RHODORA, Vol. 98, No. 893, 1-79, 1996 pp. NEW ATLAS OF THE FLORA OF ENGLAND AND GYMNOS PTERIDOPHYTES PERMS Ray Angelo New England Botanical Club, 22 Divinity Avenue, Cambridge, Massachusetts 02138-2020 David Boufford E. Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138-2020 ABSTRACT. Dot maps provided are to depict the distribution the county at level of the pteridophytes (Lycopodiophyta, Equisetophyta, Polypodiophyta) and rm gland states of the northeastern United States. The 171 taxa (species, subspecies, and mapped varieties hybrids, but not forms) arc at the county level based on New specimens major in the herbaria of Maine, Hampshire, Vermont, Massachu- Rhode and setts, Island Connecticut, with primary emphasis on the holdings of New the England Botanical Club Herbarium synonymy (nebc). Brief account to names for used in recent manuals and floras for the area, habitat and chromo- some common infonnation and names also are provided. Key New Words: Flora, England, gymnospcrms atlas, distribution, pteridophytes, This article is the first in a series that will present the New distributions of the vascular flora of England in the form of New maps dot distribution the county England com- at level. prises the six northeastern states of the United States: Maine, New Hampshire, Vermont, Rhode Massachusetts, and Island Connecticut. These range from 4 °N states in latitude to 1 67°W 47°28'N and longitude from 73^45'W. The in to elevation m of the region extends from sea level to 1,898 (6,228 the ft.) at New summit of Mt. Washington, Hampshire, moun- the highest tain in eastern North America outside of North Carolina. The km^ total area of the region 172,515 (66,608 mi.^) including is km^ bodies of water; the land area 163,157 total (62,995 mi.^). is The of which effects continental covered glaciers, the entire region several times during the Pleistocene, and the of last which did not retreat until about 15,000 years ago, are still clearly evident throughout The by the area. state state statistics on and shown size elevation are Table (Merriam-Webster, in 1 Inc. 1984). 1 Rhodora 2 98 [Vol. New Tabic Total area, land area, and range in elevation for the six 1. Eniiland states. Area and Area hlcvation Ran^e Total T m km^ km- 0- Maine 86,028 80,117 ,606 1 m New Hampshire 24,097 km^ 23,395 km^ 0-1,898 m km- knr Vermont 24,887 24,020 29 1,339 m Massachusetts 21,386 knr 20,287 knr 0-1,064 m Rhode km^ km^ 0-248 Island 3,144 2,725 m km^ km^ 725 Connecticut 12,973 12,613 The more Rhode and southerly states of Massachusetts, Island, Connecticut densely populated densely populated are (the least of these three states, Connecticut, had an average of 262 persons per km^ 1990; Hofftiian 1992). The northern states of Maine, in New Hampshire, and Vermont mostly most densely are rural (the New populated of these three Hampshire, had an average states, km" Hoffman of 48 persons per in 1990; 1992). This work encompasses vascular plants (pteridophytes and all and spermatophytes) the rank of species, subspecies, variety at New Hy- growing England outside of cultivation the in states. below brids also will be included, but forms and other ranks the level of variety will not. The dots are based primarily on voucher New specimens in the herbaria of England representing reproduc- when phmts ing populations, or persisting after cultivation is it uncertain that they are actually naturalized. This first installment includes the pteridophytes (Lycopodiophyta, Equisetophyta, Poly- gymnosperms podiophyta) and (Coniferophyta). Future accounts will treat the distribution of angiosperms. We intend to gather this series of articles, together with addi- background volume upon comple- material, into a separate tional tion of the maps, and to make available on the World Wide all it Web hope mean- as various parts are finished. our that, in the It is work time, these articles will stimulate additional field to supple- New ment maps. The England the distributions portrayed in the Botanical Club herbarium, which has proven to be the most im- portant resource for this project, is especially eager to receive We specimens documenting range extensions. also would like to be informed of such specimens other herbaria. Similarly, be- in New cause the atlas of the England flora will be continuously new becomes we updated as information available, are eager to receive notification of published corrections of cytological infor- — New Angelo and Boufford Atlas of England Flora 3 1996J mation and new, documented chromosome counts for taxa in the New England flora. AND METHODS MATERIALS New This grew out of an inventory of England Bo- atlas the tanical Club vascular plant herbarium. The purpose of the inven- conducted county was tory, at the level, to serve as a filter for number unmounted the processing of a large of specimens being considered for accession. Consequently, the core of the database for this atlas consists of specimens in the herbarium of vascular NEBC New England Club The plants of the Botanical (nebc). New herbarium, which geographical scope En- limits to the its more gland presently comprises than 251,000 specimens states, New and the largest collection of plants for the England re- is gion. In the mid 1980s the herbarium inventory was expanded into NEBC an from herbarium atlas project, since data the essentially New depict the distributions of the vascular pkmts of England. Additional data from specimens other major herbaria with large in New holdings of England plants have allowed us to portray the more distributions completely. New The data represented currently by specimens England in when herbaria best depict vascular plant distributions displayed at the county level. Attempting to display such data by the exact geographical coordinates of each collection locality, or even at By township such the level, distorts the actual distributions. de- tailed mapping, patterns tend to reflect concentrated collecting along major highways and clustering near popular resort areas or was from some botanical hot spots. This evident of the detailed maps distribution produced about 15 years ago by the plant dis- New committee (now England tribution defunct) of the Botanical Club. The distortions portrayed by such detailed mapping are re- solved by using data only at the county level. New The counties of England range in size from Bristol County, Rhode Island (65 km", or 25 mi.^), to Aroostook County, Maine — The (17,666 km^, or 6821 mi.^ land only). great majority of km counties, however, are between 1300 (500 mi.") and 3900 km To (1500 mi.*"). reduce the size disparity at the large end, and more Maine to depict distributions evenly, the largest counties are subdivided. Penobscot, Piscataquis, Somerset, and Oxford Coun- ties are divided arbitrarily into roughly equal-sized, north and Rhodora 4 98 Vol. south sectors along township boundaries. Aroostook County, the New Coos largest county, divided into three sectors. County, is The Hampshire, divided into north and south sectors. division is Coos County White Mountains, which an of isolates the harl^or New alpine flora associated mostly with England's highest peak. Mount Washington, Figure shows the location of states and 1 New counties of England and the subdivisions of the largest coun- ties. Data were collected primarily from herbarium specimens of New major herbaria in England. The herbaria used to compile data New for this work were those of the Enuland Botanical Club (ncbc). Harvard University ames, University of Maine (a, c,h). New University of Massachusetts (mass). University of (MAiNtz), Hampshire Rhode (nha), University of Island University (kiri), Brown of Vermont (vt), and University (bru). Records from the herbaria the University of Connecticut (conn), Connecticut Bo- at Society (ncbs), and Yale University (yu) were accessed tanical from data compiled by Leslie Mehrhoff. Lastly, the notebooks of Harry Ahles the University of Massachusetts were consulted. at Ahles visited these same herbaria, plus one or two others, in the 1960s and 1970s to gather locality information at the county level. We some where obtained records from systematic treatments the information on voucher specimens was specific enough to pro- name vide county data and the of the herbarium where the speci- mens were deposited. no was made In recording the collection data attempt to verify the accuracy of the identification of every specimen, but obvious Even misidentifications were cumotated with corrections. though the basis for a particular dot might be a misidentified specimen, New common for plants within England, our view that the taxon is most Specimens documenting likely occurs within the county. dis- junct or marginal occurrences (or otherwise rare, endangered or threatened taxa) usually have received greater scrutiny, loecause and of a heightened interest in rare plants nationwide, are less be likely to 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 which voucher specimen The the deposited. in letter is code herbarium where voucher specimen was indicates the a first No seen. letter codes were added for subsequent voucher speci- mens other herbaria, because our primary objective has been at to plot the distribution of the taxon, not to inventory the holdings — New Angelo cUid Boufford Atlas of England Flora 5 1996J of any particular herbarium. While such information might be of some we value for purposes, felt that our time at other herbaria was New devoted best to searching for data not found in the En- gland Botanical Club herbarium, or each subsequently searched in herbarium, gaps to in in distribution. fill The data in the database were converted to dot distribution A maps method using a devised by Angelo more (1994). detailed and mapping technical description of methodology the will appear an appendix volume. as in the final Interested parties may obtain of appendix upon a draft the request to the first author. TAXONOMY AND FORMAT The taxonomy and nomenchiturc adopted work for essen- this !/ With major for the divisions within the pteridophytes (Lycopodiophyta, Equisetophyta, Polypodiophyta), which follows Flora cind the of North America (Flora of North America Committee Editorial 1993) sequence, the families are ordered alphabetically. The gen- era are alphabetical within families, as are species within genera. Named unnamed and hybrid taxa are placed alphabetically the at Unnamed end of the genus. hybrids combine the names of the progenitors alphabetically by epithet. deemed New Species be to introduced, that not native is, in England the time of European contact with North America, at are indicated by the use of all upper case type for the scientific name. No single source of information was used determining in intro- duced For most species. taxa there is little dispute as to nativity. Where we differences of opinion exist in the literature, have used own our judgment based on evidence the available. A common name supphed when name appears be is to a in it Names New general use. used England have been given in pref- er erence. chromosome numbers Cited are taken from indices prepared by Cave (1958a, 1958b, 1959a, 1959b, 1960, 1961, 1962, 1963, 1964, 1965), Goldblatt (1981, 1984, 1985, Goldblatt and 1988), Johnson Moore (1990, 1991, 1994), (1973, 1974, Ornduff 1977), (1967, 1968, 1969), and reports by Takamiya and Kurita (1983), Taylor Tryon Tryon and Tryon and Wag- (1970), (1978), (1982), ner (1971). Very few of the counts are based on material from Rhodora 6 98 [Vol. New made from England, but instead rellect counts throughout the range of the taxon. The habitat data are distillations from a variety of sources aug- own An was made mented by our observations. attempt to field taxon indicate habitat information as applies to a particular in it New England rather than the entire range of the taxon. to Synonymy names used provided primarily with respect to in is New standard manuals covering England published from 1950 the onward, including Fernald (1950), Gleason (1952), Gleason and Seymour Cronquist (1963, 1991), and (1969a, 1982). A many selection of references, including not cited above, is provided. This consists of the standard manuals published list from 1950 regional and county floras and checklists pro- to date, we duced alter 1950, and what believe to be the key articles and New gymnosperms books on pteridophytes and pertinent the to We we may England. would appreciate being notified of papers have overlooked. LYCOPODIOPHYTA ISOETACEAE — = Isoctes acadiensis Kott Acadian Quill wort (Figure 2n 44; 2). = 44 and macro- the report of (Kott Britton 1980) for 2/7 /. spora Durieu forma hicroglyphica (A. A. Eaton) N. E. Pfeiffer was Shallow wa- laler attributed (Kott 1981) to acadiensis. /. moving ponds and slow of streams. ter — = echinospora Durieu Braun's Quillwort (Figure 2n Isoctes 2). 22, 33. Shallow water and shores of slightly acidic lakes, ponds, and streams. eclunospora var. hraiinii (Durieu) En- [/. Engelmann; gelmann; echinospora muricata (Durieu) var. /. muricata Durieu] /. — engelmannii A. Braun Engelmann's Quillwort (Figure Isoetes = ^ 2n Shallow water of ponds, streams, n 22. lakes, 2). 11; muddy or ditches. in — ^ Isoetes lacustris Linnaeus Lake Quillwort (Figure 2). 2n 110. Shores and shallow water of slightly acidic ponds, lakes, and streams. macrospora Durieu; liieroglyphica A. A. Eaton] /. [/. — New Angelo and Boufford England Atlas of Flora 7 1996J ~ M. Deep Isoetes prototypus D. Britton (Figure 2n 22. wa- 3). of ter cold, acidic lakes. — Engelmann Isoetes riparia ex A. Braun Riverbank Quillwort = Muddy (Figure 2n 44. or sandy shores (including 3). tidal and shores estuaries), usually of rivers. riparia var. cana- [/. Engelmann; Engelmann densis saccharata amesii var. /. A. A. Eaton] — Isoetes tuckermanii A. Braun ex Engelmann Tuckerman's Quill- = Pond wort (Figure 2n 44. and lake margins, shores 3). (fre- submersed quently tidal), usually in quiet water. Isoetes Fiybrids X X Dodge [^ Isoetes eatonii (Figure echinospora Durieu 3). /. X engelmannii A. Braun; gravesii A. A. Eaton; eatonii /. /. /, Dodge gravesii (A. A. Eaton) Clute] var. X was Isoetes harveyi A. A. Eaton. This hybrid recognized only recently by Britton (1991). Herbarium specimens annotated as hybrid are from southern Penobscot Co., Maine this (us); Somerset Co., Maine (maine); and Mt. Desert Island, Maine (mo). Since the holdings of nebc have not been examined criti- mapped. [= cally for this hybrid, not laciistris Lin- is it /. X naeus tuckermanii A. Braun] /. LYCOPODIACEAE — Diphasiastrum complanatum (Linnaeus) Holub Northern Run- = = ning Pine (Figure n 22-24, 23; 2n 40, 44, 46, 48. 4). Dry, open coniferous or mixed [Lycopodium com- forests. planatum Linnaeus] — Diphasiastrum digitatum (Dillenius ex A. Braun) Holub South- ^ = ern Running Pine (Figure n 23; 2n 46. Usually dry 4). woods, open [Lycopodium thickets, or fields, in acid soils. complanatum Linnaeus flahelliforme Fernald] var. — Diphasiastrum Holub Clubmoss sitchense (Ruprecht) Sitka — = (Figure n 23; 2n 46. Alpine meadows, open rocky 4). Rhodora 8 [Vol. 98 woods [Lycopodium Ru- barrens, conifer (rarely). sitchense precht] — Holub Ground Cedar Diphasiastrum tristachyum (Pursh) (Fig- — = ure n 23; 2n 46. Acid soils in open conifer or oak 4). woods, [Lycopodium tristachyum Pursh] thickets. Diphasiastrum Hybrids X Holub Diphasiastrum com- Diphasiastrum (Linnaeus) alpiniim planatum Holub (Linnaeus) (Figure 5). X complanatum Holub Diphasiastrum Diphasiastrum (Linnaeus) di^itatum (Dillcnius ex A. Braun) Holub (Figure 5). X Diphasiastrum digitatum (Dillenius ex A. Braun) Holub Dipha- X siastrum (Willdenow) Holub (Figure salfinifolium 5). X Holub [^ Diphasiastrum hahereri (House) (Figure D. 5). dif^i- X Holub tatum (Dillenius ex A. Braun) D. tristachyum (Pursh) X Lycopodium House] Holub; hahereri X Diphasiastrum (Rouy) Holub (Figure [^ D. alpi- issleri 6). X num Holub (Linnaeus) D. tristachyum (Pursh) Flolub; Ly- X copodium (Rouy) Lawalrcc] issleri — X Diphasiastrum sahiuifolium (Willdenow) Holub Savin- = Woods Clubmoss leaved (Figure 2n 46. (often subal- 6). [^ pine spruce), thickets, clearings. D. sitchense (Ruprecht) X Holub D. tristachyum (Pursh) Holub; Lycopodium sahiui- Willdenow] folium X Diphasiastrum (Rouy) Holub (Figure [= D. zeillcri 6). X complanatum Holub (Linnaeus) D. tristachyum (Pursh) Holub] & Huperzia appalachiana Mickel Beitel (Figure Cliffs, talus 6). damp, [Specimens slopes, acidic rocks in alpine zones. New mapped from England are the herbaria of the Botanical Club and Harvard University.] New Angelo and Boufford^Atlas of England Flora 9 1996] — Clubmoss Huperzia (Michaux) Trevisan Shining (Fig- lucidida = — woods and ure n 67; 2n 134. Cool, moist in rich acid 7). [Lycopodium lucidulum Michaux] soils. Marti = Damp Mountain 2n 268. 13611; woods, mountainous rocks, barrens, cold in regions. [H. sc- densa Lycopodium selago Linnaeus; Ly- lago Trevisan; var. copodium Linnaeus appressum Desvaux] selago var. Huperzia Hybrids & X Huperzia appalachiana Mickel Huperzia lucidnla Beitel (Michaux) Trevisan (Figure 7). & X Huperzia appalachiana Mickel Huperzia selago (Lin- Beitel & naeus) Bernhardi ex Schrank Martius (Figure 7). & X Gandhi Huperzia (Abbe) Kartesz (Figure buttersii 8). X [= H. lucidula (Michaux) Trevisan selago (Linnaeus) //. & Bernhardi ex Schrank Martius] — Club- Lycopodiella alopecuroides (Linnaeus) Cranfill Foxtail = moss (Figure 2n 156. Bogs, marshes, ditches, sandy or 8). {Lycopodium alope- peaty shores, wet barrens, in acid soils. Linnaeus] curoides — Bog (Chapman) Appressed Lycopodiella appressa Cranfill = Damp, Clubmoss (Figure L56. acid or peaty soils in 2/7 8). & [Lycopodium appressum (Chapman) Lloyd Un- open. the immdatum derwood; Lycopodium Linnaeus bigelovii var. Tuckerman; Lycopodium immdatum Linnaeus rohustum var. Eaton] R. J. — Bog Clubmoss immdata (Figure Lycopodiella (Linnaeus) Holuh) = ^ Damp, n 78; 2n 156. acid, sandy or peaty soils in 8). immdatum [Lycopodium Linnaeus] open. the & W. Wagner Lycopodiella margueritae G. Bruce, H. Beitel J. = 2n (Figure 312. 9). Rhodora 10 98 [Vol. — Lycopodiuni annotinum Linnaeus Clubmoss (Figure Stiff 9). - - n 34, 68TT; 2// ca. 50, 66, 68. Woods, cool shaded thick- ets, exposed rocky or peaty habitats higher elevations. at [L. ahyestre :7 C. Harlnian; annotinum van pungens L. (Bachelot de Py- la Desvaux] laie) — Lycopodiuni clavatuni Linnaeus Staghorn Clubmoss (Figure 9). = = n 22, 34, 34II, 34111, 34TV, 68 (69, 70); 2/7 34, 44, 68, Open, woods 102, 136. usually and dry, thickets, clearings, exposed rocky shuations, acid clavatum sub- in soils. [L. var. rcmotuni Victorin — Lycopodiuni dcndroideuni Michaux Clubmoss Tree (Figure 9). = 2n 68. Dry, open woods and clearings, obscuruni Lin- \L. naeus dcndroideuni (Michaux) var. D. C. Eaton] — & Lycopodiuni hickeyi W. H. Wagner, Beitel R. C. Moran Mick- = Clubmoss ey's (Figure Mainly deciduous woods 10). 2// 68. and thickets, usually acid obscuruni Linnaeus in soils. [L. var. isophyllum Hickey] Lycopodiuni lagopus (Laestadius ex C. Hartnian) G. Zinserling = ex Kuzeneva-Prochorova (Figure In Grassy 68. 10). fields and openings second-growth woods. clavatum Linnaeus in [L. & var. megastacJiyon Fernald clavatum Linnaeus Bissell; L. & Hooker nionostachyon var. Greville] Distribution data are New primarily from the herbaria of the England Botanical Club and Harvard University. — Lycopodiuni obscuruni Linnaeus Clubmoss Tree (Figure 10). = = 34, 34II; 68. Rich woods, clearings, edges of boggy 2fi // forests, usually acid in soils. Lycopodiuni Hybrids X Lycopodiuni Michaux dcndroideuni Lycopodiuni ohscurum Linnaeus (Figure 10). Pseudolycopodiella caroliniana (Linnaeus) HoIub^Carolina - = Clubmoss (Figure n 11). 35, 68, 70, 15; 2/? 70, 78, 140. 1