the flora are represented by sight records: As- paragus officinalis and Monotropa uniflora. Toxicodendron rad- 1996] Bennett and Course—Hopewell Culture Flora 151 Table Vascular plant species of Hopewell Culture National Historical Park. i: list of taxa is sorted first by major group, then alphabetically by family, genus and species. For each species, relative abundance on a three point scale (C = common, I = intermediate, R = rare) and the units nee the park the species occurs in (HB = High Banks, HL = Hopewell, = Hopeton, MC = Mound City, and SP = Seip) are given. Species preceded by a ' are new records for Ross County, and those by a ? are not native Abun- Taxon dance Park Units Pteridophytes ADIANTACEAE Adiantum pedatum L. R HL ASPLENIACEAE Asplenium platyneuron (L.) Oak I HL, MC, SP Polystichum acrostichoides ae ) Schott R HL EQUISETACEAE Equisetum arvense L. R HL ONOCLEACEAE Onoclea sensibilis L. R HL OPHIOGLOSSACEAE Botrychium dissectum Spreng. R HL Botrychium virginianum (L.) Swartz I HL, MC Gymnosperms CUPRESSACEAE Juniperus virginiana L. R HN, MC Angiosperms ACANTHACEAE Justicia americana (L.) M. Vahl R HB, MC, SP Ruellia strepens L. I HN, HL, MC, SP ACERACEAE Acer negundo Cc HB, HN, HL, MC, SP Acer sac siesta Ju: Cc HB, HN, HL, MC, SP Acer saccharum Marshall Cc HB, HN, HL, MC, SP ALISMATACE Sagittaria eee Willd. R MC ANTHACEAE *Amaranthus retroflexus L. ] HB, HN, HL Amaranthus tuberculatus (Mog.) Sauer R MC ANACARDIACEAE Rhus glabra L. I MC, SP Rhus typhina L. I HN, HL, MC, SP Toxicodendron radicans (1L.) Kuntze Cc HB, HN, HL, MC, SP ANNONACEAE Asimina triloba (L.) Dunal I HB, HL, MC, SP 1352 Rhodora [Vol. 98 Table |. Continued. Abun- Taxon dance Park Units APIACEAE Chaerophyllum procumbens (L.) Crantz ] HL 2Conium maculatum C HB, HN, HL, MC, SP C sane ey eee (L.) DC. GC HB, HN, HL, MC, SP 2>Daucus Cc HB, HN, HL, MC, SP Erigenia a (Michx.) Nutt. I HL Osmorhiza clatyonii (Michx.) C.B. Clarke ] HL 'Osmorhiza longistylis (Torr.) DC. Cc HB, HN, HL, MC, SP *Pastinaca sativa C HB, HN, HL, MC, SP Sanicula gregaria E. Bickn. Cc HB, HN, HL, MC, SP ?Torilis arvensis es Link I HB, HN, HL, MC, SP APOCYNACEAE Apocynum cannabinum L. Cc HB, HN, HL, MC, SP '2Vinca mino I MC ARACEAE Arisaema triphyllum (L.) Schott R HL, MC ARALIACEAE Panax quinquefolius L. R HL ARISTOLOCHIACEAE Asarum canadense L. I HL, MC, SP SCLEPIADACI Ampelamus Fits (Nutt.) Britton I HB, HN, HL, MC, SP Asclepias incarnata L. R HL Asclepias syriaca L Cc HB, HN, HL, MC, SP Asc sae tuberosa L. R HN, SP ASTERACEAE rene millefolium L. I HB, HN, HL, MC, SP “Ambrosia artemistifolia L. Cc HB, HN, HL, MC, SP Ambrosia trifida Cc HB, HN, HL, MC, SP 2Arctium minus Schk. I HB, HN, HL, MC, SP *Artemisia annua L. R "Artemisia vulgaris L. R MC Aster cordifolius L. I HN, HL, MC, SP Aster lanceolatus Willd. I HL Aster lateriflorus (L.) Britton I HL Aster novae-angliae L. I HN, HL, MC, SP Aster pilosus Willd C HB, HN, HL, MC, SP Aster sagittifolius Willd. R Aster shortiit Lindley I HB, HN, HL, MC, SP Bidens bipinnata L. R HL, MC Bidens cernua L. R HL Bidens frondosa L. I HL, SP Bidens vulgata Greene I HL 1996] Bennett and Course—Hopewell Culture Flora 153 Table 1. Continued. bun- Taxon dance Park Units a alia ashe _= L. '?Carduus nutan cae aren ce anthemum L. 2Cichorium intybus L *Cirsium arvense (L.) Scop. °Cirsium vulgare (Savi) Tenore Conyza canadensis (L.) Cronq. Echinacea purpurea (L.) Moench Erechtites ne (L.) Raf. Erigeron annuus Erigeron philadelphicus L. Eupatorium altissimum L. Eupatorium coelestinum L. Eupatorium perfoliatum L. Eupatorium purpureum L., Eupatorium rugosum Houttuyn 'Eupatorium serotinum Michx Euthamia graminifolia (L.) Nutt. 'Gnaphalium obtusifolium L. Helianthus tuberosus L Heliopsis felianinoides (L.) Sweet '?Hieracium ¢ caespitosum Dumort. Kuhnia eupatorioides L. 'Lactuca canadensis L Lactuca floridana (L.) Gaertner *Lactuca serriola L '2Matricaria maritima L. 2Matricaria matric arioides (Less.) Polymnia canader Ratibida pinnata (Vent ) Barnhart n 2Senecio obovatus Muhl. Silphium perfoliatum L. Solidago canadensis L. Solidago flexicaulis L. Solidago gigantea Aiton Solidago juncea Aiton Solidago rugosa Miller 2Sonchus asper (L.) Hill Taraxacum officinale Weber ex Wiggers °Tragopogon pratensis L. Verbesina alternifolia (L.) Britton ATA TT TFT WAN TWAT TATMWDONAANNIN TATTOO TT AROATTAWAINNTMANNINNNAA , HN, HL, MC, SP HN, HL, MC. SP , HN, HL, MC, SP »~ AN, Hb, MC, SP , HN, HL, MC, SP , HN, HL, MC, SP , HN, HL, MC, SP N , MC, SP , HN, HL, MC, SP L , HL, MC » HN, Bl, MC..oP , HN, HL, MC, SP , oP , HL , HL ,HL, MCP eH HL MC SP , HN, HL, MC, SP L3F , HN, HL, MC, SP » HN, HL, MC, SP , HN, HL, MC, SP , HN, HL, MC, SP eHN HL MC. sP , HN, HL, MC, SP 154 Rhodora [Vol. 98 Table |. Continued. Abun- Taxon dance Park Units Vernonia gigantea (Walter) Trel. Cc HB, HN, HL, MC, SP Xanthium strumarium L, I HL, MC, SP SAMINA Tes patians 6 capensis Meerb. C HB, HN, HL, MC, SP Impatiens ane Nutt. C HB, HN, HL, MC, SP BERBERIDACE -Jeffersonia Tinie (L.) Pers. I HL Podophyllum peltatum L. I HL, MC BETULACEAE Carpinus caroliniana Walter R SP Ostrya virginiana (Miller) K. Koch I HL, MC, SP BIGNONIACEAE Campsis radicans (L.) Seemann ‘@ HB, HN, HL, MC, SP ?Catalpa speciosa Warder R HN BORAGINACEAE -Echium vulgare L. R HB Hackelta virginiana (L.) Il. M. Johnston ] HL, MC ?Lithospermum arvense L ] HN, MC Mertensia mea (L.) Pers. ] HL, MC BRASSICA -Alliaria nee (Bieb.) Cavara & Grande “Arabidopsis thaliana (L.) Heynh. Arabis perstellata L. Braun °Barbarea vulgaris R. Br '?Berteroa incana (L.) DC. *Brassica nigra L °Capsella bursa-pastoris (L.) Medikus 'Cardamine douglassii Britton aan parviflora L. raba verna L ‘Erysinum repandum L. “Hesperis matronalis L. lodanthus pinnatifidus — Steudel *Lepidium aaa oe virginicut orippa nasturtium- reac (L.) Hayek 2Sisvmbrium officinale (L.) Sc °Thlaspi arvense °Thlaspi perfoliatum L. HB, HN, HL, MC, SP HB, HN, HL, MC, SP HB, HN, HL, MC, SP i , HL, MC, SP L On @ mae @ ae ean © asian cae 2/8 een laa an es) an Z CAESALPINIACEAE Cercts canadensis L. , SP Gleditsia triacanthos L HB, HN, HL, MC, SP Gymnocladus dioica (L. ) K. Koch R HN, HL, MC ae) Tr ve) a 4 en = ns 0 1996] Bennett and Course—Hopewell Culture Flora 155 Table |. Continued. Abun- Taxon dance Park Units CAMPANULACEAE Ss shee americana L. I HN, HL, MC, SP Lobelia at I HB, HN, HL, MC Lobelia ae riliti I HN, HL, SP Triodanis senses (L. ) Nieuwl. I HB, HN, HL, MC, SP CANNABACEAE *Humulus lupulus L. I HL, MC CAPRIFOLIACEAE *Lonicera japonica Thunb Cc HB, HN, HL, MC, SP '?Lonicera maackii (Rupr.) Maxim. C HB, HN, HL, MC, SP '2Lonicera tatarica L C MC Sambucus canadensis L I HN, HL Symphoricarpos orbiculatus Moench I HL, MC CARYOPHYLLACEAE >Cerastium vulgatum L. I HN, MC *Dianthus armeria L. I HB, HN, HL, MC, SP *Saponaria officinalis L. Cc HB, HN, HL, MC, SP Silene antirrhina L. I HL '2Silene ee ee Cc HB, HN, HL, MC, SP *Silene noctiflor R SP Silene stellata i) ee f. R HL, MC Silene eh ica L. R HL *Silene vulgaris (Moench) Garcke C HB, HN, HL, MC, SP *Stellaria media (L.) Villa Cc HB, HN, HL, MC, SP CELASTRACEAE Celastrus scandens L Cc HN, HL, MC, SP '2Euonymus alatus (Thunb.) Siebold R MC Euonymus atropurpureus Jacq. I HB, HN, HL, MC, SP Chenopodium album L. C HB, HN, HL, MC, SP CLUSIACEAE Hypericum mutilum L. R HL -Hypericum perforatum L. I HB, HN, HL, MC Hypericum punctatum Lam. I HL ELINACEAE 2Commelina communis L. I HB, HN, HL, SP CONVOL een sepium (L.) R. Br. Cc HB, HN, HL, MC, SP 2Convolvulus arvensis L I HB, HL, M "Ipomoea hederacea Jacq. I HB, HN, HL, MC, SP Ipomoea lacunosa L. I HB, HN Ipomoea pandurata (L.) G. Meyer Cc HB, HN, HL, MC, SP iG Rhodora [Vol. 98 Table |. Continued. Abun- Taxon dance Park Units CORNACEAE Cornus drummondit C. A. Meyer R HB, HN, MC Cornus florida L. I HB, HN, HL, MC, SP CUCURBITACEAE ‘Cucurbita pepo L. R HN Sicyos angulatus L. R MC CYPERACEAE 'Carex albicans Willd. Carex albursina Sheldon 'Carex amphibola Steudel 'Carex blanda Dewey 'Carex careyana Torr. Carex davisti Schwein. & Torr. Carex festucacea Schk 'Carex flaccosperma lee 'Carex gracillima Schwel Carex hirtifolia Mackenzie Carex jamesti Schw 'Carex laxiculmis aide 'Carex retroflexa Muhl. Carex rosea Schk. 'Carex shortiana Dewey 'Carex squarrosa L Carex stipata Muhl. 'Carex VANIER a Cyperus esculentus Eleoc ied ovata ae Roemer & THT TARR RAZA APRARDARAR™— en co Schu R HL 3 atrovirens Willd. I HL Scirpus lineatus Michx. R HL DIPSACACEA *Dipsacus ee Hudson C HB, HN, HL, MC, SP ELAEAGNACEAE '-Flaeagnus angustifolia L. R HN, HL EUPHORBIACEAE Acalypha rhomboidea Rat. ] HN, HL, MC Acalypha virginica R HL Euphorbia comunutata Engelm. R HL Euphorbia dentata xX. R MC Euphorbia maculata L. Cc HB, HN, HL, MC, SP FABACEAE Amphicarpaea bracteata (L.) Fern. Desmodium canescens (L.) DC ~ 7 eo = wn , SP HN, HL, MC, SP 1996] Bennett and Course—Hopewell Culture Flora 157 Table 1. Continued. bun- Taxon dance Park Units 'Desmodium nudiflorum (L.) DC. R HL 2Medic ina L HB, HN, HL, MC, SP HB, HN, HL, MC, SP HB, HN, HL, MC, SP B, HN, HL, MC, SP B, HN, HL, MC, SP B, HN, HL, MC, SP HB, HN, HL, MC, SP N B °Me. Oo. aL Meiilons er Medikus 2Melilotus lesen — ) Pallas Robinia pseudoa Trifolium eee oS ai haa hybridum L. Trifolium incarnatum L. Trifolium pratense L. *Vicia cracca =aARQQNINQIANANN o , HN, HL, MC, SP N AGACEAE Fagus grandifolia Ehrh. R HL Quercus alba L. I HN, HL, MC Quercus bicolor Willd. I HL, MC Quercus imbricaria Michx. R HL, MC 2Quercus macrocarpa Michx. R HL, MC, SP Quercus muehlenbergii Engelm. I HL, SP ales rubra L J HB, HN, HL, MC, SP FUMAR aa vile (Raf.) DC. I MC Dicentra cucullaria (L.) Bernh. R HL, MC GERANIACEAE Geranium carolinianum L. Cc HB, HN, HL, MC, SP Geranium maculatum L. I HL, MC HAMAMELID E ee virginiana L. R MC 'Liquidambar styraciflua L. I HN, HL, MC, SP HIPPOCASTANACEAE Aesculus — Willd. I HB, HL, MC, SP ACEAE ios vue appendiculatum Michx. I HL 2Phacelia purshii Buckley I HN, HL IRIDACEAE Sisyrinchium angustifolium Miller I HL JUGLAND Carya cree (Wangenh.) K. Koch R HL Carya ovata (Miller) K. Koch I HN, HL Juglans le L. Cc HB, HN, HL, MC, SP JUNCACE sae tenuis Willd. R HN, HL Juncus torreyi Cov. R HL Luzula multiflora (Retz.) Lej. I HL 158 Rhodora [Vol. 98 Table 1. Continued. Abun- Taxon dance Park Units LAMIACEAE “A gastache nepetoides (L.) Kuntze R HL Blephilia hirsuta (Pursh) Benth. I HL °Glechoma hederacea L. Cc HB, HN, HL, MC, SP Hedeoma pulegioides (L.) Pers. R HL 'Lamium album L. R MC -Lamium amplexicaule L I HN, MC -Lamium purpureum L, Cc HB, HN, HL, MC, SP *Leonurus cardiaca L. I HB, HN, HL, MC, SP Lycopus americanus Muhl. R HL Monarda fistulc ] HN, MC *Nepeta cataria “L. R HB, SP Prunella vulgaris L. ] HB, HN, HL, MC, SP Teucrium canadense L. ~ HN, HL, MC o) a0 ive) LAURACEAE Sassafras albidum (Nutt.) Nees I HN, HL, MC, SP LENTIBULARIACEA ‘Utricularia ae L: wy en rm LILIACEAE ‘Allium cernuum Roth ‘Allium tricoccum Aiton Allium vineale L. -Asparagus officinalis L. HB, HN, HL, MC, SP P HB, HN, HL, MC, SP Cc R Cc ] HN, MC Camassia scilloides (Raf.) Cory R HL, SP 'Erythronium albidum Nutt. ] HL I, a alee allis Le ] HN, HL, SP Lilium canader nse L R SP I, ene umbellatum L. Cc HB, HN, HL, MC, SP Polygonatum biflorum ene Elliott R HN, HL, MC Smilacina racemosa (L.) I HL, MC, SP Trillium sessile L | HL Uvularia out J. E. Smith R MC MAGNOLIACEAE peat tulipifera L. | MC, SP MALVACEAE ites ss aia Medikus R HN, MC 1,2 cus trionum R SP Me oo a in L. Cc HB, HN, HL, MC, SP °Sida spinosa L. I HB, HN, HL, MC MENISPERMACEAE Menispermum canadense L. Cc HB, HN, HL, MC, SP MONOTROPACEAE Monotropa uniflora L. R HL 1996] Bennett and Course—Hopewell Culture Flora 159 Table 1. Continued. Abun- Taxon dance Park Units MORACEAE °-Maclura oe (Raf.) Schneid. R HN, MC °Morus Cc HB, HN, HL, MC, SP Morus aera . I HB, HN, HL, MC, SP NYCTAGINACEAE Mirabilis nyctaginea (Michx.) MacMillan I HN, MC OLEACEAE Fraxinus americana L Cc HB, HN, HL, MC, SP Fraxinus pennsylvanica Marshall Cc HB, HN, HL, MC, SP Fraxinus quadrangulata Michx. R HL '2Ligustrum vulgare R MC ONAGRACEAE Circaea lutetiana L. I HL aacie wine Lehm. R HL Gaura bienni: I HN, MC a a ee | I HN, HL, MC Oenothera —_ 1. R HN, HL ORCHIDACE 'Aplectrum s emale (Muhl.) Torr. R HL 'Spiranthes ovalis Lindl. R MC OXALIDACEAE Oxalis stricta L. C HB, HN, HL, MC, SP Oxalis violacea L. R HL PAPAVERACEAE *Papaver dubium L. R HN 2Sanguinaria canadensis L. I HL PHYTOLACCACEAE Phytolacca americana L. Cc HB, HN, HL, MC, SP PLANTAGINACEAE *Plantago lanceolata L. Cc HB, HN, HL, MC, SP '2Plantago major L Cc HB, HN, HL, MC, SP PLATANACEAE 2Platanus occidentalis L. I HB, HN, HL, MC, SP POACEAE ‘Agrostis hyemalis (Walter) BSP. R SP Andropogon gerardii Vitman dt SP 2Bromus commutatus Schrader I HB, HN, HL, MC, SP !2Bromus inermis Cc MC °Bromus tecto I HN, MC ee voto (Michx.) Yates R SP 'Cinna ee aL I HB, HN, HL, MC, SP *Dactylis glomerata L. Cc HB, HN, HL, MC, SP *Digitaria eens (L.) Scop. I HB, HN, HL, MC, SP 160 Rhodora [Vol. 98 Table 1. Continued. Abun- Taxon dance Park Units ‘Ec hinoc hloa c seas (L.) P Beauv. Cc HB, HN, HL, MC, SP Eleusine indica (L.) Gaertn. I HB, HN, HL, MC, SP Elymus pecones . Cc HB, HN, HL, MC, SP Elymus hystrix L. I HB, HN, HL, MC, SP Elymus villosus Muhl. I HN, HL '-Elytrigia repens (L.) Nevski Cc HB, HN, HL, MC, SP 'Fragrostis capillarts (L.) Nees I HL '?Festuca elatior L. Cc HB, HN, HL, MC, SP '?Festuca ovina L R SP 'Festuca subverticillata (Pers.) E. Alexeev I HN, HL. MC, SP Glyceria striata (Lam.) A. Hitche. R HL Leersia virginica Willd ] HL, MC “Muhlenbergia schreberi J. & Gmelin I HN, HL, MC, SP Panicum capillare L. I HB, HN, HL, MC, SP Panicum dich Se ane Michx. | HB, HL Panicum lanuginosum Elliott I HB, HN, HL, MC, SP ‘Panicum latifolium L. I HL, SP Panicum virgatum L. I HN, HL, SP °Phleum pratense L. Cc HB, HN, HL, MC, SP °Poa pratensis L. I HB, HN, HL, MC, SP Poa sylvestris A. Gra I MC '2Seraria faberi R. Herrm. I HN, HL, MC, SP *Setaria glauca (L.) P. Beauv. Cc HB, HN, HL, MC, SP '2Seraria italica (L.) PR Beauv. ] HL *Setaria viridis (L.) P. Beauv. Cc HB, HN, HL, MC, SP Sorghastrum nutans (L.) Nash I HL..SP °Sorghum halepense (L.) Pers. C HB, HN, HL, MC, SP Tridens flavus (L.) A. Hitche. Cc HB, HN, HL, MC, SP POLEMONIACEAE Phlox divaricata L I HL Phlox paniculata L. I MC, SP Polemonium reptans L. I HL POLYGONACEAE °Polygonum aviculare Polygonum cespitosum Blinc Polygonum pensylvanicum L. °Polygonum persicaria L Polygonum punctatum Elliott *Polygonum scandens L Polygonum virginianum L, -Rumex acetosella L. “Rumex crispus L. *Rumex pene [es Rumex verticillatus L. HB, HN, HL, MC, SP HB, HN, HL, MC, SP HB, HN, HL, MC, SP L A et ee I ey Se ma ‘o) 1996] Bennett and Course—Hopewell Culture Flora 161 Table 1. Continued. Abun- Taxon dance Park Units PORTULACACEAE Claytonia virginica L. Cc HB, HN, HL, MC, SP POTAMOGETONACEAE Potamogeton nodosus Por. R HL PRIMULACEAE Lysimachia ciltata L. R MC *Lysimachia nummutlaria L. Cc HB, HN, HL, MC, SP Samolus floribundus HBK R HL RANUNCULACEAE Actaea alba (L.) Miller Anemone virginiana L. Anemonella thalictr cag (L.) Spach Aquilegia canadens Clematis viorna L. Clematis virginiana L. Delphinium tricorne Michx. Hydrastis canadensis L. Ranunculus abortivus L. Ranunculus hispidus Michx. HB, HN, HL, MC, SP AA OAT TART TDW en = 2Ranunculus micranthus Nutt. HL Thalictrum revolutum DC. SP ROSACEAE Agrimonia gryposepala Wallr. ] HL, SP Agrimonia parviflora Aiton HL d land, Stable. 2/786 (srh, CLEMS * Stac es byzantina C.Koch, Lamb’s-ears. Scie Disturbed land, Waif. S side US I pV. post opposite Long Wharf eee 22 Jul 1990, 21768 (srh, " First report for southeastern Connecticu a pe eno L. var. canadense, Seaside Sane Occasional, Disturbed land, Tidal marshes (above highest tide mark), Stable. 934 (vt); 9446 (srh, MARY, NY, VT); 2/6/35 (srh, CLEMS, GH, NY, NYS). Pycnanthemum muticum Mic hs .) Pers., Mountain-mint. Infrequent, Oak forest, Distu 1996] Hill—Latimer Point Flora 205 Trichostema dichotomum L., Blue-curls. Infrequent, Disturbed land, Stable. 9344 (sth, MARY, NY, TAES, VT). LAURACEAE Sassafras albidum (Nutt.) Nees, Sassafras. Occasional, Oak forest margins, Stable. Il 3 (CLEMS, GH, NY, NYS); 23098 (srh, CLEMS, GH, ILLS, MO, NY, TAES, USF). LINACEAE Linum striatum Walt., Yellow wild flax. Infrequent, Disturbed land, Stable. 2/740 CLEMS, NYS as L. virginianum L.). LYTHRACEAE *Lythrum salicaria L., Purple loosestrife. Occasional to Infrequent, Disturbed land, Freshwater marshes, Stable. 939/ (srh, MARY, MO, NY, TAES a MALVACEAE *Abutilon theophrasti Medikus, Velvet-leaf. Scarce, Disturbed land, Waif. SW quadrant of Latimer Point in hedgerow, 8 Sep 1979, 8733 (srh, MARY, NY, TAES, VT). Hibiscus palustris L., Wild hibiscus. Infrequent, Freshwater marshes, Declining. showiest flower of Latimer Point; very attractive when in full flower in mid-August. The single pond where it grows is vulnerable to herbicide runoff from the RR right-of-way and to any road or RR construction nearby. One location, pond, NW quadrant of Latimer Point, 21 Aug 1972, 924 (Ny, VT): same locality, 13 Aug 1980, 9366 (srh, GH, MARY, MO, NY, TAES, VT). Gleason Cronquist (1991) and Tucker (1995) consider this a synonym of Hibiscus refer to recognize these northern all-pink forms as spe- moscheutos L., but are never mixed with the white flowered H. ally distinct. In our area, they eee which ts more souther *Malva neglecta Wallr., Common ide Scarce, (srh). Disturbed land, Stable. 2/623 MELASTOMATACEAE Rhexia virginica L., Meadow-beauty. Infrequent, Disturbed land, Freshwater marshes, Stable. 2/750 (CLEMS); 2/796 (sth, CLEMS). MOLLUGINACEAE *Mollugo verticillata L., Carpetweed. re Disturbed land, Stable. 9398 (srh, , TAES, VT); 2/605 (CLEMS, GH, MONOTROPACEAE [within Ericaceae in Tucker 1995] sea uniflora L., Indian pipe. Scarce, Oak forest, Stable. 7/855 (CLEMS); 9 (srh, CLEMS). 206 Rhodora [Vol. 98 MYRICACEAE Comptonia peregrina (L.) Coult., Sweet-fern. Infrequent, Disturbed land, Oak forest margins, Stable. 22486 (srh, CLEMS, CM, GH, KANU, MO, NCU, NY, RSA TAES, USCH, USF, UTC, WVA,); 23097 (srh, CLEMS, GH, MO, NY, SMU, TAES, TEX). Myrica pensylvanica Loisel., Bayberry. Occasional, eet ke land, Coastal beaches and dunes (above highest tide mark), Stable. 9//7 (srh, MARY, NY, TAES, VT); Y38O (srh, MARY, NY, TAES, VT) NYMPHAEACEAE pres odorata Dryand. ex Aiton, Water-lily., Scarce, Freshwater marshes, ining. In the pond with Hibiscus, NW quadrant of Latimer Point, 4 Jul 1980, 9327 et MARY, VT). NYSSACEAE Nyssa sylvatica Marsh. var. sylvatica, Black gum. Occasional, Oak forest margins Stable. /0/00 (srh, CLEMS, GH, MARY, MO, NY, TAES, VT) OLEACEAE *Ligustrum vulgare L., European privet. Infrequent, Disturbed land, Increasing 9308 (srh); 23/705 (srh, CLEMS ONAGRACEAE Ludwigia nee ae L., Seedbox. Infrequent, Freshwater marshes, Disturbed land, Stable. 2/738 (srh, CLEMS Ludwigia ilnees (L.) Ell., Water slates Infrequent, Freshwater marshes, Sta- ble. 9374 (srh, MARY, NY, TAES, VT). Oenothera biennis L., Evening-primrose. Frequent, Disturbed land, Coastal beac es and dunes, Stable. 873/ (srh, MARY); 9388 (MARY, NY, VT as O. parviflora L.) OXALIDACEAE Oxalis stricta L., Sour-grass. Occasional, Disturbed land, Stable. 2/622 (srh). PAPAVERACEAE *Chelidonium majus L., Celandine. Scarce, Disturbed land, Increasing. Seen but ot vouchered in residential area in SE quadrant of Latimer Point, starting in 1982 (pers. obs.). It is frequent in nearby Mysti *Eschscholzia californica Cham., California poppy. Scarce, Disturbed land, Waif. S side US Rt. | near VFW post opposite Long Wharf Road on landfill, 7 Aug 1991, 225/72 (srh, CLEMS, NY). First report for southeastern Connecticut. PHYTOLACCACEAE Phytolacca americana L., Pokeweed. Occasional, Disturbed land, Stable. 937/ (srh, MARY, NY, TA VT) 1996] Hill—tLatimer Point Flora 207 PLANTAGINACEAE *Plantago aristata Michx., Buckhorn plantain. Infrequent, Disturbed land, Stable. 1688 (srh, CLEMS, GH, NYS). *Plantago lanceolata L., English plantain. Occasional, Disturbed land, Stable. 11806 — major L., Common plantain. Occasional, Disturbed land, Stable. 2/599 (srh, CLEMS, GH, NY, NYS, TAES, USF). Bee. maritima L., subsp. juncoides (Lam.) Hultén, Seaside plantain. Infre- quent, Tidal marshes, “peclinin ng. I have seen little of this in recent years. NE quadrant of Latimer Point, 28 Aug 1983, /32/4 (srh, GH, MARY, MO, NY, TAES, PLUMBAGINACEAE Limonium carolinianum (Walt.) Britton, Sea lavender. Occasional, Tidal marshes, Stable. 943/ (srh, CLEMS, MARY, MO, NY, TAES, VT); /32/7 (srh, GH, MARY, MO, NY, TAES, VT). POLYGALACEAE Polygala sanguinea L., Common milkwort. Scarce, Disturbed land, Freshwater marshes, Stable. 2/734 (srh, CLEMS, GH, NYS). POLYGONACEAE * Polygonum aviculare L., Knotweed. a Disturbed land, Coastal beaches and dunes, Stable. 8754 (srh, MA VT Beles Sal convolvulus L., Black Ser Occasional, Disturbed land, Stable. 1767 (CLEMS, NY, NYS); 2248? (CLEMS, GH, NY, TAES, USF). ee cuspidatum Sieb. & Zucc., Japanese bamboo. Infrequent, Disturbed nd, Stable. 22507 (srh, CLEMS, GH, NY, USF). *Polygonum hydropiper L., Water-pepper. Occasional, Disturbed land, Stable. 21686 (srh, CLEMS, NYS); ies (srh, CLEMS, GH, NY, TAES, USF). Polygonum jvdvopioenerdes Michx., Mild water pepe Occasional, Freshwater marshes, Stable. 9358 (MARY, NY, TAES, VT); 9369 (MARY, NY, TAES, VT Polygonum pensylvanicum L., Smartweed. Occasional, Disturbed land, Coastal beaches and dunes, Stable. 21633 (srh, CLEMS). * Polygonum persicaria L. [incl. P. dubium Soa. Lady’s-thumb. Occasional, Dis- turbed land, Stable. 22505 (srh, CLEMS, GH, NY, USF); 22509 (srh, CLEMS). Polygonum punctatum Ell., Smartweed. ‘Oecasi onal, Disturbed land, Freshwater marshes, Stable. 8732 (srh, MARY, VT). Polygonum ramosissimum Michx., Knotweed. Occasional, Tidal marshes, Stable. 9434 (srh, MARY, MO, NY, TAES, VT). Polygonum sagittatum L., Tear thumb. Infrequent, Freshwater marshes, Stable. 21754 (srh, CLEMS, NYS). Polygonum scandens L., Climbing wild buckwheat. Occasional, Disturbed land, Stable. 935 (vT) 8737 (sth, MARY, NY, TAES, VT). *Rumex acetosella L., Sour-grass. ee Penal land, Stable. 2/6// (srh, CLEMS, GH, NY, NYS). 208 Rhodora [Vol. 98 *Rumex pike L., Curly dock. Occasional, Disturbed land, Stable. 2/604 (srh CLEMS, GH, NY, NYS). PORTULACACEAE * Portulaca oleracea L., Purslane. Occasional, Disturbed land, Stable. 2/639 (srh, MS). PRIMULACEAE *Lysimachia punctata L., Garden loosestrife. Infrequent to Occasional, Disturbed land, Increasing. Only in cultivation through the early 1980s escaped and become successfully established on the Point in the late 1980s and early 1990s. Latimer Point garden, 28 Jun 1982, //S802 (srh, MARY, NY SW quadrant of La , the species has pe a 7 VE); timer Point near main beach, 17 Jul 1990, 2/644 (srh, CLEMS , LG, NCU, NY, TAES, USF). eee quadrifolia L., ares loosestrife. Occasional, Oak forest, Stable 21715 (srh, CLEMS, GH, NCU, NY PYROLACEAE [within Ericaceae in Tucker 1995] Chimaphila maculata (L.) Pursh, Spotted wintergreen. Scarce, Oak forest, Stable 21782 (srh). Pyrola sa ‘a Nutt., Common shinleaf. Scarce, Oak forest, Shrub swamps, Sta- ble. 22485 (srh, ¢ RANUNCULACEAE Anemone Linge ete L., Windflower. Occasional, Oak forest, Stable. /OO82 (srh MARY, . VT); 23088 (srh, CLEMS, GH, MO, NY, : 72) es] ). Clematis virginiana a Clematis. Scarce, Oak forest margin, Stable. 2/784 (srh, GH, MO). ss onsolida eee (L.) PW. Ball & Heywood ee ajacts L.|, Delphin- , Disturbed land, Waif. /7062 (srh, hae a ioe L., Bulbous buttercup. eae Disturbed land, Increas- ing. 2/743 (srh, CLEMS); 23/04 (srh, CLEMS). Thalictrum pubescens Pursh [T. a eas Muhl.], Tall meadow rue. Infrequent margins of Oak forest and Shrub swamps, Stable. 9309 (MO, NY, vT); 9353 (srh); 2/656 (CLEMS, GH, NY, NYS, TAES se RHAMNACEAE *Rhamnus frangula L., Alder buckthorn. Infrequent, Disturbed land, Increasing. arrival on the Point, first noted with this collection. NE quadrant of Latimer Point, 5 Aug 1991, 22473 (srh, CLEMS, NY, ROSACEAE Amelanchter canadensis (L.) aga) Shadbush. v1); 23700 (srh, CL , GH, ILLS, MO, NY, SMU, TAES, TEX, USF). Aronia arbutifolia (L.) Ell., Red hee eitvequent Freshwater marshes, Dis- turbed land, Stable. 9386 (srh, MARY, NY, Frequent, Oak forest, Stable. TAES, VT) 1996} Hill—Latimer Point Flora 209 Aronia prunifolia (Marsh.) Rehd., Chokeberry. Occasional, Oak forest margin, Stable. 9/05 (srh, MARY, VT). Crataegus pruinosa (Wendl. f.) K. Koch, Hawthorn. saperanies Shrub swamps, < forest margin, Stable. /0094 (srh, GH, MARY, MO, NY, TAES, VT). Fragaria virginiana Duchesne, Wild strawberry. Occasional, ae land, Sta- ble. 2/657 (srh, CLEMS, GH, MOR); 23/07 (srh, CLEMS). Geum canadense Jacq., White avens. Occasional, Disturbed land, Stable. 2/6/8 (CLEMS, MOR). *Malus a Sieb., Flowering crabapple. Infrequent, Disturbed land, Stable. 23096 (srh, ILLS, TAES, USF). *Malus kai Mill. [Pyrus malus auth.], Wild a Infrequent, Oak forest mar- gin, Disturbed land, Stable. 9/04 (srh, MARY, Pre argentea L., Silvery cinquefoil. ae Disturbed land, Stable. 10069 (srh). Potentilla canadensis L., Common cinquefoil. Occasional, Disturbed land, Stable. 23122 (CLEMS, GH, MO, NY, USF). Potentilla norvegica L., Rough cinquefoil. Infrequent, Disturbed land, Stable. h): 2/647 (srh). According to Gleason and Cronquist (1991) our plants may or may not be native. *Potentilla recta L., Upright cinquefoil. Scarce, Disturbed land, Stable. 2/729 (srh, CLEMS) Potentilla simplex Michx., Cinquefoil. oe Oak forest margin, Disturbed and, Stable. /0099 pe MARY, NY, TAES, VT). Prunus serotina Ehrh., d black cnet Occasional to Frequent, Oak forest margins, Disturbed a 9408 (srh, MARY, , NY, TAES, VT). *Pyrus communis L., Pear. Scarce, Disturbed land, eee 10068 (srh, GH, MARY, . NY, TAES, VT) *Rosa multiflora Thunb., Multiflora rose. Occasional to il frequent, Disturbed land, Oak forest margin, Stable. //SO4 (srh, MARY, NY, *Rosa rugosa Thunb., Beach rose. Occasional, Coastal nies and dunes, Stable. 314 (vr); 8755 (srh, MARY). The white flowered form also occurs here but is scarce: 3/5 (vt); 2/779 (srh, CLEMS, MOR). Rosa virginiana Miller, Wild rose. Occasional to Frequent, margin of Oak forest, Tidal marshes, Stable. 293 (vT); 93/3 (srh MARY, MO, NY, TAES, VT); /OSO4 (srh, MARY, N AES, VT); J/7SOS (srh, MARY, NY, TAES, VT). Rubus dileoientensis Porter ex Bailey, Northern blackberry. Occasional, Disturbed land, Stable. 2/666 (srh, CLEMS, GH, NY, MOR). Rubus flagellaris Willd., Northern ae ena Disturbed land, Stable. (srh, GH, MARY, NY, TAES, VT) Spiraea alba Duroi var. latifolia (Ait.) mies Meadowsweet. Occasional, Fresh- water marsh, Stable. 9376 (srh, MARY, NY, VT). cas rae tomentosa L., Steeplebush. Occasional, Freshwater marsh, Stable. 9370 , MARY, MO, NY, TAES, VT). RUBIACEAE *Galium aparine L., Bedstraw. Occasional, Disturbed land, Stable. 23//9 (srh, CLEMS, NY, USF). 210 Rhodora [Vol. 98 *Galium Hida L., White bedstraw. Occasional, Disturbed land, Stable. 2/752 (srh, Galium eee Bivel., Marsh bedstraw. Occasional, Freshwater marshes, Stable. vt as G. tinctorium); 9365 (srh, MARY, NY, VT SALICACEAE *Populus alba L., White poplar. Infrequent, Disturbed land, Stable. /0072 (srh, GH, MARY, MO, NY, TAES, VT). Populus ee ee Michx., Quaking aspen. agian Disturbed land, Increas- ing. 2/665 (srh, CLEMS, GH, NY, MOR); 23093 (srh, CL *Salix alba L., White willow. Occasional, Dictucbad ae Stable. 2/758 (srh, CLEMS). *Salix cinerea L., Ashy European willow. Infrequent, Shrub swamps, Stable. 9//9 (srh, MARY, MO, NY, VT). Salix eriocephala Mich, Wand willow. Infrequent, Shrub swamps, Stable. /0076 (srh, MARY, NY, TAES, VT); 23082 (srh, ILLS, NY, SMU, TAES, USF). SCROPHULARIACEAE Agalinis maritima (Raf.) Raf., Salt marsh gerardia. Occasional, Tidal marshes, Stable. 929 (NY, VT); 9433 (srh, MARY, MO, NY, TAES, VT); /32/8 (GH, MARY, MO, NY, TAES, ) Linaria canadensis (L.) Dum.-Cours., Toadflax. Occasional, Disturbed land, Sta- le. JI55 (v1); 9331 (srh, MARY, TAES, VT). *Linaria vulgaris Mill., Butter-and-eggs. Occasional, Disturbed land, Stable. 9394 (srh). Lindernia phnoaaren (Michx.) Pennell, False pimpernel. Infrequent, Freshwater marshes, Stable. 9375 (srh, MARY, NY, VT). + a tomentosa an ) Sieb. & Zucc. ex Steud., Princess tree. oe Disturbed land, Increasing. Not seen in the area until first sighting in 1991, i is likely to become much more common in the future since this individual was already fertile though only 3 m tall. Dodges Island, 8 Aug 1991, 225/6 (srh, CLEMS, GH, NY) [included in Bignoniaceae by Tucker 1995]. Scrophularia lanceolata Pursh, sae ort. Occasional, Disturbed land, Stable. 7445 (srh, TAES, VT); 7/834 (srh, GH, RY, MO, NY, TAES, VT, WS *Verbascum thapsus L., Caner. eae Infrequent to Occasional Disturbed land, Stable. 2/602 (CLEMS, ISC, NY). *Veronica arvensis L., Corn oe Occasional, Disturbed land, Stable. 23/25 (CLEMS, GH, MO, NY, TAES, USF) es: peregrina L. subeD: peregrina, Neckweed. Occasional, Disturbed land, Stable. /0246 (srh, GH, MARY, MO, NY, TAES, VT); 2/606 (CLEM SIMAROUBACEAE *Ailanthus altissima (Mill.) Swingle, Tree-of-Heaven. Infrequent to locally Fre- quent, Disturbed land, Increasing. A few trees were found already mature on Dodges Island in 1980; by 1982 this had become an almost pure grove taking the place of Prunus, Rhus and Myrica at this site. In 1991 one individual was first seen on Latimer Point opposite Dodges Island near the main beach. It is likely to become much more common in the future. Dodges Island, 16 Aug 1996] Hill—Latimer Point Flora 211 1980, 9448 (srh, MARY, MO, NY, TAES, VT); same locality, 12 Jul 1982, //83/ (CLEMS, GH, MARY, NY, TAES, VT); same locality, 7/832 (srh, GH, MARY, NY, TAES, SOLANACEAE *Datura stramonium L., Jimson-weed. Occasional, Coastal beaches and dunes, Disturbed 7 d, Stable. Most common around dead Zostera piles at the upper beach s . 791 (vT); 8738 (srh, MARY). *Physalis ate L., Chinese lanterns. Scarce, Disturbed land, Stable. 2/725 MS). *Solanum Sane L., Climbing nightshade. Occasional, Disturbed land, Stable. 21616 (srh, CLEMS, CTES, GH, NY, NYS, TAES, USF). Solanum ptychanthum Dun. ex . [S. americanum auth., often incl. within S. nigrum L.], Black pros boat Coastal beaches and dunes, Dis- turbed land, Stable. 945 (GH, NY, VT); 2/637 (srh). ULMACEAE Ulmus americana L., American elm. Occasional to Infrequent, Shrub swamps, Declining. Upper leaf surfaces rough in this population. As in most of New England, ares mature trees of the species have declined due to Dutch Elm disease, but younger trees still seem to survive and reproduce. 2/722 (CLEMS, GH, LG, NY, NYS, USF). URTICACEAE Urtica dioica L. subsp. gracilis (Ait.) Solander, Stinging nettle. Infrequent, Dis- turbed land, Increasing. /0798 (srh, MARY). VERBENACEAE Verbena hastata L., Blue vervain. Occasional, Disturbed land, Stable. 9449 (srh, MARY, NY, TAES, VT); 2/6/4 (CLEMS, GH, NY, NYS). VIOLACEAE Viola fimbriatula Sm., Northern downy violet. Infrequent, Disturbed land, Stable. 21684 (srh, CLEMS). Viola lanceolata L., Lance-leaf violet. Infrequent, Disturbed land, Stable. //65 (VT); S (MARY); 2/683 (srh). *Viola Aa Pollard [often incl. in V. sororia Willd.], Confederate violet. Infrequent, Distur a land, Stable. SE quadrant of Latimer Point, residential area ae weed, 13 May 1992, 23103 (srh). First report for southeastern Con- necticut, but eae included previously within V. soro Viola sagittata Ait. [may = V. fimbriatula Sm.|, Arrow- ce ace 23081 (srh, CLEMS, NY). *Viola sororia Willd., Common blue violet. Occasional, Disturbed land, Stable. 3121 (CLEMS, GH, NY, USF) *Viola tricolor L., oe y-jump-up. Scarce, Disturbed land, Waif. Escape from a Ae not persist for long. SE quadrant Latimer Point: 5 Jul 1980, 9333 (srh, MARY). Ze Rhodora [Vol. 98 VITACEAE *Ampelopsis cordata Michx. Scarce, Disturbed land, Increasing. First seen in 1980 along oo Road just outside of this study area, it has now spread closer the Point. S side US Rt. | near VFW post opposite Long Wharf Road in landfill, a Aug 1991, 22506 ee CLEMS, GH, NY). Parthenocissus genie (L.) Planchon, Virginia creeper. Frequent, Oak forest, Disturbed lan table. Similar to and confused with Parthenocissus vitifolia (Knerr) A. te ie which has been collected in nearby Mystic and may also occur on the Point. 2/66/ (srh, CLEMS, GH, LG, S). Vitis labrusca L., Fox grape. erie Oak forest margins, Stable. 7/807 (srh, GH, MARY, MO, NY, OKL, OSC, RENO, TAES, UTC, VT, WIS, WS) DISCUSSION Seymour (1969) reports 2882 vascular plant species present in New England, 2005 (70%) of which are native and 877 (30%) of which are non-native. Dowhan (1979) reports 2489 vascular plant species in Connecticut, 1636 (66%) of which are native and 853 (34%) of which are non-native. Tucker (1995) reports 1550 vascular plant species in southeastern Connecticut of which 1170 (75%) are native and 380 (25%) non-native. The flora of Latimer Point and vicinity includes 385 vascular plant species, 248 (64%) of which are native taxa and 137 (36%) of which are non-native. The proportion of native and non-native species in these floras remains rather constant despite major differences in coverage area. This list represents primarily the coastal element of the flora reported by Tucker (nearly 25% of the total flora of southeastern Connecticut or nearly 16% of the entire state flora). The flora of Latimer Point and vicinity was searched out and studied during the period 1969 to 1992. Four-hundred twenty- seven taxa of plants are listed in this paper, of which 30 are marine macroalgae, 12 are mosses, 10 are ferns and fern allies, 1 is a conifer, and 374 are flowering plants (107 monocots and 267 dicots). There are 92 vascular plant families represented, the five largest being Poaceae (53 taxa), Asteraceae (39), Cyperaceae (25), Rosaceae (22), and Fabaceae (20). The two largest genera represented are Carex and Polygonum, each with 11 species. Among the elements of the flora, 381 taxa have had stable pop- ulations during the period, 30 have seen their numbers increase (23 exotics and 7 natives), and 16 have seen their numbers decline (3 exotics and 13 natives). One species (Honckenya peploides) was found which has been listed as a Connecticut Species of 1996] Hill—Latimer Point Flora 213 Special Concern. One species listed as endangered in Connecticut (Ligusticum scothicum) was also found, and another (Angelica lucida) has been reported from the study site. Seven plant habitats are described in the vicinity of Latimer Point, and the numbers of all taxa most commonly found in each include 32 in the estuaries, 22 on the coastal beaches and dunes, 30 in the tidal marshes, 44 in the freshwater marshes, 25 in the shrub swamps, 63 in the oak forest, and 211 on disturbed lands. The large percentage (49%) of taxa found primarily in disturbed areas reflects the history of human and natural coastal disturbance characteristic of coastal New England today. Exotic species have increased in disturbed areas, but not in the more mature plant associations. Of the 23 exotics that have be- come more common over the past 24 years, 9 have presumably been dispersed widely by birds which eat their fleshy fruits (Am- pelopsis cordata, Berberis thunbergii, B. thunbergii var. atropur- purea, Elaeagnus umbellata, Euonymus alatus, Ligustrum vul- gare, Lonicera japonica, Lonicera morrowii, Rhamnus frangula). Seven of the increasing exotics are dispersed primarily by wind (Acer platanoides, Acer pseudoplatanus, Hypochoeris maculata, Lactuca serriola, Sonchus arvensis, Paulownia tomentosa, Ailan- thus altissima). One is a recent garden escape (Lysimachia punc- tata) and the remainder are opportunistic species that have in- creased by chance with human disturbance of the landscape (Ar- temisia vulgaris, Centaurea maculosa, Chelidonium majus, [pom- oea purpurea, Ranunculus bulbosus, Spergularia marina). Seven species (Abutilon theophrasti, Consolida ambigua, Eschscholzia californica, Panicum millaceum, Silene armeria, Stachys byzan- tina, Viola tricolor) were seen once and not again and are re- ported here as waifs; of these the first and fourth are widespread weeds which may have been introduced in bird seed or fill dirt, and the remaining five are all garden plants that were probably casually discarded and were only briefly established. The flora and fauna have been impacted by human activities resulting in a high percentage of non-native species, but the remaining native terrestrial flora has not seriously declined. Seven native species increased their numbers during the ob- servation period. The green alga Cladophora albida has possibly increased due to increased nutrient content (pollution) of the coastal waters. Cuscuta pentagona, dodder, is an opportunist with no obvious reason for its increase. Dennstaedtia punctilobula is 214 Rhodora [Vol. 98 a fern which has increased due to brush clearing near the resi- dential areas of the point because it prefers this open habitat to a closed canopy. Lycopus americanus and Phragmites australis have expanded along US Rt. | because of increased trenching of roadsides and disturbance of wetland communities in this area. Populus tremuloides and Urtica dioica may be increasing their ranges due to increased disturbance along roadsides. Sixteen species declined in numbers of individuals during the period of observation. Three of these were exotic species which apparently were not very competitive with the native flora or their fellow exotics (Anthemis cotula, Artemisia stelleriana, Berberis vulgaris). Among the 13 native species in decline, 3 are marine algae that may be decreasing due to pollution, though other fac- tors may be involved (Chondrus crispus, Dasya baillouviana, Leathesia difformis). The filling of wetlands, as well as increased herbicide use along the railroad which passes through the wet- lands, may be responsible for the decline in four species (Carex intumescens, Cyperus diandrus, Hibiscus palustris, Nymphaea odorata). It is tempting to speculate that warming of the envi- ronment is responsible in part for the decline in three generally more northern species (Honckenya peploides, Osmunda clayton- iana, Plantago maritima). Predation by gypsy moths was the ap- parent cause for a decline in Quercus bicolor, and Dutch elm disease has continued to take its toll on Ulmus americana. Betula populifolia seems to be in decline due to the increased use of herbicides along the woodland margins bordering the railroad, a habitat which it prefers. I was able to observe several other vegetation changes over time. During the 24 year observation period the trees of the oak forest became taller and more massive despite several hurricanes and storms. As shade increased, the Smilax and Vitis colonies have declined slightly in the former openings in the forest. The Zostera populations in the bay have fluctuated greatly; there was a period in the 1970s and 1980s when there were none to be found, while in the late 1980s and early 1990s it was difficult to get a boat through the massive colonies in the estuaries. The flora is in rather good condition. Since the beginning of my observa- tions, no native taxa have been lost. However, there is an influx of exotic species still taking place, and the more fragile islands and any disturbed lands soon may have forests dominated by Adanthus altissima and Paulownia tomentosa, if they do not al- 1996] Hill—Latimer Point Flora 215 ready. Furthermore, the native white-tailed deer population has increased noticeably with the lack of hunting and the increased foraging may lead to a decline of the more edible species. The deer sometimes are seen swimming from island to island. The coastal flora is a resilient one; it has developed in the presence of storms, tidal surges, erosion and other disturbance, so humankind probably cannot destroy it completely except by paving it over. However, the mature oak forests of the area have been gone for well over 150 years and are unlikely to return any time soon. Gypsy moths arrived in the area in the early 1980s and for several years in succession stripped the oaks (especially Quercus bicolor) of leaves. The infestation seems to have stabi- lized and is in decline in the early 1990s. The aquatic animals have declined. When I first started ex- ploring the area, harbour seals sometimes were seen on the rocks at Lyddy Island and the outer islands were homes to large colo- nies of common terns. In the clear shallow waters, horseshoe crabs were common, and edible fish such as flounder and black- fish were plentiful offshore. Steamer clams could be dug some- what easily in the mudflats of the estuaries. During these earlier years, sea urchins and sea cucumbers could be found in the lower tide pools, squid sometimes were found breeding in the shallow waters north of Lyddy Island, and blue mussels abounded. While this area has been spared of much of the pollution found in other parts of coastal Connecticut it has not been spared all of it. In the last decade and a half the fishing areas and shellfish beds have been closed often due to algal and bacterial blooms, which, along with silt, nearly obscure the formerly clear waters. The coastal trawlers (and, possibly, pollution) have eliminated nearly all of the edible sport fish. The mussels and steamer clams nearly are gone (probably due to over-harvesting as well as siltation), and I have not seen a sea urchin, or sea cucumber, or even a starfish there for years. On the encouraging side, the central por- tion of Latimer Point has been protected from development by The Nature Conservancy, the bird and mammal fauna seems healthy, quahogs and cherrystone clams are rather common, and a seal or two has been seen again in Fishers Island Sound. ACKNOWLEDGMENTS. I would like to thank the late Frank Gould (TAES) for assistance with the identification of some grass- es, Tony Reznicek (MICH) and Gordon Tucker (NYS, now at EIU) 216 Rhodora [Vol. 98 for assistance with sedge identifications, Craig W. Schneider (tr) for assistance with the algae, and Lewis Anderson (DUKE) for assistance with the mosses. I also thank Stacey Kingsbury of the Connecticut Natural Resources Center for information on histor- ical species records for listed Connecticut plants. Comments on the manuscript by Rick Phillippe and Geoff Levin (both ILLS) and two anonymous reviewers have been especially useful. Special thanks go to my parents Margaret E. and Richard J. Hill, to whom this paper is dedicated, for putting up with a budding botanist and the inconveniences and associated messes over the years of this study, and for constant encouragement to explore and retain a sense of wonder. LITERATURE CITED Downan, J. J. 1979. Preliminary Checklist of the Vascular Flora of Con necticut. Report of Investigations No. 8. Connecticut Geological and Natural History Survey, Hartford, CT. GLEASON, H. A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. 2nd ed. The New York Botanical Garden, Bronx, NY. Hitt, S. R. 1995. How to make a plant collection. Herbarium Supply Co., Menlo Park, CA. HoL”MGREN, P. K., N. H. HOLMGREN, AND L. C. BARNETT, eds. 1990. Index Hemartomm. Part I: The aaa of the World. 8th ed. New York Bo- tanical Garden, Bronx, SCHNEIDER, C. W. AND R. B. Soa ES. 1991. Seaweeds of the Southeastern United States. Duke University Press, Durhan SEYMOUR, FC. 1969. The Flora of New Breland: — E. Tuttle Co., Inc., Rut- land, VT. THompson, E. 1950. Draggerman’s Haul. Viking Press, New York. Tucker, G. C. 1995. The Vascular Flora of Southeastern Connecticut. Mem- oirs of the Connecticut Botanical Society No. 3. RHODORA, Vol. 98, No. 894, pp. 217, 1996 BOOK REVIEW Howard, Richard A. 1996. An Almanac of Botanical Trivia. 52 pp. $7.00 (paper). Privately published by the author, 4 Jef- ferson Drive, Acton, MA 01720. All of us doubtless have our little collections of curious trivia, produced from time to time for enlightenment, embellishment, or entertainment of classes, after-dinner audiences, friends at parties, or ceremonial presentations [‘‘there must be some great signifi- cance, my colleague, to your retiring on May 24, the anniversary of the birth of Linnaeus ...”’]. Surely few if any botanists have so large a store of trivia as Dr. Howard, and none have done so systematic a job of curating their collection. The booklet at hand is arranged chronologically, with most days of the year represented by one or more statements of birth, death, first-day postal issues, bizarre occurrences in the lives of botanists, or other historic facts. Some less precisely dated items are added and an appendix lists generic names that are anagrams of others (e.g., Filago, Gifola, Ifloga, Logfia). I learn that the day on which my review copy arrived was the 40th anniversary of the death of H. N. Ridley ‘“‘at age 100 plus 10 months.”’ The day on which I mail my review to the editor is the 181st anniversary of the shipwreck of C. S. Rafinesque off Long Island (“‘after the War of 1912” [sic]). A few of the events celebrated are the same as those on the fascinating wall calendars published by the Mis- souri Botanical Garden (sharing even their misspelling of the ees of Asa Gray on November 18, 1810). But many of the “‘trivia’’ are not easily found elsewhere. (J had not previously realized that I share a birthday with William Trelease.) The author solicits additional items, so perhaps larger—but never complete—editions are contemplated. I suggest for the next one the addition of October 18, 1993: death of Ralph R. Stewart, at the age of 103 years plus 6 months, thus well surpassing H. N. Ridley’s record as the longest-lived botanist. And I suggest adding July 1, 1917, birthdate of the energetic assembler without whose efforts we would not have this delightful compilation. —EDWARD G. Voss, University of Michigan Herbarium, Ann Ar- bor, MI 48109-1057. 217 RHODORA, Vol. 98, No. 894, pp. 218-219, 1996 BOOK REVIEW Semple, John C. 1996. A Revision of Heterotheca sect. Phyllo- theca (Nutt.) Harms (Compositae: Astereae): The Prairie and Montane Goldenasters of North America. University of Waterloo Biology Series No. 37. 164 pp., illus. $15.00 plus $5.00 shipping and handling (paper). Published by U. W. Biology Series, Department of Biology, University of Wa- terloo, Waterloo, Ontario, Canada N2L 3G1. The goldenasters (golden asters) and camphorweeds are yel- low-flowered, summer and fall-blooming, annual and perennial herbs ranging in eastern North America from the Massachusetts coastal plain south into Texas and in the west from Canada to southern Mexico. Their generic circumscriptions have been “‘in turmoil”” (Semple’s phrase) since the early 1950s. Thus the only somewhat widespread New England species, Chrysopsis falcata (Pursh) Ell. in the great mid-century floras of the region, became for a time Heterotheca falcata (Pursh) Harms and is now Pityop- sts falcata (Pursh) Nutt., a name it will seemingly retain in Flora North America. This very well done monograph would be useful alone for its introductory sections expounding the nomenclatural history of this complex group. According to Semple, most North American goldenasters are divided among Chrysopsis (9 species), Heterotheca (28 species), and Pityopsis (7 species). Pityopsis and Chrysopsis are centered in the southeastern United States while Heterotheca is most abun- dant and diverse in the west. Within the United States, Hetero- theca sect. Phyllotheca occurs largely in the prairie and montane regions west of the Mississippi, although H. camporum (Greene) Shinners var. camporum extends through the Prairie Peninsula region into western Indiana and H. camporum var. glandulissi- mum Semple has recently spread to a number of eastern sites. Perhaps surprisingly, most of the twenty species in sect. Phyl- lotheca and their varieties historically have been included within Chrysopsis villosa (Pursh) Nutt., characterized by H. A. Gleason in 1952 as “‘a complex and poorly understood aggregate species, of uncertain limits,” its often distinctive varieties “‘connected by numerous intergrades.”” Semple, in revising the group, has taken on a daunting task, and at the very least has established a coherent system for a widespread and protean assemblage, with clear de- 218 1996] Book Review 219 scriptions of its taxa. His taxonomic treatment includes keys to the three sections and 28 species of Heterotheca and their vari- eties, a general summary of section Phyllotheca, and accounts of each of its species and varieties. Included are range maps in suf- ficient detail to delight the biogeographers among us and black and white illustrations showing growth habit, leaves, and, in very fine detail, the reproductive structures of each taxon. There are also lengthy lists of cited specimens and narrative discussions with notes on aberrant or atypical forms, potentially very useful information of a sort too quickly glossed over in broad floristic treatments. Semple warns the reader early-on that the taxa within Phyllotheca are defined by combinations of traits rather than by single diagnostic features and that within most species, leaf in- dument (often featured prominently within his keys) varies in a gradient from taxa possessing numerous hairs and few or no glands to closely related forms with few hairs and many glands. Indeed, within some specimens, the indument of the basal leaves may vary sharply from that of the upper leaves, reflecting differ- ent environmental conditions at the time of leaf formation. Thus, I suspect that despite Dr. Semple’s efforts, because of its inherent variability, section Phyllotheca will continue to present difficulties for field biologists. Hence, whether or not his treat- ment, which proposes six new names and combinations within section Phyllotheca alone, will survive the inevitable revisions of the 21st Century, is very much an open question. If his structure is in time dismantled, he has nonetheless brought together here the building blocks to shape another. —C. JOHN Burk, Department of Biological Sciences, Smith Col- lege, Northampton, MA 01063. RHODORA, Vol. 98, No. 894, pp. 220-222, 1996 NEBC MEETING NEWS April 1996. Dr. Christopher Campbell of the University of Maine spoke on “Visions of Sugar Plums Dancing in My Head: An Apomictic Nightmare.’’ He was led to the study of the very dif- ficult genus Amelanchier in part because he lives in the center of diversity of the genus, and because the University thoughtfully planted research material at the door of his office. He obviously hadn’t read Ed Voss’s statement that ““The only advantage that Amelanchier has over Crataegus and Rubus is that, by being smaller, it lures us to the hope that it will be more manageable.” The shadbushes are a taxonomic nightmare, and appear to be a complex of hybrids and apomictic (asexual) taxa that are re- ferred to as “‘microspecies.’’ Dr. Campbell has been investigating whether agamospermy actually is responsible for the formation of microspecies, attempting to determine how much hybridization actually occurs, and the extent to which these two forces shape evolution in the genus. Amelanchier appears to be an old genus, represented in eastern Asia, Europe, and Asia Minor, as well as in western and eastern North America where the highest diversity occurs. One species of western North America, A. a/nifolia, is cultivated for its edible fruits known as “‘sugarplums.”’ Although not morphologically distinct, the western and eastern North American taxa are highly distinct genetically. This genetic difference is most clearly reflect- ed in fruit taste—all of the tasty species are western, with the exception of the eastern A. humilis. There is no consensus on the actual number of species in the genus; various treatments range from 6 to 33, with virtually all of the taxa polyploid and most at least facultatively agamospermic. All possible hybrids have been reported for the New England taxa. Dr. Campbell’s research has combined studies of morphological and genetic variation, as well as experimental hybridizations. A series of controlled crosses shows that inheritance of genetic markers is strongly maternal, indicating that hybridization rarely occurs and that sex is not responsible for the introduction of ge- netic variation. He compared within and among population vari- ation for Amelanchier bartramiana (a sexual species) and A. /ae- vis (an agamospermous species). This research has shown that, contrary to most hypotheses, sexual and apomictic species do not partition variation differently, and that variation is greater in the 220 1996] NEBC Meeting News 221 sexual species. Results were not conclusive on the question of whether agamospermy is responsible for the formation of micro- species. This research on hybridization has shown clearly that Amelanchier X neglecta is a hybrid of A. bartramiana and A. laevis, with the apomictic A. /aevis serving as the pollen parent. Amelanchier X neglecta is almost entirely agamospermous, with no evidence of backcrossing with its parents. Other research on a confusing plant known informally as Ame- lanchier “‘humilopsis”” suggests that it is an old hybrid of A. humilis and an unknown species, both agamospermous. This plant has an unusual multigene polymorphism that is maintained by agamospermy. Dr. Campbell has observed “‘humilopsis”’ and A. laevis hybridizing at a disturbed site, and hypothesizes that Fer- nald saw these plants as a child growing up in Orono. It appears that agamospermy by itself does not generate vari- ation in Amelanchier, but perpetuates the diversity that results from extensive hybridization. Future attempts to resolve the tax- onomy of Amelanchier should identify and focus on the sexual species that are the oldest and most stable members of the genus. May 1996. Dr. Gregory Anderson, of the University of Con- necticut at Storrs, spoke on “The Origin and Evolution of the Pepino, One of the Forgotten Domesticates of the Incas.” The Pepino (Solanum muricatum) is truly a lost crop, with no known wild progenitors. Domesticated prior to the Incas in the Ecuador/Columbia/Peru area, pepinos appear in pottery dating back 2,000—3,000 years, but were not much cultivated by Incan cultures. The name derives from the Spanish, who named it “‘pe- pino dulce” to distinguish it from the earlier-named “‘pepino” or cucumber. Now becoming more popular as a “‘dessert-quality subacid fruit” with a high vitamin C content, it is cultivated from Mexico to Chile as well as in New Zealand. Pepinos exhibit great variation in fruit morphology, consistent with the “‘first law of economic botany,’’ which states that the greatest variation occurs in that feature for which the species was cultivated. The result of recent intensive cultivation has been a substantial loss in fruit diversity. Morphological studies suggested that there were 3 candidates for the wild progenitor of pepino: Solanum tabanoense, S. cari- pense, and S. basendopogon. Studies by Greg and numerous col- laborators have examined “‘traditional’? biosystematic evidence Pen Rhodora [Vol. 98 from breeding systems, genetic relationships, flavinoids, meiotic behavior, and mitotic chromosome structure. These studies sug- gested that relationships based on morphology should be revised, and indicated that Solanum caripense was most closely related to the pepino. More recent studies have focused on molecular evidence using chloroplast and nuclear DNA sequencing. These DNA studies indicate that Solanum tabanoense, not S. caripense, is most close- ly related to the pepino. However, some collections of pepino were very similar to S. caripense. These data are best explained by the hypothesis that pepino was domesticated from S. taban- oense, and later was hybridized with S. caripense. The large range of morphological variation in the pepino is likely to be the result of this post-origin hybridization. —LISA A. STANDLEY, Recording Secretary. INFORMATION FOR CONTRIBUTORS TO RHODORA Submission of a manuscript implies it is not being considered for publication simultaneously elsewhere, either in whole or in part. GENERAL: Manuscripts should be submitted in triplicate. The text must be double-spaced throughout, including tables, figure legends, and literature citations. Use a non-proportional font throughout and do not justify the right margin. Do not indicate the style of type through the use of capitals, underscoring, or bold, except for names of genera and species which should be in italics or underscored throughout. Do not underline punctuation. All pages should be num- bered in the upper right-hand corner. For guidance in matters not addressed here, consult the editorial office by phone at (603) 862- 3205, FAX (603) 862-4756, or e-mail: janets@christa.unh.edu. Brev- ity is urged for all submissions. TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital letters. Omit authors of scientific names. Below title, include au- thor(s) name(s), affiliation(s), and address(es). If “present address”’ is different, it should follow immediately below, not as a footnote. ABSTRACT: An abstract and a list of key words should be included with each paper, except for shorter papers submitted as Notes. An abstract must be one paragraph, and should not include literature citations or taxonomic authorities. Please be concise, while including information about the paper’s intent, materials and methods, results, and significance of findings. TEXT: Main headings are all capital letters and centered on one line. Examples are: MATERIALS AND METHODS, RESULTS, and DIS- CUSSION. Do not title the Introduction. Do not combine sections of the paper (such as Results and Discussion), or use Conclusions or Summary. Second level headings should be indented, bold, upper and lower case, and end with a period. Taxonomic authorities should be cited for all species names at their first usage in the text, or in a referenced table. Cite each figure and table in the text in numerical order. Each reference cited in the text must be in the Literature Cited. Cross-check spelling of author(s) name(s) and dates of publication. Literature citations in the text should be as follows: Hill (1982) or (Hill 1982). For two or more authors, cite as follows: Angelo and Boufford (1996) or (Angelo and Boufford 1996). Within parenthesis, use a semicolon to separate different types of citations (Hill 1982; Angelo and Boufford 1996) or (Figure 4; Table 2). FLORAS AND TAXONOMIC TREATMENTS: Specimen citation should be selected critically, especially for common species of broad distribution. Keys and synonymy for systematic revisions should be prepared in the style of “‘A Monograph of the Genus Malvastrum,” 223 i) i) aN S. R. Hill, RHODORA 84: 159-264, 1982. Designation of a new taxon should carry a Latin diagnosis (rather than a full Latin descrip- tion), which sets forth succinctly how the new taxon differs from its congeners. LITERATURE CITED: All bibliographic entries must be cited in the paper, unless a special exception has been made by the Editor (such papers will be allowed a REFERENCES section). Verify all entries against original sources, paying special attention to spelling and de- tails of publication. Cite references in strict alphabetical order by first author’s surname. Do not write authors’ names in all capital letters. References by a single author precede multi-authored works of same senior author, regardless of date. Use a long dash when the author(s) are the same as in the entry immediately preceding (see recent is- sues). Refer to Botanico-Periodicum-Huntianum (B-P-H 1968) and B-P- H/Supplement (1991) for standardized abbreviations for jour- nals. TABLES: Tables must be double-spaced. Tables may be continued on an extra page, if necessary. As much as possible, the title should be self-explanatory. Do not use footnotes; instead, add notes after the end of the table title. Broadside tables should be avoided, if pos- sible. Each table should be cited in the text in numerical order. FIGURES: Illustrations must be either black and white half-tones (photograph), drawings, or graphs. Illustrations must be camera- ready; flaws cannot be corrected by the Editor or the printer. Add symbols or shading with press-on sheets. The printed plate will be 4 X 6 inches; be sure that illustrations are proportioned to reduce correctly. Allow space for a caption, if possible. Magnification/re- duction values should be calculated to reflect the actual printed size. Maps must indicate scale and compass direction. The double-spaced list of legends for figures should be provided on a separate page. Each figure should be cited in the text in numerical order THE NEW ENGLAND BOTANICAL CLUB 22 Divinity Avenue Cambridge, MA 02138 The New England Botanical Club is a non-profit organiza- tion that promotes the study of plants of North America, es- pecially the flora of New England and adjacent areas. The Club holds regular meetings, and has a large herbarium of New En- gland plants and a library. It publishes a quarterly journal, RHODORA, which is now in its 99th year and contains about 400 pages per volume. Visit our web site at http://www.herbaria. harvard.edu/nebc/ Membership is open to all persons interested in systematics and field botany. Annual dues are $35.00, including a subscription to RHODORA. Members living within about 200 miles of Boston receive notices of the Club meetings. To join, please fill out this membership application and send with enclosed dues to the above address. Regular Member $35.00 Family Rate $45.00 Student Member $25.00 For this calendar year ae For the next calendar year eee Name Address City & State Zip Special interests (optional): RHODORA, Vol. 98, No. 894, pp. 226-231, 1996 BYLAWS OF THE NEW ENGLAND BOTANICAL CLUB, INCORPORATED (To repeal and replace all previous Bylaws) Adopted by the membership 5 April 1996 Bylaws last published: Rhodora, vol. 91, no. 867, pp. 270-276 (1989) ARTICLE I NAME AND PURPOSE Section |. The name of this corporation shall be The New Eng- land Botanical Club, Incorporated. Section 2. Its purpose shall be to promote the dissemination of local and general botanical information. Section 3. The Club shall be organized and operated exclusively for its educational and scientific purposes. No part of the property or net earnings of the Club shall inure to the benefit of any in- dividual; and no part of the property of the Club shall be used directly or indirectly in carrying on propaganda, nor shall any substantial part of the activities of the Club consist of the carrying on of propaganda or otherwise attempting to influence legislation. The Club shall not participate in, nor intervene in, any political campaign on behalf of any candidate for public office, nor shall it publish or distribute any statements with respect thereto. ARTICLE II The seal of the Club shall, subject to alteration by the Council, consist of a flat-faced circular die with the words *‘Massachu- setts,” “Organized 1895, Incorporated 1920,” and the name of the corporation cut or engraved thereon. 226 1996] NEBC Bylaws 22) EE NE ie ARTICLE III OFFICERS AND COUNCILLORS Section 1. The officers of the Club shall be a President, a Vice- President, a Corresponding Secretary, a Recording Secretary, a Treasurer, a Curator of Vascular Plants, an Assistant Curator of Vascular Plants, a Curator of Nonvascular Plants, and a Librarian. There shall be five councillors, one of whom may be a student and one of whom may be the immediately preceding President. Section 2. All officers and councillors shall be elected by ballot by a majority of those members voting at the Annual Meeting of the Club, and, except in the case of death, resignation, or removal, each officer shall hold office until the next Annual Meeting or until a successor is elected. Voting by proxy shall not be allowed. Section 3. If the office of any officer or councillor becomes va- cant by reason of death, resignation, or removal, the Council may appoint a successor who shall hold the office until the next An- nual Meeting or until a successor is elected. ARTICLE IV COUNCIL Section 1. The Council shall consist of the above-named officers and councillors, the Associate Curator and the Editor-in-Chief of Rhodora. The Associate Curator shall be appointed by the Coun- cil to serve as a liaison with the Harvard University Herbaria. The Editor-in-Chief of Rhodora shall be appointed by the Coun- cil. 228 Rhodora [Vol. 98 Section 2. The Council, a majority of which shall constitute a quorum, shall have the management and control of the Club and of all its property and affairs and shall direct the expenditure of its funds. Section 3. The Council shall authorize and approve, except as the Council may generally or in particular cases authorize the execution thereof in some other manner, all deeds, transfers, and contracts; and all bonds or notes made or endorsed by the Club shall be signed by the Treasurer and countersigned by the Presi- dent or the Vice-President. All deeds, transfers, and contracts shall be signed by the President or in the President’s absence by the Vice-President. Section 4. The Council may appoint and remove such other of- ficers or agents as it may from time to time determine. It may appoint committees as it sees fit and may delegate to these com- mittees such powers for such terms as the Council deems best, subject to the power of the Council to revoke any such appoint- ment at any time. Section 5. The Council shall have the books and accounts of the Treasurer audited at least once a year by an external Auditor. ARTICLE V DUTIES OF THE OFFICERS Section |. The President and Vice-President shall perform the usual duties of their offices. Section 2. The Curator of Vascular Plants and the Curator of Nonvascular Plants shall have charge of the botanical collections of the Club. The Assistant Curator of Vascular Plants (or in that officer’s absence, the Associate Curator) shall perform the duties of the Curator of Vascular Plants in the Curator’s absence. Section 3. The Librarian shall have charge of the books and manuscripts of the Club. Section 4. The Corresponding Secretary shall give notices of all meetings of the Club and of the Council, and shall conduct the correspondence of the Club. 1996] NEBC Bylaws 229 Section 5. The Recording Secretary, who shall be the Clerk, shall be sworn before entering upon the duties of the Recording Sec- retary, and as such shall keep the minutes of all meetings of the Club and of the Council and such other records as the Council may direct. Section 6. The Treasurer shall, subject to the orders and super- vision of the Council, collect and disburse the funds of the Club, and for this purpose shall have power to endorse for deposit or collection, all funds, checks, drafts, etc. payable to the corporation or its order. The Treasurer shall keep, or cause to be kept, accurate books of account, and shall make a report of the financial con- dition of the Club at each Annual Meeting, and at such other times as the Council may request. AR TICLE-V] NOMINATING COMMITTEE On or before December fifteenth of each year, the President shall appoint a committee of three members—who shall not be offi- cers—to nominate officers for the ensuing year. The report of this committee, which shall be filed with the Corresponding Secretary and open to inspection at least three weeks prior to the Annual Meeting, shall be incorporated in the call for the Annual Meeting. Nothing herein shall restrict the right of members to offer nom- inations from the floor provided a notice listing such nominees and signed by not less than three members shall be filed with the Corresponding Secretary not later than ten days prior to the date of the meeting. ARTICLE VII MEMBERSHIP AND DUES Section 1. Membership inquiries and resignations may be made to the Corresponding Secretary. Section 2. Upon payment of annual dues, all members shall have voting privileges and shall receive Rhodora. Section 3. Dues and subscription price of Rhodora shall be de- termined at appropriate intervals by the Council. 230 Rhodora [Vol. 98 Section 4. Annual dues shall be payable the first of January. Any member whose dues remain unpaid for more than three months after same become payable shall cease to receive Rhodora. The membership of any member shall automatically terminate if dues or other indebtedness to the Club remain unpaid for thirteen months after same become payable. Section 5. In the event of the death, resignation, or other termi- nation of the membership of a member, all privileges shall cease. Dues will not be prorated and reimbursed except in instances where more than one year’s membership remains. ARTICLE VIII MEETINGS OP THE CLUB Section |. The Annual Meeting of the Club shall be held on the first Friday in March of each year, unless otherwise ordered by the Council, and regular meetings shall be held monthly, except during July, August, and September at such times as the Council may determine. If the Annual Meeting is omitted by oversight or otherwise on the day herein provided therefor, a special meeting may be held in place thereof, and any business transacted or elec- tions held at such meeting shall have the same effect as though transacted or held at the Annual Meeting. Section 2. Special meetings may be called by the President or by vote of the Council or by written request of any ten members given to the Corresponding Secretary. Every such call shall state the object for which the meeting is being called. Section 3. Notice of all meetings shall be distributed at least sev- en days before such meetings. Section 4. At any meeting of the Club, fifteen members shall constitute a quorum for the transaction of business, except the amendment of the Bylaws. ARTICLE IX AMENDMENTS OF THE BYLAWS These Bylaws may be altered, amended, or repealed in the fol- lowing manner: at any Annual, regular, or special meeting of the Club by a two-thirds vote of the members present and voting, 1996] NEBC Bylaws 231 provided that at least thirty members or two-thirds of the mem- bership (whichever is the smaller number) are present at the meet- ing, and that the subject matter of the proposed alteration, amend- ment, or repeal has been given in the call for the meeting at which the alteration, amendment, or repeal is to be considered. ARTICLE X FISCAL YEAR The fiscal year of the Club shall end on the thirty-first day of December in each year. ARTICLE XI DISSOLUTION Should the Club be dissolved for any reason, the officers shall, after paying or making provision for payment of all the liabilities of the corporation, distribute all assets, including all accrued in- come, to one or more scientific, educational and/or literary or- ganizations under Section 501(c)(3) of the Internal Revenue Code of 1986 (or the corresponding provision of any subsequent United States Internal Revenue Law). THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 1996—1997: President: W. Donald Hudson, Jr., Chewonki Foundation, RR 2, Box 1200, Wiscasset, ME 04578 Vice-President (and Program Chair): David S. Conant, Depart- ment of Natural Sciences, Lyndon State College, Lyndon- ville, VT 05851 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Bi- ology, North Adams State College, North Adams, MA 01247-4100 Recording Secretary: Lisa A. Standley Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Pamela B. Weatherbee Curator of Nonvascular Plants: Anna M. Reid Librarian: Paul Somers Councillors: C. Barre Hellquist (Past President) Garrett E. Crow 1997 Matthew Hickler (Graduate Student Member) 1997 Edward Hehre, Jr. 1998 Michael J. Donoghue 1999 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora _ a 7 ha 7 _ 7 7 7 ; : 7 - _ a a oS a . — oe , 7 7 7 a - ; 7 ; 7 a | ; 7 a SS ee _ : . Oe . —_ OS | - a 7 7 a : on Oo oe oO _ - : oo : : : ; 7 . _ : : fi : : 7 oe _ 7 —— : - : a a : : a a a a ane a a oe : a 7 _ _ 7 - : 7 _ ; > - 7 ; : _ ; a 7 ee : 1 i 7 i _ ee a a —_ : 7 ran _ ne Ss re SS iether: a a _ | a . . : : 7 | | - 7 anh ae ns on 7 7 7 RHODORA The Journal of the CONTENTS New England Botanical Club Flora Conservanda: ae England. The New England Plant Conservation ogram (NEPCoP) list of plants in need of conservation. William E. Brumback and ee J. Mehrhoff, in collaboration with Richard W. Enser, Susan C. Gawler, Robert G. Popp, Paul Somers, and Daniel D. Sperduto, with assistance from William D. Countryman and C Barre Hellquist Introduction 235) Development of the NEPCoP **Flora Conservanda: New England” ...... 238 Format of the NEPCoP List 242 Discussion 247 Recommendations 249 Flora Conservanda: New England 25 Literature Cited 332 Appendix I. State Status Codes 334 Appendix II. Global Ranks 336 Appendix III. State Ranks 337 Appendix IV. Federal Listing Designations 338 Index to the NEPCoP List 340 Vol. 98 Summer, 1996 issued: July 7, 1997 No. 895 The New England Botanical Club, Inc. 22 Divinity Avenue, Cambridge, Massachusetts 02138 RHODORA JANET R. SULLIVAN, Editor-in-Chief Department of Plant Biology, University of New Hampshire, Durham, NH 03824 MARGARET P. BOGLE, Managing Editor Department of Plant Biology, University of New Hampshire, Durham, NH 03824 Associate Editors HAROLD G. BROTZMAN THOMAS D. LEE CHRISTOPHER S. CAMPBELL LESLIE J. MEHRHOFF DAVID S. CONANT THOMAS MIONE GARRETT E. CROW LISA A. STANDLEY NANCY M. EYSTER-SMITH K. N. GANDHI—Latin diagnoses and nomenclature RHODORA (ISSN 0035-4902). Published four times a year (January, April, July, and October) by The New England Botanical Club, 810 East 10th St., Lawrence, KS 66044 and printed by Allen Press, Inc., 1041 New Hampshire St., Lawrence, KS 66044-0368. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send address changes to RHODORA, P.O. Box 1897, Lawrence, KS 66044-8897. RHODORA is a journal of botany devoted primarily to the flora of North America. Monographs or scientific papers concerned with systemat- ics, floristics, ecology, paleobotany, or conservation biology of the flora of North America or floristically related areas will be considered. SUBSCRIPTIONS: $75 per calendar year, net, postpaid, in funds pay- at par in United States currency. Remittances payable to RHO- DORA. Send to RHODORA, PO. Box 1897, Lawrence, KS 66044- 8897. MEMBERSHIPS: a Sead $35; Family $45; Student $25. Application form printed her NEBC WEB SITE: Information about The New England Botanical Club, its history, officers and councillors, herbarium, monthly meet- ings and special events, annual graduate student award, and the jour- nal RHODORA is available at http://www.herbaria.harvard.edu/nebc/ BACK ISSUES: Information on availability of back issues should be addressed to Dr. Cathy A. Paris, Department of Botany, University of Vermont, Burlington, VT 05405-0086. E-mail: cparis@ moose.uvm.edu ADDRESS CHANGES: In order to receive the next number of RHO- DORA, changes must be received by the business office prior to the first day of January, April, July, or October. INFORMATION FOR CONTRIBUTORS: See contents. Submit manu- scripts to the Editor-in-Chief. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). RHODORA, Vol. 98, No. 895, pp. 233-361, 1996 FLORA CONSERVANDA: NEW ENGLAND. THE NEW ENGLAND PLANT CONSERVATION PROGRAM (NEPCoP) LIST OF PLANTS IN NEED OF CONSERVATION WILLIAM E. BRUMBACK and LESLIE J. MEHRHOFF New England Wild Flower G. Safford Torrey Herbarium Society Box U-42 OQ Hemenway Road University of Connecticut Framingham, MA 01701 Storrs, CT 06269 in collaboration with RICHARD W. ENSER, Rhode Island Natural Heritage Program SUSAN C. GAWLER, Maine Natural Areas Program RoBERT G. Popp, Vermont Nongame and Natural Heritage Program PAUL SOMERS, Massachusetts Natural Heritage and Endangered Species Program DANIEL D. SpERDUTO, New Hampshire Natural Heritage Inventory Program with assistance from WILLIAM D. COUNTRYMAN, Aquatech, Inc. C. BARRE HELLQuiIstT, North Adams State College N es) Qo 234 Rhodora [Vol. 98 COMPLETE ADDRESSES OF COLLABORATING AUTHORS Richard W. Enser, Rhode Island Natural Heritage Program, De- partment of Environmental Management, 235 Promenade St., Providence, RI 02908. Susan C. Gawler, Maine Natural Areas Program, 93 State House Station, Augusta, ME 04000-0093. Robert G. Popp, Vermont Nongame and Natural Heritage Pro- gram, Department of Fish and Wildlife, Waterbury, VT 05676 Paul Somers, Massachusetts Natural Heritage and Endangered Species Program, Massachusetts Division of Fisheries and Wildlife, Rt. 135, Westborough, MA 01581 Daniel D. Sperduto, New Hampshire Natural Heritage Inventory Program/DRED, Box 1865, Concord, NH 03302-1856. William D. Countryman, Aquatech, Inc., R. D. 1, Box 990, Northfield, VT 05663 C. Barre Hellquist, Biology Department, Box 9145, North Adams State College, North Adams, MA 01247. 1996] Brumback and Mehrhoff, et al—NEPCoP Pe ABSTRACT. The New England Plant Conservation Program (NEPCoP) re- gional rare plant list, ““F/ora Conservanda: New England,” identifies vascular plant taxa in need of regional conservation. In 1993, NEPCoP established a Listing Committee consisting of representatives of each of the six state Nat- ural Heritage Programs (NHPs) and additional scientists. The most current information on the rare vascular flora of each state, stored at the respective NHPs, was used as the basis of the NEPCoP List. The List comprises 576 taxa in five divisions: Division |—Globally Rare Taxa (57 taxa); Division 2—Regionally Rare Taxa (273 taxa); Division 3—Locally Rare Taxa (75 taxa); Division 4—Historic Taxa (55 taxa); and Division Indeterminate (IND.)—116 taxa. Key Words: NEPCoP, New England, conservation, endangered species, plants, regional plant program, Flora Conservanda, regional conservation list The New England Plant Conservation Program (NEPCopP), a voluntary collaboration of botanists, state and federal agencies, and conservation organizations in each of the New England states, was initiated by the New England Wild Flower Society (NEWFS) in 1991. The goals of NEPCoP are to prevent the ex- tirpation and promote the recovery of the endangered flora of New England (New England Wild Flower Society 1992). The Program provides regional coordination for state-based plant con- servation efforts to make best use of limited resources. NEPCoP was initiated for several reasons: |) on a global and regional scale, plant species are under extreme threat; 2) a lack of public awareness concerning the importance of plants has contributed to plant endangerment; and 3) there is inadequate support for plant conservation activities from both the public and private sectors. The Program is administered by a Regional Advisory Council, Task Forces in the six New England states, and WES. The Council, consisting of representatives from each state Task Force, members of the Board of Trustees at NEWEFS, and other repre- sentatives from other conservation organizations, convenes as necessary to monitor and advise the overall Program. The Council sets policy for regional plant conservation, oversees development e ‘“‘Flora Conservanda: New England,” and selects priority species for conservation action throughout the region. State Task Forces are the heart of the program. Each Task Force, comprised of individuals knowledgeable of the state’s flo- ra, selects populations of priority species to be surveyed and di- 236 Rhodora [Vol. 98 rects conservation actions. The Task Forces meet at least once annually to discuss progress and set conservation agendas. NEPCoP integrates in situ with ex situ conservation methods. In situ methods. The best method for conserving plants is to pro- tect the habitats where wild populations occur. Protection of land in itself, however, may not ensure the perpetuation of plant populations. Changes in habitats over time may neces- sitate management in order to preserve rare species. Mem- bers of each state’s Task Force survey occurrences of rare plants, identify threats, and make suggestions for future man- agement. Ex situ methods. As a complement to protection and management of wild plant populations, plant propagules are collected from selected populations for seed banking, research, and public education. A seed bank of endangered species has been es- tablished at Garden in the Woods, the botanic garden of the New England Wild Flower Society in Framingham, Massa- chusetts, as a backup in the event of catastrophic loss in the wild. Seeds are collected from vulnerable populations of pri- ority species as determined by each state Task Force. The number of populations sampled depends on many factors in- cluding the size and number of occurrences within each state, the type of land ownership, and each species’ potential for successful seed banking. An important part of the seed banking process is research on the propagation and cultivation of rare plant species. As seeds are tested for banking, the optimal propagation methods for each species are researched, and any plants produced can be made available for research. Most importantly, this propagation re- search provides information that can be applied to management of wild populations as well as propagules for reintroduction if deemed necessary. In addition, plants obtained through propa- gation research become part of the collection of the New England Garden of Rare and Endangered Plants, also maintained at the Garden in the Woods. This collection is an educational display for the public and a genetic resource for conservation and re- search. Realizing that continual monitoring of all regional rare plant populations will not be accomplished by the currently limited 1996] Brumback and Mehrhoff, et al—NEPCoP Zot number of professionals, NEPCoP has instituted a pilot program of Volunteer Rare Plant Monitors. This program, at present in Massachusetts only, trains volunteers to survey rare plant occur- rences. The initial results have been impressive and the expansion of this program both within Massachusetts and throughout the region is anticipated. New England states have had a strong interest in conservation of the native flora for many years. Connecticut passed a law in 1868 to protect the Hartford Fern, Lygodium palmatum (Mehrhoff 1980), which may be the earliest legal attempt at conservation of plants for aesthetic interest rather than for utilitarian reasons. The New England Botanical Club (NEBC) formed its first con- servation committee, the Natural Areas Criteria Committee, in 1971. The Endangered Species Committee was an offshoot of this initiative and in 1975 began the preparation of individual state reports of rare and endangered vascular plants with the support of the U.S. Fish and Wildlife Service. These reports included: Maine (Eastman 1978), New Hampshire (Storks and Crow 1978), Vermont (Countryman 1978), Massachusetts (Coddington and Field 1978), Rhode Island (Church and Champlin 1978), and Connecticut (Mehrhoff 1978). These state lists served as a basis for the NEBC regional list (Crow et al. 1981). This regional list included 479 taxa, the ma- jority of which were either “‘E/T”’ (Endangered/Threatened—337 taxa) or “R”’ (Rare—133 taxa). Taxa designated as “‘E/T’’ were documented from 10 or fewer towns and “‘R”’ were documented from 10 to 20 towns. In addition, other designations, such as federal status or consideration, were included. Occurrences for each taxon within each state were noted by a variety of desig- nations. The 1981 list was ‘based, in large part, on historical records documented by specimens in herbaria’? (Crow et al. 1981). Prior to this publication there had been little concerted effort to collect or document regionally rare taxa, and the NEBC committee was faced with the daunting task of determining which taxa were rare throughout the region. This was accomplished using the first-hand knowledge of state experts, information culled from the literature, and selective checking of herbarium records. Often, there was no way to ascertain the current status of a taxon with only specimens and literature. Beginning in 1978, The Nature Conservancy facilitated the es- 238 Rhodora [Vol. 98 tablishment of state Natural Heritage Programs (or their equiva- lent) in each of the New England states. Similar programs already existed in some states. These Programs undertook and encouraged intensive field work in their respective states, resulting in many new or updated records for rare taxa. Because of this field work, many individual state lists were refined to reflect the more current status information. Some taxa remained elusive or showed a de- cline while others were shown to be more common and were removed from state lists. During the 1980s, most New England states formulated state laws regarding rare or endangered plants (see Appendix I for current state laws). DEVELOPMENT OF THE NEPCOP “‘FLORA CONSERVANDA: NEW ENGLAND’”’ Purposes. In order to guide regional efforts, NEPCoP first needed to identify species and populations of regional conser- vation concern. Towards this end, a Listing Committee of the Regional Advisory Council was formed to develop and maintain a regional list of plants known as ‘Flora Conservanda: New England” (often referred to as the NEPCoP List). In addition to identifying taxa and populations of regional con- servation concern, the NEPCoP List is intended to promote the resolution of nomenclatural and taxonomic vagaries or problems and to suggest priorities for protection at both the species and population levels. It is hoped that it will aid the development of priorities for research, protection, and recovery on a regional ba- sis, and help states to coordinate their individual species conser- vation efforts. The NEPCoP List differs from state and federal lists in two ways: first, it provides a regional (New England) perspective on the conservation status of each taxon; second, it has no legal standing. (Legal protection or status, however, may be afforded a taxon within an individual state or through the federal government.) The NEPCoP List focuses on taxa that are globally and re- gionally rare (Divisions | and 2). It also identifies taxa that may be common throughout a significant portion of the region, but that have occurrences of conservation importance owing to their biological, ecological or (potential) genetic significance (Division 3). It further identifies taxa which are considered historic in the region (Division 4) as well as those which may be rare throughout 1996] Brumback and Mehrhoff, et al—NEPCoP 239 New England, but for which taxonomic or distributional infor- mation is insufficient to determine status (Division IND.). The List is intended to be useful to the following: 1) NEPCoP State Task Forces in selecting species for conservation; 2) sci- entists in focusing efforts on critical species; 3) federal, state, and local government agencies and private land conservation organ- izations in identifying the most important taxa to protect and man- age within the region; and 4) the public in supporting conserva- tion efforts. Methods. “Flora Conservanda: New England” was devel- oped by a Listing Committee consisting of representatives of each of the six state Natural Heritage Programs (NHPs) and several other botanists familiar with the regional flora. The most current information regarding the distribution and status of the rare vas- cular plants of each state is stored in their respective NHPs in a comprehensive Biological and Conservation Database (BCD) de- veloped by The Nature Conservancy. The records maintained in the state databases constituted the basis for developing *“‘Flora Conservanda: New England.” Each NHP monitors (tracks) a large number of species consid- ered to be rare within its state. The initial compilation of the six state lists combined with the NEBC list (Crow et al. 1981) re- sulted in the identification of more than 1 100 taxa. Approximately 200 additional taxa were reviewed by the Committee, as well. By developing strict definitions for the inclusion of a taxon within one of the five Divisions of “*Flora Conservanda: New England,” the Committee succeeded in identifying 576 taxa of highest re- gional concern out of a total of approximately 3024 indigenous or partly indigenous taxa in New England (Seymour 1969). Determination for listing is based on the number of Element Occurrences (EO) within each state. The term was devised by The Nature Conservancy and is used in conservation as an alter- native to “‘population.’’ Populations of organisms often are dif- ficult to delineate without intensive research, and use of the term ‘“‘population”’ often implies that its limits are known. Somewhat broader in scope, an occurrence is defined as follows: “‘For spe- cies.... element occurrences represent the full occupied habitat (or previously occupied habitat) that contributes, or potentially contributes, to the persistence of the species at that location. EOs are separated from each other by substantial barriers to movement 240 Rhodora [Vol. 98 or dispersal, or by specific distances defined for each element across either unsuitable, or suitable but apparently unoccupied habitat.””’ (The Nature Conservancy, Conservation Science Divi- sion, 1n association with the Network of Natural Heritage Pro- grams and Conservation Data Centers 1997). The state NHPs have made every attempt to verify the records included in the NEPCoP List. In some cases, certain occurrences were revisited during the development of the List with the intent of assessing current status and updating existing NHP files. Herbarium specimens have been crucial to the preparation of “Flora Conservanda: New England” as vouchered records of occurrences included in the List. An herbarium specimen col- lected in New England exists for every taxon included in the NEPCoP List. These specimens have helped clarify taxonomic and distributional issues, and they exist as a permanent record of a plant’s existence at a particular site and time. All data included within “Flora Conservanda: New England” are current as of December 1995; in some cases, discoveries made during the 1996 field season are included. Occurrence numbers included in Divisions 1, 2, and 3 and IND. are for occurrences verified as extant since 1970. The database used for *‘Flora Con- servanda: New England” was developed by BG-BASE, Inc., Holden Arboretum, Kirtland, Ohio. The List is dynamic, and it is the intent of the Regional Ad- visory Council to update it every five years. To facilitate this process, the state NHPs are actively seeking information on the status of listed taxa. Corrections, comments, and additional in- formation pertaining to any taxon already listed, or warranting listing, are solicited by the NHPs and NEPCoP. Nomenclature. Precise nomenclature for each taxon was of paramount concern for the Listing Committee because of the di- verse audience of anticipated users and the plethora of potential identification manuals and field guides. No single reference is used by botanists, conservationists, government officials, and wildflower enthusiasts throughout New England. The late Arthur Cronquist’s recent manual (Gleason and Cronquist 1991) likely will become a standard reference, but nomenclature in this man- ual does not in all instances match names used in some recent state checklists, i.e., ““Checklist of the Vascular Plants of Maine”’ (Campbell et al. 1995). Furthermore, some state Natural Heritage 1996] Brumback and Mehrhoff, et al—NEPCoP 24] Programs use names suggested by the national office of The Na- ture Conservancy which follows Kartesz (1994). NEPCoP’s Regional Advisory Committee adopted the follow- ing policy for nomenclature (New England Wild Flower Society 1992); 1) The primary source is to be the Flora of North America (FNA), a multi-year, multi-volume endeavor. As of this writing, only the volume covering the Pteridophyta and Pinophyta has been published (Flora of North America Editorial Committee 1993). A second volume, covering the Magnoliidae and Hama- melidae of Cronquist (1981), is in press. 2) Secondary sources are to be the authors of taxonomic treat- ments for future volumes of FNA. Although changes in nomen- clature are possible through the editing and review phases of the preparation of each volume, it is expected that most of these names are likely to pass the rigorous review of the FNA Editorial Committee and their reviewers. Consequently, for difficult taxo- nomic groups, every effort was made to contact authors of future treatments of FNA (see Acknowledgments). In some instances, however, authors have not yet been selected by the FNA editorial committee. When the nomenclature to be used in future FNA treatments coincided with that of a published source, the existing publication is cited as the primary source of the name. 3) The tertiary source for nomenclature is Gleason and Cron- quist (1991), the most recent floristic manual available for New England. Although most of this work provides an acceptable tax- onomic reference, treatments such as those for Viola and Scirpus (sensu lato) pose problems. Viola novae-angliae and Scirpus an- cistrochaetus, acknowledged by other sources including the U.S. Fish and Wildlife Service, have been included in other wide- ranging, polymorphic taxa in this manual. Similarly, other treat- ments do not follow current thought: e.g., the use of Lycopodium instead of the separate genera Lycopodium, Dip! strum, Hu- perzia, Lycopodiella, and Pseudolycopodiella (as cited by Flora of North America Editorial Committee 1993) or Habenaria as opposed to the currently widely accepted Platanthera. Three important references were not chosen as standards for nomenclature, although they are often cited in the NEPCoP List. Merritt Lyndon Fernald’s Gray’s Manual of Botany (Fernald 1950), although still used by many field botanists because of its thoroughness, is not current in nomenclature, taxonomy, or de- 242 Rhodora [Vol. 98 scriptions of plant distribution. The Flora of New England (Sey- mour 1969) was derived primarily from the study of herbarium specimens, and although it is extremely helpful in visualizing plant distribution, it essentially follows the taxonomic treatments found in Gray’s Manual of Botany (Fernald 1950). A Synony- mized Checklist of the Vascular Flora of the United States, Can- ada, and Greenland (Kartesz 1994) had not been published at the time this project was started. Moreover, most of the intended au- dience for the NEPCoP List is not likely to have easy access to this work, which does not contain keys to aid in field identifi- cation. Fortunately, many of the names used in the NEPCoP List are the same as in Kartesz (1994), which is an especially valuable reference because of its nomenclatural accuracy. To increase the utility of *‘Flora Conservanda: New England,” identification manuals and widely reviewed regional floristic treatments are cited as the source of either the primary name or synonym in order to facilitate field identification. Either the pri- mary name or its synonym(s) usually can be found in at least one manual that contains a key. Each entry includes preferred syn- onyms used by one or more of the six New England states. The synonymy is not intended to be complete. An Index to all names and synonyms used in the NEPCoP List follows the Appendices. FORMAT OF THE NEPCOP LIST Divisions of the List. ‘‘Flora Conservanda: New England” is divided into five divisions: Division 1: Globally Rare Taxa occurring in New England. Taxa included in this Division are listed as Globally Rare (G1 through G3 or T1 through T3) by The Nature Conservancy (adapted from Master 1991 and The Nature Conservancy 1996; see Global Rank—GRank—explanations under Notes below or in Appendix II). Usually only a few occurrences of these taxa exist within our region, but New England does contain the ma- jority of occurrences for a few of these highly ranked taxa. In some cases, taxa with GRanks that normally would place them in this division have taxonomic or other issues that make their current status in New England unclear, and the majority of these taxa have been placed in other divisions. GRanks for taxa in this division appear in the Notes section under each taxon in the list. Division 2: Regionally Rare Taxa. These taxa have fewer than 1996] Brumback and Mehrhoff, et al—NEPCoP 243 20 current occurrences (seen since 1970) within New England. This division includes taxa which are rare throughout their range in all of New England as well as taxa that reach the edge of their distributional range in our region. It is important to conserve these edge-of-range occurrences as part of New England’s natural heritage as well as to avoid further shrinkage of these species’ entire ranges. A taxon with more than 20 occurrences in New England might also be included in Division 2 if a substantial number of occurrences contain small numbers of individuals making them more vulnerable to extirpation. These taxa are de- noted as 2(a). All taxa in Division 2 have GRanks of G4 or G5 (see Appendix II for definitions). Division 3: Locally Rare Taxa. These taxa may be common in part of New England, but have one or more occurrences of bio- logical, ecological, or possible genetic significance. For this di- vision, only selected occurrences in a particular state are listed, not the entire taxon’s occurrences throughout New England. A taxon may be listed as Division 3 in one or more states (desig- nated by an *), but is not considered to be regionally rare. An occurrence could be designated as Division 3 in a state if: 1. The occurrence is disjunct to such a degree that genetic isolation is likely (1.e., separated from other populations by more than 50 miles). . An occurrence represents an ecological anomaly for the tax- on within the New England region (for example, an acid bog occurrence of a species that normally grows under cal- careous conditions). . A significant number of a taxon’s occurrences have demon- strably declined within the state (in which case the entire State’s occurrences are considered to be in Division 3). N (oS) Note: The current distribution immediately outside New Eng- land, i.e., New York state and the Canadian provinces of Quebec and New Brunswick, was also considered in determination of disjunction. For example, an occurrence of a taxon in northern Maine that is disjunct from southern New England occurrences would not qualify for Division 3 if it was within 50 miles of an occurrence in New Brunswick or Quebec. Division 4: Historic Taxa. Taxa that once existed in New Eng- land, but that have not been seen since 1970. The purpose of this division is to generate interest in re-locating these taxa if they 244 Rhodora [Vol. 98 still exist and to illustrate the level at which species have been lost from the region. Division Indeterminate (IND.): Indeterminate Taxa. These taxa are under review for inclusion in one of the above divisions, but issues of taxonomy (at least for New England occurrences), no- menclature, or status in the wild are not clearly understood. The purpose of this division is to stimulate interest in taxonomic re- search and/or field surveys for these taxa. Structure of the List. The NEPCoP List is divided into 8 columns. Taxa are listed alphabetically by family, alphabetically by genus within each family, and alphabetically by species within each genus. Column 1 contains the name of the taxon, the author, and source of the name (number in parentheses). Synonyms are listed in italics below the taxon with the source of the name (number in parentheses) and the state using the synonym [in brackets]. For example: ALISMATACEAE Echinodorus tenellus (Martius) Buchenau (11) Echinodorus parvulus (15) [MA] Echinodorus tenellus var. parvulus (14) [CT] Column 2, with the heading DIV, contains the NEPCoP Divi- sion. This may be 1,2,3*,4, or IND. (see Divisions of the List above). Columns 3—8 contain State Data (next six columns under ab- breviated names of the states). Under each state are three blocks separated by vertical bars. If no data are present in any of the three blocks the taxon is not known to occur in that state. An asterisk is used when a state has occurrences of a taxon listed in Division 3. The first block contains the number of currently extant (seen since 1970) occurrences of the taxon in that state. If the number of occurrences is more than 20, but not precisely known, a “*+” is placed in this block. Typically, taxa with a ‘‘+”’ are considered common in the state and thus are not tracked by the NHP. Oc- currences discovered or known prior to 1970, but not verified since 1970, are not considered current (but could possibly still be extant). If the taxon once was native in a state, but is not currently 1996] Brumback and Mehrhoff, et al—NEPCoP 245 considered extant (not seen since 1970), it is designated with The second block contains the official State endangerment sta- tus of the taxon (Endangered, Threatened, etc.). Depending on the state, this status may have a legal designation. Since the same code may have different meanings in different states, refer to the State Status Codes in Appendix I for the definition of these terms in each state. Please note that the codes used by some states have been modified in this List for consistency and clarity. The third block contains the State Rank (SRank) as defined by The Nature Conservancy (adapted from Master 1991 and The Nature Conservancy 1996). This generalized ranking is based on the number of individuals, number of occurrences, and other fac- tors contributing to the vulnerability of a taxon within each state. The SRank codes used in this List are cited in Appendix III, but the most commonly used codes are: Sl = generally 1—5 occurrences in the state. S2 = generally 6—20 occurrences in the state. S3 = generally 21—100 occurrences in the state. S4 = generally 101—1000 occurrences in the state. SE = an exotic (non-native) species in the state. SH = Historic—occurred historically (as a native species) in the state, but is not currently known to be extant in the state. SU = State Unrankable—the status of the taxon is not known. In many instances where a taxon is ranked “SU,” the number of occurrences of the taxon (if any) is not known. SX = taxon is presumed extirpated in the state. Notes under a taxon. Explanatory notes and additional infor- mation are added where necessary beneath the state data blocks. Included in this section is the taxon’s Global Ranking or GRank. A species is given a Global Rank identified by a G followed by a number or symbol, and a subspecies or variety has a T followed by a number or symbol. (For example, Eupatorium leucolepis var. novae-angliae has a GRank of GS5T1, which means that the spe- cies is secure globally, G5, but that the variety is critically im- periled globally, T1). In this List, GRanks are given only for those taxa with a GRank containing Gl, G2, G3 or T1, T2, T3 (ora combination thereof). If no GRank is given, the taxon has a 246 Rhodora [Vol. 98 GRank of G4 or G5. See Appendix I for a complete list of GRank codes used in this List. Most commonly used ranks are: Gl = Critically imperiled globally (typically 5 or fewer occurrences globally). 2 = Imperiled globally (typically 6 to 20 occurrences globally). G3 = rare or uncommon but not imperiled globally (typ- ically 21 to 100 occurrences globally). G#G# = Numeric range rank: A range spanning two or more of the numeric ranks. Denotes range of uncertainty about the exact rarity (for example—G2G3). G? = Unranked, Element is not yet ranked globally. G#T# = for infraspecific taxa: The GRank applies to the full species; T = Taxonomic subdivision and the rank applies to the subspecies or variety. T1, or T2, or T3—same definitions as G1, G2, G3, but refers to a subspecies or variety. T#T# = Numeric range rank: A range spanning two or more of the numeric ranks for a variety or subspecies. Denotes range of uncertainty about the exact rarity of variety or subspecies (for example—GS5T2T3). ? = Inexact or uncertain (for example G3? or G5T3? means that the numeric ranking is uncertain). Questionable taxonomy: taxonomic status is ques- tionable; numeric rank may change with taxonomy (for example, G4T3Q means that the taxonomy, in this case of the subspecies or variety, 1s question- able). eo) | Also contained in the Notes section are codes used by the U.S. Fish and Wildlife Service (USFWS) under the provisions of the U.S. Endangered Species Act of 1973 (the Act), as amended. If a taxon is listed as Endangered or Threatened under the Act, LE (Listed Endangered) or LT (Listed Threatened) will appear in this column. Other designations include C2 for taxa that formerly were considered as candidate species for listing under the Act. This category has been discontinued under a notice of final de- cision published in 1996 (U.S. Fish and Wildlife Service 1996). Also included are codes for taxa no longer under consideration by the USFWS. These designations (3A, 3B, and 3C) have been 1996] Brumback and Mehrhoff, et al—NEPCoP 247 discontinued also. See Appendix IV for a complete listing of Federal codes used in this List. DISCUSSION Lists of this nature frequently point to the need for additional work. The process of compiling “Flora Conservanda: New Eng- land”? demonstrated to the Listing Committee the obvious need for protection and management for many taxa and their occur- rences. The process also emphasized the need for additional field work to gather data on occurrence sizes and distributions. Un- fortunately there are too few knowledgeable field botanists to cover the full extent of rare plants in New England. Volunteers must be sought and trained to help with this task. Basic botanical inventory is essential for an accurate understanding of the true rarity of listed taxa as well as for interpreting population trends over time. Two areas where additional work is needed became apparent as the Listing Committee worked on ‘Flora Conservanda: New England.” First, many taxonomic questions concerning the New England flora remain unanswered; many of these center on infra- specific taxa. Entities observable in the field (and often named by New England’s most famous student of its flora—Merritt L. Fernald) should be studied using current tools and methodologies in order to resolve taxonomic issues. For example, is Eupatorium perfoliatum var. Coe ane a ‘““good” variety or an ecomorph? Is Cardamine longii a ““good”’ species? A particularly perplexing taxonomic issue involves Bidens heterodoxa. Cronquist (Gleason and Cronquist 1991) mentions this taxon (under B. connata), but does not include it as a distinct species as he does for B. eatonii or B. hyperborea (which are present on the NEPCoP List). He states that B. heterodoxa consists of a series of rare and local populations. Should B. heterodoxa be afforded the same protec- tion as listed taxa? The resolution of these questions is not merely an academic exercise but helps to assure the best use of limited conservation resources. imilarly, hybrids and the hybrid nature of some taxa need clarification. Is a hybrid sterile or fertile? Do both sterile and fertile hybrids deserve protection? Some scientists argue for pro- tection of sterile hybrids because of the possibility of ploidy shift that will allow a polyploid to become fertile. In the NEPCoP List 248 Rhodora [Vol. 98 we have included hybrid taxa considered to be nothotaxa (a no- menclatural term that defines species of hybrid origin whose names include all the offspring including backcrosses). We also have listed sterile hybrids in some instances in order to raise the issue of their protection. Equally important is the need for nomenclatural study. Often, the Committee was faced with a choice of names for a given taxon. Frequently exacerbating this problem was the issue of un- clear synonymy. Moreover, different manuals and reference works sometimes use different names for what appears to be the same taxon. Occasionally, both taxonomy and nomenclature were unclear. For example, the taxon called Puccinellia tenella ssp. alascana is an extreme example of a taxon that was placed in the Division IND. (Indeterminate) because of unclear nomenclature and tax- onomic circumscriptions. Equally confusing are some taxa in the genera Panicum and Viola. The Listing Committee hopes that by focusing attention on these issues, Clarification by the scientific community will follow. Ideally, many of the issues regarding field status, taxonomy, and nomenclature that are highlighted by the publication of “Flora Conservanda: New England’’ will have been addressed by sci- entists before the next iteration of the List, anticipated in the year . The ongoing publication of Flora of North America also will provide guidance. The NEPCoP List contains 576 taxa in the following divisions: Div. 1—Globally rare = 57 taxa; Div. 2—Regionally rare = 273 taxa; Div. 3—Locally rare = 75 taxa; Div. 4—Historic = 55 taxa; and Div. IND.—Indeterminate = 116 taxa. In comparison, the NEBC regional list (Crow et al. 1981) in- cluded 479 taxa, the majority of which were designated *“‘E/T”’ (Endangered/Threatened—337 taxa) or ““R’ (Rare—133 taxa). Taxa designated as ‘E/T’? were documented from 10 or fewer towns and “*R”’? were documented from 10 to 20 towns. Because different criteria were used to create the 1981 NEBC list and the NEPCoP List, comparisons are difficult. The two major differ- ences are: 1) The NEBC list used frowns as a criterion in contrast with current occurrences in the NEPCoP List. Theoretically, a taxon could appear on the NEBC list because it was found in fewer than 20 towns in New England, but since there could be more 1996] Brumback and Mehrhoff, et al—NHEPCoP 249 than one occurrence in a town it theoretically might not appear on the NEPCoP List in Division | or Division 2. The taxon could, however, still appear in another NEPCoP division (i.e., Division 3 or Division IND.). 2) The NEPCoP List uses current occurrence data. Since 1970 was used as the cutoff date for current occurrences in the NEPCoP List, only occurrences actually verified since that date have been included. In contrast, the NEBC list used herbarium specimens as the primary source for deriving the list (Crow et al. 1981). Although some field investigations were conducted, many of the specimens which counted towards the inclusion (or exclu- sion) of a taxon in the NEBC list were collected prior to 1970. These historic (by NEPCoP standards) occurrences were not con- sidered as currently extant in the NEPCoP evaluation of the re- gion’s rare flora. These occurrences either have been re-located (and thus are considered current) or considered not to be extant. Discounting synonymy, as well as taxonomic and nomencla- tural changes, there are 22] taxa on the NEPCoP List that did not appear on the NEBC list. Seventy-three of these appear in Division 1 or Division 2, indicating that these taxa currently ap- pear to be more rare than previously thought. Seventy-eight taxa on the NEPCoP List as Division IND. (Indeterminate) did not appear on the NEBC list. This shows, perhaps, that as our knowl- edge about the rarity of the New England flora has increased, so has the realization that there are many taxa whose field status or taxonomy (at least in New England) is still unclear. Also, there are 14 taxa on the NEPCoP List in Division 4 as Historic in New England that did not appear on the NEBC list. Their appearance on the NEPCoP List is partly a function of the 1970 cutoff date, but also reflects our increased knowledge of the flora through field work of the state Heritage Programs, The Nature Conser- vancy, NEPCoP, NEBC, and other organizations. On the other hand, discounting synonymy as well as nomenclatural and taxo- nomic changes, there are 38 taxa on the NEBC list that do not appear on the NEPCoP List. Considered more common by NEPCoP standards, their exclusion is again the result of increased field work and data collection. RECOMMENDATIONS Because of differences in the criteria used to create the NEPCoP and NEBC lists, we cannot state with certainty that rare 250 Rhodora [Vol. 98 plant species have declined in New England in the interim be- tween the publication of the two lists. While comparisons are difficult, anecdotal information and field observations suggest a decline in the number of current occurrences for many taxa. This trend is substantiated by a number of occurrences that have not been re-located despite intensive field searches. The publication of “Flora Conservanda: New England’’ pro- vides baseline data to judge the future status of rare species. In order to protect New England’s flora the cooperation of many individuals and organizations will be necessary. The NEPCoP List is a good example of such cooperation that combines insti- tutional resources and interests to further conservation goals. NEPCoP recommends that the rare taxa (or occurrences, in the case of Division 3 taxa) be monitored at regular intervals as part of a regional conservation plan. Such a plan would also include habitat protection, management of existing occurrences, seed banking, and enhancement, reintroduction, or introduction if deemed necessary. Field investigations and/or taxonomic study of Indeterminate taxa (Division IND.) are needed to determine the conservation status of more than 100 taxa in this Division. It is hoped that the NEPCoP List will focus research attention on these problematic taxa. NAME ACANTHACEAE Justicia americana (L.) M.Vahl (14) ADIANTACEAE Adiantum aleuticum (Ruprecht) Paris (12) Adiantum viridimontanum Paris (12) ALISMATACEAE Echinodorus tenellus (Martius) Buchenau (11) Echinodorus parvulus (15) [MA] Echinodorus tenellus var. parvulus [CT] (14 Sagittaria rigida Pursh (14) FLORA CONSERVANDA: NEW ENGLAND ME NH VT MA HHI] HII |] [SH] HHI IJE|S1| HII [3] [SV HT HII HT [7|T|S2| HII GRank = Gl. HII HII HII |Hi|Sx| GRank = G3. GRank is based on synonym, £. parvulus. [3|E|S1|* || [SU [+] |S3| [6+|WL|S2| Disjunct occurrences in Sagadahoc County, Maine. RI HII HHH || |SU| HIT Hi [HE|S 1 || SU] dOOdAN— Ik 10 ‘Jyoyryapy pure yorquinig [9661 [Se NAME Sagittaria subulata (L.) Buchenau (14) Sagittaria teres S. Watson (14) AMARANTHACEA Amaranthus pumilus a (14) Amaranthus tuberculatus (Moq.) Sauer (14) APIACEAE Angelica lucida L. (14) Coelopleurum lucidum (11) [CT] Angelica venenosa (Greenway) Fern. (14 Hydrocotyle verticillata Thunb. (14) Lilaeopsis chinensis (L.) Kuntze (14) ME NH VT MA RI HH Ht HH 2|E|S1| HH I HI HT [54|SC|S3]_—3|E|S1| GRank = G3. ae {| IT | |SH| |H|SH|SH| GRank = G2; Fed. code = LT. | | ISE| | | [SU] 16] |S2| [| |SE| HII cy [14|SC|S3] |HISC *|SH| | | ISE| Considered adventive in most of New England, but apparently native in Vermont and perhaps New Hampshire [?| |SU| [?| [SU LH [9|WL|S2| Sal More field work needed. Not currently tracked in northern state HII HII HII |H] |SX| HH HII HI III] [14|SC|S2| |HJH|SH| [5|T|S1|* |6|T|S2| | [SIWLIS2| | ITI Disjunct occurrences in Sagadahoc and York Counties, Maine. 3]E|S1| [| SUI |H] |SH| [10|SC|S3} N tN vIOpouYy 86 TOA] NAME Div ME NH VT MA RI ct Osmorhiza chilensis Hook. & 2 H1+{T|S2} — |HJE|SH \H] [SHI nT HL nn . (14) Osmorhiza berteroi (1) [ME] Osmorhiza depauperata Philippi (14) a I] || HII |H]||SH} It || III lt || Osmorhiza obtusa (11) [VT] Sanicula canadensis L. (14) 2 {|| 1H] |SH| |9|T|S2| |8|T|S2| III | | [SUI Taenidia integerrima (L.) Drude 2 THI It 1| |7|T|S2| |H?| |S?| |H|H|SH| W|SC|ST] (14) Reported in 1913 from Massachusetts, but no specimen has been seen. Zizia aptera (A. Gray) Fern. (14) 2 It] II Tk It ann |H|H|SH| |3|E|S1| AQUIFOLIACEAE Ilex ambigua Torr. var. montana Z III III I II | |3|T|S2| II || HII Ahles (13 (13) Ilex montana (11) [MA] Ilex glabra (L.) A. Gray (14) ar JLJE|S1|* |H|E|SH| I |+| |S4| |+| |S3| |3|T|S1| One disjunct occurrence in Knox County, Maine. ARISTOLOCHIACEAE Aristolochia serpentaria L. (14) 2 III I ITI II HII |6|T|S2| [9661 ‘JJOUIYSIAT pue Yovquin4sg d9DdAN— TP 39 CSC NAME ASCLEPIADACEAE Asclepias purpurascens L. (14) Asclepias tuberosa L. (14) Asclepias variegata L. (14) Asclepias viridiflora Raf. (14) ASPLENIACEAE Asplenium montanum Willd. (11) Asplenium trichomanes-ramosum L. (12 Asplenium viride (14) [VT] ASTERACEAE Achillea borealis Bong. (11) Achillea millefolium var. nigrescens (14)[ME] Achillea millefolium var. borealis (15) [MA] IND. ME NH VT MA RI CT It |H] |SH] ITT [2|T|S1| |H] |SH| |H] |SH| (H] [SX INJEISH] = HITISH] = HJWELIS3]* — [8/C|S2| I+] |S4| Documented decline of native stands in Massachusetts and possibly other states. May be subject to overcollection in some areas. HN HH HI [1 HH HEIS1| I HII HT | | |H|SC *|SH| HH HT IIT |S I 3|E|S1| JIJE|S1| }6|T|S2| JIJE|S1| HI 4[T|S1| HIT HH HH 1 2|SH| [1] |SU| HH I|| SE?) HT HII Considered introduced in Massachusetts, but not currently known to be extant. Taxonomic and nomenclatural confusion with certain varieties of A. millefolium. N eLopoyy 86 1OA] NAME Arnica lanceolata Nutt. (14) Arnica mollis (11) [VT] Artemisia campestri sb. ssp. borealis (Pallas) Hall & Clem. (1) rtemisia campestris var. canadensis (14 Artemisia campestris L. ssp. caudata (Michx.) Hall & Clem. (14) Aster anticostensis Fern. (14) Aster concolor L. (14) Aster dumosus L. (14) Aster infirmus Michx. (14) Aster praealtus Poiret (14) ME NH VT MA RI CT |<6| |S2| [2/T|S1| |HI|SH) HII II II GRank = G3 |H| |SH| HI [3] |S1] [LES] | | [+] [SUI [8|T|S2| [2] |S1|* +] |S4| [3|C|S1| [+] |S4| Disjunct occurrences in Grand Isle County, Vermont |H] |SX| HII IIT LIT HII Ht GRank = G2Q. Cronquist (Gleason and Cronquist 1991) mentions this taxon, but does not formally include it in his treatment. The northeastern Maine locality is historic. HII HH man [9|E|S2| |H|SH|SH| | III 2|E|S1|* || |SU| HII [+] |S4| [+] |S4] [+] |S5| Disjunct occurrences in York and Oxford Counties, Maine. HI HH HI S]E|S1| IH|SHISH] || {SH |H?| |SU| [2] [SU HII |H] [Sx] |H] [SH || |SU| anon ° pai from other closely related taxa; more field work ne [9661 dOOdAN— IP 19° ‘JJoyuryapy pure yoequinig NAME Aster prenanthoides Muhl. (14) Aster sagittifolius Willd. (14) Bidens eatonii Fern. (14) Bidens heterodoxa (Fern.) Fern. & St. John (11) Bidens hyperborea Greene (15) Bidens hyperborea var colpophila (11) [MA] Bidens ee ealielae Greene var. svensonil Fassett (1) Bidens a var. cathancensis (11) Cacalia suaveolens L. (14) Synosma suaveolens (15) [MA] Chrysopsis mariana (L.) Elliott (14) ME NH VT MA RI i Ith HII LI BISCiS2; | II |H|SC *|SH| HT HII [| [S1| HHI HII HII [5|T|S]| nan II [2|T|S2?| HII [3|/SC|S1| GRank = G2G3 HIT HT nan | Ht EI] |SH| || GRank = G2Q. Cronquist (Gleason and Cronquist 1991) says that the proper taxonomic status is uncertain. JLVTISIS2} ||] | HII 2|E|S1| Ht Ht I?| |SU| Ht HII HII Ht an HH HII HI II|SE| aoe |H|SC *|SH| GRank = G3. GRank is for synonym, Synosma suaveolen III HII HII III! [2/T|S1| HII PIOPOYY 86 ‘19A] NAME Cirsium horridulum Michx. (14) Coreopsis rosea Nutt. (14) Erigeron acris L. var. kamtschaticus (DC.) Herder (14) rimorpha acris var. kamtschatica (1) [ME] Eupatorium album L. (14) Eupatorium aromaticum L. Eupatorium leucolepis (DC.) T. & G. var. novae-angliae Fern. (14) Eupatorium perfoliatum L. var. colpophilum Fern. & Grisc. (14) Eupatorium rotundifolium L. var. rotundifolium (14) Eupatorium sessilifolium L. (14) ME NH VT MA RI CL HII |2|E|S1| HII [8[WL|S2S3] |1|T|S1| [3] [SI More field work needed to clarify status in our region, especially on the islands off Massachusetts. HT HT HI +] |S3| [7|C|S2| [1 GRank = G3. |H||SH| HT HI HT HT | HT HT HT HT HIT 2|E|S1| HT HI 1 2|E|S1| |H|SH|SH]—[2|E|S1| HI IIT It S|E|S2| [6|E|S 1 Ht GRank = GST1; Fed. code = C2. 12] [S1| III HII ITI [111 HII Taxonomic and distributional status of this variety in New England is unclear. IIT HII HHI HII HII [2| [SUI Difficult to distinguish from closely related taxa. More field work needed to assess current status. HII [IES 1| S|E|S1|* +] [$4] [+] |S3| +] |S3| Vermont occurrences in Rutland County are disjunct. dODdAN—'TP 19 ‘Joyrys| puke yorquinig [9661 LSC NAME Euthamia galetorum Greene (14) Euthamia tenuifolia var. pycnocephala (1) [ME] Gnaphalium helleri Britton (14) Gnaphalium helleri var. micradenium (15) [MA,ME] Gnaphalium purpureum L. (14) Gamochaeta purpurea (1) Gnaphalium supinum L. (14) Omalotheca supina (15) [ME] Gnaphalium sylvaticum L. (14 Omalotheca sylvatica (15) ei Hieracium robinsonii (Zahn) ern. (14) Hieracium umbellatum L. (14) Iva frutescens L. var. orari (Bartlett) Fern. & Griscom (14) Iva frutescens ssp. oraria (1) {CT,MA,ME,NH,RI] Div IND. IND. IND. RI HH ME NH VT MA |19\|SR| || [SU| HII HII GRank = G3Q. Presence in New England is questionable. [?\|SU] [| |SU| HT |H||SH| aa cr HT Gleason and Cronquist (1991) note that this species is in New England, but we have not seen any specimens. |H||SX| III LHI [2?|E|S1| [2|E|S1| |VJE|S1| It IVI |4+||SU| [1] [SUI [A|E|S 1 II More field work needed to determine current status. ae [IJE|S1| II IT = G1G2; Fed. code = C2 I{ 1 HIJE|S1| HHI II 1 [3|T|S1|* |7+|T|S2| LI TI }+||S5| | Disjunct occurrences in Sagadahoc and Cumberland Counties, Maine. |H|SH|SH| HII HII |+||S3 |H] |SH| HI] HT HIT HT [+||S4| vlopoyy 86 1OA] NAME Krigia biflora (Walter) S. F. Blake Lactuca hirsuta Muhl. (14) Lactuca hirsuta var. sanguinea (11) [CT,MA,ME,NH] Liatris scariosa (L.) Willd. var. novae-angliae Lunell (14) Liatris borealis (11) [CT,NH] Pityopsis falcata (Pursh) Nutt. (15) Chrysopsis falcata (14) [CT,RY] Polymnia canadensis L. (14) Prenanthes boottii (DC.) A. Gray (14) Prenanthes x mainensis A. Gray (11) Prenanthes racemosa Michx. (14) Prenanthes serpentaria Pursh (14) ME NH VT MA RI CT HI HI HT Ht I |H] [SH] [HSU] II|SU| 8 T|S2|* | WL|SU} [| ISU] [H||SH| Disjunct occurrences in Chittenden County, Vermont. More field work needed to determine current range in New England. |4|T|S 1} |6|E|S 1} ann |33|SC|S3| |4|T|S1| [L1|SC|S2S3] GRank = G5?T3; Fed. code = C2. Former Federal candidate Category 2 status is for synonym L. borealis. [1 1 GRank = G3G4. Ht HT [3/T]S1| |4[T|S1| 2|E|S]| ITT I} 1 | ITI GRank = G2; Fed. code = C2. LIT [+] |S3S4]—[8|C|S2| 3|E|S1| 2|E|S]| | HII [EIS 1 || ISU HI nan HII HII HII Possibly seen recently, but verification needed. Gleason and Cronquist (1991) note that this is an apparent hybrid of P. racemosa with P. trifoliolata. [15 |S2| I} TI HII IH III LTH Cronquist (Gleason and Cronquist 1991) suggests that our plants are var. multiflora. |H|SH|SH| |3| |S1| HT |H] [SH] HI IS|E|S1| dODdHN—T® 19 ‘Jyourys] pue youquinig [9661 6ST NAME Sclerolepis uniflora (Walter) BSP. 4) Solidago x calcicola Fern. (13) Solidago calcicola (14) [ME,NH] Solidago poe L. var. subserrata (DC.) Cronq. (1) Solidago lepida var. ae (11) Solidago cutleri Fern. (14) Solidago multiradiata var. arctica (1) [ME] Solidago ptarmicoides (Nees) B. Boivin (14) Aster ptarmicoides (11) [NH, VT] Solidago rigida L. (11) IND. N ME NH VT MA RI CT III HE|S1| IT [AES 1] [IES 1 III Massachusetts and Rhode Island occurrences cross state boundaries and represent the same population. |HI|SH| |H||SH| [III Itt Ht Ht Hybrid between S. macrophylla and another species, possibly 5. canadensis. I?| SHI HI HIT HHI IT Hit Solidago canadensis var. subserrata is authored by Cronquist but does not appear in Gleason and Cronquist (1991). Fernald (1950) lists this taxon from Maine as S. /epida var. molina. State rank of "SH" is based on a 20-year cutoff date by the Maine Natural Areas Program. |6||S1S2| 9|T|S3| 1] [SI Hh HT HII I 2|E|S1| [11|S2S3|— |AITISIS2) | TT JIJE[S1| HI Hl I |H| |SX| || |SH| [4|E|S 1 RIOPOYY 86 ISA] NAME Solidago simplex HBK. ssp. randii (Porter) Ringius var. monticola (Porter) Ringius (14) Solidago simplex var. randii (1) [ME] Solidago glutinosa ssp. randii (31) [MA,NH]} Tanacetum bipinnatum (L.) Schultz-Bip. ssp. huronense (Nutt.) Breitung (1) Tanacetum huronense (14) Taraxacum ceratophorum (Ledeb.) DC. (14) Taraxacum latilobum (11) [ME] BERBERIDACEAE Podophyllum peltatum L. (14) 2(a) IND. ME NH VT MA RI cr |+||S4| [+||S4]* HII [4|E|S 1|* ITI II Disjunct at sites in southern Berkshire County in Massachusetts and in Cheshire County in New Hampshire. According to Gleason and Cronquist (1991), there are three varieties of the subspecies. The current distribution and status of New England taxa are unclear. [30| |S2| HII III [III HII II Restricted to St. John River. Small population sizes and ephemeral nature of the habitat are cause for concern. |H?||SU| HH HT I Hit TN Two old specimens of 7. /atilobum are unverified. || ISE| [12] |SU| [2| [S| | ISE| || ISE| | SUI This species has been introduced into all New England states. Although probably native in some states (Connecticut, New Hampshire, and Vermont), determining if an occurrence is native or introduced is often difficult. It is considered native at some locations in New York state. [9661 dOOdAN— TP 19 ‘ours pur yorquinig 197 NAME BETULACEAE Betula glandulosa Michx. (14) Betula nana (1) [M] Betula minor (Tuckerm.) Fern. (13) Betula borealis (11) [VT] Betula x minor (1) [ME] Betula nigra L. (14) Betula pumila L. (14) BORAGINACEAE Cynoglossum virginian mL. var. boreale (Fern.) coape aes (14) Cynoglossum boreale (11) [MA,NH, VT Cynoglossum virginianum (15) [CT] Cynoglossum virginianum L. var. virginianum (14 Cynoglossum virginianum (15) [CT] ME NH VT MA RI CT IJE|S1 JV 1}T|S1| HT I HII HH [IIE ah [9| |S2| |H||SHY HH [1 HT | | GRank = G3G4Q. Gleason and Cronquist (1991) note this as a hybrid of B. aaa var. glandulosa with a dwarf, alpine form of B. papyrifera, perhaps locally alloploid. HII |6/T|S2| HIT [S?7WLIST] {TI | | |SU| Native and introduced populations occur in New England; it is often difficult to determine which occurrences are native and which are introduced. [+] |S3] [NE|S1]* [IE|S1| |4|T|S2| an Ecological anomaly at a New Hampshire acidic fen. [/SC|S2| I?| [SH] [IE[S1| |2/T|S1| |H||SX| HI |H|SC *|SH| GRank = G5ST3? The Connecticut endangered species list includes var. boreale and var. virginianum under C. virginianum. IT HH HH HII HII |H|SC *|SH| The Connecticut endangered species list includes var. virginianum and var. boreale under C. virginianum. vIOPOUY 86 1OA] NAME Hackelia deflexa (Wahlenb. : a Gray) Fe Hackelia americana (40) [VT] Mertensia maritima (L.) S. F. Gray (14) Onosmodium virginianum (L.) (14) A.DC BRASSICACEAE Arabis drummondii A. Gray (14) Arabis laevigata (Muhl.) Poiret (14) Arabis missouriensis Greene (14) Barbarea orthoceras Ledeb. (14) Braya humilis B. L. Robinson (14) ME NH VT MA RI [NE|S 1 [EIS 1| |16|T|S2| I I I+] |S3S4| | IT || J6|E|S1|* HII Disjunct in Barnstable and Nantucket Counties, Massachusetts. Ht HII Itt [H] [S| |H|SH|SH| |+| |S4| |+?| |S3?| |2|E|S1|* J2|WLIS 1 |H|SH|SH| Vermont occurrences in Rutland and Addison Counties are disjunct. 2|E|S1|* |H] |S1| [+] |S4| [LO/T|S2| [| SU Disjunct in Franklin, Aroostook, and Piscataquis Counties in Maine. |4|T|S 1] [3|TIS1S2| |2| |S1| |9|T|S2| || |SU| Taxonomic question and difficulty in distinguishing this taxon from A. /aevigata HT HII [HES || |SU| 1+] |S3| | | |SU| into which one author (see Mulligan 1995) has recently placed this taxon. |H] [SH] [IE|S1 mn HT HI HT [1 [2/T|S 1 IH HI [11 mat dODdAN— TP 39 “JJOUTYSIA, pue Yorquinig [9661 9G NAME Cardamine bellidifolia L. (14) Cardamine concatenata (Michx.) O. Schwarz (14 Dentaria laciniata (11) [CT,MA,NH,VT] Cardamine douglassii Britt. (11) Cardamine x incisa K. Schum. (pro sp.) (15 Dentaria x incisifolia (11) [MA] Cardamine longii Fern. (14) Cardamine x maxima (Nutt.) A. Wood (14) Cardamine maxima (15) [ME] Dentaria maxima (14) [CT,MA,NH] Cardamine pratensis L. var. palustris Wimmer & Graebner (14) IND. ME NH VT MA RI IE|S1| [3|E|S1| HT LT HT [HE|S1|* 2|E|S1| [+||S3| +||S3| I Aroostook County, Maine, occurrence is disjunct. HT HT HI HES 1| HT Ht HI HT II|SU] HH Fernald (1950) suggests that this is a hybrid 7 Dentaria laciniata (C. concatenata) and Cronquist (1991) mention and Dentaria maxima (C. x maxima). Gleas CT III [+||S3] |7|SC|S2| [2| [SU this taxon under C. angustata (as D. ae but do not formally include this taxon in their treatment. le, |H|T|SH| HII 2|E|S1| [HE|S1 = G3Q; Fed. code = 3C. |Hj|SH| |H||SH| HII Taxonomic Status unclear a sterile hybrid between C. diphylla and C. concatenata. 2|WLIST] {IT HH IEIS1| [3] |S1| [3|T|S1| HI Hj |SH} |H||SH| . Gleason and Cronquist (1991) suggest this as possibly | SU] P9C vlOopoyy 86 “1OA] NAME Descurainia pinnata (Walter) Fern. Descurainia richardsonii (Sweet) O. E. Schulz (14) Descurainia incana (15) [ME] Draba arabisans Michx. (14) Draba cana Rydb. (14) Draba lanceolata (11) {ME,NH,VT] Draba glabella Pursh (14) Draba reptans (Lam.) Fern. (14) Neobeckia aquatica (Eaton) Greene (18) Armoracia lacustris (14) »VT [MA,ME, VT] Subularia aquatica L. (14) Britton var. brachycarpa (Richardson) (14) ME NH VT MA RI CE HI [Hy |SX| [2| |S1| | | ISE| HT HH |H||SH] HT [11 HT I HT [3|T|S1| HT [10] |S2S3] {ITI | LT Tracking of this species was only recently begun in Vermont. It may be more abundant than is now known (which is reflected in the $2S3 ranking). IE|S1| IE|S1| [3/T|S1| nan Hh HH HT HI [5] |S1| HII HII HI HT [1 HII |H] |SX| |H|SH|SH] —|4/SC|S2| |H?|SH7] TTI [4[T|S1| |H\|SH?| HT I Type locality for this species is in western Massachusetts, but no specimens have been seen from Massachusetts or Maine. I9| |S2| || [SU |H] [SH] HI HT HT dODdAN— TP 19 ‘ourysp] pur yorquinig [9661 S9C NAME CAESALPINIACEAE Cercis canadensis L. (14) Senna hebecarpa (Fern.) Irwin & Barneby (14) Cassia hebecarpa (11) (CT,NH,VT] CALLITRICHACEAE Callitriche hermaphroditica L. (14) Callitriche terrestris Raf. (14) CAMPANULACEAE Lobelia spicata Lam. var. hirtella A. Gray (14) CAPRIFOLIACEAE Lonicera dioica L. (14) Lonicera hirsuta Eaton (14) ME NH VT MA RI Ht HII HII [ITI HI longer extant there. Only naturalized occurrences remain. HII IH|E|SH| |H|T|SH| 2/E|S1| |IT|S1| IT HT |H] |SH| HT HT Ht HI I |H] |SH| HHT | | {SU || |SH] IIT II I} Current status and distribution of this variety is unclear. [IJE|S1\* || |SU| +] |S4] +1 [S4] 3}C|S1| Cumberland County, Maine, occurrence is disjunct. HT HT |12| |S2| 3|E|S1| I CT. |H|SC *|SH| Literature reports indicate that this was native at one site in Connecticut, but it is no 2/SC|S1| HT |H] [SH LT [+] |S3| C PIOPOYY 86 TAI NAME Lonicera sempervirens L. (14) Symphoricarpos albus (L.) S.F. Blake var. albus (14) Triosteum angustifolium L. (14) Triosteum aurantiacum E. Bickn. (14) Triosteum perfoliatum L. (14) Viburnum nudum L. var. nudum Viburnum nudum (11) [CT,RY] Viburnum prunifolium L. (14) Viburnum rafinesquianum Schultes (14) ME NH VT MA RI 0 | 2|E|S1| | [SU [| ISE| || ISE| [| ISE| || [SU| Difficult to determine which populations are native. Introduced occurrences in Vermont are no longer extant. IT TI IH [+] |S3| |IJE|S1|* HTT III Occurrence in Franklin County, Massachusetts, is disjunct from western Vermont occurrences. | HT HII LT HI |H|SC *|SH| Gleason and Cronquist (1991) distinguish two varieties, but both are historic in New England. |2|T|S1|* [2|E|S 1 [+| |S3| +| |S4| l4{T|S1| [+] [S3| Aroostook County, Maine, occurrence is disjunct. aan ITI I} || |4-S|E|S1 |4|C|S 1| || |SU| IIT | HII II || Ht [AIT|S1| |H|SC *|SH| LIT IIT 111 IT] | II [8|SC|S2| | [S|E|SE|* [+] |S3| |4[T|S2| | | [+] [S3| II | Southern Rockingham County, New Hampshire, occurrences are disjunct. [9661 dODdAN—T® 19 ‘JJourys] pue youquinig LOT NAME CARYOPHYLLACEAE Arenaria caroliniana Walter (14) Minuartia caroliniana (15) Cerastium nutans Raf. (14) Minuartia glabra (Michx.) Mattf. (15) Arenaria glabra (8) (CT,RI] Arenaria groenlandica var. glabra (14) Minuartia groenlandica (Retz.) Ostenf. Arenaria groenlandica var. groenlandica (14) Minuartia marcescens (Fern.) House (15 Arenaria marcescens (11) Minuartia rubella (Wahlenb.) Heirn (15) Arenaria rubella (14) Moehringia macrophylla (Hook.) Fenzl (15 Arenaria macrophylla (14) [CT] 2(a) ME NH VT MA RI cr M1 MI MI MI HHISHISH| || HI MII Pisa] (EISM| TT HIISU is} |S1S2}—aITIS2} IIT HII RIEISI| ——_(7ITIS2 Small population sizes of some occurrences are cause for concern. 24] |S3| [+] |S4| [2] |S1|* HII HII [III High peak occurrences in Chittenden, Lamoille, and Washington Counties in Vermont are disjunct. HT HH [1]T|S1] HI HH HT GRank = G2; Fed. code = C2. Not included in Gleason and Cronquist (1991). [IJE|S1 I [I[T|S1| HI I 1 HHI HHT |9| |S2| |4{T|S1S2| {fT 2|E|S1| 89C elopouy 86 TOA] NAME Paronychia argyrocoma (Michx.) Nutt. (14) Paronychia argyrocoma var. albimontana (11) [NH] Paronychia canadensis (L.) A. Wood (14) Paronychia fastigiata (Raf.) Fern. (14) Sagina decumbens (Ell.) T.&G Sagina nodosa (L.) Fenzl ssp. borealis Crow (15) Sagina nodosa var. borealis (14) Sagina nodosa (L.) Fenzl ssp. nodosa (14) Silene acaulis (L.) Jacq. (13) Silene acaulis var. exscapa (11) [ME,NH] IND. ME NH VT MA RI 6s) [8] |S1S2; |L6|T|S3)_— UAT |IE|S1| HI HH Small population sizes of some occurrences are cause for concern. HIT 7|T|S1| [3] |S1]* [+] |S4| [| ISU| +] |S4 One occurrence on Lake Champlain in Franklin County, Vermont, is disjunct. HII HII | || |SE?| HI || [SU Massachusetts occurrences may be adventive. || III |H] |SH] [IJWLISU] {ITI || {SU More study needed to clarify status. Is the Massachusetts occurrence truly native? [10] [S22] | IT HII [| |SU| HIT HII | | ISE| |H| [SH HHI [3|T|S2| I HII This taxon is considered introduced by most botanists, but there are some questions about the Massachusetts occurrences. Gleason and Cronquist (1991) consider this to be a glandular-hairy variety introduced from Europe. |H||SH| [2|T|S1| HT HI HT | [9661 d°DdHN— TP 19 ‘ours pue youquinig 697 NAME Silene stellata (L.) Aiton f. (14) CERATOPHYLLACEAE Ceratophyllum echinatum A. Gray (14) CHENOPODIACEAE Chenopodium foggii H. A. Wahl (15) Chenopodium pratericola (14) [VT] Chenopodium leptophyllum Nutt. (14) Chenopodium rubrum L. (14) ME NH VT MA RI HH HT |H| |SH] L1 |H|SH|SH| 2| |SH|* 2-3] [SU] [6] |S2| [+] |S3?| [| |SU| DLC Cr [3] |S1| [+] [S3] Occurrences in Penobscot and Waldo Counties, Maine, are disjunct, but were last recorded in 1971 and 1975. State rank of "SH" for Maine is based on a 20-year cutoff date used by the Maine Natural Areas Program. HI I?| |[SU| |H||SH| |H|WLISH] {III GRank = G3?Q. Gleason and Cronquist (1991) place this taxon under [ITI vlopouy C. pratericola and consider it introduced from the west. Massachusetts considers it native and historic. Vermont does not separate C. foggii from C. pratericola (which ts historic there). More field work needed. | [SUI HH HI HII 2|C|S1| HIT | More field study needed to assess status. Most authors consider the eastern U.S. populations of this taxon to be introductions. Some authors have included the taxon under C. pratericola. [4{T|S1|* [1[T]S2| HT [+] |S4] [| |SU| Lincoln and Washington Counties, Maine, occurrences are disjunct. || SU 86 TAI NAME Chenopodium standleyanum Aellen (14) Chenopodium berlanderi var. boscianum (15) [ME] Chenopodium boscianum (11) [NH] Suaeda americana (Pers.) Fern. ) Suaeda calceoliformis (1) [ME] Suaeda maritima (L.) Dumort. (11) Suaeda maritima ssp. richii (1) [(MA,ME] CISTACEAE Helianthemum dumosum (E. Bickn.) Fern. (14) Hudsonia tomentosa Nutt. (14) IND. ME NII VT MA RI Cr [1 ?|)SU| IJE|S2| HI |[WLISE?| {I || |SU| Difficult to distinguish. More field study needed. C. standleyanum is considered native and C. boscianum may be introduced. |H| [SH] HII IH [L2|SC|S3] {TT [ISU [IJE|S1| HII HI [7]WLIS3?|_— | | [SU] || |SU| GRank = G5T3?. | HI I [92|SC|S3|_—_|SJE|S1| |H|SC *|SH| GRank = G3; Fed. code = C2. Massachusetts has the largest number of occurrences globally, but small population sizes and diminishing habitat of many occurrences are cause for concern. |+| |S3S4| [14|T|S1| |4|E|S1|* |+| |S4| |10| |S2| |S|T|S2| ccurrences in Chittenden and Grand Isle Counties, Vermont, as well as New York occurrences on Lake Champlain, are disjunct from the rest of this species' range in New England. L966I dODdAN— I? 19 ‘Jourys|] pur yorquinig LLG NAME Lechea minor L. (14) CLUSIACEAE Hypericum adpressum Barton (14) Hypericum stragulum P. Adams & Robson (14) pericum hypericoides ssp. multicaule (15) [MA] CONVOLVULACEAE Calystegia spithamaea (L.) Pursh (14) Convolvulus spithamaeus (11) [NH] CORNACEAE Cornus florida L. (14) CRASSULACEAE Sedum rosea (L.) Scop. (14) ME NH VT MA RI oi HII HII |H| [SH| [LO[WL|S2?| || [SU | | |SU| Current status in most New England states is unknown. II] III IH |8|T|S2| |S|T|S1| |H|SC *|SH| GRank = G2G3; Fed. code = C2. IT II HT |8|E|S2| III IT [3|T|S1| [2|T|S2| [8|T|S2| [2|E|S1| III |H| |SH| 3|E|S1| [+|SC|S3| [S|T|S1|* [+| |S4| [+ |S4| [+] |S4| Strongly declining in Vermont due to an anthracnose fungus (Discula sp.). I+] [S3] HH [2\T|S1|* LH HII HII Occurrences in Bennington and Windsor Counties, Vermont, are disjunct. CLG elopoyy 86 OA] NAME CUPRESSACEAE Juniperus horizontalis Moench (14) CUSCUTACEAE Cuscuta coryli Engelm. (14) Cuscuta pentagona Engelm. (14) CYPERACEAE Bolboschoenus maritimus (L.) Palla (1) Scirpus maritimus (14) [MA,NH,RLVT Scirpus paludosus var. atlanticus (11) ECT Bolboschoenus novae-angliae (Britt.) S.G. Smith (1) Scirpus cylindricus (14) [CT,MA] Carex adusta F. Boott (14) ME NH VT MA RI Cr I+] |S3S4]—[2]E|S1]* [2|T|S1|* | | |SE| HII HIT Species = in Grafton County, New Hampshire, and in Bennington County, Vermon HI HII HI I6JWL|S2 [HI |SH] |H|SC *|SH| Difficult to distinguish. May be overlooked. HI |H|E|SH| nan |+?| [S3| |H| |SH| [HH] |SH| Difficult to distinguish. May be overlooked. |+| |S4| |+||S3S4] |1||S1|* |+||S4| |4|C|S1| |9|SC|S2| Disjunct in Addison County, Vermont. |H?| |SU| HII HII [6JWL|S2| || |8|SC|S2S3| 4|E|S1| | | {SUI HII I HII HIT [9661 dODdAN— TP 19 ‘ourys] pur yorquinig CLG NAME Carex albicans Willd. var. emmonsii (Dewey) Rettig (14) Carex emmonsii (11) [CT,NH,RI,VT] Carex alopecoidea Tuckerm. (14) Carex arcta F. Boott (14) Carex atherodes Sprengel (14) Carex atratiformis Britt. (14) Carex backii F. Boott (14) Carex baileyi Britt. (14) Carex barrattii Schwein. & Torr. (14) Carex bicknellii Britt. (14) Carex bigelowii Torr. (14) ME NH VT MA RI [+] |S4] |1\[S1 [1][S1]* [+] |S4| [3{[S1| Disjunct in Chittenden County, Vermont. [ESI Hl [4] [S1| |6|T|S2| HHI |+| |S4?| | ISU| 2|ES1|* HT III Disjunct occurrences in Grand Isle and Franklin Counties, Vermont. || |SH| HH Ht HH HH [10-15] |S2| || [SH] [1/T|S1| I HT |2| |S1]* || |SU| [+] [S3| HI HT Disjunct occurrences in Penobscot County, Maine. }1| [S12|* |3-5?|T|S1S2|+| |SU| |S|E|S 1| I || Disjunct occurrence in Oxford County, Maine. [III HH HT ITI HH |H| |SH| HIT |H| |SH| [+2] [S32| | | [SUI More field study needed. [13] |S2| +] |S3| [4| |S1|* HII HII || Occurrences in Chittenden, Addison, Lamoille, and Washington Counties, Vermont, are disjunct. CT [+I|S3} |H|SC *|SH| HH nan HT [1] |S1| |H|SC *|SH| 2|SC|SI || |SU| PIOpoyYy 86 OA] NAME Carex bushii Mackenzie (14) Carex buxbaumii Wahlenb. (14) Carex capillaris L. (14) Carex capillaris ssp. capillaris 27) Carex capitata L. (14 Carex capitata ssp. arctogena (15) [NH] Carex chordorrhiza L. f. (14) Carex collinsii Nutt. (14) Carex crawei Dewey (14) Carex davisii Schwein. & Torr. (14) Carex eburnea F. Boott (14) ME NH VT MA RI CE |H] |SX| HT |H] |SH| S]ES1| |H] |SH| [2| |S1| |+| |S4| |H|E|SH| |IJE|S1]* [LI|WLIS2| = [H|SH|SH] —[2|E|S1| Disjunct occurrence near Lake Champlain in Addison County, Vermont. [6] |S1| |1|T|S1| [1[T|S1| [1 HT HI IH [3||S1| HH HH I HT [+] |S4?| HII |2|E|S1|* IJE|S1|* Lt Hh Disjunct in Bennington County, Vermont, and in Berkshire County, Massachusetts. HII IIT] HII HIT [TE|S1| |H|SC *|SH| Last seen in 1979. This species has not been re-located in recent searches. [1] [ST HH 1 HT HT [SJE|S1S2| IT HI [1] [S1| IE|S1 [1 2|E|S1| |2|T|S1]* IE|S1| [+1 [S4| [+] [S3| H1 [+] [S3| Disjunct in Oxford County, Maine. 19661 dOOdAN— TP 10 ‘ourysy puv youquinig SLE NAME Carex garberi Fern. (15 Carex garberi var. bifaria (11) [NH] Carex glaucodea Tuckerm. (13) Carex flaccosperma var glaucodea (14) Carex gracilescens Steudel (14) Carex gynocrates Drej. (1) Carex dioica var. gynocrates (14) Carex livida (Wahlenb.) Willd. (14) Carex livida var. radicaulis (1) [MA,ME Carex lupuliformis Sartwell (14) Carex mitchelliana M. A. Curtis (14) Carex muhlenbergii Schk. (14) ME NH VT MA RI cl [142] |S2| |6/E|S1| |4|T|S1| aa HI Lt GRank = G4T3Q; Fed. code = C2. GRank is for synonym C. garberi var. bifaria. I IH|E|SH| 1 4{E|S1| I [1] |S1| Ht |H] |SH| |H| |SH| |IJE|S1| HII [5| |S2| [10+] {S2S3] {11 HT HII HII man Possibly overlooked. [5{T|SIS2| | ITI |AJT|S1| |IJE|S]| aa HIT HII HII [6| |S2| HII II 2|SC|S1| GRank = G3?. LHI io HII IS|WL|S2| | | |SU| HII GRank = G3G |H]| [SH] | |SU| |6/T|S2|* +] |S4S5|— | | [SU [+] |S3| Disjunct occurrences in Chittenden County, Vermont. Reportedly extant in Maine in 1996. OL6 elopoyy 86 TOA] NAME Carex nigromarginata Schwein. (14) Carex norvegica Retz. (14) Carex oligocarpa Schk. (14) Carex oronensis Fern. (14) Carex polymorpha Muhl. (14) Carex prairea Dewey (14) Carex praticola Rydb. (14) Carex rariflora (Wahlenb.) J.E. Smith (14) Carex recta Boott (15) ME NH VT MA RI CT HII 1 HII HI HT |H|SC *|SH| [IJE|S1| HI HT HII HII HT Taxonomy of this and closely related species is currently being studied. HII HII [4|E|S1| |H] |SH| II [IJE|S1| ISIE|S2| HI an HII nan IIT R | GRank = G2; Fed. code = C2. All records are from the Penobscot River watershed. [S|T|S1| |1JT]S1| LIT |2|E|S1| [AJE|S 1] [3|E|S1 GRank = G2G3; Fed. code = C2. |~8|WL|S2| || || |7|T|S2| [5{T|S1|* HII [+] |S4| Maine occurrences are disjunct in Aroostook County. |] [SX| HII 11 HII Ht HII |H] |SH| LI II [1 HII I IJE|S1| II HH [LEIS 1| HII HII This taxon is listed in Gleason and Cronquist (1991) as a synonym under C. salina, but FNA may treat this as a separate species. Thought to be a stabilized hybrid between C. aquatilis and C. pallacea. More study needed. 19661 dOOdAN— TR 19 ‘ourys pur youquinig LEG NAME Carex richardsonii R. Br. (14) Carex saxatilis L. (14) Carex schweinitzii Dewey (14) Carex scirpoidea Michx. (14) Carex siccata Dewey (14) Carex foenea (11) [(CT,MA,NH,VT] Carex sparganioides Muhl. (14) Carex sterilis Willd. (14) Carex striata Michx. var. brevis L. Bailey (14) Carex striatula Michx. (14) Carex tenuiflora Wahlenb. (14) ME NH HH HH 2|E|S1| HT IT | |2|T|S1| 6|T|S]| 1?| [SU II|SU} [I | || |SU| One occurrence in Chittenden County, Vermont, is disjunct. Note: the taxon LI 11 GRank = G3; Fed. code = 2, VT MA [2|E|S1| I HI HI |14) |S2| 3|E|S1| [11] |S2| IH J2|E|S1|* |+||S3?| RI HI |H|SH|SH} HT CT HHT [3|T|S1| HH HSH formerly known as C. foenea is now C. siccata, and what was formerly known as C. aenea is now known as C. foenea. [EISI]* ——‘|AJEJS1 Disjunct in Oxford County, Maine, but varieties in New England nee examination, [3|T|S1| II 1 [1] [S1| IH [1 [8] |S2| [I [+] |[S4 +] |S4| HT |7|T|S2| HT ISJE[S1| [4] |S1| HT }C|S]| |H| |SH| NJE}S 1 HT HI +] |S3| d {9|SC|S2| [1 [2| |S] vIOPOYY 86 “1OA] NAME Carex tetanica Schk. (14) Carex trichocarpa Muhl. (14) Carex vaginata Tausch (14) Carex wiegandii Mackenzie (14) Carex willdenowii Schk. (14) Carex woodii Dewey (14) Cyperus engelmannii Steud. (14) Cyperus houghtonii Torr. (14) Cyperus odoratus L. (14) IND. ME NH VT MA RI 64 | I HT HH [14|SC|S3} |] 7|SC|S2S3| Small population sizes of some occurrences are cause for concern. I [IEJS1| [7| |S2| [8/T|S2| HII [1] [S1| [3|T|S1| [ITI 3|E[S1| HII HII HII |7| |S2| I4{T|S1S2| HI] |SHy |H] |SH| HII HII GRank = G3. HII HII |H| [SH |H| [SH] HII IH|SC *|SH| LI |H] |SX| I HI II |H|SC *|SH| HIT || |H| |SH| J6{SC/S32] [III || |SU| || Taxonomic confusion. A future FNA treatment probably will combine C. engelmannii and C. odoratus, but Massachusetts separates the two species. These taxa are separated here pending further review. |H]| [SH] [2|T|S1| |14|T|S2| 3JE|S1| HII HI Some populations small and threatened. | HT |H] |SH| [4|WLS3?| [SH | | SU Taxonomic confusion. A future FNA treatment probably will combine C. engelmannii and C. odoratus, but Massachusetts separates the two species. These taxa are separated here pending further review. 19661 d°DdAN—TE 19 ‘Joyrysyy pur yorquinig 6LC NAME Eleocharis equisetoides (Elliott) Torr. (14) Eleocharis fallax Weatherby (14) Eleocharis microcarpa Torr. var. filiculmis Torr. ( Eleocharis microcarpa (14) [RI] Eleocharis nitida Fernald (11) Eleocharis ovata (Roth) Roemer & Schultes (14) Eleocharis obtusa var. ovata (14) [CT,MA,ME, VT] Eleocharis ovata var. heurseri (13) [ME,NH] Eleocharis pauciflora (Lightf.) Link (11) Eleocharis pauciflora (14) [VT] Eleocharis quinqueflora (15) [ME] var. fernaldii Svens. Eleocharis quadrangulata (Michx.) Roemer & Schultes ( 14) IND. ME NH VT MA RI HT HH HI |H] |SX| [8|C|S2| I HT [1 |H| |SH| | | SU HI HH HT |IJE|S 1 [IEIS1| || [SRY || [SH |] |SH| IT III SR | GRank = G3G4. Possible occurrence in Maine needs verification. ll II|SU| [5{[S1| S|E|S1| | | SU CT [IJE|S1| || [SU] |H|SC *|SH HHI [+||SU| Taxonomic confusion surrounds this taxon in New England; it is included within the common and widespread E. obtusa by some authors. 2|E|S1| |2|E|S1| [3/T|S1| {IE|S1| HH HT I HI |H] [SX man [2|E|S1| O8T eIOpOouYy 86 OA] NAME Eleocharis rostellata (Torr.) Torr. (14) Eleocharis tricostata Torr. (14) Eleocharis tuberculosa (Michx.) Roemer & Schultes (14) Fuirena pumila (Torr.) Sprengel (14) Rhynchospora capillacea Torr. (14) Rhynchospora inundata (Oakes) Fern. (14 Rhynchospora nitens (Vahl) A. Gray (14) Psilocarya nitens (11) Rhynchospora torreyana A. Gray (14) Schoenoplectus etuberculatus (Steud.) Sojak (13 Scirpus etuberculatus (14) [RI] ME NH VT MA RI CT || |SH| LH II [10+|WL|S3| ae | | [SU More field study needed to determine status in New Englan Ht HII HI 2|E|S1| |H|ISH|SH| {ITI [HJE|S1|* |HJE|SH| HT I+] [S4] [3+] |S2| [+] |S3| Since New Hampshire occurrences are now historic, the Oxford County, Maine, occurrence is disjunct. HII II HH |34|WL|S3)* se |H| |SH| The Hampden County, Massachusetts, occurrence is disjunc 2|E|S]| [IE|S1| |2\T|S1| 2|E|S]| HII [2|E|S 1 HT HI HII |6|T|S2| 4|E|S1| Ht HII HII h IH |LAIT|S2| HII HII HI II IN JLOJE|S2|AE|S1] LI L1 HI II Hh IEIS1| HT d9ODdHN— [FP 12 “‘JJOUIYO| pue yoruquintig [9661 8c NAME Schoenoplectus hallii (A.Gray) S.G. Smith (30) Scirpus hallii (11) [MA] Schoenoplectus heterochaetus (Chase) Sojak (28) Scirpus heterochaetus (14) [ 2 Schoenoplectus x steinmetzii (Fern.) S.G. Smith & A.E. Schuyler Scirpus steinmetzii (11) Scirpus ancistrochaetus Schuyler (13) Scirpus longii Fern. (14) Scirpus pendulus Muhl. (14) Scirpus polyphyllus Vahl (14) IND. ME NH VT MA RI CT I1 1 LIT LI || |H||SX| III III GRank = G2; Fed. code = C2. GRanks and Federal codes are for synonym, Scirpus hallii. Future FNA editions will probably use the name Schoenoplectus Aallii. HT HT [?||S2S3| |WL|SU| HTT IIT May be more common in Vermont than current records indicate, hence the $283 ranking. Populations are difficult to delineate. [1] [ST HII HII HIT HII HII GRank = GIQ. Sterile hybrid of S. heterochaetus x S. tabernaemontani. One isolated population persisting via vegetative reproduction. HII [5-7] |S1| ISJE|S2| IJE|S1| Ht HIT GRank = G3; Fed. code = LE. This species does not appear in any manuals. See A. E. Schuyler’s 1962 paper (Rhodora 64: 43-49) for a description. S{E|S 1 [1] |S HII 4]E|S1| [IJE|S1| GRank = G2; Fed. code = C2. [3|E|S1|* [3|T|S2| I+| ee [26[WL|S3| || {| |+| |S3| Disjunct in Penobscot County, M HII |HJE|SH| 2|E|S1| [S|WLISI] | IT |H] [SH] |H|SC *|SH| i) No vLOpOUYy 86 “19A] NAME Scleria pauciflora Muhl. (14) Scleria pauciflora var. caroliniana (11) [CT,MA] Scleria reticularis Michx. (14) Scleria triglomerata Michx. (14) Scleria verticillata Muhl. (14) Trichophorum clintonii (Gray) S.G. Smith (1) Scirpus clintonii (14) DIAPENSIACEAE Diapensia lapponica L. (14) DROSERACEAE Drosera anglica Hudson (14) Drosera linearis Goldie (14) ME NH VT MA RI CT HII |H] [SH| HII |LAJE|S2| [3T|S1| |H|SC *|SH| Massachusetts has one occurrence of S. pauciflora var. pauciflora in the state and considers this taxon as distinct from S. pauciflora var. caroliniana. HII [1] |S HII |60/WL|S4]_—[3/T|S1| JHE|S 1 GRank = G3G4. Species concept used here does not include the more southern S. muhlenbergii. JWE|S TI [2|T|S1| [1 nan HH IEIS1| HH HT HI HH LH [5-10] |S2} TTI HI HT HH HI HISC *|SH| [13] |S2| [+|T|S3| [IJE|S1|* HII HI HHI Disjunct occurrence in Chittenden County, Vermont. S|E|S]| HT HT LI [1 IT IE|S1| HI IT HT Il 1 19661 dOOdAN— TP 19 ‘yours pue yorquinig 8c NAME DRYOPTERIDACEAE Dryopteris filix-mas Schott (12) Gymnocarpium jessoense (Koidzum!) Koidzumi ssp. parvulum Sarvela (12) Woodsia alpina (Bolton) Gray (12) EBENACEAE Diospyros virginiana L. (14) ELAEAGNACEAE Shepherdia canadensis (L.) Nutt. (14) ELATINACEAE Elatine americana (Pursh) Arn. (11) IND. ME NH VT MA RI CE [2|E|S1| HI |7|T|S2| HH HT HT I I IH] [SH] I [1 I| | SEI Probably introduced in Connecticut, but currently not extant there. [3)T|S]| HII 4|E|S]| HII HII HII HI HII Ht | | ISE| LIT [IJSC|S1| There is some question as to whether Connecticut's single occurrence Is native. The point is moot, however, because only a single individual is extant and the species is dioecious. TJE|S1|* HT +] |S3| HII [III HT Disjunct in Somerset County, Maine. | | |SU| I} II [2|E|S1| [2|T|S1| IH] |SH| More field work needed to determine status. There is taxonomic confusion regarding the New England specimens. vlopouy 86 ‘1OA] NAME EMPETRACEAE Empetrum nigrum L. (14) EQUISETACEAE Equisetum x mackaii (Newman) Brichan (12) ERICACEAE Arctostaphylos alpina (L.) Sprengel (14) Harrimanella hypnoides (L.) Coville 14) Cassiope hypnoides (11) [NH] Loiseleuria procumbens (L.) Desvaux (14) Lyonia mariana (L.) D. Don (14) ME NH VT MA RI CT [+] |S4| [+[T|S3| [4] |S1|* Ht HII IIT Disjunct occurrences in Chittenden, Orleans, and Washington Counties, Vermont. || |SH?| [| |SUI || |SU| HT HII [10+] [SU More field study needed. FNA (Flora of North America Editorial Committee 1993) cites this taxon in Connecticut, Maine, New Hampshire, and Vermont, but says that specimens from Connecticut and New Hampshire have not been seen. Taxon is a hybrid between E. hyemale and E. variegatum. May be more common than previously thought. 1/T|S1] [4[T|S1| HT I HI HT 2|E|S1| [4|T|S2| I HT [1 HT [E|S1| [LO|T|S2} UIT HT I HI I HT HT HH |H|SH|SH| —|HISC *|SH| [9661 dOOdHN— IP 10 “JJourys|, pue youquinig S8C NAME Phyllodoce caerulea (L.) Bab. (14) Rhododendron lapponicum (L.) Wahlenb. (14 Rhododendron maximum L. (14) Rhododendron viscosum (L.) Torr. (14) Vaccinium boreale I.V. Hall & Aald. (14) Vaccinium vitis-idaea L. var. minus Lodd. (11) Vaccinium vitis-idaea ssp. minus (15) [MA] ERIOCAULACEAE Eriocaulon parkeri Robinson (14) ME NH VT MA RI Cr 2|E|S1| [8|T|S2| HII It III I} II J2|E|S1| |7|SC|S3| I} || ITI II ITI l6{TISI|* ——_-|S|T|S2| \7|T|S2|* 7|T|S2| \10+||S3| _-|10| |S3| Disjunct in Caledonia and Orleans Counties, Vermont. Documented decline of some occurrences in Maine. |L/T|S1]* [+|T|S3| |H] |SH| [+] |S5| +] |S4] [+] |S4 Disjunct in Oxford County, Maine. 3|E|S1| [11] |S3| [3] [S| Hh HT HH GRank = G3. [+] |S4| [+] |S4|* [4] |S1| I2|E|S1|* nan Disjunct in Berkshire County, Massachusetts, and in Cheshire County, New Hampshire. [25] |S3| IH ITT I |<4|E|S1| ITI |6|E|S1| GRank = G3; Fed. code = 3C. |H|SC *|SH| O8T vIOpoyy 86 OA] NAME EUPHORBIACEAE Crotonopsis elliptica Willd. (14) Euphorbia glyptosperma Engelm. (14) Chamaesyce glyptosperma (15) [MA] FABACEAE Astragalus alpinus L. var. brunetianus Fern. (14) Astragalus canadensis L. (14) Astragalus eucosmus B. L. Robinson (14) Astragalus robbinsii (Oakes) A. Gray var. jesupii Eggleston (14) Astragalus robbinsii (Oakes) A. Gray var. minor (Hook.) Barneby (14) Astragalus robbinsii (Oakes) A. Gray var. robbinsii (14) Desmodium canescens (L.) DC. (14) ME NH VT HT [1 HT || |SU| | | |SU| [7] |S 1 Considered introduced into several states; tracking as a native species in Vermont only recently begun. [27| |S2| |H| ae |H] |SX| GRank = G5T2T3 ITI HI |9[T|S2| {H| |SX| HII HHI HI 2|E|S1| [IJE|S1| GRank = G5T1; Fed. code = LE. |H] |SX| HHI {7| |S2| III LHI i GRank = GSTX; Fed. code = 3 HT HT HII MA LH II|SE| HI HI HH |S|WLS1| RI Ht Ht Ht HT IT Os | |H|SC *|SH| LI HT HT HH nan LT | [8] |S3] [9661 dODdHN— TP 19 ‘ouryay] puv yorquinig L8C NAME Desmodium cuspidatum (Muhl.) Loudon (14 Desmodium glabellum (Michx.) DC. (14) Desmodium humifusum (Muhl.) Beck (14) Desmodium sessilifolium (Torr.) T. & G. (14) Lathyrus ochroleucus Hook. (14) Lespedeza repens (L.) Barton (14) Lespedeza stuevei Nutt. (14) Lupinus perennis L. (14) Oxytropis campestris (L.) DC. var. johannensis Fern. (14) Div ME NH VT MA RI CT HT |] |SH| [3]E|S1| ISWLIST] AT [| SU HT [11 LT HI HI [S|SC|S1| HH 1 HII 2|E|S1 HII [2/ES1| || | GRank = G1G2Q; Fed. code = C2. Recent research suggests this species is a hybrid of D. paniculata x D. rotundifolium. Ht It HHI HII |IE|S1| |H|SC *|SH| IIT HII [8] |S2| Ht Ht II HIP HII II [12] [S| III [1JSC|S1| aan || |H] |SH| 19} |S3?| [| ISUI || [SU Current status uncertain. \H] |SX |21|T|S1|* [2\T|S1|* |}+{WL|S3|* = |8}C|S2|* |12| |S2|* Documented decline in Connecticut, Rhode Island, Massachusetts, New Hampshire, and Vermont. Note: most New Hampshire occurrences are small and isolated, primarily on roadsides and power lines. [8|TISIS2| | || HII H1 HI HII GRank = G5?T3; Fed. code = 3C. vIOPOYY 86 OA] NAME Div ME NH VT MA RI CT Phaseolus polystachios (L.) BSP. (14) 4 III tI I III III II |] |H|SC *|SH| Phaseolus polystachios var. Reported from all of New England in Gleason and Cronquist (1991), but we have 19661 aquilonius (11) [CT] seen specimens only from Connecticut. Strophostyles umbellata (Muhl.) 4 HII ane ann tI] |H|SH|SH| III Britton ( FUMARIACEAE Corydalis aurea Willd. (14) 2 It |H| |SX| |6|T|S2| III I aan I} || Corydalis flavula (Raf.) DC. (14) 2 HI LI HII HII nan 4|T|S1| Dicentra canadensis (Goldie) Walp. a |3|T|S1|* JLIJT|S2S3] — [+] |S4| |+| |S4| I ||| |6|T|S2| (14) Disjunct in Franklin and Penobscot Counties, Maine. GENTIANACEAE Gentiana andrewsii Griseb. (14) 2 Lt |2|T|S 1 [S|T|S1| |2\T|S1| |H|SH|SH| — || |SU| Specimens of this species and G. clausa should be examined closely due to similarity. Gentianella amarella (L.) Boerner (14) pl JIJE|S1} II {| |H|T|SH| I 1| Ree III] Gentiana amarella (11) [VT] Gentianella quinquefolia (L.) Small 2 |H] |SH| |H] |SH| [2\T|S1| \12+|WLIS2| ||] | |2|E|S1 (14) Gentiana quinquefolia (11) [CT,MA] dODdHN—TP 39 ‘ourys, pue yorquinig O8C NAME Lomatogonium rotatum (L.) Fries (14) Sabatia campanulata (L.) Torr. (14) Sabatia dodecandra (L.) BSP. (14) Sabatia kennedyana Fern. (14) Sabatia stellaris Pursh (14) GROSSULARIACEAE Ribes rotundifolium Michx. (14) HALORAGACEAE Myriophyllum pinnatum (Walter) BSP. (14) Myriophyllum verticillatum L. (14) IND. IND. ME NH VT MA RI Cr 9 |SIS2] IIT HI I [1 nae [1 HT HT [S|E]S1| HI HII HHT I HI HH HT |H|SC *|SH| I I [1 140|SC|S3|_|4{E|S 1 ae | GRank = G3. Massachusetts has the largest number of occurrences globally. HII HHI It IJE|S1| 4|T|S1| [2| [S| Species has not been seen in Massachusetts in recent years. It HI HII JWLISI fT |H|SC *|SH| Specimen from Massachusetts appears valid, but it is unknown whether this species is truly native there. HI ITI Ht] |6|SC|S2?| More field study needed to determine status. lI[TIS1| |HISC *|SH| [+?| |SU| || |SU| [5| |S1| L|T|S1| HII More field study needed to determine status. [| |SU| ic vlopoyy 86 TOA] NAME HAMAMELIDACEAE Liquidambar styraciflua L. (14) HIPPURIDACEAE Hippuris vulgaris L. (14) HYDROPHYLLACEAE Hydrophyllum canadense L. (14) HYMENOPHYLLACEAE Trichomanes intricatum Farrar (12) IRIDACEAE Sisyrinchium mucronatum Michx. (14) ISOETACEAE Isoétes acadiensis Kott (12) ME NH VT MA RI CT HII HI HH HI IIT] |8|SC|S2S3 This species has been introduced into some states. It is difficult to determine which Connecticut occurrences are native and which are introduced. || [SU| |2/T|S3| [2|E|S1| HT HT nae HII HI {1}T}S1| 2|E|S1| HII HII [1] [St || SUI 3|T|S1] HH [3|SC|S1| GRank = G3G4. |<52| [S12] |H] |SH| |] [SH [3|T|S1| HII || |SU| Not tracked by the Connecticut Natural Diversity Data Base. The Maine Natural Areas Program began tracking this species in 1996. HII I3|E [1] |S] |1| |S1 S]| HT IT GRank = G3?. [9661 dODdAN—'IP 19 ‘ourya| pur yorquinig 16c NAME Isoétes x eatonii Dodge (12) Isoétes eatonii (13) [NH] Isoétes x foveolata A.A. Eaton ex Dodge (12) Isoétes lacustris L. (12) Isoétes macrospora (11) [CT,MA,ME,NH,VT] Isoétes prototypus D.M. Britton (12) Isoétes riparia Engelmann ex A. Braun (12) Isoétes riparia var. canadensis (11) [RY] JUGLANDACEAE Juglans cinerea L. (14) IND. IND. ME NH VT MA RI be i [III |H||SH| HII [1] |SU| HII [4 |S GRank = G2Q; Fed. code = 3B. GRank is for synonym, /. eatonii. Sterile hybrid of J. engelmannii with I. echinospora. Ht HII HHI |] |SH| HII [NISC *|SH Current status unknown; hybrid of /. engelmannii with I. tuckermanii. | +2||SU| [ITS1| [?||S2?| JIJE|S1| || |SU| |1||SUJ More field study needed. FNA (Flora of North America Editorial Committee 1993) says that North American plants of /. /acustris have been segregated as /. macrospora by some authors, but that the two taxa cannot be distinguished reliably from each other except on the basis of geography. 1] [S1| HII HII HII HII || GRank = G2G3. |H| |SH| [2|T|S2| [2| |S1| |] [SH |4|C\S]| | | [SU +] |SU] [+] [S3?| [+] |S4] [+] |S4| [+] [SU] [+] [SU] Fed. code = C2. Declining in some New England states because of the fungus, Sirococcus clavigignenti-juglandacearum, and therefore should be monitored. Z vIOPOYY 86 TOA] NAME JUNCACEAE Juncus alpinus auct. non Vill. (13) Juncus alpinoarticulatus (14) [ME,NH] Juncus biflorus Ell. (14) Juncus debilis A. Gray (14) Juncus x oronensis Fern. (14) Juncus oronensis (11) Juncus pervetus Fern. (11) Juncus stygius L. var. americanus ch. (14) Juncus subtilis E. Meyer (14) Juncus torreyi Cov. (14) Juncus trifidus L. (14) IND. ME NH VT MA RI tl 9 [7-10|T|S2|_— {IIS [8] [S2| I HH HH HT HI I [S|E|S2| HH I Itt HT HIT |IJE|S1| [3|C|S1| |H|SC *|SH| |H| |SH?| HII HH [III II HII Name is based on two historic specimens occurring with the parents (J. tenuis x J. vaseyi). Status uncertain, Not tracked by the Maine Natural Areas Program. HII HII II |H] |SX| II 1 HI Taxonomic status unclear. Fernald (1950) says native at one site in Massachusetts, but Gleason and Cronquist (1991) include the taxon under the European Juncus subnodulosus. [9| |S2| I HH HT HI HT |H] |SU| HI HT HT HT HT || [SU] Ht [2|E|S1| | | SE} I HI |+||S3S4] [S384] [S| [SI I II HT | | Disjunct occurrences on high peaks in Chittenden, Windsor, Washington, and Lamoille Counties, Vermont. [9661 dODddN— TP 19 ‘ourysp] pue yorqunig C67 NAME Juncus vaseyi Engelm. (14) Luzula confusa Lindeberg (14) Luzula spicata (L.) DC. (14) JUNCAGINACEAE Triglochin gaspense Lieth & D. Léve (15) LAMIACEAE Agastache nepetoides (L.) Kuntze (14) Agastache scrophulariifolia (Willd.) Kuntze (14) Blephilia ciliata (L.) Benth. (14) Blephilia hirsuta (Pursh) Benth. (14) Blephilia hirsuta (Pursh) Benth. var. glabrata Fern. (11) IND. ME NH VT MA RI CT [1] [S1| HII 1} |S1| HI Itt HII JIE|S 1 |H|E|SH| HH HII HII Ht [ESI |7|T|S3| [1] |S HII HII HHI |H| |SH| ITT || | aan HT LTT GRank = G2; Fed. code = C2. This taxon has been separated from T. maritimum. (See Léve and Lieth 1961). I IH IIT |S 1 H1 HIT [I|SC|S 1 HI HH [ITS 1 |] [SH] HII 2|SC|S1| HH HI |H| [SH| JIJE|S 1 I |] [SH] HI ITI [2/T|S]| I7|E|S1| III [IISC|S1| HH aa || |SH| HH HII HA GRank = G4?T1Q. This taxon is a Vermont endemic described by Fernald. It has never been re-collected. POC vIOPOYY 86 1A] NAME Dracocephalum parviflorum Nutt. (14) Lycopus rubellus Moench (14) Monarda punctata L. var. villicaulis Pennell (14) Pycnanthemum clinopodioides T. & G. (14) Pycnanthemum torrei Benth. (14) Scutellaria integrifolia L. (14) Scutellaria leonardii Epling (14) Scutellaria parvula var. leonardi (11) [MA,ME] Scutellaria parvula Michx. var. parvula (14) Stachys hyssopifolia Michx. (14) IND. IND. 3* ME NH VT MA RI CT || |SE| HII JAJT|S1| || |SE| || |SE| | | |SE| Recent searches have not re-located the Vermont occurrence. It was last seen in 1983 [1 |H] [SH] [1] [S1| 2(T|S1| 1] |SU| [2| [SU] HT HT [2| |S} || ISE| HII || |SUI HT | II | [IJE|S 1] HII S|E|S1] | GRank = G2. Taxonomic confusion. Difficult to distinguish and suspected by some botanists to be of hybrid origin. More field and laboratory work needed. HI [IE|ST| Ht nae IIT 2/E|S 1 GRank = G2. Taxonomic confusion. Difficult to distinguish and suspected by some botanists to be of hybrid origin. More field and laboratory work needed. HI HT HT |H] [S| HT [I|SC|S1| |H||SH| II I WLISU] {III [IE|S1| || [SU HH [11] |S2| I HT LT HT HI [1 [S9|WL|S4]—|IJE|S1| 2\E|S1]* || Disjunct in Hartford County, Connecticut. 19661 dODdAN— IP 19 ‘ouryspy pure yorquinig S6C NAME Stachys pilosa Nutt. (13) Stachys palustris ssp. pilosa (1) [(MA,ME] Stachys tenuifolia Willd. (14) Stachys tenuifolia var. platyphylla (11) [NH] Trichostema brachiatum L. (15) Isanthus brachiatus (14) [CT] LEMNACEAE Lemna valdiviana Philippi (14) Wolffiella gladiata (Hegelm.) Hegelm. (17 Wolffiella floridana (14) LENTIBULARIACEAE Pinguicula vulgaris L. (14) IND. ME NH VT MA RI Cr II|SE| || [SU] || [SU| II|SU| HII HIT Status unclear for this and closely related taxa. | | ISE| II|SU| HIT] [| |SU| |H| |SH| |H|SC *|SH| More field study needed. Recorded for New England in Seymour (1969) and the Maine checklist (Campbell et al. 1995), but Gleason and Cronquist(1991) give a more western range for this and closely related taxa. HII HH [HI |SH] 3|E|S1| HII |H|SC *|SH| HII |IJE|S2| HT | | {SU HII | | |SU} Not enough information on extant occurrences. Identity of plants in certain reports has been questioned. More field work needed. [II IIT HII [1] |S1| | HIT | || Species is being recommended for state endangered species list in Massachusetts in 1] |S1| |2|E|S2| 1] {S11 III III IT Unknown from Maine until 1996. 96C vIOpOYYy 86 “1OA] NAME Utricularia biflora Lam. (14) Utricularia fibrosa Walter (14) Utricularia inflata Walter (14) Utricularia resupinata B. D. Greene (14) Utricularia subulata L. (14) LILIACEAE Chamaelirium luteum (L.) A. Gray (14) Melanthium hybridum Walter (14) Tofieldia glutinosa (Michx.) Pers. (14) Zigadenus elegans Pursh var. glaucus (Nutt.) Preece (14) Zigadenus glaucus (11) [VT] ME NH VT MA RI Cr HII It Ht [7/T|S2| 2/T|S]| |H|SC *|SH| Treated as a synonym of U. gibba in Kartesz (1994). HT HI HT |6{T|S2| HT |H|SC *|SH| | HI HT [2| |S1| HT HH Indigenous status in Massachusetts is uncertain. [2|T|S1| | | SU S|T|S 1 |~8)WL|S2| — |4{T|S1] [I|SC]S 1 Some occurrences contain small numbers of plants. HH HA HH [LO|SC|S3]—[4|C|S1| HH HI I 1 3-4|ESI] TIT S|E|S1| III HT [11 It HI |H|SC *|SH| |+| |S3S4] |4[T|S1]* |4|T|S1|* I HI LI Disjunct in Cheshire, Sullivan, and Grafton Counties, New Hampshire, and in Windsor and Caledonia Counties, Vermont. HT I |H||SH| [1 HI HT dODdHN— TP 19 ‘Jourys, pue yorquinig [9661 LOC NAME LIMNANTHACEAE Floerkea proserpinacoides Willd. (14) LINACEAE Linum medium (Planchon) Britton var, texanum (Planchon) Fern. (14) Linum medium [ME,VT] Linum sulcatum Riddell (14) LYCOPODIACEAE Diphasiastrum x sabinifolium (Willd.) Holub (12 Lycopodium sabinifolium (14) Feral sitchense (Rupr.) Holub (12) Lycopodium sitchense (14) (NH, VT] Huperzia appalachiana Beitel & Mickel (12 IND. ME NH VT MA RI ed [tl HIT || |SH| |H]| [SX HII 4|E|S1 II|SU} HH |H||SH| 4/T|S1| || [SUI || |SU HII HH |H] |SH| |H] [SX] JIE|S1| [I|SC|S]| [5|T|S1| II|SU| [3||S1 IT HI HT According to FNA (Flora of North America Editorial Committee 1993) this hybrid between D. sitchense and D. tristachyum is highly variable. |LJE|S1| {ISU |H|T|SH| [ITI HII HII |7| [S2| | | [SU 7] [S]| | 2\E|S1| HII HII GRank = G3. Recent FNA (Flora of North America Editorial Committee 1993) treatment suggests high elevation alpine occurrences are this taxon, but not all occurrences have been verified. Massachusetts’ only known occurrence is suspected to be a hybrid (H. appalachiana x H. lucidulum). vIOpOYyy 86 [OA] NAME Huperzia selago (L.) Bernhardi ex Schrank & Martius (12) Lycopodium selago (14) (CT,NH,VT] Lycopodiella alopecuroides (L.) Cranfill (12) Lycopodium alopecuroides (14) [CT,MA,RI] Pseudolycopodiella caroliniana (L.) Holub (12) Lycopodium carolinianum (14) [MA] LYGODIACEAE Lygodium palmatum (Bernh.) Swartz (12) LYTHRACEAE Cuphea viscosissima Jacq. (14) ME NH VT MA RI Cr 1] [S| [?||SU| [?|S1| IJE|S1 HII IH|SC *|SH| Recent FNA (Flora of North America Editorial Committee 1993) treatment suggests lower elevation plants are this taxon, but not all occurrences have been verified. HH HT I [E|S1| IES] | |H|SC *|SH| III nae HI |H||SX| I I || This species was last seen in New England on Mt. Toby in Massachusetts in 1976, but is believed to be extirpated from this site. HII |H] [S| [IJE|S1|* I22|SC|S3|_—J6|C|S1| [L1]SC|S2]* Documented decline in Connecticut. Disjunct occurrence in Lamoille County, ont. HII HII HII | |SX| |H] |SH| |H] [SH| Sorrie (1991) considers this species "doubtfully native" in Massachusetts. [9661 dODdHN— IP 19 ‘Jourys| pue yorquinig 6670 NAME Lythrum alatum Pursh var. alatum (14) Rotala ramosior (L.) Koehne (14) MAGNOLIACEAE Magnolia virginiana L. (14) MELASTOMATACEAE Rhexia mariana L. (14) MORACEAE Morus rubra L. (14) NAJADACEAE Najas guadalupensis (Sprengel) Magnus (1 IND. ME NH VT MA RI Gi 8 | | [SE| || ISU| | |SE| | | ISE| [| |SU| || |SH| ome occurrences in southern New England are thought to be native, but the species is adventive northward. Vermont occurrences are introduced and no longer extant. IT HII H IE}S1| IJE|S 1 [S|E|S1| HII | HT 3|E|S]| HII [111 [III HII IIH] S|E|S1| HT I IT HT |2/T|S1| S|E|S1| I [S]E|S 1 Some introduced populations occur in Vermont and other New England states. |H] [SH] ||| SU [S| |S1| ||WLISE| —|2{T|S1| More field study needed. Considered introduced into at least one state (Massachusetts). May be more common than previously recorded. IIISC|S1| OO€ vIOPOUY 86 TOA] NAME NYMPHAEACEAE Nuphar lutea (L.) Sm. ssp. advena (Ait.) Kartesz & Gandhi (1) Nuphar advena (14) [CT,NH,VT] Nymphaea leibergii Morong (13) Nymphaea tetragona (14) Nymphaea odorata Aiton ssp. tuberosa (Paine) Wiersema & Hellquist (13) Nymphaea odorata var. tuberosa [VT] Nymphaea tuberosa (11) [MA] ONAGRACEAE Epilobium anagallidifolium Lam. (15) Epilobium alpinum (14) Epilobium hornemannii Reichb. (14) Epilobium hornemanni (11) [ME,NH] Ludwigia polycarpa Short & Peter (14) Ludwigia sphaerocarpa Elliott (14) Oenothera fruticosa L. (14) IND. 2 IND. ME NH VT MA RI 64 [1-5] [S22] [AIST [H||SH| It HI |HSH| Relatively few occurrences of this taxon, but some occurrences include entire [6] |S1| HII HT IIT HII HT Ht HI |?|[SU| II| SEI HII |H|SC *|SH| More field study needed. 2|T|S1| [1] [S| HIT man HII Ht 3/T|S1| |6/T|S2| HII Ht III HII HII HII 2\E|S1| [7\T|S2| [ITI |H|SC *|SH| HIT HII IIT 2/T|S1| [IJE|S1| 2|E|S1| | | |SE| HII [ITI | | |[SE?| [1+] |S1| |H|SC *|SH| More field study needed to determine status in New England. Uncertainty exists regarding its status as a native species. dODdAN— TP 19 ‘Joyrys] puke youquinsg [9661 NAME OPHIOGLOSSACEA Botrychium lunaria (L. Swartz (12) Botrychium minganense Victorin (12) Botrychium oneidense (Gilbert) House ( Botrychium rugulosum W. H. Wagner (12) Ophioglossum pusillum Raf. (12) Ophioglossum vulgatum (14) [CT] ORCHIDACEAE Amerorchis rotundifolia Banks (13) Orchis rotundifolia (14) IND. IND. IND. ME NH Ve MA RI CT [3|E|S1| |] |SH| |HJE|SH| || |SR| HII HII FNA (Flora of North America Editorial in dines 1993) reports this species from Massachusetts, but we have not seen specimen |H?| |SH2|_ | | [SRI | [SRI HII [III Ht FNA (Flora of North America Editorial Committee 1993) sites this taxon from New Hampshire and Vermont, but we have not seen specimen || {SUI | | [SU | | [SU | | [SU | | |SU [| |SU| Difficult taxon to distinguish. FNA (Flora of North America Editorial Committee 1993) attributes this to every state in New England. HII II || |SU| ITI HII | | [SU GRank = G3. Many older Vermont collections of this taxon exist. Taxon is not in FNA (Flora of North America Editorial Committee 1993) for Connecticut, but one specimen exists from that state. Difficult to distinguish from B. multifidum or B. dissectum. More field work is needed. [+2] |S3?| | | |SU| +] |S3] |6{T|S2|* warat [3/T|S1|* Documented decline in Massachusetts, Connecticut, and Rhode Island |6|T|S1]° HT |H| |SH| HT HII HII cOe elopoyy 86 1OA] NAME Aplectrum hyemale (Muhl.) Torr. (14) Arethusa bulbosa L. (14) Calypso bulbosa (L.) Oakes (14) Corallorhiza odontorhiza (Willd.) Nutt. (14) Cypripedium arietinum R. Br. (14) Cypripedium parviflorum Salisb. var. makasin (Farwell) Sheviak (13) Cypripedium parviflorum Salisb. var. parviflorum (13) Cypripedium calceolus var. parviflorum (14) [CT,MA Cypripedium parviflorum (15) [ME,NH,RI,VT] IND. ME NH VT MA RI CT HIT Ht JIJT|S1| 3|E|S1| IT] |H|SC *|SH| [+||S3S4| [4{E|S1| 6IT|S2\* —|L6|TIS2\* —|SJEISU* {LEIS Documented decline in Connecticut, Massachusetts, Rhode Island, and Vermont. |+| |S3S4| |H|E|SX| [7|T|S2|* III I} 1 ae Documented decline in Vermont. |2|E|S1|* |3|E|S1|* |6|T|S2| J13|SC|54| [3] TIS TI [10] |S3| Disjunct occurrences in Strafford and Carroll Counties, New Hampshire and in York and Oxford Counties, Maine. |S|T|S1| \2|E|S 1 |18|T|S2| HIJE|S 1} a |H|SC *|SH]| GRank = G3; Fed. code = 3C. Ht || [SU] | | {SU [2?| |SU| || |SU| || [SUI Difficult to distinguish. Recent taxonomic work splits yellow lady's slippers into three taxa; the disposition of this taxon in New England is unclear. [+2||S3?| 2\E|S1| ||S3| 29E|S1| ISU} [III Difficult to distinguish. Recent taxonomic work splits yellow lady's slippers into three taxa; the disposition of this taxon in New England is unclear. 19661 d9DdAN— TP 19 JJOUIYOI Pue Youquins | NAME Cypripedium parviflorum Salisb. var. pubescens (Willd.) Knight (13) Cypripedium calceolus var. pubescens (14) [CT] Cypripedium pubescens (15) [MA,ME,NH,RI,VT] Cypripedium reginae Walter (14) Galearis spectabilis (L.) Raf. (15) Orchis spectabilis (14) [RI, VT] Goodyera oblongifolia Raf. (14) Isotria medeoloides (Pursh) Raf. (14) Liparis liliifolia Rich. (14) Listera auriculata Wieg. (14) Listera australis Lindl. (14) IND. ] ME NH VT MA RI Cr |+?||S3?| 16|T|S2| |+||S3| IH?|}WLIS3| [4 T|S1| |ISU| Difficult to distinguish. Recent taxonomic work splits yellow lady's slippers into three taxa; the disposition of this taxon in New England is unclear. \26| |S2S3| — [S5|SE|S1| |+| |S3| |18|SC|S3|* | | | | |3JE|S1|* Documented decline in Connecticut and Massachusetts. [2/T|S1|* 4/T|S2| +||S4| +] |S3| [I|SE|S1]| I+] |S3| Disjunct in Oxford and Somerset Counties, Maine. Recent field work seems to indicate a decline in southern New England occurrences. [S|E|S 1] IIT It I 1 I} | IT JI7|EIS2) — H{E|S2I IHJEISH} —(3|E|S1] |IJEJS1| {IJE|S1| GRank = G2G3; Fed. code = LT. The majority of occurrences of this globally rare species are in New Hampshire and Maine. HT |H] |SX| 1]T|S1| I8+{WL|S2|_— [2/T|S1| |2|E|S1| [7| |S1| |3|E|S1| IE|S1| HT HT HT GRank = G3, HT IT 2|E|S1| IT II [1 poe vlopoyuYy 86 “1OA] NAME Listera cordata (L.) R. Br. (14) Malaxis bayardii Fern. (11) Platanthera ciliaris (L.) Lindl. (15) Habenaria ciliaris (14) Platanthera cristata (Michx.) Lindl. (15 Habenaria cristata (14) Platanthera leucophaea (Nutt.) Lindl. var. leucophaea (15) Habenaria leucophaea (14) Spiranthes casei Catling & Cruise (14) Spiranthes x intermedia Ames (20) Tipularia discolor (Pursh) Nutt. (14) IND. ME NH VT MA RI cr |+| [S3S4] |8|T|S2} |+| |S3| Wiad |H|SH|SH III I Disjunct in Barnstable County, Massachusetts HII HII |H] |SH] 2-3]E|S1] tt HT] [SH GRank = G2?. This species is similar to Malaxis unifolia. Specimens should be looked at critically. HII HT HT |H] |SX 2|E|S1| [8|T}S2| HII HII HIT [E?|E|S 1 HI IIT Massachusetts occurrence has not been seen in recent years and is likely now extirpated. Aan IT HII HII HII HII = G2; Fed. code = LT. || [SUI ua [?| [SU HII HII HII Further study need | | [SU] [| SU | |SU| || |SU| || ISU || |SU Luer (1975) shows this taxon as present in all New England states. It is reportedly (Sorrie 1991) a hybrid of S. lacera var. gracilis x S. vernalis, but S. vernalis does not occur in Maine. Further study needed. HT [11 HT [7|E|S2| [1 HH [9661 dODdAN— TP 19 ‘ourysp] pur yorquinig SOe NAME Triphora trianthophora (Swartz) Rydb. (14) OXALIDACEAE Oxalis violacea L. (14) POACEAE Agrostis mertensii Trin. (14) Agrostis borealis (11) [NH] Ammophila champlainensis Seymour (29) Amphicarpum purshii Kunth (14) Aristida basiramea Engelm. (14) Aristida purpurascens Poiret (14) Aristida tuberculosa Nutt. (14) Bouteloua curtipendula (Michx.) Torr. (14) 2(a) IND. 2 ME NH VT MA RI Cr [7|T|S1S2| |10/T|S2| [3|T|S1| [2|E|S1| I ||| |H|SC *|SH| Small population sizes of some occurrences are cause for concern. Furthermore, entire occurrences of this species may not emerge every year. HII | HT [5|T|S1| IE|S1| [SSCS 1 16| |S2| [7\|S3| [5] [S| HH HII HII HIT 111 2|E|S1| HT HH HT GRank = GIQ. Taxonomic study to determine if this is a good species or a variety of A. breviligulata is in progress. HA HI HT [IE|S1| HH HH [2| |S1| [1| [SU] [2 [SUI III an HII May be more common than originally thought. More field work needed. HN HH [1 |14[T|S2S3) — [3/T|S1| [| [SH HH |2|E|S1| I ISISCIS3}— IIT [5|T|S1| HI LT HN HT HI HJE|S1| 90¢ VIOPOYUY 86 [OA] NAME Calamagrostis canadensis (Michx.) Beauv. var. langsdorfii (Link) Inman (15) Calamagrostis nubila (11) [NH] Calamagrostis pickeringii A. Gray (14) Calamagrostis stricta (Timm) Koel. ssp. inexpansa (Gray) C. W. Greene (15) Calamagrostis stricta var. inexpansa [NH,VT] Calamagrostis lacustris (14) [NH] Calamagrostis stricta (Timm) Koeler ssp. stricta Calamagrostis neglecta (11) [NH] Deschampsia atropurpurea (Wahlenb.) Scheele (14) Vahlodea atropurpurea (1) [ME] Elymus villosus Muhl. (14) Eragrostis capillaris (L.) Nees (14) IND. ME NH VT MA RI CT HIT |HI[SX| HII II It HII GRank = GHQ. GRank applies to synonym C. nubdila. Last collection at Lake of the Clouds by Boott in 1862. 2|E|S1| 9|T|S3| [H] |SH| |H| |SX| HT II [3|T|S1| [7|E|SU| [2|E|S1| IH I 11 [AJSC|S1| GRank = G3Q. GRank is for C. lacustris. S|EIS1| [5|T|S1| [1 HII HII [1 |H||SH| [3] |S2| |H| |SH| LI IT HI [1 I [3] |S1| |3[T|S2| || |SU| | | SU [IE|S1| | | [SU] ?||S2S3]— 4[WLISU] || |SU| | | [SU Adventive in gardens, roadsides, and railroads; difficult to determine which occurrences are native. 19661 dODdAN— TP 19 ‘Jouryspy pue yorquinig NAME Hierochloe alpina (Swartz) Roemer & Schultes (14) Leptochloa fascicularis (Lam.) A. Gray var. maritima (Bicknell) Gleason (14) Diplachne maritima (11) [(CT,MA,NH] Leymus mollis (Trin.) Pilger var. mollis 14 Elymus mollis [MA,NH] Muhlenbergia capillaris (Lam.) Trin. (11) Muhlenbergia richardsonis (Trin.) Rydb. (14) Muhlenbergia sobolifera (Muhl.) Trin. (14) Oryzopsis canadensis (Poiret) Trin. (14) IND. ME NH VT MA RI Le i [5|T|S]| |7| |S2| [2|T|S1| [1 I HII HT eae man |6|T|S2| |H|H|SH| 2|E|S1| GRank = G5T3T [+] |S4| |H||SX| HH I2|E|S1| HII aan Confusion with the introduced L. arenarius makes this taxon's status unclear. HII HII HT |H] |SX| HII 2|E|S1| [2| |S1| HII HT HII HII HII Lea [2|T|S1| +] |S3| +] |S4| [| ISU ut The occurrence in Oxford County, Maine, is disjunct. State rank of "SH Maine i is based on a 20-year cutoff date used by the Maine Natural . Program. |H?||SH?|——_|HJE|SH| Ht HT I | 80e vVIOPOYY 86 ‘1OA] NAME Panicum amarum Elliott (14) Panicum flexile (Gattinger) Scribn. (14) Panicum gattingeri Nash (11) Panicum dichoton mattamuskeetens al 3) Dichanthelium Ree eee (31) [MA] Panicum mattamuskeetense Ashe (11) m SSp. Panicum polyanthes Schultes (14) Dichanthelium sphaerocarpon var. isophyllum (15) [CT] Dichanthelium polyanthes [MA] Panicum rigidulum Bosc. var. pubescens (Vasey) Lelong (15) Panicum longifolium (11) (CT,MA,NH] Panicum scabriusculum Elliott (14) Dichanthelium scabriusculum (15) x IND. IND. ME NH VT MA RI CT ITT III III | | |SE} [1+] |SU| |7|T|S2| Massachusetts notes P. amarum var. amarulum as introduced in the state. I HH I2|E|S1| HT I |] [SH] Hh HI Ht [7/SC|S22| |] |H] |SH| III HII eR [72|E|SU?| [A] [SH| HII May be overlooked on Cape Cod and the islands off the coast of Massachusetts. More field work needed. HI IIT HT II SU] HT oe There is some confusion with this taxon and P. sphaerocarpon. More fi work needed |H||SH| HT |6|T|S2| || [SU] |HI|SH| IH HT HT [2|T|S1| HI [IE|S1| 19661 dODdAN— TP 19 ‘JJousrys] pue yorquinig 60¢ NAME Panicum sphaerocarpon Elliott (14) Dichanthelium sphaerocarpon (1) ME] Panicum stipitatum Nash (11) Paspalum laeve Michx. (14) Paspalum setaceum Michx. psammophilum (Nash) D. Banks (14) Phleum alpinum L. (14) Poa glauca Vahl (14) Poa laxa ae ssp. oe (Nannf.) Hylander (13 Poa nario. ([ME,NH, VT] Poa pratensis L. ssp. alpigena (Fries ex Blytt) Hiitonen (13) Poa arctica (14) IND. ME NH VT MA RI cr II|SU| |H|E|SH| [4] |S [+||SU] | |SU| [| ISU Taxonomic confusion between this taxon and P. polyanthes. Further taxonomic and field work needed. ITI III It ITI IT |H| |SH| State of Connecticut a i species list cites Panicum rigidulum var. elongatum as this taxo |H] |SX| IH HHT HH HII i2|E|S1| an HIT II [7JWLIS2|— | | [SUI [8|T|S1| |2|T|S2| Ht HT HII HI |<10||SU}— [H/T|SH| [1] [SA] I I HI [IJEIS1| 2|E|S2S3] {1 ||S1| | | HT || || GRank = G2G3. Global rank is for synonym P. fernaldiana. | | |SU| |HJE|SH| II 1 I} TI ITI HHI In the Maine checklist (Campbell et al. 1995) this taxon is included under P. pratensis, which is considered common in Maine |H|SC *|SH| Ole elopoyy 86 1OA] NAME Puccinellia tenella (Lange) Holmb. ssp. langeana (Berlin) Tzvelev (15) Puccinellia tenella (Lange) Holmb. ssp. alascana (Scribn. & Merr.) Tzvelev (15) Puccinellia langeana ssp. alascana (16) [CT,MA] Puccinellia paupercula var. alaskana (11) [NH] Sorghastrum nutans (L.) Nash (14) Spartina cynosuroides (L.) Roth (14) Sphenopholis nitida (Biehler) Scribn. 4) Sphenopholis obtusata (Michx.) Scribn. (14) Sphenopholis pensylvanica (L.) A. Hitche. (14) — nN 2 ND. ME NH VT MA RI Cr HII [ITI HII | | [SUI HII HT GRank = G4?T3T4. Cited by one source as historic in Massachusetts, but recent manuals do not list a taxon with this name for that state. Investigation needed. [?| [SU II|SE| HTT |H||SH] Ht |H|SC *|SH| Taxonomy and nomenclature confusing. More study needed in New England to determine status. May not be rare in Maine. [2| |S1|* | | ISU] +] [S3| +] |S4} [S}C|S1| [+] [S4] Disjunct in Somerset and Androscoggin Counties in Maine. HIT HII [III 8|SC|S2| 3|C|S1| |<10| |S2| HII HH [IJE|S1| |3|T|S1| |H|SH|SH| | H] |SH| |H] |SH| |H{E|SH| IJE|S1| | | |SU| |H] |S] |H| [SH] May be more common than previously thought; more field study needed to determine status. [1 [1 HT [4|T|S]| || [SU |] |SH| dODdHN— IP 19 “Jyourysy, puke youquinig [9661 Ile NAME Sporobolus clandestinus (Biehler) A. Hitche. (14) Sporobolus compositus (Poir.) Merr. var. compositus (15) Sporobolus asper (14) [CT,ME,RI, VT] Sporobolus heterolepis A. Gray (14) Sporobolus neglectus Nash (14) Tripsacum dactyloides L. (14) Trisetum melicoides (Michx.) Scribn. (14) POLEMONIACEAE Polemonium van-bruntiae Britton (14) POLYGALACEAE Polygala senega L. (14) ME HH |I|E}S1| HT A] |SH| I 2|E|S1| E|S1| [1 GRank = G3; Fed. code = 3C. I2IT|S1| NH HII HH HI JIJE|S 1 II || |SH| HH VT I 3|E|S1| I 1] |S] HII |H| |SH| |8/T|S2| 112) |S2S3| MA HII 3| WL|SE?| |H] |SX| [2|E|S1| IE|S1| |] |SX| RI HI |H|C|SH| HII Ht |6|T|S1| HT CT |H|SC *|SH| |5|SC|S2| S|E|S1| |H|SC *|SH| |5| |S2| II HII 2|E|S1| aes vIOPOUuY 86 190A] NAME Polygala verticillata L. (11) Polygala verticillata var. ambigua (11) [CT,ME,NH,RI] POLYGONACEAE Oxyria digyna (L.) Hill (14) Polygonum douglasii Greene (14) Polygonum erectum L. (14) Polygonum glaucum Nutt. (14) Polygonum puritanorum Fern. (11) Polygonum setaceum Baldw. var. interjectum Fern. (11) Polygonum iad var. seftaceum IND. IND. ME NH VT MA RI a [+?||SU| II|SU| |?| |S2| H+|WLIS3S4] [2/C|S1| I|| SU] This taxon includes var. ambigua (P. ambigua of Gleason and Cronquist 1991) and var. isocycla. Most states have not differentiated between the varieties. More field study needed. HT [3|T|S1| I [1 | nan [4|T|S1| [S{T|S1| S|E|S1| HT HT HT [+2] |SU| |H|E|SH| 1] |S4| [+] |S3?| I |H] [SH Although historically ie aie this species appears to be declining in New England. Current status unkno a | i | III [III |~40|WL|S3] [3/T|S1| on *|SH| ank = G3. Massachusetts has the majority of New England occurrences eS a I} | I} II [SO|SC|S3]_ | HY |SH| IH = G3Q. Many current treatments place this under P. persicaria (as do ee and Cronquist 1991) which is common and non-native. Treated as Persicaria maculosa in Maine checklist (Campbell et al. 1995) and considered non-native HII HII HII |6|SC|S2|— |H|SH|SH] || | Not always distinguishable from P. hydropiperoides according to Gleason and Cronquist (1991). More field study needed. 19661 dOOdHN— TP 19 ourysy pur yorquinig ELS NAME Polygonum tenue Michx. (14) Polygonum viviparum L. (14) Persicaria vivipara (1) [ME] Rumex occidentalis S. Wats. (14) Rumex fenestratus (11) [MA] POLYPODIACEAE Cheilanthes lanosa (Michx.) D. C. Eaton (12) PONTEDERIACEAE Heteranthera reniformis Ruiz & Pavon (14) Zosterella dubia (Jacq.) Small (14) Heteranthera dubia (11) [(CT,MA,ME,NH,VT] ME NH VT MA RI Cr |H| |SH| |H|E|SH| |1| |S1|* False || |SU| |+| |S3| Disjunct occurrence in Chittenden County, Verm |IE|S1| [1[TIS1 |H| |SH| HT HT HT HT HIT |H] |SH| II|SE| III HI Introduced in Massachusetts, but native populations are historic in Vermont. Reported in Gleason and Cronquist (1991) from Maine, but the specimen was misidentified. HII HII HII III HII [TJE|S1| II HII II HT Bee |H] |SH| Recent reports unverified and not re-located. [4[E|S1|* [IJE|S1| [+||S3| [3+|WL|S2S3] | ||| [+||S3| Disjunct in Penobscot and Hancock Counties, Maine. Reported to be more common than records show in Berkshire County, Massachusetts. vIOPOYY 86 TOA] NAME Div PORTULACACEAE Montia fontana L. (14) 2 POTAMOGETONACEAE Coleogeton filiformis (C.H. Persoon) 2 Les & Haynes ssp. alpinus (M.N. Blytt) Les & Haynes (18 Potamogeton filiformis var. alpinus (1) [MA,ME,NH Potamogeton filiformis var. borealis (14) 0¥ 7) Coleogeton filiformis (C.H. Persoon) 2 Les & Haynes ssp. occidentalis (J.W. Robbins) Les & Haynes (18) Potamogeton filiformis var. occidentalis (1) [ME] Potamogeton confervoides Reichb. (14) 1 Potamogeton diversifolius Raf. (14) IND. Potamogeton hillii Morong (14) ] ME NH VT [12} |S2| IT Ht [9|[S2| [ESI] [3||S1| |6||S2| IIT 1 |14|T|S2| {10+| |S2S4| | 13] |S2| GRank = G3G4; Fed. code = C2. |?| |SU| HI II II Status uncertain. It HII |30| |S3| GRank = G3. MA Hh |H|[SX| HH |5+| |SU| |22|SC|S3| RI nae HT I |H| |SH| I HIT HII |H|SC *|SH| |H|SC *|SH| EJS 1| dOOdAN-— [8 19 ‘oyrysypy pure yorquinig [9661 NAME Potamogeton ogdenii Hellquist & Hilton ( Potamogeton pusillus L. ssp. gemmiparus Robbins (14) Potamogeton strictifolius Ar. Benn. (14) Potamogeton vaseyi Robbins (14) PRIMULACEAE Primula laurentiana Fern. (14) Primula mistassinica Michx. (14) PTERIDACEAE Cryptogramma stelleri (S. G. Gmelin) 4) Prantl (1 PYROLACEAE Pterospora andromedea Nutt. (14) 3% ME NH VT MA RI CT ITI I} [2| |S1| 1] |S1| I} || [SU| GRank = Gl. |10| |SU| \6|T}S2| I |19||S3S42| || |SH | | |H|SC *|SH| GRank = G5T3T4. More field work needed to determine current status. |H] |SH| HI |?| |SU| [12] |S]| HIT |IJE|S 1 Similar to closely related species. Questions remain on the identification of certain populations. 2/E|S1| 3/T|S2| [6] |S2| JWLISI III |H|SC *|SH| [14] |S2| HIT Ht HT na HII I+] |S3| LIT [S5/T|S1|* HII HI HII Disjunct occurrences in Caledonia and Orleans Counties, Vermont. [2\T|S1|* |6|T|S1| [+] |S3| [5|T]S2| HI 2|E|S1| Disjunct in Somerset, Piscataquis and Oxford Counties, Maine. HT |] [Sx] 2|E|S1| HT HIT HH vIOPOUY 86 LOA] NAME Pyrola minor L. (14) RANUNCULACEAE Anemone multifida Poiret (14) Hydrastis canadensis L. (14) Ranunculus allegheniensis Britton (14) Ranunculus ambigens S. Wats. (14) Ranunculus gmelinii DC. var. hookeri (D. Don) L. Benson (14) Ranunculus gmelinii var. purshii (1) [ME] Ranunculus hispidus Michx. (14) Ranunculus lapponicus L. (14) IND. ME NH VT MA RI CT | | 1S3?| || |SU| |IJE|S1|* HI HII HII Occurrences in Chittenden County, Vermont, as well as those in eastern New York state, are disjunct from the rest of this species’ range. [7T|S 1 HT IEIS1| HII HH ITI LH HI 2|E|S1| 2\E|S1| HH 2|E|S1| HII [III 8|T|S2 [IWLISI] | [SU |7| |S2| |] [SH |HJE|SH| HT |] [SH |] |SH] [EJS 1 |4|T|S1| HT HI HT [1 HH HI I [2| |S1| | | [SU] HT Ht Future editions of FNA likely will show that var. hispidus and var. caricetorum are in New England. Gleason and Cronquist (1991) show only var. caricetorum here. The Maine checklist (Campbell et al. 1995) shows var. nitidus as existing statewide. Clarification needed. J6{TISIS2) TIT HI HH HT HT 19661 dOOdAN— IP 19 ‘outa pur yorquunig NAME Ranunculus micranthus Nutt. (14) Trollius laxus Salisb. (14) Trollius laxus ssp. laxus (15) [CT] RHAMNACEAE Ceanothus herbaceus Raf. (14) ROSACEAE Agrimonia parviflora Aiton (14) Amelanchier nantucketensis Bickn. (11) Crataegus mollis (T. & G.) Scheele (14) Crataegus x silvestris Sarg. (14) Crataegus bicknellii (31) [MA] Crataegus chrysocarpa var. bicknellii (11) IND. IND. ME NH ee MA RI Cr IT HT HI [4[T|S1| [1|T|S1| [6| |S2S3| HT HH HT HI HT S|E|S1| GRank = G4T3Q. GRank is for synonym T. /axus ssp. laxus. ITT ITI [IJE|S1 || |SE| IIT HT IT HIT HT 3JE|S1| IIT |6|SC|S3| [11] [S2| HII IIT ISOISCIS3|— [ITI || |SUJ GRank = G3Q; Fed. code = C2. Reported from Connecticut, but no specimens seen. Massachusetts has the majority of occurrences globally. |H?| |SU| HIT [Hj [SH| [| |SU| HII HII Difficult taxonomic group. The identity of New England records for this species needs verification. HII HII I |IJE|S1| HII HIT GRank = G1Q. GRank is for synonym C. bicknellii. Difficult taxonomic group; status unclear. Listed as a hybrid of C. pruinosa x C. punctata in Gleason and Cronquist (1991). PIOPOYY 86 1OA] NAME Geum peckii Pursh (14) Geum vernum (Raf.) T. & G. (14) Potentilla pensylvanica L. var. bipinnatifida (Douglas) T.&G. (14) Potentilla pensylvanica vat. pectinata (1) [ME,VT] Potentilla pectinata (11) [NH] Potentilla robbinsiana Oakes (14) Prunus alleghaniensis T.C. Porter (14) Prunus maritima Marsh. var. gravesii (Small) G. J. Anderson (15) Rosa acicularis Lindley ssp. sayi (Schwein.) W. H. Lewis (14) Rosa acicularis (14) [MA,NH,VT] IND. nN ME NH VT MA RI CT HII |+|T|S2 HII HII HII HII GRank = G2Q. Cronquist (Gleason and Cronquist 1991) reports this for Maine, but we have not seen specimens, nor is it in the Maine checklist (Campbell et al. 1995). HI I |] |SH| HH I HA |+||S4| II|SU] TES 1| HII HII HII Current status in New England is unclear. Some Vermont occurrences may be disjunct. Further field work needed. HT [2|E|S1| I IH II HT GRank = G1; Fed. code = LE. HT eh nan | | |SE| HT Fed. code = C2. Introduced in Massachusetts, but historic occurrences in Connecticut are thought to have been native. HI HI [I I HI AJE|SI| | GRank = G4T1Q. Correct status of this taxon is uncertain. |H?| |SU| |H|E|SH| 2\E|S1| IJE|S1| HH I |H|SC *|SH| dOOdAN— [® 19 ‘JJoysys| pure yorquin.ig [9661 NAME Rosa blanda Aiton var. glabra Crépin (15 Rosa johannensis (14) Rubus aculiferus Bailey (15) Rubus x aculiferus (14) Rubus cuneifolius Pursh (14) Sibbaldia procumbens L. (14) Waldsteinia fragarioides (Michx.) tt. (14) Tra RUBIACEAE Galium kamtschaticum Steller (14) Galium labradoricum (Wieg.) Wieg. (14) IND. ME NH VT MA RI cr |?| [S2+| HIT HIT IIT IIT HII GRank = GST3Q. Present on list because of GRank, but most authors now combine this taxon under R. blanda which is more common. Not tracked by Maine Natural Areas Program. HII || [SU HT ITI LT HT GRank = G2?. Listed as a putative hybrid of R. allegheniensis x R. setosa in Gleason and Cronquist (1991). Current status in New England is unknown. Appears on this list because of global rank. HII E}S1| HII || [SUI IIT |7|SC|S2| Possibly adventive in New Hampshire and Massachusetts. HII IJE|S1| IIT HII IIT HT |2|T|S1|* 3/T]S1| I+] [S4| I24|SC|S3] {IT [JE|S 1] Disjunct in Kennebec County, Maine. {<3} |SU| [2| |SU| |?| [S2S3| HII HT HI] Distributional status in New England is unclear. || |SU| |H|E|S1| 2/T|S1|* I9+|SC|S3/* ||| |H|SC *|SH| Species is disjunct in Bennington County, Vermont, and in southern Berkshire C ounty, Massachusetts. N oC ~~ RIOpOyuy 86 TOA] NAME Galium trifidum L. var. trifidum (14) Galium brevipes (11) [ME,NH,VT] SALICACEAE Populus heterophylla L. (14) Salix arctophila Cockerell (14) Salix argyrocarpa Andersson (14) Salix candida Fluegge (14) Salix cordata Michx. (14) Salix exigua Nutt. ssp. interior (Rowlee) Cronquist (14) Salix interior (11) [ME] Salix exigua (15) [CT,MA,NH,VT] Salix herbacea L. (14) IND. ME NH VT MA RI Cl |H||SU| II|SU| |H||SH| IH HI [1 Ht HT HI HT I|C|S1 I4|E|S1| [IE|S1| HH I HT I HT |IE|S1| [S{T|S1| Ht HT HT LT |1|T|S1|* HT I+] |S3] [35|WL|S4]_— ||| 15} [S3| | Disjunct in Aroostook County, Maine. |H| [SHI {1] |S1| HIT [| [SUI HII HII Gleason and Cronquist’s (1991) range for this species does not include Massachusetts, but it is included in Sorrie's (1991) draft county checklist for Massachusetts. This species is distinct from S. eriocephala. Distributional status in New England is unclear. |2\T|S1|* II|SU| [+||S3| [LO|SC|S3] TT [4|T|S1| Disjunct in Kennebec County, Maine. IE|S1| [SJTISIS2| {TT HT HT HI dOOdAN— [¥ 19 ‘yourysyy pue yoruquinig [9661 NAME Salix myricoides (Muhl.) J. Carey (14) Salix planifolia Pursh (14) Salix uva-ursi Pursh (14) SANTALACEAE Geocaulon lividum (Richardson) Fern. (14) SAURURACEAE Saururus cernuus L. (14) SAXIFRAGACEAE Saxifraga aizoides L. (14) Saxifraga cernua L. (14) Saxifraga foliolosa R. Br. (14) Saxifraga stellaris var. comosa (11) Saxifraga oppositifolia L. (14) ME [2| |S1| |LJE|S1| [2/T|S]| [10] |S2| IE|S1| HH NH HI |4|T|S2| {10+| |S2S3| [4[T|S2| HH I [IE|S1| HT HT VT I [I|T|S1| I2|E|S1| |H] |SX| [2 |S1| HH LT [5] |S1| |H] [Sx] HII HII HH HH HH I EJS 1| [III HH I [11 I HI BIE|S1| HH I HT ey) NM elopouy 86 1OA] NAME sa st paniculata Mill. (15) axifraga aizoon var. neogaea ° 1) [NH, VT] Saxifraga rivularis L. (14) SCROPHULARIACEAE Agalinis acuta Pennell (14) Agalinis neoscotica Greene (11) Agalinis purpurea var. neoscotica (14) Aureolaria virginica (L.) Pennell (14) Castilleja coccinea (L.) Sprengel (14) Castilleja septentrionalis Lindl. (14) Collinsia parviflora Dougl. (14) 2(a) ME NH VT MA RI CT [2|T|S1| I2|E|S1| [5|[S1| HT HT I [1 SJE|S1| HH HH HH HT HII HII HII SJEIS1| IJE|S 1 LJE|S1| GRank = G1; Fed. code = LE. 4|E|S1| HI HII HII HII HII GRank = G2?. HII |4|T|S2| [2| |S1|* +] [S42 [12] |S2| [+] |S3| Occurrence in Franklin County, Vermont, is disjunct. |H] |SX| H] [S| HII |H] [SX] |H|SH|SH] —[4/E|S1| Ha [2\T|S1| [AJT|S1| HT HIT HT mall numbers of plants at most occurrences are cause for concern. III HI |H] |SH| | | SE} ITI II Native occurrences are historic. Introduced occurrence in Massachusetts is also not extant. 19661 dODdAN— TP 19 ‘ours pue yorquinig tcc NAME Euphrasia disjuncta Fern. & Wieg. (14) Euphrasia oakesii Wettst. (14) Gratiola virginiana L. (14) Melampyrum lineare Desr. var. latifolium Barton (14) Melampyrum lineare Desr. var. lineare (14 Melampyrum lineare Desr. var. pectinata (Pennell) Fern. (14) Mimulus alatus Aiton (14) Mimulus moschatus Douglas (14) Mimulus ringens L. var. colpophilus Fern. IND. IND. ME NH VT MA |H] [S| ma I [1 TES 1| [ESI HIT HII HII] HII HT HII || |[SU| |H| |SH || || |SU| RI Cr HII HT HI 1 2|C|S1| I II |?| [SUI |SH| | II Most specimens not identified to the varietal level. Field work and specimen annotation needed. | | [SU || ISU] || |SUJ HI II [III | Most specimens not identified to the varietal level. Field work and specimen annotation needed. HT | | SU HH SU| [ISU] | LISUI || Most specimens not identified to the varietal level. Field work and specimen annotation needed. II HHI aan I3|E|S]| | | |SE| S|E|S1| (| |S2| [4|T|S1 HH [1] [S| HI |H] |SH| This species is introduced into some New England states; determining which occurrences are native is often difficult. |12| |S2| HT I HT HI I GRank = G5T2Q; Fed. code = C2. Taxonomic status unclear. eS) ib vIOPOYUY 86 ‘1OA] NAME Pedicularis furbishiae S. Wats. (14) Pedicularis lanceolata Michx. (14) Rhinanthus crista-galli L. (14) Rhinanthus minor (15) [MA] Schwalbea americana L. (14) Veronica catenata Pennell (14) Veronica anagallis-aquatica (1) [ME] Veronica wormskjoldii Roemer & Schultes (14) Veronicastrum virginicum (L.) Farw. (14) IND. IND. ME NH VT MA RI Cr [26|E|S2| HT HT HIT III III GRank = G2; Fed. code = LE. HIT aan HI [2|E|S 1| HIT [3] |S1| |?| [SUI {1} |[S3| | | [SU II|SE| HI | | |SE Difficult to determine which occurrences are native and which are introduced. Gleason and Cronquist (1991) state that our lowland plants are introduced and the alpine plants are native. HII IIT] HII |H| [S| HII |H|SC *|SH| GRank = G2; Fed. code = LE. In New England, this species was last seen in Massachusetts in 1963. II|SEI [11 [1] |S I2|E|S1| I HI Difficult to determine which occurrences are native and which are introduced. Gleason and Cronquist (1991) state that /. catenata hybridizes with V. anagallis-aquatica and is included in the latter taxon by some authors. TEIS1| 2\E|S 1 HH HII HI HII || |SE| ITH [IES 1] [1OJSC|S2| {III 9] [SU Difficult to determine which occurrences are native and which are introduced. 19661 dODdAN— TP 19 ‘ours pur yorquinig NAME Div ME NH VT MA RI ct SELAGINELLACEAE Selaginella eclipes W. R. Buck (12) IND. HHI HI III I III HII |2| |[SU| Species is not included in New England in FNA (Flora of North America Editorial Committee 1993), but a specimen from Connecticut was later annotated as this species. SMILACACEAE Smilax tamnoides L. (15) ? HII IIT HII HIT HT |H|SC *|SH| Smilax tamnoides var. hispida (ij iCTi Smilax hispida (14) SOLANACEAE Leucophysalis grandiflora (Hook.) 4 HI HI HIISH| HI il HI ydb. (14) GRank = G3G4. Physalis grandiflora (15) [VT] Physalis longifolia Nutt. var. IND. || |SE| \|SU} |H||SH} | | |SE| |H||SH| |H||SH|} subglabrata (Mackenzie & Bush) More field work needed to determine status. Difficult to determine which Cronquist (14 populations are native. ) Physalis subglabrata (11) [CT,NH,RI,VT eIOPOYY 86 ‘1OAI NAME SPARGANIACEAE Sparganium minimum (Hartm.) Fries (14) Sparganium natans (15) [MA] ULMACEAE Ulmus thomasi Sarg. (14) URTICACEAE Pilea fontana (Lunell) Rydb. (14) VALERIANACEAE Valeriana uliginosa (T. & G.) Rydb. (14) Valerianella radiata (L.) Dufr. (14) Valerianella radiata var. fernaldiana (11) [CT] IND. ME NH VT MA RI Os) |?| [SUI || {SUI |13|T|S2| |4|T|S1|* HII |H|SC *|SH| Disjunct in Berkshire County, Massachusetts. HH HT |H| [SH] HII III I HII Extant populations in Vermont are introduced. Native populations are historic. HII HII HII |?| |S3?| HII |4| [SU] More field study needed. Species has been overlooked in New England because of similarity with P. pumila. [10] |S2| JIEJS1] JES | HT I HT HT HT HHT I I |H|SC *|SH| d°DdAN— TP 19 ‘Jyoysysyy puv youquinig [9661 € NAME VERBENACEAE Verbena simplex Lehm. (14) VIOLACEAE Hybanthus concolor (T. Forster) Sprengel (14) Viola brittoniana Pollard (11) Viola pedatifida ssp. brittoniana (13) Viola hirsutula Brainerd (11) Viola novae-angliae House (11) Viola palmata L. (22) Viola triloba var. dilatata (11) [MA Viola palustris L. (14) Viola striata Aiton (14) IND. IND. ME NH VT MA RI cr HT Ht |H] [SH] [HE|S 1 I |H| |SH| HHT HII [1] [S| HII HII |H|SC *|SH] |H| |SH It || |6|T|S1| HII 2|E|S1| [SH] I Preliminary research indicates ee one Massachusetts population of this taxon is known as |”. brittoniana var. pectinata (GRank of G4G5T3Q as V’. brittoniana ssp. pectinata). It is genetically reer from the other Massachusetts occurrences. HII HT HIT HT HIT |H|SC *|SH] [15] |S2| HIT IT] HII HIT HII Fed. code = 3C. HH HI | | [SUI || |SU| da || [SUI The taxonomic relationship within this complex is very unc n. The nomenclature reflects this confusion. Further study neede i IJE|S1| 4[T|S2| Itt HIT HIT HIT || [SE] | | |SU| nan || |SE IIT |H|SC *|SH] | More field study needed to determine status. New England occurrences should be suspect since this species is cultivated and sometimes escapes. eIOPOYY 86 1OA] NAME Viola subsinuata Greene (22) XYRIDACEAE Xyris smal liana Nash (14) ME NH VT MA RI ay § HT | Ht [| |SU| || [SU] HII || |SU] The nomenclature of this taxon in New England (which may involve V. palmata and its varieties as synonyms) is hopelessly confusing. Clarification is desperately needed. [LJE|S1|* HII HI }+| [SUI [+] |S2| 4|E|S1| Since the Essex County, Massachusetts, occurrences are dated pre-1970, the York County, Maine, occurrence is considered to be disjunct. dOOddN— TP 19 ‘Jyoyryayy pue youquinig [9661 € 330 Rhodora [Vol. 98 ACKNOWLEDGMENTS. The production of a list of this magni- tude involves many people. First and foremost, we thank the NEPCoP Listing Committee members for their time, information, support, and patience. This Committee met regularly for almost four years to assure the accuracy and reliability of the List. Primary support for “Flora Conservanda: New England” has been provided by the National Fish and Wildlife Foundation and the Ellis L. Phillips Foundation. Additionally, we thank the An- drew W. Mellon Foundation, the Jessie B. Cox Charitable Trust, and the Surdna Foundation for their support of NEPCoP. Staff of the Maine Natural Areas Program, the New Hampshire Natural Heritage Inventory Program, the Vermont Nongame and Natural Heritage Program, the Massachusetts Natural Heritage and Endangered Species Program, the Rhode Island Natural Her- itage Program, and the Connecticut Natural Diversity Data Base have supported this effort by supplying information that is the basis for this List. Leslie Sneddon and Steve Buttrick of The Nature Conservancy’s Eastern Regional Office, as well as Larry Morse and Kelly Watson of the International Headquarters of TNC, have generously supplied GRanks and other information for taxa included on this List. Many botanists have contributed time and advice, especially on nomenclatural matters: Susan G. Aiken, Kelly W. Allred, Lo- ran C. Anderson, George W. Argus, Peter W. Ball, Harvey E. Ballard, Fred R. Barrie, David E. Boufford, Christopher S. Camp- bell, Judith M. Canne-Hilliker, J. Richard Carter, Paul M. Catling, Ross C. Clark, Steven E. Clemants, Garrett E. Crow, Thomas FE Daniel, Brian E. Dutton, Michael Donoghue, Patrick E. Elvander, Barbara Ertter, John E. Fairey, Wayne R. Ferren, Robert W. Freckmann, Shirley A. Graham, Craig C. Greene, Erich Haber, David W. Hall, Jon Hamer, Ronald L. Hartman, Steven L. Hatch, Robert R. Haynes, Harold R. Hinds, Peter C. Hoch, Walter C. Holmes, Rick Kesseli, Robert Kral, Sylvia Kelso, Eric Lamont, David E. Lemke, Donald H. Les, Walter H. Lewis, Niall Mc- Carter, Landon E. McKinney, John McNeill, Richard S. Mitchell, David Moore, Michael O. Moore, John K. Morton, Robert F C. Naczi, John B. Nelson, Ching-I Peng, C. Thomas Philbrick, James B. Phipps, Robert A. Price, James S. Pringle, Anton A. Reznicek, Velva E. Rudd, David Schroeder, Alfred E. Schuyler, Fayla Schwartz, John C. Semple, Charles J. Sheviak, Teresa Sho- lars, Leila M. Shultz, Neil Snow, Robert J. Soreng, Janet R. Sul- 1996] Brumback and Mehrhoff, et al—NEPCoP 331 livan, W. Carl Taylor, Edward E. Terrell, John Thieret, David M. Thompson, Rahmona A. Thompson, Gordon C. Tucker, Florence S. Wagner, Warren H. Wagner, Warren L. Wagner, Donna M. E. Ware, and Alan S. Weakley. The authors apologize to anyone who offered advice but was inadvertently omitted from the Acknowl- edgments. Steve Young of the New York Heritage Program helped greatly by providing location data from that state. Guy Jolicoeur of Le Centre de Donnees sur le Patrimonie Nature] du Quebec provided location data from the Province of Quebec. Judy Unger of the Flora of North America Project office at the Missouri Botanical Garden patiently supplied names and tele- phone numbers of many authors of treatments for future FNA volumes. Judy Warnement and other staff of the Harvard University Her- baria helped in finding many of the publications necessary for clarification of nomenclatural and taxonomic issues. Mike O’Neal of BG-Base is thanked for his patience and good humor during the many hours spent improving the database for- mat. Additional support for the production of this List has been given by the State Geological and Natural History Survey of Connecticut, the New England Wild Flower Society, and the De- partment of Ecology and Evolutionary Biology, University of Connecticut. Gregory J. Anderson and Ellie DeCarli of the Uni- versity of Connecticut, as well as Richard Hyde of the Connect- icut Department of Environmental Protection, are thanked for their patience and support. Kelly Slater of NEWFS is thanked for her tireless recording of our meetings, and for keeping the Conservation Director in line. Barbara Pryor, Sarah Schonbrun and David DeKing of NEWFS are thanked for their assistance and encouragement. George McCully is thanked for his support for NEPCoP and for devising the title for the NEPCoP List. We gratefully acknowledge Susi von Oettingen of the U.S. Fish and Wildlife Service office in Concord, New Hampshire, for al- lowing us to use the facilities at her office for many meetings. The Concord, New Hampshire, D’ Angelo’s delivery service per- sonnel who did not steal our pens are thanked; those who did are thanked for the humor they supplied during sometimes tedious meetings. — to wn £ Nn ON > o No oO — — No a & an NO Rhodora [Vol. 98 LITERATURE CITED . CAMPBELL, C. S., H. P ADAMS, P. ADAMS, A. C. DIBBLE, L. M. EASTMAN, S.C. GAWLER, L. L. GREGORY, B. A. GRUNDEN, A. D. HAINES, K. JONSON, S. C. Rooney, T. E VINING, J. E. WEBER, AND W. A. WRIGHT. 1995, Checklist of the Vascular Plants of Maine, 3rd revision. Josselyn Botan- ical Society of Maine, Bulletin 13, and Maine Agricultural and Forest Experiment ape Bulletin 844. University of Maine, Orono, CHURCH, G. L. AND R. L. CHAMPLIN. 1978. me and Endangered ‘Vas- cular Plant Species in Rhode Island. The New England Botanical Club cooperation with the U.S. Fish and Wildlife Service (Region 5, New- ton Corner, MA). » CODDINGTON, J. AND K. G. Fie_p. 1978. Rare and Endangered Vascular Plan t Species in Massachusetts. The New England Botanical Club in cooperation with the U.S. Fish and Wildlife Service (Region 5, Newton Corner, MA) . CONNECTICUT DEPARTMENT OF ENVIRONMENTAL PROTECTION. ene Con- necticut’s Endangered, Threatened and Special Concern Species. Con- necticut Natural SV SUY Data Base, Connecticut Geological an Natural History Survey. Hartfor T. . COUNTRYMAN, W. D. 1978. Rare and Endangered Vascular Plant Species in Vermont. The New England Botanical Club in cooperation with the U.S. Fish and Wildlife Service (Region 5, Newton Corner, MA) . CRONQUIST, ARTHUR. 1981. An pee A sate of Scania of w Yor Flowering Plants. Colirabia University Pre Crow, G. 3 7 D, COUNTRYMAN, G. L. Cu IRCH, L. ee EASTMAN, Cy. Bi HELLQUIST, L. [L. J.] MEHRHOFF, AND I. M. StorKs. 1981. Rare and endangered a plant species in New England. Rhodora 83:259—299, OWHAN, JOsEPH J. 1979. Preliminary Checklist of the Vascular Flora of Connecticut (Growing Without Cultivation). State Geological and Nat- ural History Survey of Connecticut, Report of Investigations No. 8. Hart- ford, CT. . EasTMAN, L. M. 1978. Rare and sept aees Vascular Plant Species in Maine. The New England Botanical Club in cooperation with the U.S. Fish and Wildlife Service (Region 5, Newton Corner, MA). . ENSER, RICHARD W. 1996. Rare native plants of aes Island. The Rhode Island Natural Heritage Program. Providence . FERNALD, MERRITT L. 1950. Gray’s Manual of een “a ed. (corrected rk printing, 1970). Van Nostrand Company, New Yo . FLORA OF NORTH crm EDITORIAL COMMITTEE, ed. 1993. Flora of North America. Vol. 1 and 2. Oxford University Press, New . Future treatments of Flora North America. Taxonomic ae shared by author(s) of future treatments of FNA with Leslie J. Mehrhof . GLEASON, HENRY AND ARTHUR CRONQUIST. 1991. Manual of tale Plants of Northeastern United States and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx Y. . KArTESZ, JOHN T. 1994. A Syannymized Checklist of the Vascular Flora of the United States, Canada, and Greenland, 2nd ed. Vol. 1 and 2. Tim- ber Press, Portland, OR 1996] Brumback and Mehrhoff, et al—NEPCoP 333 16. _ ~ —_ [o,e) AND ROSEMARIE KARTESZ. 1980. A Synonymized Checklist of the Vascular Flora of the United States, Canada, and Greenland. Volume I, The Biota of North America. The University of North Carolina Press, Chapel Hill, NC : ee E. 1986. Biosystematic investigations in the family of duck- weeds (Lemnaceae). Vol. 2. The family of Lemnaceae—a monographic He Veroffentlichungen des Geobotanischen Institutes der ETH, Stif- tung Rtibel, Zurich, Switzerland. . Les, D. H. 1994. Molecular systematics and taxonomy of lake cress (Neobeckia aquatica; Brassicaceae), an imperiled aquatic mustard. Love, D. AND H. LIkETH. 1961. oe gaspense, a new species of arrowgrass. Canad. J. Bot. 39: 1261-1272. LUER, CARLYLE A. 1975. The Native Orchids of ree United States and Canada. The New York Botanical Garden, Bronx . MASTER, LAWRENCE L. 1991. Assessing oe a a priorities for 559-563. se acne Conservation Biol. 5: NDON E. 1992. A taxonomic revision of the sa ae . McKINN blue coke ot of North America. Sida, Botanical Miscellany, No. . MEHRHOFF, L. J. 1978. Rare and Endangered Vascular Plant eee in se da The ee England Botanical Club in cooperation with the U.S. Fish and Wildlife Service (Region 5, Newton Corner, MA). ——. 1980. Connecticut’s Endangered Species Progiani. Rhodora 82: 141-144. . MULLIGAN, G. 995. Synopsis of the genus ae (Brassicaceae) in Canada, a. a Greenland. Rhodora 97: 100—16 . NEW ENGLAND WILD FLOWER SoclrEtTy. 1992. New Sad Plant Con- seats Program. Wild Flower Notes 7(1). Framingham, 3W HAMPSHIRE NATURAL HERITAGE INVENTORY. 1995. Plant tracking a » November mses Unpublished. Concord, NH. . RHOADS, A. EF AND W. M. KLEIN, JR. 1993. The Vascular Flora of Penn- sylvania: pede Checklist and Atlas. American Philosophical Soci- ety, Philadelphia, . SEYMOUR, FRANK CONKLING. 1969. The Flora of New England. Charles E. Tuttle Company, Inc., 19 Rutland, V . SMITH, S. G. 1995. New combmnone in North American Schoenoplec- tus, Bolboschoenus, Isolepis, and Trichophorum (Cyperaceae). Novon 5: 97-102 . SORRIE, BRUCE A. 1991. County Checklist of Massachusetts Plants (Draft). Unpublished. . STATE OF MASSACHUSETTS. 1992. Massachusetts Endangered Species Act Regulations. 321 CMR: 10.00—10.60. Division of Fisheries and Wildlife. pp. 100-113 . STORKS, I. M. AND G. E. Crow. 1978. eae and Endangered Vascular Plant Species in New Hampshire. The New England Botanical Club in cooperation with the U.S. Fish and Wildlife Service (Region 5, Newton Corner, MA ). . THE NATURE CONSERVANCY. 1996. Computer printout of globally rare n VA. plant taxa in New England. Unpublished. Arlingto 334 Rhodora [Vol. 98 35. THE NATURE CONSERVANCY, CONSERVATION SCIENCE DIVISION, IN ASSO- CIATION WITH THE NETWORK OF NATURAL HERITAGE PROGRAMS AND CON- SERVATION DATA CENTERS. 1997, Working Draft Element Occurrence Data Standard. Arlington, V 36. U.S. FISH AND WILDLIFE SERVICE. 1996. Endangered and Threatened Wildlife and Plants: Notice of Final Decision on Identification of Can- didates for Listing as Endangered or Threatened (SO CFR Part 17). Fed- eral Register. pp. 64481-64485. APPENDIX 1. STATE STATUS CODES. Connecticut: (Connecticut Department of Environmental Protection 1993). Public Act 89-224 E = Endangered—any native species documented by biological research and inventory to be in danger of extirpation throughout all or a significant portion of its range within Connecticut and to have no more than five occur- rences in the state, and any species determined to be an “‘endangered species” pursuant to the federal Endangered Species T = Threatened—any native species documented by biological research and inventory to be likely to become an endangered species within the fore- seeable future throughout all or a significant portion of its range within Con- necticut and to have no more than nine occurrences in the state, and an species determined to be a “‘threatened species’ pursuant to the federal En- dangered Species Act, except for such species determined to be endangered by the Commissioner in accordance with section 4 of Public Act 89-224. SC = Special Concern—any native plant species or any native nonhar- vested wildlife species documented by scientific research and inventory to have a naturally restricted range or habitat in the state, to be at a low pop- ulation level, to be in such high demand by man that its unregulated taking yuld be detrimental to the conservation of its population. SC * = extirpated from the state. Maine: Maine Revised Statutes Annotated 5 MSRA C, 383, sub C. HI, arti- cles ndangered—any native plant that is in danger of extinction through- out all or a significant portion of its range within the State or any species determined to be an endangered species pursuant to the United States En- gt oe Species Act of 1973, Public Law 93-205, as amende = Threatened—any species of native plant es to ae an endan- ee species within the foreseeable future throughout all or a significant portion of its range or any species of plant saetenre! to be a threatened species pursuant to the federal Endangered Species Act of 1973 as amended. Massachusetts: eee of Massachusetts 1992). Massachusetts Endangered Species Act, MGL c. 131A and its regulations, 321 CMR 10.00. E = En Sean y species of plant or animal in danger of extinction throughout all or a significant portion of its range and species of plants or animals in danger of extirpation as eer by biological research and inventory 1996] Brumback and Mehrhoff, et al—NEPCoP ee) T = Threatened—any species of plant or animal likely to become an en- dangered species within the foreseeable future throughout all or a significant portion of its range and any species declining or rare as determined by bio- logical research and inventory and likely to become endangered in the fore- seeable fut = Sp ecial Concern—any species of plant or animal which has been documented by biological research and inventory to have suffered a decline that could threaten the species if allowed to continue unchecked or that occurs in such small numbers or with such a restricted distribution or specialized habitat requirements that it could easily become threatened within Massachu- setts. WL = Watch List—species with no legal standing, but considered by the state botanist to be sufficiently uncommon to be monitored in the field and studied further for possible listing (or relisting in some cases) under the Mas- sachusetts ESA regulations. New Hampshire: (New ree Natural Heritage Inventory 1995). State law RSA 217-A:3, HI, passed in a = Endangered (Note: this state code is actually SE, State Endangered. It has been changed to E in this list for consistency.)—all species in New Hampshire determined to be endangered as defined by RSA 217-A:3, III, or native plants documented as having three or fewer occurrences in the state within the last 50 years, or plants with more than three occurrences which are, in the judgment of se aauaus especially vulnerable to extirpation T = Threatened (Note: this state code is actually ST, State Threatened. It has been changed to T in this list for consistency.)—all species occurring in New Hampshire determined to be a threatened species as defined by RSA 217-A:3, XII or Federal candidate species as defined by Res-N 306.01 oc- curring in New Hampshire which are not listed on the endangered species listing as contained in Res-N 306.02 or native plants documented as having ten or fewer natural occurrences within the last 20 Nail or are otherwise threatened by extirpation due to habitat loss or other factor = Special Concern—plant species not threatened or coueanere but listed under state law as Special Concern Plant Species because they may be subject to commercial exploitation or overcollecting. Rhode Island: (Enser 1996). Rhode Island State Endangered Species Act, Title 20, Chapter 37-1 of the General Laws of the State of Rhode Island. E = Endangered (Note: this state code 1s actually SE, State Endangered. It has been changed to E in this list for consistency. Federally Endangered taxa, given the code FE, and Federally Threatened taxa, given the code FT by Rhode Island, are also changed to E in this list.)—in addition to the preceding federally ranked taxa, native taxa in imminent danger of extirpation from Rhode Island. These taxa may meet one or more of the following cri- teria: taxa formerly considered by the U.S. Fish and Wildlife Service for listing as Federally endangered or threatened (former C2 category see a taxon with one or two known or estimated total populations in the state; taxon apparently globally rare or threatened, estimated to occur at ae mately 100 or fewer sites range- T = Threatened (Note: this state sede is actually ST, State Threatened. It 336 Rhodora [Vol. 98 has been changed to T in this list for consistency.)—native taxa which are likely to become State Endangered in the future if current trends in habitat loss or other detrimental factors remain unchanged. In general these taxa have three to five known or estimated populations and are especially vulnerable to habitat loss C = Concern—native taxa which do not qualify under other categories but are sionally snipe auc to various factors of rarity and/or vulnerability. tate Historical—native taxa which have been documented for Rhode Island pty ee last 150 years but for which there are no extant populatio Vermont: State status as per the Vermont Endangered Species Law 10 V.S.A. Chapter 123 passed in 1991. E = Endangered. An endangered species means any species whose contin- determined to be in jeopardy. The term shall also include any species of wildlife or plant determined to be an endangered species pursuant to the ae sa ulate Species Act. Threatened. A threatened species means any species of wild flora or a whi ce appears likely within the foreseeable future to become endan- gered. That term shall also include any species of wildlife or plant determined to be a threatened species pursuant to the Federal Endangered Species Act. 5 APPENDIX I]. GLOBAL RANKS (GRanks; adapted from The Nature Conservancy 1996 and Master 1991) Ranks issued by the Nature Conservancy’s Biological Conservation Data- base. A species is given a Global Rank of G followed by a number or symbol and a variety or subspecies has a T followed by a number or symbol. (For example, for Eupatorium leucolepis var. novae-angliae, the Global Rank GST1 means that the species is secure globally, G5, but that the variety is ca imperiled global a) Gl Critically pupedics globally because of extreme rarity or because of some factor(s) making it especially vulnerable to extinction (typically 5 or fewer occurrences or very few remaining individuals or acres). G2 = Imperiled globally because of rarity or beca of some factor(s) making it very vulnerable to extinction throughout its range (typically 6 to 20 occurrences or few remaining individuals or acres). 3 = Either very rare and local throughout its range or found locally (even abundantly at some of its locations) in a restricted range (e.g., a single state or a physiographic region in the East) or because of other factors making it vulnerable to extinction throughout its range (typically 21 to 100 occur- rences). G4 = Widespread, abundant, and apparently secure globally, though it may be quite rare in parts of its range, especially at the periphery. Thus, the taxon is of song -term concern (usually 100 or more occurrences) G5 = Demonstrably widespread, abundant, and secure globally, though it may be quite rare in parts of its range, especially at the peripher G#G# = Numeric range rank: A range spanning two or more of the nu- 1996] Brumback and Mehrhoff, et al—NEPCoP 337 meric ranks. Denotes range of uncertainty about the exact rarity (for exam- re aed G? = Unranked, Element is not yet ranked globally. G#T# = For infraspecific taxa: The GRank applies to the full species and the TRank applies to the Says taxon Subrank T = Taxonomic subdivision: rank applies to a subspecies or variety. T#T# = Numeric range rank: A range spanning two or more of the numeric ranks for a variety or subspecies. Denotes range of uncertainty about the exact rarity of variety or subspecies (for example—G5T2T3). Qualifiers ? = Inexact or uncertain. (For example, G3? or G5T3? means that the numeric ranking is uncertain.) Q = Questionable taxonomy: taxonomic status is questionable; numeric rank may canes with taxonomy. (For example, G4T means that the tax- onomy, in this case of the subspecies or variety, is questionable. ) APPENDIX III. STATE RANKS (SRanks; adapted from The Nature Conservancy 1996 and Master 1991). S1 = Critically imperiled in the state because of extreme rarity or because of some factor(s) making it especially vulnerable to extirpation from the state (typically 5 or fewer occurrences or very few remaining individuals). S2 = Imperiled in the state because of rarity or because of some factor(s) making it very vulnerable to ae from the state (typically 6 to 20 occurrences or few remaining indiv ). S3 = Rare and uncommon in i state (typically 21 to 100 occurrences). 4 = Widespread, abundant, and apparently secure in state, with many occurrences, but is of long-term concern (usually 100 or more occurrences). emonstrably widespread, abundant, and secure in the state, and 2 oeae en dee present condition S#S# = Numeric range rank: A range spanning two or more of the numeric ranks. Denotes range of a ee about the exact rarity (for example— $283). S? = Unranked: not yet ranked in the state. SU = Unrankable: status uncertain; more information needed. SE = Exotic: an exotic species established in the state SR = Reported in the state but without persuasive documentation to pro- ini a basis for either accepting or rejecting (e.g., misidentified specimen) the ort. Some of these are very recent discoveries for which first-hand eaiee has yet to be received; others are old, obscure reports that are hard to dismiss ae the habitat is now destroyed. = Historical: occurred historically in the state. SX = Extirpated: believed to be extirpated from the state. Qualifier = Inexact or uncertain. 338 Rhodora [Vol. 98 APPENDIX IV. FEDERAL LISTING DESIGNATIONS. Codes used by the U.S. Fish and Wildlife Service (USFWS) for plants in this list under the U.S. Endangered Species Act of 1973 (the Act) as amended. se ppecies = Listed Endangered (Note: this Federal code is actually E, Endan- Perr It has been changed to LE in this list to distinguish from state endan germent status.) - taxa that are in danger of extinction throughout all or a oe aes of their range in the U.S. LT = ted Threatened (Note: this Federal code is actually T, Threatened. It has au changed to LT in this list to distinguish from state threatened status.) - taxa that may become endangered in the foreseeable future through- out all or a significant portion of their range in the Former Candidate Species he U.S. Fish and Wildlife Service is required to soron! species of wild- life and plants that are endangered or threatened based on the best available scientific and commercial information. As part of the program to identify species for possible listing, the as maintained a list of species re- garded as candidates for listing. Prior to 1996, there were 18 plant taxa oc- curring in New England that were considered candidates for listing. These taxa, known as Category 2 taxa, were considered taxa for which some infor- mation indicated that they might be in danger, but insufficient data on bio- logical vulnerability and threat were available to support listing. In February 1996, the USFWS published a pnw Rule which changed the definition of Candidate species, and narrowed the Candidate list to 182 taxa of plants and animals Naren No ae taxa in New England now appear on this new Candidate list which was given a Notice of final decision on December 5, 1996 mre Fish and Wildlife Service 1996). The Forme Candidate species, designated here as ‘C2,’ technically are no longer mor tored (tracked) by the USFWS, but are included in “Flora Conservanda: Ne ngland”? where eee in order that their status can be followed. C2 = Former Candidate specie Taxa no longer under Consideration by the USFWS axa that were once considered for listing as endangered but are no longer under such consideration were historically included in Category 3. Such taxa were subdivided further into three subcategories to indicate the reason for their removal from consideration. This designation of Category 3 has been discontinued under the Final Rule published in 1996 (U.S. Fish and Wildlife Service 199 2: 3A = Taxa for which the USFWS has persuasive evidence of extinction. If iced. such taxa might acquire high priority for listing. At this i however, the best available information indicates that the taxa in _ subca egory, or the habitats from which they were known, have been los 3B = Names that, on the basis of current taxonomic under oe (usually as represented in published revisions and monographs), do not represent dis- tinct taxa meeting the Act’s definition of “‘species.’’ Such supposed taxa could be reevaluated in the future on the basis of new information. 3C = Taxa that have proven more abundant or widespread than previously 1996] Brumback and Mehrhoff, et al—NEPCoP 339 believed as well as taxa that are not subject to any identifiable threat. If further research or changes in habitat indicate a significant decline in these taxa, they may be reevaluated for possible inclusion as candidates. 340 Rhodora [Vol. 98 INDEX TO THE NEPCOP LIST NEPCoP NAME FAMILY DIVISION Achillea borealis ASTERACEAE IND. Achillea millefolium var. borealis —see Achillea borealis Achillea ie ae var. nigrescens —see Achillea borealis Peete a ADIANTACEAE 2 Adiantum viridimontanum ADIANTACEAE l Agalinis acuta SCROPHULARIACEAE | Agalinis neoscotica SCROPHULARIACEAE l Agalinis purpurea var. neoscotica —see Agalinis neoscotic Agastache nepetoides LAMIACEAE 2 Agastache scrophulariifolia LAMIACEAE 2 Agrimonia parviflora ROSACEAE 2 Agrostis borealis see Agrostis mertensil Aaa mertensil POACEAE 2 Amaranthus pumilus AMARANTHACEAE 4 Amaranthus tuberculatus AMARANTHACEAE 2 Amelanchier nantucketensis ROSACEAE l Amerorchis rotundifolia ORCHIDACEAE 2 Ammophila champlainensis POACEAE IND. Amphicarpum purshil POACEAE 2 Anemone multifida RANUNCULACEAE 2 Angelica lucida APIACEAE IND. Angelica venenosa APIACEAE 4 Aplectrum hyemale ORCHIDACEAE 2 Arabis drummondii BRASSICACEAE 3:VT Arabis laevigata BRASSICACEAE ovis Arabis missouriensis BRASSICACEAE IND Arctostaphylos alpina ERI A 2 Arenaria caroliniana CARYOPHYLLACEAE 4 Arenaria glabra —see Minuartia glabra Arenaria groenlandica var. glabra —see Minuartia Arenaria (iced var. groenlandica —see uartia groenlandica Arenaria coh tla —see Moehringia macrophylla Arenaria marcescens —see Minuartia marcescens Arenaria rubella see Minuartia rubella 1996] Brumback and Mehrhoff, et al—NEPCoP 341 NEPCoP NAME FAMILY DIVISION Arethusa bulbosa ORCHIDACEAE 3:CT|MA, RI,VT Aristida basiramea POACEAE IND. Aristida purpurascens POACEAE 2 Aristida tuberculosa POACEAE 2 Aristolochia serpentaria ARISTOLOCHIACEAE 2 Armoracia lacustris —see Neobeckia aquatica Arnica lanceolata ASTERACEAE 1 Arnica mollis —see Arnica lanceolata Artemisia campestris ssp. ASTERACEAE 2 orealis Artemisia campestris var. canadensis Artemisia campestris ssp. borealis Artemisia campestris ssp. ASTERACEAE 3:VT caudata Asclepias purpurascens ASCLEPIADACEAE 2 Asclepias tuberosa ASCLEPIADACEAE 3:MA Asclepias varieg ASCLEPIADACEAE 2 Asclepias viridiflora ASCLEPIADACEAE 4 Asplenium montanum ASPLENIACEAE 2 Asplenium trichomanes- ASPLENIACEAE 2 ramosum Asplenium viride —see Asplenium trichomanes-ramosum Aster anticostensis ASTERACEAE 4 Aster co ASTERACEAE 2 Aster dumosus ASTERACEAE 3:ME Aster in ASTERACEAE 2 Aster praealtus ASTERACEAE IND. Aster ca eae i ASTERACEAE 2 Aster ptarmicoides —see Solidago ptarmicoides Aster sagittifolius ASTERACEAE 2 Astragalus alpinus var. FABACEAE It brunetianus Astragalus canadensis FABACEAE 2 Astragalus eucosmus BACEAE 4 Astragalus robbinsii var. jesupii FABACEAE 1 Astragalus robbinsii var. minor FABACEAE 2 Astragalus robbinsii var. FABACEAE 4 robbinsii Aureolaria virginica SCROPHULARIACEAE 3:VT Barbarea orthoceras BRASSICACEAE 2 342 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Betula borealis —see Betula minor titres glandu lo BETULACEAE 2 Betula r BETULACEAE ] ena minor e Betula minor Betula nana —see Betula glandulosa Betula nigra BETULACEAE 2 Betula pumila BETULACEAE 3:NH Bidens eatonii ASTERACEAE I Bidens heterodoxa ASTERACEAE IND. Bidens hyperborea ASTERACEAE 2 Bidens eg var. cathancensis ee B s hyperborea var. svensonii Bidens hyperbore var. on —see Bidens hyperbor Bidens hyperborea var. ASTERACEAE IND. svensonil Blephilia ciliata LAMIACEAE 2 Blephilia hirsuta LAMIACEAE 2 Blephilia hirsuta var. glabrata LAMIACEAE IND. Bolboschoenus maritimus CYPERACEAE 3:VT Bolboschoenus novae-angliae CYPERACEAE 2 Botrychium lunaria OPHIOGLOSSACEAE 2 Botrychium minganense OPHIOGLOSSACEAE IND. Botrychium oneidense OPHIOGLOSSACEAE IND. Botrychium rugulosum OPHIOGLOSSACEAE IND. Bouteloua curtipendula POACEAE 2 Braya humilis arene a 2 Cacalia suaveolens Sa | Calamagrostis canadensis var. POACE 4 langsdorfii Calamagrostis lacustri —see C anagrsts stricta ssp. iInexpansa Calamagrostis neglec —see Calamaro stricta ssp. stricta Calamagrostis nubi —see Cala eee canadensis var. langsdorfii Calamagrostis pickeringii POACEAE 2 Calamagrostis stricta ssp. POACEAE e inexpansa Calamagrostis stricta var. inexpansa e Calamagrostis stricta ssp. inexpansa Calamagrostis stricta ssp. POACEAE 2 stricta 1996] Brumback and Mehrhoff, et al—NEPCoP 343 NEPCoP NAME FAMILY DIVISION Callitriche hermaphroditica CALLITRICHACEAE | eee eae CALLITRICHACEAE 4 Calypso ORCHIDACEAE 3:VT pee uae CONVOLVULACEAE 2 Cardamine bellidifolia BRASSICACEAE 2 Cardamine concatenata BRASSICACEAE 3:ME Cardamine douglassii BRASSICACEAE 2 ardamine X incisa BRASSICACEAE IND. Cardamine longii BRASSICACEAE Cardamine maxima —see Car damine x maxima Cardamine X max BRASSICACEAE IND Cardamine eesti var. BRASSICACEAE 2 palustris Carex adusta CYPERACEAE Carex albicans var. emmonsii CYPERACEAE 3:VT Carex alopecoidea CYPERACEAE 2 Carex arcta CYPERACEAE 3:VT Carex atherodes CYPERACEAE 4 Carex a0 lel CYPERACEAE 2 Carex bac CYPERACEAE 3:ME Carex aie CYPERACEAE 3:ME Carex barrattii CYPERACEAE 2 Carex bicknellii CYPERACEAE IND. Carex bigelowii CYPERACEAE 3:VT Carex bushii CYPERACEAE 2 Carex buxbaumili CYPERACEAE 3:VT Carex capillaris CYPERACEAE 2 Carex capillaris ssp. capillaris —see Carex capillaris Carex capitata CYPERACEAE 2 Carex capitata ssp. arctogena arex Capitata Carex chordorrhiza CYPERACEAE 3:MA,VT Carex collinsi CYPERACEAE 2 Carex crawel CYPERACEAE 2 Carex davisii CYPERACEAE 2 Carex dioica var. gynocrates Carex gynocrates Carex eburnea CYPERACEAE 3:ME Carex emmonsit —see Carex albicans var. emmonsii Carex flaccosperma var. glaucodea —see Carex glaucodea Carex foenea —see Carex siccata 344 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Carex garberi CYPERACEAE I Carex Eger ean var. bifaria ee Carex garberi Carex glaucodea CYPERACEAE 2 Carex gracilescens CYPERACEAE 2 Carex gynocrates CYPERACEAE IND Carex livida CYPERACEAE 2 Carex livida var. radicaulis Carex livi Carex lupuliformis CYPERACEAE | Carex mitchelliana CYPERACEAE 1 Carex muhlenbergil CYPERACEAE 3:VT Carex nigromarginata CYPERACEAE 4 Carex norvegica CYPERACEAE 2 Carex oligocarpa CYPERACEAE 2 Carex oronensis CYPERACEAE l Carex polymorpha CYPERACEAE I Carex CYPERACEAE 3:ME Carex praticola CYPERACEAE 4 Carex rariflora CYPERACEAE 4 Carex recta CYPERACEAE IND. Carex richardsonii CYPERACEAE 2 Carex saxatilis YPERACEAE 2 Carex schweinitzil CYPERACEAE 1 Carex scirpoidea CYPERACEAE 2 Carex siccé CYPERACEAE aa be Carex ie aaa CYPERACEAE 3:ME Carex sterili CYPERACEAE 2 Carex striata var. brevis CYPERACEAE 2 Carex striatula CYPERACEAE 2 Carex tenuiflora CYPERACEA 2 Carex tetanica CYPERACEAE 2(a) Carex trichocarpa YPERACEAE 2 Carex vaginata CYPERACEAE 2 Carex wiegandii YPERACEA l Carex willdenowii CYPERACEAE 4 Carex woodli CYPERACEAE 4 Cassia hebecarpa ee Senna hebecarpa ie hypnoides —-- arrimanella hypnoides Castilleja coccinea SCROPHULARIACEAE 2 Castilleja septentrionalis SCROPHULARIACEAE 2(a) Ceanothus herbaceus RHAMNACEAE 2 erastium nutan CARYOPHYLLACEAE 2 Ceratophyllum echt CERATOPHYLLACEAE 3:ME 1996] Brumback and Mehrhoff, et al—NEPCoP 345 NEPCoP NAME FAMILY DIVISION Cercis canadensis CAESALPINIACEAE 4 Chamaelirium lute LILIACEAE 2 Chamaesyce elprsperna see a glyptosperma Cheilanthes faa OLY PODIACEAE 2 Chenopodium berlanderi var. boscianum e Chenopodium standleyanum Chenopodium boscianum —see ener standleyanum Chenopodium fog CHENOPODIACEAE IND Chenopodium i consiii CHENOPODIACEAE IND Chenopodium pratericola enopodium foggii Chenopodium rubrum CHENOPODIACEAE 3:ME Chenopodium standleyanum CHENOPODIACEAE IND Chrysopsis falcata —see Pityopsis falcata Chrysopsis mariana ASTERACEAE 2 Cirsium horridulum ASTERACEAE IND Coelopleurum lucidum —see Angelica lucida ap nce filiformis ssp. POTAMOGETONACEAE 2 alpin oe filiformis ssp. POTAMOGETONACEAE 2 occidentalis Collinsia parviflora SCROPHULARIACEAE 4 Convolvulus spithamaeus —see Calystegia de Corallorhiza odontorhiz ORCHIDACEAE 3:ME,NH Coreopsis rosea ASTERACEAE Cornus florida CORNACEAE 3:VT Corydalis aurea FUMARIACEAE 2 Corydalis flavula FUMARIACEAE 2 Crataegus bicknellii ee aegus x silves Crataegus chrysocarpa vat. ie knellii e Crataegus x silvestris Crataegus mollis ROSACEAE IND. Crataegus * silvestris ROSACEAE IND. Crotonopsis elliptica EUPHORBIACEAE 4 Cryptogramma stelleri PTERIDACE 3:ME Cuphea viscosissima LYTHRAC 4 Cuscuta coryli CUSCUTACEAE Z Cuscuta pentagona CUSCUTACEAE IND. — boreale —see C ynoglossum virginianum var. boreale 346 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Cynoglossum virginianum —see Cynoglossum virginianum var. boreale ape var. virginianum BO l Cynoglossum virginianum var. AGINACE oreale Cynoglossum virginianum var. BORAGINACEAE 4 virginianum Cyperus engelmannii CYPERACEAE IND. Cyperus houghtonii CYPERACEAE 2 Cyperus odoratus CYPERACEAE IND. Cypripedium arietinum ORCHIDACEAE I Cypripedium calceolus var. parviflorum —see Cypripedium parviflorum var. parviflorum Cypripedium calceolus var. pubescens —see Cypripedium parviflorum var. pubescens Cypripedium parviflorum —see Cypripedium parviflorum var. parviflorum ORCH Cypripedium parviflorum var. IDACEAE IND. makasin Cypripedium parviflorum var. ORCHIDACEAE IND. parviflorum Cypripedium parviflorum var. ORCHIDACEAE IND. ubescens Cypripedium pubescens —see Cypripedium parviflorum var. pubescens Cypripedium reginae ORCHIDACEAE 3:CT.MA Dentaria * incisifolia —see Cardamine * incisa Dentaria laciniata Cardamine concatenata Dentaria maxima —see Cardamine * maxima Deschampsia atropurpurea POACEAE 2 Descurainia incana e Descurainia richardsonii ent pinnata var. BRASSICACEAE 2 brachycarpa Descurainia richardsonii BRASSICACEAE 4 Desmodium canescens FABACEAE 2 Desmodium cuspidatum FABACEAE 2 Desmodium eee FABACEAE 2 Desmodium humifusur FABACEAE | Desmodium ol tiie FABACEAE 2 Diapensia lapponi DIAPENSIACEAE 3:VT Dicentra seem FUMARIACEAE 3:ME Dichanthelium mattamuskeetense —see Panicum mattamuskeetense 1996] Brumback and Mehrhoff, et al—NEPCoP NAME FAMILY 347 NEPCoP DIVISION Dichanthelium polyanthes —see Panicum polyanthes Dichanthelium scabriusculum —see Panicum scabriusculum Dichanthelium ase arpon Panicum sphaerocarpon Dichanthelium sphaerocarpon var. isophyllum se icum polyanthes ss ae virginiana EBENACEAE hs asiastrum X sabinifolium LYCOPODIACEAE Diphasiastrum sitchense LYCOPODIACEAE Diplachne maritima —see Leptochloa fascicularis var. maritima AS Draba arabisans BRASSICACEAE Draba cana BRASSICACEAE Draba glabella BRASSICACEAE Draba lanceolata —se raba cana Draba reptans BRASSICACEAE Dracocephalum parviflorum LAMIACEAE Drosera anglica DROSERACEAE Drosera linearis ROSERACEAE Dryopteris filix-mas DRY OPTERIDACEAE Echinodorus parvulus —see Echinodorus tenellus Echinodorus tenellus ALISMATACEAE Echinodorus tenellus var. parvulus —see Echinodorus tenellus Elatine americana ELATINACEAE Eleocharis equisetoides CYPERACEAE Eleocharis fallax CYPERACEAE Eleocharis microcarpa —see Eleocharis microcarpa var. filiculmis —_ Eleocharis microcarpa var. CYPERACEAE filiculmis Eleocharis nitida CYPERACEAE Eleocharis obtusa var. ovata —see Eleocharis ovata Eleocharis ovata CYPERACEAE Eleocharis ovata var. heurseri ee Eleocharis ovata oe haris pauciflora Eleocharis pauciflora var. fernaldu Eleocharis pauciflora var. CYPERACEAE fernaldii NN Nv Nw bd NNWYN LY 348 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Eleocharis aces ee Ele ris pauciflora var. fernaldii Eleoc ee aa CYPERACEAE 2 Eleocharis rostellata CYPERACEAE IND. Eleocharis tricostata CYPERACEAE 2 eocharis tuberculosa CYPERACEAE 3:ME Elymus mollis —see Le oe mollis var. mollis Elymus villosus POACEAE 2 Empetrum nigrun EMPETRACEAE 3:VT naa alpinum —see Epilobium anagallidifolium Epilobium anagallidifoliu ONAGRACEAE 2 Epilobium hornemanni —see Epilobium hor nemannii Epilobium hornemanni ONAGRACEAE 2 Equisetum te EQUISETACEAE IND. Eragrostis capillaris POACEAE IND. Erigeron acris var. ASTERACEAE 4 kamtschaticus Eriocaulon parkeri oo I Eupatorium album ASTERACEA 2 Eupatorium aromaticum eee 2 pene leucolepis var. ASTERACEAE ] vae-angliae Pica ene Var. ASTERACEAE IND. colpophilu Eupatorium anes var ASTERACEAE IND. rotundifolium Eupatorium sessilifolium ASTERACEAE 3:VT Euphorbia aa EUPHORBIACE IND. Euphrasia disjunc Score coe 4 Euphrasia oakesii (ce ys Euthamia galetorum ASTERACE IND. Euthamia tenuifolia var. jes noc an —see Euthamia galetor Floerkea See des LIMNANTHACEAE 2 Fuirena pumila CYPERACEAE 3:MA Galearis cocci Bilis ORCHIDACEAE 3:ME Galium brevipes —see Galium trifidum var. trifidum Galium kamtschatic RUBIACEAE IND. Galium labradoricum RUBIACEAE 3:MA,VT Galium trifidum var. trifidum RUBIACEAE Gamochaeta purpure ee si ea purpureum 1996] Brumback and Mehrhoff, et al—NEPCoP 349 NEPCoP NAME FAMILY DIVISION Gentiana amarella see Gentianella amarella Gentiana andrewsii GENTIANACEAE 2 Gentiana quinquefolia —see Gentianella quinquefolia Gentianella amarella GENTIANACEAE 2 Gentianella quinquefolia GENTIANACEAE 2 Geocaulon lividum ANTALACEAE 2 il ROSACEAE 1 Geum vernum ROSACEAE 4 Gnaphalium helleri ASTERACEAE IND. Gnaphalium ane var. mic cradenium see Gnaphalium heller Gnaphalium purpureum ASTERACEAE 2 Gnaphalium supinum ASTERACEAE 2 Gnaphalium sylvaticum easel iets IND. Goodyera oblongifolia ORCHIDACE 2 Gratiola virginiana SCR oar 2 Gymnocarpium jessoense ssp. DRYOPTERIDACEAE 4 parvulum Habenaria ciliaris —see Platanthera ciliaris Habenaria cristata —see Platanthera cristata Habenaria leucophaea —see Platanthera leucophaea var. leucophaea Hackelia americana —see Hackelia deflexa var. americana Hackelia deflexa var. americana BORAGINACEAE 2 Harrimanella hypnoides ERICACEAE 2 Helianthemum dumosum CISTACEAE 1 Heteranthera du —see Zost se ‘dubia Heteranthera reniformis arr ae AE 4 Hieracium robinsonii ASTER 1 Hieracium umbellatum ee 2 Hierochloe alpina POACEAE 2 Hippuris vulgaris HIPPURIDACEAE 2 Hudsonia tomentosa CISTACEAE 3:VT Huperzia appalachiana LYCOPODIACEAE IND. Huperzia selago LYCOPODIACEAE IND. Hybanthus concolor VIOLACEAE 2 Hydrastis canadensis a 2 Hydrocotyle verticillata APIACE 2 Hydrophyllum canadense anon eG: 2 350 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Hypericum adpressum CLUSIACEAE | Hypericum hypericoides ssp. multicaule pericum stragulum Hypericum stragulum CLUSIACEAE 9 Ilex ambigua var. montana AQUIFOLIACEAE 2 Hex glabra AQUIFOLIACEAE 3:ME Ilex montana ee Ilex ambigua var. montana Isanthus brachiatus —see Trichostema brachiatum Isoétes acadiensis ISOETACEAE | Isoétes eatonti —see SOsIes < eatoni Isoétes X eatonii ISOETACEAE IND Isoétes x aie ISOETACEAE IND. Isoétes lacustris ISOETACEAE IND. Isoétes macrospora —see Isoétes lacustris Isoétes prototypus ISOETACEAE l Isoétes riparia ISOETACEAE 2 Isoétes riparia var. canadensis —see Isoétes riparia Isotria medeoloides ORCHIDACEAE I Iva frutescens ssp. oraria —see Iva frutescens var. oraria Iva frutescens var. oraria ASTERACEAE 3:ME uglans cinerea JUGLANDACEAE IND. Juncus alpinoarticulatus —see Juncus alpinus Juncus alpinus eerie 2 Juncus biflorus JUNCACE 2 Juncus debilis satay peraies 2 Juncus oronensis —see Juncus X oronensis Juncus X oronensis JUNCACEAE IND. Juncus pervetus JUNCACEAE IND. Juncus stygius var. americanus JUNCACEAE 2 Juncus subtilis JUNCACEAE IND. Juncus torreyi JUNCACEAE 2 Juncus trifidus JUNCACEAE 3:VT Juncus vaseyl UNC a 2 Juniperus horizontalis CUPRESSACEAE 3:NH,VT Justicia americana ACANTHACEAE 4 Krigia biflora ASTERACEAE 4 Lactuca hirsuta ASTERACEAE 3:VT 1996] Brumback and Mehrhoff, et al—NEPCoP 35] NEPCoP NAME FAMILY DIVISION Lactuca hirsuta var. sanguinea —see Lactuca hirsuta Lathyrus ochroleucus FABACEAE 2 Lechea mi CISTACEAE IND Lemna valdivian EMNACEAE IND. nen fascicularis var. POACEAE 1 maritim cee repens FABACEAE 2 Lespedeza stuevei FABACEAE IND. Leucophysalis grandiflora SOLANACEAE 4 Leymus mollis var. mollis POACEAE IND. Liatris borealis —see Liatris scariosa var. novae-angliae Liatris scariosa var. novae- ASTERACEAE I angliae Lilaeopsis chinensis APIACEAE 3:ME Linum seus see Linum medium var. texanum Linum ae var. texanum LINACEAE 2 Linum sulcatum LINACEAE 2 Liparis liliifolia ORCHIDACEAE 2 Liquidambar styraciflua HAMAMELIDACEAE 2 Listera auriculata ORCHIDACEAE 1 Listera australis ORCHIDACEAE 2 Listera cordata ORCHIDACEAE 3:MA Lobelia spicata var. hirtella CAMPANULACEAE IND Loiseleuria procumbens ERI 2 Lomatogonium rotatum GENTIANACEAE 2 Lonicera dioica CAPRIFOLIACEAE 3:ME Lonicera hirsuta CAPRIFOLIACEAE 2 Lonicera pies lag CAPRIFOLIACEAE IND. Ludwigia polyca ONAGRACE 2 Ludwigia Seis ONAGRACEAE 2 Lupinus perennis FABACEAE 3:CT MA NH,RI,VT sae confusa JUNCACEAE 2 uzula spicata JUNCACEAE 2 See a alopecuroides LYCOPODIACEAE 2. Lycopodium alopec uroides —see Lycopodiella alopecuroides Lycopodium pear ei m —see Pseudolycopodiella caroliniana Lyc pares sabinifolium e Diphasiastrum * sabinifolium eee it selago —see Huperzia selago 302 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Lycopodium sitchense see Diphasiastrum sitchense Lycopus rubellus LAMIACEAE 2 ee odin palmatum LYGODIACEAE 3:CT,VT Lyonia mariana ERICACEA 4 Lythrum alatum var. alatum LYTHRACEAE IND. Magnolia virginiana MAGNOLIACEAE 2 Malaxis bayardili ORCHIDACEAE l Melampyrum lineare var. SCROPHULARIACEAE IND. latifolium Melampyrum lineare var. SCROPHULARIACEAE IND. lineare Melampyrum lineare var. SCROPHULARIACEAE IND. pectinata Melanthium hybridum LILIACEAE 4 Mertensia maritima BORAGINACEAE 3:MA Mimulus alatus SCROPHULARIACEAE 2 Mimulus moschatus SCROPHULARIACEAE 2 Mimulus ringens var. SCROPHULARIACEAE IND. colpophilus Minuartia car oliniana a caroliniana Minuartia nee CARYOPHYLLACEAE 2(a) Minuartia groenlandica CARYOPHYLLACEAE 3:VT Minuartia marcescens CARYOPHYLLACEAE I Minuartia rubella CARYOPHYLLACEAE 2 Moehringia macrophylla CARYOPHYLLACEAE 2 Monarda oe var. LAMIACEAE y) villicaulis Montia pee PORTULACACEAE 2 Morus rubra MORACEAE 2 Muhlenbergia capillaris POACEAE 2 Muhlenbergia richardsonis POACEAE 2 Muhlenbergia sobolifera 3:ME Myriophyllum pinnatu HALORAGACEAE IND. Myriophyllum verticillatum HALORAGACEAE IND. Najas guadalupen NAJADACEAE IND. Neobeckia aquatica BRASSICACEAE Zz Nuphar advena e Nuphar lutea ne advena Nuphar lutea ssp. adve YMPHAEACEAE IND. Nymphaea leibergii Cee 2 Nymphaea odorata ssp. NYMPHAEACEAE IND. tuberosa Nymphaea odorata var. tubero —see Nymphaea odorata ssp. Suiberse 1996] Brumback and Mehrhoff, et al—NEPCoP 353 NEPCoP NAME FAMILY DIVISION Ae la tetragona e Nymphaea leibergil Noma tuberosa e Nymphaea odorata ssp. tuberosa en fruticosa ONAGRACEAE IND. Omalotheca supina —see Gnaphalium supinum Omalotheca sylvatica —see Gnaphalium sylvaticum Onosmodium virginianum BORAGINACEAE 2 Ophioglossum pusillum OPHIOGLOSSACEAE 3:MA,RI,CT Ophioglossum vulgatum —see Oph se as pusillum Orchis rotundifolic —see Amer bone rotundifolia Orchis oe tabilis alearis spectabilis Oryzopsis canadensis POACEAE 4 Osmorhiza berteroi —see Osmorhiza chilensis Osmorhiza chilensi APIACEAE 2 Osmorhiza ene APIACEAE 4 aie obtusa —see Osmorhiza depauperata rac violacea OXALIDACEAE 2 Oxyria digyna POLY GONACEAE 2 Oxytropis campestris var. FABACEAE 1 johannensis Panicum amarum POACEAE 2 Panicum dichotomum ssp. mattamuskeetense nicum mattamuskeetense Panicum flexile POACEAE 2 Panicum gattingeri POACEAE 2 Panicum reek aaa —see Panicum Sadie var. pubescens Panicum aa cae POACEAE IND Panicum polyanthes POACEAE IND Panicum rigidulum var. POACEAE 2 pubescens Panicum scabriusculum POACEAE 2 Panicum sphaerocarpon POACEAE IND Panicum stipitatum P “EAE 4 Paronychia argyrocoma CARYOPHYLLACEAE 2(a) Paronychia argyrocoma var. albimontana e Paronychia argyrocoma Paronychia canadensis CARYOPHYLLACEAE 3:VT 354 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Paronychia fastigiata CARYOPHYLLACEAE IND. Paspalum laeve POACEAE 2 Paspalum orn var. POACEAE 2 psammophi Pedicularis mere lae SCROPHULARIACEAE I Pedicularis lanceolata SCROPHULARIACEAE 2 os vivipara —see Polygonum viviparum Phaseo ae polystachios FABACEAE 4 Phaseolus polystachios var. aquilonius —see Phaseolus polystachios Phleum alpinum POACEAE 2 hyllodoce caerulea ERICACEAE 2 Physalis grandiflora —see Leucophysalis grandiflora Physalis longifolia var. SOLANACEAE IND. ubglabrata Physalis subglabrata —see oo longifolia var. subglabrata Pilea fonta IND. Pinguicula agai LENTIBULARIACEAE 2 Pityopsis falca ASTERACEAE ] Platanthera ser ORCHIDACEAE 2 Platanthera cristata ORCHIDACEAE 2 Platanthera leucophaea var. ORCHIDACEAE ] leucophaea Poa arctica —see Poa pratensis ssp. alpigena Poa fernaldiana oa laxa ssp. fernaldiana Poa glauca POACEAE 2 ae laxa ssp. fernaldiana POACEAE ] a pratensis alpigena POACEAE IND. ae sic BERBERIDACEAE ye Polemonium van-bruntiae POLEMONIACEAE l Polygala senega POLY GALACEAE 2 Polygala verticillata POLYGALACEAE IND. ik aa hs as var. ambigua lygala verticillata Sees a douglas POLYGONACEAE 2 Polygonum erectum POLYGONACEAE IND. Polygonum glaucum POLYGONACEAE 1 Polygonum hydropiperoides var. setaceum see Polygonum setaceum var. interjectum Polygonum puritanorum POLY GONACEAE IND. 1996] Brumback and Mehrhoff, et al—NEPCoP 555 NEPCoP NAME FAMILY DIVISION Polygonum setaceum var. POLY GONACEAE IND. interjectum Polygonum tenue POLYGONACEAE 3:VT Polygonum viviparum POLY GONACEAE 2 Polymnia canadensis ASTERACEA 2 Populus heterophylla SALICACEAE 2 Potamogeton confervoides POTAMOGETONACEAE 1 Potamogeton diversifolius POTAMOGETONACEAE _— IND. Potamogeton ener var. alpin —see Coleo n filiformis ssp. iais Potamogeton tn var. borealis —see Coleogeton filiformis ssp. alpinus aah es filiformis var. occidentalis —see Coleogeton filiformis ssp. eon is ee hillii POTAMOGETONACEAE ] Potamogeton eee ey scrtceye 1 asses es ssp. POTAMOGETONACEAE — IND. gemmipat Be acon strictifolius POTAMOGETONACEAE _ IND. Potamogeton vaseyi POTAMOGETONACEAE 2 Potentilla pectinata —see Potentilla pensylvanica var. bipinnatifida Potentilla se ata var. ROSACEAE IND. bipinnatifida ee hie aa var. pectinata a pensylvanica var. bipinnatifida Potentilla rsa ROSACEAE I Prenanthes boo ASTERACEAE l Prenanthes x ac ASTERACEAE IND. Prenanthes racemosa ASTERACEAE 2 Prenanthes serpentaria ASTERACEAE 2 Primula laurentiana PRIMULACEAE 2 Primula mistassinica PRIMULACEAE 3:VT Prunus alleghaniensis ROSACEAE 4 runus maritima var. gravesil ROSACEAE IND. Peeucolycopodiella ee LYCOPODIACEAE 4 Psilocarya nitens — hynchospora nitens Pterospora andromedea PYROLACEAE 2 Puccinellia eae ssp. alascana uccine enella ssp. alascana Bei ee var. alaskana —see Puccinellia tenella ssp. alascana Puccinellia tenella ssp. alascana POACEAE IND. POACEAE IND. Puccinellia tenella ssp. eana 356 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Pycnanthemum clinopodioides LAMIACEAE IND. sie aaa torrei LAMIACEAE IND. Pyrola minor PYROLACEAE 3:VT a seeee ere allegheniensis RANUNCULACEAE 2 Ranunculus ambigens RANUNCULACEAE 2 Ranunculus gmelinit var. RANUNCULACEAE 2 Ranunculus gmelinii var. purshit —see Ranunculus gmelinii var. ae Ranunculus hispidus NUNCULACEAE IND. Ranunculus lapponicus Ure 2 Ranunculus micranthus RANUNCULACEAE 2 Rhexia mariana MELASTOMATACEAE 2 Rhinanthus crista-galli SCROPHULARIACEAE IND. Rhinanthus | minor ‘ nanthus crista-galli Ricdnaendion lapponicum ERICACEAE Rhododendron maximum ERICACEAE 3:ME,VT Rhododendron viscosum ERICACEAE 3:ME Rhynchospora capillacea CYPERACEAE 2 Rhynchospora inundata CYPERACEAE 2 Rhynchospora nitens CYPERACEAE 2 Rhynchospora penal CYPERACEAE 2 Ribes rotundifolium GROSSULARIACEAE IND. Rosa acicularis —see Rosa acicularis ssp. sayi Rosa acicularis ssp. sayl ROSACEAE 2 Rosa blanda var. glabra ROSACEAE IND. Rosa johannensis —see Rosa blanda var. glabra Rotala ramosior LYTHRACEAE 2 Rubus aculiferus ROSACEAE IND. Rubus X aculiferus —see Rubus aculiferus Rubus cuneifoliu ROSACEAE 2 Rumex fenestratus —see Rumex occidentalis Rumex occidentalis POLY GONACEAE 4 Sabatia campanulata GENTIANACEAE 2 Sabatia dodecandra GENTIANACEAE 4 Sabatia kennedyana GENTIANACEAE 1 Sabatia stellaris GENTIANACEAE 2 Sagina decumbens CARYOPHYLLACEAE IND. Sagina nodosa ssp. borealis CARYOPHYLLACEAE 2 Sagina nodosa var. borealis —see Sagina nodosa ssp. borealis 1996] Brumback and Mehrhoff, et al—NEPCoP a7 NEPCoP NAME FAMILY DIVISION Sagina nodosa ssp. nodosa ee IND. Sagittaria rigida ALISMATACEAE 3:ME Sagittaria subulata eens 2 Sagittaria teres ALISMATACEAE 1 Salix arctophila SALICACEAE 2 Salix argyrocarpa SALICACEAE 2 Salix candida SALICACEA 3:ME Salix cordata SALICACEAE IND. Salix exigua —see Salix exigua ssp. interior Salix exigua ssp. interior SALICACEAE 3:ME Salix herbacea SALICACEAE 2 Salix interior —see Salix exigua ssp. interior Salix myricoides SALICACEAE 2 Salix planifolia SALICACEAE 2 Salix uva-ursi SALICACEAE 2 Sanicula canadensis APIACEAE 2 Saururus cernuus SAURURACEAE 2 Saxifraga aizoides SAXIFRAGACEAE 2 Saxifraga aizoon var. neogaea e xifraga paniculata Saxifraga cernua SAXIFRAGACEAE 2 Saxifraga foliolosa SAXIFRAGACEAE 2 Saxifraga oppositifolia SAXIFRAGACEAE 2, Saxifraga paniculata SAXIFRAGACEAE 2 Saxifraga rivularis SAXIFRAGACEAE 2 eee stellaris var. comosa = axifraga foliolosa Schoe i etuberculatus CYPERACEAE 1 Sc cc: hallu CYPERACEAE 4 Schoenoplectus heterochaetus Seat oteies IND. Schoenoplectus * steinmetzti CYPE IND. Schwalbea americana oe eee 4 Scirpus ancistrochaetus CYPERACEAE I Scirpus clintonti —see Trichophorum clintonii Scirpus cylindricus —see Bolboschoenus novae-angliae Scirpus etuberculatus —see Schoenoplectus etuberculatus Scirpus hallii —see nia hallu Scirpus heterochaetu —see See anceus heterochaetus 358 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Scirpus longii CYPERACEAE | Scirpus maritimus —see Bolboschoenus maritimus Scirpus paludosus var. atlanticus —see Bolboschoenus maritimus Scirpus pendulus CYPERACEAE 3:ME Scirpus polyphyllus CYPERACEAE 2 Scirpus steinmetzit e Schoenoplectus * steinmetzii Scleria pauciflora CYPERACEAE 2 Scleria pauciflora var. caroliniana —see Scleria pauci Scleria reticularis CYPERACEAE | Scleria triglomerata CYPERACEAE 2 Scleria verticillata CYPERACEAE 4 Sclerolepis uniflora ASTERACEAE 2 Scutellaria integrifolia LAMIACE 2 Scutellaria leonardii LAMIACEAE ps Scutellaria parvula var. leonardi —see Scutellaria leonardii Scutellaria parvula var. parvula LAMIACEAE 2 Sedum rosea CRASSULACEAE 3:VT Selaginella eclipes SELAGINELLACEAE IND. Senna hebecarpa CAESALPINIACEAE 2 Shepherdia canadensis ELAEAGNACEAE 3:ME Sibbaldia procumbens ROSACEAE 2 Silene acaulis CARYOPHYLLACEAE 2 Silene acaulis var. exscapa —see Silene acaulis Silene stellata CARYOPHYLLACEAE 2 Sisyrinchitum mucronatum IRIDACEAE 2 Smilax eager ‘ Smilax tamnoides ae tamnoides SMILACACEAE 4 Smilax tamnoides var, hispida —see Smilax tamnoides Solidago calcicole —see So enti * calcicola Solidago X calcicola ASTERACEAE 4 Solidago canadensis var. ASTERACEAE IND. Solidago cutleri ASTERACEAE 2 Solidago glutinosa ssp. randii —see Solidago simplex ssp. randii var. monticola Solidago lepida var. molina —see Solidago canadensis var. subserrata 1996] Brumback and Mehrhoff, et al.—NEPCoP 359 NEPCoP NAME FAMILY DIVISION ines multiradiata var. arctica olidago cutleri i. ptarmicoides ASTERACEAE 2 Solidago rigida ASTERACEAE 2 Solidago simplex ssp. randii ASTERACEAE 3:MA,NH var. monticola —— simplex var. randii olidago ca ssp. randii var. monticola Sod nutan POACEAE 3:ME Sparganium minimum SPARGANIACEAE 3:MA Sparganium natan see Sparganium minimum Spartina cynosuroides POACEAE pi Sphenopholis nitida POACEAE 2 Sphenopholis obtu POACEAE IND. Sphenopholis aie POACEAE ps Spiranthes casei ORCHIDACEAE IND. Spiranthes < intermedia ORCHIDACEAE IND. a sancebdens as see Seen compositus var. compositus Spore: clandestinus POACEAE 4. Sporobolus compositus var. POACEAE compositus Sporobolus heterolepis POACEAE 2 Sporobolus neglectus POACEAE 2 Stachys hyssopifolia LAMIACEAE 3CL Stachys palustris ssp. pilosa —see Stachys pilosa Stachys pilosa LAMIACEAE IND. Stachys tenuifolia LAMIACEAE IND. sink pea ae var. platyphylla —see Stachys tenuifolia Seo umbellata FABACEAE 4 Suaeda americana CHENOPODIACEAE IND. Suaeda calceoliformis e Suaeda americana Saaeds maritima CHENOPODIACEAE IND. Suaeda maritima ssp. richii —see Suaeda maritima Subularia aquatica BRASSICACEAE 2 Symphoricarpos albus var. CAPRIFOLIACEAE 3:MA albus Synosma suaveolens —see calia suaveolens Taenidia renin APIACEAE 2 360 Rhodora [Vol. 98 NEPCoP NAME FAMILY DIVISION Tanacetum bipinnatum ssp. ASTERACEAE 2(a) huronense Tanacetum huronense anacetum bipinnatum ssp. huronense axacum ceratophorum ASTERACEAE IND. Taraxacum latilobum —see Taraxacum ceratophorum Tipularia discolor ORCHIDACEAE 2 Tofieldia glutinosa LILIACEAE 3:NH,VT Trichomanes intricatum HY MENOPHYLLACEAE l Trichophorum clintonii CYPERACEAE 2 Trichostema brachiatum LAMIACEAE 2 Triglochin gaspense JUNCAGINACEAE 4 sale acris var. kamtschatica Erigeron acris var. kamtschaticus ieetcan ona CAPRIFOLIACEAE Triosteum aurantiacum CAPRIFOLIACEAE 3:ME Triosteum perfoliatum CAPRIFOLIACEAE Z Triphora trianthophora ORCHIDACEAE 2(a) Tripsacum dactyloides POACEAE 2 Trisetum melicoides POACEAE 2 Trollius lax RANUNCULACEAE l Trollius roe ssp. laxus Trolhius laxus Ulmus thomasi ULMACEAE 4 Utricularia biflora LENTIBULARIACEAE 2 Utricularia fibrosa LENTIBULARIACEAE 2 Utricularia inflata LENTIBULARIACEAE IND. Utricularia resupinata LENTIBULARIACEAE 2(a) Utricularia subulata LENTIBULARIACEAE 2 Vaccinium boreale ERICACEAE l Vaccinium vitis-idaea ssp. minus —see Vaccinium vitis-idaea var. minus Vaccinium vitis-idaea var. RICACEAE 3:NH,MA minus Vahlodea atropurpurea —see Deschampsia atropurpurea Valeriana uliginosa VALERIANACEAE 2 Valerianella radiata VALERIANACEAE 4 Valerianella radiata var. ees —see faleri anella r Verbena simplex VERBENACEAE 2 Veronica Sahara -aquatica —see Veronica catenata Veronica catenata SCROPHULARIACEAE IND. Veronica wormskjoldii SCROPHULARIACEAE 2 1996] Brumback and Mehrhoff, et al—NEPCoP 361 NEPCoP NAME FAMILY DIVISION Veronicastrum virginicum SCROPHULARIACEAE IND. Viburnum nudum —see Viburnum nudum var. nudum Viburnum nudum var. nudum CAPRIFOLIACEAE 2 Viburnum prunifolium CAPRIFOLIACEAE 2 Viburnum rafinesquianum CAPRIFOLIACEAE 3:NH Viola brittoniana VIOLACEAE 2 Viola hirsutula VIOLACEAE 4 Viola novae-angliae VIOLACEAE 2 Viola palmata VIOLACEAE IND. Viola palustris VIOLACEAE 2 Viola pedatifida ssp. brittoniana —see Viola brittoniana Viola striata VIOLACEAE IND. Viola subsinuata VIOLACEAE IND. Viola triloba var. dilatata —see Viola palmata Waldsteinia fragarioides ROSACEAE 3:ME Wolffiella floridana —see Wolffiella gladiata Wolffiella gladiata LEMNACEAE 2 Woodsia alpina DRY OPTERIDACEAE 2 Xyris smalliana XYRIDACEAE 3:ME Zigadenus pris var. glaucus LILIACEAE 4 Zigadenus glau —see Zigadenus elegans var. yea Zizia aptera PIACEAE 2 Zosterella dubia Soe ecu 3:ME THE NEW ENGLAND BOTANICAL CLUB 22 Divinity Avenue Cambridge, MA 02138 The New England Botanical Club is a non-profit organiza- tion that promotes the study of plants of North America, es- pecially the flora of New England and adjacent areas. The Club holds regular meetings, and has a large herbarium of New En- gland plants and a library. It publishes a quarterly journal, RHODORA, which is now in its 99th year and contains about 400 pages per volume. Visit our web site at http://www.herbaria. harvard.edu/nebc/ Membership is open to all persons interested in systematics and field botany. Annual dues are $35.00, including a subscription to RHODORA. Members living within about 200 miles of Boston receive notices of the Club meetings. To join, please fill out this membership application and send with enclosed dues to the above address. Regular Member $35.00 Family Rate $45.00 Student Member $25.00 For this calendar year For the next calendar year ae Name Address City & State Zip Special interests (optional): THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 1996—1997: President: W. Donald Hudson, Jr., Chewonki Foundation, RR 2, Box 1200, Wiscasset, ME 04578 Vice-President (and Program Chair): David S. Conant, Depart- ment of Natural Sciences, Lyndon State College, Lyndon- ville, VT 05851 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Bi- ology, North Adams State College, North Adams, MA 01247-4100 Recording Secretary: Lisa A. Standley Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Pamela B. Weatherbee Curator of Nonvascular Plants: Anna M. Reid Librarian: Paul Somers Councillors: C. Barre Hellquist (Past President) Garrett E. Crow 1997 Matthew Hickler (Graduate Student Member) 1997 Edward Hehre, Jr. 1998 Michael J. Donoghue 1999 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora The Journal of the New England Botanical Club CONTENTS Heuchera micrantha var. macropetala (Saxifragaceae), a new variety. B. G. Shipes and E. F. Wells 365 A comparison of insular seaweed floras from Penobscot Bay, Maine, and other northwest Atlantic islands. Arthur C. Mathieson, Edward Hehre, Julie Hambrook, and James Gerweck 369 Survey of old-growth forest in Massachusetts. Peter W. Dunwiddie and Robert T. Leverett 419 Comments on the occasion of the centenary of the New England Botanical Club. Richard A. Howard 445 BOOK REVIEWS Flora of Berkshire County, Massachusetts 459 Michigan Flora, Part II 460 NEBC MEETING NEWS 463 Reviewers of Manuscripts 468 Information for Contributors 469 NEBC Membership Form 471 Index to Volume 98 473 NEBC Officers and Council Members inside back cover Vol. 98 Autumn, 1996 No. 896 Issued: August 13, 1997 The New England Botanical Club, Inc. 22 Divinity Avenue, Cambridge, Massachusetts 02138 RHODORA JANET R. SULLIVAN, Editor-in-Chief Department of Plant Biology, University of New Hampshire, Durham, NH 03824 MARGARET P. BOGLE, Managing Editor Department of Plant Biology, University of New Hampshire, Durham, NH 03824 Associate Editors HAROLD G. BROTZMAN THOMAS D. LEE CHRISTOPHER S. CAMPBELL LESLIE J. MEHRHOFF DAVID S. CONANT THOMAS MIONE GARRETT E. CROW LISA A. STANDLEY NANCY M. EYSTER-SMITH K. N. GANDHI—Latin diagnoses and nomenclature RHODORA (ISSN 0035-4902). Published four times a year (January, April, July, and October) by The New England Botanical Club, 810 East 10th St., Lawrence, KS 66044 and printed by Allen Press, Inc., 1041 New Hampshire St., Lawrence, KS 66044-0368. Periodicals postage paid at Lawrence, KS. POSTMASTER: Send _ address changes to RHODORA, P.O. Box 1897, Lawrence, KS 66044-8897. RHODORA is a journal of botany devoted primarily to the flora of North America. Monographs or scientific papers concerned with systemat- ics, floristics, ecology, paleobotany, or conservation biology of the flora of North America or floristically related areas will be considered. SUBSCRIPTIONS: $75 per calendar year, net, postpaid, in funds pay- able at par in United States currency. Remittances payable to RHO- DORA. Send to RHODORA, P.O. Box 1897, Lawrence, KS 66044- 8897. MEMBERSHIPS: a $35; Family $45; Student $25. Application form printed her NEBC WEB SITE: Information about The New England Botanical nal RHODORA is available at http://www.herbaria.harvard.edu/nebc/ BACK ISSUES: Information on availability of back issues should be addressed to Dr. Cathy A. Paris, Department of Botany, University of Vermont, Burlington, VT 05405-0086. E-mail: cparis@ moose.uvm.edu. ey Be cial acca In order to receive the next number of RHO- DORA, changes must be received by the business office prior to the first of January, April, July, or October. INFORMATION FOR CONTRIBUTORS: See contents. Submit manu- scripts to the Editor-in-Chief. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). RHODORA, Vol. 98, No. 896, pp. 365-368, 1996 HEUCHERA MICRANTHA VAR. MACROPETALA (SAXIFRAGACEAE), A NEW VARIETY B. G. SHIPES Department of Biological Sciences, Hampton University, Hampton, VA 23668 E. EF WELLS Department of Biological Sciences, The George Washington University, Washington, DC 20052 RACT. This newly described variety of Heuchera micrantha is char- acterized by the presence of widely ovate petals, a cyathiform calyx, and in general, a taller inflorescence than the other varieties of H. micrantha. Clearly different from the other varieties of H. micrantha in the shape of the flower, in the greater width of the petal, and in having fewer flowers per lateral branch of the inflorescence, it appears to be distinct at the varietal level. The cyathiform calyx is not found in any other H. micrantha variety. The exserted stigmas and stamens, the size of the flower, the leaf shape, and indumentum are characters held in common with the other varieties of H. micrantha, which place this taxon within our circumscription of the specie Key Words: Heuchera micrantha var. macropetala, Micranthae, var. nov., cyathiform calyx In their monograph on Heuchera, Rosendahl, Butters, and La- kela (1936) treated H. micrantha as a polymorphic species with five varieties and two forms. From locations in the Siskiyou (Or- egon and California) and Klamath (California) Mountains they reported the existence of a number of specimens which fit no varietal description and which they identified as transitional forms. Since the publication of their work, specimens from the same populations within these locations collected in different years have been identified as H. micrantha var. micrantha, H. micrantha var. pacifica, H. micrantha var. erubescens, and H. pilosissima. Some of these populations share a stable set of mor- phological characteristics and, in a recent revision of Micranthae and Pilosissimae of Heuchera in the Saxifragaceae (Shipes 1988), a new variety of Heuchera micrantha has been recognized in this region of northern California and southern Oregon. For clarity and bibliographic convenience, the new variety is validated, de- scribed, and briefly explained here. 365 366 Rhodora [Vol. 98 Heuchera micrantha Doug]. ex Lindl. var. macropetala Shipes & Wells, var. nov. Type: Oregon, Curry Co., Big Craggy, 12 Jun 1933, L. Leach s.n. (HOLOTYPE: ORE!; ISOTYPE: OSU!) Calycibus cyathiformibus usque ad non profunde turbinatis; pe- talis late ovatis, vix unguiculatis, cum tribus typis venationis: (1) una vena principalis cum 2—4 venis lateralibus; (2) una vena mi- dia; (3) venatio imperfecta suprabasalaris acrodroma (3-5). Leaf blade semi-orbicular to pentagonal, at least 3 leaf forms on each plant: (1) blade semi-orbicular with 7 very shallow lobes, each lobe with lateral lobes; (2) blade pentagonal, the length equal to the width, with 5—7 lobes, each lobe with lateral lobes; (3) blade pentagonal, the length greater than the width, with lobes, blade 3.5—7.0 cm from base to apex, 2.4—7.0 cm wide, main lobes somewhat spreading with crenate sublobes. In all three forms the lobe apices and teeth are cuspidate to acute apiculate (1.0 mm vein extension); leaf bases cordate; margins sparingly ciliate; upper leaf surface glabrous to sparsely strigose; lower leaf surface sparsely villous to strigose on veins; petioles 4.2—-15.0 cm long, puberulent, sometimes sparsely villous. Inflorescences 6.0—57.0 cm tall, somewhat lax, 2.3—11.0 cm broad, lateral axes with short, even puberulence, sometimes sparingly villous; pe- duncle bracts scarious, lanceolate to linear, occasionally large, similar to basal leaves in shape but never as large; bracts sub- tending floral branches herbaceous, margins divided, divisions with aristate tips. Calyces cyathiform (shaped like a small, shal- low bowl) to shallowly turbinate, 1.0—3.4 mm long and 1.3—3.6 mm wide at anthesis; free hypanthia 1.1 mm long; sepal lobes usually white with green or red tips, rounded, 0.5—1.5 mm long. Petals white, widely ovate, barely clawed, with three types of venation: (1) one main vein and 2—4 almost parallel lateral veins; (2) single mid-vein; (3) 3-5 secondary veins branching supra- basally from one main vein; 1.5—3.5 mm long, 0.4—1.4 mm wide with puberulent patches in the upper 1/3 of the blade; margins entire. Stamens at anthesis exserted 1.0—3.2 mm beyond calyx. Styles at anthesis exserted 0.1—2.5 mm from calyx, length of style and portion of the ovary above the hypanthium at anthesis 1.2— 3.9 mm. Capsules exserted, 2.5—5.0 mm long excluding persistent styles; seeds dark brown, echinate, 0.7 mm long, 0.4 mm wide. PHENOLOGY. Flowering from middle of April through middle of June. 1996] Shipes and Wells—Heuchera 367 HABITAT. Mixed evergreen forest on either limestone or serpen- tine base, in the Klamath Mountains of California and the Sis- kiyou Mountains of California and Oregon. Latitude 40—43°N. This newly described variety is characterized by the presence of widely ovate petals, a cyathiform calyx, and in general, a taller inflorescence than the other varieties of Heuchera micrantha. Based on floral, fruit, and inflorescence characters, it appears to be distinct at the varietal level. It is clearly distinct from the other varieties of H. micrantha in the shape of the flower, in the greater petal width, and in having fewer flowers per lateral branch of the inflorescence. The cyathiform calyx is not found in any other H. micrantha variety. However, the exserted stigmas and stamens, the size of the flower, the leaf shape, and indumentum are char- acters held in common with the other varieties of H. micrantha, which place this taxon within our circumscription of the species. REPRESENTATIVE SPECIMENS. OREGON: Curry Co., Bald Mt. Creek, NW ex- posure, McGribble Forest Service Camp, T33S, R24W, Sect. 20, Detling 5880 (ORE); Rogue River Canyon, rock ledges along Rogue River W of Brushy Bar, Baker 3832 (JEPS, UC); Squirrel Camp, Leach 1889 (ORE, OSC); Big Crag- gy, 6/12/33, Leach s.n. (ORE, OSU); Forest Service Rd. #333 (Powers-Agness mi N of the jct with #330, Agness vicinity, Crosby 1224 (ORE); Siskiyou National Forest, 16-18 mi W of jct of Hwy. 199 with Forest Service Rd. 3807 on 3807 (Rd. to Kalmiopsis Wilderness), Denton 2350 (osc, UW); Josephine Co., Mouse Creek on Rogue River, 2.3 mi upstream from bridge at mouth of Graves Creek, Crosby 1154 (DH, ORE, OSC); Graves Creek Rd., 1 mi E of jct of Graves Creek with Rogue River, T33S; R7W, Sect. 31, Crosby 1153 (osc); meee National Forest, 20—23 mi NW of Selma on the Bee River Rd. or 8-11 mi N of jct with Rt. 374 on Forest Service Rd. 3504, Denton 2475 (Uw); Lane Co. Ollalie Trail in Willamette National Hora T16S, RSE, Ireland 1117 (ORE); CALIFORNIA: Humboldt Co., Horse Mt N R4E, Sect. 34, Klipfel 676, 680 (usc); Little Horse Linto Creek below Grizzly Camp, Tracy 18976 (uc); Willow Creek, E fork, T6N, R4E, Sect. 15, 34 mi from US 101 along State Rt. 299, Reed 66 (Hsc); Shasta Co., Redding, 25 mi E, Hitchcock 6481 (poM, Uw); Siskiyou Co., State Rt. 299, 0.2 mi E of jet with Hennessey Rd., 4/30/72, Sawyer, s.n. (HSC); rd. between Forks of Salmon & Somes Bar, Smith 8695 (HSC); i aneice Mt. above Toehead Lake, T18N, R12W, Sect. 16, Smith, Sawyer, & O’Mear 5675 (usc); Salmon River, Creek, Klamath National Forest, Detling 6098 (ORE); Trinity Co., Burnt Ranch, 8 mi W on Underwood Mt. Rd., Spellenberg 454 (Hsc); Canyon Creek above McKay Camp, Ferlatte 65 (HSC, RSA); Gray Falls, trails above the Trinity River, Humboldt Meridian, TON, R7W, Sect. 34, Smith 8098 (HSC, RSA); Gray Falls, New River Trail, TON, ROE, Sect. 35, Lundquist 58 (Hsc); 368 Rhodora [Vol. 98 Gray Falls Campground above ty Trinity River, DeTomaso 924 (usc); Hobo Gulch Camp and vicinity (18 mi NW of Weaverville, along N fork of trail at the washout below Struce Cabin), Carter 546 (CAS); Ironside Mt. Qua TON, R6E, Sect. 35, Sawyer 2499 (usc), Sawyer 260] (HSC); White’s Bar, above State Rt. 299, Sawyer & Cole (HSC). LITERATURE CITED ROSENDAHL, C. O., EF K. BUTTERS, AND O. LAKELA. 1936. A monograph on the genus Heuchera. Minnesota Studies in Plant Sciences 2: 1—180. SuHIpes, B. G. 1988. A systematic survey of subsections Micranthae and Pilosissimae section Heuchera in the genus Heuchera (Saxifragaceae). Ph.D. dissertation, The George Washington University, Washington, DC. RHODORA, Vol. 98, No. 896, pp. 369-418, 1996 A COMPARISON OF INSULAR SEAWEED FLORAS FROM PENOBSCOT BAY, MAINE, AND OTHER NORTHWEST ATLANTIC ISLANDS ARTHUR C. MATHIESON, EDWARD J. HEHRE, JULIE HAMBROOK, AND JAMES GERWECK Department of Plant Biology and Jackson Estuarine Laboratory, University of New Hampshire, Durham, NH 03824 ABSTRACT. Ninety-seven taxa of seaweeds were collected from nine is- lands within the Penobscot Bay area of central Maine, including 25 Chloro- phyceae, 34 Phaeophyceae, and 38 Rhodophyceae. The patterns of species richness and longevity Nioaalar ond oe ares per rane are highly variable, with species richness ranging from only 4 taxa on Allen I. to 65 on Wooden Ball I. (mean = 30.8 + 23.0 taxa) Ga longevity ratios varying from 0.33— 1.0 (0.64 + 28.1). A total of 48 annuals and 49 perennials is recorded throughout the Bay, giving a longevity ratio of 0.98. This value is lower than that found at many open coastal habitats within the Gulf of Maine. An ex- amination of historical collections by J. Hooper and E S. Collins from the 1800s revealed three unusual coastal records (Stilophora rhizoides, Gracilaria tikvahiae, and Cladophora ruchingeri) and sixteen additional seaweeds that were not encountered in our study. Based upon these historical and present collections, as well as extensive sampling from more than 70 nearshore sites, a total of 139 seaweed taxa is known from the Penobscot Bay area, including 40 Chlorophyceae, 47 Phaeophyceae and 52 Rhodophyceae. In a comparison of seaweed floras from fourteen individual islands, from South Wolf I., New Brunswick, to Penikese I., Massachusetts (including a “‘composite’’ of Pe- nobscot Bay Islands), 216 seaweed taxa are documented, including 52 Chlo- rophyceae, 63 Phaeophyceae, and 101 Rhodophyceae. The highest numbers of taxa occur at Smuttynose I., Maine (136), and Penikese I., Massachusetts (131), and the lowest on Wooden Ball I., Maine (65), Monhegan I., Maine (64), and Kent I., New Brunswick (63). Floristic affinities of individual is- lands tend to mimic patterns of species richness, with Smuttynose having the highest number of shared taxa aN = 92.4 + 20.2) and Kent and Wooden Ball Islands the lowest numbers (m = 50.8 + 5.2 and 53.9 + 6.2, re- spectively). The seaweed diversity or Penikese Island and nearby environs (187 taxa) is comparable to the entire Gulf of Maine. Key Words: eos coastal islands, Penobscot Bay, Maine, New Bruns- ick, New Hampshire, Massachusetts, Gulf of Maine As noted by Hill (1923), and Pike and Hodgdon (1963), com- parative floristic investigations of islands have been grossly ne- glected within North America, even though there have been many fine studies of individual islands. Although their statement applies 369 370 Rhodora [Vol. 98 Table |. Dates and locations of insular collections within the Penobscot Bay area of mid-coastal Maine. Symbols: X = intertidal; X* = subtidal; X“" = intertidal and subtidal. Islands 1985 6/2 6/6 6/9 6/11 6/12 Allen I. Hurricane I. x x ong I. Marshall I. x McGlathery I. x** Monhegan I. Pond I. Two Bush I. x Wooden Ball I. xX to terrestrial vegetation, it is equally true for marine floras (Bow- ers 1942; Collins 1900; Doty 1948; Koetzner and Wood 1972; Lewis 1924; Scagel 1970; Sears and Wilce 1975). For example, along Maine’s extensive and indented coastline there are some two thousand islands (Platt 1996; Simpson 1987), but little atten- tion has been paid to their marine algal floras. Johnson and Skutch (1928a, b) conducted an early ecological study of inter- tidal algal associations on Mount Desert Island in Bar Harbor, Maine, including a synopsis of species composition, zonation, and ecology. It is still one of the most significant studies of its kind for any Maine island. The only floristic synopsis of multiple in- sular sites within the Gulf of Maine was conducted at the Isles of Shoals in southern Maine and New Hampshire (Mathieson and Penniman 1986b). We are aware of only one other comparative study of insular marine floras in North America, namely the Channel Islands in Southern California (Murray and Littler 1981; Murray et al. 1980). In the present study, we evaluate the floristic composition and affinities of seaweed populations from nine islands within the Penobscot Bay area of mid-coastal Maine, including contiguous nearshore Jericho Bay and offshore habitats (Table 1; Appendix). We compare these records with previous documentations of spe- cies composition from thirteen other islands, from the Bay of Fundy to southern Massachusetts (Figures 1—3; Table 2, 3). Data from 20 of these 22 islands represent our collections (Hehre et al. 1970; Mathieson and Penniman 1986b; Stone et al. 1970), 1996] Mathieson et al.—Insular Seaweed Floras 371 Table 1. Extended. 1986 1994 5/29 5/30 6/1 6/2 6/5 6/6 6/7 6/8 6/9 6/12 TAS 7/16 7/17 7/18 x Xx Me ae x x xX x X XxX x mee made in accordance with a uniform protocol for collecting and identifying seaweeds, including the establishment of extensive voucher collections (Druehl 1981). Information for the other two islands (Kent Island, New Brunswick, and Penikese Island, Mas- sachusetts) is based upon the work of others (Bowers 1942; Doty 1948; Koetzner and Wood 1972; Lewis 1924). In summary, the objectives of our study were four-fold as follows: (1) to assess the numbers and types of species from nine islands within the Penobscot Bay area of mid-coastal Maine (Appendix); (2) to eval- uate the longevity pattern of seaweed populations from the Pe- nobscot Bay area; (3) to compare the patterns of species richness among 22 individual islands, including nine from Penobscot Bay and thirteen other sites, from the Bay of Fundy, Canada, to Buz- zards Bay, Massachusetts; and (4) to compare a “composite” of Penobscot Bay taxa from nine islands with thirteen other North- west Atlantic islands. MATERIALS AND METHODS Collections of intertidal and shallow subtidal seaweeds were made at nine islands within the Penobscot Bay area (Figure 2, 3a, b). The selection of islands was based upon a variety of fac- tors, including accessibility by boat, variation in size, degree of wave exposure, and proximity to other insular and nearshore en- vironments. Four of the nine islands are located within Penobscot Bay proper (Hurricane, McGlathery, Two Bush, and Wooden Rhodora [Vol. 98 r re 69 CANADA (| SOUTH WOLF |. < CAMPOBELLO I. ° Ke: leeds EASTPORT 7 p Pa ’ MAINE ' J Ins Ss \ wy) | \ . \ 0. Pa) ‘ & a : _--"”” PENOBSCOT BAY CAPE SABLE \ \ L 44 6 PORTLAND \ ‘ GULF OF MAINE KITTERY ~ <2] 4 BOON |. z e—— |SLES OF SHOALS N.H. ae) MASSACHUSETTS CAPE COD 20km _ < ~~ BUZZARDS \ RL \Ar MARTHA'S VINEYARD CONN. ee SD \___/PENIKESE |. Figure The northeastern coast of North America, from South Wolf Island, New Brunswick, Canada, to Buzzards Bay, Massachusetts, including the mid-coastal Penobscot Bay area of Maine. Ball); three others (Long, Marshall, and Pond) occur within the contiguous Jericho Bay to the east (Figure 3b), and two (Allen and Monhegan) lie just W of Penobscot Bay proper (Figure 2, 3a). As noted by Conkling (1996), the seaward boundary for Pe- nobscot Bay is somewhat vague but it is approximated by the 50 Mathieson et al.—Insular Seaweed Floras SEARSPORT cs. PENOBSCOT RIVER . - 1996] BELFAST CASTINE —~ 3.0 km eS HARBORSIDE a MAINE CAPE ROSIER 45°N a ?, ~0 PENOBSCOT BAY 6 6 Ww BAO 6 6 Ss esd a . 4 > ROCKPORT & @ , NORTH HAVEN. Og ie) a rea oe & ° oO oa ° 8 70 & I?OP w & A 2 = lm oe +5 3 MoGLATHERY |. 20 ° FOX |. THOROFARE FOX ISLANDS ° 8 GREEN I. HURRICANE |. & *<-TWO BUSH I. rod PENOBSCOT BAY os" v MATINICUS |. @ WOODEN BALL I. y b-— 43°50'N BANTAM LEDGE [pa I. (CRIEHAVEN) GF NoNHecan I a ~ MATINICUS ROCK 69°10'W | | | Figure 2. The mid-coastal Penobscot Bay area of Maine, showing the location of Monhegan, Hurricane, Two Bush, and Wooden Ball Islands. 68°S0'W 374 Rhodora [Vol. 98 = rot (3 Tf porrciype . so pe () 43°57' Se : a 7 HOOPER |. "e % © MOSQUITO! oO =I ig THOMPSON . nN a Nag D & | L 1. . GEORGESISLANDS PECL 7 we eC & BURNT | 3.0 km 69°19" 69°16" 69°13 T I . é ' U MT. DESERT I. %, % DEER ISLE y a ° YL BASS HARBOR GREAT GOTT. e SY Sees a aw ce, "be BLACK | 94% Sy * o ° ke KH 8 44°09’ 4 of i, QO Zoe

4 x x xX x x 92.9 Choreocolax polysiphoniae Reinsch x x Xx x x x x x x x 71.4 (64.3) Clathr ae circumscriptum (Strémfelt) Foslie x xX KB &K& BK RK KX K K K X x 85.7 Coccotylus truncatus (Pallas) M. Wynne et Heine x xX X XK jK XK K XK X x x 78.6 SeIO[Y PposMeag Je[Nnsuy—'[e® Jo UOSoOTUeAI [9661 C8E Table 2. Continued. SW CA PB WB BO AP CE DU LU MA SM ST WH PI %o Corallina officinalis L x x x x x x x x x x Xx x x x x 100 Cystoclonium eae ea Batters X xX xX xX xX x xX xX xX xX xX x xX x x 100 Dasya baillouviana (S. G = Monta x eI 1 amentacea . . xX x x x xX x Xx x xX x x x 85.7 Dumontia contorta (S. Gmelin) ee x x xX x xX xX xX x xX xX xX xX xX 92.9 (85.7) rmis traillii poeta ex Batters) aime et Garbary x XxX xX XxX x x 42.9 Erythrotrichia carnea (Dillwyn) J. Agardh xX xX x x xX x 42.9 Erythrotric ‘hopeltis ciliaris Dic hael ex Hooker) Kor x x KX XxX x x x 50.0 Finbrtin dichotomum epee) ans x x 14.3 Eee iaes ae (Hudson) armichael ex Berkele x x x x xX 35.7 (28.6) Griffithsia selec Harvey ex Kiit x 7A Grinnellia ame na (C. Agardh) Harvey x 7.1 Gymnogongrus peer (Turner) J. Agar x xX x x “UX X x x 57.1 Halosacciocolax kjellmanii Lund x Wak rveye ella as (Reinsch) E Schmitz et x x x 21.4 a rubra (Sommerfelt) eneghini x x x x x x x XxX XK x x 78.6 Lithothamnion glaciale Kjellman x xX Xx XK K & XK K K K XK XK X x 92.9 Lomentaria baileyana (Harvey) Farlow x Tk Lomentaria clavellosa (Turner) Gaillon D« 7A Lomentaria orcadensis (Harvey) Collins ex W. R. Taylor x x x 21.4 P8E elopoyy 86 TOA] Table 2. Continued. SW CA PB WB BO AP CE DU LU MA SM ST WH _ PI % Mastocarpus stellatus (Stackhouse in Withering) Guiry in Guiry et al. x x xX x x xX X X K K X x 85.7 Membranoptera alata (Hudson) Stack- ouse x x x x x x xX x Xx x 71.4 Nemalion helminthoides (Velley in Withering) Batters x Xx x Xx x 35.7 Palmaria palmata (L.) Kuntze x x x > x x xX xX x xX x x x x 100 Pantoneura baerii ee et Ruprecht) Kylin x 7.1 *Petrocelis cruenta J. Agard xX xX xX xX xX xX xX xX X xX xX xX 85.7 (78.6) Peyssonnelia rosenvingit Schmitz in Rosenvinge x x x xX 28.6 Phycodrys rubens (L.) ers xX xX xX xX xX xX x x xX x xX x xX 100 (92.9) ge aa peudoceraoies (Gmelin) Ne et aylo x x Xx x x x xX xX x x 71.4 (64.3) Bas joze aoe (Kjellman) iiwel et Wegeberg x xX xX XxX x xX x 50.0 Phymatolithon lae nee (Foslie) Foslie xX xX xX x xX x 42.9 ymatolithon lenormandii (Areschoug in J. Agar x x x xX x x x x xX xX x x 85.7 (78.6) hymatolithon polsmorphm (L.) Foslie xX? 7.1 Phymatolithon rugulos dey x x 14.3 Phymatolithon tenue eee Diiwel et Wegeber X KX j%K& X XK XK X x 57.1 en see borreri (J. E. Smith) ageli ex Hauck x 7.1 Plumaria plumosa (Hudson) Kuntze xX x x x x x xX xX xX x x x xX xX 92.9 Pneophyllum fragile Kiitzing x xX xX XxX x XxX x x 571 SPIO] poamvas Ie[Nsu]— Je 1a UOSaTyIepy [9661 S8e Table 2. Continued. SW CA KI PB WB BO AP CE DU LU MA SM ST WH PI % Polyides rotundus (Hudson) Greville x x x x xX x x x x x 71.4 Polysiphonia denudata (Dillwyn) Greville arvey in Hooker x 7.1 Polysiphonia pee (Hudson) Sprengel x 7.1 Polysiphonia fibrillosa (Dillwyn) Sprengel x tA Polysiphonia flexicaulis (Harvey) Collins xX x xX x xX xX xX xX xX xX Xx 78.6 (71.4) Polysiphonia fucoides (Hudson) Greville x xX xX xX xX x x x xX x x xX 85.7 Polysiphonia harveyi J. Bailey x x x x x xX x xX x x x x 85.7 (78.6) Polysiphonia lanosa (L.) Te x x x x x x XxX X x xX XxX x Xx x 100 Polysiphonia nigra (Hudson) x x 14.3 Polysiphonia stricta (Dillwyn) Greville x x x x x x KX xX x xX x x x xX 100 Polysiphonia subtilissima Montagne x 7 Porphyra amplissima (Kjellman) Setchell et Hus x x x x xX x 42.9 (35.7) Porphyra leucosticta Thuret in Le Jolis x x x x x x x x x 64.3 Porphyra linearis Greville x xX xX 21.4 Porphyra miniata (C. Agardh) C. Agardh xX > xX xX xX xX xX xX 57.1 (50.0) Porphy ra umbiltcalis (L.) J. Agardh > xX xX Xx X x x xX xX x x xX x 100 Pterothamnion lite (Ellis) Nageli x x x 21.4 Ptilota serrata Kiitz Xx x x x x Xx x xX x XX x x x 92.9 Rhodomela cor gaa. (Hudson) Silva x x x x x Xx x x XX x XX x x x 100 hodophysema elegans (P. Crouan et rouan ex J. Agardh) P Dixon xX x xX x 28.6 Rhodophysema georgii Batters x x : Scageli. aei (Mont.) M. Wynne x x x xX x x xX x x x x 78.6 (71.4) Scinaia forcellata Bivona x 7.1 Seirospora interrupta (J. E. Smith) F Schmitz x 7.1 O8E elopoyuYy 86 TOA] Table 2. Continued. SW CA KI PB WB BO AP CE DU LU MA SM ST WH PI % Spermothamnion repens (Dillwyn) osenving x x x 23.1 Spyridia filamentosa (Wulfen) Harvey in Hooker xX 7.1 Styvlonema alsidii (Zanardini) Drew x x x 21.4 Titanoderma corallinae (P. Crouan et H. Crouan) Woelkerling, Chamberlain et ilva x x xX xX xX xX x 50.0 (42.9) Titanoderma pustulatum (Lamouroux) oelkerling, Chamberlain et Silva x Xx -X& xX XK X K XK K X x 78.6 “Trailliella intricata Batters” x xX xX x xX x x 50.0 (42.9) Total Rhodophyta taxa = 101 36 40 2] 38 21 43 55 54 48 52 44 62 51 48 67 Grand total seaweed taxa = 216 85 107 63 97 65 94 123 112 110 116 94 136 113 106 131 SPIO] PeeMvas Ie[Nsu[— |e Ja UOSaTYe [9661 L8e 388 Rhodora [Vol. 98 Table 3. Summary of seaweed taxa from nine islands within the Penob- scot Bay area of mid-coastal Maine. Key to symbols: X = a taxon’s presence; % = percent occurrence based upon all nine islands. pie to abbreviations: Islands: A I = Allen I.; H I = Hurricane I.; LI = ; M I = Marshall I., Mc I = McGlathery I.; MO I = Monhegan L.; aye Sa BI = Two Bush I.; WB I = Wooden Ball I. Longevity: Ann. = seasonal cael AAnn = aseasonal annual; Per. = perennial; PPer. = pseudoperennial. Al HI LI MI Mc!I Chlorophyta Acrochaete viridis Blidingia minima Capsosiphon fulvescens erea Chaetomorpha haetomorpha linu Chaetomorpha melagonium x x x Chaetomorpha picquotiana orochytrium cohnii ladophora sericea xX ‘Codiolum petrocelidis”’ Enteromorpha flexuosa ssp. paradoxa Enteromorpha intestinalis Enteromorpha lin Enteromorpha prolifera Monostroma grevillei Protomonostroma undulatum f. pulchrum Pseudendoclonium submarinum Rhizoclonium riparium Rhizoclonium tortuosum ongomorpha arcta Spongomorpha spinescens Ulothrix flacca Ulva lactuca Ulvaria obscura Urospora penicilliformis Total Chlorophyta taxa = 25 0 Phaeophyta ‘ ~ x a ~ * xx KM ORK MMM OX mK \O Agarum clathratum Alaria esculenta Ascophyllum nodosum x Chorda tomentosa Chordaria flagelliformis Dictyosiphon foeniculaceus ctocarpus fasciculatus ctocarpus siliculosus mmx mK x eK ~ mx xX ~*~ 1996] Mathieson et al.—Insular Seaweed Floras 389 Table 3. Extended. MOI PI TBI WBI Longevity %o xX AAnn iy ee x x AAnn Pipes x Ann 11.1 xX Per. 11.1 xX Per. 11.1 x x Per. 55.6 x x Per. 22.2 x Ann. 11.1 xX x PPer. 33.3 x Ann.? 11.1 xX Ann. 11.1 x x AAnn 22.2 xX AAnn 22.2 X AAnn 22.2 x Ann 11.1 xX Ann. 44.4 x AAnn 11.1 x AAnn 11.1 xX x xX x AAnn.? 77.8 x x Ann 33.3 x x Ann 55.5 x Ann. 22.2 x x PPer. 66.7 x x Ann 55.6 x Ann 22:2 14 1 1 21 Per. 33.3 xX xX Per. 44.4 x x xX Per. 77.8 Ann 33.3 x x Ann 33:3 xX x Ann 44.4 x Ann 11.1 xX x Ann 33.3 390 Rhodora [Vol. 98 Table 3. Continued. Al HI LI MI McI Phaeophyta Elachista fucicola x Fucus distichus ssp. distichus Fucus distichus ssp. edentatus Fucus distichus ssp. evanescens ucus spiralis Fucus vesiculosus Isthmoplea a ee minaria digitata Laminaria de Leathesia difform Melanosiphon cual Myrionema corunnae Myrionema strangulans Petalonia fascia x x Petroderma maculiforme Pilayella littoralis x x xX x x Pseudolithoderma extensum Punctaria latifolia Punctaria plantaginea x “Ralfsia clavata” Ralfsia verrucosa Saccorhiza dermatodea xX Scytosiphon simplicissimus x xX Sphacelaria cirrosa Sphaerotrichia divaricata Xx xX en rhizophorum ae KR KM OM ~ ~ KK x KK Total Phaeophyta taxa = 34 3 18 14 8 =) Rhodophyta Ahnfeltia plicata x Antithamnionella ase x Audouinella alar mx ISPUS Choreocolax polysiphoniae Clathromorphum circumscriptum Corallina o j Cystoclonium purpureum Devaleraea ramentacea a oa ne 2S xr OM ax 1996] Mathieson et al.—Insular Seaweed Floras 39] Table 3. Extended, Continued. MOI PI TBI WBI Longevity Yo x xX x x Per. 100 xX Per. 11.1 xX x Per. 44.4 xX xX Per. 44.4 xX xX Per. 22.2 x xX xX x Per. 88.9 xX xX Ann 22.2 x xX Per. 55.6 x x Per. 44.4 x Ann Kal on | Ann 11.1 xX Ann 11.1 x Ann 11.1 xX x x Ann 55.6 xX Per. 11.1 xX xX x x Ann 100 x Per. 11.1 xX Ann 11.1 Ann 11.1 xX Per.? 11.1 xX Per. 11.1 Ann 11.1 X x xX Ann 55.6 xX Per. 11.1 x Ann 33.3 xX Ann. 2222 21 7 4 23 xX x Per. 33.3 x AAnn 22.2 x Ann 11.1 x Per. 11.1 x x AAnn 22.2 x Ann 22.2 xX Per. 22.2 xX xX Per.? 22.2 xX xX Per. 66.7 xX xX xX Xx Per. 88.9 xX Per. 33:5 xX X Per. 33.3 xX xX Per. 66.7 xX xX Per. 44.4 xX xX Per. 44.4 392 Rhodora [Vol. 98 Table 3. Continued. Al HI LI MI McI Rhodophyta Dumontia contorta Gloiosiphonia capillaris Hildenbrandia Lithothamnion glaciale Mastocarpus stellatus Palmaria palmata ‘“‘Petrocelis cruenta” Phycodrys rubens Phyllophora pseudoceranoides Phymatolithon lenormandii Plumaria a Polysiphonia flexicaulis Polysiphonia harveyi Polysiphonia lanosa x Polysiphonia stricta Porphyra amplissima Porphyra miniata Porphyra umbilicalis ee mela confervoides Rhodophysema elegans Scagelia pylaisei Titanoderma corallinae 4 “Trailliella intricata” Total Rhodophyta taxa = 38 1 Grand total seaweed taxa = 97 4 45 36 24 16 > > > > > ~ xX ~ mx x ~ eK RM OM ~ xX Total no. annual taxa 1 21 17 7 3 Total no. perennial taxa 3 24 19 17 16 Longevity ratio 0.33 0.88 0.90 041 0.19 (annual/perennial taxa) % Annual taxa 25.0 47.0 47.2 29.2 18.8 % Perennial taxa 75.0 53.0 52.8 70.8 81.2 of Harvard University (FH). Most of Hooper’s collections are in the Herbarium at the Brooklyn Botanic Garden (BKL), with a few occurring at the University of New Hampshire (NHA). An exam- ination of all Penobscot Bay specimens from each of these his- torical collections was made and pertinent details are discussed in a later section. Specific information regarding the locations and shoreline di- 1996] Mathieson et al.—Insular Seaweed Floras 393 Table 3. Extended, Continued. MOI PI TBI WBI Longevity % x x x Ann. 77.8 Ann. 11.1 x x Per. 22.2 x x Per. 22.2 x x x Per. 66.7 x x x Per. 66.7 xX Per. 11.1 x Per. 11.1 x Per. 11.1 x Per. 11.1 x x Per. 22.2 x Per. 22.2 x Ann. 11.1 x x xX Per. 77.8 x x Per. 44.4 X Ann. 22.2 Ann. 22.2 x Ann. 55.6 x Per. 444 Per. 11.1 AAnn. 22.2 x er. 22.2 x Per. 11.1 29 5 5 21 64 13 10 65 26 5 5 29 (Mean = 12.7 + 10.7) 38 8 5 36 (Mean = 18.4 + 12.5) 0.68 0.63 1.0 0.81 (Mean = 0.65 + 0.28) 40.6 38.5 50.0 44.6 (Mean = 37.9 + 11.0) 59.4 615 500 55.4 (Mean = 62.1 + 11.0) mensions (at high tide) of the nine Penobscot Bay islands (Allen, Hurricane, Long, Marshall, McGlathery, Monhegan, Pond, Two Bush, and Wooden Ball) is given in the Appendix. The latter values were enumerated by tracing each island’s outline with a piece of thread on a National Oceanic Survey chart and convert- ing it via the map’s scalar (Mathieson and Penniman 1986b). As noted previously, thirteen other islands from the Bay of Fundy to 394 Rhodora [Vol. 98 southern Massachusetts also were compared (Table 2). Detailed characterizations of nine of these islands (Appledore, Cedar, uck, Lunging, Malaga, Smuttynose, Star, and White within the Isles of Shoals, plus Boon) are given by Mathieson and Penniman (1986b). Hehre et al. (1970) and Stone et al. (1970) describe South Wolf and Campobello Islands, New Brunswick; Doty (1948) and Lewis (1924) characterize Penikese Island (Buzzards Bay, Massachusetts); Koetzner and Wood (1972) describe Kent Island, New Brunswick, at the mouth of the Bay of Fundy. Two comparisons of the Penobscot Bay floras and the thirteen other Northwest Atlantic sites are given here (Tables 2—4). A ‘““composite”’ of the nine islands from the Penobscot area is used in One comparison, due to limited seasonal and subtidal collec- tions of individual islands (Table 1); a comparison with Wooden Ball Island, which shows the most diverse flora within Penobscot Bay, also is given. The number and percentage of taxa in common to these different sites are presented (Table 4), with the percent similarity (C) determined using Czekanowski’s coefficient (Bray and Curtis 1957): 2w C= a+b where ‘“‘w’’ = the number of taxa in common to both sites, ‘‘a”’ = the number of taxa at one site, and ‘‘b’’ = the number of taxa at the second site. Longevity characteristics of all seaweed taxa from the Penob- scot Bay area are enumerated according to the scheme outlined by Knight and Parke (1931). The different taxa are designated as annuals (aseasonal or seasonal), perennials, or pseudoperennials, depending upon their life span, growth, and reproductive char- acteristics (Mathieson 1989). Longevity delineations are based on a number of field studies throughout New England (Coleman and Mathieson 1975; Hehre and Mathieson 1970; Hehre et al. 1970; Mathieson and Hehre 1982, 1983, 1986; Mathieson et al. 1993; Mathieson, Hehre, and Reynolds 1981; Mathieson and Penniman 1991; Mathieson, Reynolds, and Hehre 1981; Reynolds and Ma- thieson 1975; Sears and Wilce 1975; Stone et al. 1970). Habitat description. Penobscot Bay (Figure 1) is the second largest embayment on the Atlantic Coast of the United States (Platt et al. 1996) and the largest within Maine (Apollonio 1979; 1996] Mathieson et al.—Insular Seaweed Floras 395 Conkling 1996; Doyle et al. 1970; Platt 1996; Porter 1966; Rich 1975). The Bay is located approximately 43 km northeast of Port- land. It extends northward nearly 38 km from the edge of the Gulf of Maine to the mouth of the Penobscot River (near Sears- port), encompassing hundreds of insular habitats (Figure 2, 3). In plan view, Penobscot Bay is triangular in shape, with its apex being near the mouth of the Penobscot River at Searsport and its base stretching about 19 km east-west between Port Clyde and Isle Au Haut (offshore of these sites). As noted by Conkling (1996), the precise delineation as to where the Bay ends and the Gulf of Maine begins is somewhat arbitrary, but a physical ocean- ographic boundary of sorts is the 50 fathom line that is just sea- ward of Matinicus Rock. Overall, the Bay embraces about 186 km? and has an irregular perimeter of 93 km (Doyle et al. 1970). Approximately twenty percent of the total river water entering the Gulf of Maine passes through the Penobscot watershed, which originates some 200 miles inland from the mouth of the Penob- scot River (Platt et al. 1996). Historically, the Penobscot Bay occupied the lower channel of a preglacial river (the Penobscot River), which flowed out onto the continental shelf at a time when sea levels were considerably lower than today. A rise in sea level since the retreat of the Wis- consin ice sheet (approximately 10,000 years ago) drowned the ancestral valley to form the Penobscot Bay and estuary (Kelly 1992). Because submergence affected the land surface which was modified by glacial activity, the Bay should be classified as a fjord-type habitat. The coastline displays a narrow, glacially scoured channel, a relatively uniform nearshore bottom, and a seaward shelf. As noted by Collins (1899), the Penobscot Bay area is seldom exposed to the open sea. It is guarded by a thick fringe of both large and small islands, which provides a variety of sheltered habitats; only the outer islands are exposed to heavy seas. Overall, there are hundreds of islands and seven major harbors (Belfast, Camden, Castine, Deer Isle Harbor, Rockland, Rockport, and Searsport) within the Bay (Figure 2, 3). The two largest islands are Vinalhaven and Deer Isle, which are in the central and north- eastern portions, respectively. The Fox Islands (Green, Hurricane, North Haven, and Vinalhaven, plus many smaller ones) cover an area of 6.2—7.4 km’. The islands were so named by an explorer in 1603 who was intrigued by the number of silver-gray foxes 396 Rhodora [Vol. 98 Table 4. Floristic affinities of fourteen individual islands from the Ba of Fundy, Canada, to Buzzards Bay, Masssachusetts, plus a ‘‘composite’’ of nine islands from the Penobscot Bay area of mid-coastal Maine, expressed as the number and % of shared taxa/location. Key to abbreviations: see Table 2. SW CA KI PB South Wolf > 100% Campobello 80 107 83% 100% Kent 42 52 63 57% 61% 100% Penobscot Bay 64 78 47 97 ““composite”’ 710% 16% 59% 100% Wooden Ball 52 59 42 69% 69% 66% 80% Boon 62 73 47 68 69% 713% 60% 71% Appledore 66 85 at 63% 74% 61% 73% Cedar 70 78 53 11% 71% 61% 75% Duck 67 80 52 73 69% 74% 60% 71% Lunging 69 77 53 69% 71% 59% 71% Malaga 61 73 50 68 68% 13% 64% 84% Smuttynose 76 55 69% dae 55% 713% Star 71 83 51 72% 76% 58% 74% White 67 78 49 75 70% 713% 58% 74% Penikese a1 65 49 59 47% 55% 51% 52% Mean (#) 65.5 cee | 50.8 Tat +10.8 +12.4 2 +113 Similarity 69.7 oS 62.0 135 (all islands) (%) iia +9.4 210 =102 Mean (#) 66.6 78.0 50.9 Toe +10.4 2123 +5.4 ne Glee Similarity 71.4 74.8 62.8 yj | (except Penikese) (%) +9.5 +8.3 +10.7 +8.7 1996] Mathieson et al.—Insular Seaweed Floras 397 Table 4. Extended. WB BO AP CE DU LU MA 65 100% 46 94 58% 100% 53 85 123 56% 18% 100% 56 97 112 63% 19% 83% 100% 53 93 eal L110 61% 78% 80% 82% 100% 54 1 102 94 116 60% 718% 84% 89% 83% 100% 53 85 85 88 94 67% 72% 16% 83% 83% 87% 100% 57 89 11 104 97 104 88 57% 17% 85% 84% 19% 83% 717% 56 77 94 81 54% 74% 81% 84% 82% 83% 718% 52 92 l 81 61% 76% 81% 84% 84% 83% 81% 45 72 60 46% 60% 57% 59% 56% 53% 53% 53.9 73.0 86.1 84.3 81.7 84.5 T4.5 +6.2 +13.3 +18.2 +16.4 +15.9 +18.0 +13.3 64.5 73.5 I35 77.9 76.1 76.7 76.2 Ae +918 +11.6 +10.8 +10.9 +12.2 +10.6 47.6 73.4 87.1 85.2 82.6 85.9 75.5 +18.4 #£13.7 +18.5 + 16.7 + 16.0 +17.8 +13.2 65.8 74.5 76.8 79.2 717.6 78.4 77.9 +11.5 +9.4 +10.8 +99 +99 +10.8 +8.9 398 Rhodora [Vol. 98 Table 4. Extended, continued. Grand SM ST WH PI Mean South Wolf Campobello Kent Penobscot Bay “composite” Wooden Ball Boon Appledore Cedar Duck Lunging Malaga Smuttynose 136 100% Star 106 113 85% 100% White 9 81% 82% 100% Penikese 77 69 63 131 58% 57% 53% 100% Mean (#) 92.4 83.3 80.2 67.7 Ta:2 +20.2 +16.5 +16.2 +19.1 +11.4 Similarity 75.7 76.0 76.1 57.1 72.3 (all islands) (%) +11.8 +117 +12.1 +12.5 +6.0 Mean (#) 92.4 84.3 81.4 76.8 +20.2 +16.7 +16.1 +12.7 Similarity T10 7714 Tad 74.7 (except Penikese) (%) 122 + 10.9 +10:1 +4,7 1996] Mathieson et al.—Insular Seaweed Floras 399 that lived freely in their woods and roamed the shores to catch fish (Simpson 1987). The Thorofare, a passage between North Haven and Vinalhaven (Figure 2), was a prehistoric deep valley between mountains. Ragged (Criehaven), Wooden Ball, and Ma- tinicus Islands, plus Matinicus Rock, represent the most exposed habitats seaward of the Fox Islands. Matinicus Rock (commonly called The Rock), with its lighthouse, is the outermost marker for Penobscot Bay and the coast of Maine. It consists of about 13 hectares of precipitous granitic ledges arising from the continental shelf 3.1 km seaward of Matinicus Island. The Rock serves as a rookery for Atlantic Puffin, and it is one of only a few islands in the western Atlantic where these birds nest (Porter 1966; Simpson 1987). The Fox Islands (Figure 2) divide Penobscot Bay into western and eastern basins (Doyle et al. 1970). On the northern portion of West Penobscot Bay, two deep channels (over 52 m) extend along both sides of Islesboro Island and then coalesce further seaward (to the south). Deep water habitats, therefore, are in close proximity to the land in West Penobscot Bay. Seaward of Isles- boro, several channels breach the sill between the two Bays, al- lowing east-west access via Two Bush Island Channel and be- tween Matinicus and Wooden Ball Islands near Bantam Ledge. The Bay’s western shore has numerous coastal irregularities that mark the harbor towns of Searsport, Belfast, Camden, Rockport, and Rockland. The upper half of the eastern shore is very irreg- ular, with Castine, Harborside, and Cape Rosier forming major harbors; the mid-eastern shore is characterized by numerous is- lands, the most prominent being Deer Isle and Isle Au Haut. Together, they form the boundary of East Penobscot Bay, i.e., offshore to the 50 fathom line (Conkling 1996). Penobscot Bay is underlain primarily by rocks originating from the lower Paleozoic (Doyle et al. 1970). These former sedimen- tary rocks are highly metamorphosed into complex schists and gneisses. Granitic outcrops are most common on the north and east sides. The mean tidal range varies spatially from about 1.2—3.6 m (Anonymous 1985). Tidal currents average 2.5 knots, except within upper Bay channels (between islands) where they can range from 4 to 7 knots. Because of these strong currents, water masses below 9.0 m are remarkably homogeneous, with salinities varying from 29.9—31.8 %o. As noted previously, the Penobscot River is the major source of fresh water to the Bay; it 400 Rhodora [Vol. 98 is also the second largest drainage basin within New England (Anonymous 1989), draining Hancock, Penobscot, and Waldo Counties. Its tidal limits extend inland about 22.0 km to a tidal dam in Bangor. A series of minor rivers and streams drains into the area, including the Passagassawakeag River at Belfast, the Union and Megunticook Rivers from Camden Hills, and the Ba- gaduce River from the Castine and Cape Rosier basins on the east side of the estuary. In characterizing environmental quality within Penobscot Bay, Larsen (1992) states that nickel and zinc concentrations within some sediments are moderately high (29 and 88 ppm dry weight, respectively). The concentrations of other contaminants (chro- mium, copper, lead, polychlorinated biphenyls, and polycyclic ar- omatic hydrocarbons) are elevated locally and well above prein- dustrial levels (Larsen et al. 1984a, b). For example, sediment polycyclic aromatic hydrocarbons in Penobscot Bay averaged 2,600 ppb (Johnson et al. 1985) versus 4,300 ppb for Casco Bay (Larsen et al. 1983). Surprisingly, such values are comparable to some of the most industrialized regions of the world (Larsen 1992). Typically, industrial contamination is highest near the head of the Bay, indicating that river flow is the major pathway into this marine environment. The occurrence of uniform sediment chromatographs also suggests that the major source of contami- nation is via runoff of compounds deposited within the Penobscot Bay watershed. RESULTS Species composition, distribution, and longevity patterns. Ninety-seven seaweed taxa, which include 25 Chlorophyceae, 34 Phaeophyceae, and 38 Rhodophyceae, were collected from the nine islands within the Penobscot Bay area (Table 3). Species richness per island is highly variable (Figure 4) and appears to be a function of collecting frequency (Table 1). The highest num- bers of taxa and percentages of the total flora were recorded from Wooden Ball (65 taxa, 67%), Monhegan (64 taxa, 66%), and Hurricane Islands (45 taxa, 46%) where 4 or 5 collections per site were made; the lowest numbers were recorded from Allen (4 taxa, 4%) and Two Bush Islands (10 taxa, 10%), where single collections were made. Green, brown, and red algae are most diverse on Wooden Ball (21, 23, and 21 taxa, respectively) and 1996] Mathieson et al.—Insular Seaweed Floras 401 TOTAL TAXA SW CA KI PB WB BO AP CE DU LU MA SM ST WH PI Figur The number of seaweed taxa (green, brown and red) at fourteen individual islands from the Bay of Fundy, Canada, to Buzzards Bay, Mas sachusetts, plus a “‘composite’”’ of nine islands from the Penobscot Bay area of Maine. Abbreviations: SW = South Wolf I., New Brunswick; CA = Cam- pobello I., New Brunswick; KI = Kent I., New Brunswick; PB = a ‘‘com- of nine islands from the Penobscot Bay area of Maine A Allen, Hurricane, Long, Marshall, McGlathery, Monhegan, Pond, Two Bush, and Wooden Ball Islands; WB = Wooden Ball I., Penobscot aa ME; BO = Boon I., ME; AP = Appledore I., ME; CE = Cedar I., ME; DU = Duck I., ME; LU = Lunging I., NH; MA = Malaga I., ME; SM = Smuttynose I., ME; ST = Star L, NH; WH = White I., NH; PI = Penikese I., Monhegan (14, 21, and 29 taxa), intermediate on Hurricane (9, 18, and 18 taxa) and Long Islands (8, 14, and 14 taxa), and lowest on Allen (0, 3, and | taxa). Overall, the mean number of taxa per island is very low, being 30.8 + 23.0 SD; the green algae contribute 6.9 + 7.0, the brown algae 11.4 + 7.7, and the red algae 12.4 + 9.0. As outlined in Figure 5 and Table 3, 34 of the 97 total taxa from the Penobscot Bay area (10 green, 14 brown, and 10 red algae) are restricted to a single island (11.1% occurrence), while 23 taxa (7 green, 3 brown, and 13 red algae) occur on two islands (22.2% occurrence). Thirty seaweeds occur on 4—9 islands (44.4— 100% occurrence); this includes 6 greens, 12 browns, and 12 reds. The most ubiquitous taxa are Elachista fucicola and Pilayella littoralis (100% occurrence); Fucus vesiculosus and Chondrus crispus occur at 88.9% of the sites, while Rhizoclonium tortuo- 402 Rhodora [Vol. 98 30 O green brown 20 Wired NUMBER OF TAXA oad 11.1 22.2 33.3 44.4 55.6 66.7 77.8 88.9 100 FREQUENCY (%) e 5. Frequency distribution patterns of 97 seaweed taxa recorded ae nine islands within the Penobscot Bay area. Taxa found on only one of the nine islands (11.1% occurrence) are represented by the left-most bar of the se those found on only two islands (22.2%) are represented by the next bar, etc. sum, Ascophyllum nodosum, and Dumontia contorta occur at 77.8% (Table 3). The longevity patterns of seaweeds at each of the nine Penob- scot Bay islands are summarized in Table 3. The numbers of annuals range from 29 taxa on Wooden Ball to only | on Allen Island (mean = 12.7 + 10.7 taxa); perennials vary from 38 taxa on Monhegan to 3 on Allen Island (mean = 18.4 + 12.5 taxa). The longevity ratios (annuals/perennials) at the various sites range from a maximum of | on Two Bush Island to 0.19 on McGlathery Island (mean = 0.65 + 0.28). As shown in Figure 6, the ‘“‘com- posite” flora consists of 48 annuals and 49 perennials, giving an overall longevity ratio of 0.98. With respect to individual groups, the green algae are predominantly annuals (19/25 or 76%), the brown algae are intermediate (18/34 or 53%), and the red algae exhibit the lowest percentage of annuals (11/38 or 29%). Floristic comparisons. A total of 216 seaweed taxa (52 Chlorophyceae, 63 Phaeophyceae, and 101 Rhodophyceae) ts re- corded from fourteen individual islands, ranging from the Bay of Fundy (South Wolf Island, New Brunswick) to Buzzards Bay 1996] Mathieson et al.—Insular Seaweed Floras 403 (Penikese Island), Massachusetts, including a “‘composite”’ of Pe- nobscot Bay’s insular flora (Table 2; Figure 4). The ‘“‘composite”’ data are presented due to limited seasonal and subtidal collections from individual islands within the Penobscot Bay area; collections from Wooden Ball provide the most realistic representation of an individual island within this embayment. Overall, the highest numbers of species are recorded from Smuttynose (136 taxa) and Penikese Islands (131 taxa), with Kent, Monhegan, and Wooden Ball Islands having the lowest numbers (63—65 taxa). Penobscot Bay’s “‘composite”’ insular flora (97 taxa) exceeds the last three sites, as well as those of South Wolf (85 taxa), Boon, and Malaga Islands (94 taxa each). The last-named site is the smallest island within the Shoals archipelago. The two islands having the most diverse floras within the Penobscot Bay area (Table 3), Wooden Ball (65 taxa) and Monhegan (64 taxa), have approximately the same number of taxa as Kent Island (63 taxa), while the other thirteen individual islands (Table 2) have more diverse floras. Based upon the data in Table 2, the following values for species richness (mean + SD) are calculated for the fourteen individual islands, plus the “‘composite’’ data for the Penobscot Bay area: (1) 103.9 = 21.9 taxa for fourteen individual islands, excluding Penobscot Bay’s “‘composite”’ value; (2) 106.2 + 19.0 taxa for thirteen individual islands and Penobscot Bay’s ‘“‘composite,”’ ex- cluding Wooden Ball Island as an individual; (3) 106.9 + 19.5 taxa for thirteen individual islands, excluding Penobscot Bay’s ‘““composite’” value and Wooden Ball Island; (4) 101.8 + 21.3 taxa for thirteen individual islands, excluding Penobscot Bay’s ‘““composite’’ and Penikese Island; (5) 104.3 + 18.3 taxa for twelve individual islands and Penobscot Bay ‘‘composite,”’ ex- cluding Wooden Ball and Penikese Islands; (6) 104.9 + 19.0 taxa for twelve individual islands, excluding Penobscot Bay ‘‘com- posite,’’ Wooden Ball and Penikese Islands. Thus, the values are very uniform. Figure 7 summarizes the distributional patterns of the 216 total seaweed taxa in Table 2 and is expressed as percent occurrence on fourteen individual islands, excluding the Penobscot Bay ‘““composite.”” A bimodal pattern is evident, with peak numbers occurring at one and all fourteen islands. Fifty-one seaweeds are restricted to a single island (7.1% occurrence), while 25 occur on all fourteen. The most ubiquitous taxa (100% occurrence) consist of 5 green algae (Blidingia minima, Chaetomorpha melagonium, 404 Rhodora [Vol. 98 m= 5_( SS yC€Uc_»sg SSSI Sia PENOBSCOT BAY TAXA (# & %) g SSVsggsss_o&j# 7) = CA KI PB WB BO AP CE DU LU MA SM ST WH PI >< Figure 10. Number and percentage of Penobscot Bay taxa (“‘composite’’) at fourteen individual islands from the Bay of Fundy, Canada, to Buzzards Bay, Massachusetts, including overall mean values. See Figure 4 for abbre- viations. similarity values for the other fourteen islands given in Table 4. As noted above, floristic affinities of individual islands tend to ‘““mimic”’’ patterns of species richness. The highest numbers of shared taxa occur on Smuttynose (85 taxa) and Appledore Islands (80 taxa), while Kent and Penikese Islands show the lowest num- bers (47 and 59 taxa, respectively). By contrast, the percent sim- ilarity values are highest on Malaga (84%) and lowest on Peni- kese (52%) and Kent Islands (59%). The mean number and per- cent similarity of Penobscot Bay taxa on all fourteen islands are 72.7 + 11.3 taxa and 73.5 + 10.2%, respectively. An analogous calculation for thirteen islands (minus Penikese) gives slightly higher mean values of 73.7 + 11.1 taxa and 75.1 + 8.7%. DISCUSSION Although Penobscot Bay is the largest embayment on Maine’s irregular coastline (Apollonio 1979; Doyle et al. 1970; Platt 1996; Porter 1966), its algal flora has been documented previously by only a few published records of individual taxa. For example, Taylor (1957) gives no specific records for the Bay, simply citing Maine. However, in a discussion of historical data (p. 22), he mentions the early Maine collections of Franklin W. Hooper that are deposited in BKL. A recent examination of these collections 408 Rhodora [Vol. 98 revealed that they were, in fact, collected by John Hooper during September 1851 and bequeathed to the Long Island Historical Society upon his death in 1869. Most of John Hooper’s Penobscot Bay collections are from inshore sites near Camden, Maine, while the precise locations of others are not given. The collections at BKL also include several isotypes. The fact that EK W. Hooper was born just seven months prior to the Camden collections in 1851 confirms this misinterpretation. Perhaps FE W. Hooper’s affiliation with Harvard University and W. G. Farlow, and his subsequent directorship (1889) at the Brooklyn Institute contributed to this confusion (Anonymous 1887, 1906). Eaton (1873) mentions Ceramium hooperi from Penobscot Bay, noting that it was collected by Hooper and named by W. H. Harvey for its discoverer. The name now is considered conspe- cific with C. deslongchampii (South and Tittley 1986; Taylor 1957). Seven additional taxa documented by Hooper (in NHA and BKL) have not been observed by us: Cladophora rupestris, En- teromorpha clathrata, Stilophora rhizoides, Myriotrichia clavae- formis, Gymnogongrus crenulatus, Polyides rotundus, and Grac- tlaria tikvahiae (Tables 2 and 3). With the exception of S. rhi- zoides and G. tikvahiae, which are warm-water disjuncts in the northwest Atlantic (South and Tittley 1986; Taylor 1957), the other five taxa are likely to occur on nearshore islands within Penobscot Bay (Appendix). The coastal occurrence of G. tikva- hiae and S. rhizoides in Maine is particularly noteworthy (Bour- que 1996). We have collected Gracilaria only from shallow es- tuarine habitats in northern New England: the Great Bay Estuary System of Maine/New Hampshire (Mathieson and Hehre 1986), the Salt Bay area near Newcastle, Maine, and within the upper reaches of the New Meadow River near Bath, Maine. Stilophora rhizoides has been found only within the same Salt Bay area in Newcastle. The NHA specimen of G. tikvahiae from Penobscot Bay was initially identified by J. Hooper as Gymnogongrus nor- vegicus,; there are, however, correctly identified specimens of both taxa within Hooper’s collections (BKL). Collins (1891, 1896a, b, 1899, 1901) collected at several in- shore (Camden and Cape Rosier) and insular habitats within the Penobscot Bay area (Eagle, Great Duck, Spectacle, and York Is- lands). We have found most of the taxa documented by Collins, with the following exceptions: Cladophora albida, C. ruchingeri, Gononema aecidioides, ““Hecatonema maculans,’’ Leptonematel- 1996] Mathieson et al.—Insular Seaweed Floras 409 la fasciculata, Ralfsia fungiformis, Streblonema fasciculatum, Ceramium strictum, and Erythrodermis traillii. As noted by Hoek (1982), Cladophora ruchingeri was identified initially by Collins (1899) as Cladophora gracilis forma elongata, based upon col- lections from Spectacle Island. An isotype of this taxon is de- posited in the Farlow Herbarium (Phycotheca Boreali-Americana #725). Although there are several other records of C. ruchingeri from Maine’s open coast (Hoek 1982, p. 20), our New England records are restricted to a single estuarine tidal rapids site at Do- ver Point, New Hampshire. Aside from the 97 insular (Table 3) and 16 historical records noted above, we have recently documented 26 additional taxa from more than 70 inshore Penobscot Bay habitats (Mathieson and Hehre, unpubl. data). These include 11 green algae (Bryopsis plumosa, Capsosiphon groenlandicum, Codiolum pusillum, En- teromorpha compressa, E. torta, Epicladia flustrae, Microspora pachyderma, Percursaria percursa, Prasiola stipitata, Ulothrix speciosa, and Ulvaria oxysperma), 6 brown algae (Ascophyllum nodosum ecad scorpioides, Chorda filum, Desmarestia viridis, Fucus vesiculosus ecad limicola, Hincksia granulosa, and ‘‘Ralf- sia bornetii’’), and 9 red algae (Audouinella dasyae, A. membra- nacea, Erythrotrichia carnea, Erythrotrichopeltis ciliaris, Peys- sonnelia rosenvingil, Porphyra leucosticta, Ptilota serrata, Sac- cheria fucina, and Stylonema alsidii). Thus, a total of 139 sea- weed taxa is recorded from the Penobscot Bay area, which is slightly greater than floristically diverse islands like Smuttynose (136) and Penikese Islands (131 taxa) but less than that of nearby Casco and Passamaquoddy Bays within southern Maine and New Brunswick, respectively (Collins 1911; Tittley et al. 1987). The reduced mean number of taxa per island within Penobscot Bay (30.8 + 23.0) versus the Isles of Shoals (114.0 + 11.4) should be noted (Mathieson and Penniman 1986b). While riverine contamination within Penobscot Bay (Larsen 1992; Larsen et al. 1984a, b) might be a contributing factor to this low species di- versity, several islands are considerably offshore (Appendix), di- minishing the effects of nearshore pollution. More likely, the low diversity is associated with the very sheltered nature of Penobscot Bay (Collins 1899), which would allow a dominance of selected taxa (Mathieson et al. 1991; Tittley et al. 1987; Vadas et al. 1976). Further, the occurrence of limited seasonal and subtidal collec- tions is no doubt a contributing factor (Mathieson 1979; Mathie- 410 Rhodora [Vol. 98 son and Penniman 1986a), as well as the fact that only 9 of several hundred islands were studied (Hill 1923). For the above reasons, it is premature to make any detailed statistical inferences of insular species richness within Penobscot Bay as a function of area, shoreline length or exposure to wave action (Mathieson and Penniman 1986b). For example, the lowest number of taxa 1s found on Allen Island (4 taxa) having a shoreline of approxi- mately 7620 m, while the highest number of taxa (65) is found on Wooden Ball with a smaller shoreline of about 5560 m (Table 3; Appendix). As noted previously, we are aware of only two comparative studies of insular marine floras in North America, those involving the Channel Islands in Southern California (Murray and Littler 1981; Murray et al. 1980) and the Isles of Shoals in southern Maine/New Hampshire (Mathieson and Penniman 1986b). Al- though both archipelagos contain eight islands, this is where the analogies end. The maximum distance between islands at the Shoals is very small (4.8 km) and individual islands exhibit a wide range of wave exposure and relatively uniform hydrography and substrata (Sze 1982). By contrast, the distances between the individual Channel Islands are much greater and they collectively extend over a much broader geographical range. They occur just offshore and south of Point Conception and their oceanographic and meteorological conditions are complex and spatially variable due to the proximity of the cold California Current and the warm Southern California Countercurrent (Liining 1990). Not supris- ingly, the floristic similarities of the Channel Islands are relatively low (mean = 45.4%) and the different islands contain a mixture of both northern and southern elements. Even so, the islands have their greatest affinities with the mainland south of Point Concep- tion, while individual islands exhibit a pattern of variable affini- ties to the central California flora. In contrast to such low floristic affinities and spatially variable and complex oceanographic pa- rameters within the Southern California Bight, oceanographic conditions within the Gulf of Maine are more stable spatially (Mathieson and Penniman 1986b; Mathieson et al. 1991). Such uniformity may contribute to the relatively high floristic affinities of Gulf of Maine islands as a whole (mean = 72.3 + 6.0%; Table 4). Surprisingly, an analogous comparison of the percent similar- ity of Gulf of Maine sites, minus the southernmost Penikese Is- land, shows only a slightly higher mean value of 74.7 + 4.7%. 1996] Mathieson et al.—Insular Seaweed Floras 411 Thus, floristic similarities within these Northwest Atlantic islands are much higher than those of the Channel Islands, which show maximum and minimum interisland values of 64.2 and 32.6%, respectively. In contrast to the latter situation, Scagel (1963) em- phasizes that the northeast Pacific flora is relatively uniform and exhibits a gradual transition in species composition from northern California to Sitka, Alaska. Such patterns are more comparable to those of the Gulf of Maine and Bay of Fundy (Figure 9). A comparison of the floras north and south of Cape Cod (the Gulf of Maine and Bay of Fundy versus Buzzards Bay) reveals differences in species richness and numbers of warm-temperate taxa (Liining 1990). For example, of the different insular sites discussed above (Figure 7; Table 2), only Smuttynose has more taxa (136) than Penikese (131). Amazingly, the Penikese records are based almost entirely upon summer collections from the in- tertidal and shallow subtidal zones (Doty 1948; Jordan 1874; Lewis 1924), while those from Smuttynose are based upon de- tailed seasonal collections of both intertidal and subtidal seaweeds (to the lower limits of vegetation). No doubt, enhanced seasonal and subtidal collections on Penikese would increase the number of known taxa. In a detailed seasonal study of subtidal seaweeds (from 1.2 to 41.0 m) within nearby Martha’s Vineyard, Waquoit Bay, and Buzzards Bay, Sears and Wilce (1975) record 132 taxa, consisting of 13 Chlorophyceae, 52 Phaeophyceae, and 67 Rho- dophyceae. Thus, a total of 187 taxa is recorded from Penikese and adjacent subtidal sites, which is comparable to the entire Gulf of Maine (Mathieson and Hehre 1986; Mathieson and Penniman 1986b; South and Tittley 1986). Seventy-six of these taxa are common throughout the region (6 Chlorophyta, 24 Phaeophyta, 46 Rhodophyta), while 55 are restricted to Penikese (19 Chlo- rophyta, 14 Phaeophyta, 22 Rhodophyta) and 56 are recorded only by Sears and Wilce (7 Chlorophyta, 28 Phaeophyta, 21 Rho- dophyta). Overall, the intertidal flora at Penikese has a greater number of green algae, while crustose browns are more common at the adjacent subtidal sites. In a comparison of multiple nearshore sites north and south of Cape Cod, Massachusetts, Coleman and Mathieson (1975) found an enhancement of annuals to the south, while perennials were more abundant to the north. They state that greater numbers of annuals occur where large temperature amplitudes are evident south of Cape Cod, while the opposite pattern exists with reduced 412 Rhodora [Vol. 98 temperature variations north of the Cape. Similarly, Littler and Littler (1981) state that annuals or opportunistic forms dominate in seasonally disturbed environments due to catastrophic environ- mental fluctuations, while long-lived perennials are more con- spicuous in environmentally constant habitats. Mathieson and Penniman (1986b) summarize longevity ratios (#annual/#perennial taxa) for 11 island floras and 4 nearshore habitats, from South Wolf Island, New Brunswick, to Woods Hole, Massachusetts; the values vary from 1.29 (Woods Hole) to 0.66 (Scituate), with an overall mean of 0.85 (greater # of peren- nials than annuals). The latter value (0.85) is intermediate be- tween Penobscot Bay’s “‘composite’’ flora (0.97) and the mean for the nine individual islands (0.65 + 0.28). Mathieson and Pen- niman (1986b) also note that similar values may result from dif- ferent combinations of perennials and annuals. They find a ratio of 0.95 for Smuttynose Island, an exposed offshore habitat within the Isles of Shoals, and at Dover Point, a mid-estuarine site, with the two calculations based upon values of 61 annuals/64 peren- nials and 45 annuals/47 perennials, respectively. Comprehensive seasonal and spatial collections (intertidal and subtidal) are ob- viously necessary before any meaningful comparisons of such ratios can be made. The wide range of values in Table 3 confirms the preliminary nature of the individual insular collections within Penobscot Bay. ACKNOWLEDGMENTS. We acknowledge the support of The Is- land Institute staff of Rockland, Maine, who provided logistical and boat support to one of us (JH) via their Outward Boun Program on Hurricane Island. We are indebted also to Kerry Bar- ringer, Steve Clemants, and Linda Marschener (BKL) for their in- valuable assistance with the John Hooper algal collections and for information on both John and Franklin Hooper, and to Don Pfister and Gennaro Caccavio at FH for their help with the col- lections of W. G. Farlow. Published as Scientific Contribution Number 1905 from the New Hampshire Agricultural Experiment Station; also issued as Contribution Number 294 from the Jackson Estuarine Laboratory and the Center for Marine Biology. LITERATURE CITED AbEY, W. H. AND P. J. AbEy. 1973. Studies of the biosystematics and ecology of the epilithic crustose Corallinaceae of the British Isles. Brit. Phycol. J. 8: 343-407. 1996] Mathieson et al.—Insular Seaweed Floras 413 ANONYMOUS. 1887. Appleton’s Cyclopedia of American Biography, Vol. II. pleton and Co., New York. p. 252. . 1906. The National Cyclopedia of American Biography, Vol. XIII. James T. White and Co., New York. p. 46. 1985. Tide “Tables 1985, High and Low Water Predictions, East Coast of North and South America, Including Greenland. United States Department of Commerce, NOAA, Washington, DC . 1988. National Status and Trends Program for Marine Environmen- tal Quality: Progress Report—A summary of selected data on chemical contaminants in sediments eo during 1984, 1985, 1986, and 1987. NOAA Techn. Mem. NOS OM APOLLONIO, S. 1979. The Gulf of Mee Courier-Gazette, Inc., Rockland, ME Birp, C. J. AND J. L. MCLACHLAN. 1992. Seaweed Flora of the Maritimes 1. Rhodophyta—the Red Algae. paca Ltd., Bristol, England. Biair, S. M. 1983. Taxonomic treatment of the Chaetomorpha and Rhizo- clonium species ‘Cae. Ene in New England. Rhodora 85: 175-211 BLIDING, C. 1963. A critical survey of European taxa in Ulvales, aa I. Cap- sosiphon, teed ria, Blidingia, oe Opera Bot. 8: 1—60. 1968. ritical oe of Eur ae taxa in Ulvales, Part a Ulva, (le ria, ee Kor . Bot. Not. 121: 535-629. BOURQUE, B. 1996. Pichistorie eee responses to changes in the Gulf of Maine. Absieacts: Gulf of Maine Ecosystem Dynamics, St. Andrews, B, Canada. Bowers, C. H. 1942. Algae of Kent Island. Bowdoin Sci. St. Bull. 8: 36-37. BRAY, J, KR. AND J, J. Corrs . An ordination of the upland forest communities of eae Wisconsin. Heol, Monogr. 27: 325-349. Burrows, E. M. 1991. Seaweeds of the British Isles. Vol. 2, Chlorophyta. Natural pone erie Londo COLEMAN, D. C. AND A. C. MATHIESON. 1975. Investigations of New England marine algae VI seasonal occurrence and reproduction of marine algae ar Cape Cod, Massachusetts. Rhodora 77: 76—104. ear LINS, E S. 1891. Notes on New England marine algae. V. Bull. Torrey Bot. Club 18: 335-341. ——— ee as on New England marine algae. VI. Bull. Torrey Bot. Club 23: ee es ae on New England marine algae. VII. Bull. Torrey Bot. Club 23: 458—462. 1899. A seaweed colony. Rhodora |: 69-71 . 1900. The marine aa. of Great Duck iiawa. Maine. Rhodora 2 209-211. . 1901. Notes on algae, IV. Rhodora 3: 289-293. . 1911. The marine algae of Casco Bay. Proc. Portland Soc. Nat. Hist. 2: 257-282. CONKLING, P. W. 1996. Marine Ecology of Penobscot Bay, pp. 33—44. In: D. D. Platt, ed., Penobscot: The Forest, River and Bay. Island Institute, Rockland, ME. Dixon, P. S. AND L. M. IRVINE. 1977. Seaweeds of the British Isles. Vol. 1, 414 Rhodora [Vol. 98 Rhodophyta. Part 1, Introduction, Nemaliales, Gigartinales. British Mu- seum (Natural History), London Doty, M.S. 1948. The flora of Penikeds seventy-four years after. I. Penikese Island marine algae. Rhodora 50: 253-269. Doy_e, R. G., W. A. ANDERSON, AND FRANK M. BLACKETT, JR. 1970. Pe- nobscot Bay physical resources report (preliminary). Maine Dept. Eco- nomic Development, Augusta, ME. DruenuL, L. D. 1981. Geographical distribution, pp. 306-325. In: C. S. Lob- ban and M. J. Wynne, eds., The Biology of Seaweeds. Botanical Mono- graphs, Vol. 17. Blackwell Scientific Publications, London DuweL, L. AND S. WEGEBERG. 1996. The typification and status of Lepto- phytum (Corallinaceae, Rhodophyta). Phycologia 35: 470—483. Eaton, D. C. 1873. List of marine algae collected near Eastport, Maine in August and September, 1872, in connection with the work of the U.S. Fish Comm. under Prof. S. E Baird. Trans. Connecticut Acad. Arts 2: 343-350. FARLOw, W. G. 1881. Marine algae of New England. Rep. U.S. Comm. Fish and Fisheries for 1879, Appendix A-1:; 1-210. FLETCHER, R. L. 1987. Seaweeds of the British Isles. Vol. 3, Fucophyceae (Phaeophyceae), Part 1. British Museum (Natural History), London HEHRE, E. J . CONWAY, AND R. A. Stone. 1970. Flora of the Wolf Istande Part IH: The marine neve Rhodora 72: 115-118. AND A. C. MATHIESON. 1970. Investigations of New England marine algae II. Composition, seasonal occurrence and reproductive aap of the marine Rhodophyceae in New Hampshire. Rhodora 72: | HILL, A. F 1923. The vegetation of the Penobscot Bay region, Maine. se Portland Soc. Nat. Hist. 3: 305—438. Hoek, C. VAN DEN. 1963. Revision of the European species of Cladophora. E. J. Brill, Leiden, Netherlands. 1982. A taxonomic revision of the American species of Cladophora (Chlorophycese in the North Atlantic Ocean and their geographic dis- ribution. North-Holland Publ. Co., Amsterdam. ne L. M. ae Seaweeds of the British Isles. Vol. 1, Rhodophyta. Part 2A, Cryptonemiales (sensu stricto), Palmariales, Rhodymeniales. British Museum (Natural History), London. AND Y. M. CHAMBERLAIN. 1994. Seaweeds of the British Isles. Vol. 1, Rhodophyta. Part 2B, Se Pile peace Natural History Museum, London. JOHNSON, A. C., P. EF Larsen, D. EF Gappois, AND A. W. HUMASON. 1985. The distribution of polycyclic aromatic hydrocarbons in the surficial sed- iments of Penobscot Bay ese , U.S.A.) in relation to possible sources and to other sites eth Mar Environ. Res. 15: |—16 JOHNSON, D. S.A a) SKUTCH. 1928a. Littoral vegetation on a headland of Mt. oe ee Maine. Submersible or strictly littoral vegetation. Ecology 9: 188-215. AND . 1928b. Littoral vegetation on a headland of Mt. Desert Island, Maine. II. Tidepools and the aac and classification of submersible plant communities. Ecology 9: 307-338 — 1996] Mathieson et al.—Insular Seaweed Floras 415 JorDAN, D. S. 1874. The flora of Penikese Island. Amer. Naturalist 8: 193-197. KELLy, J. T. 1992. Sea- ae ene and coastal erosion in the western Gulf of Maine, pp. 27—44. In: D. D. Townsend and P. E Larsen, eds., T ite of Maine. on Coie Ocean Program, Regional Synthesis Se- s No. 1, Washington, D.C. ee J. M. 1969. The Seaweeds of Cape Cod and the Islands. Chat- ham rae Inc., Chatham, MA. KNIGHT, M. AND M. W. Parke. 1931. nae algae: an algal survey of the south aa — Isle of Man. L. M. B. C. Memoirs 30: 1-155. KOkETZNER, K. L. AND R. D. Woop. 1972. os the marine algae of Kent Island, Bay of Bundy, Rhodora 74: 264-271 LARSEN, P. F 1992. An of the environmental quality of the Gulf of Maine, pp. 71-95. In: D. D. Townsend and P. EF Larsen, eds., Th Gulf of Maine. rave Coaeel Ocean Program, Regional Synthesis Se- ries No. 1, Washington, D.C D. E Gapsols, AND A. C. JOHNSON. 1984a. Trace metal distributions in the surficial sediments of Penobscot Bay, Maine. Bull. Environ. Con- tam. Toxicol. 31: 566-573. , AND . 1984b. Sediment PCB distribution in the Pe- nobscot Bay region _ re Gulf of Maine. Mar. Poll. Bull. 15: 34-35. —— » L. E Doccetr. 1983. Distribution of poly- evelie aromatic hydrocarbons in the surficial sediments of Casco Bay, Maine. Bull. Environ. Contam. Toxicol. 30: 530-535. Lewis, I. E 1924. The flora es Penikese, fifty years after. Rhodora 26: 181-195. Litter, M. M. AND D. S. LirtLer. 1981. Intertidal macrophyte communities from Pacific Baja California and the upper Gulf of California: Relatively constant versus environmentally fluctuating systems. Mar. Ecol. Prog. Ser. 4: 145-15 LUNING, K. 1990. Scawesde Their Environment, Biogeography and Eco- ae Wiley-Interscience Publ., New York. Maaas, C. A. AND M. OMMERSAND. 1993. Seaweeds of the British Isles. ol. I, nin Part 3A, Ceramiales. Natural History Museum, Lon- Matuieson, A. C. 1979. Vertical distribution and longevity of subtidal sea- weeds in northern New England, U.S.A. Bot. Mar. 22: 511-520. 989. Phenological patterns of northern New England seaweeds. Bot. Mar. 32: 419-438. AND E. J. HEHRE. 1982. The composition, seasonal occurrence and reproductive periodicity of the marine Phaeophyceae in New Hampshire. Rhodora 84: 4 Se AND 1983. The composition and seasonal periodicity of the marine Chionephycese in New Hampshire. Rhodora 85: 275—299 AND ————. 1986. A synopsis of New Hampshire seaweeds. Rho- dora 88: 1— 139. AND M. Costa. 1993. Algal vegetation of the York River Estuary and the adjacent open coast of southern Maine. Rhodora 95: 285-324. 416 Rhodora [Vol. 98 > N. B. REYNOLDS. 1981. Investigations of New England marine algae t floristic and nr ak ecological study of the marine algae at ae Point, New Hampshire. Bot. Mar. 24: 521-532. AND . PENNIMAN. 1986a. ae composition and seasonality of New ae seaweeds along an open coastal-estuarine gradient. Bot. Mar. 29: 161-176. AND ———. 1986b. A She aus interpretation of the marine flora from the ices of Shaals, U.S.A . Mar. 29: 413-434. AND ———. 199]. Floristic sere a numerical classification of New En agland< estuarine and open coastal seaweed populations. Nova Hedwigia 52: 453-485 : , AND L. G. Harris. 1991. Northwest Atlantic rocky shore ecology, pp. 109-191. In: A. C. Mathieson and P. H. Nienhuis, eds., Intertidal ae Littoral Ecosystems. Elsevier, Amsterdam. , N. B. REYNoLpbs, AND E. J. HEHRE. 1981. Investigations of New England marine algae I: The species composition, distribution and zo- nation of seaweeds in the Great Bay Estuary System and the adjacent open coast of New Hampshire. Bot. Mar. 24: 521-532. Murray, S. N. AND M. M. Litter. 1981. Biogeographical analysis of in- tertidal macrophyte floras of southern California. J. Biogeogr. 8: 339-— — > 1. A. Asporr. 1980. Biogeography of the California marine igae ce emphasis on the Southern California Islands, pp. 325— 339. In: D. M. Powers, ed., The California Islands: Proceedings of a Multidisciplinary Symposium. Santa Barbara Museum of Natural His- tory, ae hee CA. PIKE, R. B. AN - R. HopGpon. 1963. The flora of the Wolf Islands, New Baek Se 1. Rhodora 65: 82-96. Piatt, D. D., ed. 1996. Penobscot: The Forest, River and Bay. Island Insti- tute, Rockland, ME. » P. W. CONKLING, A. NAEGEL, S. DICKERSON, T. AMES, AND C. OLD- HAM. 1996. Introduction, pp. 8-9. In: D. D. Platt, ed., Penobscot: The Forest, River and Bay. Island Institute, Rockland, ME. Porter, E. 1966. Summer Island: Penobscot County. Sierra Club, Ballantine Books, oe pegs CA. REYNOLDS, N. AND A. C. MATHIESON. 1975. Seasonal occurrence and ecology . marine i in a New Hampshire tidal rapid. Rhodora 77: 512- Ricu, L. D. 1975. The Coast of Maine: An Informal History and Guide. Down East Books, Camden, ME. SCAGEL, R. E 1963. Distibution of attached marine algae in relation to oceanographic conditions in the northeast Pacific, pp. 37-50. In: M. J. bar, ed., Marine Distributions. Roy. Soc. Canada Spec. Publ. No. 5, University of Toronto Press, Toronto . 1970. a algae of Bowie Ssnonnt Syesis 3: 15-16 SCHNEIDER, C. W. AND R. B. SEARLES. 1991. Seaweeds of the Southensieni United States. one. Univ. Press, Durham SEARS, J. R. AND R. T. Witce. 1975. Sublittoral ‘henthie marine algae of 1996] Mathieson et al.—Insular Seaweed Floras 417 southern Cape Cod and adjacent islands: seasonal periodicity, associa- Sout, G. R. AND I. TirrLey. 1986. A Checklist and Distributional Index of the Benthic Marine Algae of the North Atlantic Ocean. Huntsman Mar. Lab. and British Museum (Natural Hist.), St. Andrews, New Brunswick and London. STONE, R. A., E. J. HEHRE, J. M. Conway, AND A. C. MATHIESON. 1970. A preliminary checklist of the marine algae of Campobello Island, New Brunswick, Canada. Rhodora 72: 313-338. Sze, P. 1982. Distribution of macroalgae in tidepools on the New England coast. Bot. Mar. 25: 269-276. Tay_Lor, W. R. 1957. The Marine Algae of the Northeastern Coast of North America. Univ. Michigan Press, Ann Arbor, MI. TittLey, I., W. EF FARNHAM, G. R. SOUTH, AND D. KEATS. 1987. Seaweed communities of the Passamaquoddy region, southern Bay of Fundy, Can- ada. Brit. Phycol. J. 22: 313. Vapas, R. L., M. KESER, AND P. C. RuSANOWSKI. 1976. Influence of thermal loading on the ecology of intertidal algae, pp. 202-212. In: G. W. Esch MacFarlane, eds., Thermal Ecology II. ERDA Symposium Series, Augusta, GA. VILLALARD-BOHNSACK, M. 1995. Illustrated Key to the pane of New England. Se Island Natural History Survey, Kings WEBBER, E. E. AND R. T. WILCE. 1971. Benthic salt ae ee at Ipswich, oe Rhodora 73: 262— 291, WOELKERLING, W. J. 1973. The morphology and systematics of the Audoui- nella complex Pcrochastncsae, Rhodophyceae) in northeastern United States. Rhodora 75: 529-621. WYNNE, M. J. AND J. N. HEINE. 1992. Collections of marine red algae from St. Matthew and St. Lawrence Islands, the Bering Sea. Nova Hedwigia 55: 55-97. APPENDIX: STUDY SITES WITHIN THE PENOBSCOT BAY (MAINE) AREA 1. Allen Island: one of the Georges Islands at the mouth of the St. George River on the southwesterly portion of Penobscot Bay, due north of Mon- hegan Island and south of Port Clyde (43°52’'N, 69°19.8°W); ~7620m of shoreline. . Hurricane Island: located in the mid-central part of Penobscot Bay (44°02'N, 68°54.6'W), offshore from the southwesterly side of Vinalhaven Island and near Green Island; ~2930m of shoreline 3. Long Island: located within Jericho Bay (44°07'N, 68° 21.5'W), offshore (SE) from Swans Island and northeast from Isle Au Haut; ~16770m of shoreline. Collections were made on the NW side near Frenchboro. . Marshall Island: located within Jericho Bay (44°06.5'N, 68°30.5'W), off- shore (S) from Swans Island and northeast from Isle Au Haut; ~11940m of shoreline. Collections were made in Baxam Cove. N eo.) £ 418 Rhodora [Vol. 98 n ~) ioe) \o - McGlathery Island: located within Merchants Row of Penobscot Bay (44°08.5'N, 68°37'W), due north of Isle au Haut and south of Deer Isle: ~4020m of shoreline. . Monhegan Island: located seaward of Penobscot Bay (43°46'N, 69°19'W): due south of Allen Island and almost equidistant from New Harbor to the northwest and Port Clyde to the north; ~8567m of shoreline. . Pond Island: located within Jericho Bay (44°13.3'N, 68°28.6'W) due north of Swans Island; ~4318m of shoreline. Collections were made on the sheltered SE and NE sides. . Two Bush Island: located within the western part of Penobscot Bay, due east (and somewhat south) of Tenants Harbor, as well as SW of Hurricane Island (43°58.2'N, 69°04.3’W): ~1700m of shoreline. . Wooden Ball Island: located within the south central (exposed) part of the Penobscot Bay (43°51.3'N, 68°49.05'W), offshore (S) from Vinalhaven Island and Isle Au Haut (SW), and near Matinicus and Ragged Islands; ~5560m of shoreline. Collections were made on the sheltered NW and exposed eastern sides. RHODORA, Vol. 98, No. 896, pp. 419-444, 1996 SURVEY OF OLD-GROWTH FOREST IN MASSACHUSETTS PETER W. DUNWIDDIE Massachusetts Audubon Society, Lost Farm, Hummock Pond Rd., Nantucket, MA 02554 Current Address: The Nature Conservancy, 217 Pine St., Suite 1100, Seattle, WA 98101 ROBERT T. LEVERETT 52 Fairfield Ave., Holyoke, MA 01040 ABSTRACT. We surveyed forests in Massachusetts to identify any remain- ing aid. growth stands. We located twenty-eight tracts in western Massachu- setts that met our criteria, with one additional site east of the Connecticut River. Hemlock and northern hardwoods in excess of 150—200 years dominate most sites, which range in area from 3—28 ha. Most stands occur on steep slopes and may have escaped cutting due to their inaccessibility. Several potential sites remain to be investigated, but we do not expect that the total area of old-growth forests in Massachusetts will greatly exceed 260 ha. How- ever, less seangen definitions of old growth may include additional sites and eee acreages Dat Sollesied from 26 permanent vegetation monitoring plots Sele comparisons with other forests. Basal areas in the old-growth stands are s ilar to other old-growth forests in New England, from 34—42.8 m’/ha; va aM are only slightly higher than basal areas fon nearby second growth forests of the same type. Densities of stems >10 cm dbh in the old-growth plots are 347-480 trees/ha, 25-40% lower than in second growth forests. No vascular plant species were encountered that were unique to the old-growth stands. Key Words: old-growth, northern hardwood forests, eastern hemlock, Mas- sachusetts Old-growth forests in the eastern United States have attracted considerable attention in the last decade, and increasingly are threatened in much of North America (Davis 1996). Such forests are important because they may contain unique assemblages of species, offer significant dendrochronological information on past tree growth, and provide valuable baseline information on forest composition and dynamics for comparison with other areas (Whitney 1987). Definitions vary among researchers, but forests that frequently are referred to as old growth are generally regard- ed to have had continuous forest growth over a long period with 419 420 Rhodora [Vol. 98 little human disturbance (Leverett 1996; Whitney 1987). Exam- ples have been reported in several New England states (Dunwid- die et al. 1996), including Maine (Maine Critical Areas Program 1983), Vermont (Bormann and Buell 1964), New Hampshire (Cline and Spurr 1942; Carbonneau 1986; Leak 1987), and New York (Leopold et al. 1988; Woods and Cogbill 1994). In Mas- sachusetts, however, Egler (1940) reported finding no ‘‘pre-co- lonial” forests in his study of the vegetation in the Berkshires. One unpublished reference (Hosier 1969) presents data from a purported virgin stand in northwestern Massachusetts, but no ev- idence is provided to support the claim that the stand is, indeed, old growth. Thus, a primary purpose of our study was to deter- mine whether any old-growth forest remains in Massachusetts. Once we determined that old-growth forests existed in the state, three specific objectives were defined: (1) to identify and map existing old-growth forests in Massachusetts, (2) to compile a descriptive summary of the location, physical characteristics, flo- ra, and structure of these sites, and (3) to determine how the Massachusetts old-growth stands compare with data from other forests in the northeastern United States. Detailed reports docu- menting features of the individual stands have been submitted to the Massachusetts Natural Heritage and Endangered Species Pro- gram (Dunwiddie 1991; Dunwiddie 1993). In this paper we pro- vide an overview of all the known old-growth sites in Massa- chusetts, summarize some of the tree and understory data col- lected in this study, and compare results with other old-growth sites in New England. The first step in identifying old-growth forests was to develop criteria that distinguish such stands from regrowth forests. We modified criteria used elsewhere in the Northeast (Maine Critical Areas Program 1983) to identify stands that would be considered by most researchers to be old growth. Using less conservative criteria, others might add to the acreage of old-growth forest in Massachusetts. However, our intention was to provide an inven- tory of fairly unequivocal sites that could be useful as a starting point for other investigations. To qualify as old-growth forest, a stand must (1) be a relatively homogeneous area of at least 3 ha, (2) exhibit minimal evidence of human influence or other catastrophic disturbance of the stand, (3) show evidence of tree regeneration, especially of late-succes- sional species, resulting in a relatively stable forest composition, 1996] Dunwiddie and Leverett—Old-growth Forest 421 and (4) include dominant canopy trees of an age >50% of the maximum age for those species (Fowells 1965). These criteria eliminated small or fragmented sites with little undisturbed forest interior, as well as stands dominated by ear- ly-successional species such as Betula papyrifera Marsh. (paper birch) and Acer rubrum L. (red maple). The second and fourth criteria eliminated areas where widespread blowdowns, landslips, and other disturbances removed old trees from dominance in stands. The determination of a minimum age for trees to be con- sidered old growth is arbitrary, but application of the last criterion included stands dominated by Tsuga canadensis (L.) Carr. (east- ern hemlock) >200—225 years old, and hardwoods such as Acer saccharum Marsh. (sugar maple) and Fagus grandifolia Ehrh. (American beech) >175—200 years old. MATERIALS AND METHODS Forests considered to be probable old growth were identified by extensive field searching, as well as by information provided by the Massachusetts Natural Heritage and Endangered Species Program, foresters, and other individuals. Although we consid- ered potential sites throughout the state, our work fairly quickly became focused in western Massachusetts. Numerous sites were rejected due to small size, evident disturbance, or lack of old trees. Areas that appeared to meet our criteria were selected for more detailed study. Beginning in 1991, assessments were made of composition, signs of disturbance, area, and likelihood of suf- ficient age; selected increment cores were gathered from domi- nant trees to more accurately determine the age of the older in- dividuals. General information was recorded on the species, slope, aspect, and elevation of each stand. A compass and altim- eter were used to map the perimeter of old-growth stands on enlarged U.S. Geological Survey topographic maps. Stand acre- age was calculated from these maps. Even with well-defined criteria for identifying old-growth stands, mapping boundaries and determining acreages is still sub- ject to considerable interpretation. The density of older trees nec- essary to qualify a stand as old growth is ultimately a subjective decision, and different perspectives adopted by other researchers could significantly change acreage estimates of old-growth forest in the state. Because of our desire to identify reasonably unequiv- 422 Rhodora [Vol. 98 ocal sites, we only considered stands that included at least three older trees per hectare throughout the stand. In the field, this necessitated identifying probable old trees, and estimating their density on a site. Where old individuals were common and readily recognizable, decisions on the boundaries of old-growth stands were clear. However, where densities of old trees were low, or where trees lacked characteristics often associated with great age, such as deeply fissured bark, snag tops, or heavy limbs, mapping decisions were more difficult. Detailed quantitative data on the canopy and understory veg- etation were collected from within the twelve stands that had been identified by 1992. A standardized methodology developed for sampling old-growth forests was adapted from Shifley et al. (1991). Permanent 0.1 ha fixed-radius plots were located within each stand to collect tree data, with several nested plots for sam- pling other vegetation strata. Time constraints precluded installing the large number of randomly placed plots (often >30 for many parameters recommended by Shifley et al. 1991) to derive statis- tically valid descriptive samples of the entire stands. Instead, from one to four plots were established in locations that included sev- eral trees that were typical of the older trees in the stand. Where several different forest types were represented within a single stand, attempts were made to place plots to capture this variability in composition. The plot layout follows Shifley et al. (1991) and consists of a 0.1 ha circular plot for trees, a 0.01 ha circular subplot for sap- lings, and four 2.5 m? subplots for ground vegetation. All living and dead trees >10 cm diameter at breast height (dbh) were re- corded in the large plot, and saplings from 2—10 cm dbh were sampled in the 0.01 ha subplot. Each plot center was marked with a 2.5 cm plastic pipe. Data collected for each tree included azi- muth and distance from the plot center, species, dbh, crown class and ratio, and decay stage. At least two trees, estimated to be among the oldest trees in the plot based on size, bark, and crown characteristics, were cored. These cores were glued into mounts and sanded, and growth rings were counted to provide minimum ages. Actual ages were estimated by determining growth rates for the innermost several centimeters of the cores, calculating the distance to the center based on diameter measurements, and adding the appro- priate number of years based on the extrapolated growth. At sev- 1996] Dunwiddie and Leverett—Old-growth Forest 423 eral sites, heights were measured for individual trees using a tape measure and inclinometer. Understory vegetation was sampled in four square subplots 3 m from the center of the main plot. Woody stems at least 1 m tall and less than 2 cm dbh were counted by species, and the abundance of each species in the subplots was recorded on a three-point scale. A species list also was compiled for each 0.1 ha plot and for each site as a whole. In this paper, we present summaries of physical site character- istics, basal area and density data from living and dead trees in the different forest types, and understory compositional data. Vas- cular plant names follow Gleason and Cronquist (1991); bryo- phytes follow Conard and Redfearn (1979). RESULTS Twenty-eight old-growth stands in western Massachusetts were identified and described in this study (Tables | and 2). One ad- ditional site, Wachusett Mountain, was identified in central Mas- sachusetts. Collectively, they include a total area of 255.2 ha. In addition, we identified ten other small stands of old growth in western Massachusetts, but did not include them here because the individual stands are less than 3 ha. We have yet to confirm whether Chesterfield Gorge, a 7 ha hemlock stand belonging to The Trustees of Reservations, is an old-growth forest that meets our criteria. Quantitative data were gathered from 26 plots in 13 of the stands (Table 1). Most of the stands are small, generally less than 12 ha, and occur on steep slopes (av.=35°). Many stands are on northwest- to northeast-facing slopes and, with the exception of the Mt. Greylock sites, most occur at elevations from 300—500 m. Canopy trees ranged from about 24 m to 36 m tall. The data on the total area of old-growth forest from the Mt. Greylock stands represent minimum estimates, and are likely to understate the true amount. The stands all are scattered within the area known as the Hopper, within which exists a complex disturbance matrix resulting from fire, landslides, blowdowns, and land use. The interspersion of old trees through much of the Hop- per, individually and in larger stands, makes it difficult to delin- eate precise boundaries of areas that meet all the criteria of old growth defined in this study. 424 Rhodora [Vol. 98 Table 1. Characteristics of old-growth stands with permanent vegetation monitoring plots in Massachusetts. Different stands within each site are dis- tinguished by letters or local names. H = Hemlock, NHW = Northern hard- oods. All sites are owned by the Massachusetts Department of Environ- mental Management with the exception of Bash Bish A, which is private. Stand Size Lat. Long. Site Name (ha) (N) (W) Bash Bish A-Grinder 6.9 42°6' 73°29' B-Falls 5.6 42°7' 73°30' Cold River A 7.6 42°38' 72°58' B-Black Bk. 4.5 42°37'30" 72°58'30" C 16.6 42°38’ 72°58'30" D 28.2 42°38’ 72°59' E-Manning Bk. 5.3 42°38'30" 72°59'30" Dunbar Bk. 13.1 42°42’ 72°58’ Parsonage 4.2 42°43’ 72°59' Fife Bk. 9.5 42°41'30" 72°59' Mt. Greylock Deer Hill 11.6 42°38’ Wo Ll Money Bk. 9 42°39'30" 73°10'30" Mt. Everett 9.8 AQ" 73°25'30" Tower Bk. 11.3 42°44’ 72°56 In 1995, we confirmed the presence of a large old-growth . on Wachusett Mountain (Dunwiddie 1995; Cogbill 1996). T stand is surprising for its size, as well as for its previously over- looked location in the east-central part of the state. While detailed 1996] Dunwiddie and Leverett—Old-growth Forest 425 Table 1. Extended. Plot Elev. Aspect Slope Forest No. (m) (deg.) (deg.) Type l 421 70 45 H-NHW 2 442 35 40 H 1 367 10 45 H 1 308 360 32 NHW 2 413 30 45 H-NHW 3 457 320 42 H-NHW 4 396 310 40 NHW 1 384 310 43 H-NHW 2 390 310 38 H-NHW 3 396 320 40 H-NHW 1 372 90 39 NHW 1 381 320 37 H 2 381 40 35 H-NHW 1 440 90 16 H-NHW 2 430 100 15 NHW 1 392 58 28 NHW 2 468 46 26 H-NHW I 494 290 25 1 500 190 17 NHW 2 497 190 10 NHW 3 369 200 38 H-NHW 4 378 210 35 H-NHW 1 611 360 26 H 2 611 340 45 NHW 3 659 360 45 H-NHW 1 502 28 30 H 0 400-488 90-120 25-30 H-NHW data on the stand have not been collected yet, our initial surveys suggest that it extends from about 420—600 m elevation on the north and east sides of the mountain, with an area of about 28 ha. The site is dominated by several forest types, including hem- 426 Rhodora [Vol. 98 Table 2. Characteristics of old-growth stands in Massachusetts in which permanent plots have not been established. Different stands within each site are distinguished by local names. Areas are approximate. H = Hemlock, HW = Northern hardwoods, BB = Black Birch, RO = Red Oak, RP = Red Pine, WP = White Pine, RS = Red Spruce. DEM = Mass. Department of Environmental Management, TTOR = The ee of Reservations, AMC = Appalachian Mountain Club. Size Lat. Long. Site Name (ha) (N) (W) egus a 42 39' gb oe Si Cold River Todd-Clark 24 42°38'45" Jo. 30) Wheeler Bk. 3 4238’ 72°58'45" Upper Cold R. 3 42°38’ voy Trout Bk. 3 42°37’ e312 Bryant Estate 3 42 30" 72°57! Dunbar Brook Bear Swamp 10 4242’ 72:57'40" Upper Dunbar 6 42 42' 72:58'45" Ice Glen 4 42°16'20" 72 18'45" Sages Ravine 6 4231! 7227'15” Mt. Greylock Paris Bk./Mt. Fitch 4 42°39’ 73 10' Money BK. tribut. 6 4238’ 7310' Roari 6 42°37' Fk Spruce Mtn 4 42°42’ 72°59’ 15" Windsor Jams 3 42°32’ 73 59' Wachusett Mtn. 28 42 30/30" 71-53'30" Total (Tables 1 & 2) 259.2 lock, Betula alleghaniensis Britton (yellow birch), beech, other hardwoods, and Picea rubens Sarg. (red spruce). Ages of some of the trees have been documented in excess of 300 years (Dun- widdie 1995). Forest composition and structure. The old-growth stands found in this study represent a spectrum of hemlock and northern hardwood forest types common in the Berkshires. These include stands dominated almost entirely by hemlock, such as Bash Bish and Mt. Greylock-Deer Hill, forests with a mixture of hemlock and northern hardwoods including yellow birch, beech, and sugar maple, and forests consisting primarily of northern hardwoods. The plot data were grouped within these three forest types for most analyses. Several sites that were documented after 1992, and 1996] Dunwiddie and Leverett—Old-growth Forest 427 Table 2. Extended. Elev. Forest (m) Aspect Type Owner 300—S500 South NHW-RO NE Power 240-520 Varied H-NHW, H-BB-RO DEM 300-390 Varied H-NHW DEM 360—450 Varied H-NHW-RS DEM 330—480 East H-NHW DEM 400—450 Varied H-NHW TTOR 330—540 Varied NHW-RO-H DEM 420-510 Varied NHW, H-NHW DEM 220—330 Varied H-WP-RP-NHW Laurel Hill Assoc. 330-470 North H DEM-AMC 450-750 Varied H-NHW-RS DEM 450-850 West H-NHW-RS DEM 480—670 Varied H-NHW-RS DEM 600—730 Varied RS-NHW DEM 425-450 Varied -RS DEM 420—600 N-E NHW-H-RO DEM in which no plot data were gathered, presented an unusual mix- ture of hemlock, Betula lenta L. (black birch), and Quercus rubra L. (northern red oak) that might be considered another forest type. Seven plots fell within hemlock-dominated stands (Tables 3 and 4). Hemlock comprises an average of 75% of the basal area (65-90%) and 71% of the stem density. Red spruce and yellow birch are frequent but minor associates in the canopy, and Acer pensylvanicum L. (striped maple) is common in the subcanopy. These plots contain the highest average basal area of living trees of any of the three old-growth forest types examined in this study (42.75 m?/ha), with a maximum value of 52.22 m*/ha recorded at one site. Average live stem densities are also higher in the hemlock stands than in the other two forest types (480 trees/ha). The species richness of understory vascular plants is the lowest 428 Rhodora [Vol. 98 Table 3. Average basal area and density of living and standing dead trees >10 cm dbh in seven plots in Berkshire old-growth hemlock forests. Live Dead BA Density BA Density (m*/ha) (trees/ha) (m*/ha) — (trees/ha) Acer pensylvanicum 0.6 28.6 0.3 10.0 Acer rubrum 0.02 4.3 Acer saccharum 0.1 1.4 Betula alleghaniensis 1.9 30.0 0.2 a7 Betula lenta 1.5 27.1 Fagus grandifolia 0.7 11.4 0.3 4,3 Picea rubens 4.0 22.9 0.7 10.0 Pinus strobus 1.5 10.0 0.5 4.3 Quercus rubra 0.3 1.4 Tsuga canadensis 32.1 342.9 1.4 31.4 Unknown 0.5 1.4 Total 42.8 480.0 3.9 67.1 in this forest type, with an average of 10.6 species in the 0.1 ha plots. Besides seedlings of hemlock and yellow birch, the only commonly occurring understory species is Dryopteris carthusi- ana (Vill.) H. P. Fuchs, found in 50% of the subplots. Seven old-growth plots were placed in northern hardwood for- ests (Tables 5 and 6). Sugar maple is the primary species in these stands, with beech as a secondary dominant. The lowest average basal areas and stem densities occur in this forest type, but the understory species richness is high (25.3 spp./0.1 ha). The most common understory taxa include Dryopteris carthusiana (68%), Arisaema triphyllum (L.) Schott (64%), and seedlings of striped maple (54%) and beech (54%). Twelve plots were established in hemlock-northern hardwood forests (Tables 7 and 8). This forest type is intermediate in most measures between the hemlock and northern hardwood stands: both the basal area and stem density values fall between the other two forest types. Hemlock has the highest basal area and stem densities, but beech, sugar maple, and yellow birch also are prom- inent in the canopy. Understory species richness averaged 15.5 spp./0.1 ha. Taxa frequently encountered in the subplots include Dryopteris carthusiana (79%), Viburnum alnifolium Marsh. (60%), Lycopodium lucidulum Michx. (35%), and Oxalis aceto- sella L. (33%). 1996] Dunwiddie and Leverett—Old-growth Forest 429 Table 4. Percent frequency of understory species in old-growth hemlock forest Bash Bash Cold ae Parson- Grey- Ever- Bish Bish River River age lock ett A2 Bl A4_ Dt Acer pensylvanicum 25 100 Acer rubrum 50 25 Acer spicatum 25 25 Acer sp. 75 Betula ris alae 100 50 25 25 25 Betula len 75 25 Dryopteris as 50 50 Dryopteris carthusiana 25 20 75 50-100 75 Fagus grandifolia 50 25 Lycopodium lucidulum 25 50 Mitchella repens 50 Oxalis acetosella 100 Picea rubens 75 Polypodium vulgare 100 25 Polystichum acrostichoides 50 runus serotina 25 uga canadensis 100 75 25-100 75 25 100 Viburnum alnifolium 75 No. of Vascular Species 8 10 14 7 10 16 6 Bazzania trilobata 100 25 75 50 50 Dicranum sp. 50 50 50 Hypnum sp. 75 25 25 100 Leucobryum cf. albidum 25 Leucobryum glaucum 100 75 1D 25 25 Lichens 50 25 Nowellia sp. 25 Odontoschisma denudatum — 25 ella sp. 23 25 pee capania nemorosa 25 Tetraphis pellucida 50 25 Unidentified bryophytes 50. 100 =100-— 100 100 25 No. of Bryophytes and Lichens 8 9 I 5 6 1 8 In all three forest types, about 50% of the trees >10 cm dbh were recorded in the canopy, with the other half classified as either overtopped or midstory. The density and basal areas of standing dead trees varied widely among all the plots. However, both the average stem density and basal area values are close to 430 Rhodora [Vol. 98 Table 5. Average basal area and density of living and standing dead trees in northern hardwood forests. All data are from old-growth forest sites with the exception of the 22 Berkshire Continuous Forest Inventory plots (CFI); Berk = Berkshires, MA (this study; n = 7); Adir = Adirondacks, NY (Le- opold et al. 1988); Bowl = The Bowl, NH; Mt. Pd = Mountain Pond, NH; Wms = Williams, NH (Leak 1987). Live BA (m7?/ha) Berk Adir CFI Bowl Mt. Pd Wms >10cm >S5cecm >13 cm >5 ecm >S5cm >5 cm Acer pensylvanicum 0.1 6) 0 1.4 0.8 0.5 Acer rubrum 0.8 0 3.4 0 0 0 Acer saccharum 21.4 30.0 6.1 10.6 15.8 6.5 Betula alleghaniensis 1.4 2.2 3.6 7.7 2.3 3.4 Betula lenta 0.2 0 0 0 0 0 Fagus grandifolia 6.8 2.0 5.1 10.3 8.4 5.8 Fraxinus americana 2.5 0 L.1 1.2 0.8 Ostrya virginiana 0.1 0.6 0 0 0 0) cea rubens 0 6) 0) 0.4 0.3 0 Pinus strobus 10) 0 0 0 0 0 Quercus rubra 0 0 1.3 0 0 0 Tsuga canadensis 0.7 2.8 1.7 0.1 0 0.9 Other 0.1 0 5.3 0.1 1.7 75 Total 34.1 37.6 27.7 30.6 30.6 25.3 10% of the total (living and dead) in all three forest types (Tables 3, 3, and 7). Age estimates for selected older trees were obtained from ring counts on increment cores collected at many of the sites. These figures may underestimate the true ages by up to 50 years or so for many of the trees due to heart rot or cores not reaching the center. Some of the hemlock cores contained over 300 rings, and a few of these trees may have attained ages of 400 years or more. Fewer hardwoods were cored, but sugar maples of 200-250 years were encountered in several plots. All the plots contain trees in excess of 200-225 years, confirming the assessment that the stands were of an age to be considered old growth. A graph of the estimated ages of all the cored trees in the study plots shows a modal age of 220—240 years (Figure 1). The size of old trees in Massachusetts old-growth stands varies greatly with the growing conditions. In many sites studied here, 1996] Dunwiddie and Leverett—Old-growth Forest 43] Table 5. Extended. Live Dead Density BA Density (Tr/ha) (m*/ha) (Tr/ha) Berk Adir CFI Berk Adir Berk Adir >10 cm >5 cm >13 cm >10cm >5 cm >10 cm >5 cm 11.4 0 0 0 0 10) 0 0 0 53.7 0 0 0 0 184.3 180 0 1.9 0.5 11.5 50 10 0 0.5 0 2.8 0 1.4 0 0 0 0 0 0 110 110 97.9 0.8 0 8.7 0 12.9 0 0 0 0 0 0 1.4 60 0) 0 0 0 0 0) 0) 0 0) 0.2 6) 20 0 0 0) 0 0 0) 0 0 0 10.7 0 0 0 0 7.1 70 30.4 0 0 0 10 0 276.8 0.02 0 2.8 0 347.1 430 469.3 3.2 0.8 25.7 70 old trees were neither particularly tall nor of exceptional girth due to rigorous growing conditions, a factor which may have contributed to these stands not being cut for lumber. However, since maximum dimensions of trees frequently are of interest in comparing growth in different regions, data are included sum- marizing the upper size limits measured for twelve species in Massachusetts old-growth sites in the most favorable growing conditions (Table 9 DISCUSSION The stands identified as old growth in this study consist of later-successional species that have been undisturbed for over 200 years. If limited disturbances, such as selective cutting, occurred centuries ago, they might be difficult to detect. However, we con- sider any such activity at these sites unlikely to have been sig- 432 Rhodora [Vol. 98 Table 6. Percent frequency of understory species in old-growth northern hardwood forests. Cold Cold Cold Dun- Fife Fife Grey- River River River bar Bk. Bk. lock Al Cl E2 I | 2 2 Acer pensylvanicum 25 25 50 50. 100 100 25 Acer rubrum 25 Acer saccharum 50 25 25 50 75100 Acer spicatum 50 25 25 r sp. 25 75 50 Actaea pachypoda 25 25 Allium tricoccum 100 100 Arisaema triphyllum 100 50 100 100 100 Aster acuminatus 75 Aster divaricatus 50 Aster sp. 50 25 Athyrium ie -femina 50 25 Betula len 25 ene rhe virginianum 23 Carex sp. 25 25 Caulophyllum thalictroides 100 50 Claytonia virginica 50 Dicentra canadensis 25 100 Dryopteris marginalis 25 25 25 Dryopteris carthusiana 100-100 75 100 LOO Erythronium americanum 25 75 100 75 Fagus grandifolia 75 25 50 25 100 100 raxinus americana 25 25 25 25 Galium triflorur 75 Impatiens capensis 100 25 portea canadensis 25 Lonicera canadensis 25 Lycopodium lucidulum 50 100 Osmorhiza clayton 50 Oxalis acetosella 50 25 Panax trifolius 25 Polygonatum pubes 25 Polystichum ire 50 is) 75 75 Prunus serotina 25 25 Ranunculus sp. 25 Ribes sp. 25 Rubus sp. 25 Streptopus roseus 50 Thelypteris noveborascensis 25 Tiarella cordifolia 25 22 25 Trillium cernuum 25 Trillium erectum 25 25 75 25 25 1996] Dunwiddie and Leverett—Old-growth Forest 433 Table 6. Continued. Cold Cold Cold Dun- Fife Fife Grey- River River River bar Bk. Bk. lock Al Cl E2 1 1 2 2 Tsuga canadensis 25 Viburnum acerifolium 25 Viburnum alnifolium 72 75 50 100 Viola canadensis 100 Viola rotundifolia 50 25 50 25 25 Viola sp. 23 23 WD 25 No. of Vascular Species 16 18 29 20 30 38 26 Bazzania trilobata 25 Dicranum sp. 25 25 Unidentified bryophytes 25 50 50 75 pes) 75 ~=100 No. of Bryophytes 3 l 2 1 l 1 nificant. Although settlements existed in Berkshire County in the mid-1700s, forest cutting at that time would have focused on clearing the better agricultural land and on harvesting high quality timber, especially spruce and pine, in accessible locations. While factors such as steep slopes, remoteness from logging roads, and lesser quality wood combined to favor the survival of these rem- nants of old growth, chance no doubt contributed as well. This survey of old-growth forest in Massachusetts emphasizes its extreme rarity in the state. After more than 300 years of in- tensive land clearance, agriculture, forestry, and development, only about thirty documented sites remain. Furthermore, these scattered remnants are small patches in a matrix of regrowth for- est. The largest old-growth site is only about 28 ha, considerably smaller than the stands of several thousand hectares in the Adi- rondacks of New York (Leopold et al. 1988) and the White Mountains of New Hampshire (Leak 1987). Data from some other New England old-growth forests are presented in Tables 5 and 7 for comparison. In addition, data are included from selected Continuous Forest Inventory (CFI) sites in Massachusetts (W. Rivers, Massachusetts Department of En- vironmental Management, pers. comm.). These variable-radius tree plots are located every half mile on most state lands in the Berkshires. Data included in the tables come from all the CFI plots in the vicinity of the old-growth stands (generally sites with- 434 Rhodora [Vol. 98 Table 7. Average basal area and density of living and standing dead trees in hemlock-northern hardwood forests. All data are from old-growth forest sites with the exception of the 22 Berkshire Continuous Forest Inventory plots (CFD; Berk = Berkshires, MA (this study; n = 12); Adir = Adirondacks, NY (Leopold et al. 1988). Live BA (m?*/ha) Berk Adir CFI >10 cm >5 cm >13 cm Acer pensylvanicum 0.2 0.5 0.0 Acer rubrum 0.0 0.0 7.1 Acer saccharum 4.6 3.2 0.2 Betula es 3.6 0.0 4.4 Betula lente 0.3 0.0 0.0 aes grandifolia 5.8 3,5 22 F us americana 0.2 0.0 0.0 Gis virginiana 0.01 0.0 0.0 Picea rubens 0.3 2.2 0.0 Pinus strobus 0.0 0.0 1.8 Quercus rubra 1.6 0.0 0.7 Tsuga canadensis 20.5 30.9 14.1 ther 0.3 0.0 4.6 Total 37.4 42.3 35.2 in the same valley) that were from the same forest types. These figures provide comparative data for second growth hemlock- northern hardwood and northern hardwood forest types; no CFI data were available for the hemlock forest type. It is important to note, however, that due to the small sample sizes and non- random placement of plots in the Berkshire old-growth stands, differences between these data sets should be considered only as general trends. In addition, different authors used different min- imum diameters, which also limits data comparability. The hemlock old-growth stands in the Berkshires (Table 3) had a similar total basal area (42.8 m*/ha) to that reported by G. Whit- ney from Heart’s Content, Pennsylvania (42.4 m2/ha), and an old- growth hemlock-yellow birch forest in New Hampshire (Tritton and Siccama 1990). Heart’s Content had a lower stem density (366 stems/ha) than the average from the Berkshire sites (480 stems/ha). Standing dead stems in the Berkshire old-growth hem- lock forests represented 8% of the total basal area, and 12% of 1996] Dunwiddie and Leverett—Old-growth Forest 435 Table 7. Extended. Live Dead Density BA Density (Tr/ha) (m?/ha) (Tr/ha) Berk Adir CFI Berk Adir Berk Adir >10 cm >5 cm >13 cm >10cm >S5cm 10cm >5 cm 10.8 93.0 0.0 0.1 0.1 6.7 18.0 0.0 0.0 119.7 0.0 0.0 0.0 0.0 61.7 25.0 0 0.4 0.3 5.8 3.0 46.7 0.0 0.0 0.0 3.2 0.0 15.0 10.8 0.0 0.0 0.1 0.0 1.7 0.0 109.2 238.0 41.2 1.2 1.4 15.8 23.0 1.7 0.0 0.03 0.0 0.8 0.0 0.8 0.0 0.0 0.0 0.0 0.0 0.0 1.7 28.0 0.0 0.0 0.9 0.0 15.0 0.0 0.0 12.3 0.0 0.0 0.0 0.0 14.5 0.0 0.0 0.0 0.0 163.3 240.0 328.3 2.6 4.2 11.7 25.0 204.2 0.04 2 1.7 415.8 622.0 720.2 4.5 10.4 44.7 102.0 the density. These values closely match those reported by Tritton and Siccama (1990) for this forest type (8 and 9%, respectively). Data from northern hardwood forests in New England are com- pared in Table 5. Basal areas of old growth in Massachusetts are similar to values from the Adirondacks and New Hampshire, but averaged about 23% higher than in the CFI second growth plots. Although somewhat different minimum stem diameters were used in different studies, stem densities were higher in the Adirondacks and in the CFI plots than in the Berkshire old growth; this pattern also was observed in the hemlock-northern hardwood types (Ta- ble 7). Standing dead stems in the Berkshire old-growth northern hardwood forests represented 8% of the total basal area, and 7% of the stem density, less than the average of figures reported by Tritton and Siccama (1990) for this forest type in New Hampshire and Vermont (12% of the total basal area, 13% of the total stem density). Average live basal areas in the twelve hemlock-northern hard- 436 Rhodora [Vol. 98 Table 8. Percent frequency of understory species in hemlock-northern hardwood old-growth. Bash Bish Cold River Al A2 A3 Acer pensylvanicum 25 Acer saccharum 25 Acer spicatum 50 Acer sp. 25 Aralia nudicaulis 25 25 i triphyllum ster sp. ewes alleghaniensis 50 Betula lenta 25 Carex s Clintonia borealis 25 Dryopteris marginalis Dryopteris carthusiana 100 100 Erythronium americanum Fagus grandifolia 75 Fraxinus americana Gymnocarpium dryopteris sai latifolia 25 Lycopodium lucidulu 25 75 Maianthemum enone Mitchella repens 25 Oxalis acetosella 100 25 Picea rubens Polypodium vulgare 25 Polystichum acrostichoides 25 runus serotina Sambucus sp. Streptopus roseus Taxus canadensis Trientalis borealis Tsuga canadensis 50 25 Viburnum acerifolium Viburnum alnifolium 50 100 No. of Vascular Species 8 21 6 Atrichum undulatum Bazzania trilobata 25 Dicranum sp Hypnum sp. 100, Tetraphis pellucida 25 Thuidium delicatulum Unidentified bryophytes 100 50 100 No. of Bryophytes 5 1 i 1996] Dunwiddie and Leverett—Old-growth Forest 437 Table 8. Extended. Grey- Cold River Dunbar Fife Bk. lock Bl B2 B3 D2 El 2 3 - 3 25 fis) 23 75 50 50 25 25 1D 25 25 25 25 LOO 100 25 50 100 25 2 100 25 ZS 25 50 75 2 25 75 100 100 100 100 100 50 25 100 100 50 25 50 25 50 25 75 50 75 25 25 100 100 25 25 iD 100 75 25 25 50 25 25 2D 25 25 25 25 25 25 25 50 50 25 50 50 50 75 100 75 75 100 25 75 10 9 15 13 32 11 23 17 21 25 50 25 25 25 25 i 75 75 75 50 LOO 100 75 438 Rhodora [Vol. 98 | No. of Trees 3 8 7 6 5 4 3 2 alll = a Age Class ure |. Distribution of ages of old trees in the study plots, determined by ring counts on increment cores and estimates of additional rings to tree center. wood plots sampled within old-growth forests in the Berkshires are about 10% lower than in the Adirondack sites (Table 7), many of which appear to be in richer soils and on more level ground (pers. obs.). The CFI plot live basal areas are very similar to the nearby old-growth stands. However, live stem densities are nearly twice as high in the CFI sites. Stem densities are also 50% higher Table 9. Maximum tree sizes in old- are stands. Absolute maxima rep- resent the greatest values found over all site e maximum diameters and heights for a species seldom apply to the same tree. The average maxima apply to sites exhibiting favorable growing conditions. All units are in meters. Absolute Absolute Average Average Maximum Maximum Maximum Maximum Species Diameter Height Diameter Height Acer saccharum 1.26 41.0 0.97 33.5 Acer rubrum 0.99 32.9 0.77 28.1 Acer pensylvanicum 0.34 18.0 0.19 11.6 Betula alleghaniensis 1.24 29.9 1.02 27.1 Betula lenta 1.00 33.2 0.74 26.2 Fagus grandifolia 0.98 35.7 0.81 31.4 Fraxinus americana 1.35 39.3 1.01 35.1 Picea rubens 0.78 39.0 0.60 31.7 Pinus strobus 1.24 46.5 0.95 38.7 runus serotina 0.90 34.2 0.70 29.5 Tilia americana 0.85 35.1 0.68 30.1 Tsuga canadensis 1.30 41.2 1.04 34.2 1996] Dunwiddie and Leverett—Old-growth Forest 439 Table 10. Average basal area and density of large trees (>35 cm dbh) in Berkshire old-growth forests. % of BA % of Density Total (m?/ha) Total BA (Trees/ha) Density Hemlock 30.9 72 159 33 Hemlock-Northern Hardwood 27.8 74 114 27 Northern Hardwood 26.6 78 99 29 in the Adirondack old-growth sites than in the Berkshires. The composition of all these sites is similar, with the exception of red maple instead of sugar maple in the second growth CFI plots. The basal area and density of standing dead trees in this forest type were similar to both the hemlock and northern hardwood types (10-11% of the total), but considerably less than values reported from the Adirondacks by Leopold et al. (1988). Under- story species composition was similar in old-growth hemlock- northern hardwood stands from both the Berkshires and Adiron- dacks, with Dryopteris carthusiana a common dominant herb. Species richness was also similar to the Adirondacks (15.8 vs. 17.3 spp./0.1 ha). The density of old trees in Massachusetts old-growth forests varies widely both within and between sites. Individuals are often highly clumped, and large areas may have few old trees. Data on the denser, more clumped areas of this old tree distribution can be extracted from the study plots, which were chosen to charac- terize areas with higher densities of large and generally old trees. Large trees, arbitrarily defined here to include individuals >35 cm dbh, have an average density of 120 trees/ha in the 26 old- growth plots (Table 10). The lowest density occurs in northern hardwood stands (99 trees/ha); hemlock stands have the highest (159 trees/ha). These trees comprise an average of about one- third of the stems, but three-fourths of the basal area. In the west- ern Adirondacks, similar statistics are available in historical re- ports from uncut hardwood forests (cited in Leopold et al. 1988). Average densities of ““sound”’ canopy trees (>25.4 cm dbh) were reported at 195 (Graves 1899) and 211 stems/ha (McCarthy and Belyea 1920). Values from the Berkshire plots using this lower minimum size cutoff are 186 stems/ha for the plots in hemlock- 440 Rhodora [Vol. 98 northern hardwood forest types, and 147 stems/ha for northern hardwoods. Comparisons of these data suggest that the Massachusetts old-growth forests may have lower average densities of large trees than what was typical in similar forest types in the Adiron- dacks. These lower values in the Berkshires may result from a number of causes, including (1) slower growth rates, (2) a larger average size for the big trees, which generally is correlated with a lower average density, (3) higher probability of tree fall or other mortality in large trees, or (4) differences in sampling methods. The first and third explanations appear to be most likely, but future research should be directed towards this question for two reasons. First, the delineations of old-growth forests could be quite different, depending on what minimum density of old trees is included. Thus, determinations of the area of old growth in the state could vary considerably on this basis alone. Second, the nature of disturbances and gaps in these forests is not well un- derstood. Viable old-growth ecosystems must include areas big enough to accommodate the scale of disturbances characteristic of these stands. It is not yet clear how large these disturbances typically are in the Massachusetts forests. The data presented here help identify old-growth forest types that appear to be absent from the state today, but which may have occurred in the past. Notably, Pinus strobus L. (white pine) is missing from nearly all the old-growth stands. This species was prized by early loggers, and most stands that included significant numbers of large trees almost certainly would have been cut quickly (Pike 1967). White pine also tends to be an early-suc- cessional species and is more prone to wind damage than hem- lock; thus it is less likely to persist in great numbers in many later-successional forests. Old northern hardwood stands also are relatively uncommon. These were cut for fuel, potash, and lumber (Pike 1967), and, like the white pines, the hardwoods tend to be more prone to breakage and windthrow than the hemlocks. Thus, the probability of a stand of hardwoods reaching 250—300 years with most of its canopy trees intact approaches zero in this area. Old-growth stands of any type on level ground also are scarce. Such sites generally would have been quite accessible, and hence were likely to have been cut. No vascular plant taxa were found that were unique to the old- growth sites. The only taxon included in the list of rare species 1996] Dunwiddie and Leverett—Old-growth Forest 44] tracked by the Massachusetts Natural Heritage and Endangered Species Program that was seen occasionally was Ribes lacustre (Pers.) Poiret (Special Concern). This species also was observed in forests that were not old growth. However, other groups of organisms that were not examined extensively, including fungi, lichens, bryophytes, and invertebrates that inhabit the canopy, bark, rotted wood, and soil, may have taxa that are found pri- marily in old-growth habitats. For example, recent studies by Cooper-Ellis (1994), in several of the Massachusetts sites iden- tified in this study, noted 24 bryophyte species that occurred exclusively in old-growth forests. Various characteristics of the forest floor, including accumulations of rotted wood, microtopog- raphy such as tip-up mounds and pits, and the development of soil horizons, may also reveal some unique qualities in the old- growth sites. Whitbeck (1995) reported significantly higher amounts of woody debris in the Massachusetts old-growth hard- wood stands he studied than in second growth forests. Future research should focus on these poorly understood aspects of old-growth forests. The criteria we used to identify old-growth forests were ade- quate for providing a first approximation of the minimum extent and distribution of old-growth forest in Massachusetts. However, these criteria clearly excluded areas within old-growth ecosystems where widespread blowdowns and other disturbance processes eliminated most of the old trees. Since disturbance is certainly a component of healthy old-growth ecosystems, future work is needed to develop criteria that more adequately incorporate dis- turbance processes in definitions of these forests types, and to quantify the range of densities of old trees characteristic of dif- ferent old-growth forest types. The methods we employed to quantify most parameters were insufficient to provide statistically valid descriptions of each stand. A much larger number of randomly placed plots would be needed to adequately characterize most of the sites we visited. An expanded array of plots would not only provide a more ac- curate description of the various components of the stands, but also would serve as an excellent basis for evaluating future changes in these forests. Only about 30 old-growth stands are known in Massachusetts, with a total area of approximately 257 ha. The extremely small size of these stands, together with their small number, emphasizes 442 Rhodora [Vol. 98 the importance in preserving these few fragments that remain of the original forest that once covered much of the state. Efforts can be made to ensure that these stands are not lost to human activities in the future; however, these stands can, and will, even- tually fall to storms and other natural disturbances. The loss of large areas of several old-growth stands within the last few de- cades in the Northeast due to windthrow testifies to the ephemeral nature of this resource. Clearly, if forests that capture many o the attributes and characteristics of old-growth stands are to be conserved into the future, protection efforts must include more than the small fragments that remain. Preservation of old-growth forest in Massachusetts will require the establishment of reserves of sufficient size to accommodate natural processes of disturbance and regeneration. ACKNOWLEDGMENTS. This study was supported by contracts with the Massachusetts Natural Heritage and Endangered Species Program, for which we are extremely grateful. Field assistance was provided by Jeanne Anderson, Karen Harper, Sarah Coo- per-Ellis, Sarah Jennings, William Dunwiddie, Rob Leverett, Jr., and Phil Robakievicz. Bill Rivers provided the CFI plot data. Charlie Cogbill contributed much to the methodological and def- initional portions of this work. We also thank Pat Swain, Bill Patterson, David Foster, Jack Lash, and the many foresters and staff of the Massachusetts Department of Environmental Man- agement, who assisted in many ways with this work. LITERATURE CITED BORMANN, FH. AND M. FE BuELL. 1964. Old-age stand of hemlock-northern hardwood forest in central Vermont. Bull. Torrey Bot. Club 91: 451— CARBONNEAU, L. E. 1986. Old-growth forest stands in New Hampshire: a preliminary investigation. M.S. thesis, Univ. of New Hampshire, Dur- ham, NH. CLINE, A. C. AND S. H. Spurr. 1942. The virgin upland forest of central New England. Harvard Forest Bull. No. 21 CocBILL, C. V. 1996. An assessment of the historical ecology of the forests n the northeast slope of Wachusett Mountain, acy Report prepared ne ees Hangen Brustlin, Inc., Watertown, MA. CONARD, H. S. > P. L. REDFEARN, JR. 1979. How to Know the Mosses and Li egos "and ed. Wm. C. Brown Co., Dubuque, IA. Cooprr-E.uis, S. 1994. Ecology and distribution of bryophytes in old- 1996] Dunwiddie and Leverett—Old-growth Forest 443 growth forests of western Massachusetts. M.A. thesis, Smith College, n, MA Davis, M. B. 1996. Extent and Location, pp. 18-32. In: M. B. Davis, ed., Eastern Old-Growth Forests: Prospects for Rediscovery and Recovery. Island Press, Washington DuNWIDDIE, P. W. 1991. Survey of old-growth forest in Massachusetts. Un- published progress report to the Massachusetts Natural Heritage and En- dangered Species Program, Nantucket, MA. Survey of old-growth forest in Massachusetts. Unpublished report to the Massachusetts Natural Heritage and Endangered Species Program, Nantucket, M 1995. An evaluation “ot forest age on Mt. Wachusett. Unpublished report to Massachusetts Department of Environmental Management. Nantucket, MA. Dunwippik, P., D. Foster, D. LEopoLD, AND R. T. LevereTT. 1996. Old- Growth Forests of ae New England, New York, and Pennsylvania, pp. 126-143. In: M. B. Davis, ed., Eastern Old-Growth Forests: Pros- pects for Rediscovery and ae ees Island Press. Washington, DC. EGLeR, FE E. 1940. Berkshire Plateau vegetation, ae nee Ecol. Mono- gr. 10: 145-192. FowELLs, H. A. 1965. Silvics of forest trees of the United States. Agriculture Handbook 271. U.S. Department of Agriculture, Washington, DC. GLEASON, H. A. » A. CRoNnQuistT. 1991. Manual of Vascular Plants of Northeastern ae States and Adjacent Canada, 2nd ed. The New York Botanical Garden, Bronx, NY. GRAVES, H. S. 1899. Saal forestry in the Adirondacks. USDA. Division of Forestry Bulletin No. 26, Washington, DC. Hosier, P. E. 1969. The structure and composition of a virgin hemlock- northern hardwoods forest in northwestern Massachusetts. M.A. thesis, University of Massachusetts, Amherst, MA. oBy 1987: LEAK, haracteristics of five climax stands in New Hampshire. USDA Forest Service, Northeastern Forest Experiment Station, Research Note NE-336. LEopoLp, D. J., C. RESCHKE, AND D. S. SMiTH. 1988. Old- alas forests of Adirondack Park, New York. Nat. Areas J. 8: 166-18 LeveRETT, R. 1996. Definitions and History, pp. 3-17. In: - B. Davis, ed., Eastern Old-Growth Forests: Prospects for Rediscovery and Recovery. Island Press, Washington, DC. MAINE CRITICAL AREAS PROGRAM. 1983. Natural Old-Growth Forest Stands in aes and its [sic] Relevance to the Critical Areas — Planning Report No. 79. Maine pu e Hiannine Office, August McCarthy, 2 F AND H.C. BeLyea. 1920. Yellow birch au its relation to the Adirondack ara a mks State College of Forestry Technical Publication No. 12. Syra PIKE, R. E. 1967. Tall Trees, on aon W. W. Norton and Co., New York. SHIFLEY, S. R., EF R. THOMPSON III, R. C. SCHLESINGER, EF Conon. Jr. G.R PARKER, AND M. A. SpeticH. 1991. Study Plan: Composition and struc- ture of old-growth hardwood forests in the Midwest. Unpublished. 444 Rhodora [Vol. 98 USDA Forest Service, North Central Forest Experiment Station, Colum- bia, MO. Tritton, L. M. AND T. G. SICCAMA. 1990. What proportion of standing trees in forests of the Northeast are dead? Bull. Torrey Bot. Club 117: 163— 166. WHITBECK, M. 1995. Accumulation of fallen woody debris in old growth stands of western Massachusetts: a search for a definition of eastern old- growth. Senior project, Prescott College, Prescott, AZ. WHITNEY, G. G. 1987. Some reflections on the value of old-growth forests, scientific and otherwise. Nat. Areas J. 7: 92—99. Woops, K. D. AND C. V. CoaBILL. 1994. Upland old-growth forests of Ad- irondack Park, New York, USA. Nat. Areas J. 14: 241-257 RHODORA, Vol. 98, No. 896, pp. 445-458, 1996 COMMENTS ON THE OCCASION OF THE CENTENARY OF THE NEW ENGLAND BOTANICAL CLUB' RICHARD A. HOWARD The Arnold Arboretum of Harvard University, 22 Divinity Avenue, Cambridge, MA 02138 In December 1895, William G. Farlow wrote to a number of friends suggesting a meeting at his home on Quincy Street in ambridge to discuss the formation of a botanical club. A second meeting agreed to such an organization, and at a third meeting on February 5, 1896, the New England Botanical Club was of- ficially founded with seven professional botanists and ten amateur botanists as the charter members. We have no record of others who might have been invited or whether there were refusals. At the fourth meeting in March 1896, Messrs. Fernald, Fuller, Green- man, Jack, John Robinson, and Rich from Boston were elected members; Sears and Bailey from Providence and Jackson from Worcester became the first nonresident members. Since Fernald was a Student at the time, he became the first student member of the NEBC. With some knowledge of organizations then active, I have wondered why the New England Botanical Club was organized. Asa Gray had died in 1888 and Louis Agassiz before him, so there was no leader in the natural sciences in the Boston area. Sereno Watson, Gray’s immediate successor at the Botanic Gar- den, and his successor, Benjamin Robinson, had no major impact in Boston science or society. Charles Sargent was concerned only with the Arnold Arboretum. The Darwin controversy had subsid- ed in the Boston area. Farlow, Gray’s protégé for nonvascular plant collections, was outshone by George Goodale, who handled Gray’s teaching chores and areas of physiology, morphology, and, eventually, economic botany. Both Farlow and Goodale had sep- arated their activities and collections from the Botanic Garden and the herbarium, creating their own departments at the Agassiz ' The following is derived from the remarks delivered to The New England Botanical Club at its Centennial Banquet on November 1, 1996. Readers inter- ested in more information about the history of the Club and its founding mem- bers should refer to Dr. Howard’s paper *‘A partial history of The New England Botanical Club,” published in Rhodora 75: 493-516 (1973).—Ed. 445 446 Rhodora [Vol. 98 Museum in the campus area in Cambridge. Since Goodale was a better speaker and a more successful fundraiser for the more ap- pealing collections he curated, he was better known in the Boston area. He was one of the charter members of the NEBC. The oldest of the learned societies in America was the Amer- ican Philosophical Society, founded in Philadelphia in 1771. The American Academy of Arts and Sciences was organized in 1780. Asa Gray was an active member after his arrival in the Boston area in 1842. He served as president for 10 years (1863-1873) and published frequently in the Proceedings of the Academy. In 1791 the Massachusetts Historical Society was founded, the ear- liest in the United States, and the following year, 1792, the Mas- sachusetts Society for Promoting Agriculture was organized. The Boston area botanists were not involved in either of these. Other learned societies in New England were represented by athenaeums, Linnaean societies, lyceums, natural history socie- ties, horticultural societies, and scientific clubs. I want to com- ment briefly on each group. ATHENAEUMS The Athenaeum was“‘an institution or society for the promotion of literary and/or scientific learning.’ The first one in New Eng- land was founded in 1753 in Providence, Rhode Island. The Bos- ton Athenaeum, founded in 1807, was to have ‘‘a reading room, a library, a museum, and a laboratory.”’ It remains active as a membership organization to this day. The ‘“‘museum and a labo- ratory’’ were never developed, but an art collection begun in 1827 preceded the Boston Museum of Fine Arts by 43 years LINNAEAN SOCIETIES The Linnaean Society of London was founded in 1788. Simi- larly named societies with comparable interests developed world- wide, but without official linkage to the London group. In 1815, the New England Society for the Promotion of Natural History changed its name to the Linnaean Society of New England. This was dissolved in 1823 but re-established seven years later in 1830 as the Boston Society of Natural History. Later it was renamed the Boston Museum of Natural History and maintained its head- quarters, museum, library, series of publications, and an active 1996] Howard—NEBC Centenary Address 447 program until 1941. It then became the Museum of Science, dis- posing of its collections and the Berkeley Street building in favor of new quarters and programs in its present location on the Charles River Dam. During its most active period the Boston Museum of Natural History offered the free Lowell lectures, for which George Good- ale was a speaker six times. The Museum’s Teacher’s School of Science was instrumental in early teaching of natural history in Boston schools. Farlow gave “‘botany lectures”’ in the series, and Goodale gave ‘ta comprehensive course of twenty-one lectures on morphological, physiological, ae systematic botany to an au- dience averaging 120 students.’’ Goodale introduced the tech- nique of printed synopses of his lectures, which developed into the published Guides for Science Teaching. He supplied dried as well as fresh specimens in sets for the students to use in their teaching. A survey of Massachusetts plants was requested by the Commonwealth of Massachusetts, and the resulting work by Prof. Chester Dewey of the Berkshire Medical Institute (herbaceous plants) and George Emerson (trees and shrubs) was published by the Society between 1839 and 1846. Walker prizes for essays on designated themes or topics in science were offered between 1864 1876 and continued in recent years by the Museum of Sci- ence as awards for outstanding scientists, especially local Nobel prize winners. The Linnaean Society of New York was founded in 1878 and continues to the present with the name intact. The organization’s primary interest is in “natural science, conservation, especially ornithology, including sponsored field trips.’ It is open to ama- teurs and professionals. LYCEUMS Lyceums were created ‘‘in remembrance of the school founded by that sublime genius, Aristotle, at Athens.’’ Originally, they were debating societies or discussion groups for the ‘“‘dissemi- nation of useful knowledge.’ There is an estimate that 3,000 lyceum programs existed at one time. The Lyceum of Natural History of New York was founded in 1817. In 1833 John Torrey was a member, and in 1835 Torrey’s associate, Asa Gray, was appointed curator of the Lyceum’s botanical collections. In Mas- sachusetts there was a lyceum in Worcester in 1825 which has 448 Rhodora [Vol. 98 been renamed many times: the American Lyceum of Science and Arts for Worcester County (1826), the Worcester County Lyceum (1829), the Lyceum and Library Association (1866), the Worces- ter Natural History Society (1884), and the Worcester Science Center, before becoming the New England Science Center in 1986. In the decade between 1829 and 1838 there were active Lyceums in Millbury, Littleton, Shrewsbury, Marlborough, Pe- tersham, Barre, and at Williams College. The Littleton Lyceum remains active to the present, no longer presenting mainly lec- tures, debates, and discussions, but now offering also programs of music, mime, and drama. A cadre of speakers developed that moved from city to city presenting programs at local lyceums. Henry David Thoreau was the most noteworthy of the naturalists on the lecture circuit. No members of the future NEBC appear on program lists available. The Chautauqua Library and Scientific Society, organized in 1878 in New York state, essentially repre- sented a travelling lyceum program. HORTICULTURAL SOCIETIES The New York Horticultural Society was incorporated in 1822, and the Pennsylvania Horticultural Society was organized in 1827 and chartered in 1831. The Massachusetts Horticultural Society, founded in 1829, offered a very active and diverse program be- fore the founding of the NEBC. Flower shows were at various times offered weekly, and lectures were scheduled regularly. The Harvard Botanic Garden (with contributions from Asa Gray), the Bussey Institution (with contributions from Jackson Dawson), Charles Sargent, and, later, the Arnold Arboretum were strong supporters and exhibitors. In 1877 the Massachusetts Horticul- tural Society began a scientific lecture series. Assistant Professor George Goodale told of poet Goethe’s conclusions that parts of flowers were modified leaves. The records also report that Darwin and his writings were discussed, and surprise was expressed that Goodale differed from Darwin on many points. B. D. Halsted of the Bussey Institution talked on injurious fungi, Rev. J. V. Russell lectured on cryptogamic botany, and William Farlow talked on diseases of forest trees. In 1841, before coming to Harvard, Asa Gray presented a talk and was elected a corresponding member. He was listed as “‘professor of botany and plant physiology”’ at the Society in 1860—62, although it is noted: “‘His interest lay in 1996] Howard—NEBC Centenary Address 449 scientific study rather than cultivation and he has been of great service to the library.”’ Charles Sargent was listed as professor of botany in 1891-92. NATURAL HISTORY SOCIETIES There were many active natural history societies in New Eng- land. The Providence (Rhode Island) Franklin Society was formed in 1821, held weekly meetings, and in 1877 established a botanical section to create a catalogue of Rhode Island plants. The secretary recorded that ‘‘a feature of the meetings so far has been the attendance of a number of ladies and we can safely affirm an increasing interest in our beloved science in this com- munity.” Essex County had a Natural History Society (1833) and the Essex Institute (1834); the two merged in 1848 and created de- partments of natural history, history, and horticulture. Flower shows were held regularly, especially exhibitions of vegetables. In 1854 J. EF Allen had a fine garden in Salem and must have had greenhouses, for he published an elaborate elephant folio on Victoria regia, which had been brought into flower. George Pea- body was born in Danvers, Massachusetts, in 1795. He amassed a fortune in Anglo-European trade and made generous gifts to New England. In 1853 he made a gift of $100,000 to Yale, where his nephew was the paleontologist, Prof. Marsh. In 1862 Peabody promised Harvard President Edward Everett $100,000 in support of astronomy and art, but the local scientific circles “‘helped di- rect’ this gift to the design of a museum of archeology and an- thropology. Peabody died in 1867, leaving a bequest of $140,000 to support the study of natural history in Essex County. The Pea- body Academy of Sciences was formed in 1868 with this money and with Asa Gray as one of its trustees. Its publications had few articles on botany. The outstanding were Robinson’s lists of ferns of North America north of Mexico, later revised by D. C. Eaton, and an article on the strength and fuel value of native woods, which Robinson had extracted from Sargent’s report on the forests of North America. The lecture series and courses at the Peabody Academy featured offerings in botany by Goodale, Robinson, John Sears, C. E. Bessey, and D. P. Penhallow. Between 1876 and 1881 the Academy offered a Summer School in Biology pat- terned after Agassiz’s school on Penikese Island. Dr. Packard of 450 Rhodora [Vol. 98 Peabody was the director, and John Robinson, Goodale, Penhal- low, and Bessey were the instructors. In 1880 the Academy de- cided not to compete with the Boston Museum of Natural History or the Museum of Comparative Zoology in acquiring general col- lections, but to concentrate on works of man and the flora and fauna of Essex County. In 1922 the complex was renamed the Peabody-Essex Museum. An herbarium is maintained to the pres- In 1834 the Yale Institute of Natural Sciences was formed and was to become the Yale Natural History Society. The Natural History Society of Hartford (1835) became, in 1845, the Con- necticut Society of Natural History. The Maine Institute of Nat- ural History (1836) became the Portland Society of Natural His- tory (1843). The Cuvierian Society of Wesleyan University was alternately known as the Natural History Society of Wesleyan University. In Cambridge, Massachusetts, students organized the Harvard Natural History Society in 1837, creating a cabinet and housing specimens in Massachusetts Hall. Asa Gray lectured to this group on April 18, 1878, on forest geography and archeology. Benjamin Watson, later a charter member of the NEBC, was a member when a student at Harvard. Little more has been located of this club or its collections. Other natural history groups were formed in Lynn, Massachu- setts (1842), Concord, New Hampshire (1846), and Springfield, Massachusetts (1859); other similar organizations were the Bos- ton Amateur Scientific Society (1877), Middlesex Field Club (1881), Natural History Club of West Roxbury (1882), Natural History Club of Boxford, Massachusetts (1882), and the Newport Natural History Society of Rhode Island (1883). Perhaps the most ambitious amateur natural history undertak- ing was the Agassiz Association, developed in 1875 by Harlan Hage Ballard of the Lenox Academy in western Massachusetts. The Association was formed while Louis Agassiz was alive, and with his permission. To encourage the study, discussion, collec- tion, and exchange of natural history objects, Ballard encouraged the formation of chapters. Four people could form a chapter and name it. They could be of the same sex or age, as adults, families, or children. At its peak the Association had 1200 ‘‘chapters”’ of 4 to 120 members, with a total of 20,000 members in the United States, Canada, England, Ireland, France, Chile, New Zealand, and Japan! Chapters had meetings, conducted field trips, ex- 1996] Howard—NEBC Centenary Address 451 changed artifacts, and published newsletters. Correspondence with Ballard was encouraged, and he would redirect the letter to a specialist for a personal reply and future contact. With permis- sion, specimens or artifacts could be submitted for identification, at which time the enclosure of five two-cent stamps to cover postage was requested. An Association pin was available through the manufacturer, who would supply a price list. A chapter cer- tificate was priced at 75 cents. A handbook of the Agassiz As- sociation called Three Kingdoms was published in at least four editions. The fourth, in 1897, mentioned need for an endowment to maintain the organization with a paid staff, but I can find no reference to dues or of payment to Ballard. Ballard maintained contact with scientists in several countries to help answer ques- tions submitted or to conduct correspondence courses. Marcus Jones, E. L. French, W. O. Crosby, Alexander Wight, W. Whitman Bailey, William B. Werthner, John Davis, Jacob Bigelow, and Francis Gay were among the specialists who cooperated with identifications, and who offered correspondence courses. In 1887 the Asa Gray Memorial Botanical Chapter was formed, which eventually had five geographic divisions and a total membership of 44. The chapter issued a publication which became the Asa Gray Bulletin, as well as Reports and the Observer. Also in 1887 there is casual mention of the Asa Gray Botanical Club in Utica, New York, which purchased the Hunt Herbarium and gave it to the Oneida Historical Society. The Linnaean Fern Chapter was also a spinoff of the Agassiz Association in 1893. This group eventually became the American Fern Society and later spawned the Bryological Society. SCIENTIFIC CLUBS In 1854 Asa Gray assembled his scientific colleagues, mostly all professors, to regular meetings for professional discussions. No records of a club have been located. The most successful and longest lasting of the scientific clubs is certainly the Torrey Botanical Club of New York. It began in 1867 with informal meetings with Dr. Torrey. When William H. Leggett compiled a modest four-page monthly news sheet, he named it the Bulletin of the Torrey Botanical Club, and the name was adopted. The Bulletin became the first botanical periodical in America, and was dedicated to the study of the flora of the 452 Rhodora [Vol. 98 New York City area. There followed a series of noteworthy con- tributions. In 1873 a Botanical Directory of North America was the first such publication. In 1886 the Index to Recent American Botanical Literature began under the editorship of Elizabeth Brit- ton and Frederick Merrill. It also appeared as a card index in 1890 but was stopped by the Club in 1996 and is being continued in Brittonia by editors at the New York Botanical Garden. The Memoirs of the Torrey Botanical Club met the need, in 1889, for a regular series of publications in monographic form. The first number had an article by Liberty Hyde Bailey entitled ‘Studies of types of various species of Carex.”’ In 1901 Torreya was cre- ated to handle brief communications and those of a popular char- acter. This was combined with the Bulletin in 1946. Beginning in 1997 the Torrey Botanical Club will be renamed a Society and the Bulletin will become the Journal of the Torrey Botanical So- ciety. Sometime in the 1870s ‘ta few ladies of this city (Syracuse, New York) have formed a club named the Rust Botanical Club, of which Mrs. Rust has been elected president. At first we shall make the study of ferns our specialty, hoping afterwards to study general botany.”’ In 1879 it is reported that the president objected to the eponym, and it was changed to the Syracuse Botanical Club. The Club was “exclusively” for ladies. Numbering 32, they had field meetings twice a week in season, weekly meetings for study, and a business meeting on the third Monday of eac month. The Club reported regularly in the Bulletin of the Torrey Botanical Club and by correspondence to Cambridge botanists Gray, Eaton, Peck, Robinson, and Davenport. They were proud of tracing the routes of Frederick Pursh and rediscovering Scol- opendrium. In 1912 A Flora of Onondaga County as Collected by the Members of the Syracuse Botanical Club was published. A photograph in the Gray Herbarium archives shows the group of ladies in the field on July 16, 1915. The demise of the Club is not known. In 1874 there was a Connecticut Valley Botanical Society which had ‘‘a very animated meeting at Mt. Holyoke Seminary Wednesday, June 10th.’? At a meeting on October 6, 1875, Pro- fessor Asa Gray was in attendance. The American Botanical Club was formed in 1883 by members of the American Association of Arts and Sciences for the study of plants by correspondence. In 1892 this became the Botanical 1996] Howard—NEBC Centenary Address 453 Society of America, which “‘limited its membership to those who have published worthy work and are actively engaged in botanical investigations.” Eighteen ninety-five proved to be an exceptional year for the formation of botanical clubs. On July 4th a group gathered to explore a mountain plateau in Vermont and to form an association which became the Vermont Botanical Club. Ezra Brainerd of Middlebury and L. R. Jones of Burlington were Vermont resi- dents. Summer visitors included Rev. James A. Bates of South Royalston, Massachusetts, and Marshall Howe, W. W. Eggleston, and A. J. Grout of New York City. Cyrus Pringle and George Perkins of Burlington joined immediately, as did Barnhart of New York. The group planned one summer meeting for a field trip and one winter meeting for papers. Between 1906 and 1914 the Club published a Bulletin. Fernald wrote an article “‘“Some Northern Plants Possibly to be Found in Vermont’’ for the third bulletin, issued in 1908. On July 12 another group of amateur botanists assembled in Portland, Maine, to organize a botanical club. Asa L. Lane was the first president; Kate Furbish, then 61, was elected vice pres- ident, and Merritt Fernald was the first recording secretary. It was Fernald who proposed the name Josselyn Botanical Society to honor John Josselyn’s Rarities, the first written account of the Maine flora. It was suggested that ‘“‘Club’”? was an overworked title; hence the name Josselyn Botanical Society of Maine was adopted. This group has published a Bulletin at irregular intervals, the last being Number 13 in 1995. There is no clear answer to the question of why the New Eng- land Botanical Club was proposed in 1895 and organized in 1896. There were active organizations in the Boston area involving both professional and amateur botanists. Not all clubs admitted wom- en, and the NEBC began as a gentlemen’s organization. Charter members comprised seven professional botanists and ten ama- teurs. Brief biographies of them are supplied in previous articles cited in the closing suggestions for additional reading. The group had little in common socially, and varied in professional roles. Nevertheless, the Club flourished for a century, and shows no signs of weakening in its goals or accomplishments. 454 Rhodora [Vol. 98 TRIVIA FROM THE NEBC’S FIRST HUNDRED YEARS M. L. Fernald was the first student member of the NEBC, being elected at the fourth meeting in 1896. He edited Rhodora for 23 years. When he died in 1950 he had been an active Club member for 54 years. David Pearce Penhallow of McGill University was possibly the first foreign member of the Club, having joined November 6, 1896. Mary Day, librarian of the Gray Herbarium, published in volume 3 of Rhodora, in 1901, a list of 66 private herbaria and a list of 258 local floras as an aid to Robinson in compiling distribution data for the 7th edition of Gray’s Manual. The choice of Rhodora for the title of the Club’s publication was to indicate a primary interest in plants of the same range. Initially, Club members were to concentrate on plants of the Boston region and of the mountains in New England. Advertisements in Rhodora in 1902 drew published criticism from other botanical journals and had to be defended in editorials. The advertisements by the Bangor and Aroostook Railroad stated, “If you are looking for the Best Botanizing in the eastern states, you should save up pennies enough to visit The Aroostook.” Among the plants which could be collected easily was Pedicu- laris furbishii, which in recent years was declared to be an en- dangered species. The editor replied: “‘Indeed the flora of no other area east of the Rocky Mountains and south of British America seems to us less in need of concealment or special protection.” The NEBC may well be the first to drop page charges to its publication on January 27, 1996 The NEBC herbarium was started immediately upon the Club’s organization. For years it was fumigated with carbon bisulfide, fortunately with no casualties. This dangerous, poisonous, and flammable chemical was proposed for insect control in the Bul- letin de la Société Botanique de France in 1877. 1996] Howard—NEBC Centenary Address 455 In 1996 an agreement was made to integrate the NEBC herbarium with the general collections of the Harvard University Herbaria. Currently the NEBC records 251,000 specimens. Fellowship prizes were offered by the NEBC for the first time in 1985 For many years candidates for admission were nominated and supported by statements to the membership. “‘Scrutineers’’ were appointed and written votes counted and reported. When there were negative votes one such blackball disqualified the candidate. Women were elected to the Club for the first time on December 6, 1968. Alice Tryon was the first woman elected and the first to serve as President of the NEBC in 1978. The early meetings of the Club were held in members’ homes and subsequently at Young’s Hotel and the Twentieth Century Club, and fancy meals were served. Pease reported: ‘“‘Ample col- lation—e.g., chicken croquettes, cold joints, salads, escalloped oysters, and ice cream replenished by one or two bustling wait- ers.”” With food supplies limited during World War I, the refresh- ments dropped to crackers and cheese. The oldest member in longevity was Harold St. John (1911) who died December 12, 1991, at the age of 99 years and five months. The oldest current member is Wayne Manning (1934), who at 97 years answered the telephone in his office. Lyman Smith (1923) and Aaron Jack Sharp (1944) are both 92 at this writing (January 1997). Bill Benninghoff, elected in 1940, died in 1993 and is buried in Arlington National Cemetery. The gavel used by the President of the NEBC was turned from a “‘corner post’? of Asa Gray’s home on Garden Street. The first field day of the NEBC was on May 30, 1911, to Prov- idence, Rhode Island. 456 Rhodora [Vol. 98 George Cooley (1953) fell into a volcano on St. Vincent, but his slide was stopped by grasping a bromeliad. He received an hon- orary Doctor of Science degree from the new University of South Florida before the school had awarded any degree. Six members of the Club were presidents or chancellors of seven universities. Edward Rand was called the Bard of the NEBC for his poem, ‘In the days of Walter Deane.” It was sung to the tune of John Brown’s Body. The text and other poems are printed in Rhodora (75: 493-516. 1973). Perhaps the most quoted poem in the ar- chives is this joke on Arthur Stanley Pease (1902): Here lies the body of Stanley Pease Buried beneath the venerable trees What lies here is only the pod Pease shelled out and went to God. No poet laureate for the NEBC has surfaced in recent years, but I felt this centenary deserved some poetic recognition. I’m not a poet, so I asked a friend in Acton who has notable talents in this area to prepare an ode for this occasion, the 100th anni- versary. I close with these words supplied by Maurice Sagoff: If you’ve enjoyed the historical show You'll surely be pardoned for feeling a glow Of real satisfaction and well-deserved pride In what’s been portrayed by each story and slide. So here’s to our Club! Though a hundred years old, Its greater achievements have yet to be told. We count in our membership leaders in botany, Proven achievers; some clubs haven't got any. Members of fishing clubs may not have caught any. Hunt clubs don’t care if you never have shot any. Teachers in unions may hardly have taught any. Even some legionnaires may not have fought any! But, to our credit, our workers in botany Truly have shown they are worthy of fame. I'll mention a dozen, at random, by name: There’s Fernald and Merrill and Harris and Bean, 1996] Howard—NEBC Centenary Address 457 There’s Bogle and Hodgdon and Walker and Pease, Barrington, Weatherby, Nickerson, and Crow, Eaton, and Knowlton and Ray Angelo; If your name is not here with our sisters and brothers, I’ll add to the list the addendum—‘“‘and others.” Our members’ professional writings abound In volumes of vital research: we have found Botanical treasures the world never knew As well as a gaggle of trivia, too; For instance, one member has found that Linnaeus Once grumbled, “‘No matter how much they may pay us, You can be sure that the wicked economy Shortly will lay a tremendous Tax-on-o-me!”’ (That derivation is open to question, But someone will find a more valid suggestion.) Now, is there a club that can mark its centenary So free of corruption and scandal and venery? Minds so devoted to pleasures botanic That some of our spouses regard us as manic! Some of this group are revisiting Harvard, Some come to find out what’s been lately ‘“‘discarvered;”’ Some meet their old colleagues and thump on their shoulder And tell them they don’t look a single bit older! Some come to relax and perhaps get a snoot-full, But for most, the occasion is sober and fruitful,— A time to reflect on the glorious past, And a time when our eye to the future is cast; The next hundred years for the students of botany Surely will chase any thoughts of monotony— Challenges, problems, there will be no dearth Of issues that threatened the health of the earth, But well in the forefront of progress will be The men and the women of the N.E.B.C. ADDITIONAL READING DEANE, WALTER. 1899. The herbarium of the New England Botanical Club. Rhodora 1: 56-57. HOwaArp, R. - 1973. A partial history of the New England Botanical Club. Rhodora 75: 493-516. 458 Rhodora [Vol. 98 . 1986. The New England meee Club 800th meeting. [A tribute o M. L. Fernald.] Rhodora 88: 159-228. eon C. H., M. L. FERNALD, AND FE G. Ftoyp. 1912. Field excursions of the New England Botanical Club. Rhodora 14: 71—76 Pease, A. S. 1951. The New England Botanical Club a half-century ago and later. Rhodora 53: 97-105. WILLIAMS, E. E 1899. The New England Botanical Club. Rhodora |: 37-39. RHODORA, Vol. 98, No. 896, p. 459, 1996 BOOK REVIEW Flora of Berkshire County, Massachusetts by Pamela B. Weather- bee. 1996. 135 pp. 48 color photos. ISBN 0-964821 1-0-9. $20.00 (paper). Berkshire Museum, Pittsfield, MA. This highly readable and accessible modern flora provides a wealth of information on the history, geology, floristics, vegeta- tion communities and vascular plant species of Berkshire County. The habitat diversity of this 940-square mile region, which ex- tends from Connecticut to Vermont, includes calcareous, serpen- tine, and metamorphic substrates and has a 2900-ft. elevation range, contributing to the occurrence of the 37 distinct plant com- munities for which Ms. Weatherbee provides comprehensive and detailed descriptions. The book contains considerable information on each of the 1,675 taxa recorded from the county: whether native or intro- duced, frequency and abundance, habitats, flowering and fruiting dates and distribution within the county. Genera and species are in alphabetical order and nomenclature generally follows Kartesz. Eight color plates illustrate rare and uncommon species. Ms. Weatherbee also provides statistics on floristic change; exotics account for 27 percent of the taxa, and 35 native taxa and 107 aliens have migrated into the county since the last comprehensive survey in 1922. The book also includes a list of species which are occasionally observed but are nonpersistent. I highly recommend this volume as a model for modern flo- ristic treatments in New England. It provides a comprehensive picture of the flora of this complex segment of the region, and an engaging and well-written introduction to the habitat charac- teristics that account for species distributions and the occurrence of plant communities in western Massachusetts. —LIsA A. STANDLEY, VHB, Inc., PO. Box 9151, 101 Walnut Street, Watertown, MA 02272. 460 Rhodora [Vol. 98 RHODORA, Vol. 98, No. 896, pp. 460-462, 1996 BOOK REVIEW Michigan Flora, Part II, Dicots (Pyrolaceae—Compositae) by Edward G. Voss. 1996. xix + 622 pp. 771 maps, 266 line drawings, 8 color plates. $15.00 (cloth). Cranbrook Institute of Science Bulletin 61 and University of Michigan Herbari- um, Ann Arbor, MI. Part of a project begun more than 40 years ago, this book completes a monumental study of the seed plants of Michigan. Michigan Flora, Part III, by Edward G. Voss, is an excellent book in an excellent series. Part J (Voss 1972) covered the gym- nosperms and monocots, Part // (Voss 1985), the apetalous and polypetalous dicots, Part I/II, the sympetalous dicots. Save for the absence of the pteridophytes from the Michigan series, its three parts parallel both in coverage and family sequence the three volumes of Gleason’s (1952) New Britton and Brown Illustrated Flora. In the course of the requisite research for the three books, Voss studied more than a quarter million specimens collected from Michigan during the 19th and 20th centuries. He estimates that the number of Michigan specimens in the state’s several herbaria has more than doubled over the past 40 years. A similar estimate would likely be valid for most states of the Midwest. In all, the three books contain county dot distribution maps for ca. 2,450 species, each dot substantiated by at least one of the herbarium specimens. The maps are rich with dots, and the distribution pat- terns they display are interesting and in some cases provocative. They remind one of the need for an atlas of plant distribution covering all the states of the Gray’s Manual range (Fernald 1950 Michigan Flora, Part III treats the plants of 33 dicot families, those of the large family Compositae or Asteraceae occupying about 40% of the taxonomic text. The text includes thorough, usually multi-charactered, dichotomous keys for species identifi- cation. The keys are often augmented by discussion of additional character-state differences, combinations, and nuances that further serve to distinguish the various species, hybrids, and infraspecific taxa. The comprehensive habitat statements will also, in some 1996] Book Review 461 cases, aid in the identification of unknowns. About one-third of the species are illustrated by habit sketches and/or drawings of diagnostic morphological details. The taxonomic section opens with eight plates of attractive color photographs of some 50 of the species. The section closes with a set of master keys to fam- ilies and to special groups, e.g., aquatic plants, woody plants, covering all three parts of the Michigan Flora. Nuggets abound in the discussion of individual genera and spe- cies. A few examples are: a brief history of the harvesting of floss from the seeds of common milkweed during World War II by the “Milkweed Floss Division of War Hemp Industries,” headquartered at Petoskey, Michigan; the speculation that Cyno- glossum boreale was the blue-flowered borage noted by Henry David Thoreau during his travels in Michigan in 1861; a discus- sion of the commercial growing of peppermint in Michigan, in- cluding notes on the establishment of the town of Mentha— whose post office closed in 1954; the observation that in most Michigan species of Viburnum a fruit is purple-black when ma- ture, ‘although it may be red when it is green”? (ouch!); some thoughts on the gender of the generic name Bidens, a discourse on taxonomic problems in the genus Aster; and the comments on the ‘‘tragic spread”’ of the European thistle Cirsium palustre into natural wetlands. The three-part Michigan Flora will be of great value to pro- fessional botanists and botany students dealing with plants of the Midwest, the Northeast, and Ontario. It will be of great utility to all individuals whose interest or work brings them into contact with the region’s plant life, such as those concerned with the identity of plants growing in wetlands or lakes, the occurrence of trees or wildflowers, the relationship between plants and insects or between plants and soils, and the preservation of rare species or important natural areas. ‘‘A flora,’ as Voss writes, “is an in- ventory or census of a basic resource.” LITERATURE CITED FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book o., New York. GLEason, H. A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. The New York Botan- ical Garden, Bronx, NY. 3 vol. Voss, E. G. 1972. Michigan Flora, Part I, Gymnosperms and Monocots. 462 Rhodora [Vol. 98 Cranbrook Institute of Science Bulletin 55 and University of Michigan Herbarium, Ann Arbor, MI 1985. Michigan Flora, Part II, Dicots (Saururaceae—Cornaceae). Cranbrook Institute of Science Bulletin 59 and University of Michigan Herbarium, Ann Arbor. —Tom S. CooPeRRIDER, Department of Biological Sciences, Kent State University, Kent, OH 44242 Note: Parts I and II are available for $12.50 each from the Cranbrook Institute of Science, 1221 N Woodward Ave., PO. Box 801, Bloomfield, MI 48303-0801 (phone 810-645-3239 or 810- 645-3203 ).—Ed. RHODORA, Vol. 98, No. 896, pp. 463-467, 1996 NEBC MEETING NEWS September 1996. Pamela Weatherbee spoke on “Mount Grey- lock, Haven for the Berkshire Flora.’’ Although it may be the world’s smallest mountain, Greylock has a long and interesting history and an unusual flora. The mountain rises abruptly 2800 feet in elevation from the valley floor, resulting in sharp distinc- tions between vegetation zones and severe summit weather. The geology of Mount Greylock is responsible for much of the di- versity of the flora. The hard, resistant and somewhat acidic schists on the summit provide a very different substrate from the calcareous marbles along the mountain’s flanks. After reviewing the history of the mountain, Pam led the Club on an armchair tour of the diverse plant communities on Mount Greylock, starting at the summit where northern species such as Sorbus decora, Amelanchier bartramiana, Solidago macrophylla, Euphrasia, Streptopus amplexicaulis, Ledum groenlandicum, Bet- ula cordifolia, and the state’s only Vaccinium vitis-idaea occur. Boggy areas near the summit support Milium effusum and Gaul- theria hispidula. In the Northern Hardwoods zone, species include Platanthera macrophylla, P. grandiflora, P. lacera and their hybrid, P. x andrewsii, in addition to more typical herbaceous and shrub spe- cies. The rich sugar maple forests on marble support a very di- verse herbaceous flora, including wild leeks, Dicentra, Hepatica, Viola canadensis and V. rostrata, Disporum, Caulophyllum, Uvu- laria grandiflora, Orchis spectabilis, Panax quinquefolius, Hy- drophyllum canadense, Conopholis, Botrychium matricariifolium, and Camptosorus, as well as the hairy woodmint. Steep ravines are a high-disturbance zone, and support Ribes lacustre, R. triste, Conioselenium chinense, and Polystichum braunii along the cold, rushing brooks. Cliffs provide habitat for Woodsia glabella, Cryptogramma, and Potentilla tridentata. September Field Trip. On Saturday, September 14th, 20 NEBC members and guests met at the summit of Mount Grey- lock, in the fog, to experience the flora described at the Club meeting. Led by Pam Weatherbee, we started with “parking lot”’ botany at the summit where, accompanied by a cellist, we saw Sorbus decora in fruit (and compared it to S. americana), Ame- 463 464 Rhodora [Vol. 98 lanchier bartramiana, wind- and ice-shaped balsam firs, and Eu- Phrasia in bloom. The sole site of Vaccinium vitis-idaea in Mas- sachusetts was observed on the drive to the ‘“‘boreal forest’? zone, where the group was introduced to Betula cordifolia, Dryopteris campyloptera, Solidago macrophylla, Carex intumescens, and As- ter acuminatus. Carex gynandra, C. trisperma, Chelona glabra, and Nemopanthus mucronatus in fruit were highlights of a small summit bog, along with two sharp-shinned hawks. A wet glade in the “northern hardwoods” forest lower down the mountain provided Glyceria melicaria, Carex scabrata, Milium effusum, Laportea, Ribes triste, and Cinna latifolia. The final highlights of the trip were along a steep stream ravine, where we found Solidago flexicaulis, Ribes lacustre, Caulophyllum thalictroides, Allium tricoccum, Asarum canadense, Phegopteris hexagonop- tera, a two-lined salamander, and (growing together) Polysticum acrostichoides, P. braunii, and their hybrid, P. X potteri. October 1996. Dr. Thompson Webb III of Brown University spoke on “‘Vegetational History of New England and Eastern North America: a 20,000 year perspective.’’ The talk was based on a recently published book, Global Climate since the Last Gla- cial Maximum, produced by the Cooperative Holocene Mapping Group and edited by Herb Wright. The book integrates vegetation data derived from pollen analyses with climatic data from lake levels to produce computer simulations of climate that are useful both in documenting the past and providing a test for predictive climate models. Pollen analyses have some severe limitations for understanding vegetation. Since these are based on anemophilous species, pollen records show only a small portion of the flora and may represent species from several plant communities. Recent work has shown that the relative abundance of tree species and pollen deposition correlate best at the continental scale, where climate is the most important factor controlling plant distribution. A quick overview of the history of New England flora shows that spruce and pine dominated the vegetation before about 9000 years ago, with oak and pine more recent dominants. Ragweed pollen appeared fol- lowing the European colonization and land clearing, and is prob- ably the first source of New World air pollution. Pollen data show that plant associations were very different at past times than we know them today, and that distributions have 1996] NEBC Meeting News 465 changed substantially. Spruce/fir forests, for example, have oc- curred only recently. Following the last glaciation, species did not simply migrate south and north in response to the advancing and retreating glaciers, but individually responded to changes in climate. Some species were rare everywhere at the glacial max- imum but increased in abundance after glaciation, spreading along the glacial front. Dr. Webb concluded the presentation with dynamic represen- tations of the change in plant distributions over time, using a video film of a computer 3-dimensional model. November 1996. The Centennial Banquet was held at the Har- vard Faculty Club with 85 members and guests present, including six who had been members for over 50 years. Dr. Richard Howard, a member since 1940, spoke on “An Illustrated History of the New England Botanical Club.’’ Former Director of the Arnold Arboretum and Science Director at the New York Botanical Garden, as well as Past President of the NEBC, Dr. Howard was the featured speaker at the 700th and 800th meetings of the Club and holds the record for speaking at commemorative Club milestones. Dr. Howard provided an over- view of the environment in which the Club emerged. Many of the extant scientific organizations have their roots in this period, including the American Philosophical Society, AAAS, the Mas- sachusetts Horticultural Society, and the Massachusetts Audubon Society. The NEBC is unusual in that it has operated for a century without a change in name or organization. Key prior organizations included the Linnaean Society of New England, which became the Boston Museum of Natural History and eventually was re-established as the Museum of Science. Nat- ural history societies were founded in several locations, including the Essex County Natural History Society which merged with the Essex Institute, and later became the Peabody-Essex Museum. Others included the Agassiz Association, which began in Lenox and comprised 20,000 members worldwide at its peak but ex- pelled members who failed to supply exchange specimens, the Cambridge Science Club, the Torrey Botanical Club (the oldest botanical club), the Connecticut Valley Botanical Society, the American Botanical Club (now the Botanical Society of Ameri- ca), and the Josselyn Botanical Society, founded in 1895 with Kate Furbish as Vice President. This was an era of intense interest 466 Rhodora [Vol. 98 in the local flora. Mary Day, the legendary Gray Herbarium li- brarian, identified 61 private herbaria in New England in the early 1900s, and Robinson cited 365 individual local floras in his edi- tion of Gray’s Manual. The New England Botanical Club was founded a year later, in 1896. The Club’s founders, particularly George Goodale, had all been involved in other local scientific clubs and organizations. The goals of the founders were to organize a club to study the New England and alpine flora. The name of the Club’s journal, Rhodora, was deliberately chosen to correspond with the goal of studying the flora in the natural range of Rhododendron cana- dense. Dr. Howard provided brief sketches of each of the Club’s founding members. Among these gentlemen were: J. R. Chur- chill, a judge and banker; W. G. Farlow, to whom we owe the current old herbarium building (he left his collection to Harvard on the condition that it be moved to a fireproof building); C. E. Faxon, who drew the plates for Sargent’s Silva of North America, despite his training as an engineer; G. L. Goodale, who was a highly successful fundraiser and apparently tireless teacher; E. L. Rand, who focused on Mt. Desert; C. S. Sargent, who served as Director of the Arnold Arboretum for 54 years; R. Thaxter, the first Club President to offer travel talks; and B. M. Watson, cred- ited with introducing the Japanese honeysuckle. Dr. Howard concluded with an eclectic list of NEBC trivia and doggerel, to the appreciation and amusement of those present. December 1996. Dr. David Barrington of the University of Ver- mont concluded the Centennial Year presentations with a talk entitled ““What We Don’t Know About the New England Flora and Why We Want To Know It.”” Some of the major unknown areas of the New England flora are those that first interested the founders of the Club—disjunctions in distributions and the ori- gins of the flora. Using examples from research in his lab, Dave showed how modern techniques are expanding our knowledge of the evolution of New England taxa through both reticulate and divergent evolution, resolving old taxonomic problems and shed- ding light on evolutionary origins. Examples that Dave discussed include the tetraploid/diploid complex within Asplenium trichomanes, the two allotetraploids Cystopteris tenuis and C. fragilis, the serpentine endemic allote- traploid within the Adiantum pedatum complex, the as-yet unre- 1996] NEBC Meeting News 467 Ll M tr solved relationships within the Dip complanatum/dig- itatum complex, and the evolation of ecnetcally distinct groups within the Ammophila breviligulata complex. In each of these cases the application of new theories of systematics and new and powerful molecular tools has, in combination with detailed mor- phological studies, allowed us to understand the evolutionary re- lationships and history of the taxa in the New England flora. In several instances, however, this has created problems for the con- servation of biological diversity, when science recognizes taxa that are evolutionarily distinct but morphologically cryptic. —Lisa A. STANDLEY, Recording Secretary. REVIEWERS OF MANUSCRIPTS 1996 The editors of Rhodora are grateful to each of the following Lawrence A. Alice David S. Barrington William Brumback Ann Bruneau Charles Cogbill Tom S. Cooperrider Howard Crum Alison C. Dibble Joseph J. Dowhan Arthur Haines Matthew Hickler Larry Hufford specialists for their participation in the review process. Susana Magallon Steven R. Manchester T. Lawrence Mellichamp Norton Miller Robert Naczi Donald Padgett John Pruski Paul E. Rothrock Gordon C. Tucker Lowell Urbatsch Thomas E Vining Warren H. Wagner, Jr. INFORMATION FOR CONTRIBUTORS TO RHODORA Submission of a manuscript implies it is not being considered for publication simultaneously elsewhere, either in whole or in part. GENERAL: Manuscripts should be submitted in triplicate. The text must be double-spaced throughout, including tables, figure legends, and literature citations. Use a non-proportional font throughout and do not justify the right margin. Do not indicate the style of type through the use of capitals, underscoring, or bold, except for names of genera and species which should be in italics or underscored throughout. Do not underline punctuation. All pages should be num- bered in the upper right-hand corner. For guidance in matters not addressed here, consult the editorial office by phone at (603) 862- 3205, FAX (603) 862-4757, or e-mail: janets@christa.unh.edu. Brev- ity is urged for all submissions. TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital letters. Omit authors of scientific names. Below title, include au- thor(s) name(s), affiliation(s), and address(es). If ‘‘present address”’ is different, it should follow immediately below, not as a footnote. ABSTRACT: An abstract and a list of key words should be included with each paper, except for shorter papers submitted as Notes. An abstract must be one paragraph, and should not include literature citations or taxonomic authorities. Please be concise, while including information about the paper’s intent, materials and methods, results, and significance of findings. TEXT: Main headings are all capital letters and centered on one line. Examples are: MATERIALS AND METHODS, RESULTS, and DIS- CUSSION. Do not title the Introduction. Do not combine sections of the paper (such as Results and Discussion), or use Conclusions or Summary. Second level headings should be indented, bold, upper and lower case, and end with a period. Taxonomic authorities should be cited for all species names at their first usage in the text, or in a referenced table. Cite each figure and table in the text in numerical order. Each reference cited in the text must be in the Literature Cited. Cross-check spelling of author(s) name(s) and dates of publication. Literature citations in the text should be as follows: Hill (1982) or (Hill 1982). For two or more authors, cite as follows: Angelo and Boufford (1996) or (Angelo and Boufford 1996). Within parentheses, use a semicolon to separate different types of citations (Hill 1982; Angelo and Boufford 1996) or (Figure 4; Table 2). FLORAS AND TAXONOMIC TREATMENTS: Specimen citation should be selected critically, especially for common species of broad distribution. Keys and synonymy for systematic revisions should be prepared in the style of ““A Monograph of the Genus Malvastrum,”’ 469 470 S. R. Hill, RHODORA 84: 159-264, 1982. Designation of a new taxon should carry a Latin diagnosis (rather than a full Latin descrip- tion), which sets forth succinctly how the new taxon differs from its congeners. LITERATURE CITED: All bibliographic entries must be cited in the paper, unless a special exception has been made by the Editor (such papers will be allowed a REFERENCES section). Verify all entries against original sources, paying special attention to spelling and de- tails of publication. Cite references in strict alphabetical order by first author’s surname. Do not write authors’ names in all capital letters. References by a single author precede multi-authored works of same senior author, regardless of date. Use a long dash when the author(s) is the same as in the entry immediately preceding (see recent issues). Refer to Botanico-Periodicum-Huntianum (B-P-H 1968) and B-P- H/Supplement (1991) for standardized abbreviations for journals. TABLES: Tables must be double-spaced. Tables may be continued on an extra page, if necessary. As much as possible, the title should be self-explanatory. Do not use footnotes; instead, add notes after the end of the table title. Broadside tables should be avoided, if pos- sible. Each table should be cited in the text in numerical order. FIGURES: Illustrations must be either black and white half-tones (photograph), drawings, or graphs. Illustrations must be camera- ready; flaws cannot be corrected by the Editor or the printer. Add symbols or shading with press-on sheets. The printed plate will be 4 X 6 inches; be sure that illustrations are proportioned to reduce correctly. Allow space for a caption, if possible. Magnification/re- duction values should be calculated to reflect the actual printed size. Maps must indicate scale and compass direction. The double-spaced list of legends for figures should be provided on a separate page. Each figure should be cited in the text in numerical order. THE NEW ENGLAND BOTANICAL CLUB 22 Divinity Avenue Cambridge, MA 02138 The New England Botanical Club is a nonprofit organization that promotes the study of plants of North America, especially the flora of New England and adjacent areas. The Club holds regular meetings, and has a large herbarium of New England plants and a library. It publishes a quarterly journal, RHO- DORA, which is now in its 99th year and contains about 400 pages per volume. Visit our web site at http://www.herbaria. harvard.edu/nebc/ Membership is open to all persons interested in systematics and field botany. Annual dues are $35.00, including a subscription to RHODORA. Members living within about 200 miles of Boston receive notices of the Club meetings. To join, please fill out this membership application and send with enclosed dues to the above address. Regular Member $35.00 Family Rate $45.00 Student Member $25.00 For this calendar year ee For the next calendar year ese Name Address City & State Zip Special interests (optional): INDEX TO VOLUME 98 New scientific names are in bold face. Angelo, Ray and David E. Boufford. Atlas of the flora of New England: pteridophytes and gymnosperms. 1-7 ANNOUNCEMENTS: NEBC Graduate Student Research Award 110 1996 Jesse M. Greenman Award 111 Ronald L. Stuckey Endowment d-Ad12 Arctostaphylos uva-ursi 117-145 Asteraceae 85-93 Atlas of the flora of New England: pteridophytes and gymnosperms. 1-79 Bennett, James P. and Jennifer E. J. ourse. The vascular flora of Hopewell eur National Histor- ical Park, Ross County, Ohio. 146-167 Bolboschoenus 168-179 BOOK REVIEWS: An Almanac of Botanical Trivia. 217 Flora of Berkshire County, Mas- sachusetts. 459 Michigan Flora, Part III, Dicots Compositae). Native Tice: Shrubs and Woody Vines of Cape Cod and the Is- lands. 105—106 A Revision of Heterotheca sect. Phyllotheca (Nutt.) arms (Compositae: Astereae): The Prairie and Montane Goldenas- rs of North America. 218— 219, The Vascular and Non-Vascular Flora of Nantucket, Tuckernuck, and Muskeget ea 103-104 Boufford, David E. Brazil, Eremanthus species in 86—93 Brumback, William E. and Leslie J. Mehrhoff, in collaboration with Richard W. Enser, Susan C. Gaw- ler, Robert G. Popp, Paul Somers, and Daniel D. Sperduto, with as- sistance from William D. Country- man and C. Barre Hellquist. Flora — New England. The New England Plant Conservation sae (NEPCoP) list of plants n need of conservation. 233-361 eee 188 Bylaws of the New England Botani- cal Club, Inc. 226-231 ia Cod, coastal heathlands and andplain grasslands on 117-14 Gia 186-187, 369- 418 Classification of coastal heathlands and sandplain grasslands in Ma sachusetts. A, 117-145 Coastal heathlands 117-145 Coastal islands, seaweeds of 369— Colombia, Uncinia species in 80-84 Comments on the occasion of the secre of New England Bo- Club. —458 as an 117-145 Comparison of insular seaweed floras from Penobscot Bay, Maine, and other northwest Kuan islands. Compositae 85—93 Coniferophyta 25-28, 66-72, 151, 189, 273 Connecticut 29-72, 180-216, 251- Conservation, New England plants in need of 233-361 Sain Le William D. 233-361 C ennifer E. J. 146-167 amen 187 473 474 Cyperaceae 80-84, 99-102, 168— 179 adeap rates flexuosa = 145 istribution maps Dane. Peter W. a ea T. Leverett. Survey of old-growth forest in Massachusetts. 419-444 Dunwiddie, Peter W. and Bruce A. Sorrie. A flora of the vascular and non-vascular plants of Nantucket, Tuckernuck, lands. 94-98 (New eid Note) Dunwiddie, Peter W., Robert E. Za- remba, and Karen A. Harper. A classification of coastal heathlands and sandplain grasslands in Mas- ae 117-145 Endangered species 233-361 Enser, Richard W. 233-361 Equisetophyta 11-12, 40—42, 188, 285 Eremanthus 85-93; bicolor 86-88; har > reflexo- \ o the Paralychnopho- ra group of Eremanthus 86 First report of Uncinia macrolepis and U. tenuis (Cyperaceae) in Co- lombia and new Colombian sites for U. paludosa. 80-84 Flora 1-79, 9 , 146-167, 16, 369-418 Flora Obie ronda! New England. w England Plant Conser- vation nites (NEPCoP) list of plants in need of conservation. 233- Flora of Latimer Point and vicinity, New London County, Connecticut. The, 180-216 Flora of the vascular and non-vas- cular plants of Nantucket, Tuck- ernuck, and Muskeget Islands. A, 94—98 (New England Note) 180— Gawler, Susan C. 233-361 Rhodora [Vol. 98 ie ae ta baccata 117-145 wick, James 369-418 aa of cea seaweed flora of 369-418 Gymnosperms 1—79, 151, 189, 273 Hambrook, Julie 369-418 Harper, Karen A. 117-145 Hehre, Edward J. 369-418 Hellquist, C. Barre 233-361 Hemlock (Tsuga canadensis) 419— 444 Heuchera micrantha var. macrope- tala (Saxifragaceae), a new varie- ty. 365-3 Hill, Steven R. The flora of Latimer Hopewell Culture National Historical Park 146-167 Howard, Richard A. Comments on the occasion of the centenary of the New England Botanical Club. 445-458 Isolepis 168-179 Leverett, Robert T. 419-444 Liliopsida 151-163, 189-195, 251- 329 (families in alphabetical or- der) Lycopodiophyta 6—11, 30—39, 291- 292, 298-299, 326 Magnoliophyta 151-163, 189-212, 251-329 (families in alphabetical order) Magnoliopsida 151-163, 195-212, 251-329 (families in alphabetical order) Maine 29-72, 251-329, 369-418 Marine algae 186-187, 369-418 Martha’s Vineyard, coastal heath- lands and sandplain grasslands on 117-145 i 29-72, 94-98, 117— 145, —329, 369-418, 419— 444 a. Arthur C., Edward J. ehre, Julie Hambrook, and James 1996] Gerwick. A comparison of insular seaweed floras from Penobscot Bay, Maine, and other northwest Atlantic islands. 369-418 Mehrhoff, Leslie J. 233-361 Missouri, Scirpus sensu lato in Muskeget Island, flora of 94-98 Nantucket Island, flora of 94-98, 117-145 National park flora 146—167 NEBC Meeting News 107-109, 220-222, 463-467 NEPCoP (New England Plant Con- servation Program) 233-361 New Brunswick, Canada, seaweeds of 369-418 New England flora 1-79, 233-361 New Hampshire 29-72, 99-102, 251-329, 369-418 New Hampshire. Scleria reticularis (Cyperaceae) new to 2 Northern hardwood forests 419-444 Notes on the genus Scirpus sensu lato in Missouri. 168—179 Ohio, vascular flora of Hopewell Culture National Historical Park 46-167 Old-growth forest in Massachusetts. Survey of, 419-444 Paralychnophora group of Ereman- h nonieae: Asteraceae). The, 85—93 Penobscot Bay, seaweed flora of Phaeophycophyta 187, 369-418 Pinophyta 25-28, 66-72, 151, 189, 273 Polypodiophyta 12—25, 42-66, 151, 188-189, 251-329 (families in al- phabetical order) Popp, Robert G. 233-361 Pteridophytes 1-79, 151, 188-189, 251-329 (families in alphabetical order) Index to Volume 98 Quercus ilicifolia 117-145 Reviewers of manuscripts, 1996. 468 Rhode Island 29-72, 251-329 Rhodophycophyta 187-188, 369- 418 Robinson, Harold. The Paralych- nophora f Eremanthus (Vernonieae: Asteraceae). 85—93 Sandplain grasslands 117-145 Schizachyrium scoparium 117-145 ge ace 168-179; key to the ri members of sections Ac- ne eon oan 177 Scirpus 168-179; key to the genera eee ne Scirpus sensu lato n Missouri 170-171 Seer haa oan (Cyperaceae) new Hampshire. 99—102 (N eae Note) Shipes, B. G. and E. E Wells. Heu- chera micrantha var. macropetala (Saxifragaceae), a new variety Smith, S. Galen and George Yat- skievych. Notes on the genus Scir- pus sensu lato in Missouri. 168— L79 Somers, Paul. 233-361 Sorrie, Bruce A. 94—98 Sperduto, Daniel D. 233-361 Sperduto, Daniel D. Scleria reticu- laris (Cyperaceae) new to New Hampshire. 99-102 (New England Note) Survey of old-growth forest in Mas- sachusetts. 419-444 Trichophorum 168-179 Tuckernuck Island, flora of 94-98 Two-way indicator eee analysis (TWINSPAN) 117-14 Uncinia 80-84; macrolepis 80-81; paludosa 81-82; tenuis 82-83; key to the Uncinia species of Co- lombia 83 476 Rhodora Vascular flora of Hopewell Culture ational Historical Park, Ross 8 County, Ohio. The, 146—167 eon 29-72, 251-329 Vernonieae §5—93 Wells, E. F 365-368 Wheeler, Gerald A. First report of Uncinia macrolepis and U. tenuis (Cyperaceae) in Colombia and new Colombian sites for U. palu- dosa. 80-84 Yatskievych, George 168-179 Zaremba, Robert E. 117—145 [Vol. 98 THE NEW ENGLAND BOTANICAL CLUB Elected Officers and Council Members for 1996—1997: President: W. Donald Hudson, Jr., Chewonki Foundation, RR 2, Box 1200, Wiscasset, ME 04578 Vice-President (and Program Chair): David S. Conant, Depart- ment of Natural Sciences, Lyndon State College, Lyndon- ville, WT 05851 Corresponding Secretary: Nancy M. Eyster-Smith, Department of Natural Sciences, Bentley College, Waltham, MA 02154- 4705 Treasurer: Harold G. Brotzman, Box 9092, Department of Bi- ology, North Adams State College, North Adams, MA 01247-4100 Recording Secretary: Lisa A. Standley Curator of Vascular Plants: Raymond Angelo Assistant Curator of Vascular Plants: Pamela B. Weatherbee Curator of Nonvascular Plants: Anna M. Reid Librarian: Paul Somers Councillors: C. Barre Hellquist (Past President) Garrett E. Crow 1997 Matthew Hickler (Graduate Student Member) 1997 Edward Hehre, Jr. 1998 Michael J. Donoghue 1999 Appointed Councillors: David E. Boufford, Associate Curator Janet R. Sullivan, Editor-in-Chief, Rhodora