Full text of "Rhodora"
RHODORA
Journal of the
New England Botanical Club
CONTENTS
Atlas of the Flora of New England: Monocots except Poaceae and Cypera-
c Ray Angelo and David E. Boufford ]
NEBC MEETING NEWS 120
Information for Contributors 126
Statement of Ownership 128
NEBC Officers and Council Members inside back cover
Vol. 102 Winter, 2000 No. 909
Issued: February 23, 2000
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
ANTOINETTE P. HARTGERINK, Managing Editor
Department of Plant Biology, University of New Hampshire,
Durham, NH 03824
Associate Editors
HAROLD G. BROTZMAN STEVEN R. HILL
DAVID S. CONANT THOMAS D. LEE
GARRETT E. CROW THOMAS MIONE
Kk. N. GANDHI—Latin diagnoses and nomenclature
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RHODORA, Vol. 102, No. 909, pp. 1-119, 2000
ATLAS OF THE FLORA OF NEW ENGLAND: MONOCOTS
EXCEPT POACEAE AND CYPERACEAE
RAY ANGELO
New England Botanical Club, 22 Divinity Avenue,
ambridge, MA 02138-2020
e-mail: rangelo@oeb.harvard.edu
Davip E. BOUFFORD
Harvard University Herbaria, 22 Divinity Avenue,
Cambridge, MA 02138-2020
e-mail: boufford @oeb.harvard.edu
ABSTRACT. Dot maps are provided to depict the distribution at the county
level of the families of Monocotyledons except Poaceae and Cyperaceae
growing outside of cultivation in the six New England states of the north-
eastern United States. The 325 of the 329 taxa (species, subspecies, varieties,
and hybrids, but not forms) treated are mapped based on specimens in the
major herbaria of Maine, New Hampshire, Vermont, Massachusetts, Rhode
Island, and Connecticut, with primary emphasis on the holdings of the New
England Botanical Club Herbarium (NEBC). Brief synonymy to account for
names used in recent manuals and floras for the area, habitat and chromosome
information, and common names are also provided.
Key Words: sie New England, atlas, distribution, Juncaceae, Liliaceae,
idaceae, Potamogetonaceae, aquatic plants, rushes, lilies,
nee
This article is the third in a series that will present the distri-
butions of the vascular flora of New England in the form of dot
distribution maps at the county level (Figure |). The atlas is post-
ed on the internet at http://www.herbaria.harvard.edu/~rangelo/
NeatlasO/WebIntro.htm] where we will attempt to keep it updated.
This project encompasses all vascular plants (pteridophytes and
spermatophytes) at the rank of species, subspecies, and variety
growing outside of cultivation in the six New England states.
Hybrids are also included, but forms and other ranks below the
level of variety are not. The dots are based primarily on voucher
specimens in the herbaria of New England representing repro-
ducing populations, or plants persisting long after cultivation
when it is uncertain that they are actually naturalized. This third
installment includes the families of the Monocotyledoneae except
the Poaceae and Cyperaceae. The number of taxa treated is 329,
I
N
Rhodora [Vol. 102
of which 325 are mapped. Of these 329 taxa, 56 (mostly in Lil-
laceae) are not native to the region. Future accounts will treat the
distribution of the rest of the 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. It is our hope, in the meantime, that these
articles will stimulate additional field work to supplement the
distributions portrayed in the maps. The New England Botanical
Club herbarium, which has proven to be the most important re-
source 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, because the atlas
of the New England flora will be continuously updated as new
information becomes available, we are eager to receive notifica-
tion of published corrections of cytological information and new,
documented chromosome counts for taxa in the New England
flora.
MATERIALS AND METHODS
Materials and methods are as outlined in Angelo and Boufford
(1996) and are not repeated here.
TAXONOMY AND FORMAT
The taxonomy and nomenclature adopted for this work essen-
tially follow that of the Flora of North America project in pro-
gress, except that families, genera, and species are arranged al-
phabetically. Named and unnamed hybrid taxa are placed alpha-
betically at the end of the genus. Unnamed hybrids combine the
names of the progenitors alphabetically by epithet. Taxa that are
not native to New England are indicated by uppercase text. Un-
published names are not used, even if publication is pending.
Cited chromosome numbers are taken from indices prepared
by Cave (1958a, b, 1959a, b, 1960, 1961, 1962, 1963, 1964,
1965), Goldblatt (1981, 1984, 1985, 1988), Goldblatt and John-
son (1990, 1991, 1994, 1996), Léve and Live (1975), Moore
(1973, 1974, 1977), and Ornduff (1967, 1968, 1969). Very few
of the counts are based on material from New England, but in-
stead reflect counts made from throughout the range of the taxon.
Synonymy is provided primarily with respect to names ac-
2000] Angelo and Boufford—Atlas of New England Flora 3
cepted in standard manuals covering New England published
from 1950 onward, including Fernald (1950), Gleason (1952),
Gleason and Cronquist (1991), and Seymour (1982). Synonyms
have not been provided where the distribution for the synony-
mized name does not include New England.
The following list will aid readers in finding familiar names
that have been transferred to other taxa:
ACORACEAE
LILIACEAE
Coeloglossum
ARACEAE (Acorus) =>
AMARYLLIDACEAE =>
Habenaria (in part) =
Habenaria (in part) = Platanthera
LILIACEAE (Smilax) = SMILACACEAE
LILIACEAE (Yucca) => AGAVACEAE
Lophotocarpus = Sagittaria
Orchis (in part) = Amerorchis
Orchis (in part) = Galearis
Potamogeton (in part) => Stuckenia
Smilacina = Maianthemum
The following species are reported from our area in manuals,
but no specimens were seen, or the substantiating specimens were
misidentified:
Melanthium hybridum Walter (no specimen seen)
Smilax bona-nox Linnaeus [misidentified: = S. rotundifolia
Linnaeus (Sorrie 1987)]
ANGIOSPERMAE (MAGNOLIOPSIDA )—
ANGIOSPERMS
MONOCOTYLEDONEAE (LILITIDAE)
ACORACEAE
Acorus americanus (Rafinesque) Rafinesque—Sweet Flag (Figure
2n = 24. Marshes, shores, wet meadows. [A. calamus
misapplied; The mapped distribution may include specimens
of the introduced sterile triploid, A. CALAMUS, which is not
generally distinguished in herbaria. ]
ACORUS CALAMUS Linnaeus—Sweet Flag (Figure 2). 2n = 36.
Marshes, shores, wet meadows. From Europe. [This sterile
4 Rhodora [Vol. 102
triploid is not separated from the native species in most her-
baria. |
AGAVACEAE
YUCCA FILAMENTOSA Linnaeus— Yucca (Figure 2). 2” = 60.
Roadsides. From farther south.
ALISMATACEAE
Alisma gramineum Lejeune—(Figure 2). 27 = 14, 16. Muddy
shores and shallow water of basic lakes and streams.
Alisma subcordatum Rafinesque—Southern Water-plantain (Fig-
ure 3). 2n = 14. Muddy or sandy shores, marshes, ditches,
shallow water. [A. plantago-aquatica Linnaeus var. parviflo-
rum (Pursh) Torrey]
Alisma triviale Pursh—Northern Water-plantain (Figure 3). 2” =
14, 28. Muddy shores, marshes, ditches, shallow water. [A.
plantago-aquatica Linnaeus var. americana J. A. Schultes &
Schultes]
Echinodorus tenellus (Martius) Buchenau—(Figure 3). 2n = ?.
Sandy shores of streams and lakes. [E. parvulus Engelmann]
Sagittaria cuneata Sheldon—Wapato (Figure 3). 2n = 22. Al-
kaline waters of muddy shores and shallow water of rivers.
Sagittaria engelmanniana J. G. Smith—(Figure 4). 2n = 22.
Acidic waters of shores, marshes, and bogs.
Sagittaria filiformis J. G. Smith—(Figure 4). 27 = ?. Deep water
of streams and in rapids. [S. subulata (Linnaeus) Buchenau
var. gracillima (S. Watson) J. G. Smith]
Sagittaria graminea Michaux subsp. graminea—(Figure 4). 2n =
22. Muddy or sandy shores, shallow water. [S. eatonii J. G.
Smith]
Sagittaria latifolia Willdenow—(Figure 4). 2n = 22. Muddy
shores, ditches, bogs. [$. /atifolia Willdenow var. obtusa
(Muhlenberg) Wiegand]
Sagittaria montevidensis Chamisso & Schlechtendahl subsp.
2000] Angelo and Boufford—Atlas of New England Flora 5
spongiosa (Engelmann) Bogin—(Figure 5). 2n = 22. Tidal
mud flats of estuaries and salt marshes. (8. spatulata (J. G.
Smith) Buchenau; Lophotocarpus spongiosus (Engelmann) J.
G. Smith]
Sagittaria rigida Pursh—(Figure 5). 2n = 22. Alkaline or brack-
ish shores and shallow water.
Sagittaria subulata (Linnaeus) Buchenau—(Figure 5). 21 = 22.
Tidal mud.
Sagittaria teres S. Watson—(Figure 5). 2n = 22. Acid sandy
pond shores.
ARACEAE
Arisaema dracontium (Linnaeus) Schott—Green Dragon (Figure
6). 2n = 28, 56. Rich or alluvial soil.
Arisaema triphyllum (Linnaeus) Schott—Jack-in-the-pulpit (Fig-
ure 6). 2n = 28, 36, 56. Rich damp-to-wet woods, boggy
places. [A. triphyllum var. pusillum Peck; A. triphyllum var.
stewardsonii (Britton) Stevens ex Wiegand & Eames; A.
atrorubens (Aiton) Blume; A. pusillum (Peck) Nash; A. ste-
wardsonii Britton]
—Arisaema hybrids—
Arisaema dracontium (Linnaeus) Schott * Arisaema triphyllum
(Linnaeus) Schott—(Figure 6).
Calla palustris Linnaeus—Wild Calla (Figure 6). 21 = 36, 60,
72. Bogs, marshes, swampy woods, pond margins, shallow
water.
Orontium aquaticum Linnaeus—Golden Club (Figure 7). 2” =
26. Shallow water of ponds, sandy, muddy, or sphagnous
shores.
Peltandra virginica (Linnaeus) Schott—Arrow Arum (Figure 7).
= 112. Shallow water or mud at margins of ponds and
streams, swamps, bogs, damp meadows.
Symplocarpus foetidus (Linnaeus) Nuttall—Skunk Cabbage (Fig-
6 Rhodora [Vol. 102
ure 7). 2n = 26, 60. Swamps, damp thickets and woods, wet
meadows, shores.
BUTOMACEAE
BUTOMUS UMBELLATUS Linnaeus—Flowering Rush (Figure
7). 2n = 16, 20, 24, 26, 30, 39. Muddy shores and marshes.
From Eurasia.
COMMELINACEAE
COMMELINA COMMUNIS Linnaeus—Asiatic Dayflower (Fig-
ure 8). 2n = 16, 22, 28, 32, 36-90. Waste places, roadsides,
disturbed moist soil in shade. From eastern Asia. [C. COM-
MUNIS var. LUDENS (Miquel) C. B. Clarke]
COMMELINA DIFFUSA Burman f.—Creeping Dayflower (Fig-
ure 8). 27 = 18, 28-60. Waste places. From the Old World.
TRADESCANTIA BRACTEATA Small—Sticky Spiderwort (Fig-
ure 8). 2n = 12, 18, 24. Roadsides. From farther west.
Tradescantia ohiensis Rafinesque—Smooth Spiderwort (Figure
8). 2n = 12, 24. Roadsides, waste places, thickets.
Tradescantia virginiana Linnaeus—Widow’s Tears (Figure 9). 2
= 12, 18, 24, 24 + 6B. Roadsides, waste places, thickets.
—Tradescantia hybrids—
Tradescantia ohiensis Rafinesque X TRADESCANTIA SUBAS-
‘A Ker Gawler—(Figure 9).
Tradescantia ohiensis Rafinesque < Tradescantia virginiana Lin-
naeus—(Figure 9)
DIOSCOREACEAE
DIOSCOREA BATATAS Decaisne—Cinnamon-vine (Figure 9).
2n = ca. 140-144. Thickets, waste places. From China.
Dioscorea villosa Linnaeus—Wild Yam (Figure 10). 2” = 60.
Damp thickets, wet woods, roadsides.
2000] Angelo and Boufford—Atlas of New Engiand Flora 7
ERIOCAULACEAE
Eriocaulon aquaticum (Hill) Druce—White-buttons (Figure 10).
2n = 32, 64. Acid shores, shallow water, and bogs. [E. sep-
tangulare Withering]
Eriocaulon parkeri B. L. Robinson—(Figure 10). 2n = 48. Tidal
mud and estuaries.
HAEMODORACEAE
Lachnanthes caroliniana (Lamarck) Dandy—Redroot (Figure
10). 2n = 24, 48. Sandy or sphagnous pond shores. [L. tinc-
toria (J. E Gmelin) Elliott]
HYDROCHARITACEAE
EGERIA DENSA Planchon—(Figure 11). 2n = 46, 48. Ponds.
From Brazil and Argentina. [ELODEA DENSA (Planchon)
Caspary; ANACHARIS DENSA (Planchon) Marie-Victorin]
Elodea canadensis Michaux—(Figure 11). 2” = 24, 48. Ponds,
lakes, and quiet streams, mostly basic. [Anacharis canadensis
(Michaux) Richardson]
Elodea nuttallii (Planchon) St. John—(Figure 11). 2n = 48.
Ponds, lakes, and streams, acidic to moderately basic. [An-
acharis nuttallii Planchon]
HYDRILLA VERTICILLATA (Linnaeus f.) Royle—(Figure 11).
2n = 16, 24, 32. Ponds, lakes, and streams. From the Old
World.
Vallisneria americana Michaux—Water-celery (Figure 12). 2n =
20. Ponds and quiet streams.
IRIDACEAE
BELAMCANDA eens (Linnaeus) de Candolle—(Figure
28, 30, 32, 128. Fields, roadsides, thickets,
open aa. elias eastern Asia.
CROCUS VERNUS (Linnaeus) J. Hill subsp. VERNUS—Dutch
8 Rhodora [Vol. 102
Crocus (Figure 12). 21 = 8, 10, 12, 16, 16 + 2B, 18, 19,
20, 22, 23, 32. Grasslands. From Europe.
IRIS CRISTATA Aiton—Dwarf Crested Iris (Figure 12). 2n = 24,
32. Rich woods, in acid soils. From farther south and west.
IRIS GERMANICA Linnaeus—Fleur-de-lis (Figure 13). 2” = 28,
36—48. Roadsides, waste places. From Europe.
IRIS KAEMPFERI Siebold ex Lemaire
13). 2n = 24. Habitat? From Japan.
Japanese Iris (Figure
Iris prismatica Pursh—Slender Blue Flag (Figure 13). 2n = 42.
Marshes, meadows, swamps, sands, shores, seacoasts.
IRIS PSEUDACORUS Linnaeus—Yellow Iris (Figure 13). 2 =
24—34. Swamps, wet meadows, marshes, brooksides, waste
places. From Europe.
IRIS PUMILA Linnaeus subsp. PUMILA—(Figure 14). 2n = 20,
24, 30, 31, 32. Dry rocky knolls. From Eurasia.
[ris setosa Pallas—Beachhead Iris (Figure 14). 2n = 34-38.
Rocky slopes, upper borders of beaches, moist fields, always
near salt water. [/ris hookeri Penny ex G. Don]
IRIS SIBIRICA Linnaeus—Siberian Iris (Figure 14). 2” = 28. Wet
meadows, waste lots. From Eurasia.
IRIS TECTORUM Maximowicz—Wall Iris (Figure 14). 2n = 28,
36. Habitat? From China.
Iris versicolor Linnaeus—Blue Flag (Figure 15). 27 = 108.
Swamps, marshes, meadows, shores, ditches.
—Iris hybrids—
[ris prismatica Pursh X Tris versicolor Linnaeus—(Figure 15).
Sisyrinchium albidum Rafinesque—(Figure 15). 2n = 32, 64.
Dry, sandy, open soil and thin woodlands.
Sisyrinchium angustifolium Miller—Stout Blue-eyed Grass (Fig-
ure 15). 2n = 48. Meadows, fields, low woods, thickets,
damp shores. [S. bermudiana misapplied; S. graminoides
Bicknell]
2000] Angelo and Boufford—Atlas of New England Flora 9
Sisyrinchium atlanticum Bicknell—Eastern Blue-eyed Grass (Fig-
ure 16). 2n = 16, 32, 96. Meadows, marshes, low woods.
Sisyrinchium fuscatum Bicknell—(Figure 16). 2” = 32. Grass-
lands, mostly sandy soils. [S. arenicola Bicknell]
Sisyrinchium montanum Greene var. crebrum Fernald—Common
Blue-eyed Grass (Figure 16). 2n = 32, 96. Fields, meadows,
open woods.
Sisyrinchium mucronatum Michaux—Slender Blue-eyed Grass
(Figure 16). 2n = 30, 32. Meadows, fields, sandy places,
open woods.
JUNCACEAE
Juncus acuminatus Michaux—(Figure 17). 2n = 40. Tidal mud
flats, salt marsh borders, ditches, shores, meadows.
Juncus alpinoarticulatus Chaix—Alpine Rush (Figure 17). 2n =
40. Shores, meadows, usually basic. [J. alpinus Villars; J.
alpinus Villars var. fuscescens Fernald; J. alpinus Villars var.
rariflorus Hartmann]
Juncus anthelatus (Wiegand) R. E. Brooks—(Figure 17). 2” =
80. Moist sandy or clay soils. [J. tenuis Willdenow var. an-
thelatus Wiegand]
Juncus arcticus Willdenow var. balticus (Willdenow) Trautvet-
ter—Wire Rush (Figure 17). 2” = 40, 80. Sandy shores,
sphagnous meadows. [J/. balticus Willdenow var. littoralis
Engelmann]
Juncus articulatus Linnaeus—(Figure 18). 2n = 40, 80. Shores,
springy spots, ditches. [J. articulatus var. obtusatus Engel-
Juncus brachycarpus Engelmann—(Figure 18). 2n = 44. Salt
marshes, ocean beaches.
Juncus brachycephalus (Engelmann) Buchenau—(Figure 18). 2n
= 80. Basic shores, marshes, meadows, swamps.
Juncus brevicaudatus (Engelmann) Fernald—(Figure 18). 2n =
Shores, bogs, marshes.
Juncus bufonius Linnaeus—Toad Rush (Figure 19). 2n = 27-37,
10 Rhodora [Vol. 102
40, ca. 54, 58-81, 106, 1O8—115, 120. Shores, salt marshes,
roadsides, moist to wet borrow pits. [J. bufonius var. halo-
philus Buchenau & Fernald]
Juncus canadensis J. Gay—(Figure 19). 2n = 80. Shores,
swamps, marshes, meadows. [/. canadensis var. sparsiflorus
Fernald]
JUNCUS COMPRESSUS Jacques—(Figure 19). 2n = 40, 44.
Disturbed wet ground, especially ditches, roadsides, fre-
quently saline or basic soils. From Europe. [J. bu/bosus Lin-
naeus |
Juncus debilis A. Gray—(Figure 19). 2n = ?. Ditches, pools,
shores.
Juncus dichotomus Elliott—(Figure 20). 27 = 80. Sandy or
sphagnous shores, salt marsh borders. [/. platyphyllus (Wie-
gand) Fernald; J. tenuis Willdenow var. dichotomus (Elliott)
A. Wood]
JUNCUS DIFFUSISSIMUS Buckley. 2n = ?. Disturbed open bar-
rens. From farther south. [Voucher discovered while mss. in
press. South Windsor, Hartford County, CT (CONN). ]
Juncus dudleyi Wiegand—(Figure 20). 2n = 42, 80, ca. 84.
Shores, boggy meadows, in basic soils. [/. tenuis Willdenow
var. dudleyi (Wiegand) E J. Hermann]
Juncus effusus Linnaeus—Tufted Rush (Figure 20). 2n = 5, 40,
42. Marshy ground, low spots. [/. effusus var. compactus
Lejeune & Courtois; J. effusus var. conglomeratus (Linnaeus)
Engelmann; J. effusus var. costulatus Fernald; J. effusus var.
decipiens Buchenau; J. effusus var. pylaei (Laharpe) Fernald
& Wiegand; J. effusus var. solutus Fernald & Wiegand; J.
conglomeratus Linnaeus; J. pylaei Laharpe]
Juncus filiformis Linnaeus—(Figure 20). 2n = 40, 70, 80, 84.
Shores, swamps, bogs, alpine meadows.
Juncus gerardii Loiseleur-Deslongchamps—Black Grass (Figure
21). 2n = 80, 84. Salt marshes. [J. gerardii var. pedicellatus
Fernald]
Juncus greenet Oakes & Tuckerman—(Figure 21). 2n = 80. Dry
2000] Angelo and Boufford—Atlas of New England Flora 11
open places, usually well-drained, sandy soil. In pine lands
near lake shores, dunes.
JUNCUS INFLEXUS Linnaeus—(Figure 21). 2n = 38, 40. Mead-
ows, damp roadsides. From Eurasia and northern Africa.
Juncus marginatus Rostkovius—Grass Rush (Figure 21). 2n =
38, 40. Sandy pond margins, wet meadows. [J. biflorus El-
liott]
Juncus militaris Bigelow—Bayonet Rush (Figure 22). 2n = ?.
allow water of ponds and streams with sandy, gravelly, or
sphagnous margins.
Juncus nodosus Linnaeus var. nodosus—(Figure 22). 2n = 40.
Shores, marshes, meadows, swamps, especially basic soils.
Juncus pelocarpus E. Meyer—(Figure 22). 2n = 40. Sandy
shores, marshes, ditches.
Juncus secundus Beauvois ex Poiret—(Figure 22). 2n = ca. 80.
Ledges, dry open sterile soils.
Juncus stygius Linnaeus var. americanus Buchenau—(Figure 23).
2n = ?. Bogs, bog pools.
Juncus subcaudatus (Engelmann) Coville & S. EK Blake—(Figure
= ?. Swamps, bogs, shaded spring-heads, mossy
woade
JUNCUS SUBNODULOSUS Schrank—(Figure 23). 2n = 40.
Salt marsh borders. From Europe. [J. PERVETUS Fernald]
Juncus subtilis E. Meyer—Creeping Rush (Figure 23). 2n = 40.
Muddy shores.
Juncus tenuis Willdenow—Path Rush (Figure 24). 2n = 40, 80,
84. Roadsides, paths. [J. tenuis var. williamsii Fernald]
JUNCUS TORREYI Coville—(Figure 24). 2n = 40. Shores, ditch-
es, roadsides. From farther west and south.
Juncus trifidus Linnaeus—Highland Rush (Figure 24). 2n = 20,
0, 40. Exposed, rocky, or sterile summits. [J. trifidus var.
monanthos (Jacquin) Bluff & Fingerhuth]
Juncus vaseyi Engelmann—(Figure 24). 2n = 42, ca. 80.
wamps, shores, thickets.
12 Rhodora [Vol. 102
—Juncus hybrids—
Juncus articulatus Linnaeus < Juncus brevicaudatus (Engel-
mann) Fernald—(Figure 25).
Juncus brevicaudatus (Engelmann) Fernald * Juncus nodosus
Linnaeus var. nodosus—(Figure 25).
Juncus X oronensis Fernald—(Figure 25). Alder swamps, thick-
ets. [J. tenuis Willdenow X J. vaseyi Engelmann ?]
Juncus secundus Beauvois ex Poiret * Juncus tenuis Willden-
ow—(Figure 25).
Juncus tenuis Willdenow X Juncus vaseyi Engelmann—(Figure
26).
Luzula acuminata Rafinesque var. acuminata—(Figure 26). 2n =
18. Thickets, clearings, woods.
Luzula acuminata Rafinesque var. carolinae (S. Watson) Fer-
ald—2n = 18. Limy wooded slopes. [Reported (Flora of
North America, in press); no voucher seen. ]
Luzula bulbosa (Wood) Rydberg—(Figure 26). 2n = 12. Dry,
sandy, open woods and fields.
LUZULA CAMPESTRIS (Linnaeus) de Candolle—(Figure 26). 2n
= 12, 12 + 1B, 13, 14, 16. Lawns. From Europe.
Luzula confusa Lindeberg—Northern Woodrush (Figure 27). 2”
= 36, 44-48. Alpine areas.
Luzula echinata (Small) Hermann—(Figure 27). 2n = 12. Woods,
thickets, clearings. [L. campestris (Linnaeus) de Candolle
var. echinata (Small) Fernald & Wiegand]
LUZULA LUZULOIDES (Lamarck) Dandy & Wilmott subsp.
LUZULOIDES—Forest Woodrush (Figure 27). 20 = 12.
Rocky woods, roadsides, lawns. From Europe.
Luzula multiflora (Ehrhart) Lejeune subsp. mu/tiflora—Common
Woodrush (Figure 27). 2n = 12, 24, 28, 36. Open woods,
dry fields, meadows. [L. multiflora var. acadiensis Fernald:
L. campestris (Linnaeus) de Candolle var. multiflora (Ehr-
hart) Celakovsky]
Luzula multiflora (Ehrhart) Lejeune subsp. frigida (Buchenau)
2000] Angelo and Boufford—Atlas of New England Flora Ts)
Kreczetowicz—(Figure 28). 2” = 36. Sphagnous barrens,
clearings, fields. [L. multiflora var. fusconigra Celakovsky]
LUZULA PALLIDULA Kirschner—(Figure 28). 27 = 12-18.
Meadows, open woods, clearings, rocky places. From Eura-
sia. [L. pallescens (Wahlenberg) Besser]
Luzula parviflora (Ehrhart) Desvaux var. melanocarpa (Michaux)
Buchenau—(Figure 28). 2n = 24. Mossy wooded banks, dry
woods, damp thickets, often rocky places.
Luzula spicata (Linnaeus) de Candolle—Alpine Woodrush (Fig-
ure 28). 2n = 12, 14, 18, 24, 36. Alpine areas.
JUNCAGINACEAE
Triglochin gaspense Lieth & D. Léve—(Figure 29). 2n = 96.
Salt marshes.
Triglochin maritima Linnaeus—(Figure 29). 2n = 12, 24, 30, 36,
48, 96, 120. Salt, brackish, and freshwater marshes.
Triglochin palustre Linnaeus—(Figure 29). 2n = 24, 28, 36. Salt
marshes and river shores.
LEMNACEAE
Lemna minor Linnaeus—(Figure 29). 27 = 20, 30, 40, 42, 50,
63, 80, 126. Floating on quiet water of ponds and streams,
muddy shores.
Lemna perpusilla Torrey—(Figure 30). 2n = 20, 40, 42, 50, 60,
80. Floating on quiet water of ponds and streams.
Lemna trisulca Linnaeus—(Figure 30). 27 = 20, 40, 42, 44, 56—
60, 63, 80. Surface of quiet often basic water of ponds and
streams.
Lemna turionifera Landolt—(Figure 30). 21 = 40, 42, 50, 80.
Quiet waters.
Lemna valdiviana Philippi—(Figure 30). 2n = 36, 40, 42. Quiet
waters or swift currents of streams.
Spirodela polyrrhiza (Linnaeus) Schleiden—Water Flaxseed (Fig-
14 Rhodora [Vol. 102
ure 31). 2n = 30, 32, 38, 40, 50, 80. Surface of quiet water
of lakes, ponds, ditches, and streams; muddy shores.
SPIRODELA PUNCTATA (G. E W. Meyer) C. H. Thompson—
2n = 40, 43—44, 46, 50. Quiet waters. From subtropical re-
gions. [S. OLIGORRHIZA (Kurtz) Hegelmaier; reported from
Massachusetts (Flora of North America, in press); no vouch-
er seen|
Wolffia borealis (Engelmann ex Hegelmaier) Landolt—(Figure
2n = 20, 22, 30, 40. Floating on quiet waters of ditches,
ponds, lakes, and streams. [W. punctata misapplied]
WOLFFIA BRASILIENSIS Weddell—(Figure 31). 2n = 20, 40,
42, 50, 60, 80. Floating on quiet waters. From farther south.
[W. PAPULIFERA C. H. Thompson]
Wolffia columbiana Karsten—(Figure 31). 2n = 30, 40, 42, 50,
70. Quiet mostly basic waters of lakes, ponds, ditches, and
streams.
Wolffiella gladiata (Hegelmaier) Hegelmaier—(Figure 32). 2n =
40, 42. Quiet acidic waters. [W. floridana (J. D. Smith) C.
H. Thompson]
LILIACEAE
Note: The Liliaceae are here treated mostly in their traditional
sense with the understanding that numerous smaller families
will be recognized within the very near future.
Aletris farinosa Linnaeus—Unicorn-root (Figure 32). 2n = 26.
Moist, open, sandy soil and barrens.
Allium canadense Linnaeus var. canadense—Wild Garlic (Figure
32). 2n = 14, 21, 28, 82. Low woods, alluvial thickets, mead-
OWS.
ALLIUM CEPA Linnaeus—Onion (Figure 32). 2n = 14, 16, 28,
32, 64. Waste ground. From southwest Asia.
ALLIUM OLERACEUM Linnaeus—Wild Onion (Figure 33). 2n
= 24, 32, 40, 48. Wood borders, thickets, roadside banks.
From Europe.
2000] Angelo and Boufford—Atlas of New England Flora 15
ALLIUM SATIVUM Linnaeus—Garlic (Figure 33). 2n = 12, 16.
Roadsides, pastures, fields. From western Asia.
Allium schoenoprasum Linnaeus—Chives (Figure 33). 2n = 16,
16 + (1—18)B, 24, 32. Gravelly and rocky shores. [A. schoen-
oprasum var. sibiricum (Linnaeus) Hartman]
Allium tricoccum Aiton var. tricoccum—Wild Leek (Figure 33).
2n = 16. Rich basic woods and bottoms.
Allium tricoccum Aiton var. burdickii Hanes—(Figure 34). 2n =
16. Dry soil in upland woods.
ALLIUM VINEALE Linnaeus—Field Garlic (Figure 34). 2n = 16,
32, 32 + (0-2)s, 40. Dry grasslands, fallow fields, lawns,
waste places. From Europe.
ASPARAGUS OFFICINALIS Linnaeus—Asparagus (Figure 34).
2n = 20, 40. Roadsides, near buildings, fields, fence rows.
From Europe.
Chamaelirium luteum (Linnaeus) Gray—Blazing-star (Figure 34).
2n = ?. Meadows, thickets, rich woods.
Clintonia borealis (Aiton) Rafinesque—Yellow Clintonia (Figure
35). 2n = 28, 28 + 2B, 32. Woods, usually moist, thickets.
COLCHICUM AUTUMNALE Linnaeus—Autumn Crocus (Figure
35). 2n = 36, 38, 42. Meadows, fields. From Europe.
CONVALLARIA MAJALIS Linnaeus var. MA/JALIS—European
Lily-of-the-Valley (Figure 35). 2m = 36, 38. Roadsides, old
house sites, thickets, open woods. From Europe.
Erythronium americanum Ker—Trout Lily (Figure 35). 2n = 48.
Rich moist woods and thickets.
GALANTHUS NIVALIS Linnaeus subsp. NJVAL/JS—Snowdrop
(Figure 36). 2n = 18, 24, 24 + (1-10)B, 26. Abandoned
gardens. From Europe.
HEMEROCALLIS FULVA (Linnaeus) Linnaeus—Orange Day-
lily (Figure 36). 2n = 22, 33, 36. Roadsides, waste places.
From Asia.
HEMEROCALLIS LILIOASPHODELUS Linnaeus— Yellow Day-
lily (Figure 36). 27 = 22. Roadsides, waste places. From
Asia. [H. FLAVA (Linnaeus) Linnaeus]
16 Rhodora [Vol. 102
HOSTA LANCIFOLIA Trattinnick—(Figure 36). 2n = 60. Thick-
ets, roadsides, waste places. From eastern Asia. [H. JAPON-
ICA Voss]
HOSTA VENTRICOSA (Salisbury) Stearn—Blue_ Plantain-lily
(Figure 37). 2n = 60, 120. Rich woods along streams, moist
banks. From eastern Asia.
Hypoxis hirsuta (Linnaeus) Coville—Common Stargrass (Figure
37). 2n = 28. Open woods, fields.
LEUCOJUM AESTIVUM Linnaeus—Summer Snowflake (Figure
7). 2n = 22, 24. Meadows, low woods. From Europe.
Lilium canadense Linnaeus—Canada Lily (Figure 37). 2n = 24.
Meadows, low woods, thickets.
LILIUM LANCIFOLIUM Thunberg—tTiger Lily (Figure 38). 2n
= 24, 36. Old house sites, roadsides, dry thickets. From east-
ern Asia. [L. TIGRINUM Ker]
Lilium philadelphicum Linnaeus—Wood Lily (Figure 38). 2” =
24. Dry open woods, thickets, clearings.
Lilium superbum Linnaeus—Turk’s-cap Lily (Figure 38). 2n =
24. Meadows, damp thickets, swampy woods.
Maianthemum canadense Desfontaines—Canada Mayflower
(Figure 38). 2n = 36. Woods, clearings. [M. canadense var.
interius Fernald]
Maianthemum racemosum (Linnaeus) Link subsp. racemosum—
False Solomon’s-seal (Figure 39). 2n = 36, 72, 144. Woods,
clearings. [Smilacina racemosa (Linnaeus) Desfontaines;
Smilacina racemosa (Linnaeus) Desfontaines var. cylindrata
Fernald |
Maianthemum stellatum (Linnaeus) Link—(Figure 39). 2n = 36,
54. Sandy soil of shores, hillsides, fields, thickets. [Smilacina
stellata (Linnaeus) Desfontaines; Smilacina stellata (Linnae-
us) Desfontaines var. crassa Victorin]
Maianthemuna trifolium (Linnaeus) Sloboda—(Figure 39). 2n =
36. Sphagnum bogs, sphagnous shores and woods. [Smila-
cina trifolia (Linnaeus) Desfontaines]
2000] Angelo and Boufford—Atlas of New England Flora 17
Medeola virginiana Linnaeus—Indian Cucumber Root (Figure
39). 2n = 14. Rich woods.
MUSCARI BOTRYOIDES (Linnaeus) Miller—Grape-hyacinth
(Figure 40). 2n = 18, 36, ca. 40, 48. Pastures, fields, road-
sides, waste places. From Europe.
MUSCARI NEGLECTUM Gussone ex Tenore—Blue-bottle (Fig-
ure 40). 2n = 18-72. Fields, roadsides, lawns, waste places.
From Europe. [M. ATLANTICUM Boissier & Reuter; M. RA-
CEMOSUM Lamarck & de Candolle]
NARCISSUS POETICUS Linnaeus—Poet’s Narcissus (Figure
40). 2n = 14, 21, 28. Fields, moist meadows. From Europe.
NARCISSUS PSEUDONARCISSUS Linnaeus—Daffodil (Figure
40). 2n = 14, 14 + (1-2)B, 15, 21, 26-30, 35, 43. Fields,
open groves, moist meadows. From Europe.
ORNITHOGALUM NUTANS Linnaeus—(Figure 41). 2n = 14,
15, 40, 41, 45. Low meadows, fields. From western Asia.
ORNITHOGALUM UMBELLATUM Linnaeus—Star-of-Bethle-
hem (Figure 41). 27 = 18-108. Old house sites, roadsides,
thickets, fields, meadows. From Europe.
Polygonatum biflorum (Walter) Elliott var. biflorum—Great Sol-
omon’s-seal (Figure 41). 2” = 20, 40. Rocky woods.
Polygonatum biflorum (Walter) Elliott var. commutatum (J. A. &
J. H. Schultes) Morong—Giant Solomon’s-seal (Figure 41).
2n = 40. Rich woods, alluvial thickets, riverbanks, hedge-
rows. [P. canaliculatum misapplied; P. commutatum (J. A.
& J. H. Schultes) A. Dietrich]
POLYGONATUM LATIFOLIUM (Jacquin) Desfontaines—(Fig-
ure 42). 2n = 18, 20. Roadside thickets. From Europe.
Polygonatum pubescens (Willdenow) Pursh—Small Solomon’s-
seal (Figure 42). 2n = 20. Dry-to-rich woods.
SCILLA SIBERICA Haworth ex Andrews—(Figure 42). 2n = 12,
2+ B, 12 + 4B, 18, 24, 30. Roadsides, pastures. From
Eurasia.
Streptopus amplexifolius (Linnaeus) de Candolle—Twisted Stalk
18 Rhodora [Vol. 102
(Figure 42). 2n = 16, 32. Rich damp woods and thickets. [S.
amplexifolius var. americanus Schultes]
Streptopus lanceolatus (Aiton) Reveal—Rose Mandarin (Figure
43). 2n = 16, 48. Rich, damp, cool woods and thickets. [S.
roseus Michaux var. longipes (Fernald) Fassett: S. roseus Mi-
chaux var. perspectus Fassett]
—Streptopus hybrids—
Streptopus X oreopolus Fernald—(Figure 43), 2n = 24. Subal-
pine woods and meadows. [S. amplexifolius (Linnaeus) de
Candolle x S. lanceolatus (Aiton) Reveal; S$. amplexifolius
(Linnaeus) de Candolle var. oreopolus (Fernald) Fassett]
Tofieldia glutinosa (Michaux) Persoon—(Figure 43). 2n = 30.
Marshes (basic), bogs, shores.
TRICYRTIS HIRTA (Thunberg) Hooker—Toad Lily (Figure 43).
2n = 24, 26, 37. Open woods. From eastern Asia.
Trillium cernuum Linnaeus—Nodding Trillium (Figure 44). 21 =
10. Rich damp woods, most often in acid soil. [T. cernuum
var. macranthum A. J. Eames & Wiegand]
Trillium erectum Linnaeus—Purple Trillium (Figure 44). 21 =
. Rich woods. [7. erectum var. blandum Jennison]
Trillium grandiflorum (Michaux) Salisbury—Snowy_ Trillium
(Figure 44). 27 = 10. Rich usually basic woods and thickets.
Trillium undulatum Willdenow—Painted Trillium (Figure 44). 2
= 10. Rich usually wet woods in acidic soils.
TULIPA GESNERIA Linnaeus—Garden Tulip (Figure 45). 2n =
24, 25, 26, 36, 48. Waste areas. From Eurasia.
TULIPA SYLVESTRIS Linnaeus—(Figure 45). 2n = 24, 48.
Meadows. From Europe.
Uvularia grandiflora J. E. Smith—Big Merry-bells (Figure 45).
2n = 14. Rich moist woods and thickets, chiefly basic.
Uvularia perfoliata Linnaeus—Perfoliate Bellwort (Figure 45).
2n = 14. Rich usually dry woods and thickets, preferring
acid soils.
2000] Angelo and Boufford—Atlas of New England Flora 19
Uvularia sessilifolia Linnaeus—Wild-oats (Figure 46). 2n = 14.
Woods, thickets, clearings.
Veratrum viride Aiton—White Hellebore (Figure 46). 27 = 32.
Swampy woods, wet meadows.
Zigadenus elegans Pursh var. glaucus (Nuttall) Preece—White
Camass (Figure 46). 2 = 32. Basic gravel, cliffs, shores,
bogs. [Z. glaucus Nuttall]
NAJADACEAE
Najas flexilis (Willdenow)—Northern Water-nymph (Figure 46).
2n = 24. Quiet shallow water of ponds and streams, usually
rooting in mud.
Najas gracillima (A. Braun ex Engelmann) Magnus—(Figure
47). 2n = 12. Muddy, sandy, or sphagnous ponds and shores.
Najas guadalupensis (Sprengel) Magnus subsp. guadalupensis—
Southern Water-nymph (Figure 47). 2m = 12, 36, 48, 54, 60.
Ponds and streams.
Najas guadalupensis (Sprengel) Magnus subsp. olivacea (Rosen-
dahl & Butters) R. R. Haynes & Hellquist—(Figure 47). 2n
= ?, Habitat? [N. olivacea Rosendahl & Butters]
NAJAS MINOR Allioni—(Figure 47). 2n = 12, 24, 46, 56. Quiet
basic water of lakes and streams. From the Old World.
ORCHIDACEAE
Amerorchis rotundifolia (Banks) Hultén—Small Round-leaved
Orchis (Figure 48). 2n = 42. Bogs, swamps, boggy woods,
in basic soil. [Orchis rotundifolia Banks]
Aplectrum hyemale (Muhlenberg ex Willdenow) Nuttall—Putty-
root (Figure 48). 27 = ?. Rich rocky woods.
Arethusa bulbosa Linnaeus—Arethusa (Figure 48). 2n = 40.
Sphagnous bogs, meadows, and swamps.
Calopogon tuberosus (Linnaeus) Britton, Sterns & Poggenberg
Grass-pink (Figure 48). 2n = 26, 42. Sphagnous bogs,
swamps, and meadows. [C. pulchellus (Salisbury) R. Brown]
20 Rhodora [Vol. 102
Calypso bulbosa (Linnaeus) Oakes var. americana (R. Brown ex
Aiton f.) Luer—Calypso (Figure 49). 21 = 28. Thuja
swamps.
Coeloglossum viride (Linnaeus) Hartman var. virescens (Muhl-
enberg ex Willdenow) Luer—Bracted Orchis (Figure 49), 2n
= 40, 42. Rich moist woods, thickets, meadows. [Habenaria
viridis (Linnaeus) R. Brown var. bracteata (Muhlenberg) A.
Gray ]
Corallorhiza maculata (Rafinesque) Rafinesque—Spotted Coral-
root (Figure 49). 2n = 42, 84. Dry-to-moist woods.
Corallorhiza odontorhiza (Willdenow) Poiret—Autumn Coral-
root (Figure 49). 2n = ?. Rich dry woods, in basic soil.
Corallorhiza trifida Chatelain—Early Coral-root (Figure 50). 2n
= 38, 40, 42. Rich wet woods, swamps with Thuja. [C. tri-
fida var. verna (Nuttall) Fernald]
Cypripedium acaule Aiton—Pink Lady’s-slipper (Figure 50). 2n
= 20. Dry woods, acid soils.
Cypripedium arietinum R. Brown—Ram’s-head Lady’s-slipper
(Figure 50). 22 = 20. Rich damp woods, usually on hillsides,
usually acid soils in coniferous woods.
Cypripedium parviflorum Salisbury var. parviflorum—Small Yel-
low Lady’s-slipper (Figure 50). 27 = 20. Rich usually
swampy basic woods, bogs (chiefly basic), shores. [C. cal-
ceolus Linnaeus var. parviflorum (Salisbury) Fernald]
Cypripedium de lags Salisbury var. makasin (Farwell) Shev-
lak. 2n = ?. Thuja bogs and fens. [There are no distribution
data for this relatively newly described taxon since it is not
yet distinguished in New England herbaria. The author of
this combination reports that ‘‘ Virtually everything reported
from NE as var. parviflorum is in fact var. makasin”’ (pers.
comm.). |
Cypripedium parviflorum Salisbury var. pubescens (Willdenow)
Knight—Large Yellow Lady’s-slipper (Figure 51). 2n = 20.
Rich woodlands. [C. calceolus Linnaeus var. pubescens
(Willdenow) Correll]
Cypripedium reginae Walter—Showy Lady’s-slipper (Figure 51).
2n = 20. Bogs, swamps, swampy woods, in basic soils.
—
2000] Angelo and Boufford—Atlas of New England Flora 2
EPIPACTIS HELLEBORINE (Linnaeus) Crantz—Helleborine
(Figure 51). 2n = 36-44. Woods, thickets, roadsides. From
Europe.
Galearis spectabilis (Linnaeus) Rafinesque—Showy Orchis (Fig-
ure 51). 2n = 42. Rich woods, mostly in basic soils. [Orchis
spectabilis Linnaeus]
Goodyera oblongifolia Rafinesque—Giant Rattlesnake-plantain
(Figure 52). 2n = 22, 30. Dry coniferous and mixed woods.
Goodyera pubescens (Willdenow) R. Brown—Downy Rattle-
snake-plantain (Figure 52). 2n = 26. Dry-to-moist woods.
Goodyera repens (Linnaeus) R. Brown var. ophioides Fernald—
Dwarf Rattlesnake-plantain (Figure 52). 2n = 30, 40. Damp
mossy woods, especially under conifers.
—Goodyera hybrids—
Goodyera X tesselata Loddiges—(Figure 52). 21 = 59-61. Rich
woods, often pine. [G. oblongifolia Rafinesque x G. repens
(Linnaeus) R. Brown var. ophioides Fernald]
Isotria medeoloides (Pursh) Rafinesque—Small Whorled Pogonia
(Figure 53). 2n = 18. Open second growth, rich woods, often
near Fagus.
Isotria verticillata (Muhlenberg ex Willdenow) Rafinesque—
Large Whorled Pogonia (Figure 53). 2m = 18. Acidic woods,
usually damp, often with Medeola.
Liparis liliifolia (Linnaeus) Richard ex Lindley—Large Tway-
blade (Figure 53). 2n = ?. Rich moist woods.
Liparis loeselii (Linnaeus) Richard—Loesel’s Twayblade (Figure
2n = 26, 32. Swamps, bogs, damp thickets, sphagnous
meadows, ditches.
Listera auriculata Wiegand—Auricled Twayblade (Figure 54). 2n
= ?. Moist woods and thickets.
Listera australis Lindley—Southern Twayblade (Figure 54). 2n
= ?,. Sphagnous thickets and bogs.
Listera convallarioides (Swartz) Elliott—Broad-lipped Tway-
i)
i)
Rhodora [Vol. 102
blade (Figure 54). 27 = 36. Wet or swampy woods, often
with Thuja, shores.
Listera cordata (Linnaeus) R. Brown—Heartleaf Twayblade (Fig-
ure 54). 2n = 34, 34 + (1-9)B, 36-42, 44. Mossy knolls in
wet woods.
Malaxis bayardti Fernald—(Figure 55). 2” = ?. Dry sandy woods
and clearings.
Malaxis monophyllos (Linnaeus) Swartz var. brachypoda (A.
Gray) EF Morris & Eames—(Figure 55). 2n = 28, 30. Thuja
swamps and thickets. [M. brachypoda (A. Gray) Fernald]
Malaxis unifolia Michaux—Green Adder’s-mouth (Figure 55). 2
= ?. Woods, borders of swamps or bogs.
Platanthera_ blephariglottis (Willdenow) Lindley var. blephari-
glottis—White Fringed Orchis (Figure 55). 2n = 42. Sphag-
num bogs, wet sphagnous soil. [Habenaria blephariglottis
(Willdenow) Hooker]
Platanthera ciliaris (Linnaeus) Lindley—Yellow Fringed Orchis
(Figure 56). 217 = ?. Swampy woods, wet thickets, bogs.
[Habenaria ciliaris (Linnaeus) R. Brown]
Platanthera clavellata (Michaux) Luer—-Green Woodland Orchis
(Figure 56). 2n = 42. Swampy woods, bogs, spring-heads,
shores, typically sphagnous. [Habenaria clavellata (Mi-
chaux) Sprengel; H. clavellata var. ophioglossoides Fernald]
Platanthera cristata (Michaux) Lindley—Crested Yellow Orchis
(Figure 56). 2n = ?. Damp acid woods, low moist meadows.
[Habenaria cristata (Michaux) R. Brown]
Platanthera dilatata (Pursh) Lindley ex Beck var. dilatata—Bog
Candle (Figure 56). 2n = 42. Springy woods, bogs, shores,
meadows. [Habenaria dilatata (Pursh) Hooker]
Platanthera flava (Linnaeus) Lindley var. herbiola (R. Brown ex
Aiton f.) Luer—Tubercled Orchis (Figure 57). 2n = 42.
Springy meadows, shores. [Habenaria flava (Linnaeus) R.
Brown var. herbiola (R. Brown) Ames & Correll]
Platanthera grandiflora (Bigelow) Lindley—Large Purple
Fringed Orchis (Figure 57). 2n = 42. Rich swampy woods,
2000] Angelo and Boufford—Atlas of New England Flora = 23
spring-heads, along streams, thickets. [Habenaria fimbriata
(Aiton) R. Brown; H. psycodes (Linnaeus) Sprengel var.
grandiflora (Bigelow) A. Gray]
Platanthera hookeri (Torrey ex A. Gray)—Hooker’s Orchis (Fig-
ure 57). 2n = 42. Rich dry woods. [Habenaria hookeri Tor-
rey]
Platanthera hyperborea (Linnaeus) Lindley—Northern Green Or-
chis (Figure 57). 2n = 42, 84, 84 + 1. Springy woods, sphag-
num bogs, ditches. [P. hyperborea var. huronensis (Nuttall)
Luer; Habenaria hyperborea (Linnaeus) R. Brown; H. hy-
perborea (Linnaeus) R. Brown var. huronensis (Nuttall) Far-
well]
Platanthera lacera (Michaux) G. Don—Ragged Orchis (Figure
58). 2n = 42. Meadows, damp fields, alluvial or wet woods.
[Habenaria lacera (Michaux) Loddiges]
Platanthera leucophaea (Nuttall) Lindley—Prairie Fringed Or-
chis (Figure 58). 21 = 42. Bogs, open tamarack swamps.
[Habenaria leucophaea (Nuttall) A. Gray]
Platanthera obtusata (Banks ex Pursh) Lindley—Blunt-leaf Or-
chis (Figure 58). 2n = 42. Sphagnum bogs, damp woods,
especially coniferous or mixed. [Habenaria obtusata (Banks
ex Pursh) Richards]
Platanthera orbiculata (Pursh) Lindley var. orbiculata—Round-
leaved Orchis (Figure 58). 2n = 42. Rich woods. [Habenaria
orbiculata (Pursh) Torrey |
Platanthera orbiculata (Pursh) Lindley var. macrophylla (Goldie)
Luer—(Figure 59). 2n = ?. Rich woods. [Habenaria macro-
phylla Goldie; H. orbiculata (Pursh) Torrey var. macrophylla
(Goldie) B. Boivin]
Platanthera psycodes (Linnaeus) Lindley—Small Purple Fringed
Orchis (Figure 59). 2n = 42. Wet woods, damp thickets,
along streams. [Habenaria psycodes (Linnaeus) Sprengel]
—Platanthera hybrids—
Platanthera X andrewsii (M. White) Luer—(Figure 59). 2n =
42. [P. lacera (Michaux) G. Don X P. psycodes (Linnaeus)
Lindley ]
24 Rhodora [Vol. 102
Platanthera grandiflora (Bigelow) Lindley Platanthera hyper-
borea (Linnaeus) Lindley—(Figure 59).
Platanthera grandiflora (Bigelow) Lindley < Platanthera lacera
(Michaux) G. Don—(Figure 60).
Platanthera X media (Rydberg) Luer—(Figure 60). [P. dilatata
(Pursh) Lindley ex Beck var. dilatata X P. hyperborea (Lin-
naeus) Lindley |
Pogonia ophioglossoides (Linnaeus) Ker—Rose Pogonia (Figure
= 18. Sphagnum bogs, swamps, wet meadows, pond
shores, sphagnous thickets.
Spiranthes casei Catling & Cruise—(Figure 60). 2n = 60, 75.
Sandy acid soils, roadsides, fields.
Spiranthes cernua (Linnaeus) Richard—Nodding Ladies’ -tresses
(Figure 61). 2n = 30, 45, ca. 50, 60, 61. Damp banks, mead-
ows, bogs, shores, low thickets, open moist sandy places.
Spiranthes lacera (Rafinesque) Rafinesque var. Jacera—Northern
Slender Ladies’-tresses (Figure 61). 217 = 30. Open sandy
places.
Spiranthes lacera (Rafinesque) Rafinesque var. gracilis (Bigelow)
Luer—Southern Slender Ladies’-tresses (Figure 61). 2n =
30. Sterile open soils, thickets, and open woods. [S. gracilis
(Bigelow) Beck]
Spiranthes lucida (H. H. Eaton) Ames—(Figure 61). 2n = 44.
Damp rocky shores, meadows, rich damp thickets, usually in
basic soils.
Spiranthes ochroleuca (Rydberg) Rydberg—(Figure 62). 2n =
Sterile fields, dry barrens, rocky slopes, open woods,
pages [S. cernua (Linnaeus) Richard var. ochroleuca
(Rydberg) Ames]
Spiranthes romanz ps Chamisso—Hooded Ladies’ -tresses
(Figure 62). = 30, 44, 60. Swampy places, often along
rivers and an thickets.
Spiranthes tuberosa Rafinesque—Little Ladies’-tresses (Figure
62). 2n = ?. Dry sandy fields, woodland borders, cemeteries,
roadsides. [S. tuberosa var. grayi (Ames) Fernald]
2000} Angelo and Boufford—Atlas of New England Flora = 25
Spiranthes vernalis Engelmann & A. Gray—Spring Ladies’ -tress-
es (Figure 62). 2n = 30. Grasslands, sandy fields, clearings.
—Spiranthes hybrids—
Spiranthes X intermedia Ames—(Figure 63). [S. lacera (Rafin-
esque) Rafinesque var. lacera X S. vernalis Engelmann & A.
Gray |
Spiranthes lacera (Rafinesque) Rafinesque var. /acera x Spiran-
thes romanzoffiana Chamisso—(Figure 63).
Spiranthes lacera (Rafinesque) Rafinesque var. gracilis (Bigelow)
Luer X Spiranthes tuberosa Rafinesque—(Figure 63).
Tipularia discolor (Pursh) Nuttall—Crane-fly Orchid (Figure 63).
2n = ?. Rich, damp, oak-holly-beech woods.
Triphora trianthophora (Swartz) Rydberg subsp. triant
Nodding Pogonia (Figure 64). 2n = 44. Deep cianic ie -
moist woods with Fagus, often on rotten logs.
PONTEDERIACEAE
EICHHORNIA CRASSIPES (Martius) Solms-Laubach—Water-
hyacinth (Figure 64). 2n = 30, 32, 58. Floating in ponds and
quiet streams. From South America.
Heteranthera dubia (Jacquin) Mac argrass (Fig-
ure 64). 2n = 30. Shallow, quiet, basic water. [Zosterella
dubia (Jacquin) Small]
Heteranthera reniformis Ruiz & Pav6n—(Figure 64). 2n = 48.
Muddy river shores (in mud or floating in shallow water).
Pontederia cordata Linnaeus—Pickerelweed (Figure 65). 2n =
16. Shallow water of ponds, lakes, and river shores, rooting
in mud.
POTAMOGETONACEAE
Potamogeton alpinus Balbis—Red Pondweed (Figure 65). 21 =
52. Slow-moving streams, shallow water of ponds and
26 Rhodora [Vol. 102
lakes. [P. alpinus var. subellipticus (Fernald) Ogden; P. al-
pinus var. tenuifolius (Rafinesque) Ogden]
Potamogeton amplifolius Tackerman—(Figure 65). 2n = 52.
Deep water of lakes and river coves.
Potamogeton bicupulatus Fernald—(Figure 65). 2n = ?. Acidic
shallow water of rivers and ponds. [P. diversifolius Rafin-
esque var. trichophyllus Morong; P. capillaceus Poiret]
Potamogeton confervoides Reichenbach—Alga Pondweed (Fig-
ure 66). 2n = ?. Acidic sandy or sphagnous ponds, mountain
pools, and lakes.
POTAMOGETON CRISPUS Linnaeus—Curly Pondweed (Figure
66). 2n = 26, 36, 42, 50, 52, 72, 78. Shallow basic or brack-
ish water. From Europe.
Potamogeton diversifolius Rafinesque—Common Snailseed
Pondweed (Figure 66). 27 = ?. Acidic shallow water. [P.
capillaceus Poiret]
Potamogeton epihydrus Ratinesque—Ribbonleaf Pondweed (Fig-
ure 66). 27 = 26. Shallow quiet and moving water. [P. epihy-
drus var. nuttallii (Chamisso & Schlechtendahl) Fernald: P.
epihydrus var. ramosus (Peck) House]
Potamogeton foliosus Rafinesque subsp. foliosus-—Leafy Pond-
weed (Figure 67). 2n = 26, 28. Basic still water of ponds,
lakes, and streams. [P. foliosus var. macellus Fernald]
Potamogeton friesii Ruprecht—(Figure 67). 2n = 26. Alkaline
deep water of lakes and ponds.
Potamogeton gramineus Linnaeus—(Figure 67). 2n = 52. Shal-
l
water. [P. gramineus var. maximus Morong; P. grami-
neus var. myriophyllus Robbins]
Potamogeton hillii Morong—(Figure 67). 2n = 26. Alkaline wa-
ter of ponds and streams.
Potamogeton illinoensis Morong—(Figure 68). 2n = 104. Alka-
line water of ponds and streams.
Potamogeton natans Linnaeus—Floating Pondweed (Figure 68).
= 42, 52. Ponds and slow streams.
2000] Angelo and Boufford—Atlas of New England Flora = 27
Potamogeton nodosus Poiret—Longleaf Pondweed (Figure 68).
= 52. Streams.
Potamogeton oakesianus Robbins—Oakes Pondweed (Figure
2n = ?. Acidic ponds and lakes.
Potamogeton obtusifolius Mertens & Koch—(Figure 69). 2n =
26. Quiet basic water, usually cold.
Potamogeton ogdenii Hellquist & R. L. Hilton—Ogden’s Pond-
weed (Figure 69). 21 = ?. Basic water of ponds and lakes.
Potamogeton perfoliatus Linnaeus—Redhead-grass (Figure 69).
2n = 14, ca. 40, 52, 78. Shallow water. [P. perfoliatus var.
bupleuroides (Fernald) Farwell]
Potamogeton praelongus Wulfen—White-stem Pondweed (Fig-
u 2n = 52. Moderately alkaline still water, usually
deep.
Potamogeton pulcher Tuckerman—Spotted Pondweed (Figure
= ?. Acidic shallow water of ponds and muddy
shores.
Potamogeton pusillus Linnaeus subsp. pusillus—(Figure 70). 2n
Alkaline water of ponds and streams. [P. pusillus var.
minor (Bivona-Bivardi) Fernald & B. G. Schubert]
Potamogeton pusillus Linnaeus subsp. gemmiparus (Robbins) R.
Haynes & Hellquist—(Figure 70). 2n = 26. Quiet acidic
water of ponds and streams. [P. gemmiparus (Robbins) Rob-
bins ex Morong]
Potamogeton pusillus Linnaeus subsp. tenuissimus (Mertens &
Koch) R. R. Haynes & Hellquist—(Figure 70). 2n = 26.
Quiet shallow water of ponds and streams. [P. berchtoldii
Fieber; P. berchtoldii Fieber var. acuminatus Fieber; P. ber-
chtoldii Fieber var. lacunatus (Hagstr6m) Fernald; P. ber-
chtoldii Fieber var. polyphyllus (Morong) Fernald; P. ber-
chtoldii Fieber var. tenuissimus (Mertens & Koch) Fernald]
Potamogeton richardsonii (A. Bennett) Rydberg—Redhead
Pondweed (Figure 71). 2m = 26, 52. Alkaline water of lakes
and streams.
Potamogeton robbinsii Oakes—Fern Pondweed (Figure 71). 2n
= 52. Ponds, lakes, and slow streams.
28 Rhodora [Vol. 102
Potamogeton spirillus Tuckerman—Northern Snailseed Pond-
weed (Figure 71). 2n = ?. Neutral-to-acidic quiet water of
ponds and streams.
Potamogeton strictifolius A. Bennett—(Figure 71). 2n = 52. Al-
kaline water of lakes and slow streams. [P. strictifolius var.
rutiloides Fernald]
Potamogeton vaseyi Robbins—Vasey’s Pondweed (Figure 72). 2
= 28. Quiet water of ponds and streams, of low-to-moderate
alkalinity. [P. lateralis Morong]
Potamogeton zosteriformis Fernald—Flatstem Pondweed (Figure
72). 2n = 52. Quiet often alkaline water of ponds and slow
streams.
—Potamogeton hybrids—
Potamogeton alpinus Balbis X Potamogeton epihydrus Rafin-
esque—(Figure 72).
Potamogeton amplifolius Tuckerman * Potamogeton illinoensis
Morong—(Figure 72).
Potamogeton amplifolius Tuckerman X Potamogeton praelongus
Wulfen—(Figure 73).
Potamogeton X argutulus Hagstrom—(Figure 73). [P. gramineus
Linnaeus X P. nodosus Poiret]
Potamogeton X faxonii Morong—(Figure 73). [P. illinoensis Mo-
rong X P. nodosus Poiret]
Potamogeton X haynesii Hellquist & G. E. Crow—(Figure 73).
[P. strictifolius A. Bennett X P. zosteriformis Fernald: P.
longiligulatus misapplied]
Potamogeton illinoensis Morong X Potamogeton perfoliatus Lin-
naeus—(Figure 74),
Potamogeton illinoensis Morong * Potamogeton richardsonii (A.
Bennett) Rydberg—(Figure 74).
Potamogeton X mysticus Morong—(Figure 74). [P. perfoliatus
Linnaeus X P. pusillus Linnaeus subsp. tenuissimus (Mertens
& Koch) R. R. Haynes & Hellquist]
2000] Angelo and Boufford—Atlas of New England Flora = 29
Potamogeton X nericius Hagstt6m—(Figure 74). [P. alpinus Bal-
bis X P. gramineus Linnaeus]
Potamogeton X nitens G. Weber—(Figure 75). [P. gramineus
Linnaeus X P. perfoliatus Linnaeus]
Potamogeton perfoliatus Linnaeus * Potamogeton richardsontii
(A. Bennett) Rydberg—(Figure 75).
Potamogeton praelongus Wulfen xX Potamogeton richardsonii
(A. Bennett) Rydberg—(Figure 75).
Potamogeton X prussicus Hagstro6m—(Figure 75). [P. alpinus
Balbis X P. perfoliatus Linnaeus]
Potamogeton X spathuliformis (Robbins)—(Figure 76). [P. gra-
mineus Linnaeus X P. illinoensis Morong]
Stuckenia filiformis (Persoon) Borner subsp. alpina (Blytt) R. R.
Haynes, Les & M. Kral—(Figure 76). 2n = 78. Highly al-
kaline water of cold springs and lakes. [Potamogeton filifor-
mis Persoon var. borealis (Rafinesque) St. John; P. filiformis
Persoon var. macounii Morong]
Stuckenia filiformis (Persoon) Borner subsp. occidentalis (J. W.
Robbins) R. R. Haynes, Les & M. Kral—(Figure 76). 2n =
?. Highly alkaline water of cold streams.
Stuckenia pectinata (Linnaeus) Borner—Sago (Figure 76). 2m =
42, ca. 66, 70-87. Brackish or alkaline waters. [Potamogeton
pectinatus Linnaeus]
RUPPIACEAE
Ruppia maritima Linnaeus—(Figure 77). 2n = 14, 16, 20, 24,
28, 40. Saline or brackish water. [R. maritima var. longipes
Hagstr6m; R. maritima var. obliqua (Schur) Ascherson &
Graebner; R. maritima var. rostrata Agardh; R. maritima var.
subcapitata Fernald & Wiegand]
SCHEUCHZERIACEAE
Scheuchzeria palustris Linnaeus—(Figure 77). 2n = 22. Sphag-
num bogs. [S. palustris var. americana Fernald]
30 Rhodora [Vol. 102
SMILACACEAE
Smilax glauca Walter—Sawbrier (Figure 77). 21 = 32. Sandy
thickets, open woods. [S. glauca Walter var. leurophylla
Blake]
Smilax herbacea Linnaeus—Carrion-flower (Figure 77). 2n = 26.
Rich thickets, low woods.
Smilax pulverulenta Michaux—(Figure 78). 2n = ?. Rich mostly
basic woods and thickets. [S. herbacea Walter var. pulveru-
lenta (Michaux) A. Gray]
Smilax rotundifolia Linnaeus—Common Greenbrier (Figure 78).
2n = 32. Woods and thickets.
Smilax tamnoides Linnaeus—China-root (Figure 78). 27 = 32.
Low woods and thickets. [S. tamnoides var. hispida (Muhl-
enberg) Fernald; $. hispida Muhlenberg ex Torrey]
SPARGANIACEAE
Sparganium americanum Nuttall—(Figure 78). 2n = ?. Muddy
shores, shallow water.
Sparganium androcladum (Engelmann) Morong—(Figure 79). 2
= ?. Muddy shores, marshes, shallow water.
Sparganium angustifolium Michaux—(Figure 79). 2n = 30. Shal-
low-to-deep water, shores. [S. mu/tipedunculatum (Morong)
Rydberg]
Sparganium emersum Rehmann—(Figure 79). 2n = 30. Muddy
shores, marshes, shallow water. [S. chlorocarpum Rydberg;
S. chlorocarpum Rydberg var. acaule (Beeby) Fernald]
Sparganium eurycarpum Engelmann—(Figure 79). 2n =
Mostly basic muddy shores, marshes, shallow water.
Ww
=
Sparganium fluctuans (Engelmann ex Morong) B. L. Robinson—
(Figure 80). 2n = ?. Nonbasic lakes and ponds.
Sparganium natans Linnaeus—(Figure 80). 21 = 30. Shallow
pools, streams, shores, bogs. [$. minimum (Hartman) Fries]
2000] Angelo and Boufford—Atlas of New England Flora 31
—Sparganium hybrids—
Sparganium americanum Nuttall < Sparganium fluctuans (En-
gelmann ex Morong) B. L. Robinson—(Figure 80).
Sparganium angustifolium Michaux X Sparganium emersum
Rehmann—(Figure 80).
TYPHACEAE
ch angustifolia Linnaeus—(Figure 81). 2n = 30. Salt marsh-
, inland marshes, mostly alkaline, and near highways.
Typha latifolia Linnaeus—Common Cat-tail (Figure 81). 21 =
30. Marshes, shores, roadside ditches.
—Typha hybrids—
Typha X glauca Godron—(Figure 81). [7. angustifolia Linnaeus
x T. latifolia Linnaeus]
XYRIDACEAE
Xyris difformis Chapman var. difformis—Common Yellow-eyed
Grass (Figure 81). 2n = 18. Sandy shores of acid ponds,
lakes, and bogs. [X. caroliniana misapplied]
Xyris montana Ries—Small Yellow-eyed Grass (Figure 82). 2n
= ?. Sandy mostly acidic shores and bogs.
Xyris smalliana Nash—(Figure 82). 2n = 18. Muddy shores,
bogs, swamps, sandy or sphagnous shallows. LX. caroliniana
Walter; X. congdoni Small]
Xyris torta Smith—(Figure 82). 2n = 18. Sandy shores, sphag-
nous depressions, meadows.
ZANNICHELLIACEAE
Zannichellia palustris Linnaeus—Horned Pondweed (Figure 82).
2n = 12, 24, 28, 32, 34, 36, 48. Fresh-to-brackish water of
lakes, streams, and estuaries. [Z. palustris var. major (Hart-
man) W. D. J. Koch]
ee)
N
Rhodore [Vol. 102
ZOSTERACEAE
Zostera marina Linnaeus—Common Eel-grass (Figure 83). 27 =
hallow waters of coastal estuaries, coves, and bays. [Z.
marina var. stenophylla Ascherson & Graebner]
ACKNOWLEDGMENTS. We thank the curators and directors of
the herbaria of Harvard University, the University of Maine, the
University of Massachusetts, and the University of Vermont for
allowing us access to their collections. We particularly 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 and for verifying some voucher spec-
imens there. We also appreciate the research into voucher speci-
mens by Arthur Haines. Barre Hellquist gave especially gener-
ously of his time and knowledge to provide much information on
the aquatic groups. Kancheepuram Gandhi provided valuable as-
sistance in settling nomenclatural issues. Les Mehrhoff also was
very helpful in reviewing our Connecticut data and providing
many additional records. Janet Sullivan verified records at NHA.
Charles Sheviak provided information relative to Cypripedium
parviflorum. Anthony Reznicek searched for a voucher specimen
at MICH.
ADDENDUM. As this paper went to press an article in Rhodora (101:419—
423, 1999) by Donald H. Les and Robert S. Capers reported the collection
of Limnobium spongia (Bosc) Steudel (Hydrocharitaceae) from Tolland
County, Connecticut. The voucher specimen is deposited at CONN
2000] Angelo and Boufford—Atlas of New England Flora
Qo
es)
AROOSTOOK AROOSTOOK
NW NE
PISCATAQUIS
NEW
AROOSTOOK
HAMPSHIRE doanpin-e s
tae paainiCn
PISCATAQUIS
s
\ PENOBSCOT WASHINGTON
FRANKLIN Ss
SOMERSET
GRAND( ] FRANKLIN) ORLEANS coos S
ISLE N
OXFORD
N
KENNEBEC
MT. OESERT ISL
SAGADAHOC
MAINE
MIODLE-
SEX
ES ~ _., MASSACHUSETTS
ee DUKES a
CONNECTICUT _s ISLAND
re 1, Key map for counties of the New England states (and Mt. Desert
Island, Maine; Block Island, Rhode Island; arbitrary divisions of larger Maine
counties and of Cods County, New Hampshire)
34 Rhodora [Vol. 102
Acorus americanus ACORUS CALAMUS
oy
YUCCA FILAMENTOSA Alisma gramineum
Figure 2. Distribution maps for Acorus americanus, A. CALAMUS, YUCCA
FILAMENTOSA, and Alisma gramineum.
2000] Angelo and Boufford—Atlas of New England Flora = 35
Alisma subcordatum Alisma triviale
Echinodorus tenellus Sagittaria cuneata
Figure 3. Distribution maps for Alisma subcordatum, A. triviale,
Echinodorus tenellus, and Sagittaria cuneata.
36 Rhodora [Vol. 102
Saguttarta graminea Sagittarta latifolia
subsp. graminea
Figure 4. Distribution maps for Sagittaria engelmanniana, S. filiformis,
S. graminea subsp. graminea, and S. latifolia.
2000] Angelo and Boufford—Atlas of New England Flora = 37
Sagittaria montevidensis Sagittaria rigida
subsp. spongiosa
Sagittaria subulata Sagittaria teres
Figure 5. Distribution maps for Sagittaria montevidensis subsp. spongiosa,
S. rigida, S. subulata, and S. teres.
38 Rhodora [Vol. 102
Arisaema dracontium Arisaema triphyllum
© 'e@e
LL eet
is =
®@) 5 3
Arisaema dracontium Calla palustris
X A. triphyllum
Figure 6. Distribution maps for Arisaema dracontium, A. triphyllum,
A, dracontium X A. triphyllum, and Calla palustris.
2000] Angelo and Boufford—Atlas of New England Flora 39
Orontium aquaticum Peltandra virginica
Symplocarpus foetidus BUTOMUS UMBELLATUS
Figure 7. Distribution maps for Orontium aquaticum, Peltandra virginica,
Symplocarpus foetidus, and BUTOMUS UMBELLATUS.
40 Rhodora [Vol. 102
COMMELINA COMMUNIS COMMELINA DIFFUSA
TRADESCANTIA BRACTEATA Tradescantia ohiensis
Figure 8. Distribution maps for COMMELINA COMMUNIS, C. DIFFUSA,
TRADESCANTIA BRACTEATA, and T. ohiensis.
2000] Angelo and Boufford—Atlas of New England Flora 4]
Tradescantia virginiana Tradescantia ohiensis
Tradescantia ohiensis DIOSCOREA BATATAS
X T. virginiana
Figure 9. Distribution maps for Tradescantia virginiana, T. ohie sx
T. SUBASPERA, T. ohiensis X T. virginiana, and DIOSCOREA BATATAS
42 Rhodora [Vol. 102
Dioscorea villasa Eriocaulon aquaticum
Eriocaulon parkeri Lachnanthes caroliniana
Figure 10. Distribution maps for Dioscorea villosa, Eriocaulon aquaticum,
E. parkeri, and Lachnanthes caroliniana.
2000} Angelo and Boufford—Atlas of New England Flora 43
EGERIA DENSA Elodea canadensis
Elodea nuttallii HYDRILLA VERTICILLATA
Figure 11. Distribution maps for EGERIA DENSA, Elodea canadensis,
E. nuttallii, and HYDRILLA VERTICILLATA.
44 Rhodora [Vol. 102
Vallisneria americana BELAMCANDA CHINENSIS
me
Me
CROCUS VERNUS IRIS CRISTATA
subsp. VERNUS
Figure 12. Distribution maps for Vallisneria americana, BELAMCANDA
CHINENSIS, CROCUS VERNUS subsp. VERNUS, and IRIS CRISTATA.
2000] Angelo and Boufford—Atlas of New England Flora = 45
IRIS GERMANICA IRIS KAEMPFERI
Iris prismatica IRIS PSEUDACORUS
Figure 13. Distribution maps for JRJS GERMANICA, 1. KAEMPFERI,
I, prismatica, and I. PSEUDACORUS
46 Rhodora [Vol. 102
m3
IRIS PUMILA Iris setosa
subsp. PUMILA
IRIS SIBIRICA IRIS TECTORUM
Figure 14. Distribution maps for JRJS PUMILA subsp. PUMILA,
I. setosa, I. SIBIRICA, and I. TECTORUM.
2000] Angelo and Boufford—Atlas of New England Flora 47
Tris versicolor Tris prismatica
I. versicolor
Sisyrinchium albidum Sisyrinchium angustifolium
Figure 15. Distribution maps for Jris versicolor, I.. prismatica X
L versicolor, Sisyrinchium albidum, and S. angustifolium.
48 Rhodora [Vol. 102
Sisyrinchium atlanticum Sisyrinchium fuscatum
Sisyrinchium montanum Sisyrinchium mucronatum
var. crebrum
Figure 16. Distribution maps for Sisyrinchium atlanticum, S. fuscatum,
S. montanum var. crebrum, and S. mucronatum.
2000] Angelo and Boufford—Atlas of New England Flora 49
Juncus acuminatus Juncus alpinoarticulatus
Juncus anthelatus Juncus arcticus
var. balticus
Figure 17. Distribution maps for Juncus acuminatus,
J. alpinoarticulatus, J. anthelatus, and J. arcticus var. balticus.
50 Rhodora [Vol. 102
Juncus articulatus Juncus brachycarpus
Juncus brachycephalus Juncus brevicaudatus
Figure 18. Distribution maps for Juncus articulatus, J. brachycarpus,
J. brachycephalus, and J. brevicaudatus.
2000] Angelo and Boufford—Atlas of New England Flora — 51
Juncus bufonius Juncus canadensis
e* x oy
JUNCUS COMPRESSUS Juncus debilis
Figure 19. Distribution maps for Juncus bufonius, J. canadensis,
J. COMPRESSUS, and J. debilis.
52 Rhodora [Vol. 102
Juncus dichotomus Juncus dudleyi
Juncus effisus Juncus filiformis
Figure 20. Distribution maps for Juncus dichotomus, J. dudleyi,
J. effusus, and J. filiformis.
2000] Angelo and Boufford—Atlas of New England Flora = 53
Juncus gerardii Juncus greenei
JUNCUS INFLEXUS Juncus marginatus
igure 21. Distribution maps for Juncus gerardii, J. greenei,
J. INFLEXUS, and J. marginatus.
54 Rhodora [Vol. 102
Juncus militaris Juncus nodosus
var, nodosus
Juncus pelocarpus Juncus secundus
Figure 22. Distribution maps for Juncus militaris, J. nodosus
var. nodosus, J. pelocarpus, and J. secundus.
2000] Angelo and Boufford—Atlas of New England Flora 55
Juncus stygius Juncus subcaudatus
var. americanus
JUNCUS SUBNODULOSUS Juncus subtilis
Figure 23. Distribution maps for Juncus stygius vat. americanus,
J. subcaudatus, J. SUBNODULOSUS, and J. subtilis.
56 Rhodora [Vol. 102
om
=
Y
TAS >
Juncus tenuis JUNCUS TORREYI
& 3
Juncus trifidus Juncus vaseyi
Figure 24. Distribution maps for Juncus tenuis, J. TORREYI,
J. trifidus, and J. vaseyi.
2000] Angelo and Boufford—Atlas of New England Flora = 57
Juncus articulatus Juncus brevicaudatus
X J. brevicaudatus X J. nodosus var. nodusus
Juncus X oronensis Juncus secundus
Figure 25. Distribution maps for Juncus articulatus X J. brevicaudatus,
J. brevicaudatus X J. nodosus var. nodosus, J. X oronensis, and J. secundus
X J. tenuis
58 Rhodora [Vol. 102
Juncus tenuis Luzula acuminata
J. vaseyi var. acuminata
@ 2
Luzula bulbosa LUZULA CAMPESTRIS
Figure 26. Distribution maps for Juncus tenuis X J. vaseyl, Luzula
acuminata var. acuminata, L. bulbosa, and L. CAMPESTRIS.
2000] Angelo and Boufford—Atlas of New England Flora 59
Luzula confusa Luzula echinata
LUZULA LUZULOIDES Luzula multiflora
subsp. LUZULOIDES subsp. multiflora
Figure 27. Distribution maps for Luzula confusa, L. echinata,
L. LUZULOIDES subsp. LUZULOIDES, and L. multiflora subsp. multiflora.
60 Rhodora [Vol. 102
Luzula multiflora LUZULA PALLIDULA
subsp. frigida
Luzula parviflora Luzula spicata
var. melanocarpa
Figure 28. Distribution maps for Luzula multiflora subsp. frigida,
L. PALLIDULA, L. parviflora var. melanocarpa, and L. spicata.
2000] Angelo and Boufford—Atlas of New England Flora 61
Triglochin gaspense Triglochin maritima
Triglochin palustre Lemna minor
Figure 29. Distribution maps for Triglochin gaspense, T. maritima,
T. palustre, and Lemna minor.
62 Rhodora [Vol. 102
oy
Lemna perpusilla Lemna trisulca
Lemna turionifera Lemna valdiviana
Figure 30. Distribution maps for Lemna perpusilla, L. trisulca,
L. turionifera, and L. valdiviana.
2000] Angelo and Boufford—Atlas of New England Flora = 63
Spirodela polyrrhiza Wolffia borealis
~ 3
WOLFFIA BRASILIENSIS Wolffia columbiana
Figure 31. Distribution maps for Spirodela polyrrhiza, Wolffia borealis,
W. BRASILIENSIS, and W. columbiana.
64 Rhodora [Vol. 102
Wolffiella gladiata Aletris farinosa
Allium canadense ALLIUM CEPA
var. canadense
Figure 32. Distribution maps for Wolffiella gladiata, Aletris farinosa,
Allium canadense var. canadense, and A. CEPA.
2000] Angelo and Boufford—Atlas of New England Flora = 65
ALLIUM OLERACEUM ALLIUM SATIVUM
Allium schoenoprasum Allium tricoccum
var. tricoccum
Figure 33. Distribution maps for ALLIUM OLERACEUM, A. SATIVUM,
A. schoenoprasum, and A. tricoccum var. tricoccum.
66 Rhodora [Vol. 102
ALLIUM VINEALE
ASPARAGUS OFFICINALIS Chamaelirium luteum
Figure 34. Distribution maps for Allium tricoccum var. burdickii,
A, VINEALE, ASPARAGUS OFFICINALIS, and Chamaelirium luteum.
2000] Angelo and Boufford—Atlas of New England Flora 67
Clintonia borealis COLCHICUM AUTUMNALE
CONVALLARIA MAJALIS Erythronium americanum
var. MAJALIS
Figure 35. Distribution maps for Clintonia borealis, COLCHICUM
AUTUMNALE, CONVALLARIA MAJALIS var. MAJALIS, and Erythronium
americanum.
68 Rhodora [Vol. 102
GALANTHUS NIVALIS HEMEROCALLIS FULVA
subsp. NIVALIS
any
HEMEROCALLIS LILIOASPHODELUS HOSTA LANCIFOLIA
Figure 36. Distribution maps for GALANTHUS NIVALIS subsp. NIVALIS,
HEMEROCALLIS FULVA, H. LILIOASPHODELUS, and HOSTA
LANCIFOLIA.
2000] Angelo and Boufford—Atlas of New England Flora 69
4
any
HOSTA VENTRICOSA
LEUCOJUM AESTIVUM Lilium canadense
Figure 37. Distribution maps for HOSTA VENTRICOSA, Hypoxis hirsuta,
LEUCOJUM AESTIVUM, and Lilium canadense.
70 Rhodora [Vol. 102
LILIUM LANCIFOLIUM Lilium philadelphicum
Lilium superbum Maianthemum canadense
Figure 38. Distribution maps for LILIUM LANCIFOLIUM,
L. philadelphicum, L. superbum, and Maianthemum canadense.
2000] Angelo and Boufford—Atlas of New England Flora 7]
Maianthemum racemosum Maianthemum stellatum
subsp. racemosum
Maianthemum trifolium Medeola virginiana
Figure 39. Distribution maps for Maianthemum racemosum
subsp. racemosum, M. stellatum, M. trifolium, and Medeola virginiana.
We Rhodora [Vol. 102
oa
MUSCARI BOTRYOIDES MUSCARI NEGLECTUM
NARCISSUS POETICUS NARCISSUS PSEUDONARCISSUS
Figure 40. Distribution maps for MUSCARI BOTRYOIDES,
M. NEGLECTUM, NARCISSUS POETICUS, and N. PPEUDONARCISSUS.
2000] Angelo and Boufford—Atlas of New England Flora = 73
ORNITHOGALUM NUTANS ORNITHOGALUM UMBELLATUM
Polygonatum biflorum
Polygonatum biflorum
var. biflorum var. commutatum
Figure 41. Distribution maps for ORNITHOGALUM NUTANS,
O. UMBELLATUM, Polygonatum biflorum var. biflorum, and P. biflorum
var. commutatum.
74 Rhodora [Vol. 102
ay
POLYGONATUM LATIFOLIUM Polygonatum pubescens
os
SCILLA SIBERICA Streptopus amplexifolius
Figure 42. Distribution maps for POLYGONATUM LATIFOLIUM,
P. pubescens, SCILLA SIBERICA, and Streptopus amplexifolius.
2000] Angelo and Boufford—Atlas of New England Flora 13
Streptopus lanceolatus Streptopus X oreopolus
oa
Tofieldia glutinosa TRICYRTIS HIRTA
e 43. Distribution maps for Streptopus lanceolatus, S. X oreopolus,
ee glutinosa, and TRICYRTIS HIRTA.
76 Rhodora [Vol. 102
Trillium cernuum Trillium erectum
Trillium grandiflorum Trillium undulatum
Figure 44. Distribution maps for Trillium cernuum, T. erectum,
T. grandiflorum, and T. undulatum.
2000] Angelo and Boufford—Atlas of New England Flora = 77
TULIPA GESNERIA TULIPA SYLVESTRIS
Uvularia grandiflora Uvularia perfoliata
Figure 45. Distribution maps for TULIPA GESNERIA, T. SYLVESTRIS,
Uvularia grandiflora, and U. perfoliata.
78 Rhodora [Vol. 102
Uvularia sessilifolia Veratrum viride
Zigadenus elegans Najas flexilis
var. glaucus
Figure 46. Distribution maps for Uvularia sessilifolia, Veratrum viride,
Zigadenus elegans var. glaucus, and Najas flexilis.
2000] Angelo and Boufford—Atlas of New England Flora 79
Najas gracillima Najas guadalupensis
subsp. guadalupensis
Najas guadalupensis NAJAS MINOR
subsp. olivacea
Figure 47. Distribution maps for Najas gracillima, N. guadalupensis
subsp. guadalupensis, N. guadalupensis subsp. olivacea, and N. MINO
80 Rhodora [Vol. 102
Amerorchis rotundifolia Aplectrum hyemale
Arethusa bulbosa Calopogon tuberosus
Figure 48. Distribution maps for Amerorchis rotundifolia, Aplectrum
hyemale, Arethusa bulbosa, and Calopogon tuberosus.
2000] Angelo and Boufford—Atlas of New England Flora 81
Calypso bulbosa Coeloglossum viride
var. americana var. virescens
Corallorhiza maculata Corallorhiza odontorhiza
Figure 49. Distribution maps for Calypso bulbosa var. americana,
Coeloglossum viride var. virescens, Corallorhiza maculata, and
C. odontorhiza.
82 Rhodora [Vol. 102
Corallorhiza trifida Cypripedium acaule
Cypripedium arietinum Cypripedium parviflorum
var. parviflorum
Figure 50. Distribution maps for Corallorhiza trifida, Cypripedium
acaule, C. arietinum, and C. parviflorum var. parviflorum.
2000] Angelo and Boufford—Atlas of New England Flora 83
os
Cypripedium parviflorum Cypripedium reginae
var. pubescens
ran
EPIPACTIS HELLEBORINE Galearis spectabilis
Figure 51. Distribution maps for Cypripedium parviflorum var. pubescens,
C. reginae, EPIPACTIS HELLEBORINE, and Galearis spectabilis.
84 Rhodora [Vol. 102
Goodyera oblongifolia
Goodyera repens Goodyera X tesselata
var. ophioides
Figure 52. Distribution maps for Goodyera oblongifolia, G. pubescens,
G. repens var. ophioides, and G. X tesselata.
2000] Angelo and Boufford—Atlas of New England Flora = 85
Isotria medeoloides Isotria verticillata
Liparis liliifolia Liparis loeselii
Figure 53. Distribution maps for /sotria medeoloides, I. verticillata,
Liparis liliifolia, and L. loeselii.
86 Rhodora [Vol. 102
aan
Listera auriculata Listera australis
os
Listera convallarioides Listera cordata
Figure 54. Distribution maps for Listera auriculata, L. australis,
L. convallarioides, and L. cordata.
2000] Angelo and Boufford—Atlas of New England Flora = 87
Malaxis bayardii Malaxis monophyllos
var. brachypoda
Malaxis unifolia Platanthera blephariglottis
var. blephariglottis
Figure 55. Distribution maps for Malaxis bayardii, M. monophyllos
var. brachypoda, M. unifolia, and Platanthera blephariglottis
var. blephariglottis.
88 Rhodora [Vol. 102
Platanthera ciliaris Platanthera clavellata
ony
Platanthera cristata Platanthera dilatata
var. dilatata
Figure 56. Distribution maps for Platanthera ciliaris, P. clavellata,
P. cristata, and P. dilatata var. dilatata.
2000] Angelo and Boufford—Atlas of New England Flora 89
Platanthera flava Platanthera grandiflora
var. herbiola
~ 3
Platanthera hookeri Platanthera hyperborea
Figure 57. Distribution maps for Platanthera flava var. herbiola,
P. grandiflora, P. hookeri, and P. hyperborea.
90 Rhodora [Vol. 102
Platanthera lacera
~ 3
Platanthera obtusata Platanthera orbiculata
var. orbiculata
Figure 58. Distribution maps for Platanthera lacera, P. leucophaea,
P. obtusata, and P. orbiculata var. orbiculata.
2000] Angelo and Boufford—Atlas of New England Flora 9]
Platanthera orbiculata Platanthera psycodes
var. macrophylla
Platanthera X andrewsii Platanthera grandiflora
X P. hyperborea
Figure 59. Distribution maps for Platanthera orbiculata var. macrophylla,
P. psycodes, P. X andrewsii, and P. grandiflora X P. hyperborea.
92 Rhodora [Vol. 102
Platanthera grandiflora Platanthera X media
cera
Pogonia ophioglossoides Spiranthes casei
Figure 60. Distribution maps for Platanthera grandiflora X P. lacera,
P. X media, Pogonia ophioglossoides, and Spiranthes casei.
2000] Angelo and Boufford—Atlas of New England Flora 93
Spiranthes cernua Spiranthes lacera
var. lacera
Spiranthes lacera Spiranthes lucida
var. gracilis
Figure 61. Distribution maps for Spiranthes cernua, S. lacera var. lacera,
S. lacera var. gracilis, and S. lucida.
94 Rhodora [Vol. 102
a
3
Spiranthes ochroleuca Spiranthes romanzoffiana
Spiranthes tuberosa Spiranthes vernalis
Figure 62. Distribution maps for Spiranthes ochroleuca,
S. romanzoffiana, S. tuberosa, and S. vernalis.
2000] Angelo and Boufford—Atlas of New England Flora = 95
Spiranthes X intermedia Spiranthes lacera var. lacera
X S. romanzoffiana
Spiranthes lacera var. gracilis Tipularia discolor
X S. tuberosa
Figure 63. Distribution maps for Spiranthes X intermedia, S. lacera
var. lacera X S. romanzoffiana, S. lacera var. gracilis X S. tuberosa,
and Tipularia discolor.
96 Rhodora [Vol. 102
ee
oy
Triphora trianthophora EICHHORNIA CRASSIPES
subsp. trianthophora
on oy
Heteranthera dubia Heteranthera reniformis
Figure 64. Distribution maps for Triphora trianthophora
subsp. trianthophora, EICHHORNIA CRASSIPES, Heteranthera dubia,
and H. reniformis.
2000] Angelo and Boufford—Atlas of New England Flora 97
Pontederia cordata Potamogeton alpinus
Potamogeton amplifolius Potamogeton bicupulatus
Figure 65. Distribution maps for Pontederia cordata, Potamogeton
alpinus, P. amplifolius, and P. bicupulatus.
98 Rhodora [Vol. 102
Potamogeton confervoides POTAMOGETON CRISPUS
oo
Potamogeton diversifolius Potamogeton epihydrus
Figure 66. Distribution maps for Potamogeton confervoides, P. CRISPUS,
P. diversifolius, and P. epihydrus.
2000] Angelo and Boufford—Atlas of New England Flora 99
Potamogeton foliosus Potamogeton friesti
subsp. foliosus
Potamogeton gramineus Potamogeton hillii
Figure 67. Distribution maps for Potamogeton foliosus subsp. foliosus,
P, friesii, P. gramineus, and P. hillii.
100 Rhodora [Vol. 102
o 3
Potamogeton illinoensis Potamogeton natans
Potamogeton nodosus Potamogeton oakesianus
Figure 68. Distribution maps for Potamogeton illinoensis, P. natans,
P. nodosus, and P. oakesianus.
—
2000] Angelo and Boufford—Atlas of New England Flora 101
Potamogeton obtusifolius Potamogeton ogdenii
Potamogeton perfoliatus Potamogeton praelongus
Figure 69. Distribution maps for Potamogeton obtusifolius, P. ogdenit,
P. perfoliatus, and P. praelongus.
102 Rhodora [Vol. 102
m2
Potamogeton pulcher Potamogeton pusillus
ubsp. pusillus
sale lal pusillus Potamogeton pusillus
subsp. gemmiparus subsp. tenuissimus
Figure 70. Distribution maps for Potamogeton pulcher, P. —
subsp. pusillus, P. pusillus subsp. gemmiparus, and P. pusi
subsp. tenuissimus.
2000] Angelo and Boufford—Atlas of New England Flora 103
Potamogeton richardsonii Potamogeton robbinsii
Potamogeton spirillus Potamogeton strictifolius
Figure 71. Distribution maps for Potamogeton richardsonii, P. robbinsit,
P. spirillus, and P. strictifolius.
104 Rhodora [Vol. 102
Potamogeton vaseyi Potamogeton zosteriformis
Potamogeton alpinus Potamogeton amplifolius
X P. epihydrus X P. illinoensis
Figure 72. Distribution maps for Potamogeton vaseyi, P. zosteriformis,
P. alpinus X P. epihydrus, and P. amplifolius X P. illinoensis.
2000] Angelo and Boufford—Atlas of New England Flora 105
Potamogeton amplifolius Potamogeton X argutulus
X P. praelongus
Potamogeton X faxonii Potamogeton X haynesii
Figure 73. Distribution maps for Potamogeton amplifolius X P. praelongus,
P. X argutulus, P. X faxonii, and P. X haynesii.
106 Rhodora [Vol. 102
Potamogeton illinoensis Potamogeton illinoensis
X P. perfoliatu X P. richardsonii
~ 4
Potamogeton X mysticus Potamogeton X nericius
Figure 74. Distribution maps for Potamogeton illinoensis
X P. perfoliatus, P. illinoensis X P. richardsonii, P. X mysticus, and
P. X nericius.
—
2000] Angelo and Boufford—Atlas of New England Flora 107
Potamogeton X nitens Potamogeton perfoliatus
X P. richardsonii
Potamogeton praelongus Potamogeton X prussicus
P. richardsonii
Figure 75. Distribution maps for Potamogeton X nitens, P. perfoliatus
X P. richardsonii, P. praelongus X P. richardsonii, and P. X prussicus.
108 Rhodora [Vol. 102
& 3
Potamogeton X spathuliformis Stuckenia filiformis
subsp. alpina
Stuckenia filiformis Stuckenia pectinata
subsp. occidentalis
Figure 76. Distribution maps for Potamogeton X spathuliformis,
Stuckenia filiformis subsp. alpina, S. filiformis subsp. occidentalis, and
S. pectinata.
2000] Angelo and Boufford—Atlas of New England Flora 109
Ruppia maritima Scheuchzeria palustris
Smilax glauca Smilax herbacea
Figure 77. Distribution maps for Ruppia maritima, Scheuchzeria
palustris, Smilax glauca, and S. herbacea.
110 Rhodora [Vol. 102
Smilax pulverulenta Smilax rotundifolia
Smilax tamnoides Sparganium americanum
Figure 78. Distribution maps for Smilax pulverulenta, S. rotundifolia,
S. tamnoides, and Sparganium americanum.
2000] Angelo and Boufford—Atlas of New England Flora’ 111
Sparganium androcladum Sparganium angustifolium
Sparganium emersum Sparganium eurycarpum
Figure 79. Distribution maps for Sparganium androcladum,
S. angustifolium, S. emersum, and S. eurycarpum.
112 Rhodora [Vol. 102
Sparganium fluctuans Sparganium natans
Sparganium americanum Sparganium angustifolium
X S. fluctuans X S. emersum
Figure 80. Distribution maps for Sparganium fluctuans, S. natans,
S. americanum X S. fluctuans, S. angustifolium X S. emersum.
2000] Angelo and Boufford—Atlas of New England Flora’ 113
Typha angustifolia Typha latifolia
Typha X glauca Xyris difformis
var. difformis
Figure 81. Distribution maps for Typha angustifolia, T. latifolia,
T. X glauca, and Xyris difformis var. difformis.
114 Rhodora [Vol. 102
oy
Xyris montana Xyris smalliana
Xyris torta Zannichellia palustris
Figure 82. Distribution maps for Xyris montana, X. smalliana,
X. torta, and Zannichellia palustris.
2000] Angelo and Boufford—Atlas of New England Flora
Zostera marina
Figure 83. Distribution map for Zostera marina.
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Bene a M. 1980. sate UA of the Arisaema aoe complex. Ph.D.
dissertation, Univ. North Carolina, Chapel Hill,
Tucker, G. C. 1989. The genera of Commelinaceae in aie southeastern Unit-
ed States. J. Arnold Arbor. 70: 97-130.
Urecu, FE H. 1973. A biosystematic study of the genus Clintonia Raf. (Lili-
aceae: Polygonatae). Ph.D. dissertation, Washington Univ., St.
WILBUR, R. Le 1963. A revision of the North American genus Uvularia (Lil-
aceae). Rhodora 65: 158—
WiGomEN: J. W. 1973. Taxonomy of seven species of Sagittaria from eastern
orth America. Brittonia 25: 64-74.
ZIKA, PE AND J. Jenkins. 1995. Contributions to the flora of Vermont. Rho-
dora 97: 291-327.
ZOMLEFER, W. B. 1996. The o_o in the southeastern United States.
vard Pap. Bot. 9: 91—
"1997. The genera 7 Mamie in the southeastern United States.
aevaid Pap. Bot. 2: —177.
. 1997. The genera i ieee in the southeastern United States.
Harvard Pap. Bot. 2: 179-16
RHODORA, Vol. 102, No. 909, pp. 120-125, 2000
NEBC MEETING NEWS
September 1999. Dr. Les Mehrhoff, curator of the G. Safford
Torrey Herbarium at the University of Connecticut, addressed the
topic, ““The Non-native Invasive and Potentially Invasive Flora
of New England: A Regional Perspective.”’ He explained first that
his concern was primarily about non-native species that are out-
competing native flora in portions of the “‘minimally managed”’
landscape. Where once we had old fields succeeding to Rubus,
now we are likely to see the ubiquitous Rosa multiflora con-
quering the abandoned opening. Although early American bota-
nist John Bartram complained in print about the proliferation of
Narcissus cultivars into the natural environment around Phila-
delphia, there was litthe mention in the scientific literature of
problematic plant introductions until a pair of papers authored by
M. L. Fernald in 1905 and 1939 described certain “fugitives that
had escaped.”’ Reflective of the change in sentiment toward cer-
tain species, Fernald commented that the common name for Hier-
aceum aurantiacum in his native Maine changed from Venus’
paintbrush to Devil’s paintbrush. In the later paper, he noted a
number of Asian taxa, such as Lonicera japonica, that were
“crossing the landscape like a horde of huns.’’ Mehrhoff pointed
out that the term “invasive” should apply only to non-indigenous
species, because technically a species cannot invade its own ter-
ritory. When natives behave aggressively, he recommends we use
the word “‘explosive.”” So as not to malign any species as inher-
ently ““bad,”’ Les cited a line from an Aldo Leopold paper in the
1940s that says that “the invasive species problem ts an attribute
of numbers, not species.’ If not for herbarium records, Mehrhoff
pointed out, it would be hard to know whether certain taxa were
native or weedy introductions. An example given was the wide-
spread Dusty Miller, Artemesia stelleriana, a species for which
specimen collected by William Farlow in Nahant, Massachusetts.
Mehrhoff then discussed current efforts by himself and others
in New England to develop definitions and criteria to be used for
determining which species are invasive or potentially invasive in
the New England region. He listed the following as characteristics
of invasive species: 1) being non-native, 2) having high seed
production, 3) being capable of rapid dispersal, 4) having the
ability to establish easily, 5) having a competitive advantage over
L200
2000] NEBC Meeting News 12]
associated plants, and 6) being persistent on the landscape. He
then recognized two categories of non-native invasive species:
‘“Widespread and Invasive,” and “‘Restricted and Invasive.’ He
and New England colleagues are calling a third “‘watch-list”
group ‘Potentially Invasive Species.”’ To determine to which of
these three groups a species might belong, Mehrhoff, being a
good taxonomist, has developed a work sheet that progresses
through a set of criteria like a dichotomous key. The four primary
questions are: (1) Is it naturalized? (2) Is it capable of rapid and
widespread dispersion and establishment? (3) Is it capable of dis-
persing over spatial gaps? (4) Is it capable of existing in high
numbers away from artificial habitats? Four additional basic cri-
teria are: (5) Is it currently widespread or at least common in the
region or in one or more habitat types? (6) Is it known to have
numerous individuals in many populations in the region? (7) Does
it out-compete other species in the same natural plant commu-
nity? (8) Does it have the potential for rapid growth, high seed
or propagule production, dissemination, and establishment in nat-
ural areas? To be considered truly invasive the answer must be
‘“‘yves”’ to the eight questions or criteria. To distinguish between
Widespread and Restricted Invasives, one must determine wheth-
er the species is widespread with many occurrences in minimally
managed natural areas or simply common in part of the region
or in one or more habitat types in the region. The species qualifies
as Potentially Invasive if the answer is “yes” to the first four
questions, but “‘no”’ to one or more of the remaining four ques-
tions. Over 50 slide images were used to illustrate taxa being
considered for status as Invasive or Potentially Invasive. A draft
list for New England was also distributed along with a request
for feedback on it.
What else is needed to deal with the problem of invasive spe-
cies? Among Mehrhoff’s answers to this question was to do further
inventory and research, including that aimed at getting a better
understanding of species biology for many of the purported invad-
ers. Early detection and removal of new invaders would be another
important action, he said. Another avenue he encouraged was
working with the nursery and landscaping industries to find native
alternatives, to test and monitor new exotic introductions, and to
educate their customers. A final item fitting into Mehrhoff’s so-
lutions would be a computerized ‘‘Atlas of Non-native Invasive
Species in New England” that would serve as a database of current
i)
NO
Rhodora [Vol. 102
and historical records. This, along with other information about
invasive plants, he envisioned being available at a web site. Cur-
rently, the web site for the ner earn at ae ey of Con-
necticut (www eeb.uconn.ed ium/herbarium.
html) has a draft of Mehrhoff’s list for the state.
October 1999. Rick Enser, State Botanist with the Rhode Island
Natural Heritage Program, presented a slide lecture entitled ‘‘The
Flora of Block Island, Rhode Island.”’ Introducing his talk, Enser
explained that in his overview of Block Island’s flora, he would
attempt to answer three questions about the island: how it came
to be, what habitat types are present, and what factors are influ-
encing the current vegetation and flora? Addressing the first ques-
tion, he stated that Block Island’s flora and fauna relate to the
island’s glacial history. The island is a product of glaciation, re-
sulting from an ice sheet pushing up ocean sediments consisting
primarily of clay to form much of the island’s mass and charac-
teristic bluffs. Glacial till and debris cap the sediments in places
and bedrock is absent. The island is about 4.5 miles wide at its
widest point and possesses 6030 acres. Using maps of the coastal
islands and the continental shelf as they appeared following the
retreat of the glaciers, he suggested that the flora and fauna of
Block Island became isolated from the mainland about 8—9000
YBP, much earlier than occurred on Martha’s Vineyard and Nan-
tucket, which he dated at about 4000 YBP. This long isolation,
he feels, is a primary reason for the depauperate flora and fauna
on the island relative to some other islands of its size in the
region. The indigenous fauna of the island, he pointed out, in-
cludes only two mammals, four amphibians, and six reptiles. An-
other reason suggested for the depauperate biota is the island’s
lack of a glacial outwash-plain.
Documentation of the early flora and vegetation of the island
is limited. Livermore, in 1875, cited historic records from the
1600s about certain trees being used as boundary markers and
surmised from this that forest must have been present on the
island previously, even though it and the boundary trees were
absent by this time. Pollen and wood fragment data suggest that
a deciduous forest was once present on the island, too. From the
time of the Revolutionary War to the present, the island has been
largely an open, agricultural landscape grazed by livestock such
as horses, cows, and sheep. Early botanical exploration by Rob-
i)
W
2000] NEBC Meeting News
bins in 1829 documented two Rhode Island records, Arenaria
caroliniana and Hydrocotyle verticillata, not seen since on the
island. Other botanical explorers of the island to follow were
Henri Young, Stephen T. Olney, and William W. Bailey. In an
1893 article in the Bulletin of the Torrey Botanical Club, Bailey
reported an island flora of 294 species. An interesting record by
Bailey was Ranunculus cymbalaria, which he described as abun-
dant around the perimeter of Great Salt Pond. This once-brackish
pond has since been permanently breached and the buttercup can
no longer be found there, Enser said. Botanists following on Bai-
ley’s heels included Arthur Hollick, James F Collins, and M. L.
Fernald. Interestingly, in 1913 Fernald was the first to collect
Liatris scariosa var. novae-angliae on Block Island. “‘Could this
conspicuous species have been overlooked by the earlier bota-
nists?”’ Enser asked. Grazing by livestock and deer, he added, is
now widespread on the island and threatens this species and oth-
ers. Fenced exclosures are being used by conservationists to pro-
tect selected sites for the species. Some species such as Platanth-
era lacera, he suspects, are gone due to the heavy deer browsing
activity. Despite some losses, the number of species documented
for the island has increased to around 760, Enser said, but pri-
marily due to intensive exploration and the increase in the number
of nonindigenous species present. Block Island’s flora includes
most of the region’s most invasive taxa, and Enser estimated that
about 30% of its current flora is introduced. Populus alba, for
instance, was introduced about 1850 and now has dense colonies
established about the island.
Habitats on the island include extensive dunes; salt flats and
marshes; freshwater ponds; sedge meadows; shrub thickets; in-
land shrub communities dominated by Viburnum dentatum, Myr-
ica pensylvanica, and Amelanchier spp.; managed open grass-
lands used for grazing and hay fields; and a depression area with
a pocket of forest possessing Nyssa sylvatica and Fagus grandi-
folia. Also in this depression are the island’s only woodland wild-
flowers such as Maianthemum canadense, first found by Fernald,
and Anemone quinquefolia, discovered for the first time on the
island only a couple of years ago. Perhaps the most important
plant community on the island in Enser’s eyes are small areas of
open moraine with maritime grasses and herbs, located on slopes
and tops of knolls. Here, indigenous species prevail and rare plant
taxa for the island and state persist. In these areas one can find
124 Rhodora [Vol. 102
Liatris, Helianthemum dumosum, Aristida purpurascens, and
Chrysopsis mariana.
In closing remarks, Enser said that the future for natural veg-
etation on the island appears dim despite 25% of the land base
being in conservation management. Nonindigenous species, in-
cluding many invasive species, comprise more than 50% of the
island’s vegetative cover, and development pressures for addi-
tional housing and recreation are on the rise.
October 1999 Field Trips. On Friday afternoon, preceding the
Club’s evening meeting, about 15—16 enthusiastic Club members
and friends explored the south shoreline of Worden Pond in South
Kingstown, Rhode Island. Rick Enser led the outing. The water
level was relatively low, so walking was mainly on mudflats
among beds of exposed wetland plants that are typically emergent
from standing water. The area was scoured for plants, both fa-
miliar and new. Dominant species included Juncus militaris, Eu-
thamia tenuifolia, and Gratiola aurea. Locally dense patches of
basal rosettes belonging to Sabatia kennedyana could be found
below the taller vegetation, but not a single flowering stem was
seen; it was suspected that this was due, in part, to two successive
high water years. Some of the more uncommon finds of the day
were occasional fruiting stems of Ludwigia sphaerocarpa, a plant
or two of Glyceria obtusa bearing dense panicles of long spike-
lets, and a few clumps of Rhynchospora macrostachya. Aquatic
finds included Elatine minima and Vallisneria americana, the lat-
ter possessing mature fruits attached to elongated scapes, coiled
as they do following anthesis.
On Saturday a small group of 8—10 Club members, led by Dr.
Keith Killingbeck of the University of Rhode Island and by Rick
Enser, toured Ell Pond in Hopkinton, Rhode Island. The site is
owned by The Nature Conservancy and is the only designated
National Natural Landmark in Rhode Island. The vertical relief
of the site surprised many as we scrambled upward along a trail
of granitic bedrock and boulders. Alongside the trail we examined
a mature, mixed hardwood/conifer forest where we found Cha-
maecyparis thyoides, Pinus strobus, Tsuga canadensis (some in-
fested with hemlock adelgid), Nyssa sylvatica, Pinus rigida,
Quercus coccinea, and Q. prinus: an uncommon mixture of spe-
cies characteristic of either wet, mesic, or dry, well-drained sites.
Another special feature of the site was the abundance of Rho-
2000] NEBC Meeting News 125
dodendron maximum in the understory, giving one the feeling of
being in the Southern Appalachians. The group could not resist
a quick foray to the edge of Ell Pond where some typical bog
species such as Rhynchospora alba, Vaccinium macrocarpon, V.
oxycoccos, and Sarracenia purpurea were seen on a narrow
Sphagnum mat that edged the pond.
—PAUL SOMERS, Recording Secretary.
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THE NEW ENGLAND BOTANICAL CLUB
Elected Officers and Council Members for 1999—2000:
President: David S. Conant, Department of Natural Sciences,
Lyndon State College, Lyndonville, VT 05851
Vice-President (and Program Chair): Lisa A. Standley, Vanasse
Hangen Brustlin, Inc., 101 Walnut St., P.O. Box 9151, Wa-
tertown, MA 02272
Corresponding Secretary: Nancy M. Eyster-Smith, Department
of Natural Sciences, Bentley College, Waltham, MA 02154-
Treasurer: Harold G. Brotzman, Box 9092, Department of Bi-
ology, Massachusetts College of Liberal Arts, North Adams,
MA 01247-4100
Recording Secretary: Paul Somers
Curator of Vascular Plants: Raymond Angelo
Assistant Curator of Vascular Plants: Pamela B. Weatherbee
Curator of Nonvascular Plants: Anna M. Reid
Librarian: Leslie J. Mehrhoff
Councillors: W. Donald Hudson, Jr. (Past President)
Arthur V. Gilman 2000
Karen B. Searcy 2001
David Lovejoy 2002
Jennifer Forman (Graduate Student Member) 2000
Appointed Councillors:
David E. Boufford, Associate Curator
Janet R. Sullivan, Editor-in-Chief, Rhodora
RHODORA
Journal of the
New England Botanical Club
CONTENTS
Five new species of Verbesina from the northern Andes (Heliantheae;
Asteraceae). Harold Robinson and José Panero 2.0.00... cee 129
Do i aes Bstony traits relate to seed maturation in a clonal herb?
ichael T. Ganger 142
Long-term vegetation dynamics of the lower strata of a western Massachu-
setts oxbow swamp forest. Marjorie M. Holland, C. John Burk, and
David McLain 154
Vascular flora of beaver wetlands in western Massachusetts. Robert T.
Me Master and Nancy D. McMaster LS
NEW ENGLAND NOTES
Rediscovery of Symphyotrichum anticostense in the United States. Arthur
Haines 198
Aegagropilous Desmarestia aculeata from New Hampshire. Arthur C.
Mathieson, Edward J. Hehre, and Clinton J. Dawes ...........006. 202
Two moresweeds in Maine Petensh: Aikaec ne eaten ans ts ee ee rcs 208
New records for Scirpus ancistrochaetus in New Hampshire. Joshua L.
Royte and John P. Lortie 10
NOTES
Sagina (Caryophyllaceae) in Illinois: An update. Gordon C. Tucker 214
ae on the Lentibulariaceae in Bolivia: A new genus record (Genlisea)
r the country, with two additional ae records in the genus
as icularia. Nur P. Ritter and Garrett E. Crow ........000 0.0 c cee 2G
BOOK REVIEWS
A Guide to the Algae of New England as Reported in the Literature from
1829-1984, Parts I and I] 225
Thoreau’s Country: Journey Through a Transformed Landscape ........ 227
NEBC MEETING NEWS 230
Information for Contributors 237
NEBC Officers and Council Members inside back cover
Vol. 102 Spring, 2000 No.
Issued: June 6, 2000
910
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
ANTOINETTE P. HARTGERINK, Managing Editor
Department of Plant aerial University of New Hampshire,
Durham, NH 03824
Associate Editors
HAROLD G. BROTZMAN STEVEN R. HILL
DAVID S. CONANT THOMAS D. LEE
GARRETT E. CROW THOMAS MIONE
kK. 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 paya-
ble at par in United States currency. Remittances payable to RHO-
DORA. Send to RHODORA, P.O. Box 1897, Lawrence, KS 66044-
8897.
MEMBERSHIPS: — $35; Family $45; Student $25. Application
form printed here
NEBC WEB SITE: Information about The New England Botanical Club,
its history, officers and councillors, herbarium, monthly meetings and
special events, annual graduate student award, and the journal RHO-
DORA is available at http://www.herbaria.harvard.edu/nebc/
BACK ISSUES: Questions on availability of back issues should be ad-
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DORA, changes must be received by the business office prior to the
first day of January, April, July, or October.
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
RHODORA, Vol. 102, No. 910, pp. 129-141, 2000
FIVE NEW SPECIES OF VERBESINA FROM THE
NORTHERN ANDES (HELIANTHEAE; ASTERACEAE)
HAROLD ROBINSON
Department of Botany, National Museum of Natural History,
Smithsonian Institution, Masai ngton, DC 20560-0166
e-mail: robinson.harold @nmnh.si.edu
JOSE PANERO
Department of ae The University of Texas, Ci TX 78713
mail: panero @uts.cc.utexas.ec
ABSTRACT. Five new species of Verbesina are described, V. biserrata, V.
clarkae, V. maldonadoensis, and V. pichinchensis all from Ecuador, and V.
perijaensis from Colombia.
Key Words: Verbesina, new species, Heliantheae, Asteraceae, Ecuador, Co-
lombia, Northern Andes
Studies in the Heliantheae of the northern Andes, mostly for
the Flora of Ecuador, have resulted in the discovery of the fol-
lowing five undescribed species of Verbesina. The species are in
addition to the over 200 species already known in the New World
genus Verbesina (Olsen 1985). The four Ecuadorian species are
in addition to the 18 listed for Ecuador in Jorgensen and Leén-
Yanez (1999). The Colombian species is in addition to the ca. 25
already known from that country (including those of Diaz-Pied-
rahita 1985). A number of the following species are distinguished
by rays being white, yellow, or lacking, and they would poten-
tially fall within the artificial sections Leucactinia, Ochractinia,
or Lipactinia, but these sections are not recognized here. For a
partial discussion of some of past artificial segregates in Verbes-
ina see Blake (1925). Most of the true diversity of the genus
seems to be centered in Mexico.
Verbesina biserrata H. Rob. & Panero, sp. nov. TYPE: ECUADOR.
Prov. Cotopaxi: approximately 10.9 km W of Pilalé along
road to Latacunga, 3200 m, 15 Jul 1992, J. Panero & B. L
Clark 3000 (HOLOTYPE: TEX; ISOTYPE: QCA). Figure 1.
In habitus ad Verbesinam Iloensem superficialiter simila sed in
foliis oppositis valde biserratis et floribus radii nullis differt. In
129
130 Rhodora [Vol. 102
UNIVERSITY OF
HERBARIUM
PLANTS OF ECUADOR
—e
Verivsina Hoensis Hieron.
Vv 10.9 Kee Wood Plakd abeerg the
4 mm tall, cere pel hog rty ti:
ras suilfument with Dac aiming
hm! Lamers pod 18 be?
“penis firmer Herb aceen
‘igure 1. Verbesina biserrata H. Rob. & Panero, holotype, Herbarium of
the University of Texas (TEX)
foliis oppositis ad V. rivettii et V. pichinchensem proximius sed
in caulibus puberulis et marginis foliorum distincte biserratis et
foliis subtus sparce tomentellis et floribus radii omnino nullis et
2000] Robinson and Panero—Verbesina 131
floribus disci 20—25 et corollis sordide albis praeter in lobis ni-
grescentibus distincta.
Trees 3—6 m tall; stems brownish, terete, striate, without wings,
minutely brownish puberulous. Leaves opposite, with petioles 1—
8 cm long, upper leaves sometimes narrowly winged to base;
blades of lower leaves oblong-ovate, to 30 cm long, 14 cm wide,
with rounded base, upper leaves ovate-elliptical, 6-15 cm long,
3—6 cm wide, with base acute or acuminate, margins doubly ser-
rate, often serrulate or with minute mucronate dentations in upper
leaves, apex acute, upper surface sparsely pilose, lower surface
thinly tomentellous, pale yellowish, not obscuring green surface
except on immature leaves, denser and brownish on veins; veins
pinnate or slightly subpinnate, with 6—9 ascending veins on each
side, basal veins of larger leaves more spreading. Inflorescence
terminal on leafy branches, broadly corymbiform; primary
branches mostly opposite, ascending, densely puberulous with
sordid hairs. Heads 10-12 mm high, ca. 7-9 mm wide; involucral
bracts ca. 18, dark green to brown, narrowly oblong, flat, 4—8
mim long, 1.5—2.5 mm wide, rounded at tip, puberulous with pale
hairs outside; pales similar to inner involucral bracts, acute, ca.
8 mm long. Ray florets lacking. Disk florets 20—25; corollas sor-
did white, blackish on lobes, ca. 7 mm long, pilosulous on tube,
with few hairs on lobes, tubes ca. 1.8 mm long, throat ca. 4.5
mm long, lobes ca. | mm long, with fringe of long papillae on
inner margins; anther thecae black, ca. 2.5 mm long; apical ap-
pendage black, ca. 0.6 mm long, 0.35 mm wide. Cypselas im-
mature, ca. 4 mm long, glabrous on sides, few small setulae along
upper margins, with wings not expanded; awns of pappus straight,
ca. 6 mm long.
Verbesina biserrata is known only from the type collection.
The plant was found in Cotopaxi at 3200 m in disturbed Andean
forest. The species superficially resembles V. //oensis Hieron., but
differs by the opposite, strongly serrate leaves and lack of ray
florets. The species is probably more closely related to other pri-
marily opposite-leaved species such as V. rivettii S. FE Blake to
the north and V. pichinchensis to the south, both of which have
dense tomentum on the leaf undersurfaces and have some ray
florets.
Verbesina clarkae H. Rob. & Panero, sp. nov. TYPE: ECUADOR.
Prov. Carchi: 10.4 km E of Julio Andrade along road to El
132 Rhodora [Vol. 102
Carmelo and El! Aljun, around town of El Aljtin, 2880 m, 21
Jul 1992, J. Panero & B. L. Clark 3040 (HOLOTYPE: US; ISO-
TYPES: QCA, TEX). Figure 2.
In radiis albis brevibus et caulibus puberulis vel glabrescenti-
bus ad Verbesinam maldonadoensem simila sed in capitulis ma-
joribus in floribus radiis ca. 8 et in bracteis involucri ca. 22 dif-
tert:
Shrubs or small trees 4—10 m tall, moderately branched; stems
terete, unwinged, brownish, puberulous, glabrescent. Leaves al-
ternate, petioles distinct, unwinged, 3.5—9.0 cm long; blades
ovate, 18-28 cm long, 10—15 cm wide, base obtuse with short
acumination, margins remotely and minutely denticulate, apex
broadly acuminate, upper surface minutely scabridulous, lower
surface densely pale hirtellous, denser and more brownish on
veins; venation pinnate, with 8—10 veins on each side, lowest
secondary veins narrower, closer, spreading at ca. 90°. Inflores-
cence from apices and upper axils of leafy branches, broadly cor-
ymbiform cymes with many heads, ca. 30 cm wide and high;
peduncles 3—18 mm long, densely brownish hirtellous with broad-
sharp hairs. Heads campanulate, 10—12 mm high; involucre 7—8
mm wide; involucral bracts ca. 22 in ca. 3 graduated series, nar-
rowly oblong, 2—7 mm long, |.0—1.3 mm wide, apices obtuse,
densely puberulous with pale hairs outside; pales similar to inner
involucral bracts, ca. 8 mm long. Ray florets ca. 8; corollas white,
sometimes with tinge of yellow, tube and tip of limb pilosulous,
tube 1.3 mm long, limb ca. 3.5 mm long, 1.3 mm wide. Disk
florets ca. 20; corollas whitish yellow, 6.0—6.8 mm long, pilo-
sulous throughout, sparsest on middle of throat, tube ca. 1.8 mm
long, throat 3.8-4.0 mm long, lobes ca. | mm long, with dense
fringe of long papillae on inner margin; anther thecae black, 2.5
mm long; apical appendage black, 0.5 mm long, ca. 0.3 mm wide.
Cypselas immature; ray cypselas ca. 3 mm long, with single outer
awn ca. 3 mm long; disk cypselas ca. 3.8 mm long, essentially
glabrous, wings not expanded, pair of pappus awns ca. 6 mm
long.
The new species is named for the co-collector on the 1992
expedition, Bonnie L. Clark, previously a student of Ted Barkley
at Kansas State University, more recently of Shawnee Mission,
Kansas.
Verbesina clarkae is known only from the type locality. The
2000] Robinson and Panero—Verbesina 133
u }
Ooi Ype
a el arnKet
a? nero
PLANT F ECUADOR
Verbesina
ARCHIE: 10.4 km E of Julio Andrade along the road to El
c ‘armelo and El Alin around the town of El Aljin. Shrubs or
mall trees 4-10 y corollas few and white sometimes
seitha ty anythat isk corollas whitish yellow, herbage
covered with aban nish pubescence. Growing in disturbed
Andean forest
2880 m
Occasional
UNITED STATES
3363405
Panero 3040 Jul 21 1992
aah ee Clark
NATIONAL HERBARIUM
Michigan State University Herbarium
Hiokfeurk support! by NSP grant DEBS H>
ure 2. Verbesina clarkae H. Rob. & Panero, holotype, United States
National Herbarium (Us)
134 Rhodora [Vol. 102
species occurs in easternmost Carchi at 2880 m in disturbed An-
dean forest. It is one of the few species in Ecuador in which
white rays are present. The closest relative is probably V. mal-
donadoensis of eastern Carchi, but the present species has larger
heads with more numerous florets, ca. 8 ray florets and ca. 20
disk florets. The rays have small limbs, and their number is dis-
tinctive within Ecuador. A possibly closely related Colombian
species from Dept. Valle is V. barragana Cuatrec., but the latter
has lanuginose pubescence on the stems, more lanceolate leaves,
and five or fewer ray florets with limbs 7 mm long.
Verbesina maldonadoensis H. Rob. & Panero, sp. nov. TYPE:
ECUADOR. Prov. Carchi: 9 km E of Maldonado, ca. 55 km W
of Tufino along road Maldonado—Tufino—Tulcan, 2100 m,
small trees to 7 m tall, corollas whitish, growing in disturbed
Andean forest, 20 Jul 1992, J. Panero & B. L. Clark 3035
(HOLOTYPE: US; ISOTYPES: QCA, TEX). Figure 3.
In radiis albis brevibus et caulibus puberulis vel glabrescenti-
bus ad Verbesinam clarkae simila sed in capitulis minoribus in
floribus radiis ca. 3 et bracteis involucri ca. 14 distincta.
Shrubs to 7 m tall, moderately branched, with brownish hairs;
stems brown, terete to slightly angled, without wings, slightly
deflected at nodes, minutely puberulous or with sparse arachnoid
hairs; internodes mostly 2—3 cm long. Leaves alternate, petioles
2-8 cm long, 1/5 to more than half as long as blade, unwinged;
blades elliptical, 10-17 cm long, 3-8 cm wide, base acute to
slightly acuminate, margins minutely mucronate-serrulate except
near base, apex short and narrowly acuminate, upper surface dark,
minutely pilosulous, densely hirtellous on veins, lower surface
densely pilosulous, subtomentellous on veins; secondary veins
pinnate, ca. 8 on each side, spreading at ca. 55°, arched before
margins. Inflorescence from apices and upper axils of leafy
branches, broadly flat-corymbiform, with many heads, ca. 25 cm
high and wide; peduncles 1-12 mm long, densely hirtellous.
Heads narrowly campanulate, S—6 mm high; involucre 3—4 mm
wide; involucral bracts ca. 14, in ca. 3 graduated series, narrowly
oblong, 1.5—4.0 mm long, 0.5—1.2 mm wide, apices acute, thinly
herbaceous, pilosulous outside; pales greenish on keel and dis-
tally, elliptical-lanceolate, 5.0—5.5 mm long, ca. | mm wide, pi-
losulous distally on keel. Ray florets ca. 3; corollas white, tube
ca. 1.5 mm long, pilosulous, limb oblong, ca. 4 mm long, 1.5—
2000] Robinson and Panero—Verbesina [25
| O& bee hd
| | ice
B of Maldonado, ca. $$ km W of Tutito along
H wlohe
CARCHL 9 ide
the read Maldonado-Tediho-Tulcin, Senall trees to 7 m tall,
ot ent attacked by insects. Growing in
tiga
UNITED STATES
José L. Pamero 3005 ful 2301092
with Bonnie Click
3363420
Michigan State Lint errelny Pevbartem
MATIONAL HERBARIUM
Figure 3. Verbesina rae H. Rob. & Panero, holotype, United
States National Herbarium
136 Rhodora [Vol. 102
2.0 mm wide. Disk florets 8-12; corollas yellowish, ca. 4.5 mm
long, pilosulous on tube and lobes, tube ca. | mm long, throat
ca. 2.5 mm long, lobes ovate-oblong, ca. 0.8 mm long, with dense
fringe of long papillae on inner margin; anther thecae black, ca.
1.8 mm long; apical appendage yellowish distally, ca. 0.35 mm
long, ca. 0.2 mm wide. Cypsela immature, ca. 2 mm long, with
wings to 1.5 mm wide, glabrous; awns of pappus straight, 2.0—
3.5 mm long.
PARATYPE: Ecuador. Prov. CARCHI: above Maldonado, in secondary cloud
forest and pastures, 2500 m, | Aug 1989, van der Werff & Gudifio 10852
(US).
Verbesina maldonadoensis is known only from secondary
cloud forests at 2100—2500 m elevation in the western part of the
Province of Carchi near the Colombian border. The species prob-
ably also occurs in the immediately adjacent areas of Colombia.
The rather small heads with white ray florets, alternate leaves,
and the sometimes very long petioles distinguish this species.
Verbesina pichinchensis H. Rob., sp. nov. TYPE: ECUADOR. Prov.
Pichincha: Paramo and shrub vegetation on eastern slopes of
Cerro Pichincha, 3600—4000 m, 26 Jan 1977, G. Harling, U.
Eliasson, & L. Andersson 14832 (HOLOTYPE: US; ISOTYPE: GB).
Figure 4.
Ad Verbesinam ecuatorianam simila sed in foliis suboppositis
in foliis plus ovatis et in ramis inflorescentium densius tomentosis
distincta.
Shrubs to 2 m tall, moderately branching; stems pale brownish,
terete, unwinged, densely lanulose with pale slender hairs, inter-
nodes between pairs of leaves 3—5 cm long, internodes between
leaves of pairs ca. 0.5—1.0 cm long. Leaves subopposite, petioles
1—2 cm long; blade narrowly ovate, 8-14 cm long, 2.5—4.5 cm
wide, base short-acute, slightly acuminate on petiole, margins
closely serrulate from below widest part, apex acute, upper sur-
face dark green, densely scabridulous with sharp hairs of many
sizes, lower surface paler, almost completely covered with bases
of hairs, subtomentose to sublanulate with long and short hairs,
main secondary veins 5 or 6 on each side, ascending at 25—40°,
with a few smaller closely spreading secondaries nearer base.
Inflorescence terminal and from upper axils on leafy stems,
broadly corymbiform with many heads, to 13 cm wide, with
2000] Robinson and Panero—Verbesina 137
"t rh ers
| =; | i 1
%
Holot :
\ type
Verdesina pichinchens ts
|
H Rb s lanero
UNITED STATES NATIONAL HERBARIUM L
tr bteina <t+a fevieeree_ dager ae
pa Jeu A Farce (99
FLORA OF ECUADOR
Verbesina arberea HBk
UNITED STATES
2851819
NATIONAL HERGARIUM sah G, HARLING, U. ELL
Figure 4. Verbesina pichinchensis H. Rob., holotype, United States Na-
tional Herbarium (Us).
138 Rhodora [Vol. 102
branches ascending, tomentose; peduncles 0-12 mm, covered
with dense pale tomentum. Heads campanulate, 9-11 mm high,
7-8 mm wide; involucral bracts ca. 15, in 2—3 series, oblong to
oblong-lanceolate, 4-6 mm long, 1.5—2.0 mm wide, apices ob-
tuse, densely puberulous to sublanulate outside and near tip in-
side, inner bracts acute, densely puberulous outside; pales similar
to inner involucral bracts, to 8 mm long. Ray florets 0—3; corolla
fertile, yellow, tube ca. 2.5 mm long, pilosulous, limb oblong, ca.
5 mm long, 2.5 mm wide, sparsely puberulous. Disk florets ca.
15; corolla yellow, 4.5—5.0 mm long, tube ca. 1.5 mm long, pi-
losulous, limb sparsely puberulous, throat ca. 3.5 mm long, lobes
0.5 mm long and wide, with dense fringe of long papillae on
inner margins; anther thecae black, ca. 2 mm long; apical ap-
pendage mostly yellowish, partially black. Cypsela (immature) 6
mm long, ca. 1.5 mm wide, with few slender hairs on sides, wings
present; pappus with 2 awns, straight, 4.5-5.0 mm long.
PARATYPE: Ecuador. Prov. PICHINCHA: Pichincha, 30 Oct 1930, R. Benoist
sn. (P, US).
Verbesina pichinchensis is known from paramo and shrub veg-
etation at 3600-4000 m on Mt. Pichincha in north-central Ec-
uador. The specimens were determined at one time as V. ecuato-
riana Sagast. (Sagastegui-Alva 1970), to which they seem closely
related, but the leaves are subopposite with long internodes be-
tween the pairs, the leaf blades are more ovate, and the branches
of the inflorescence have denser and longer tomentum. The habit
is also similar to the common V. sodiroi Hieron., but the latter
has 10-15 ray florets in a head.
Verbesina perijaensis H. Rob. sp. nov. Type: COLOMBIA. Dept.
esar: ““Magdalena’’, Sierra de Perija, east of Manaure, Que-
brada de Floridablanea, Andean forest and bushes, 2700—
2800 m, 11 Nov 1959, J. Cuatrecasas & R. Castaneda 25236
(HOLOTYPE: US). Figure 5.
In caulibus et foliis glabris ad Verbesinam laevifoliam et V.
negrensis et V. simulans simila sed in bracteis involucri exterior-
ibus 6—8 erectis obovatis et floribus radii ca. 14 in corollis longis
et albis vel lilacinis differt.
Trees 2—3 m tall, with pendulous branchlets; stems brownish,
glabrous, weakly angled. Leaves alternate, petioles 5-7 mm long;
in
ce
ches
peas
ers,
|
139
Robinson and Panero—Verbesina
hula
Holotype
loeb Wes
Verbesina per jaensis 7 /
COLOMBIA
“abit colombianae
TRECASAS
on
9 2593 j eee
25236- Tree ooo z with pendulous Caer eps
ari
Leaves subcoriaceous, shining, hit Masdelena: Siera de Petit
iguies white t, Perish. astern of Manaure: Quebrada d
green apres Involucte green. s ean the ecoming Floridablance, forest and bushes, 7700-2800 m. alt,
ceous white, wi wl 523 Geanfhs SUATRECASAS
R, ROMERG CASTANEDA Pedy | Fe, 1959
or Lila
lilac).
Verbesina perijaensis H. Rob., holotype, United States National
Figure 5.
Herbarium (Us).
140 Rhodora [Vol. 102
blades subcoriaceous, oblanceolate, 6-9 cm long, 2—3 cm wide,
base slightly acuminate, margins serrulate, apex acute, surfaces
glabrous, shining, dark yellowish green above, veinlets forming
dark reticulum below; venation pinnate, 6—8 upwardly curving
secondary veins on each side. Inflorescence from apex and upper
axils of leafy branches, laxly corymbiform with few large heads;
branches glabrous at base, branchlets sparsely hirsute on one or
more sides; peduncles 1—4 cm long. Heads broadly campanulate,
1.2—1.5 cm high, 2—3 cm wide; involucre of 6—8 spreading round-
ed herbaceous outer bracts, to 10 mm long, ca. 8 mm wide, most-
ly glabrous with some hairs near base and distal margins, apex
rounded to obtuse, ca. 8, more erect, darker, obovate, glabrous,
inner bracts with rounded thinly herbaceous often slightly re-
curved tips, 8-9 mm long, 5 mm wide above middle, with pale
margins below; pales narrowly oblong, ca. 9 mm long, paler base
clasping floret, distal 3 mm flat, darker, oblong, shortly acute.
Ray florets ca. 14 in a head; corollas white or lilac, tube ca. 2
mm long, hirsutulous, limb narrowly elliptical, ca. 19 mm long,
4 mm wide. Disk florets ca. 40; corollas ca. 6.8 mm long, tube
ca. 2 mm long, hirsutulous, throat ca. 3.5 mm long, lobes ca. 0.8
mm long, with fringes of long papillae on inner margins; anther
thecae ca. 2.8 mm long; apical appendage rather pale, ca. 0.45
mm long, ca. 0.35 mm wide. Cypsela ca. 5.5 mm long, glabrous,
wing immature; pappus awns 2—3 mm long.
Verbesina perijaensis is known only from the type, collected
in the Perija region along the Venezuelan border. It appears to be
related to some species in the Venezuelan Andes with glabrous
leaves such as V. /aevifolia S. F Blake and V. negrensis Steyerm.
of Venezuela (Aristeguieta 1964) which lack ray florets, and V.
simulans S. F Blake which has small white rays. The new species
differs from all of these by the large white rays and the very large
foliose involucral bracts.
ACKNOWLEDGMENTS. Field work of José Panero and Bonnie L.
Clark in Ecuador was funded by NSF Grant DEB 91-14798, and
it was in cooperation with the Pontificia Universidad Catolica del
Ecuador (QCA) and the Michigan State University Herbarium. The
type photographs were taken by John Steiner and Don Hurlbert
of the National Museum of Natural History Photographic Labo-
ratory.
2000] Robinson and Panero—Verbesina 141
LITERATURE CITED
ARISTEGUIETA, L. 1964. Compositae, Vol. 10. /n: T. Lasser, dir, Flora de
Venezuela. Instituto Botanico y Universidad Central de Venezuela, Cara-
ees S. E 1925. On the status of the genus Chaenoc ieee with a review
of the section Lipactinia of Verbesina. Am . Bot. 12: 625-
DIAZ-PIEDRAHITA, S. 1985. Aported a la Flora - Onan estudios en Cone
estas—V. Mutisia 61: 1-11.
in P. M. AND S. LEGN-YANEZ, eds. 1999. calogie of the Vascular
Plants of oe Monogr. Syst. Bot. Missouri Bot. Gard. 75: i—viii, 1—
OLSEN, J. 1985. Synopsis of Verbesina sect. Ochractinia (Asteraceae). Pl.
Syst. Evol. 149: 47-63.
SAGASTEGUI-ALVA, A. 1970. Tres Compuestas Austroamericanas nuevas o
criticas. Bol. Soc. Bot. La Libertad 2: 63-75.
RHODORA, Vol. 102, No. 910, pp. 142—153, 2000
DO REPRODUCTIVE HISTORY TRAITS RELATE TO
SEED MATURATION IN A CLONAL HERB?
MICHAEL T. GANGER
Department of Plant Biology, Rudman Hall,
University of New Hampshire, Durham, NH 03824
Current Address: Division of Math and Science, ae College,
147 Sycamore St., Pikeville, KY 415
e-mail: mganger@pc.edu
ABSTRACT. A field experiment tested whether seed maturation in Canada
mayflower (Maianthemum canadense) was pollen limited and whether ramets
flowering for the first time differed from ramets flowering for the second time
in the number of seeds that they matured. Addition of pollen increased the
number of seeds matured by ramets and therefore seed maturation was pollen
limited. No difference in the number of seeds matured by ramets flowering
for the first time and ramets flowering for the second time was detected and
thus previous flowering did not appear to affect current seed maturation.
There was a positive relationship between the number of seeds matured b
ramets and the total weight of these seeds. This relationship did not differ
etween ramets flowering for the first and second times
Key Words: ramet, genet, Canada mayflower, Maianthemum canadense
Plants must allocate limited resources to structures associated
with growth, defense, and reproduction (Bazzaz et al. 1987). Over
a plant’s lifetime, allocation patterns must balance survivorship
and fecundity (Bazzaz et al. 1¢
A tradeoff between current fecundity and subsequent growth
and fecundity has been reported for the orchids Cypripedium
acaule (Primack and Hall 1990), Epidendrum ciliare (Ackerman
and Montalvo 1990), and Tipularia discolor (Snow and Whigham
1989). Current fecundity also reduced growth in mayapple (Podo-
phyllum peltatum; Sohn and Policansky 1977, although see Ben-
ner and Watson 1989) and reduced the probability of producing
females in jack-in-the-pulpit (Arisaema triphyllum, Bierzychudek
1984)
Current fecundity in self-incompatible, insect-pollinated plants
may be influenced by a number of variables including the avail-
ability of compatible pollen (Bierzychudek 1981; Thompson and
Stewart 1981) and also by a plant’s reproductive history. This is
not to suggest that a plant shown to be pollen limited in a given
142
2000] Ganger—Seed Maturation 143
year is not ultimately resource limited (Bierzychudek 1981) as
the addition of pollen in one year may increase current fecundity
at the expense of future fecundity (Janzen et al. 1980).
Canada mayflower (Maianthemum canadense Desf.; Gleason
and Cronquist 1991) is a rhizomatous, perennial herb with genets
that consist of dimorphic ramets. Flowering ramets have 2-3
leaves with a terminal inflorescence consisting of 4-35 perfect
flowers, while vegetative ramets have only one leaf. Fruits con-
tain 1-4 seeds. Canada mayflower is self-incompatible (Worthen
and Stiles 1986).
In the fall, ramets “‘die back,” leaving an overwintering bud
that will become the ramet in the following year. At this time it
is possible to determine “‘by touch”? whether a bud is vegetative
or flowering (Ganger 1998). The status of this bud appears to be
determined much earlier, and flower primordia are visible under
the microscope as early as May (Kana 1982). With the dieback
of the ramets, an abscission scar is left. The scars left by vege-
tative and flowering ramets are distinctive so it is possible to
determine the age of a ramet and whether this ramet has been
vegetative or flowering for each of its previous years (Ganger
1997; Silva et al. 1982). It is possible to make this determination
in the field with a hand lens, but not as reliably as with a dis-
secting scope in the lab. It is sometimes possible to identify a
ramet that has flowered previously by the presence of a dead
flowering stalk.
The flowering ramets of Canada mayflower vary with respect
to the number of times they have flowered previously and the
number of years spent vegetative prior to flowering or since flow-
ering (Ganger 1997). It is not known whether ramets that remain
vegetative longer mature more seeds. It is also not known whether
ramets experience a cost of previous flowering. If there were such
a cost, then ramets flowering for the second time may mature
fewer seeds than ramets flowering for the first time. This cost
may also be reflected in the resources allocated to these seeds.
Specifically, previously flowering ramets may allocate fewer re-
sources to a similar number of seeds than other ramets.
A field experiment was undertaken to address the following
questions: (1) Do ramets that remain vegetative for a longer pe-
riod of time mature more seeds? (2) Is there evidence of a cost
of flowering such that ramets flowering for the first time mature
more seeds than ramets flowering for the second time? (3) Is
144 Rhodora [Vol. 102
Canada mayflower pollen limited in local populations? (4) Is
there a relationship between the number of seeds matured by
ramets and the total weight of these seeds? and (5) Does this
relationship differ between ramets flowering for the first and sec-
ond time?
MATERIALS AND METHODS
Field work was conducted in a mixed coniferous-hardwood
forest in the University of New Hampshire woodlands, Durham,
NH. Sixty ramets that were flowering for the first time were iden-
tified as well as 60 ramets that were flowering for the second
time. Ramets flowering for the first and second time occurred
together, often within millimeters of one another. These two
‘‘flowering”’ treatments represented one of the factors in a two-
factor experiment. Seed maturation in Canada mayflower has
been demonstrated to be pollen limited in New Jersey populations
(Worthen and Stiles 1988) and in previous years in the University
of New Hampshire woodlands (Ganger 1997). Therefore, a sec-
ond factor, “‘pollination,’ was included. Half of the flowering
ramets in each of these treatments were randomly assigned to
either of two treatments: hand pollinated or open pollinated. In-
cluding this factor reduced the likelihood that differences in seed
maturation between first- and second-time flowering ramets
would be obscured by low levels of pollination.
Each of the flowers of the hand-pollinated ramets were polli-
nated each day for the life of the flowers. Pollen was collected
immediately prior to use from 10—20 flowering ramets not more
than 20 m away and applied to stigmas with a wooden toothpick.
Flowers from the open-pollinated ramets were not manipulated
and presumably received natural levels of pollination. At the end
of the fruiting season, all of the flowering ramets were excavated
and taken to the laboratory. The numbers of fruits and seeds
matured by each ramet was noted. Seeds were removed by gently
pushing on the fruits. Seeds were dried for 72 hours at 80°C in
a drying oven and then weighed to the nearest | < 10° g using
a Mettler AE 63 balance. The total weight of all seeds matured
by ramets was calculated. For each of the ramets that flowered
for the first time, the number of years prior to flowering was
noted. For each of the ramets that flowered for the second time,
the number of years since flowering was noted.
2000] Ganger—Seed Maturation 145
In order to determine if ramets that remained vegetative for a
longer period of time were able to mature more seeds, two re-
gressions were performed. For hand-pollinated, first-time flow-
ering ramets, the number of seeds they matured was regressed on
the number of years prior to flowering. For hand-pollinated, sec-
ond-time flowering ramets, the number of seeds that they matured
was regressed on the number of years since flowering.
If ramets that were flowering for the first and second time
differed in the amount of resources that were allocated to seed
maturation, then this difference may have been apparent either as
a difference in the number of seeds matured or as a difference
between the two types of ramets, in the resources allocated to a
similar number of seeds.
In order to test whether the number of seeds matured by ramets
flowering for the first time differed from ramets flowering for the
second time and whether seed maturation in ramets overall was
pollen limited, a two-way analysis of variance (ANOVA) was
performed. The statistical model consisted of two factors: flow-
ering (first-time flowering vs. second-time flowering ramets) and
pollination (hand pollinated vs. open pollinated). Both flowering
and pollination were fixed, categorical variables. The dependent
variable was the number of seeds matured. If there was a differ-
ence in the number of seeds matured between ramets flowering
for the first and second time, then this effect would more likely
be observed in ramets that received an overabundance of pollen.
In other words, if resource limitation was occurring to a greater
extent in either the first- or second-time flowering ramets, then
the difference in seed maturation would more likely be detected
statistically if pollen-limited ramets were not included. Therefore
one a priori contrast was considered: the number of seeds ma-
tured by hand-pollinated, first-time flowering ramets was com-
pared to the number of seeds matured by hand-pollinated, second-
time flowering ramets.
In order to determine if ramets that were flowering for the first
and second time differed in their allocation of resources to seeds,
two analyses of covariance (ANCOVA) were performed. AN-
COVA allowed for a determination of whether there was a rela-
tionship between the number of seeds matured by ramets and the
total weight of these seeds, and whether this relationship varied
between first- and second-time flowering ramets. Two separate
OVAs were performed, one for open-pollinated ramets and
146 Rhodora [Vol. 102
another for hand-pollinated ramets. In order to determine what
variables influenced seed maturation, the number of seeds was
the dependent variable. The number of times flowering was the
independent, categorical variable and the total weight of seeds
per ramet was the covariate. In these analyses, only those ramets
that matured seeds were considered.
RESULTS
Forty-six of 60 open-pollinated ramets matured one or more
seeds while 51 of 60 hand-pollinated ramets matured one or more
seeds. Six ramets were excluded from the analyses because they
were found to have flowered more than twice, they could not be
accurately aged, or their reproductive history could not be deter-
mined due to decay.
The average number of years prior to flowering for ramets
flowering for the first time was 3.9 years (SD = 1.81) and thus
the average age of these ramets was 4.9 years (SD = 1.81). For
ramets flowering for the second time, the average number of years
since flowering was 2.9 years (SD = 0.75) and the average age
of these ramets was 8.4 years (SD = 1.87). The number of seeds
matured by hand-pollinated ramets was not correlated with the
number of years prior to flowering for first-time flowering ramets
(Fi 50005 = 0.53, p = 0.47) nor with the number of years since
flowering for second-time flowering ramets (F, 5.995 = 2.34, p =
Ramets that were hand pollinated matured more seeds than
ramets that were open pollinated (F, ,9;99; = 127.22, p < 0.005,
r’ = 0.11; Figure 1). There was no difference in the number of
seeds matured by ramets flowering for the first and second time
(F 1070.05 = 0.50, p = 0.48; Figure 1). There was no interaction
between the pollination and flowering factors (F, ,o799; = 0.90, p
= 0.35). The a priori contrast was also not significant; there was
no difference between the number of seeds matured by hand-
pollinated, first-time flowering ramets and hand-pollinated, sec-
ond-time flowering ramets (F, j¢90; = 0.63, p = 0.43; Figure 1).
In concluding that there was no difference in the number of seeds
matured between these two treatments, there is an associated
probability of being wrong. This is Type II error or 8. Following
Winer et al. (1991) it was possible to determine 8 and therefore
the statistical power (1 — 8) of this comparison, given specific
2000] Ganger—Seed Maturation 147
127
10 =)
T
8 =
5
T
Number of seeds matured per ramet (+/— 1 SD)
op
!
First Second First Second
Open Open Hand Hand
Figure 1. Mean number of seeds matured by Matanthemum canadense
ramets in each of the four experimental treatments. First = first-time flow-
ering ramets; Second = second-time flowering ramets; Open = open-polli-
nated ramets; and Hand = hand-pollinated ramets.
alternative hypotheses. The statistical power of the test, assuming
a true difference between the treatment means of |, 2, and 3
seeds, was determined to be 0.21, 0.51, and 0.80, respectively.
Seeds varied in weight from 3 to 18 mg. For open-pollinated,
first- and second-time flowering ramets, there was no difference
in the relationship between the number of seeds matured and the
total weight of these seeds with respect to either the slopes of
these relationships (F, 4.99; = 1.37, p = 0.25; Figure 2A) or the
y-intercepts (F, 4,995 = 1.89, p = 0.17; Figure 2A). There was a
positive relationship between the number of seeds matured and
148 Rhodora [Vol. 102
A
Open
e First
o Second
Number of seeds matured per ramet.
U T T T T T 1
0.00 0.02 0.04 0.06 0.08 0.10 0.12
Total weight of seeds per ramet (g)
Hand
e First
o Second
Number of seeds matured per ramet
S
0.00 0.02 0.04 0.06 0.08 0.10 0.12
Total weight of seeds per ramet (g)
ure 2. Plots of oad number of seeds matured by Maianthemum cana-
dense ramets and the total weight of these seeds for (A) o — i ena
ramets and (B) hand- Se ramets. First = first-time flowering ramets;
econd = second-time flowering ramets; Open = open- ae ramets; and
Hand = hand-pollinated ramets.
2000] Ganger—Seed Maturation 149
the total weight of these seeds (F, 49995 = 176.25, p < 0.001, r?
= 0.83). For hand-pollinated, first- and second-time dowerng
ramets, there was no difference in the relationship between the
number of seeds matured and the total weight of these seeds with
respect to the slopes (F, 4,995 = 2.57, p = 0.12; Figure 2B) or
the y-intercepts (F, 45995 = 0.55, p = 0.46; Figure 2B). There was
a relationship between the number of seeds matured and the total
weight of these seeds (F, 4; 99; = 290.78, p < 0.001, r? = 0.90).
Seed number was used as the dependent variable in both the
ANOVA and the ANCOVAs in order to focus on what factors
influenced seed maturation in Canada mayflower. It would also
be valid in the ANCOVAs to use seed number as the independent
variable and the total weight of seeds per ramet as the dependent
variable. This, however, would change the focus from what fac-
tors influenced seed maturation to what influence did seed number
have on seed weights. The results of such analyses did not differ
from those presented above and this is likely due to an overall
correlation between seed number and the total weight of seeds (p
< 0.001, r? = 0.92).
DISCUSSION
The number of years prior to flowering for first-time flowering
ramets and the number of years since flowering for second-time
flowering ramets were not related to the number of seeds matured
by ramets. Therefore ramets that waited longer to flower did not
necessarily mature more seeds. Watson (1984, 1990) has referred
to ramets as “‘mouths’’ which accumulate resources for use by
the genet and for the ramet itself in the future. It is possible that
with Canada mayflower, a vegetative ramet continues to accu-
mulate resources until a specific threshold is reached and that this
triggers the transition to a flowering ramet. The number of years
since flowering may then be related to two variables: (1) the cost
incurred by the ramet for past flowering and seed maturation (this
may be influenced by the degree of ramet integration) and (2) the
quality of the habitat. In other words, a ramet that waited a greater
number of years to flower again may have incurred a great cost
of previous reproduction, one that the genet was not able to offset,
or the quality of the habitat was such that resources were able to
be replenished only very slowly, or even some combination of
the two. It may also be that another part of the genet was flow-
150 Rhodora [Vol. 102
ering during the intervening years between flowering and that the
ramet in question was subsidizing the seed maturation of another
part of the genet.
Ramets matured comparable numbers of seeds regardless of
whether this was their first- or second-time flowering. Thus the
act of flowering did not appear to influence the success of future
seed maturation. This was true as well for ramets that received
an overabundance of pollen. For this contrast, statistical power
was low for a hypothesized difference of one seed. However, at
a hypothesized difference of three seeds (approximately the dif-
ference between the pollination treatment means) power was quite
high.
The addition of pollen resulted in an increase of, on average,
2.7 seeds per ramet. This suggests that seed maturation in ramets
was pollen limited during the experiment. Moreover, the addition
of pollen had a similar effect on ramets flowering for the first and
second time.
There was a strong relationship between the number of seeds
matured and the total weight of these seeds for both open- and
hand-pollinated ramets. Furthermore this relationship did not dif-
fer between ramets flowering for the first and second time. This
is further evidence that there was not a cost of having flowered
in the past.
While it was possible to determine if ramets had previously
flowered, nothing is known about the success of previous flow-
ering. It could be that seed maturation in the past was low and
therefore the costs incurred by ramets at that time was also low.
The cost of having flowered and reproduced in the past may also
have been mitigated by other ramets at the time of flowering.
Canada mayflower ramets are known to have increased seed mat-
uration if they receive ample pollen and their rhizomes are left
intact, but not if ramets receive ample pollen and their rhizomes
are severed (Ganger 1997). This suggests that resources may be
translocated to the flowering ramet, either from the rhizome or
from other ramets, and may offset the cost of reproduction in-
curred by the ramet. It is important to note that second-time flow-
ering ramets are much rarer than first-time flowering ramets and
that second-time flowering ramets are a subset of ramets that have
flowered before. Therefore these results should be viewed with
caution.
Canada mayflower is not the only example of a plant in which
2000] Ganger—Seed Maturation 15]
a cost of previous flowering was not detected. In fact some plants
show an increased likelihood of future flowering with greater cur-
rent reproductive allocation. In the orchid Spiranthes cernua, An-
tlfinger and Wendel (1997) found that individuals producing few-
er flowers were less likely to flower in the following year than
individuals producing more flowers. This was despite the fact that
individuals producing many flowers had a tendency to decrease
in size in the following year. In the early spider orchid Ophrys
sphegodes, Hutchings (1987) found that individuals flowering in
the current year were: (1) more likely to flower in the following
year than either vegetative or dormant individuals, (2) less likely
to enter dormancy than either vegetative or dormant individuals,
and (3) had greater survivorships than dormant individuals.
Other species show no effect of current flowering on future
reproduction. Horvitz and Schemske (1988) experimentally cre-
ated high and low levels of reproductive effort in Calathea ovan-
densis and found no difference in the growth, survival, and re-
production of individuals between treatments in the following
year. Smith and Young (1982) found that individuals of Senecio
keniodendron that had high levels of reproduction were more
likely to die than other individuals. However those that did sur-
vive had higher levels of reproduction than other individuals in
the following year.
The notion that previous reproduction should negatively affect
present reproduction seems plausible. Since in plants, size and
reproduction and size and survivorship tend to be positively cor-
related, allocation of resources to reproduction in the current year
may not only reduce growth in the current year, but may also
reduce future fecundity and survivorship due to decreased size
(Lovett Doust 1989; Stephenson 1981). This relationship in plants
such as Canada mayflower appears to be difficult to establish, in
part because of the ability of Canada mayflower to be vegetative
for a number of years between flowering and also due to Canada
mayflower’s clonal habit. These would allow Canada mayflower
ramets to mitigate the cost of previous and current reproduction.
It is possible then that the cost of previous reproduction, while
not apparent at the level of the ramet, would be evident at the
level of the genet.
ACKNOWLEDGMENTS. The author thanks Tom Lee for assis-
tance during the experiment and Tom Lee, Lenny Lord, and two
Rhodora [Vol. 102
Nn
N
anonymous reviewers for their helpful comments on the manu-
script. This work was supported in part by a Summer Teaching
Fellowship from the Graduate School at the University of New
Hampshire.
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RHODORA, Vol. 102, No. 910, pp. 154-174, 2000
LONG-TERM VEGETATION DYNAMICS OF THE LOWER
STRATA OF A WESTERN MASSACHUSETTS OXBOW
SWAMP FOREST
MARIJORIE M. HOLLAND
Department of Biology, University of Mississippi, University, MS 38677
e-mail: mholland@olemiss.edu
C. JOHN BURK
Department of Biological eee Smith College,
nei es
e-mail: cbu eee edu
Davip McLAIN
Massachusetts Audubon Society, Arcadia Wildlife Sanctuary,
127 Combs Rd., Easthampton, MA 01027
e-mail: DAVEMCLAIN @aol.com
ABSTRACT. The structure, composition, and floristics of understory swamp
d’s
forest vegetation in Ne itch, a segment of a regularly flooded oxbow in
Northampton, Massachusetts, has been investigated at intervals fro
through 1996. The forest canopy is dominated by Acer ee nie in as-
sociation with Quercus palustris and Fraxinus pennsylvanica; these sae
are regenerating despite the deaths of a number of trees between 1975 a
1985. The shrub stratum, dominated by Cephalanthus occidentalis, has re-
mained relatively eanemas In the herbaceous stratum, species abundances
fluctuate from year to year in relation to flooding and pine aspects of hy-
drology with ae such as Lemna minor panini during wet years and
annuals, particularly Bidens spp., growing to maturity in times of drought.
The abundance of Osmunda regalis and one sensibilis has remained rel-
atively constant over the 23 year period, but tree seedlings have become
increasingly important though few have been recruited to the upper strata.
verall, emergent and floating hydrophytes in the herb stratum have tended
to decline although the composition of the flora of the herb stratum continues
to strongly resemble the flora of adjacent marshes. These observations suggest
that preserving and successfully managing Ned’s Ditch and similar floodplain
forests will require the maintenance of species of diverse ecological require-
ments adapted to a range of habitat conditions.
Key Words: swamp forest, floodplain vegetation, herbaceous flora, forested
wetlands, Connecticut River, Massachusetts
Although the importance of forests along floodplains within
river ecosystems has been increasingly recognized (Decamps
154
2000] Holland et al—Oxbow Swamp Forest Understory 155
1996; Gregory et al. 1991), the dynamics of floodplain forest
vegetation remain poorly understood. Not surprisingly, investi-
gations of economically important floodplain forest trees (Jones
et al. 1994; Megonigal et al. 1997; Robertson et al. 1978) have
been more common than studies of floodplain herbs and shrubs
(Menges and Waller 1983). Kearsley (1999a) has recently inven-
toried Massachusetts floodplain forests and devised a classifica-
tion that includes understory species; however long-term studies
of the composition and structure of lower floodplain forest strata
continue to be scarce.
In western Massachusetts, studies of vegetation within Con-
necticut River oxbows have been in progress since 1969 (Robin-
ton and Burk 1971). Four of these oxbows form an apparent
chronosequence, a “‘spatial array”’ of sites of different ages (Bar-
bour et al. 1987) in various stages of seral development ranging
from open water through freshwater marsh to mature swamp for-
est (Holland and Burk 1990). The most thoroughly studied of
these sites, Ned’s Ditch, is the northwestern segment of an oxbow
that was cut off from the main stream of the Connecticut River
around 710 (+130) YBP as determined by stratigraphy and ra-
diocarbon dating (Holland and Burk 1982). Now owned by the
Massachusetts Audubon Society as a part of Arcadia Wildlife
Sanctuary, Ned’s Ditch contains one of the largest stands of flood-
plain swamp forest in New England. The vegetation of the area
within the old river bed includes this forest, stands of buttonbush
(Cephalanthus occidentalis), and relic marsh communities sur-
rounding ponds and remnants of the abandoned channel of an
adjacent stream (Holland and Burk 1984, 1990). Ned’s Ditch is
regularly flooded during periods of heavy precipitation and/or
high water on the Connecticut River with floodwaters entering its
eastern section from an adjacent oxbow that was cut off from the
main stem of the Connecticut River in 1840 (Holland and Burk
1982). During unusually heavy flooding, floodwaters also enter
from the channel of the Mill River on the western margin of
Ned’s Ditch. Studies conducted during the period 1973-1977
(Holland and Burk 1984) indicated that the canopy trees, partic-
ularly the most prominent species, Acer saccharinum, Quercus
palustris, and Fraxinus pennsylvanica, were well represented by
seedlings in the understory and apparently replacing themselves.
Hence the Ned’s Ditch forest community was thought to be in a
state of “hydric disclimax’’ (Daubenmire 1968) or “‘pulse stabil-
156 Rhodora [Vol. 102
ity’ (Odum 1969) that might persist indefinitely (Holland and
Burk 1984).
A resampling of the Ned’s Ditch canopy in 1985, however
(Holland and Burk 1986), indicated that a number of trees of all
species had died since 1975, with highest mortalities in the lower
size classes (3.0—13.5 cm diameter). In addition, recruitment from
the herb stratum had been extremely low. Comparable tree deaths
and low recruitment were also observed by studies in a regularly
flooded oxbow in Hatfield, Massachusetts, a “‘younger’”’ stage in
the oxbow chronosequence (Holland 1998; Holland and Burk, in
press), but not in an “older” rarely flooded stage on a higher
Connecticut River terrace in Whately, Massachusetts (Holland
and Burk 1986, in press, and unpubl. data). Field studies a decade
later in Ned’s Ditch indicated that few canopy trees died between
1985 and 1995/96. Nonetheless, although seedlings of Acer sac-
charinum, Quercus palustris, and Fraxinus pennsylvanica were
abundant, few trees had been recruited into the smaller stem clas-
ses of the canopy (Burk et al. 1996).
This study compared the shrub and herb strata of Ned’s Ditch
during the interval 1973-1977 with their state in 1985 and again
in 1995/96. Specific goals of the study were:
(1) to document the abundance and distribution of the vascular
plant species comprising the shrub and herb strata of the
Ned’s Ditch swamp forests throughout this 23 year interval;
to assess changes in the abundance and distribution of vas-
cular plant species in these strata throughout the study period;
to attempt to identify long-term changes in vegetation be-
tween 1973 and 1995/96, and to distinguish long-term chang-
es from changes that reflect short-term fluctuations in hydro-
logical conditions, including drought and flooding;
to examine relationships between the vegetation of the lower
strata of the swamp forest and the vegetation of adjacent
marshes.
—
N
a
—
ive)
—
S
MATERIALS AND METHODS
Quantitative sampling of the canopy, shrub, and herb strata of
the Ned’s Ditch forest was initiated in the summer of 1973 and
repeated during 1974, 1975, 1977, 1985, and 1995-1996. Quan-
2000] Holland et al—Oxbow Swamp Forest Understory — 157
titative sampling of marsh vegetation in Ned’s Ditch was con-
ducted in 1973, 1974, 1975, 1984, and 1994.
As a part of the 1973 sampling, five transects were laid out at
285 m intervals across Ned’s Ditch, beginning near its eastern
margin. Each transect extended from the old Connecticut River
bank through the former river bed to the opposite bank, a distance
of approximately 306 m. Along each transect, swamp forest veg-
etation occurred within the former Connecticut River bed, adja-
cent to the old Connecticut River bank on both sides but at a
lower elevation. Stands of buttonbush swamp and ponds with
relict marsh communities occurred farther towards the centers of
the transects; transitions between swamp forest and the button-
bush swamp and marsh communities were abrupt and boundaries
between each vegetation type were easily delimited.
To sample swamp forest vegetation, ten 10 m xX 10 m per-
manent plots were established, two on each transect beginning
3.3 m in from the edge of the forest closest to the bank at each
end. In 1996, the four corners of each plot were marked with
buried rebar which could be re-located by means of a metal de-
tector during successive sampling periods. The shrub stratum in
each plot was sampled within a 5 m X 5 m quadrat located in
one corner of each plot. Presence and coverage were estimated
for each woody vascular plant species occuring as a shrub or
sapling. To sample the herbaceous stratum within each 10 m X
10 m plot, ten | m <* | m quadrats were laid out, five at regular
intervals along the transect and five at regular intervals along a
baseline perpendicular to the transect. Presence and coverage as
determined by visual estimate were noted for all vascular plants
including herbs, vines, and woody seedlings under 60 cm height
in each of the 100 smaller quadrats. In addition the presence and
cover of the aquatic moss Amblystegium riparium (Hedw.) BSG.
(Amblystegiaceae) was also recorded for each smaller quadrat.
Sampling of the ten 5 m X 5 m shrub quadrats was conducted
during July and August of 1973, 1975, and 1985. Because of
time constraints, six shrub quadrats were sampled in July and
August of 1995 and the remaining four in 1996. Sampling of all
100 1 m X | m quadrats was conducted during July and August
of 1973, 1974, 1975, 1977, and 1985. Sampling of 60 smaller
quadrats within six 10 m X 10 m plots was conducted in 1995
and the remaining 40 smaller quadrats within the four other 10
m xX 10 m plots were sampled in 1996. During each sampling
158 Rhodora [Vol. 102
Centimeters
)
1950 1954 1958 1962 1966 1970 1974 1978 1982 1986 1990 1994 1998
Year
Figure |. Total annual precipitation (cm) in the study area, 1950 through
1998. See text for sources
period, studies of the shrub and herb strata were coordinated with
sampling of the canopy within each plot.
In some instances, particularly in years when growth was de-
layed or disrupted by late spring or summer flooding, a number
of very small plants in the quadrats could be identified only at
the generic level. Because of this, immature specimens of several
regularly or sporadically encountered genera that might be rep-
resented in the sampling by more than one species are grouped
together under their respective genera. Nomenclature of fern spe-
cies follows Flora of North America North of Mexico (Flora of
North America Editorial Committee 1993). Nomenclature of seed
plants follows Fernald (1950) except for Bidens tripartita L. to
include B. comosa (A. Gray) Wieg. and B. connata Muhl. Vouch-
er specimens have been deposited in SCH.
For analysis and comparison, average percent cover and fre-
quency were recorded for each species with cover values rounded
off to the nearest whole number. Summed cover values for each
year’s sampling were calculated using actual rather than rounded
data. Species richness, the total number of species present in each
stratum, was calculated by counting all species present. Species
grouped to genera as a single unit were counted only once in
determining richness. For example, seedlings listed as Cornus
spp. were counted as such, even though mature specimens of C.
2000] Holland et al——Oxbow Swamp Forest Understory 159
alternifolia L.f., C. amomum Miller, and C. stolonifera Michx.
were present at the site as potential seed sources. Discussions of
the value of cover and frequency data in assessing vegetational
change are in Daubenmire (1968) and Mueller-Dombois and E]-
lenberg (1974), and are summarized with reference to the adjacent
oxbow marshes in Holland and Burk (1990).
The permanent plots and all quadrats were systematically
placed, and we have attempted to re-locate each individual quad-
rat at each sampling period; hence probability statistics cannot be
used as aids in interpreting the data (Barbour et al. 1987; Mueller-
Dombois and Ellenberg 1974). As in earlier studies of changes
in floristic composition over a series of collecting periods (Hol-
land and Burk 1990; Holland and Sorrie 1989; Lauermann and
Burk 1976), the Simpson Index of Resemblance (Simpson 1965)
was used to compare the taxonomic composition of the herb stra-
tum in successive years of sampling. Simpson’s Index of Resem-
blance (100 c/nl, in which c is the number of species common
to the two floras and nl is the number of species in the smaller
flora) is helpful in comparing floras of approximately equal sizes
in a common area. In addition, to assess interrelationships be-
tween the composition of the Ned’s Ditch swamp forest and floras
of adjacent Ned’s Ditch marshes, Simpson’s Index was also used
to compare the swamp forest in 1977, 1985, and 1995/96 with
the composition of the Ned’s Ditch marshes. The marshes were
sampled in 1974 and 1984 (Holland and Burk 1990) and again
in 1994, one year prior to the sampling of the forest (Holland and
Burk, unpubl. data).
Data on yearly precipitation (Figure 1) were obtained from
the website www.ncdc.noaa.gov/onlineprod/drought/temp/
drought_15006 for western Massachusetts from 1950 through
1998. Data on the times of major flooding were obtained from
our own observations of the site.
RESULTS
As in earlier studies (Holland and Burk 1984, 1990) tables
were prepared listing all vascular plant species in any quadrat for
each sampling period. Data from 1973 are available in Sackett
(1974) and data from 1973, 1974 and 1975 in Sackett (1977).
The 1975 data are also included in Holland and Burk (1984) with
an indication of species sampled earlier but not in 1975 and spe-
Table 1. Composition of the herb stratum in Ned’s Ditch in 1977, 1985, and 1995/96. Numbers represent mean and standard
deviation (in parentheses) of percent cover (C), frequency (F), and total number of species sampled. 'mostly B. tripartita L. and
B. cernua L., *mostly S. nigra Marshall.
1977 1985 1995/96
Species Cc F Cc F S F
Herbs and vines
Agrostis alba L. <0.5 (0.83) 4 — a — _—-
Agrostis perennans (Walter) Tuckerman I .G25) 11 — — — —
Alisma subcordatum <0.5 (0.88) 5 <0.5 (0.06) 1 <0.5 (0.66) 4
Apios americana Medikus 3 (8.14) 11 7 (14.68) 19 2 (4.34) 16
Arisaema triphyllum (L.) Schott <0.5 (0.48) 5 <0.5 (0.13) 5 <0.5 (0.72) 8
Bidens frondosa L. 4 (4.85) 61 3. (6.27) 34 <0.5 (0.68) 14
Bidens spp.! <0.5 (0.70) 18 7 (5.24) 75 <0.5 (0.16) 8
Boehmeria cylindrica (L.) Swartz 1 (1.35) 46 4 (7.97) 50 2 (4.46) 19
Cardamine pensylvanica Muhl. <0.5 (0.28) 6 <0.5 (0.02) 1 — —
Carex tribuloides Wahlenb. 3 (10.25) 10 <0.5 (0.20) <0.5 —
rex spp. 1 (1.12) 18 <0.5 (0.27) <O0.5 1 (1.24) 7
Celastrus scandens L. — — <0.5 (0.03) 1
Cicuta bulbifera — es <0.5 (0.17) 6 — -=
Cuscuta cas Willd. — a <0.5 (0.03) 1 — —
s carthusiana (Villars) H. P. Fuchs 1 (2.14) 3 1 (2.37) 4 —
ie at arundinaceum (L.) Britt <0.5 (0.09) 5 <0.5 (0.13) 4 — oo
Echinochloa crusgalli (L.) P. Bea — — <0.5 (0.03) 1 --
Eleocharis acicularis (L.) snraeoes & Schultes 6 (13.4) 25 <0.5 (0.16) 16 1 (1.26) 5
Erechtites hieracifolia (L.) Raf. — — <0.5 (0.52) 9
Eupatorium spp. <0.5 (0.32) 1 - a —
—
a
Oo
vlopoyYy
TOI IPA)
Table 1. Continued
1977 1985 1995/96
Species C set c F Cc E
Galium spp. 1 (0.54) 31 1 (1.52) a2 <0.5 (0.47) 9
Geum virginianun — <0.5 (0.22) 6 — —
es St. John == 12 (14.56) 54 3 (6.25) 20
Hieracium spp. — — <0.5 (0.04) 2 = —
Hypericum virginicum L. <0.5 (0.09) | <0.5 (0.88) 7 <0.5 (0.19) 2
Impatiens capensis Meerb <0.5 (0.09) 3 <0.5 (0.68) 6 <0.5 (0.16) 4
Leersia ¢ er 1 (1.44) 11 <0.5 (0.90) 15 — —
Lemna minor 4 (7.71) 36 <0.5 (0.14) 3 <0.5 (0.22) 26
Ludwigia es (L.) Ell. 1 (2.35) 25 <0.5 (0.14) 17 —
Ludwigia polycarpa Short & Peter <0.5 (0.69) 2 <0.5 (0.40) 2 — —
Lycopus uniflorus Michx — — —_ <0.5 (0.68) 10
Lycopus virginicus L. <0.5 0. 19) > <0.5 (1.10) 5 <0.5 (0.47) I
Lysimachia ciliata L <0.5 (0.13) 5 <0.5 (0.13) l — —
Lysimachia nummularia L. <0.5 (0.06) 2 <0.5 (0.13) 3 1 (0.89) 9
Lysimachia terrestris (L.) BSP. 1 (0.84) 13 1 (1.09) 25 1 (1.09) 11
Onoclea sensibilis L 6 (13.97) 13 5 (9.27) 27 6 (14.72) 18
Osmunda regalis L 5 (10.47) 15 8 (16.27) 13 6 (10.94) 14
Panicum clandestinum L. <0.5 (0.54) 3 —_ — — —
Parthenocissus quinquefolia (L.) Planchon — — — — <0.5 (0.22) 6
Penthorum sedoides 1 (2.74) 8 <O0.5 (0.14) 3 —
Phalaris arundinacea L. — — <0.5 (0.28) 2 — —
Pilea pumila (L <0.5 (0.68) 16 — — —
Polygonum amphibium L. — — | (1.64) 6 — —
Polygonum hydropiperoides Michx. _— — <0.5 (1.22) 7 — —
L000
Alo\siopuy) jsa10o,, duewmsg mogxQ—'[R 19 purl][oy
Table 1. Continued.
1977 1985 1995/96
Species Cc F ie F Cc F
ee punctatum Elliott <0.5 (0.82) 9 — — — —
Polygonum spp. <0.5 (0.03) 1 — — <0.5 (0.16) 2.
Prunella as L. — — <0.5 (0.02) ] — —
Ranunculus flabellaris Raf. 8 (10.54) 38 1 (0.67) 40 <0.5 (0.82) 12
Rhus radicans L <0.5 (0.04) Z <0.5 (0.18) 9 1 (2.78) 6
Sagittaria latifolia Willd. — = <0.5 (0.19) I — —
Scutellaria pi La 1 (0.75) 23 1 (1.02) 18 <0.5 (0.44) 6
Sium suave Wal 1 (0.97) 17 1 (1.25) 43 <0.5 (1.00) 7
Smilax herbacea 7 <0.5 (0.28) 3 <0.5 (0.35) 3 <0.5 (0.32) l
Solanum dulcamara L. — — <0.5 (0.25) 6 <0.5 (0.16) 2
Solidago spp. 5 (0.03) | — — — —
coaane polyrhiza (L.) Schleiden <0.5 (0.32) 8 — a — —
Thelypteris palustris Schott — — — — 2 (4.60) BS)
Woody seedlings
Acer saccharinum L. 9 (6.03) 77 8 (8.90) 78 7 (9.64) 60
ie faa eee Sprengel — = <0.5 (0.06) 1 — —
a Warder — — <0.5 (0.02) 1 — —
eh on penton L. 7 (5.36) 62 5 (4.92) 60 3. (2.62) 37
Cornus spp. 1 (0.99) 20 <0.5 (0.73) 4 <0.5 (0.81) 4
Fraxinus pennsylvanica ot 2 (3.06) 51 2 (3.00) 36 3 (3.94) 42
Ilex aie snes (L.) A. Gray 1 (1.24) 13 <0.5 (0.33) 5 <0.5 (0.47) 1
Populus deltoides sect — — <0.5 (0.05) 2 — —
ee palustris Muenchh. 1 (2.36) 7 <0.5 (0.28) 19 1 (1.28) 17
Robinia pseudoacacia L. <0.5 (0.38) 14 — = <0.5 (0.06) 11
Salix spp.’ <0.5 (0.35) 7 <0.5 (0.03) 1 <0.5 (0.02) 1
co
vlopoyy
JOA]
cOl
Table 1. Continued.
1977 1985 1995/96
Species Cc F C F Cc F
Ulmus rubra Muhl. <0.5 (0.05) 3 <0.5 (0.09) 10 <0.5 (0.25) 8
Viburnum recognitum Fern. — — — <0.5 (0.06) 1
Vitis spp 1 (1.39) 27 2 (2.67) 24 1 (2.20) 5
Total species 50 a7 39
LO00T
Alo\siopup) 1sa104 duremg mogxQ— [P 19 pur[[oH
C9]
164 Rhodora [Vol. 102
cies not in any quadrat but present elsewhere in swamp forest at
the site. Data from 1977, 1985, and 1995/96, all previously un-
published, are included here in Table 1.
Despite the changes in the canopy of the Ned’s Ditch swamp
forest that were evident by 1985 (Burk et al. 1996: Holland and
Burk 1986), the shrub stratum has remained relatively unchanged
during the interval 1973—1995/96 (Table 2). Coverage of the
dominant Cephalanthus occidentalis declined between 1973 and
1975 but returned to near 1973 levels by 1985. A decline of Acer
saccharinum and Fraxinus pennsylvanica observed in the 1995/
96 sampling of the shrub stratum probably reflects the recruitment
of saplings of these tree species from the shrub stratum into the
canopy.
Changes in the herb stratum for the period 1973 through 1977
were presented in Holland and Burk (1984). Within the herb stra-
tum, total vegetative cover rose from a low of 14% in 1973 to
71% by 1975 and continued at a slightly higher level through
977. The very low cover in 1973 resulted from damage caused
by an atypical flood that had completely inundated the herb stra-
tum for at least ten days in early July, less than a month before
the sampling period (Holland and Burk 1984). Species richness
rose from a low of 27 in 1973 to 47 by 1975 (Holland and Burk
1984) and continued at a somewhat higher level through 1977
(Table 1).
Total cover in the herb stratum remained relatively constant
from 1975 through 1985 and then declined over the next ten year
period (Table 3). Overall cover was lowest in years with late
spring or summer floods and highest in wet years without late
spring or early summer flooding. Species richness reached a high
of 57 species in 1985 but had declined to 39, little above its 1974
level by 1995/96 (Table 1). The higher diversity in 1985 resulted,
in part, from the presence of weedy species such as Erechtites
hieracifolia and Prunella vulgaris, which had become established
in small numbers under the unusually dry conditions of that sea-
son, and possibly, in part, because of the higher light levels in
lower strata resulting from the extensive mortalities among can-
opy trees. Most invaders of this sort did not persist through 1995/
96 (Table 1).
From 1977 through 1995/96, the abundance of the prominent
rhizomatous ferns, Onoclea sensibilis and Osmunda regalis re-
mained relatively constant, presumably in large part because the
Table 2. Variation in the cover of shrubs in Ned’s Ditch. Numbers represent mean and standard deviation (in parentheses) of
percent cover of each vascular plant species and percent total cover.
Year
Species 1973 1975 1985 1995/96
Acer saccharinum 8 (15.22) 11 (31.35) 8 (15.10) 1 (1.75)
Alnus rugosa — <0.5 (0.32) <0.5 (0.63) <0.5 (0.95)
Cephalanthus occidentalis 37 =(32.65) 20 (26.38) 33 (31.24) 35 (39.08)
Cornus spp. 5 (5.77) 4 (4.86) <0.5 (0.95) 2 (4.83)
Fraxinus pennsylvanica — 2 (4.69) 8 (23.68) <0.5 (0.34)
Ilex verticillata 1 (2.71) 2 (5.25) 4 (6.34) 5 (11.07)
Quercus palustris 1 (1.63) <0.5 (0.42) — a
i — 1 (3.48) — —
Viburnum recognitum — <0.5 (0.32) — a
Vitis spp. 0.5 (1520) 1 (1.90) — 4 (11.01)
Total cover (%) 52 45 54 48
LO00T
AlO\sIopuy) Isao dureMmsg mogXOQ— [PR 19 pur[[OH
Table 3. Variation in percent relative cover of vascular plant species in the herb stratum of Ned’s Ditch. Relative cover of
species at 5% or more total cover at any sampling period and total cover—see discussion of Amblystegium riparium in text.
Year and Hydrology
1973 1974 1975 1977 1985 1995/96
Species July Flood Dry Wet Wet Dry Dry/Wet
Acer saccharinum 7 1] 6 12 1] 17
Apios americana ] <0.5 ] 3 10 5
Bidens spp. <0.5 | ] 4 14 2
Boehmeria cylindrica <0.5 <0.5 <0.5 ] 5 4
Cephalanthus occidentalis ] 8 6 10 7 6
Dulichium arundinaceum 29 17 10 <0.5 <0.5 6)
Eleocharis acicularis l 3 3 8 <0.5 1
Fraxinus pennsylvanica ! 1 ] 2 3 6
Glyceria fernaldii — — — oe 16 7
Lemna minor 7 3 48 7 <0.5 <0.5
Onoclea sensibilis 14 11 ] 8 7 15
Osmunda regalis 14 14 4 7 1] 14
Ranunculus flabellaris ] ) 3 1] ] ]
Total cover (%) 14 36 71 73 74 43
991
viopoyy
JOA]
cOl
2000] Holland et al—Oxbow Swamp Forest Understory 167
same clones have been encountered at each sampling period. In
addition, three major trends were noted within the vegetation of
the herb stratum:
(1) The relative abundances of the more important species con-
ae
No
~—
os
ies)
Ser!
tinued to fluctuate from year to year, apparently in relation to
hydrology (Figure |; Table 3). Hydrophytes increased during
wet years such as 1975 and 1977 (Holland and Burk 1984).
During dry seasons, annuals, particularly Bidens spp. grew to
maturity and, while flowering, achieved “‘aspect dominance”
(Oosting 1956) as in 1985 (Table 3 and field observations
during dry years when sampling was not conducted).
The relative cover of tree seedlings, particularly seedlings of
Acer saccharinum and Fraxinus pennsylvanica increased (Ta-
ble 3). In 1973, Dulichium arundinaceum, Onoclea sensibilis,
and Osmunda regalis were the most prominent species, in
descending order of abundance. By 1995/1996, A. sacchar-
inum was most abundant, followed by Onoclea sensibilis and
Osmunda regalis.
Hydrophytes generally declined from previous levels of abun-
dance. These included both ‘“‘errant hydrophytes” (Mueller-
Dombois and Ellenberg 1974) such as Lemna minor, Ranun-
culus flabellaris, and Spirodela polyrhiza (Table 1), and
emergents. Potamogeton pectinatus L. and Utricularia vul-
garis L. were sampled in 1975 (Holland and Burk 1984) but
not later. A nonvascular errant hydrophyte, the moss Am-
blystegium riparium was present at every sampling period in
the 1970s, reaching a peak cover of 21% of the total sub-
stratum in 1977, occurring in 81% of the quadrats sampled.
The peak abundance of A. riparium appeared to be inversely
related to the abundance of L. minor, which had reached its
maximum in 1975 and declined by 1977 (Table 3). Amblys-
tegium riparium was not seen during 1985, a dry season, nor
identified in 1995; it was encountered again in nearly half
the plots sampled in 1996 with an average cover of 2% of
the substratum.
Several emergent hydrophytes also found in the high- and mid-
marsh zones of the adjacent ponds declined markedly as well.
Dulichium arundinaceum, the most abundant species of the herb
stratum in 1973 and 1974, was present only at very low levels
168 Rhodora [Vol. 102
Table 4. Simpson’s Index of Resemblance comparing the flora of the
Ned’s Ditch swamp forest herb stratum at each sampling period to the flora
of successive samplings of that stratum.
Year of Years of Successive Sampling
Initial
Sampling 1974 1975 1977 1985 1995/96
1973 88.9 81.5 77.8 70.4 63.0
1974 77.8 $3.3 1. 63.9
1975 722 70.2 74.4
1977 72.9 81.6
1985 81.6
in 1977 and 1985 and not sampled at all in 1995/96, Eleocharis
acicularis persisted at generally low levels throughout the period,
reaching its greatest abundance in 1977 and then declining.
Table 4 compares the herb flora of Ned’s Ditch at each sam-
pling period from 1973 to 1985 to the floras of successive sam-
pling periods at the specific level. Floras from sampling periods
closer in time tended to be more similar than floras from longer
intervals, and the least similar floras were those of the first and
last collections of data, 1973 and 1995/96. When Ned’s Ditch
marshes were compared with the swamp forest herb stratum in
successive years of sampling, similarities tended to be higher
when comparisons were made between marsh and forest floras at
the closer time intervals (Table 5). Levels of forest/marsh simi-
larity fall within the range of forest/forest similarities. The Simp-
son Index comparing the composition of the Ned’s Ditch swamp
forest in 1974 with the forest in 1995/96 is identical with Simp-
son’s Index comparing the Ned’s Ditch marsh in 1974 with the
forest in 1995/96.
Table 5. Simpson’s Index of Resemblance comparing the flora of the
Ned’s Ditch marshes to the herbaceous flora of the adjacent swamp forest
herb stratum at each successive sampling.
Year of Years of Swamp Forest Sampling
arsh
Sampling Lo7373 oT? 1985 1995/96
1974 75.8 75.0 72.2 63,9
1984 74.1 71.1
1994
79.5
2000] Holland et al—Oxbow Swamp Forest Understory 169
DISCUSSION
Fluctuations in the composition of the herb stratum of Ned’s
Ditch in response to seasonal variations in hydrology may rep-
resent a community response to conditions associated with a par-
ticular form of hydric disclimax, i.e. a persistent state maintained
by recurrent but irregular flooding. During wet seasons, hydro-
phytic species become abundant; during dry seasons these are
largely replaced by mesophytic forms (Holland and Burk 1984).
Together the understory flora represents a spectrum of life forms
including annuals, woody seedlings, emergent perennial grami-
noids, perennial forbs, errant hydrophytes, annuals, and vines.
Previous comparisons of the herb stratum of the Ned’s Ditch
swamp forest to the floras of adjacent marshes have shown a
stronger resemblance than that of other oxbow swamp forests and
their marshes (Holland and Burk 1990). While the low marsh
dominants Nuphar variegata Durand and Potamogeton pectinatus
did not extend into the forest, many species were shared by both
communities (Table 5), and despite the decline of hydrophytes in
the swamp forest over the last two decades of the study, the floras
of the marshes and the herb stratum of the forest continued to be
strongly similar. In part this similarity reflects dispersal during
periods of flooding when high water extended throughout the
oxbow, connecting ponds and forest. In addition, the ponds oc-
casionally dried out and species more frequently found in the
forest became established on exposed pond bottoms.
The increased relative abundance of woody seedlings and over-
all decline in hydrophytes in Ned’s Ditch may have resulted, in
part, from drier conditions associated with changes in elevation
resulting from sedimentation and from reduced available light on
the forest floor associated with the growth of the canopy trees.
Studies of comparable Wisconsin floodplain forests have char-
acterized Bidens spp. and Lemna spp. as high-light specialists at
the lowest elevations of the herb strata and Acer saccharinum and
Onoclea sensibilis as light generalists at somewhat higher ele-
vations; seedlings of A. saccharinum have been shown to ger-
minate well in either light or shade (Peterson and Bazzaz 1984).
Scouring through heavy flooding and high mortalities of the dom-
inant trees before 1985 (Burk et al. 1996) may have slowed or
reversed these trends since long periods of flooding are particu-
170 Rhodora [Vol. 102
larly harmful to young seedlings of A. saccharinum (Peterson and
Bazzaz 1984).
Similar effects of different flood regimes have also been ob-
served in Louisiana swamp forests that contain understory species
in common with Ned’s Ditch (Conner and Day 1976). Stands that
were flooded to a depth of 0.6 m much of the year possessed
constantly saturated soil and herb strata containing only floating
and emergent aquatics, including Lemna minor and Spirodela po-
lyrhiza. Forests that were inundated for two to three months in
the spring but surface-dry in summer supported more diverse
lower strata of “‘briars, grasses and annual herbs” along with
seedlings, saplings, and woody vines.
Related studies (Conner et al. 1981) contrasted the effects of
altered flood regimes in swamp forests that had shared similar
vegetation and seasonal flooding through the 1950s. In an undis-
turbed control site, growth of aquatics and understory woody veg-
etation was limited by shading and periodic flooding. In a site
now permanently flooded, the resultant deaths of canopy trees,
particularly Fraxinus spp., increased light penetration and al-
lowed the spread of aquatics, including Lemna minor and Spi-
rodela_ polyrhiza. After permanant inundation, flood-tolerant
shrubs also invaded, including Cephalanthus occidentalis, the
seeds of which are capable of germinating underwater (see ref-
erences in Conner et al. 1981). In a site now managed by con-
trolled winter/spring flooding followed by an annual summer/fall
drawdown, aquatic species were absent. The managed site was
increasingly dominated by Acer rubrum var. drummondii and
Fraxinus, taxa with seeds that germinate during the dry period to
produce seedlings that become established before flooding.
Since the initial sampling of vegetation in Ned’s Ditch, the
concept of a vegetation type that may persist indefinitely through
“pulse stability’’ (Odum 1969) has been explored with reference
to floodplain forests in particular. Odum et al. (1979), using data
from Conner and Day (1976) for examples, have suggested that
moderate or seasonal flooding may result in increased tree growth
in mature floodplain forests composed of species already well
adapted to the flooding regime. Continuous high levels of flood-
ing, however, may result in impounded and stagnant conditions
that stress canopy vegetation and reduce productivity. More re-
cent studies (Megonigal et al. 1997) conclude that the results of
flooding are complex and affected by the timing and length of
2000] Holland et al—Oxbow Swamp Forest Understory 171
floods and the relative strength of their flow. In addition, stresses
resulting from drought and anaerobic soils may offset any benefits
of flooding (Mitsch and Rust 1984).
CONSERVATION
The integrity of the Ned’s Ditch swamp forest community is
largely dependent on dynamic hydrological conditions resulting
from periodic flooding on the Connecticut River. In 1973, at the
time of the initial sampling, much of the site was privately owned
and subject to intermittent logging, dumping of wastes, and other
disruptive human activities. The entire Northampton oxbow in-
cluding the Ned’s Ditch forest is now owned by the Massachu-
setts Audubon Society and preserved as a natural area with a
management plan (McGuire 1988) that includes control of poten-
tially invasive exotic species. With the exception of Lysimachia
nummularia, non-native species have not been regularly encoun-
tered in Ned’s Ditch; although Solanum dulcamara, Prunella vul-
garis, and seedlings of Catalpa speciosa were sampled in 1985,
only S. dulcamara persisted until 1995. Nonetheless, a number
of potentially invasive species, several of which are now fre-
quently encountered in other Massachusetts floodplain forests
(Kearsley 1999b) are well established nearby. These include Acer
platanoides L., Berberis thunbergii DC, Celastrus orbiculatus
Thunb., Polygonum cuspidatum Sieb. & Zucc., Rhamnus frangula
L., and Catalpa speciosa, which is spreading along the Mill River
at the western margin of Ned’s Ditch (Burk and Prabhu 1988).
Spread of these species should be monitored and, if necessary,
curtailed because of their potential harm to native species (Weath-
erbee et al. 1998). Of particular concern is Ludwigia polycarpa,
currently threatened in Massachusetts (Massachusetts Natural
Heritage and Endangered Species Program 1997) and not seen at
its former locations in Ned’s Ditch since 1985 (Holland and Burk
1990).
Sustaining biological diversity may be particularly critical in
ecosystems such as floodplain forests that experience seasonal or
longer fluctuations in hydrology (Grime 1997; Keddy and Rez-
nicek 1982). Our studies, along with other investigations of flood-
plain forest vegetation, suggest that preserving and successfully
managing these communities will require the maintenance of spe-
cies of diverse ecological requirements adapted to a range of hab-
eg? Rhodora [Vol. 102
itat conditions. In these dynamic systems, individual species tend
to fluctuate in abundance and may sometimes disappear com-
pletely from a given floodplain forest site. Hence a range of sim-
ilar protected sites within the region may be essential in providing
reservoirs from which species may be recruited as habitat con-
ditions change. Since our earliest vegetation studies in Ned’s
Ditch, we have argued for the conservation of floodplain forests
and other wetland habitats (Holland and Burk 1984, 1990); and
Kearsley’s recent inventory (Kearsley 1999a) reinforces the ne-
cessity for maintaining these increasingly scarce plant commu-
nities, particularly transitional and ‘‘Small-river’’ floodplain for-
ests, few of which are now protected, on a broader scale within
the state and region.
ACKNOWLEDGMENTS. As the Ned’s Ditch research moves along
through its third decade, we are increasingly grateful to the staff
of Arcadia Wildlife Sanctuary and to the Massachusetts Audubon
Society for their encouragement and support, and to those who
helped in the initial stages of this work and were acknowledged
in earlier publications (Holland and Burk 1982, 1984, 1990). In
addition we thank our more recent field assistants, particularly
Molly Kornblum, Jessie Lang, Alyssa Lovell, and Laurie Sanders,
as well as Bibiana Garcia Bailo, who assembled the precipitation
data.
LITERATURE CITED
Barsour, M. G., J. H. BurK, AND W. D. Prrrs. 1987. aaa o. Ecol-
ogy, 2nd ed. Benjamin/Cummings one Co., Menlo
Burk, C. J.. M. M. HOLLAND, AND D. McLain. 1996. High shea ai low
recruitment in a New England Soci forest Bull. Ecol. Soc. Amer.,
denen Program and Abstracts: Part 2: 61.
V. PRABHU. 1988. Growth and expansion of a naturalized western
hie stand of Catalpa speciosa Warder. Rhodora 90: 457—460.
CONNER, W. H. AND J. W. Day, Jr. 1976. Productivity and composition of a
baldcypress-water tupelo site | a bottomland hardwood site in a Loui-
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RHODORA, Vol. 102, No. 910, pp. 175-197, 2000
VASCULAR FLORA OF BEAVER WETLANDS IN
WESTERN MASSACHUSETTS
ROBERT T. MCMASTER! AND Nancy D. MCMASTER
Department of Biological Sciences,
Smith College, Northampton, MA 01063
e-mail: rmcmaste @science.smith.edu
ABSTRACT. The composition and structure of vegetation in beaver wet-
lands located in Franklin and Hampshire Counties of western Massachusetts
were studied from 1980 to 1995. A flora of 231 vascular plants was recorde
within fifteen selected field sites including seven new county records and one
state-listed species (Ophioglossum vulgatum var. pseudopodum). Relatively
few species (5.6% of the total flora) were introduced. Factors contributing to
high vascular plant diversity included 1) steep hydrological gradients created
by beaver dams; 2) spatial eames caused by beaver artifacts, human
structures and geological features; and 3) temporal heterogeneity resulang
from beaver activity and hydrological fluctuations. While beaver activity has
helped maintain regional vascular plant diversity in western Massachusetts
since the reestablishment of beaver populations in the 1920s, current increases
in beaver densities may pose a threat to later successional wetland species.
Key Words: beaver (Castor canadensis), vascular flora, wetlands, Massa-
chusetts
The beaver (Castor canadensis) has had a profound impact on
the landscape of western Massachusetts. Trapped to regional ex-
tinction in the mid-18th century, it returned in 1924 when a col-
ony immigrated into West Stockbridge (Berkshire County). Aug-
mented by reintroductions by the Massachusetts Department of
Conservation, the state-wide population expanded rapidly, total-
ing approximately 52,000 in 1999 (S. Langlois, pers. comm.).
Beavers alter the structure and composition of both upland and
riparian vegetation. They forage on woody vegetation 60 m or
more from a stream course, girdling and felling trees, shrubs, and
vines (Hall 1960; Rutherford 1955) and utilizing over 90 woody
food species throughout their range (McMaster 1989).
Beaver dams in Massachusetts are constructed of branches,
vines, shrubs, herbaceous plant material, boulders, sand, gravel,
and mud. They range from 0.3 to 2 m in height and 3 to 300 m
in length (Shaw 1948). Often placed against stone walls or cul-
verts, they span stream channels and extend to the edges of the
adjacent uplands.
L/S
176 Rhodora [Vol. 102
New dams create impoundments up to 8 ha in area extending
upstream and into the uplands along the stream margins. Above
dams, soils are saturated, stream velocity is reduced, and sediment
accumulates. Below dams, discharge rates decline and sediment
loads are reduced (Naiman et al. 1988). These hydrological al-
terations in turn influence stream biogeochemistry including car-
bon cycling (Naiman et al. 1986), methane evasion rate (Ford and
Naiman 1988), soil redox potential (Naiman et al. 1988), pH and
acid-neutralizing capacity (Smith et al. 1991), and biological ox-
ygen demand and dissolved oxygen concentration (Naiman et al.
1988).
Following inundation, most woody vegetation is killed within
one or two growing seasons. Within five years shallower areas,
particularly along the pond margins or adjacent to the dams or
lodges, are colonized by floating-leaved and submerged aquatics
and emergent hydrophytic plants.
When beaver colonies are removed by trapping or predation or
when they migrate to another site as food supply dwindles, un-
maintained dams deteriorate and sites gradually drain. The up-
stream end of a site and the lateral margins usually drain first
while the areas immediately above the dam and surrounding the
lodge are often among the last to drain. Eventually the stream
cuts a channel in the bottom sediments before exiting through
one or more openings in the abandoned dam.
An unmaintained beaver dam may impound water during
spring meltout and after periods of heavy precipitation, thus af-
fecting site hydrology for years, sometimes decades, after aban-
donment. Factors that influence the frequency, extent, and dura-
tion of reflooding include the condition of the old dam, the to-
pographical gradient of the site, and the size of the watershed
upstream of the site (McMaster 1997).
The goals of the study initiated by Nancy D. [Mosher] Mc-
Master in 1980 were to describe the composition and structure of
the vascular vegetation in beaver-impacted wetlands, to analyze
vegetational changes over the duration of the study, and to de-
velop a model for succession in beaver-impacted wetlands. She
initially sampled all vascular vegetation in fifteen sites over the
period from 1980 to 1984 and began resampling in 1985, a pro-
cess that continued until her death in 1990 (McMaster 1989). In
1991 Robert T. McMaster resumed the sampling schedule, em-
2000] McMaster and McMaster—Flora of Beaver Wetlands 177
ploying the transects and methodology of the previous work (Mc-
Master 1997).
This paper presents a combined list of vascular plant species
found in all fifteen sites over the period 1980-1995 from both
studies, as well as abundance data, descriptions of plant com-
munities, county and state records, and changes in species diver-
sity. A more detailed analysis of the structure and dynamics of
vascular vegetation over the fifteen-year interval will be presented
in a forthcoming paper.
MATERIALS AND METHODS
Eighty sites in Franklin and Hampshire Counties were initially
identified from topographic maps and aerial photographs as likely
beaver-impacted wetlands. After field reconnaissance, sites cur-
rently occupied by beavers and sites subject to human disturbance
such as burning, logging, and excavation were eliminated. The
fifteen sites thus selected (Figure |; Table 1) ranged in area from
0.6 to 8.0 ha and in elevation from 164 to 465 m above sea level.
All were located on the eastern slope of the Berkshire Plateau in
an area of finely corrugated muscovite schist interbedded with
layers of gneiss and white quartzite (McMaster 1989). Soils were
a combination of glacial tills, alluvium, and muck (Egler 1940).
Five sites were adjacent to paved roads, six to unpaved roads,
and four were accessible only by footpath.
Site histories were reconstructed by examination of USDA Soil
Conservation Service maps and panchromatic black and white
aerial photographs (scale 1:24,000), US Geological Survey maps,
deeds, town histories, by communication with property owners,
and by field observation. Sites were monitored regularly for pres-
ence or absence of active beaver colonies throughout the period
of the study. Site age (i.e. number of years since last beaver
occupation of a site) was determined from communication with
property owners or local residents for activity previous to 1980
and from direct observation in the field for sites occupied since
1980.
Vegetation was sampled along transects established across each
site parallel] to the remains of the beaver dam. Along each tran-
sect, 0.5 m X 0.5 m quadrats were located at 2 m intervals. Five
categories were used to estimate abundance based on criteria de-
Rhodora
e
} }
Hi &
/ \
i \
nee \
j ~
‘ ae
A
/
Ashfield |
i
; Conway
f
<tr ee, A
{ | Williamsburg |
\ “Chesterfield ei
\ Cee ee | \
’ | -
rent tay |
{ }
| | Ww
Figure 1. Location of field sites.
[Vol. 102
2000] McMaster and McMaster—Flora of Beaver Wetlands 179
Table 1. County, watershed, elevation above sea level (m), area (ha), and
age (years) since last beaver occupation (as of 1995) for the fifteen study
sites. 'Refers to the Mill River with headwaters in Conway, which enters the
Connecticut River at Hatfield. "Refers to the Mill River with headwaters in
Goshen and Ashfield, which enters the Connecticut River at Easthampton.
Eleva-
tion Area Age
Site County Watershed (m) (ha) (years)
Ashfield 1 Franklin Westfield River 417 3.8 20
Ashfield 2 Franklin Westfield River 417 1.4 40
Ashfield 3 Franklin Westfield River 393 2.3 l
Ashfield 4 Franklin Mill River' 453 6.2 l
Ashfield 5 Franklin Mill River! 453 25 l
Ashfield 6 Franklin Deerfield River 465 1.8 16
Conway | Franklin Mill River! 369 1.8 25
Conway 2 Franklin Mill River' 362 2.5 35
Conway 3 Franklin Mill River! 338 3.2 18
Conway 4 Franklin Mill River! 329 1.0 1S
Conway 5 Franklin Mill River! 243 35 16
Goshen 1 Hampshire Mill River? 417 2.1 18
Williamsburg | Hampshire Mill River! 220 7.8 26
Williamsburg 2 Hampshire Mill River? 164 0.6 16
Worthington | Hampshire Westfield River 375 8.0 10
scribed by Palmer et al. (1995): abundant, frequent, occasional,
infrequent, and rare.
All sites were originally sampled in mid-summer between 1980
and 1984 (McMaster 1989; Mosher 1981). Three sites, Conway
1, 2, and 3, were resampled at five year intervals. The remaining
twelve sites were to be resampled at ten year intervals when pos-
sible. Seven of these were resampled on schedule. Ashfield 3,
which had been reoccupied by beavers in 1992, was resampled
in 1993 after the site was once again abandoned. Four sites, Ash-
field 6, Conway 5, Goshen |, and Worthington |, which were
first sampled in 1984, could not be resampled in 1994 or subse-
quently due to continued beaver occupation but were revisited at
regular intervals.
Percent species turnover for a site (T) was calculated as fol-
lows:
T = (E+ RYS X 100
where,
E = Number of extinctions between samplings;
180 Rhodora [Vol. 102
Table 2. Floristic summary of plant taxa in fifteen beaver-impacted wet-
lands, Franklin and Hampshire Counties, Massachusetts
Native Introduced Total
Group Families Genera Species Species Species
Pteridophytes 7 8 12 0 12
Gymnosperms 2 ic) 3 0 3
Angiosperms
Dicots 45 89 138 10 148
Monocots 14 32 65 S: 68
TOTAL 68 132 218 ie zo
R = Number of recruitments between samplings;
S = Total number of species present in both samplings of the
site,
The species list presented includes all vascular plant taxa found
in systematic sampling of the fifteen sites as well as in general
reconnaissance of the sites from 1980 to 1995. Nomenclature for
all species and identification of introduced species are based on
Gleason and Cronquist (1991). State and county records are based
on Magee and Ahles (1999) and lists obtained from the Massa-
chusetts Natural Heritage and Endangered Species Program.
Voucher specimens for all taxa are held in the Smith College
Herbarium (SCH).
RESULTS
The flora of the 15 study sites included 231 vascular plant taxa
in 68 families and 132 genera (Table 2; Appendix). The distri-
bution of taxa by life-form was as follows: 108 forbs, 55 gra-
minoids, 30 shrubs, 22 trees, 12 ferns and fern allies, and 4 vines.
Plant families best represented in the flora were Cyperaceae (30
species), Asteraceae (19 species), Rosaceae (16 species), and Po-
aceae (15 species). All but four species were perennials. Two
species are county records for Hampshire County and five are
county records for Franklin County (Table 3). One species is cur-
rently listed as threatened in Massachusetts, Ophioglossum vul-
gatum var, pseudopodum, the northern adder’s-tongue fern (Mas-
sachusetts Natural Heritage and Endangered Species Program
1997). The population in this study was the largest in the state
with approximately 900 sporophytes counted in 1991 (McMaster
2000] McMaster and McMaster—Flora of Beaver Wetlands 181
Table 3. County records among the flora of the fifteen study sites.
Species Site
Hampshire County
Galium trifidum Williamsburg 1 and 2, Worthington |
Juncus conglomeratus Williamsburg |
Franklin County
Carex eburnea 7
Carex lacustris Ashfield 3, 4, 5
Juncus brachycephalus Conway 3, 4
Oenothera fruticosa Conway 3
Potentilla palustris Ashfield 2
1994, 1996). Currently, O. vulgatum var. pseudopodum is also
state-listed in Connecticut (Connecticut Department of Environ-
mental Protection 1995) and Rhode Island (Enser 1998).
Species richness per site ranged from 32 species in Conway 2
(1995) to 79 species in Conway 5 (1984). Number of families
represented in each site ranged from 19 families in Conway 2
(1995) to 34 families in Conway 5. Over one-third (84) of the
species occurred in only one site. Ten species occurred in all
fifteen sites: Acer rubrum, Carex stipata, Impatiens capensis,
Leersia oryzoides, Ludwigia palustris, Onoclea sensibilis, Salix
sericea, Scirpus cyperinus, Spiraea alba var. latifolia, and S. to-
mentosa (Appendix).
Thirteen species or 5.6% of the total vascular flora were intro-
duced species: Achillea millefolium, Agrostis gigantea, Berberis
thunbergii, Cirsium arvense, Conium maculatum, Echinochloa
crusgalli, Hypericum perforatum, Juncus conglomeratus, Loni-
cera X bella, Myosotis scorpioides, Potentilla norvegica, Rorippa
nasturtium-aquaticum, and Solanum dulcamara. This compared
to 20% for eastern and central North America according to Fer-
nald (1950). Eight of the thirteen introduced species occurred in
only one site. The most common introduced species was Agrostis
gigantea, which occurred in nine of the fifteen sites. Surprisingly,
the site with the most introduced species was Conway 3, the most
remote of all the sites.
A typical beaver-impacted wetland may best be characterized
as a complex mosaic of five physiographic zones, each with its
own characteristic plant community. The precise arrangement of
182 Rhodora [Vol. 102
the zones varied from site to site depending on geomorphology,
beaver activity, and human disturbance
Zone 1, open water. This zone included standing water
above the dam and moving water in the stream channel that per-
sisted throughout the growing season. Areas of deeper water were
dominated by floating-leaved aquatics such as Lemna minor and
Utricularia vulgaris or rooted hydrophytes such as Brasenia
schreberi, Najas flexilis, Nuphar variegata, Polygonum amphi-
bium, and Potamogeton spp. while shallower waters were colo-
nized by emergents such as Sparganium spp., Carex spp., and
Scirpus spp. The stream channel was often vegetated by Pota-
mogeton spp. and Ludwigia palustris.
Zone 2, mud flats. Upstream of the open water, saturated
bottom sediments were exposed through most of the growing sea-
son but were frequently reflooded for short periods. Mud flats
were often dominated by stands of Leersia oryzoides, Lysimachia
terrestris, or Eleocharis acicularis.
Zone 3, wet meadows. Farther upstream, the partially
drained soils of wet meadows occasionally were reflooded during
the growing season. They were most often dominated by species
of Carex, Scirpus, Juncus, Solidago, and Eupatorium as well as
by /mpatiens capensis. In some sites tussock-forming graminoids
such as Calamagrostis canadensis, Carex stricta, Phalaris arun-
dinacea, and Scirpus cyperinus created a hummock-and-hollow
microtopography (McMaster 1989).
Zone 4, drier meadows. Still farther upstream and adjacent
to the site margins were drier meadows, areas of well-drained
soils which rarely were reflooded during the growing season.
Here emergent hydrophytic graminoids such as Carex crinita,
Eleocharis spp., Glyceria canadensis, Juncus effusus, and Typha
latifolia occurred in association with herbaceous species includ-
ing Aster puniceus, Eupatorium maculatum, Galium tinctorium,
and Impatiens capensis. In some sites Typha latifolia formed
large pure stands in this zone.
Zone 5, dry meadows and upland margins. Along the wet-
land/upland boundaries of sites where reflooding seldom oc-
2000] McMaster and McMaster—Flora of Beaver Wetlands 183
Table 4. First and last sampling years, species richness at first sampling,
number of recruitments and extinctions, species richness at last sampling, net
change in species richness, and percent species turnover for eleven study sites.
Species Species
Rich- Re- Rich- Species
Years ness cruit- Extine- ness Net Turn-
Site Sampled First ments tions Last Change over
Ashfield 1 82-92 47 20 22 45 = 63%
Ashfield 2 82-92 41 2 19 34 a | 58%
Ashfield 3 82-93 40 24 19 45 es) 67%
Ashfield 4 84-94 72 18 36 54 = 13 60%
Ashfield 5 84-94 68 12 41 39 =29 66%
Conway | 80-95 57 10 15 a2 = 37%
Conway 2 80-95 44 13 25 ops ={2 67%
Conway 3 80-95 70 16 2 59 a 50%
onway 84-94 ne) 38 23 70 a 1S 66%
Williamsburg 1 84-94 49 19 23 45 =f 62%
Williamsburg 2 84-94 52 22 20 54 mat? 579%
curred, the emergent hydrophytes and herbaceous species found
in Zone 4 mixed with a variety of woody wetland species in-
cluding Spiraea alba var. latifolia, Ilex verticillata, and Alnus
incana. Also occasionally found in this zone were saplings of
woody species common in the surrounding upland, especially
Acer rubrum, Betula alleghaniensis, B. lenta, B. papyrifera, B.
populifolia, and Prunus serotina.
Disturbed areas. Some areas of a site including beaver
dams, lodges, and food caches; stone walls; roadway embank-
ments; and outcrops of bedrock were above high water but were
subject to frequent disturbance from beaver or human activity.
Vegetation in these areas included woody upland species such as
Fagus grandifolia, Fraxinus americana, Pinus strobus, and Tsuga
canadensis, shrubby pasture species such as Juniperus communis
and Berberis thunbergii, and herbaceous species including Co-
nium maculatum, Achillea millefolium, and Solanum dulcamara.
Table 4 lists the number of vascular plant species occurring in
sampled plots over ten-to-fifteen-year intervals between sampling
in eleven sites (the other four sites were not resampled due to
reoccupation). Recruitments exceeded extinctions (i.e. species
richness increased) in three sites over the interval between sam-
pling while extinctions exceeded recruitments in eight sites. All
184 Rhodora [Vol. 102
but one of the eleven sites had a percent turnover of 50% or
more. Mean percent turnover for the eleven sites was 67%.
DISCUSSION
Total vascular plant diversity of the fifteen field sites, 231 taxa,
was relatively high considering the small area (48.7 ha) repre-
sented. In a study of the herb strata of three Connecticut River
oxbows in western Massachusetts, Holland and Burk (1984) iden-
tified 130 taxa of herbs, vines, and woody seedlings over a single
growing season in three sites totaling 64 ha. M. Hickler reported
177 vascular species in a three-year survey of fifteen oxbows of
the Nashua River (Massachusetts) totaling 12.5 ha (pers. comm.).
In a botanical survey of a 162 ha natural area in Franklin County,
McMaster (1987) identified 532 vascular plant species in a single
season.
High plant diversity in the study sites resulted from several
factors. Spatial variability within a site may be attributed to the
steep hydrologic gradient created by the dam and to the topo-
graphical heterogeneity created by beaver artifacts (dams, lodges,
food caches, canals, etc.), human structures (walls, roadways, util-
ity lines, etc.), and geological features (glacial erratics and out-
crops of bedrock). Temporal variability within a site resulted from
beaver activity (building of dams, lodges, and other artifacts;
maintenance of beaver structures; eventual abandonment), human
impacts (destruction of dams, trapping, filling, dredging, burning,
grazing cattle), and hydrological effects (reflooding, draining, and
drought). Invasion of dominant forest species that could shade
out herbaceous species was controlled by the saturated soils over
most of the area of a site. Finally, vascular plant diversity among
the fifteen sites appeared to be enhanced by variations in soil and
water chemistry (McMaster 1997).
The small number of introduced species may be attributed to
two factors. Many of the non-native plants of eastern North
America are associated with agriculture, adapted to dry, disturbed
habitat such as fields or road edges, and are unlikely to germinate
in the saturated soils found in beaver-impacted wetlands. Those
able to germinate as soils drained in late spring or early summer
may not have been able to reach maturity in the short period
before reflooding occurred and conditions again became intoler-
able.
2000] McMaster and McMaster—Flora of Beaver Wetlands 185
Two wetland plant species, Lythrum salicaria and Phragmites
australis, are of particular concern in the northeastern United
States due to their aggressiveness and ability to drive out native
wetland species. Both are common in disturbed areas and appear
to follow major transportation arteries. While L. salicaria is found
in abundance in many wetlands of eastern and central New Eng-
land, it has not yet appeared in any of the study sites. This may
be due in part to elevation, climate, soil type, or to the relative
remoteness of these sites from major highways. Phragmites aus-
tralis was present in only one of the fifteen study sites where it
occurred in a very limited area along an abandoned utility right-
of-way.
Decline in species richness observed in eight sites was pri-
marily among shrubs and trees, probably the result of repeated
beaver occupation and reflooding during the 10—15 year interval
(while all sites were inactive at the onset of the study, at least
one reoccupation occurred in ten of the fifteen sites over the pe-
riod of this study). Increase in species richness observed in the
other three sites was due primarily to recruitment of woody spe-
cies from the surrounding upland or from the seed bank as drain-
ing proceeded in the absence of reoccupation. Vascular plant di-
versity began to decline again in sites unoccupied for more than
twenty years, however, probably due to the loss of herbaceous
species unable to compete with invading shrubs and trees. Ad-
ditional data on changes in species composition, zonation, and
diversity in the fifteen sites over the duration of this study will
be presented in a forthcoming paper.
Most of the vascular plant species found in beaver-impacted
wetlands depend on the presence of saturated soils and full sun,
a rare combination of conditions in New England in the absence
of beaver activity. As beaver population density increases and
available habitat declines, reoccupation of abandoned sites is like-
ly to occur at shorter intervals. Under these conditions plant spe-
cies such as Ophioglossum vulgatum var. pseudopodum that favor
older successional sites may be expected to decline. Active man-
agement of beaver populations may be necessary to promote re-
gional vascular plant diversity. Any anthropogenic activities that
alter hydrology such as draining or filling wetlands, and diverting
or channelizing streams may also have a negative impact on vas-
cular plant diversity in these habitats.
186 Rhodora [Vol. 102
ACKNOWLEDGMENTS. I wish to thank C. John Burk of Smith
College, research advisor to both authors, for his guidance
throughout this project and for his patient review of this manu-
script. I am also indebted to my other dissertation committee
members, Paul Jeff. Godfrey, William A. Patterson III, and Karen
B. Searcy of the University of Massachusetts, Amherst, and the
other members of Nancy D. McMaster’s dissertation committee,
Margaret E. B. Bigelow and David L. Mulcahy of the University
of Massachusetts, Amherst, and Allen H. Curran and Stephen G.
Tilley of Smith College, for their guidance and assistance at many
stages in this project, to Smith College students Alison Coppola,
Marilyn M. Bekech, Jessie Gunnard, and N’Goundo Magassa for
their assistance both in the field and in the laboratory, and to Paul
Somers of the Massachusetts Natural Heritage Program for assis-
tance with county records. Funding for various phases of this
project from the Department of Biological Sciences at Smith Col-
lege, the Biology Department at the University of Massachusetts,
Amherst, the Sigma Xi Scientific Research Society, the B. Eliz-
abeth Horner Research Fellowship at Smith College, the Con-
necticut River Watershed Council, and the Massachusetts Natural
Heritage Program is gratefully acknowledged. This paper is ded-
icated to the memory of Nancy D. McMaster.
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GLEASON, H. A. » A. CRONQUIST. 1991. Manual of Vascular Plants of
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MAGEE, D. W. AND H. E. AHLEs. 1999. Flora of the Northeast: A Manual of
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——. 1989. The floristics and synecology of fifteen Gee ae
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McMaster, R. T. 1994. Ecology, reproductive biology and population ge-
netics of Pes Gea vulgatum (Ophioglossaceae) in Massachusetts.
Rhodot 259-286.
19 ; Weenie reproduction observed in Ophioglossum pusillum
Rafinesque Amer. Fern J. 86: 58—60
— . Floristics, zonation and succession of vascular vegetation in
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188 Rhodora [Vol. 102
APPENDIX
ANNOTATED LIST OF THE VASCULAR PLANTS OF BEAVER WETLANDS IN
FRANKLIN AND HAMPSHIRE COUNTIES, MASSACHUSETTS.
Species name is followed ie habitat preference (Zone | = open water, Zone
2 = mud flats, Zone 3 = wet meadow, Zone 4 = drier meadow, Zone 5
dry meadow and mae margin, or disturbed areas), abundance laine
frequent, aes infrequent, or rare; based on criteria described by Palmer
et al. 1995), and the number of sites in which it occurred (e.g. 4/15 indicates
the species es in 4 of the 15 sites). Introduced species are indicated by
an asterisk (*). Nomenclature follows Gleason and Cronquist (1991)
LYCOPODIOPHYTA (Clubmosses)
SELAGINELLACEAE
Selaginella apoda (L.) Spring — Zone 3, rare, 1/15
EQUISETOPHYTA (Horsetails)
EQUISETACEAE
Equisetum arvense L. — Zone 5, occasional, 6/15
Equisetum sylvaticum L. — Zones 3,4,5, rare, 4/15
POLYPODIOPHYTA (Ferns)
ASPLENIACEAE
Dryopteris carthusiana (Villars) Fuchs — Zones 4,5, rare, 2/15
Thelypteris noveboracensis (L.) Nieuwl. — Zone 5, rare, 1/15
Thelypteris palustris Schott — Zones 3,4,5, frequent, 14/15
DENNSTAEDTIACEAE
Dennstaedtia punctilobula (Michx.) T. Moore — Zone 4, rare, 1/15
ONOCLEACEAE
Onoclea sensibilis L. — Zones 2,3,4,5, frequent, 15/15
OPHIOGLOSSACEAE
Ophioglossum vulgatum L. var. pseudopodum (S. E Blake) Farw. — Zones
, rare, |
OSMUNDACEAE
smunda cinnamomea L. — Zones 3,4,5, rare, 6/15
Donnas claytoniana L. — Zone 4, rare, 1/15
2000] McMaster and McMaster—Flora of Beaver Wetlands
Osmunda regalis L. — Zones 4,5, occasional, 8/15
PINOPHYTA (Gymnosperms)
CUPRESSACEAE
Juniperus communis L. — Zone 5, disturbed areas, rare, 1/15
PINACEAE
Pinus strobus L. — Zones 4,5, disturbed areas, rare, 5/15
Tsuga canadensis (L.) Carriere — Zones 4,5, disturbed areas, rare, 3/15
MAGNOLIOPHYTA (Flowering Plants)
MAGNOLIOPSIDA (Dicots)
ACERACEAE
Acer pensylvanicum L. — Zone 5, rare, 1/15
Acer rubrum L. — Zones 4,5, occasional, 15/15
Acer spicatum Lam. — Zone 5, rare, 2
ANACARDIACEAE
Toxicodendron radicans (L.) Kuntze — Zones 4,5, rare, 2/15
APIACEAE
Cicuta bulbifera L. — Zones 3,4,5, rare, 1/15
*Conium maculatum L. — Zone 4, disturbed areas, rare, 1/15
ydrocotyle americana L. — Zones 2,3,4,5, occasional, 11/15
Stum suave Walter — Zones 1,2,5, rare, 4/15
Zizia aurea (L.) Koch — Zone 3, occasional, 1/15
AQUIFOLIACEAE
flex verticillata (L.) A. Gray — Zones 4,5, frequent, 11/15
Nemopanthus mucronatus (L.) Loes. — Zone 5, rare, 2/15
ARALIACEAE
Aralia nudicaulis L. — Zone 5, rare, 1/15
ASCLEPIADACEAE
Asclepias incarnata L. — Zone 3, infrequent, 1/15
ASTERACEAE
*Achillea millefolium L. — Zone 3, disturbed areas, rare, 1/15
189
190 Rhodora [Vol.
Aster novae-angliae L. — Zone 4, rare, 1/15
Aster puniceus L. — Zones 3,4, occasional, 11/15
Aster racemosus Elliott — Zones 3,4,5, rare, 3/15
Aster umbellatus Miller — Zone 4, rare, 1/15
Bidens cernua L. — Zones 4,5, frequent, 6/15
Bidens connata Muhl. — Zones 2,5, frequent, 6/15
Bidens frondosa L. — Zone 3, rare, 1/15
*Cirsium arvense (L.) Scop. — Tone 3, rare, 1/15
Eupatorium maculatum L. — Zones 2,3,4,5, frequent, 12/15
Eupatorium perfoliatum L. — Zones 2,3,4,5, frequent, 14/15
Eupatorium purpureum L. — Zones 2,3,4, infrequent, 3/15
Euthamia graminifolia . ) Nutt. — Zones 2,4, occasional, 13/15
Senecio aureus L. — Zones 3,4,
Senecio eas Mubl. = Zone o. rare, 1/15
Solidago caesia L. — Zone 5, rare, 1/15
Solidago canadensis L. — Zones 2,3,4, occasional, 11/15
Solidago patula Muhl. — Zones 3,4,5, infrequent, 3/15
Solidago rugosa Miller — Zones 2,4, occasional, 11/15
BALSAMINACEAE
Impatiens capensis Meerb. — Zones 2,3,4,5, frequent, 15/15
BERBERIDACEAE
*Berberis thunbergii DC — Zone 3, disturbed areas, rare, 1/15
BETULACEAE
Alnus incana (L.) Moench — Zones 2,3, aes a 11/15
Alnus serrulata (Aiton) Willd. — Zone 5, 5
Betula ee Britton — Zones 0's, en, TAS
Betula lenta L. — Zones 4,5, infrequent, °
Betula nae Marshall — Zone 5, i aieauedt: 2/15
Betula populifolia Marshall — Zones 4,5, rare, 7/15
BORAGINACEAE
*Myosotis scorpioides L. — Zone 4, rare, 1/15
BRASSICACEAE
Cardamine pensylvanica Muhl. — Zone 1, rare, 2/15
*Rorippa nasturtium-aquaticum (L.) Hayek — Zones 2,3, rare, 1/15
CALLITRICHACEAE
Callitriche palustris L. — Zone 1, rare, 3/15
2000] McMaster and McMaster—Flora of Beaver Wetlands 191
CAPRIFOLIACEAE
*Lonicera X bella Zabel — ee 4,5, rare, 1/15
Lonicera canadensis Marshall — ne 4, rare, 1/15
Sambucus canadensis L. — Ge 7 ,., disturbed areas, rare, 6/15
Viburnum ue Marshall — Zone 5, rare, 1/15
Viburnum dentatum L. var. lucidum Aiton — Zones 2,3,4,5, disturbed areas,
occasional, 9/15
Viburnum lentago L. — Zone 5, rare, 2/15
Viburnum nudum L. var. cassinoides (L.) T. & G. — Zone 5, occasional, 4/15
CLUSIACEAE
Hypericum canadense L. — Zones 2,5, occasional, 4/15
Hypericum ellipticum Hook. — Zones 2,3, occasional, 12/15
Hypericum majus (A. Gray) Britton — Zone 4, rare, 1/15
Hypericum mutilum L. — Zones 2,4, infrequent, 12/15
*Hypericum perforatum L. — Zones 2,3,4, rare, 3/15
Triadenum virginicum (L.) Raf. — Zones 2,3,4,5, frequent, 2/15
CORNACEAE
Cornus amomum Miller — Zones 2,3,4, disturbed areas, occasional, 4/15
DROSERACEAE
Drosera rotundifolia L. — Zones 4,5, infrequent, 7/15
ERICACEAE
Lyonia ligustrina (L.) DC — Zones 3,4,5, infrequent, 2/15
Vaccinium corymbosum L. — Zone 5, occasional, 8/15
FABACEAE
Amphicarpaea bracteata (L.) Fern. — Zone 4, rare, 1/15
Apios americana Medikus — Zone 5, rare, 1/15
FAGACEAE
Fagus grandifolia Ehrh. — Zone 5, Ri 1/15
Quercus rubra L. — Zone 5, rare, 1/15
GENTIANACEAE
Gentiana clausa Raf. — Zones 2,3, infrequent, 1/15
HAMAMELIDACEAE
Hamamelis virginiana L. — Zones 4,5, occasional, 2/15
192 Rhodora [Vol. 102
JUGLANDACEAE
are, 1/15
en
Juglans cinerea L. — Zone 5,
LAMIACEAE
Lycopus americanus Muhl. — Zones 2,4,5, infrequent, 6/15
Lycopus unifolorus Michx. — Zones 2,3,4, occasional, 6/15
Lycopus virginicus L. — Zones 2,4,5, infrequent, 12/15
Mentha arvensis L. — Zone 2, disturbed areas, infrequent, 7/15
Prunella vulgaris L. — Zone 3, rare, 1/15
Scutellaria galericulata L. — Zones 2,3,4, 5/15
Scutellaria lateriflora L. — Zones 3,4,5, rare, wnt
LAURACEAE
Lindera benzoin (L.) Blume — Zone 5, rare, 1/15
LENTIBULARIACEAE
Utricularia vulgaris L. —'Zone 1, occasional, 3/15
MYRICACEAE
Myrica gale L. — Zones 2,3,4,5, occasional, 1/15
NYMPHAEACEAE
Brasenia schreberi J. F Gmelin — Zone 1, rare, 1/15
Nuphar variegata Durand — Zone |, infrequent, 1/15
OLEACEAE
—
Fraxinus americana L. — Zone 5, disturbed areas, rare, 1/15
ONAGRACEAE
Circaea alpina L. — es 3,4, rare, 1/15
Epilobium inne Bichler ~ Zones 3.4, rare, 7/15
Epilobium glandulosum Lehm. — Zone 2, be ae 2/15
Epilobium leptophyllum Raf. — Zone 2, rare, 1/15
Epilobium strictum M Muh ~ Zones 3,4,5, eset TAS
Ludwigia palustris (L.) Elliott — Zone 2, occasional, 15/15
Oenothera fruticosa L. — Zones 4,5, rare, 1/15
OXALIDACEAE
Oxalis acetosella L. — Zones 3,4,5, infrequent, 5/15
2000] McMaster and McMaster—Flora of Beaver Wetlands 193
Oxalis stricta L. — Zones 3,4,5, rare, 2/15
POLYGONACEAE
Polygonum amphibium L. — Zones 1,2, rare, 1/15
Polygonum arifolium L. — Zones 3,4,5, rare, 4/15
Polygonum hydropiper L. — Zones 2,3, infrequent, 9/15
Polygonum pensylvanicum L. — Zone 1, rare
Polygonum sagittatum L. — Zones 2,3,4, disturbed areas, occasional, 13/15
PRIMULACEAE
Lysimachia ciliata L. — Zones 3,4, rare, 1/15
Lysimachia terrestris (L.) BSP. — Zones 2,3,4,5, frequent, 9/15
Trientalis borealis Raf. — Zone 5, rare, 1/15
RANUNCULACEAE
Caltha palustris L. — Zones 2,3,4, occasional, 4/15
Clematis virginiana L. — Zones 3,4,5, occasional, 7/15
Coptis trifolia (L.) Salisb. — Zones 4,5, rare, 2/15
Ranunculus tricophyllus Chaix — Zone |, occasional, 1/15
Thalictrum pubescens Pursh — Zones 1,3,4, disturbed areas, rare, 5/15
ROSACEAE
see Sieh Nie Elliott — Zones 3,4,5, ene 3/15
virginiana F V. Duchesne — Zone 3, 2/15
um eae hia Zones 3,4, rare, 15
Coun rivale L. — Zones 3,4, rare, 2/15
Potentilla fruticosa L. — Zones 3,4,5, rare, 2/15
*Potentilla norvegica L. — Zone 4, rare, 1/15
Potentilla aries (L.) Scop. — Zone 3, rare, 1/15
Potentilla simplex Michx. — Zones 2,3,4,5, rare, 10/15
Prunus pea Ehrh, — Zones 4,5, infrequent, 7/15
Prunus virginiana L. — Zone 4, rare, |
Rubus hispidus L. — Zones 2 ah 5, dicnubed areas, occasional, 13/15
/15
w — Zone 4,
Spee a alba DuRoi var. latifolia pee es Zones 2,3,4,5, disturbed
areas undant, 15/15
Panty tomentosa L. — Zones 2,3,4,5, occasional, 15/15
RUBIACEAE
Cephalanthus occidentalis L. — Zones 4,5, rare, 1/15
Galium asprellum Michx. — Zones 2,3, occasional, 6/15
Galium palustre L. — Zone 2, occasional, 1/15
194 Rhodora [Vol
Galium tinctorium L. — Zones 2,3,4,5, abundant, 14/15
Galium trifidum L. — Zones 2,3,4,5, occasional, 9/15
Galium triflorum Michx. — Zone 5, rare, 1/15
SALICACEAE
Populus balsamifera L. — Zone 3, rare, 1/15
Populus tremuloides Michx. — Zone 5, infrequent, 3/15
Salix bebbiana Sarg. — Zones 3,4,5, rare, 3/15
Salix discolor Muhl. — Zones 2,5, infrequent, 5/15
Salix eriocephala Michx. — Zones 4,5, infrequent, 2/15
Salix lucida Muhl. — Zones 3,4,5, infrequent, 2/15
Salix nigra Marshall — Zone 5, rare, 1/15
Salix sericea Marshall — Zones 3,4,5, occasional, 15/15
SAXIFRAGACEAE
Penthorum sedoides L. — Zone 2, infrequent, 4/15
Saxifraga penysylvanica L. — Zone 4, rare, 1/15
Tiarella cordifolia L. — Zone 2, rare, 1/15
SCROPHULARIACEAE
Chelone glabra L. — Zones 3,4,5, infrequent, 4/15
Mimulus ringens L. — Zones 2,3, infrequent, 11/15
Veronica americana (Raf.) Schwein. — Zones 2,3,4, rare, 2/15
Veronica scutellata L. — Zone 3, rare, 1/15
SOLANACEAE
*Solanum dulcamara L. — Zones 3,4,5, disturbed areas, rare, 3/15
ULMACEAE
Ulmus rubra Muhl. — Zone 4, rare, 1/15
URTICACEAE
Boehmeria cylindrica (L.) Swartz — Zones 2,3,4, infrequent, 1/15
VERBENACEAE
Verbena hastata L. — Zones 2,3,4, rare, 3/15
VIOLACEAE
Viola cucullata Aiton — Zone 5, rare, 1/15
Viola lanceolata L. — Zones 3,4, rare, 2/15
Viola macloskeyi Lloyd — Zones 2,4,5, infrequent, 14/15
. 102
2000] McMaster and McMaster—Flora of Beaver Wetlands 195
VITACEAE
Parthenocissus quinquefolia (L.) Planchon — Zone 3, rare, 1/15
LILIOPSIDA (Monocots)
ALISMATACEAE
Sagittaria latifolia Willd. — Zones 2,4, occasional, 4/15
ARACEAE
Arisaema triphyllum (L.) Schott — Zone 3, rare, 1/15
CYPERACEAE
Carex atlantica L. Bailey — Zones 2,3,4,5, occasional, 11/15
Carex canescens L. — Zones 2,4, pear 2/15
Carex comosa FE Boott — Zones 2,3,4,5, abundant, 8/15
Carex crinita Lam. — Zones 2,4, feanene TAS
Carex debilis var. debilis Michx. — Zones 3,4, rare, 2/15
Carex eburnea F. Boott — Zones 4,5, rare, 1/15
Carex flava L. — Zones 2,4,5, infrequent, 10/15
Carex intumescens Rudge — Zones 2,3, abundant, 4/15
Carex lacustris Willd. — Zones 3,4, occasional, 3/15
Carex lupulina Muhl. — Zone 3, rare
Carex lurida Wahlenb. — Zones 2,3,4, occasional, 14/15
Carex scabrata Schwein. — Zones 3,4, rare 5
Carex scoparia Schkuhr — Zones 2,4, occasional, 11/15
Carex stipata Muhl. — Zones 2,3,4,5, occasional, 5/15
Carex stricta Lam. — Zones 3,4,5, abundant, 13/15
Carex vulpinoidea Michx. — Zones 4,5, occasional, 4/15
Cyperus bipartitus Torr. — Zone 5. rare, 5
Cyperus strigosus L. — Zone 3, rare, 1/15
Dulichium arundinaceum (L.) Britton — Zones 3,4,5, abundant, 9/15
Eleocharis acicularis (L.) Roemer & Schultes — Zones 2,3,5, occasional, 8/15
Eleocharis ovata (Roth) Roemer & Schultes — Zone 2, Gecnent 13/15
Eleocharis palustris L. — Zones 2,5, occasional, 6/15
Eleocharis heii (Willd. ) an — Zones 2,4,5, frequent, 12/15
Rhynchospora capitellata (Michx.) Vahl — Zone 1, rare, 3/15
Scirpus atrovirens Willd. — Zones 2,4, occasional, 5/15
Scirpus cyperinus (L.) Kunth — Zones 2,3,4,5, occasional, 15/15
Scirpus microcarpus C. Presl. — Zones 2,4,5, occasional, 9/15
Scirpus polyphyllus Vahl — Zone 1, rare, 2/15
Scirpus smithii A. Gray — Zones 2,3,4, rare, 5/15
Scirpus validus Vahl — Zones 2,4, infrequent, 8/15
ERIOCAULACEAE
Eriocaulon aquaticum (Hill) Druce — Zone 2, rare, 1/15
196 Rhodora [Vol. 102
IRIDACEAE
Tris versicolor L. — Zones 1,2,3,4, rare, 10/15
Sisyrinchium montanum Greene — Zone 1, rare, 1/15
JUNCACEAE
Juncus brachycephalus (Engelm.) Buchenau — Zones 3,4, rare, 2/15
Juncus brevicaudatus (Engelm.) Fern. — Zones 2,4, occasional, 11/15
Juncus canadensis J, Gay — Zone 4, parry 6/15
* JUNCUS conglomeratus L. — Zone 4, rare, 1/15
Juncus effusus L. — Zones 2,3,4, cea 14/15
Juncus marginatus Rostk. — Zone 1, rare, 1/1
LEMNACEAE
Lemna minor L. — Zone 1, occasional, 6/15
Spirodela polyrhiza (L.) Schleiden — Zone 1, rare, 2/15
LILIACEAE
Lilium canadense L. — Zone 2, rare, 1/15
NAJADACEAE
Najas flexilis (Willd.) Rostkov & Schmidt — Zones 1,2,3, frequent, 2/15
ORCHIDACEAE
Habenaria psycodes (L.) Sprengel var. psycodes — Zone 3, rare, 1/15
Liparis loeselii (L.) Rich. — Zones 2,3, rare, 1/15
POACEAE
*Agrostis gigantea Roth — Zones 2,3,4, frequent, 10/15
Agrostis hyemalis (Walter) BSP. — Zone 3, rare, 1/15
Brachyelytrum erectum (Schreber) Beauv. — Zone 3, rare, 1/15
Calamagrostis canadensis (Michx.) P. Beauv. — Zones 2,3, abundant, 10/15
*Echinochloa crusgalli (L.) P. Beauv. — Zone 5, rare, 1/15
Glyceria canadensis (Michx.) Trin. — Zones 2,3,4,5, occasional, 14/15
Glyceria grandis S. Wats. — Zone 4, rare, 1/15
Glyceria melicaria (Michx.) C. E. Hubbard — Zone 4, rare, 2/15
Glyceria striata (Lam.) A. Hitche. — Zones 2,3,4, infrequent, 9/15
Leersia oryzoides (L.) Swartz — Zones 2,3,4,5, abundant, 15/15
Muhlenbergia mexicana (L.) Trin. — Zones 2,5, infrequent, 4/15
Panicum lanuginosum Elliott — Zones, 3,4, eo 2/15
Panicum rigidulum Nees — Zones 2,3,4, , W/15
Phalaris arundinacea L. — Zones 2,3,4, Eee TNS
Paes australis (Cav.) Trin. — Zone 3, rare, 1/15
2000] McMaster and McMaster—Flora of Beaver Wetlands 197
SPARGANIACEAE
Sparganium americanum Nutt. — Zones 1,2,3,4,5, oe TNS
Sparganium androcladum (Engelm.) Morong — Zones 2,3,5, occasional, 9/15
Sparganium chlorocarpum Rydb. — Zones 2,3,5, aa 6/15
TYPHACEAE
Typha latifolia L. — Zones 2,3,4,5, abundant, 12/15
ZOSTERACEAE
Potamogeton epihydrus Raf. — Zone 1, rare, 3/15
Potamogeton natans L. — Zone 1, rare, 2/15
RHODORA, Vol. 102, No. 910, pp. 198-201, 2000
NEW ENGLAND NOTE
REDISCOVERY OF SYMPHYOTRICHUM ANTICOSTENSE
IN THE UNITED STATES
ARTHUR HAINES
Woodlot Alternatives, Inc., 122 Main Street, Number 3,
Topsham, ME 04086
e-mail: ahaines @ woodlotalt.com
Symphyotrichum anticostense (Fern.) Nesom (synonyms: Aster
anticostensis Fern., A. gaspensis Victorin) was first described by
Merritt Lyndon Fernald from Anticoste Island, Quebec (Fernald
1915). Since its discovery, this species has had a complicated
taxonomic history. It has been considered by various authors to
be a distinct species (under each of the above names), a variety,
or a form unworthy of recognition. In the third case, it has been
placed with Symphyotrichum novi-belgii (L.) Nesom (synonym:
Aster novi-belgii), which it closely resembles. Studies by Brouil-
let and Labrecque (1987) have shown S. anticostense (as Aster)
to be an allopolyploid derivative of S. novi-belgii and S. boreale
(Torr. & Gray) Nesom.
Recent work by Xiang (1994), Xiang and Semple (1996), and
Nesom (1993, 1994) has shown that the genus Aster, as treated
by Fernald (1950) and Gleason and Cronquist (1991), is poly-
phyletic. Some of the plants traditionally considered to be asters
are more closely related to Erigeron, the fleabanes, than to Aster
sensu Stricto. In addition, the genus Solidago is nested within
Aster sensu lato and necessitates name changes (i.e., renaming all
the goldenrods as Aster, or subdividing Aster into smaller, mono-
phyletic groups). Their approach, after carefully weighing the
data, has been to split Aster into a number of small, morpholog-
ically similar groups to create a nomenclatural system that match-
es the phylogeny of these plants.
Identification of Symphyotrichum anticostense is somewhat
problematic because of its intermediate morphology between S.
boreale and S. novi-belgii, and because it is known to hybridize
with the latter. In many respects it resembles a robust S. boreale,
with firm, ascending leaves that barely clasp the stem, erect
branches of the capitulescence, and appressed phyllaries that are
herbaceous at the apex and chartaceous at the base. However,
198
2000] New England Note Loo
unlike S. boreale, it has a thicker stem and rhizome (>2.0 mm
wide) and larger leaves (width and length). Symphyotrichum novi-
belgii is distinguished from both of these species by its phyllaries,
that are usually foliaceous and squarrose, its open-branched cap-
itulescence, and herbaceous to fleshy, clasping leaves. A detailed
description of this species’ morphology, a photo of the type spec-
imen, and key to related species are provided in Labrecque and
Brouillet (1990; as Aster).
Symphyotrichum anticostense is endemic to northeastern North
America, known only from Anticoste Island, the Gaspé, and the
St. John and Aroostook River watersheds (Labrecque and Brouil-
let 1990). It was first collected in the United States by M. L.
Fernald in 1901 in Fort Fairfield, Maine (under the name Aster
junceus Ait.). It was found in circumneutral gravel on the shore
of the Aroostook River, near the U.S.—Canada border. Unfortu-
nately, this site was destroyed by a rise in water level caused by
Tinker Dam in the early part of the century. Though field surveys
have revealed new populations in Quebec and New Brunswick
(Luc Brouillet, pers. comm.), this aster has not been collected
from Maine for nearly a century. It is listed in Flora Conservan-
da: New England (Brumback and Mehrhoff, et al. 1996) as his-
toric and presumed extirpated in the state (SX). Recent reports
of this aster in Maine in Gleason and Cronquist (1991) are based
on annotations of historic material by Labrecque and Brouillet
(1990).
Utilizing natural community and historic range information
gathered from articles (Fernald and Wiegand 1910; Labrecque
and Brouillet 1990) and herbarium labels (at MAINE), I surveyed
sites along river shores on 5 and 6 September 1998 in seven
townships on the St. John and Aroostook Rivers that are known
presently, or were known historically, to possess the appropriate
substrate and/or associated species. Symphyotrichum anticostense
is usually found on well-drained, circumneutral, cobble shores
that are kept open by water and ice scour. It often grows with
other well-known species that favor this habitat, such as Anemone
multifida Poiret, Oxytropis campestris (L.) DC var. johannensis
Fern., and Tanacetum bipinnatum (L.) Shultz-Bip. ssp. huronense
(Nutt.) Breitung (Labrecque and Brouillet 1990). Searches at most
of these areas revealed only common asters, including Doellin-
geria umbellata (P. Mill.) Nees, S. novi-belgii, S. puniceum (L.)
Nesom, and S. lanceolatum (Willd.) Nesom.
200 Rhodora [Vol. 102
On a section of scoured, cobble shore on the Aroostook River
in Caribou, Maine, I located twelve stems of Symphyotrichum
anticostense growing in a 0.5 X 0.5 m area. This site seemed
least likely for discovery of this aster due to lack of circumneutral
plant indicators and its invasion by field species, such as Phalaris
arundinacea L. and S. lanceolatum. Additionally, Prunus pumila
L. var. depressa (Pursh) Gleason, Rosa blanda Ait. var. blanda,
and Salix interior Rowlee were found in the area. The plant’s
identity was confirmed by Luc Brouillet and a specimen deposited
at Herbier Marie-Victorin (MT). This find indicates that additional
riverbank habitats may harbor populations of S. anticostense.
Symphyotrichum anticostense is considered globally imperiled
(ranked G2; Labrecque and Brouillet 1990). Three major threats
to its survival in the northeast are: (1) water flow alteration due
to damming; (2) riverbank disturbance; and (3) crowding by in-
vasive plants. Dams are likely the most significant threat to S.
anticostense. In addition to inundation of the plants, dams de-
crease the effect of spring ice scour by regulating a river’s flow,
thereby allowing woody species to invade the riverbanks. Sym-
phyotrichum anticostense appears to be similar to Pedicularis fur-
bishiae S. Wats. in its need for cyclical river shore disturbance
to remove woody competitors. Riverbank disturbance by motor-
ized vehicle traffic and development on the river shores have
impacted populations of the aster on the Bonaventure River in
Quebec (Labrecque and Brouillet 1990). Symphyotrichum anti-
costense is apparently unable to tolerate crowding by invasive
plants. At the Caribou station, where much of the region is uti-
lized for agricultural purposes, Phalaris arundinacea and S. lan-
ceolatum form a dense cover on the shore. Further survey effort
is needed to determine the full extent of S. anticostense’s presence
in Maine and identify methods of mitigating its threats.
LITERATURE CITED
BROUILLET, L. AND J. LABRECQUE. 1987. Aster gaspensis Victorin: Nombre
chromosomique et hybridation naturelle avec 1’A. novi-belgii. Naturaliste
Canadien 114: 150-165.
BRUMBACK, W. E. AND L. J. MEHRHOFF, in collaboration with R. W. ENSER,
S. C. GAWLER, R. G. Popp, P. SomERS, AND D. D. SPERDUTO, with assis-
tance from W. D. COUNTRYMAN AND C. B. HELLQuIST. 1996. Flora Con-
servanda: New England. The New England Plant Conservation Pr pot
(NEPCoP) list of plants in need of conservation. Rhodora 98: 233-36
2000] New England Note 201
FERNALD, M. L. 1915. Some new or unrecorded Compositae chiefly of north-
eastern North America. Rhodora 17: 1—20.
. 19 Gray’s Manual of Botany, 8th ed. Van Nostrand Reinhold
ee New Yor
——— AND K. M. WIEGAND. 1910. A summer’s botanizing in eastern Maine
and western New Brunswick. Rhodora 12: a —121.
GLEASON, H. A. AND A. CRoNQuisT. 1991. Manual of Vascular Plants of
Narheadten United States and Adjacent phe 2nd ed. The New York
Botanical Garden, Bron pe
LABRECQUE, J. AND L. eoitEe 1990. Aster anticostensis, an endemic of
northeastern North America: Biology and conservation. Rhodora 92: 129—
141
Nesom, G. L. 1993. Taxonomy of Doellingeria (Asteraceae: Astereae). Phy-
tologia 75: 452—462.
——.. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: As-
eeieuey, emphasizing the New World species. Phytologia 76: 141-297.
XIANG, C. S. 1994. Molecular systematic study of Aster sensu lato and related
genera (Asteraceae: Astereae) based on as oplast DNA restriction site
analysis. Ph.D. dissertation, Univ. of Waterloo, Waterloo, ON, Canada.
AND J. C. SEMPLE. 1996. Molecular systematic study of on sensu
lato and related genera (Asteraceae: Astereae) based on chloroplast DNA
restriction site analyses of mainly North American taxa, pp. 393-423. In:
N. Hinds and H. Beetje, eds., Proceedings of the International Com-
positae Conference, Kew, 1994. Vol. 1. Systematics.
RHODORA, Vol. 102, No. 910, pp. 202—207, 2000
NEW ENGLAND NOTE
AEGAGROPILOUS DESMARESTIA ACULEATA FROM
NEW HAMPSHIRE
ARTHUR C. MATHIESON! AND EDWARD J. HEHRE
Department of Plant Biology and Jackson Estuarine Laboratory,
University of New wy Elan Snine, pura, NH 03824
‘e-mail: arthur@hopper.unh
CLINTON J. DAWES
Department of Biology, University of South Florida, Tampa, FL. 33620
Although most seaweeds are anchored solidly by their hold-
fasts, unattached populations are common in calm bays, fjords,
salt marshes, and estuaries throughout the world (Benz et al.
979; Dawes et al. 1985; Josselyn 1977; Orth et al. 1991; Phillips
1961; Zobell 1971). Five somewhat arbitrary and overlapping
categories of unattached seaweeds can be recognized (Norton and
Mathieson 1983): (1) entangled: highly branched and intertwined
plants [e.g., Bonnemaisonia hamifera Har., Gracilaria tikvahiae
McLachlan, and Hypnea musciformis (Wulfen in Jacq.) J. V. La-
mour.] that occur among other drifting seaweeds and may include
multiple plants or taxa; (2) loose-lying: completely unattached
plants such as the saltmarsh fucoids Ascophyllum nodosum (L.)
Le Jol. ecad scorpioides (Hornemann) Reinke and Fucus vesi-
culosus L. ecad volubilis (Hudson) Turner; (3) embedded: plants
that lack a holdfast and are partially buried in sand or mud (e.g.,
Fucus cottonii Wynne et Magne); (4) free-floating: entangled and
drift plants like Sargassum natans (L.) Gaillon; and (5) aegagro-
pilous: spherical masses of radially arranged branches that are
composed of either single or multiple plants held together by
interlocking branches [e.g., Pilayella littoralis (L.) Kjellm. and
Spermothamnion repens (Dillwyn) Rosenv.].
Of the 328 species of unattached seaweeds recorded by Norton
and Mathieson (1983), 146 were loose-lying, 62 free-floating, 58
entangled, 53 aegagropilous, and 9 embedded. The aegagropilous
taxa included 25 red, 18 green, and 10 brown algae. Some of the
best known ball-forming algae are produced by freshwater and
marine species of Cladophora (Hoek 1963; Newton 1950; Sakai
202
2000] New England Note 203
Figure 1. Two aegagropilous specimens of Desmarestia aculeata, with
the sample on the left being more compacted and spherical than the one on
the right; scalar = 5 cm.
1964), with the famous Japanese “‘lake-balls’’ designated as nat-
ural monuments (Kurogi 1980). According to Newton (1950),
Cladophora balls occur sporadically, sometimes cast up in enor-
mous numbers and at other times totally absent. Aegagropilous
species, like most other unattached seaweeds, originate from at-
tached plants, and they usually become infertile and reproduce
entirely by vegetative means (Fritsch 1935, 1945).
The present paper reports the occurrence of two aegagropilous
specimens of Desmarestia aculeata (L.) J. V. Lamour. (Desma-
restiales, Phaeophyceae; Figure 1) that were collected at Concord
Point (43°01'00"N, 70°43’55”W), Rye, New Hampshire on Feb-
ruary 6, 1998. It is a common perennial (cf. Mathieson and Hehre
1982; Taylor 1957) that grows attached to solid substrata within
subtidal environments throughout the eastern (Portugal to Iceland
and Greenland) and western North Atlantic (New Jersey to the
Canadian Maritimes), the North Pacific from Oregon to the Aleu-
tian Islands of Alaska, the Bering Sea, Kurile Islands, and Russia
(Mathieson 1979; Scagel et al. 1986; South and Tittley 1986).
We believe this represents the first record of D. aculeata growing
in an aegagropilous habit, as only free-floating or entangled mas-
204 Rhodora [Vol. 102
ses have been previously reported (Norton and Mathieson 1983).
Drift material of D. aculeata is common in the Gulf of Maine,
where biomass values in excess of 11 kg wet wt. m* may occur
after major winter storms (Mathieson, unpubl. data).
One of the two Desmarestia balls was found within a deep
tidal pool, while the other occurred within a tidal channel that
was closed at one end and open to strong wave action at the
other. The specimens (Figure 1) were approximately the size of
a tennis ball (7.5 X 7.5 cm and 7.5 X 8.5 cm). They were com-
posed primarily of entangled, ‘“‘wiry”’, and very spiny Desmares-
tia fronds, which were flatter than most terete, attached speci-
mens. A diverse assemblage of plants, animals, and shell frag-
ments was associated with the Desmarestia balls: the cord grass
Spartina alterniflora Lois.; the seaweeds Chaetomorpha linum
(O. E Mill.) Kiitz., Rhizoclonium tortuosum (Dillwyn) Kiutz.,
Chondrus crispus Stackh., Polysiphonia fucoides (Huds.) Grev.,
and Prilota serrata Kiitz.; plus the invertebrates Dynamena pum-
ila (L.), Membranipora membranacea (L.), Tubularia sp., and
Mytilus edulis L.
As noted by several investigators (cf. Fritsch 1935, 1945; Gibb
1957; Nakazawa and Abe 1973; Sakai 1964; Yoshida 1963), the
aegagropilous morphology results from a variety of factors, in-
cluding detachment/breakage, oscillating movement of fragment-
ed materials, meristematic injury, scar tissue development, and
extensive regeneration and/or proliferation of new growth. Sub-
sequent rolling and further injury causes pruning, proliferation,
and compaction, resulting in a dense, spherical structure.
In the case of Cladophora, aegagropilous specimens are com-
posed of entangled masses of filaments bound by rhizoids. In old
Cladophora balls the center often decays, leaving a cavity; youn-
ger balls may form several concentric layers. The “beach form”’
of the temperate North Atlantic Ascophyllum nodosum [1.e., either
A. mackaii (Turner) Holmes et Batters or A. nodosum ecad mack-
aii (Turner) Cotton of different investigators] is one of the most
highly modified spherical forms (Gibb 1957; South and Hill
1970). Two temperate aegagropilous seaweeds form extensive
blooms, including the persistent nuisance brown alga Pilayella
littoralis in Massachusetts and the ceramialean “‘‘red tide”’ alga
Spermothamnion repens in southern New England (Wilce et al.
1982). Tropical and subtropical seaweeds form similar structures,
detached Caulerpa racemosa (Forsskal) J. Agardh and Bryotham-
2000] New England Note 205
nion seaforthii (Turner) Kititz. form ball-shaped masses after ex-
posure to gentle water motion near coral reefs and within man-
grove canals (Almodovar and Rehm 1971). Nuisance populations
of lagunal ball-forming Cladophora prolifera (Roth) Kiitz. occur
in Bermuda (Bach and Josselyn 1978)
The crustose coralline red alga Lithothamnion glaciale Kjellm.
may form free-living balls or rhodoliths on sandy or gravelly
substrata in quiet bays in Newfoundland (Hooper 1981). Ice is
instrumental in these habitats as it breaks off the calcareous
crusts, allowing small fragments to roll around and acquire a dis-
tinctive ball-shaped configuration. Extensive populations of rho-
doliths (Lithophyllum and Lithothamnion spp.) also occur in the
Gulf of California (cf. Bosence 1983; Foster and Riosmena-Rod-
riguez 1999; Steller and Foster 1995) where they constitute major
sources of carbonate sediment and habitats of high diversity (Fos-
ter et al. 1997). In contrast to the production of ball-shaped struc-
tures from living, photosynthetic seaweeds, balls are also pro-
duced from dead leaf and rhizome materials of the Mediterranean
seagrass Posidonia oceanica (L.) Delile. In a series of experi-
mental evaluations, Cannon (1979) demonstrated the importance
of oscillating water motion, clumping, compaction, and disinte-
gration of detrital materials in the formation of Posidonia balls.
Apparently, frond detachment in Desmarestia is followed by
injury to its intercalary meristem, resulting in the loss of trichoth-
allic hairs and a lack of proliferations. Its ball-shaped morphology
probably develops because of rolling and compaction of residual
branches, plus the incorporation of ‘‘foreign’’ materials. The re-
tention of detached Desmarestia fragments within deep tide pools
or semi-enclosed tidal channels may provide a vehicle for con-
sistent movement (rolling) and compaction.
ACKNOWLEDGMENT. We are indebted to Ms. Jenna Wanat and
Ms. Kim Mayer who collected the two samples of Desmarestia
balls during field investigations in association with a Marine Bot-
any class in the spring of 1998. The paper is issued as Contri-
bution Number 347 from the Jackson Estuarine Laboratory and
the Center for Marine Biology.
LITERATURE CITED
ALMODOVAR, L. R. AND A. REHM. 1971. Marine algal balls at La Parguera,
Puerto Rico. Nova Hedwigia 21: 255-259.
206 Rhodora [Vol. 102
Bacu, S. D. AND M. N. JOSSELYN. eee Mass blooms of the alga Cladophora
in ese Mar. Pollut. Bull. 9: 34—37.
Benz, M. C., N. J. EISEMAN, AND E. “| GALLAGHER. 1979. Seasonal occur-
rence and variation in standing eg of a drift algal community in the
Indian River, Florida. Bot. Mar. 22: 413-420.
BosENCE, D. W. J. 1983. Description aa classification of rhodoliths (rho-
doids, rhodolites), pp. 217-224. In: M. Peryt, ed., Coated Grains.
Springer-Verlag, Berlin.
CANNON, J. E M. 1979. An experimental investigation of Posidonia balls.
Aquatic mae 6: 407—410
Dawes, C. J., M. O. HALL, AND R. REICHERT. 1985. Seasonal biomass and
energy content in seagrass communities on the west coast of Florida. J.
Coastal Res. 1: 255-262.
Foster, M. S. AND R. RIOSMENA-RODRIGUEZ. 1999. Rhodolith beds in the
Gulf of California. J. Phycol. Sed ar 35: 10-11.
RIOSMENA-RODRIGUEZ, D. L. STELLER, AND W. J. WOELKERLING.
1997, Living rhodolith beds in the Gulf of California and their implica-
tions for paleoenvironmental interpretations, pp. 127—140. In: John-
son and J. Ledezma-Vazguez, eds., Pliocene Carbonates and Related Fa-
cies Flanking the Gulf of California, Pale California Sur. Mexico. The
Geological Society of America, Boulder, CO
FrirscH, EK E. 1935. The Structure and Reproduction of the Algae, Vol. I.
sae Univ. Press, Cambridge, aaa
. 1945. ibid., Vol. Il. Cambridge Univ. Press, Cambridge, Engl
Gipp, D. . 1957. The free-living forms a. Ascophyllum nodosum cee Le
Sie J. Ecol. 45: 49-83.
, C. VAN DEN. 1963. Revision of the European Species of Cladophora.
Bull Leiden, The Netherlands.
Hooper, R. 1981. Recovery of Newfoundland benthic marine communities
from sea ice, pp. 360-366. /n: G. E. Fogg and W. E. Jon
ceedings of the VIII International Seaweed Symposium, Ba, angor, North
les. Mar. Sci. Lab., Menai Bridge, University College, Anglesey, North
Wales, U.K.
JossELYN, M. N. 1977. Seasonal changes in the distribution and growth of
Laurencia ae — Ceramiales) in a subtropical lagoon.
Aquatic Bot. —229.
KuRoaI, M. 1980. ane ball “‘Marimo”’ in Lake Akan. Jap. J. Phycol. 28:
168-169.
MATHIESON, A. C. 1979. Vertical distribution and longevity of subtidal sea-
weeds in northern New England, U.S.A. Bot. Mar. 30: 511-520.
AND E. J. HEHRE. 1982. The composition, seasonal occurrence, and
reproductive periodicity of the marine Phaeophyceae in New Hampshire.
Rhodora 84: 411—437.
NAKAZAWA, S. AND M. Abe. 1973. Artificial globing of algae. Bull. Jap. Soc.
Phycol. 21: 53-56
Newron, L. M. 1950. A beach-ball mystery at Torbay. Illustrated London
News 216: 98.
Norton, T. A. AND A. C. MATHIESON. 1983. The biology of unattached sea-
weeds, pp. 333-386. /n: E E. Round and D. J. Chapman, eds., Progress
2000] New England Note pLNe|
in Phycological Research, Vol. 2. Elsevier Scientific Publ. Co., Amster-
OrtH, R. J., K. L. Heck, Jr., AND R. J. Diaz. 1991. Littoral and intertidal
systems in the mid- jee coast of the United States, pp. 193-214. In:
. Mathieson and P. H. Nienhuis, eds., Ecosystems of the World, Vol.
24: Intertidal and Littoral ee Elsevier Scientific Publ. Co., Am-
sterdam, The Netherlan
PHILLIPS, R. C. 1961. cee aspect of the marine algal flora of St. Lucie
Inlet and adjacent Indian River, Florida. J. Florida Acad. Sci. 24: 135-
SAKAI, Y. 1964. The species of Cladopora from Japan and its vicinity. Sci.
Pap. Inst. Algol. Res. Fac. Sci. Hokkaido Imp. Univ. 5: 1-104.
SCAGEL, R. FE, D. J. GARBARY, L. GOLDEN, AND M. W. HAwkKEs. 1986. A
Synopsis of the Benthic Marine Algae of British Columbia, Northern
Washington and Southeast Alaska. fe aoeccae Contribution No. 1. Dept.
of Botany, Univ. British Columbia, Vanco nada.
SOUTH, : R. AND R. D. Hie. 1970. cinieee on marine ie of Newfound-
land. I. Occurrence and distribution of free-living Ascophyllum nodosum
in are Canad. J. Bot. 48: 1697-1701.
AND I. TrrrLtey. 1986. A Checklist and Distributional Index of the
Benthic Algae of the North Atlantic Ocean. Spec. Publ. Huntsman Mar.
Lab. and British Museum (Natural Hist.), St. Andrews, NB, Canada and
London, caeee
STELLER, D. L. AN . S. Foster. 1995. Environmental factors influencing
distribution oad norpholos of rhodoliths in Bahia Concepcion, B.C.S.,
Mexico. J. Exp. r. Biol. Ecol. 194: 201-212.
TAYLOR, W. R. 1957. The Marine Algae of the oe Coast of North
America. Univ. Michigan Press, Ann Arbor
WILCE, R. T., C. W. SCHNEIDER, A. V. QUINLAN, AND K. VAN DEN BOSCH.
1982. The life history and morphology of free-living Pilayella littoralis
(L.) Kjellm. tg ia a Ectocarpales) in Nahant Bay, Massachusetts
Phycologia 21: 336-354.
YOsHIDA, T. 1963. Studies on the distribution and drift of the floating sea-
eds. Bull. Tohoku Reg. Fish Res. Lab. 23: 141-I§
ZOBELL, Cc. E. 1971. Drift seaweeds on San Diego County beaches, pp. 269-—
In: W. J. North, ed., The Biology of Giant Kelp Beds (Macrocystis)
in California. Nova Hedwigia Beih., Verlag von J. Cramer, German
RHODORA, Vol. 102, No. 910, pp. 208-209, 2000
NEW ENGLAND NOTE
TWO MORE WEEDS IN MAINE
PETER F. ZIKA
Herbarium, Dept. of Botany, Box 355325,
University of Washington, Seattle, WA 98195-5325
e-mail: zikap@aol.com
Senecio viscosus L. (Asteraceae) was discovered in Maine in
August 1969 by Frank Seymour on a field trip of the Josselyn
Botanical Society. His specimen (Seymour 27556 v7) is from a
roadside at the foot of Boarstone Mt., Eliotville, Piscataquis Co.
(Seymour 1982). This collection was overlooked by recent au-
thors in Maine (e.g., Campbell et al. 1995; Haines and Vining
1998). Senecio viscosus was found again in Maine in July 1999,
as a weed at both ends of a culvert on Route 162, where McLean
Brook enters the southwestern arm of Long Lake (Zika 1392]
NEBC, WTU). The site is dry, gravelly, and sunny, at an elevation
of 180 m, in T17 R4 WELS, Aroostook Co. About 100 plants
were seen, growing with Plantago major L. Senecio viscosus 1s
a European native, known as an occasional adventive in Nova
Scotia and in the northeastern United States.
A second addition to the flora of Maine is Corynephorus ca-
nescens (L.) Beauv. (Poaceae), collected in July 1999 as a rare
weed in a cultivated cranberry crop on Route 1A about 2 km east
of the Millbridge town line (Zika 13946 NEBC). The inconspicuous
plants were in sunny, moist, and sandy soil, at an elevation of 12
meters, in the town of Harrington, Washington Co. Angelo and
Boufford (1998) recorded this European grass from southeastern
Massachusetts, but nowhere else in New England. The species is
similar to Festuca ovina L. in general habit, and perhaps is over-
looked.
LITERATURE CITED
ANGELO, R. AND D. E, Bourrorb. 1998. Atlas of the flora of New England:
Poaceae. Rhodora 100: 101—233.
CAMPBELL, C. S. et al. 1995. Checklist of the Vascular Plants of Maine, 3rd
rev. Bull. 844, Maine Agricultural and Forest Experiment Station, Univ.
Maine, pee ME.
HAINES, A. AND T. E VINING. 1998. Flora of Maine, a Manual for Identifi-
208
2000] New England Note 209
cation of Native one Naturalized Vascular Plants of Maine. V. K Thomas
, Bar Harbor,
Sheu PA, 1982. ae Flora of New England, 2nd ed. Phytologia Mem-
oirs V, Plainfield, NJ.
RHODORA, Vol. 102, No. 910, pp. 210-213, 2000
NEW ENGLAND NOTE
NEW RECORDS FOR SCIRPUS ANCISTROCHAETUS IN
NEW HAMPSHIRE
JOSHUA L. ROYTE
The Nature Conservancy, 14 Maine Street, Suite 401,
Brunswick, ME 04011
e-mail: jroyte@tnc.org
JOHN P. LORTIE
Woodlot Alternatives, Inc., 122 Main Street, Topsham, ME 04086
In 1992, the New Hampshire Natural Heritage Inventory
(NHNHI) procured a Section 2 grant from the U.S. Fish and
Wildlife Service (USFWS) to perform de novo inventories for
northeastern bulrush (Scirpus ancistrochaetus Schuyler) in New
Hampshire. While the site location for this bulrush was in Ver-
mont, across the Connecticut River from Walpole, New Hamp-
shire (see Schuyler 1962), it had not been found at that time in
New Hampshire. Northeastern bulrush was listed by USFWS as
endangered in 1991 and a recovery plan was prepared and signed
in 1993. Listing was based on the lack of protection for nearly
all of the known sites and the high degree of threat from imminent
development pressure (United States Fish and Wildlife Service
1991). The Natural Heritage Program lists this species as globally
rare (G3) since its discovery at over 50 sites along the Appala-
chians between western Virginia and southwest New Hampshire
(A. E. Schuyler pers. comm. 1996). It is listed as S1 (critically
imperiled with five or fewer occurrences) in four states, S2 (im-
periled with six to 20 occurrences) in three states, extirpated (SX)
in One state, and as a Division | species in the Flora Conservanda
(Brumback and Mehrhoff, et al. 1996).
The NHNHI hired Woodlot Alternatives, Inc. to perform
searches for northeastern bulrush during three-to-four days each
in 1992, 1993, and 1994. Aerial photographs were reviewed be-
fore initiating searches in the towns of Acworth, Charlestown,
Langdon, and Unity in Sullivan County, and the towns of Alstead,
Gilsum, Surry, and Walpole in Cheshire County, New Hampshire.
Fifty-four sites were selected out of 125 potential sites and seven
210
2000] New England Note 214
of these contained populations of northeastern bulrush. A sample
from the first population found was verified by A. E. Schuyler in
the field the following year. Each population was found growing
in wetlands where natural water levels had been altered by either
beaver or human-caused draining of small ponds by culverts and
beaver dam removal.
In 1992 likely sites along the Connecticut River were visited
by canoe but no new populations were found there, although it
was re-located at the type locality originally described by Schuy-
ler. Other sites were visited on foot in 1992 and the following
two years. When an area was found to have the appropriate hab-
itat (open graminoid/sedge swales, evidence of fluctuating water
levels, and bulrushes) the wetland was searched intensively for
Scirpus ancistrochaetus.
Scirpus ancistrochaetus was found in wetlands with the fol-
lowing characteristics: saturated to slightly inundated (to 18 in.
deep) emergent benches found next to slightly deeper emergent
zones (e.g. areas too shallow for floating-leaf emergent plant spe-
cies, and too deep for woody plants); fluctuating water levels
(stagnant water levels allow succession to shrub and forested nat-
ural communities); associated plant species included S. cyperinus
(L.) Kunth, §. atrovirens Willd., Leersia oryzoides (L.) Swartz,
Schoenoplectis tabernaemontani (C. C. Gmel.) Palla, and Spar-
ganium androcladum (Engelm.) Morong, which was almost al-
ways found close by in an “‘off shore” zone.
The most useful field characteristics for identification of the
species were: (1) drooping, glomerular fruiting heads (these are
similar looking to the fruiting heads of Schoenoplectis tabernae-
montani—drooping heads are supported by curved inflorescence
rays that rarely project upwards or straight out, as the inflores-
cence of Scirpus cyperinus may, and as S. atrocinctus Fern. and
S. hattorianus Makino almost always do); (2) dark, chocolate-
brown florets (S. cyperinus has tawny brown florets); and (3)
broad bracts (close to 3/4 in. wide, while S. cyperinus has narrow
acuminate bracts).
Once identified, the number of fruiting heads and the number
of vegetative shoots per clone were counted in small patches of
plants (less than 100), and estimated from counts of subsamples
in large patches of plants. Collections of Scirpus ancistrochaetus
were made if more than 20 fruiting culms were present, otherwise
portions of the inflorescence were sampled for a voucher speci-
pi Rhodora [Vol. 102
men and photographs were taken. Voucher specimens have been
deposited in the Hodgdon Herbarium (NHA) at the University of
New Hampshire in Durham. Wetland communities were de-
scribed using The Nature Conservancy Eastern Heritage Task
Force 1991 Site Survey Summary and Special Plant Forms, and
using Natural Community Forms developed by NHNHI. Inven-
tories of associated plants and their relative abundance were re-
corded for the field forms. In addition, the population’s location
in relation to present and former water levels, topography, and
juxtaposition to other vegetation zones was mapped and/or de-
scribed. All data forms and site specific information reside at
NHNHI in Concord, New Hampshire and with the USFWS in
Hadley, Massachusetts.
SPECIMEN CITATION: NEW HAMPSHIRE: Sullivan Co., Langdon, (elev. ca. 235
m), 22 foliose culms and 7 fruiting stems, growing on the edge of a breached
headwater beaver pond in four clonal clumps, 9 Oct 1992, Royte s.n. (NHA);
Charlestown, western bay of a pond, 119 fruiting stems and 150 leafy shoots
were found in five patches, 18 Aug 1993, Royte, von Oettingen, Schuyler &
Schuyler s.n. (NHA); (elev. 235 m) a large beaver flowage with beaver-im-
pounded meadows in two tributaries (elev. 244 m and 250 m) there were a
total of 400+ foliose shoots and 250 fruiting stems, found growing with
Scirpus cyperinus, Leersia oryzoides, and — arundinaceum, 19 Aug
1993, Royte s.n. (NHA); aes on the west side road, (elev. 260 m), 115
fruiting stems and 75 foliose shoots were found growing on the wetland edge
2—5 cm above a large emergent zone dominated by Sparganium androcladum,
19 Aug 1993, Royte s.n. (NHA); headwater wetland (elev. ca. 260 m), 23
foliose culms and 15 fruiting stems were found in two clumps in two areas,
th clumps were isolated islands near larger island clumps of Scirpus cy-
perinus and Leersia oryzoides, the water depth of 38-61 cm appeared to be
higher than normal, 15 Aug 1994, Royte s.n. (NHA); wetland, along the north-
ern shore of a beaver Sneath (elev. ca. 319 m), 40—SO foliose culms and 23
oo stems in an emergent bench of Leersia oryzoides with 38—46 cm of
ater, 19 Aug 1994, Royte s.n. (NHA)
LITERATURE CITED
BRUMBACK, W. E. AND L. J. MEHRHOFF, in collaboration with R. W. ENSER,
S. C. GAWLER, R. G. Popp, P. SOMERS, AND D. D. SPERDUTO, with assis-
tance pa W. D. COUNTRYMAN AND C. B. HELLQuIST. 1996. Flora Con-
servanda: New England. The New England Plant Conservation Program
(NEPCoP) ae of plants in need of conservation. Rhodora 98: 223-361.
2000] New England Note pA Fe.
SCHUYLER, A. E. 1962. A new species of Scirpus in the northeastern United
States. Rhodora 64: 43-49.
UNITED STATES FISH AND WILDLIFE SERVICE. 1991. Endangered and threatened
wildlife and plants; determination of endangered status for Scirpus ancis-
trochaetus (northeastern bulrush). Federal Register 56: 21091—21096.
RHODORA, Vol. 102, No. 910, pp. 214-216, 2000
NOTE
SAGINA (CARYOPHYLLACEAE) IN ILLINOIS:
GORDON C. TUCKER
Department of Biological Sciences,
Eastern Illinois University, Charleston, IL 61920
e-mail: cfgct@eiu.edu
The genus Sagina L. is circumboreal and was most recently
monographed by Crow (1978). Only one species, §. decumbens,
was noted by Mohlenbrock and Ladd (1978) and Mohlenbrock
(1986) from Illinois until S$. procumbens was reported from the
Chicago area (Swink and Wilhelm 1994). Examination of speci-
mens in Illinois herbaria revealed an additional adventive species
in the state, S. japonica. Also, the occurrence of S. apetala in
Illinois noted by Crow (1978), seems to have been overlooked
by recent workers on the Illinois flora.
KEY TO SAGINA SPECIES IN ILLINOIS
|. Flower parts in 4s (rarely 5s on the same plant); sepals reflexed
in fruit; matted wiry perennials, spreading by offshoots
Uh Eas RE eh eeenewade Pee eeeeau eng es S. procumbens
|. Flower parts primarily in 5s (rarely 4s on the same plant);
sepals erect or appressed in fruit; annuals with erect-as-
cending (or decumbent) often capillary stems and slender
taproots, not strongly tufted, not spreading by offshoots
(
Caer we ie er Se a aR ee oes You Sues feat Suse, Rae YOR ON coe as Se SR er Sr Se ee Toe STR Tae a alr GT a ee
NoN
iy
Q
pay)
<
oO
n
=
Se
=a
QO
g.
=r
ie)
2)
=
Sy
jee)
”
io)
nH
iS
—
~
iS
=
S
=~
g
» MseAVeS-2labTOUS Al DASS 2.2 cctv bu waves deus eeeeu ars
3. Seeds pale brown, triangular with a dorsal groove; cap-
sules longer than broad; pedicels and sepals glan-
dular or glabrous; leaves not succulent ......
Fh Se gme Bae oo bee eee Sea oer ee S. decumbens
3. Seeds dark brown to black, plump, ellipsoid-ovoid, lack-
ing a dorsal groove; capsules globose; pedicels and
sepals glandular; leaves succulent .... $. japonica
214
2000] Note 215
NOTES ON THE SPECIES
Sagina apetala Ard. This species was first recorded from IlIli-
nois (Crow 1978) as follows: Union Co., no date, Forbes s.n.
(MICH). I have not located duplicates in any of the herbaria I have
checked. Crow also cited this European species from New Jersey,
Maryland, and Lousiana, as well as Washington, Oregon, and
California. Cronquist (Gleason and Cronquist 1991) did not in-
clude S. apetala in the revised manual, apparently unaware of
Crow’s records from the Manual range.
Sagina decumbens (Sw.) Ohwi. Our only native species was
noted from 25 counties by Mohlenbrock and Ladd (1978). The
following specimens appear to be additional county records: Ef-
fingham Co., Wildcat Hollow NE of Mason, 6 Jun 1967, Evers
90778 (ILLS); Washington Co., Nashville, 2 Jun 1990, Shildneck
16361 (ISM).
Sagina japonica (Sw.) Ohwi. The widespread occurrence of
this eastern Asian species in the northeastern states was noted by
Mitchell and Tucker (1991) and Mitchell (1993). In the Midwest,
it has only been reported from northwestern Ohio (Rabeler 1996).
The following specimen appears to be the first Ilinois record:
Sangamon Co., near Oak Ridge Cemetery, damp soil in grass,
rare, 21 Sep 1951, Lola Carter 15667 (1sM). In 1997, I noted this
species in Charleston, Coles County in a limited area covering
several dozen blocks, between the Eastern Illinois University
Campus and the county courthouse approximately one km to the
north. Searches of herbaria (noted below) and field work in east-
ern and central Illinois have not turned up any additional records.
Specimens: Coles Co., Charleston, 1049 11th St., 28 May 1997,
Tucker 11273 (€1U); 1400 block of 7th St., 14 Jun 1997, Tucker
11275 (EIU, ILLS, ISM); 7th St. and Buchanan Ave., 24 Jun 1999,
Tucker 11731 (EIU).
Sagina procumbens L. This native of Europe (Crow 1978) is
widely naturalized in the northeastern states. It was first noted in
Illinois from the Chicago area by Swink and Wilhelm (1994).
The Morton Arboretum has specimens from Cook and Kane
counties. The following record suggests it may be more widely
distributed: Peoria Co., Peoria, 200 block North Garfield St.,
growing with moss in brick sidewalk cracks, 15 Jun 1955, Chase
14252 (EIU; ILL det. R. K. Rabeler, 1993).
216 Rhodora [Vol. 102
ACKNOWLEDGMENTS. _ I thank the curators of the following her-
baria for lending specimens or providing access to collections
under their care: Hlinois State Museum; [linois Natural History
Survey; Morton Arboretum; University of Illinois at Urbana-
Champaign. I also thank Richard Rabeler for providing infor-
mation on Illinois specimens of Sagina in the University of Mich-
igan Herbarium.
LITERATURE CITED
Crow, G. E. 1978. A taxonomic revision of Sagina (Caryophyllaceae) in
North caer Rhodora 80: 1-91
GLEASON, H. A. AND A. CRONQuUIST. 1991. Manual of Vascular Plants of
Northeastern United States and Adjacent Canada. The New York Botanical
Garden, Bronx
MircHELL, R. S. 1993. Portulacaceae through Caryophyllaceae of New York
State. New York State Museum Bulletin 486. Albany, NY.
D C. Tucker. 1991. Sagina japonica (Sw) Ohwi (Caryophyl-
laceae), an Overlooked adventive in the northeastern United States. Rho-
ora 93: 192-194,
MOHLENBROCK, R. H. 1986. Guide to the Vascular Flora of Illinois, revised
and enlarged ed. Southern Illinois Univ. Press, Carbondale, IL.
AND D. M. Lapp. 1978. Distribution of Illinois Vascular Plants.
Southern Illinois Univ. Press, Carbondale, IL.
RABELER, R. K. 1996. Sagina japonica (Caryophyllaceae) in the Great Lakes
Region. Michigan Bot. 35: 43-44.
SWINK, E AND G. WILHELM. 1994. Plants of the Chicago Region, 4th ed.
Indiana Academy of Science, Indianapolis, IN
RHODORA, Vol. 102, No. 910, pp. 217-224, 2000
NOTE
NOTES ON THE LENTIBULARIACEAE IN BOLIVIA: A
NEW GENUS RECORD (GENLISEA) FOR THE COUNTRY,
WITH TWO ADDITIONAL SPECIES RECORDS IN THE
GENUS UTRICULARIA
Nur P. RITTER! AND GARRETT E. CROW
Department of Plant Biology, University of New Hampshire,
Durham, NH 03824
‘e-mail: npr@cisunix.unh.edu
As part of our investigations in the wetlands of Parque Na-
cional Noel Kempff Mercado in eastern Bolivia, we have en-
countered two species of Lentibulariaceae that were previously
not known for the country’s flora: Utricularia nana A. St.-Hil. &
Girard and Genlisea guianensis N. E. Brown. The latter consti-
tutes the first record of this genus in Bolivia. Furthermore, while
studying specimens at the Missouri Botanical Garden (MO), we
encountered an unidentified specimen of Utricularia that had
been collected from a large inselberg (granitic outcropping) just
outside of the western border of Parque Noel Kempff. We were
able to determine this specimen as U. oliveriana Steyermark, a
third member of the Lentibulariaceae that had not previously been
known for Bolivia.
The presence of Genlisea guianensis and Utricularia nana in
eastern Bolivia represents only a small extension of their previ-
ously known distribution, as both are known to occur in the near-
by state of Mato Grosso, Brazil (Taylor 1989). On the other hand,
the population of U. oliveriana at Cerro Pelao represents an im-
pressive disjunction from other known populations.
Parque Nacional Noel Kempff Mercado (Figure 1) is situated
in the northeastern corner of the Department of Santa Cruz, in
the Province of Velasco. As currently delineated, the park encom-
passes an area of approximately 15,300 km?’ (Killeen and Schu-
lenberg 1999). Genlisea guianensis and Utricularia nana were
encountered growing in seeps in a small clearwater stream on top
of the Serrania de Huanchaca, a massive, steep-sided plateau sit-
uated along the eastern border of the park. The stream was fairly
narrow (1—2 m), widening in a few areas to form small, still
pools, with numerous wet seepy habitats present along the edges.
210
218 Rhodora [Vol. 102
Brazil
Parque Nacional
Noel Kempff Mercado
e Parque Nacional Noel Kempff Mercado. A: Cerro Pelao. B:
The stream on the Meseta. (Redrawn from Killeen and Schulenberg 1999)
Typical species in the seeps were U. amethystina Salzmann ex
A. St.-Hil. & Girard and U. pusilla Vahl, and small, ephemeral,
semi-aquatic herbs, such as Polygala microspora Blake (Poly-
galaceae) and Burmannia flava Mart. (Burmanniaceae).
Cerro Pelao (Figure 1) is a large inselberg located just outside
the western border of the park (Killeen 1996), in the Reserva
Forestal Bajo Paragua. As with some areas of the Meseta, large
expanses of exposed rock (lajas) are present. Where small sea-
sonal streams and springs flow over these outcroppings, hydro-
philic species in the genera Urricularia and Rhynchospora, and
the family Eriocaulaceae can become established on the rocks and
in the shallow sand and sediments that accumulate in pockets in
the lajas. It was in this type of habitat that the third new record,
U. oliveriana, was observed.
Genlisea guianensis is an erect herb with a rosette of elongate,
2000] Note 219
strap-shaped leaves and purple flowers. This species is considered
to be relatively large for the genus (Taylor 1991), attaining
heights up to about 30 cm (Brown 1900). Prior to its discovery
in Bolivia, G. guianensis was thought to be limited to Venezuela,
Guyana, and Brazil (Fromm-Trinta 1984; Taylor 1991).
Although superficially quite similar to Utricularia, Genlisea
can be clearly differentiated on the basis of sepal number and
trap morphology. The calyx is composed of five sepals, as com-
pared to the typical 2 sepals in Utricularia (Cook 1990; Taylor
1991), while the traps are forked and possess two helically twist-
ed branches, in contrast to the globose and bladder-like traps of
Utricularia (Cook 1990; Reut 1993; Taylor 1991). Although the
traps have long been thought to be involved in some form of
carnivory, their function in trapping protozoa was only recently
identified by Barthlott et al. (1998), who demonstrated that the
traps lured protozoa through a chemical attractant.
The fruits of Genlisea are capsular, with the capsules of some
species said to be unique among flowering plants in possessing
an unusual circumscissile dehiscence that ruptures along three
different planes (Taylor 1991). In describing this pattern of de-
hiscence, Taylor (1991) likened the capsule to a globe and de-
picted the planes of dehiscence as occurring not only at the equa-
tor, but also at two additional latitudes between the equator and
one pole.
Utricularia nana is typically a diminutive herb, although the
species is quite variable, ranging in height from 1.5 to 12.0 cm
(Taylor 1989). All individuals we observed were extremely small
(2.0—2.5 cm tall). We had never before encountered a Utricularia
of such small stature, and it wasn’t until we were able to ascertain
that what at first appeared to be grains of sand trapped among
the base of the plant were actually miniscule traps, that we be-
came convinced that this was, indeed, a species of Utricularia.
According to Taylor (1989), U. nana has a fairly wide distribution
in South America, and was previously known from Venezuela,
Guyana, Surinam, French Guiana, Brazil, and Paraguay.
Utricularia oliveriana is a small rheophytic, apparently peren-
nial herb (Taylor 1989). This species is extremely similar to U.
neottioides A. St.-Hil.—another rheophyte that is frequently en-
countered in streams and in water flowing over expanses of rock
on the Meseta. These two species—which are the sole members
of Section Avisicaria Kamiénski—are most easily distinguished
Table 1. Species of Utricularia known for Bolivia. Species names in parenthesis are as given by Foster (1958).
Parque Noel
Species Foster (1958) Taylor (1989) Kempff M.
U. alpina Jacq. a Me
U. amethystina Salzmann ex A. St.-Hil. & Girard (U. velascoénsis O. Ktze.) + + +
U. breviscapa Wright ex Grisebach + +
U. cornuta Michx. (misidentified: almost certainly U. meyeri Pilg.) a3
U. erectiflora A. St.-Hil. & Girard a5
U. foliosa L. + +
U. gibba L. (U. obtusa Sw.) + + +
U. cf. guyanensis A. DC +
U. hispida Lam +
U. hydrocarpa Vahl +
U. lloydii Merl +
U. meyeri Pil + +
U. myriocista ae - -Hil. & Girard + +
U. nana A. +
U. retoides A. a -Hil. (U. Herzogii Liitzelberg) + + +
U. a G. Weber ex Benj. + +
U. ea. Steyermark +
U. poconensis Fromm-Trinta +
U. pusilla Vah + + =f
U. simulans Pilger + +
O7@7c
vlopouYy
JOA]
col
Table 1. Continued.
Parque Noel
Species Foster (1958) Taylor (1989) Kempff M
U. subulata ks + +
U. ricophyla Spruce ex Oliver (U. globulariaefolia Mart. ex Benj.) +
U. tricolor A. St.-Hil. + + +
U. triloba ex Mart. +
U. unifolia Ruiz & Pav6én + +
U. warmingii Kamiénski +
L000
ION
[Gc
a7. Rhodora [Vol. 102
on the basis of leaf shape, with U. oliveriana possessing tiny (2—
8 mm total length) leaves with obovate simple laminae, while the
leaves of U. neottioides are finely divided into pinnately arranged
capillary segments and range in length from a few mm to several
cm (Taylor 1989). The two species also differ in stature, with U.
oliveriana characteristically possessing shorter inflorescences and
thinner rhizoids than U. neottioides. Dimensions of the specimens
from Cerro Pelao were slightly smaller than the lower limits listed
for the species by Taylor (1989), with inflorescences from 12—17
mm in height (vs. 2 cm, Taylor), and leaves scarcely reaching 2
mm in length. Utricularia oliveriana was previously thought to
be restricted to the Guyana Highland region, with populations
known from Venezuela, Colombia, and Brazil (Taylor 1989).
To date, the most comprehensive floristic account of Bolivia is
the checklist published over 40 years ago by Foster (1958). In
his checklist, Foster listed nine species of Utricularia for Bolivia.
In contrast, based on distributional information included by Tay-
lor (1989) in his monograph of Utricularia, 22 of the 214 species
that he recognized world-wide are known from Bolivia. With the
additions of U. nana and U. oliveriana, 14 species of Utricularia
are now known for Parque Noel Kempff Mercado, with a pro-
visional fifteenth species awaiting confirmation (Table 1). Ac-
cording to Taylor (1989) 70 species occur within the entirety of
South America, therefore, this one small corner of Bolivia con-
tains one fifth of the continent’s species of Utricularia. Based on
Taylor’s (1989) monograph, augmented by these new records,
there are now 24 (possibly 25) species of Utricularia known for
Bolivia (Table |). Therefore, greater than half (58%) of the coun-
try’s Utricularia species are now known to occur in Parque Na-
cional Noel Kempff Mercado. Furthermore, this level of diversity
exceeds the number of species known for a number of other Neo-
tropical countries, such as Panama (13 species; D’Arcy 1987),
Peru (12 species; Brako and Zarucchi 1993), Ecuador (11 species;
Jgrgensen and Le6n-Yanez 1999), and Costa Rica (10 species;
Crow 1992).
EXSICCATAE
Genlisea sebecladlaale N. E. Brown. sae Nacional Noel Kempff Mercado.
6
‘La Meseta’’; east of Los Fierros. Elev. ca. 760 m. Semi-aquatic, emergent
herb. Giatine along the edges of the stream. Common. Corolla purple-blue.
2000] Note 223
Fruits present. The stream bottom varies from exposed bedrock to sand. Nu-
merous small pools are present. Samar Been Cerrado. 16 Au
1996, N. Ritter, G. Crow, M. Garvizu, M. Ritter & J. Crow 3614. (MO, NHA,
USZ).
Utricularia nana A. St.-Hil. & Girard. Parque Nacional Noel Kempff Mer-
cado. ‘‘La Meseta’’; east of Los Fierros. Elev. ca. 760 m. Diminutive herb.
Growing in small seeps along the edges of the stream. Only a small number
of individuals were noted. Corolla yellow; subtended by red sepals. The
stream bottom varies from exposed bedrock to sand. Numerous small pools
are present. sacar re ae Cerrado. 16 Aug. eae N. Ritter, G.
Crow, M. Garvizu, M. Ritter & J. Crow 3600. (MO, NHA, US
Utricularia aes ees Santa Cruz, Provincia Velasco. Parque
Nacional Noel Kempff Mercado. Cerro Pelao. Bosque Seco con Talisia, Di-
lodendron, Amburana, Hymenaea, Anadenanthera, Chorisisa, Luehea, Me-
trodorea, Rhamnidium, Sebastiana, Spondias, Astronium, Aspidosperma .. .
Substrato con poco suelo sobre roca sence 14°32'23" S 61°29'53” W.
300m. Hierba; sobre roca himeda, inclinada. | Apr 1994, A. Jardim, with
Saldias, Guillen. Ramos, Jensen, & Surubt 484 (MO, USZ).
ACKNOWLEDGMENTS. We are grateful to the New York Botan-
ical Garden and the Missouri Botanical Garden for providing ad-
ditional herbarium specimens of South American Genlisea for
study, with further thanks due to the latter for making their fa-
cilities available to us during our time in St. Louis. We also wish
to thank Peter Jorgensen, Susana Le6n-Yanez, and the Missouri
Botanical Garden for providing us with an electronic copy of their
checklist of Ecuadorian plants. Additionally, Timothy Killeen of
Museo Noel Kempff Mercado in Santa Cruz, Bolivia deserves
special thanks for his assistance in facilitating our fieldwork in
the Park and for his suggestions regarding potential study areas.
Appreciation is also due to all those individuals who assisted us
with the fieldwork: Marisol Garvizu, Juan Surubi, Martha Ritter,
and Jason Crow. Additional appreciation is due to the following:
the park guards and administrators of Parque Nacional Noel
Kempff, the Direcciédn General de Biodiversidad, the Servicio
Nacional del Areas Protegidas (SERNAP), and the Vice-Minis-
terio del Areas Protegidas. This research was supported in part
by a grant from the Vice President for Research and Public Ser-
vice, University of New Hampshire. This paper is Scientific Con-
tribution Number 2037 from the New Hampshire Agricultural
Experiment Station.
LITERATURE CITED
BARTHLOTT, W., S. POREMBSKI, E. FISCHER, AND B. GEMMEL. 1998. First pro-
tozoa-trapping plant found. Nature 392: 447
224 Rhodora [Vol. 102
BRAKO, L. AND J. L. ZARUCCHI. 1993. Catalogue of the cra elas Plants and
Gymnosperms of Peru. Missouri Botanical Garden, St. Louis, MO.
Brown, N. E. 1900. Genlisea guianensis. Hooker’s eae Plantarum. Fourth
Series. Vol. VII—Part I. Plate 2629. Arthur Felix, Leipzig.
Cook, C. D. K. 1990. ah Plant Book. SPB Academic Publishing, The
, The Netherlan
ae. °G. E. 1992. The genus Urricularia (Lentibulariaceae) in Costa Rica.
Brenesia 38: I-18.
D’Arcy, W. G. 1987. Flora of Panama: Checklist and Index. Part I: The
Introduction and Checklist. Missouri Botanical Garden, St. Louis, MO.
Foster, R. C, 1958. A Catalogue of the Ferns and sed toa Plants of Bo-
livia. The Gray Herbarium of Harvard Univ., Cambridge, M
FROMM-TRINTA, E. 1984. Genliseas eet. Sellowia 36: 55-62
J@RGENSEN, P. M. AND S. LEON-YANEZ, eds. 1999 eat of the Vascular
Plants ‘ Ecuador. Missouri Botanical Garden, St. Loui
KILLEEN, T. J. 1996. Historia natural y biodiversidad - Paraue Nacional
“Noel Kempff Mercado’, Santa Cruz, Bolivia. Plan de Manejo, Com-
ponente Cientifico. Museo re Historia Natural Noel ea Mercado and
Missouri Botanical Garden, Santa Cruz de la Sie
— AND T. S. SCHULENBERG. 1999. A Rapid Biological Assessment of the
Parque Nacional Noel Kempff Mercado, Bolivia. RAP Working Paper
#10, Univ. Chicago, IL.
Reut, M. S. 1993. Trap structure of the carnivorous plant Genlisea (Lenti-
bulariaceae). Bot. Helv. 103: 101-111.
TAYLOR, P. 1989. The Genus Urricularia—A Taxonomic Monograph. Kew,
tee ene Gardens on.
———.. 199]. The Genus Genlisea Carniv. Pl. Newslett. 20: 20-59.
RHODORA, Vol. 102, No. 910, pp. 225-226, 2000
BOOK REVIEW
A Guide to the Algae of New England as Reported in the Liter-
ature from 1829-1984, Parts I and II (in 2 volumes), by
LeBaron C. Colt, Jr. 1999. vi+1019 pp. maps. $100.00 plus
shipping (softbound). Available from L. C. Colt, 61 Philip
St., Medfield, MA 02052.
The two volume compilation by Barry Colt represents an ex-
haustive and important documentation of New England’s algal
flora, covering both microalgae and seaweeds. The value of such
work is priceless, providing critical information regarding histor-
ical records, biodiversity patterns, biogeographic comparisons,
potential environmental impacts, etc. The compendium represents
a “labor of love”? from a very talented and committed phycolo-
gist. Its dedication to Hannah T. Croasdale and the late Gerald
W. Prescott is particularly fitting, as they both produced analo-
gous and exhaustive publications on freshwater microalgae. Hav-
ing followed the author’s progress for more than a decade I can
only imagine the difficulties, frustrations, effort, and many, many
years involved! Certainly he is to be commended for finalizing
such a major synopsis on New England algae.
The two-volume compendium contains a general Introduction
(3 pages) describing the total project, plus four other major sec-
tions. The first section (the New England Region, 3 pages) de-
scribes the area, including all of the counties where collections
have been made; some special geographical areas (e.g. Narragan-
sett Bay, Rhode Island) are designated when individual counties
are not easily ascribed. Generally the format follows that of a
vascular plant flora, with listings of algal taxa given by counties
and a standardized format utilized for authors, dates, etc. Two
maps describe specific site identifications and locations within the
region. The second section (The Algae of New England 1829-—
1984, 775 pages) gives a detailed listing of all known algae (..e.
freshwater and marine microalgae, plus seaweeds) from New
England during this 155 year period; it is by far the largest part
of the compendium, being approximately three-quarters of the
entire text. A standardized format is used for the listings of algal
taxa, being initially arranged alphabetically by genus and then by
descending hierarchy (i.e. species, variety, forma, or other taxo-
nomic levels as appropriate). Collection data for each taxon is
229
N
i)
On
Rhodora [Vol. 102
also reported alphabetically by state and then chronologically by
date of publication. A series of fourteen examples is given, using
a uniform set of abbreviations for different states and counties.
A third section (Authors and Contributors, 43 pages) gives a syn-
opsis of author name(s), plus dates of publication(s) and pages
on which the listed species are reported. Three levels of author
citations are given: (1) individual, (2) coauthored, and (3) con-
tributed material cited by other authors; all of these listings ate
arranged chronologically according to publication dates. The
fourth section (Literature Cited, 175 pages) gives an exhaustive
synopsis of supporting literature that is arranged alphabetically.
It is also annotated to provide a variety of specific information:
(1) site(s) of collections (i.e. state, county, or specific geograph-
ical area); (2) habitat (brackish, freshwater, marine, and terrestri-
al); (3) the presence of maps; (4) the presence or absence of
descriptive materials; (5) the occurrence of figures and plates; and
(6) the numbers of genera, species, varieties, forma, etc.
Barry Colt has synthesized an exhaustive set of information,
providing direct citations to diverse taxa, authors, and the poten-
tials for detailed cross-referencing. I’ve already found it very
helpful in identifying several references and geographical data;
the volumes also provide a logical tool for diverse searches, etc.
While I am genuinely impressed with the author’s efforts, no
work of such a magnitude can be finalized without minor errors.
For example, some typos and grammatical errors are evident
within the text, including a few sentences that are incomplete or
have mixed tenses. I assume that the term alpha really means
alphabetical, but it is not clarified in several places. A few of the
descriptive sections seem to have been run-on, while some hand-
written parts also seem to be present. Lastly, the second part (i.e.
Volume IT) might be improved by a brief transitional paragraph
showing its context and interrelationships to the first part (Volume
I). No doubt these minor points can be rectified if there are future
revisions or updates of literature. In summary, my few construc-
tive comments should in no sense take away from the importance
and value of this compendium. Congratulations to the author for
a fine job!
—ARTHUR C. MATHIESON, Professor of Plant Biology, Jackson
Estuarine Laboratory, University of New Hampshire, Durham,
NH, 03824
RHODORA, Vol. 102, No. 910, pp. 227-229, 2000
BOOK REVIEW
Thoreau’s Country: Journey Through a Transformed Landscape,
by David R. Foster. 1999. xiv + 270 pp. illustrations, bib-
liographic essay, index. ISBN 0-674-88645-3 $27.95 (hard-
cover). Harvard University Press, Cambridge, MA.
If you have ever crossed paths with a lichen-encrusted stone
wall or a lonely cellar hole on a walk through a New England
forest, you may have wondered how such things came to be in
the middle of the woods. Who built them, and why? New Eng-
land’s stone walls have maintained a constant presence across the
land over the centuries, but what changes have they witnessed in
their lifetimes? How have the forests changed’? How has the hu-
man impact on the landscape changed?
Such are the questions that can be answered by combining the
observations of a nineteenth century naturalist with those of a
twentieth century ecologist. In Thoreau’s Country: Journey
Through a Transformed Landscape, David R. Foster provides us
with a window through which we can look back on New Eng-
land’s past. This window is actually the eye of one of New Eng-
land’s most keen 19th century observers, Henry David Thoreau.
Though Thoreau is probably most well known for his philosoph-
ical writings, his daily journals afford an invaluable source of
insight on daily life and nature in the mid-1800s. Foster, Director
of the Harvard Forest since 1990, masterfully weaves Thoreau’s
observations into a tapestry that illustrates the origins of today’s
forests. Through his thoughtful introductions to each chapter, Fos-
ter makes a convincing case that it is necessary to know the
cultural and natural history of New England in order to more
fully understand the present day ecology of our own landscape.
Foster has tastefully selected and organized hundreds of Tho-
reau’s journal entries into chapters that illustrate many aspects of
the mid-1800s, when New England was at its peak in agricultural
production. In Thoreau’s time, more than 60 percent of the land
in southern New England had been cleared and tamed, while the
remaining sections were constantly subject to the axe and saw to
supply an increasing demand for firewood and timber. Foster uses
Thoreau’s talent for imagery and detail to conjure up a striking
picture of the 19th century countryside: pockets of isolated wood-
lands in a matrix of cultivated fields, meadows, and pastures,
221
N
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Rhodora [Vol. 102
much the opposite of what we see today. The land was kept in
this unnatural state by the endless toil of the New England farm-
ers, who Thoreau described as heroic in their year-round work to
improve their land and their livelihoods. The book contains a
wealth of information about 19th century land use practices,
which are ultimately responsible for determining the destiny of
the forests that followed.
The reader will be amazed to discover that Thoreau, through
his years of very detailed observations, was able to describe mod-
ern ecological concepts years before scientists would actually
publish them. It was Thoreau who coined the term “‘forest suc-
cession”’ after he watched different species of trees reclaim the
land that was increasingly being abandoned by farmers as they
moved west or to the cities. He also recognized that each kind of
tree had a situation in which it grew best, in essence that each
species had its own niche. Foster points out that if 20th century
ecologists had paid more attention to the works of Thoreau, much
time and effort might have been saved in the rediscovery of these
concepts. The fruitless attempts to grow white pine on clear-cut
pine stands in the early part of this century could have been
avoided if anyone had studied Thoreau; he anticipated that the
dominance of white pine in New England would be a direct result
of 19th century land use practices, and that hardwoods naturally
recruit underneath the pines.
Although the changes that brought about the forests of today
were well under way by the mid-1800s, the landscape was a
vastly open one compared to today, and the communities of or-
ganisms were very different. Thoreau’s journal entries are in-
valuable in that they describe an ecosystem that is completely
outside of the experience of anyone living in our time. In his
journals, Thoreau romanticized about the songs of bobolinks and
meadowlarks, but bewailed the loss of the “‘noble’’ animals such
as deer and bear. Today the situation is reversed, and we are
concerned with preserving the diversity of the once common field
species, which are declining at an alarming rate. Thoreau also
observed the passenger pigeon and the American chestnut ful-
filling their original ecological roles. Today the pigeon is extinct
and the chestnut has been reduced to a shadow of its former
stature by the chestnut blight.
Thoreau’s Country: Journey Through a Changing Landscape
would be of great interest to both the beginning naturalist and the
2000] Book Review 229
experienced ecologist. Detailed and often romantic accounts of
19th century life and nature are combined with ecological prin-
ciples and thoughtful reflections in such a way that readers gain
a new appreciation for the hard life of their ancestors and for
their ancestors’ legacy, the woods through which we walk today.
The book is well organized and enjoyable to read, complete with
beautiful line drawings that depict life in a time gone by.
Having read the book, one can find new meaning in the forest
landscape and be able to picture where it may have come from.
You will find that the wall you discovered on your walk was
most likely built from stones removed from a plowed field, and
positioned to divide a pasture on the hill from the fertile soils in
the lowland. The stones for the wall were relocated by the strain
and sweat of a farmer who once lived in the house that sat upon
the mossy foundation. The old oak tree with the spreading lower
branches, another witness to more than a century of change, prob-
ably once shaded his livestock in the sweltering heat of a summer
day. The other trees are the products or a predictable and ongoing
succession of species that started when the farmer and his family
abandoned their New England home to seek gold in California.
You will also realize that as you stand on the same spot as the
old farmer and look upon the same wall, you are just taking
another snapshot, observing an instant in the continuous story of
natural change that shapes and forms our reality.
—LAUREN E Howarb, Department of Plant Biology, University
of New Hampshire, Durham, NH 03824.
RHODORA, Vol. 102, No. 910, pp. 230-236, 2000
NEBC MEETING NEWS
November 1999. Bruce A. Sorrie, former botanist for the Mas-
sachusetts Natural Heritage and Endangered Species Program and
now a botanical consultant in the southeastern United States,
spoke on the topic, “Diversity and endemism in the Coastal Plain
Flora.”’ Sorrie defined the coastal plain as the exposed portion of
the continental shelf that extends from Cape Cod, Massachusetts
to a portion of eastern Mexico and northward into the area known
as the Mississippi embayment. It is an area composed of Creta-
ceous age and younger deposits, which are mostly oceanic but
augmented by materials derived from the older adjacent physio-
graphic provinces. Its inland boundary is defined by “‘the Fall
Line,’ where one encounters rocks of Paleozoic age. *“The coast-
al plain occupies about 8% of the North American landmass,”’ he
said. The geologic boundaries of the coastal plain match the
boundaries of what Sorrie considers to be the Coastal Plain Flo-
ristic Province. To put the coastal plain flora in perspective, Sorrie
compared it to the Appalachian Floristic Province, a much older
area geologically, and one regarded as a refuge for plants during
periods of widespread inland seas and global climate change.
While it has long been considered a major center of evolution
from which most eastern North American species evolved, the
Appalachian Floristic Province, Sorrie points out, has only seven
endemic genera: Cymophyllus, Galax, Rugelia, Diamorpha, Am-
phianthus, Jamesianthus, Nestronia, and Rugelia. Many other
genera often thought of as endemic to the Appalachian Province,
e.g., Astilbe, Disporum, Jeffersonia, and Menziesia, are actually
ret: -Tertiary disjuncts with species also occurring in eastern
Asia or elsewhere. Sorrie also mentioned a number of genera
centered in the Appalachians which have spread well beyond the
borders of the Province, e.g., Chamaelirium, Clintonia, Epigaea,
and Liriodendron. He estimated that there might be about 200—
300 endemic species in the Province, but he has not seen a figure
on this,
Using a quote from the late Alwyn Gentry, Sorrie explained
that the southeastern coastal plain is ‘‘a conspicuous but often
overlooked center of endemism in temperate North America.”
Gentry said, ‘It is remarkable that Florida, only 152,000 km? and
with virtually no topographic relief, should rank second only to
California in number of endemic species; it is even more re-
230
2000] NEBC Meeting News 2o1
markable when we consider that the endemic plant species are
concentrated in northern and central Florida, not in the subtropical
southern part.’’ According to Sorrie, there are 215 species wholly
confined to Florida, and another hundred or so that extend but a
short distance from its borders into Alabama and/or Georgia. For
the coastal plain as a whole, there are two endemic plant families,
48 endemic genera, 35 of which are monotypic, and about 1400
endemic species, he said. Endemism occurs in many other coastal
plain plant genera, 98 of which have five or more endemic spe-
cies, he added. He felt that 60 million years of partial exposure
of the coastal plain had allowed for considerable in situ plant
colonization and evolution of new taxa. Sorrie’s slides illustrated
many of the endemic genera. Among them were: Balduina, Cer-
atiola, Dicerandra, Franklinia, Harperocallis, Lachnanthes, Ma-
cranthera, Pinckneya, Pyxidanthera, Schwalbea, Sclerolepis, Sto-
kesia, Warea, and Zenobia.
Sorrie joked about the seemingly monotonous, pine-dominated
landscape of the coastal plain. He quoted Roland Harper, a pio-
neering botanist in the southeastern coastal plain, who described
a 700-mile train trip from Augusta, Georgia to Richmond, Vir-
ginia, where he did “‘not remember seeing any rocks, bluffs, es-
carpments, hills, ravines, gullies, springs, or hammocks, or pass-
ing through any railroad cuts deep enough to obstruct the view.”’
Sorrie commented that some topograpic maps for eastern North
Carolina even lack topographic contour lines! Why then, he rhe-
torically asked, does the coastal plain support such botanical di-
versity? Answering his own question, he gave seven possible rea-
sons: (1) subtle shifts in soil composition and chemistry with
eight of ten global soil orders represented; (2) subtle shifts in soil
moisture; (3) subtle elevational differences that have profound
effects on plant communities; (4) high humidity and percentage
of sunshine; (5) the highest frequency of lightning strikes in the
U.S., which results in many fire-adapted communities with high
herb diversity; (6) up to 60 million years of vegetational history
that has provided, at least, some localized refuges for temperate
species during times of maximum glacial advance; and (7) the
derivation of the flora from multiple source areas, including the
tropics, subtropics, prairies and deserts, as well as from in situ
speciation. A summary of ten different geographic patterns of
floristic endemism in the coastal plain, followed by some ques-
tions from the audience, ended the meeting.
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Rhodora [Vol. 102
December 1999. Dr. Leila Shultz, Research Associate Professor
at Utah State University, presented a lecture entitled “Breaking
new ground in floristics: Using geographic information systems
to predict species distributions in western North America.” For
the neophytes on Utah geography and flora, she started by de-
scribing 4—6 floristic provinces of Utah, the exact number de-
pending on one’s interpretation. The Colorado Plateaus define the
southeastern portion of the state while the southwestern corner is
considered an eastern extension of the Mojave Desert floristic
province. A western fifth of the state consists of the Great Basin
(formerly occupied by the pleistocene Lake Bonneville) and 35
associated mountain ranges including the calcareous Wasatch
Mountains that form the Basin’s eastern border. The northwestern
corner has mountains of igneous origin and a flora influenced by
migrations from the Pacific Northwest. This leaves one or more
provinces in the east that include the Uinta Mountains, with flo-
ristic affinities to the Rocky Mountains; the Uinta Basin, consid-
ered by some to be part of the Colorado Plateaus; and the La Sal
Mountains along the border with Colorado. The Utah flora in-
cludes 2602 native and 682 introduced species, and these num-
bers are increasing due to new discoveries and introductions. Be-
tween 1974 and 1994, 88 new species were described from Utah,
a number of them by Shultz herself. Also, newly naturalized in-
troductions have contributed about 100 new taxa to the flora since
1987, she said. It is a state where 10—15% of the flora is consid-
ered endemic and about 250 species have been proposed for fed-
eral listing.
Dr. Shultz described the collaborative efforts between herself,
Martha Aiken, and other researchers at Utah State University to
develop and test a geographic information system (GIS) for flo-
ristic data that would have the capability of predicting new lo-
cations for the state’s rare plants. A first step toward this end was
to create a rare species specimen database from which geographic
coordinates could be extracted. A rare species appendix to the
Atlas of Vascular Plants of Utah published in 1988 by Albee,
Shultz, and Goodrich helped with this effort. Herbarium speci-
mens that could be mapped at a 10 X 10 km scale or finer were
selected and digitized, so that each mapped species represented a
data layer in the GIS. The predictive modeling research was
largely that of Aiken who completed a Master’s thesis entitled
“Predictive modeling of rare plant habitat in the eastern Great
2000] NEBC Meeting News 235
Basin,”’ a project funded by the Bureau of Land Management and
the Hill Air Force Base. A field key was developed from envi-
ronmental attributes and associated species data collected at 467
site plots. Approximately 20% were presence plots for rare spe-
cies. Four rare plant species were selected for their representation
of different kinds of habitats: Sphaeralcea caespitosa (valley &
foothill sites), Penstemon concinnus (pinyon-juniper woodland),
Primula domensis (faces of dolomite cliffs), and Jamesia tetra-
petala (granite canyons). New data layers with site-specific data
were then added to the baseline information provided by the
coarse grid-distributions provided by the Atlas. Additional data
for the GIS models came from four existing geographic databas-
es: one elevational, both state and national soil databases, and a
surficial geology database. Probability of occurrence maps were
then developed from the GIS data containing 13 environmental
variables encompassing slope, elevation, aspect, soil, and geolog-
ic data.
The predictive model used a tree-classification system to sort
data using binary recursive partitioning. The attribute data for
each variable were examined sequentially to identify the optimal
partition resulting in the most homogeneity within classes and the
most heterogeneity between classes. The procedure was repeated
for each branch of the key. The result was a dichotomous key
that was then incorporated into a computer program for extrap-
olation of the classification over large areas. The dichotomous
key produced in S-Plus was written as a series of conditional
statements for GRID, such that each variable in the model was
represented by a unique grid coverage. GRID is a cell-based geo-
processing software that is integrated with ARC/INFO. As GRID
reads the conditional statement, each grid cell is analyzed and
simultaneously a new grid is generated in which each cell reflects
the predictions of the terminal leaves of the conditional statement.
The new grid is then converted to polygon coverage and the pre-
dictions are mapped using ARCPLOT.
Models were evaluated for total percentage of correct predic-
tions and analyzed using two statistical tests for utility and bias.
Both field-based and GIS-based models performed well for all
four species of plants tested. For the GIS, based on 12 different
models, mean accuracy was 97% for all predictions; for the Field
Key, based on 16 models, the mean for correct predictions ex-
ceeded 95%. The models with the highest utility and lowest bias
234 Rhodora [Vol. 102
used elevation and aspect in predicting distributions. Over-pre-
diction occurred for all species but was considered less of a prob-
lem than under-prediction.
The presentation included habitat pictures for a number of rare
species from the vast remote areas of Utah. Most of the species
shown were discovered and described in the 1970s—80s. Although
the rate of new discoveries has declined, a respectable number of
new finds occurred in the 1990s, demonstrating a need for con-
tinued botanical exploration in remote areas of the intermountain
west. Shultz emphasized the importance of using separate fields
for spatially explicit data (e.g., latitude and longitude) in herbar-
ium databases, thus providing a means for transporting floristic
data to geographic information systems. She encouraged the em-
ployment of different spatial scales depending on the data source,
e., 10 km grids for the generalized localities provided by most
herbarium collections and | km grids for records with latitude
and longitude given in seconds. Databases developed from site-
intensive studies such as those used in the Utah predictive model
can serve the dual role of providing floristic information for her-
barium vouchers and ecological data for mathematical models
that investigate the relationship of plant distributions to climate
and ecology.
January 2000. The first program of the year 2000 was titled
‘“‘First Friday Foray into Fantastic Flora,’ better known as the
annual “‘show and tell,’’ where members are invited to make short
presentations that typically involve showing and narrating a small
number of slide images. As exemplified by first presenter, Donald
Lubin, however, slides are not a necessary prerequisite. Don ex-
plained that he had prepared 62 laminated fronds of fern taxa and
would have them available for examination after the meeting. He
also spoke of fern exploration with Ray Abair that resulted in
three wood fern hybrid taxa being discovered in the Blue Hills
south of Boston and the verification of 39 pteridophyte taxa at
Wachusett Mountain in Worcester County, including eight that
had not been reported previously. Lisa Standley started the slides
with images from the Okavango Delta in Botswana. We were
shown a relatively flat landscape with enormous wetlands that
resemble, according to Lisa, marshes of Manitoba. Here she saw
many familiar genera such as Typha, Phragmites, Nymphaea,
Eleocharis, and Scirpus, but mixed with them stems of Papyrus.
No
es)
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2000] NEBC Meeting News
Several trees from upland habitat were featured including the sau-
sage tree, Kigelia (with bat-pollinated orange flowers and large,
sausage-shaped fruits), and baobabs, all large trees up to 10-12
ft. in diameter at their bases. Not seeing any immature baobabs,
Lisa expressed concern about whether or not they were repro-
sues Close-ups of African large game, including one showing
a group of side-by-side lionesses in crouched position lapping
water, ended the brief glimpse of Africa.
Nancy Eyster-Smith brought us back to the U.S. for a look at
vegetation management activities witnessed in national parks on
a family trip across country this past summer. At Glacier National
Park, she saw propagated native species being planted along
walkways as part of a revegetation project, and new metal board-
walks that had been installed to prevent further trampling near
Logan Pass. She also observed sites where exotic taxa had been
spot-sprayed with herbicides. At Little Bighorn Battlefield Na-
tional Memorial, people were seen pulling an invasive species of
Hypericum by hand. Jumping to the Caribbean, Richard Falcona
illustrated some arid landscapes and scenic views from the island
of St. John in the U.S. Virgin Islands. Plant taxa shown were
turk’s cap cactus, Melocactus intortus, and century plant, Agave
missionum, both native to the island. Paul Somers illustrated a
few nonindigenous species encountered on a trip to the islands
of Nevis and St. Kitts. Examples shown were Momordica char-
antia (Cucurbitaceae) and Calotropis procera (Asclepiadaceae),
both indigenous to the Old World tropics, and cashew trees, An-
acardium occidentale, a native of northern South America. Paul
also showed a few shots of wetland plants taken in Massachusetts,
including Utricularia cornuta and U. inflata from Plymouth
County, Potamogeton ogdenii from Berkshire County, and a pos-
sible new record of Lycopodiella alopecuroides from northern
Worcester County. Sticking with the Massachusetts theme, Pam
Weatherbee illustrated some habitats and plants encountered dur-
ing a biological survey of the Hop Brook Wildlife Management
Area in southern Berkshire County. Despite a long history of land
utilization, Pam reported finding some relatively natural wetlands
with species such as /ris versicolor, Galium palustre, Salix can-
dida, and Salix serissima; forest communities containing an in-
teresting association of Quercus bicolor and Carpinus carolini-
ana; and even a couple of rare plant species. Also catching Pam’s
eye during the survey was a beautiful Baltimore checkerspot, a
236 Rhodora [Vol. 102
butterfly species thought to be switching from Chelone to Plan-
tago as a food plant.
Andy Finton then took us across the Berkshires to the Hudson
River Valley of New York for a presentation on plant community
inventory work recently completed there by himself and col-
leagues at the New York Natural Heritage Program. We learned
about remnant serpentine barrens on Staten Island; oak-dominated
forests with heath understories in the river valley and rocky sum-
mit communities; and beech-maple and spruce-dominated old
growth forests in the Catskills, where one conifer swamp yielded
a black gum aged at 485 yrs. Other communities highlighted were
calcareous cliff communities with calciphile ferns in Albany and
Greene Counties and bog, sedge meadow, and spruce flat com-
munities of the Rensselaer Plateau.
The closing presentation was by George Newman who visually
transported us to the Gaspé Peninsula for a preview of sites to
be visited and things to do during the NEBC summer field trip
in July, 2000. George emphasized the extensive serpentine bar-
rens above timberline on Mont-Albert and the many calciphiles
that could be found in sea cliffs around Mont Ste. Pierre. At
Forillon National Park, options of boating to watch sea lions and
seals or botanizing the talus slopes of Cap-Bon-Ami were offered
as enticements. In the Percé vicinity, exploring calcareous con-
glomerate formations of Mont Ste-Anne, sea cliffs occupied by
gannets and puffins on Ile Bonaventure, or limestone river beds
of Grand Riviere were presented as interesting options.
—PAUL Somers, 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
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do not justify the right margin. Do not indicate the style of type
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TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital
letters. Omit authors of scientific names. Below title, include au-
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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). Cite several refer-
ences alphabetically by first author, rather than chronologically. With-
in 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
237
238 INFORMATION FOR CONTRIBUTORS
distribution. Specimen citations should include collector(s) and col-
lection number in italics or underscored, and herbarium acronym in
capital letters. Keys and synonymy for systematic revisions should
be prepared in the style of ““A Monograph of the Genus Malvas-
trum,” S. R. Hill, RHODORA 84: 159-264, 1982. Designation of a
new taxon should carry a Latin diagnosis (rather than a full Latin
description), 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
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author’s surname. Do not write authors’ names in all capital letters.
References by a single author precede multi-authored works of same
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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
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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 102nd 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 subscrip-
tion 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 eres
For the next calendar year _
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Special interests (optional):
THE NEW ENGLAND BOTANICAL CLUB
Elected Officers and Council Members for 1999-2000:
President: David S. Conant, Department of Natural Sciences,
Lyndon State College, Lyndonville, VT 05851
Vice-President (and Program Chair): Lisa A. Standley, Vanasse
Hangen Brustlin, Inc., 101 Walnut St., P.O. Box 9151, Wa-
tertown, MA 02272
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, Massachusetts College of Liberal Arts, North Adams,
MA 01247-4100
Recording Secretary: Paul Somers
Curator of Vascular Plants: Raymond Angelo
Assistant Curator of Vascular Plants: Pamela B. Weatherbee
Curator of Nonvascular Plants: Anna M. Reid
Librarian: Leslie J. Mehrhoff
Councillors: W. Donald Hudson, Jr. (Past President)
Arthur V. Gilman 2000
Karen B. Searcy 2001
David Lovejoy 2002
Jennifer Forman (Graduate Student Member) 2000
Appointed Councillors:
David E. Boufford, Associate Curator
Janet R. Sullivan, Editor-in-Chief, Rhodora
RHODORA
Journal of the
New England Botanical Club
CONTENTS
CENTENNIAL SYMPOSIUM: The Dynamics of the New England Flora
Dedication. 2A]
Opening remarks: Plant conservation globally and locally. Peter H. Raven.. 243
Post-glacial changes in ed and climate in northern New England.
George L. Jacobson, Jr. 246
ate ae of post-glacial eee in climate and ee on the flora
e White Mountains, New Hampshire. Ray W Spear ............ 248
Vegetation of the presettlement forests of northern New England and
w York. Charles Cogbill 250
Fifty ees of ee in Rhodora and the New England flora. Warren H.
Wag 2
Linking the We and recent past to the modern New England landscape.
vid R. Foster 278
Immigration and expansion of the New England flora. Leslie J Mehrhoff.. 280
Rates of success in the reintroduction by four methods of several perennial
plant species in eastern Massachusetts. Brian D) oe and Richard B.
Primack 299
The myth of the resilient forest: Case study of the invasive Norway maple
Acer platanoides). Sara L. Webb, Marc Dwyer, Christina K.
Kaunzinger, and Peter H. Wyckoff B32
Closing remarks. W Donald Hudson, Jr: 355
NOTE
Callery pear (Pyrus calleryvana—Rosaceae) naturalized in North Carolina.
uy Nesom 361
NEW ENGLAND NOTE
Snow algae in the northeastern U.S.: Photomicrographs, observations,
and distribution of € chloromonas spp. (Chlorophyta). Brian Binal
and Ronald W. Hoha 365
NEBC MEETING NEWS 373
ANNOUNCEMENT
New England Botanical Club Graduate Student Research Award ........ 380
Information for Contributors 381
NEBC Membership Form 383
NEBC Officers and Council Members inside back cover
Vol. 102 Summer, 2000 No. 911
Issued: September 29, 2000
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
ANTOINETTE P. HARTGERINK, Managing Editor
Department of Plant Biology, University of New Hampshire,
Durham, NH 03824
Associate Editors
HAROLD G. BROTZMAN STEVEN R. HILL
DAVID S. CONANT THOMAS D. LEE
GARRETT E. CROW THOMAS MIONE
kK. 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 paya-
ble at par in United States currency. Remittances payable to RHO-
DORA. Send to RHODORA, P.O. Box 1897, Lawrence, KS 66044-
8897.
MEMBERSHIPS: oo" $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 meetings and
special events, annual graduate student award, and the journal RHO-
DORA is available at http://www.herbaria.harvard.edu/nebc/
BACK ISSUES: Questions on availability of back issues should be ad-
dressed 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-
D , changes must be received by the business office prior to the
first day of January, April, July, or October.
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
DEDICATION
WARREN HERBERT WAGNER, JR.
1920—2000
This issue of Rhodora is dedicated to Dr. Warren Herbert Wag-
ner, Jr—botanist and friend to botanists, professor, and student.
Club members will long remember his spirited defense of field
botany and whole-plant biology given at the symposium.
Largely responsible for rejuvenating the systematic study of
North American pteridophytes, Herb considered thorough field
knowledge of the plants in question a prerequisite to any labo-
ratory investigation. He demonstrated the efficacy of this policy
with major advances in the systematics of spleenworts (Asplen-
ium), woodferns (Dryopteris), and moonworts (Botrychium).
Herb’s legacy extends beyond the study of pteridophytes, how-
ever, since his students and others have applied his lessons across
the botanical spectrum. In particular, his demonstration of retic-
241
242 Rhodora [Vol. 102
ulate evolution in many fern genera led to a general inquiry about
the significance and evolutionary consequences of natural hy-
bridization. Many of us will remember him also for the devel-
opment of ‘‘Wagner Trees” during the infancy of cladistics,
which is ironic since clades can be obscured by hybrid reticula-
tion.
In his later years Herb returned increasingly to field work, and
ended his career immersed in the discovery and naming of new
species. Within the past 20 years he named more than a dozen
new species of Botrychium from the Great Lakes region and west-
ern North America, and during his last summer he traveled to
Alaska to investigate a new moonwort there. In addition, he re-
cently named numerous species of Hawaiian ferns, on which he
was an expert, based on his many years of field experience there.
Young at heart, always curious and inquisitive, never accepting
dogma (old or new), Herb was a most enthusiastic teacher, a
prolific writer, and a superb field companion. He was always
ready to talk about the nature of plants, either on a broad scale
or a narrow one, and he enlivened any field trip with his broad
range of knowledge.
Although his main research efforts lay outside New England,
we all have benefitted from his work and his teachings. The
NEBC is especially grateful that Herb gave freely of his time and
spirit to the symposium.
—ARTHUR V. GILMAN, Rhodora Centennial Symposium Committee.
RHODORA, Vol. 102, No. 911, pp. 243-245, 2000
OPENING REMARKS
PLANT CONSERVATION GLOBALLY AND LOCALLY
PETER H. RAVEN
Missouri Botanical Garden, P O. Box 299, St. Louis, MO 63166
What I want to do here is to paint the broadest picture possible
of the current crisis in biological conservation around the world,
and in particular in the United States. This crisis is so extreme,
and so important to us all, that I want to stimulate thinking about
the issues involved and invite dialogue about strategies for com-
bating the problem.
Our planet is four and a half billion years old and life has
existed for at least 3.8 billion years, as far as we can tell. During
that time, five great extinction events have occurred. The first
three were restricted entirely to marine life. The fourth occurred
at the end of the Permean, approximately 280 million years ago.
This event changed the character of life on Earth, and led into
the Mesozoic, the era of dinosaurs and cycads. During this era,
angiosperms and other modern groups evolved and life on Earth
became more and more complex. The fifth great extinction event
occurred at the end of the Mesozoic, 65 million years ago.
he fifth extinction was probably precipitated by a meteorite,
which landed somewhere in the vicinity of the Yucatan in Mex-
ico. The result was an opaque cloud that restricted photosynthesis
and disrupted life; two-thirds of the terrestrial species became
extinct and the character of life on land again changed complete-
ly. Approximately ten million years went by before recovery of
the evolutionary pathways that led to modern groups. It’s esti-
mated that, after the extinction, there were 500,000 to 1,000,000
terrestrial species remaining. Currently, the Earth is estimated to
ouse between 7 and 10 million species of eukaryotic organisms.
Of these, only one in four has a valid name. In the tropics, the
ratio is much less, around one in twenty. Even for the described
species of organisms, our knowledge of relationships and eco-
system dynamics is extremely limited.
Homo sapiens appeared on Earth approximately 200,000 years
ago. During the past 200,000 years, our species has developed
agriculture, migrated around the world, and increased in popu-
243
244 Rhodora [Vol. 102
lation to approximately six billion. Much of this population in-
crease has occurred during the past century. Human population
growth has been fueled by the use of oil, gas, and coal, causing
tremendous atmospheric destruction. In the coming century, we
are faced with the important questions: can we achieve sustain-
ability and can we maintain the Earth’s biological diversity?
These questions are two sides of the same coin; the results of the
past 50 years are not encouraging.
Americans first became aware of the concept of biodiversity in
the 1980s. At this time, biodiversity was thought of as a kind of
inventory of species, but gradually, the definition expanded to
include all relationships and biological variations within com-
munities and within ecosystems. In 1992, Americans became
acutely aware of this situation, when President George Bush de-
clined to sign the treaty ‘International Convention on Biological
Diversity” at the World Summit held in Rio de Janeiro. At this
convention, the United States was one of only seven countries to
decline ratification.
Over the last fifty years, the population in the United States
has increased from 135 million to 270 million. Also during this
time, approximately 25% of the world’s topsoil has been lost
permanently, and 15-20% of the world’s agricultural land has
been lost to forces such as urban sprawl and deforestation. In
addition, carbon dioxide levels have increased more than 30%
and the stratospheric ozone layer has been depleted by 8%. Most
seriously, there has been a drastic increase in the proportion of
biological extinction in the past fifty years. We know from doc-
umented extinctions of birds and mammals that we can expect
any given species to last about two million years. Since the Ren-
aissance, extinction rates have increased hundreds of times. Cur-
rent extinction rates are 1,000 X background rate and are accel-
erating towards 10,000 X background rate in the next century.
Stuart Pimm of the University of Tennessee, using island bioge-
ography models, has calculated that 1/3 of all tropical species
will be extinct or nearly extinct within the next 25 years, and that
3/4 of all tropical species will be extinct or nearly extinct by the
end of the next century.
It has been projected that, worldwide, 2/3 of all organisms will
be extinct by the end of the next century. One species, Homo
sapiens, is driving an extinction event comparable in scope and
intensity to the one 65 million years ago, which completely
2000] Raven—Opening Remarks 245
changed the character of life on Earth. Nothing we could do could
be more short-sighted or damaging to our own future, since we
depend on the 350,000 species of photosynthetic organisms for
all productivity in the biosphere.
We say that the 21st century will be the “Age of Biology,”
but how can it be if we are eliminating species at a rate compa-
rable to the great extinctions of the past? How can we harness
the power of genetic variability of organisms, which we are only
just beginning to understand, when we are driving extinction rates
to such phenomenal levels? Doesn’t it make sense for us to be
interested in sustainability? Doesn’t it make sense for us to be
interested in the rest of the world? Over the past 50 years, the
industrialized world’s population has fallen from 30% to 20%,
but that 20% controls 85% of the world’s economy and creates
80-90% of the world’s pollution. The United States, as the
wealthiest nation, can and must make a difference in reversing
this trend for the benefit of ourselves and the rest of the planet.
What can we, as citizens, as scientists, do to maintain a world
in which beauty, music, poetry, philosophy, literature, biological
diversity, and all the things that we cherish can thrive? What do
we have to do to create a world that will go on? It is important
that we create a world that can continue to function sustainably.
It is important that we not lose the biological diversity that would
sustain us in the future. Our future depends on how we live now,
on the choices we make. If we want to be optimistic about the
world’s future, we need to exercise our personal determination to
do something about it. We should: (1) be leaders in developing
new forms of energy conservation and alternative sources of en-
ergy; (2) pay attention to internationalism and acknowledge that
the people living in non-industrialized countries are of profound
importance to the earth’s future; (3) vote, and encourage others
to do so; (4) support like-minded conservation groups that are
devoted to sustainability and internationalism; and (5) make wise
choices and decrease our personal levels of consumption.
New Englanders have always looked to the world far beyond
conventional boundaries. New Englanders have traditionally un-
derstood the world from a perspective of trading, knowledge, cul-
ture, and art. This is a great and traditionally defined region of
our country, and one that has made wonderful contributions, and
is set to make wonderful contributions for the future. In that vein,
I hope that the remarks I have offered here have given you some-
thing to think about and to act on.
RHODORA, Vol. 102, No. 911, pp. 246-247, 2000
POST-GLACIAL CHANGES IN VEGETATION AND
CLIMATE IN NORTHERN NEW ENGLAND
GEORGE L. JACOBSON, JR.
Institute for Quaternary Studies, University of Maine, Orono, ME 04469
e-mail: jacobson @ maine.maine.edu
SUMMARY. Quaternary research in several disciplines has pro-
duced strong independent evidence about the post-glacial vege-
tation and climate of northern New England and adjacent Canada.
Late-glacial environments in the region included extensive areas
of treeless tundra—more so than was the case in glaciated areas
of mid-continental North America. Tree taxa spread gradually
from the south, with most current forest elements present by the
early Holocene. Subsequent changes in climate have greatly af-
fected the distribution and abundance of those taxa.
Stratigraphic changes in physical and biological characteristics
of lake sediments indicate that early to middle Holocene temper-
atures were as much as 2°C warmer and that the moisture balance
(precipitation minus evaporation) was considerably lower than
today. These reconstructions are consistent with known orbital
variability (especially precession of the equinoxes) which resulted
in as much as 8% more summer insolation than at present.
Several lines of paleoecological data corroborate this paleocli-
matic reconstruction. White pine (Pinus strobus L.) was wide-
spread and abundant in the early to middle Holocene, probably
because frequent fires created conditions favorable for seedling
establishment. During that same time, both white pine and hem-
lock [Tsuga canadensis (L.) Carriére] were present at elevations
as much as 300 to 400 m higher than their present upper limit in
the White Mountains of New Hampshire and the Adirondack
Mountains of New York.
Conditions changed considerably during the past few thousand
years, however, as the climate became cooler and moister. Fossil-
pollen evidence shows that the distribution of white pine, which
had been so extensive during the drier early and middle Holocene,
has diminished consistently during the past 4000 years. This de-
cline appears to have resulted from a reduction in frequency of
forest fires during the late-Holocene shift toward a cooler, moister
climate.
Within the past 1000 years, populations of several boreal forest
246
2000] Jacobson—Changes in Vegetation and Climate 247
taxa, including spruces (Picea spp.) and balsam fir [Abies bal-
samea (L.) Miller] expanded along the southern margins of their
distribution in Canada and the northern tier of the United States—
from Minnesota to Maine. The strong expansion of spruce in the
Great Lakes-New England region, especially the past 500 years,
appears to have been associated with summer cooling of about
1°C during the Little Ice Age.
What can be said about the future? General Circulation Model
(e.g., NCAR CCM3) projections for a future with twice the pre-
sent atmospheric concentration of CO, suggest that both summer
and winter conditions in northern New England may be as much
as 3°C warmer than at present and that precipitation may also be
greater. If the models are correct, the summer conditions may be
aS warm as or warmer than those 6000 to 8000 years ago, but
possibly also wetter.
RHODORA, Vol. 102, No. 911, pp. 248-249, 2000
IMPLICATIONS OF POST-GLACIAL CHANGES IN
CLIMATE AND VEGETATION ON THE FLORA OF THE
WHITE MOUNTAINS, NEW HAMPSHIRE
RAY W. SPEAR
State University College of New York, Geneseo, NY 14454
e-mail: spear@uno.cc.geneseo.edu
SUMMARY. Steep environmental gradients along the slopes of
the White Mountains support a diverse flora in several distinct
vegetation zones. Deciduous hardwood forests with white pine
and hemlock occur at low elevations. Coniferous forests of spruce
and fir with paper birch grow at mid-elevations. Subalpine fir
forests are found at tree-line (~1500 m). The alpine meadows on
the highest ridges and peaks have a flora that includes 73 species
of vascular plants and a number of species of bryophytes that are
essentially alpine.
Paleoecological studies in the mountains have recorded the
broad-scale postglacial climate and vegetation changes over the
past 13,000 years. These changes fit the regional patterns de-
scribed for northern New England. However, because of the
mountains’ elevational range, two paleoecological changes are
unique to the White Mountain region. Pollen and plant macro-
fossil evidence indicates that tree-line reached the valleys 11,500
years ago and reached its modern position on slopes 10,000 years
ago. The contemporary alpine flora is widely assumed to be the
remnant of the arctic flora that followed the retreating ice sheet.
The mid-elevation coniferous forest of spruce, fir, and paper birch
is relatively recent in origin, expanding several thousand years
ago. Elevational range extension of white pine and hemlock and
the mixtures of deciduous tree species in mid-elevational forests
from 10,000 to roughly 4000 years ago indicate warmer climatic
conditions than today. Tree-line may have also stood above its
current elevation during the early to mid-Holocene 9000 to 4000
years ago.
While the patterns of vegetation and climate change are well
known on the scale of thousands of years, short-term changes on
the order of tens or even hundreds of years are poorly docu-
mented. Evidence from the Greenland ice-cores, the North Atlan-
tic marine record, the Atlantic Canada terrestrial record (midges,
pollen, and plant macrofossils), and the southern New England
248
2000] Spear—Flora of the White Mountains 249
pollen record show dramatic short-term climatic oscillations dur-
ing the late-glacial (14,000 to 10,000 years ago). The ice-core
and marine records show that these oscillations may have ex-
tended into the Holocene (the last 10,000 years). High (temporal)
resolution studies are underway in the White Mountains to find
evidence of these climatic oscillations. Preliminary loss-on-igni-
tion (percent organic matter in the sediments), midge, and pollen
studies document that two of these short-term oscillations, the
Killarney Oscillation (KO) and the Younger Dryas (YD) did oc-
cur in the White Mountains. The mean annual temperature is
estimated to have changed as much as 5—10°C within a period of
less than 50 years during the YD 10,700 to 10,000 years ago.
The magnitude and abruptness of these events may have created
bottlenecks that reduced the size of the alpine and lower elevation
floras in the White Mountains.
RHODORA, Vol. 102, No. 911, pp. 250—276, 2000
VEGETATION OF THE PRESETTLEMENT FORESTS OF
NORTHERN NEW ENGLAND AND NEW YORK
CHARLES V. COGBILL
ie Walker Lane, Pisinnield, VT 05667
mail: ccogbill @y ypass.com
ABSTRACT. The lotting surveys from northern New England and New
York provide a unique opportunity to derive quantitative documentary evi-
dence on past forests. Due to the distinctive ‘‘proprietor * land tenure
system, northern New England has an extensive and virtually untapped ar-
chive of land division surveys done prior to settlement (1763-1820). I
searched archives throughout Vermont, New Hampshire, and northern New
York and located records from 179 towns documenting 48,260 individual
trees across the region. Surveyors used 131 separate vernacular names rep-
resenting at least 49 recognizable species. This collection of tow -wide wit-
ness tree relative frequencies is a consistent and unbiased em cal estimate
of the composition of the natural vegetation before eonfenaiiag land use.
Five ubiquitous taxa (beech, ees maples, hemlock, birches) comprised
79% of the witness trees. Beech (32%) consistently enero the region
with greater than 60% of the aa some towns. pence %) was the
e Maple
(12%) were consistently distributed with peak abundance in Ver mont. Hem-
lock (12%) had a patchy distribution with ses of abundance, including
the eastern Adirondacks. Birches (9%) were a species group with higher
abundance in the mountains or to the no Hes White pine was consistently
uncommon with very <1%) abundance on the uplands. A dramatic
“‘oak—beech” tension zone or ecotone separated the oak—pine vegetation in
the major southern valleys from the spruce—maple—beech composition north-
ard. The eee tendency was toward spruce—hardwoods with distinctive
variants in the north, the Taconics, and the Champlain Valley. Major fires and
lowdowns were pee rare and affected only 0.5% of the region. Fire was
frequent only in the Hudson-Champlain corridor and windthrow was a low-
level background denubanee: The most dramatic changes documented over
the past 200 years have been the decline of beech and the profound effects
of human land use.
Key Words: Adirondacks, historical ecology, New Hampshire, northern
hardwood forest, plant biogeography, presettlement vegetation,
proprietory town, surveyor’s records, Vermont, witness tree
The first Europeans in northeastern North America saw the
forest as “daunting terrible . . . infinite thick woods” (Josselyn
1675). Historical views of the land are subjective and anecdotal,
but we are still influenced by their lasting metaphors: ‘‘This is
250
2000] Cogbill—Vegetation of Presettlement Forests 251
the forest primeval. The murmuring pines and the hemlocks/
Bearded with moss, and in garments green, indistinct in the twi-
light/ Stand like Druids of eld, with voices sad and prophetic”’
(Longfellow 1854). Indeed the 18th century was le grand dé-
rangement for both the people and forests of Acadia and New
England. The 1700s began the clearing and harvesting of upland
New England and within a century the land was profoundly
changed (Whitney 1994). Since much of the region is forested
today, we often assume a similarity to, if not continuity with,
forests of the past. Was the forest dominated by the pines and
hemlocks of Longfellow or the “‘hemlock—white pine—northern
hardwoods” of Braun’s (1950) classic treatise? Was the vegeta-
tion the thick woods of Josselyn or filled with decadent behe-
moths of ‘“‘old growth” stands? Were the “‘bearded”’ trees long
undisturbed and waiting to die of old age? Reconstructing the
nature of the “‘primeval’’ forests is not just an academic exercise
in historical ecology, but is necessary to establish an empirical
baseline for ecological, educational, and management activities.
The characteristics of present forests can be projected back-
ward to elucidate the composition and structure of historic forests.
The response of trees to environmental factors or the current com-
position of long undisturbed forests are potential models for past
conditions. Each of these models has limitations. Correlation with
environmental variables is usually linear and deterministic. It
tends to produce broad zones of vegetation responding to a single
factor (e.g., climate) or stereotyped vegetation types based on
distinctive topography or substrates. More appropriate models
would be more sophisticated (i.e., non-linear, multivariate, sto-
castic) and be spatially explicit (e.g., Pacala et al. 1993). In ad-
dition, environmental conditions, especially in glaciated regions,
are not a constant background and shift (e.g., climate) or develop
(e.g., soils) in the interim. Paleoecological studies show the veg-
etation in northern New England is in flux due to long- and short-
term environmental changes, land use history, and stochastic fac-
tors (Jacobson 2000).
A more practical approach to extrapolation is to use surviving
forest remnants, especially those unconfounded by human activ-
ities. Even the largest of these areas in the Northeast (e.g., The
Nature Conservancy’s Big Reed Forest Reserve in northern
Maine, The Bowl Research Area in the White Mountain National
Forest of New Hampshire, and the Five Ponds Wilderness of the
pays Rhodora [Vol. 102
Adirondack Park in New York) are few in number, have been
repeatedly naturally perturbed (i.e., wind, ice, insects, fire), and
escaped human activities exactly because they were ‘‘odd-balls”’
or “‘out-of-the way” (Cogbill 1996). Remnants, by definition,
have escaped expected disturbances and are necessarily atypical
of the “normal”’ or prevailing ‘“‘common”’ landscape at any time.
Since the nature of the primeval forest is shrouded by myths,
unrealistic models, and atypical remnants, historical methodology
must be used to discover their reality. Actual ‘‘eye-witness”’ ac-
counts are needed to document the details, variation, and dynam-
ics in the landscape. Contemporary observations of explorers,
naturalists, diarists, authors, and publicists abound, but they are
very subjective and generally qualitative ney 1994). Paleo-
historical studies are more scientific and ‘‘see”’ the forest through
reconstruction. They examine the physical evidence from earlier
forests (e.g., dead wood, charcoal, pollen, macrofossils) at a site,
testifying of the past occupants. Regional paleoecological synthe-
ses give a long-term and relatively low resolution history of re-
gional vegetation (Jacobson 2000; Spear 2000). We do not, how-
ever, have to resort only to scientific studies of remains to dis-
cover aspects of the vegetation that covered the historic land-
scape. Although the trees are no longer available for scientific
inquiry, there is a contemporary and empirical documentary re-
cord. Surveyors at the time actually saw and recorded forest com-
position during land division and survey (Whitney 1994). Early
surveys from northern New England, generally 1763 to 1820,
clearly document the actual, not theoretical, abundance of partic-
ular trees in the forest before human improvements.
A distinctive land tenure system arose in the 18th century in
the northern English colonies in North America (Clark 1983;
Price 1995; Woodard 1936). The unsettled lands in northern New
England were divided into areas, ideally six miles square on a
side (= 100 km*), called “towns.” The “‘outlines”’ of the towns
were commonly surveyed in atl estos of settlement and then
the land was erence by the crown or the colonial government to
a group of persons “‘in common,” so-called ‘‘proprietors.’’ The
main occupation of the absentee proprietors was to subdivide the
town into “lots,” survey those (typically 40-60 ha) lots, and
transfer ownership by means of a “lottery”? among shareholders.
Surveyors traditionally documented distances and corners of the
outlines and lot boundaries by blazing and recording trees (‘‘wit-
2000] Cogbill— Vegetation of Presettlement Forests 253
ness trees”). An unintended consequence of this “‘lotting survey”
was a sample of the trees in the town on a predetermined grid.
Significantly, the surveyors also often recorded general forest
conditions, the suitability for settlement, and unusual character of
each lot.
The proprietory town system was continued by the New Eng-
land states after independence until disposal of ungranted land
was completed in the mid-1800s. In northern New York much of
the land was not granted to proprietors, but starting about 1791
the various units (i.e., tracts, purchases, patents) were transferred
to individuals by the state (McMartin 1994). Significantly, in the
Adirondack region, many units or subdivisions called “town-
ships’? were nominally the same size (100 km?) as the New Eng-
land town. Although most of the New York tracts do not corre-
spond to towns today or were never settled, many were surveyed
by the state into lots in the proprietory town tradition. Taken all
together, these surveys inadvertently produced a systematic and
widespread sample of the forest of northern New York and New
England in the late 1700s and early 1800s (“‘presettlement sur-
veys’’). These records are official documents, but the local and
transitory nature of the proprietors resulted in the manuscript lot-
ting surveys, typically recorded in the ‘Proprietors’ Book,”’ or
the resultant maps being scattered in various repositories. The
New England town lotting methods were incorporated into the
Land Ordinance of 1785, which mandated the ‘rectangular sys-
tem” for land division in the western United States (Price 1995;
White 1984). The resultant federal General Land Office (GLO)
surveys have been the primary resource for numerous studies of
the historical landscape (e.g., Delcourt and Delcourt 1996;
Schwarz 1994; Whitney 1994).
Not as formalized as the GLO surveys, the unstandardized and
dispersed town proprietors’ surveys have received remarkably lit-
tle interest (cf. Bourdo 1956; Irland 1999; Whitney 1994). In the
Northeast, studies of the presettlement surveys have been done
in northern Vermont (Siccama 1971), northern Maine (Lorimer
1977), upstate New York (Marks and Gardescu 1992; McIntosh
1962: Seischab 1990, 1992) and eastern Canada (Lutz 1997;
Moss and Hosking 1983). All these studies used town outline
surveys, except Siccama (1971) who used lotting surveys within
6 towns in northern Vermont to look at local vegetation patterns.
In contrast, there are several towns in both New Hampshire and
254 Rhodora [Vol. 102
Connecticut with readily available manuscript maps or other sum-
maries of lotting surveys (Cogswell 1880: Hamburg and Cogbill
1988; Torbert 1935; Winer 1955). In addition, several researchers
at Harvard Forest have begun to analyze town-wide lotting sur-
veys in over 40 towns in southern New England (Foster et al.
1998; M. Burgi, pers. comm.). Whitney (1994) integrated many
of these surveys into maps depicting the pattern of species’ abun-
dances over the northeastern quarter of the United States. His
small-scale maps show broad continental distributions within the
Northeast. The wealth of information from the underutilized and
humerous town surveys is an unparalleled opportunity to fill in
geographic gaps in coverage and display details of species dis-
tributions. Thus this study’s purpose is to locate, collate, and sum-
marize the available town witness tree surveys to derive a quan-
titative empirical database on the presettlement vegetation and its
variation over northern New England and New York.
MATERIALS AND METHODS
I searched archives throughout Vermont, New Hampshire, and
New York to locate manuscripts, maps, and published records of
lotting surveys before settlement. The collation of witness trees
in the presettlement surveys resulted from three separate projects.
The Vermont collection was commissioned by the Vermont Bio-
diversity Project to provide background for the classification of
vegetation in the state. The majority of the recorded surveys
found were in the Proprietors’ Books typically housed with the
town land records in their respective Town Halls. Copies of many
of the early town records are on microfilm at the Public Records
Office in Middlesex. In addition, surveys of the towns granted
by colonial New York within the present borders of Vermont were
found in the New York State Archives (NYSA) in Albany. In the
1790s, the leased lots in the Rensselaer Manor towns adjacent to
Vermont were also surveyed using similar techniques (Rensse-
laerwyck Papers, NYSA). The New Hampshire surveys were col-
lected in a collaboration between the Hubbard Brook Long-Term
Ecological Research (LTER) and the Harvard Forest LTER pro-
jects to create a baseline for historical land use studies. New
Hampshire surveys were also usually recorded in the Proprietor’s
Books, commonly housed in the town office. Microfilm copies of
many town records in New Hampshire were found at the New
2000] Cogbill—Vegetation of Presettlement Forests 25)
Hampshire State Library in Concord. I collected the New York
surveys in a follow-up project to document further the character-
istics of old growth forests in the Adirondacks (Woods and Cog-
bill 1994). The New York records were found in the state’s col-
lection (NYSA) of Field Books (also available from the LDS
[Mormon] Family History Centers) or Surveyor General’s Books.
Other sundry surveys and summary maps were collated from
published papers and from manuscripts in various repositories
(Town offices, State Historical Societies, State Archives, State
offices) in all three states.
Records containing witness tree data or descriptions of the for-
est were carefully read, noting all trees cited by name and any
descriptions of the forest composition or its disturbance (6.2%
“open”, “burnt”, ‘‘fallen’’). Throughout proprietary lotting sur-
veys, virtually all witness tree citations were of a single tree at
each sample point, so species associates were only known from
supplemental line or lot descriptions. Whenever possible the trees
were located on a map of the original lots (‘‘lotting map” or
“town plot’’), and a special effort was made to avoid duplication
of trees on shared boundary lines or the corners of adjacent lots.
All witness tree “‘mentions’’ within each town were classified and
tallied by the most exact taxa inferred by the surveyor’s name.
When possible, appropriate taxa were combined and frequencies
summed into functionally similar groups (e.g., soft maples, white
oaks, hard pines, wet ashes, white birches). In order to maintain
a consistency in the identifications across all towns, the taxa and
their frequencies were further lumped into 26 exclusive genera
groups. For each town with more that 50 witness trees, the rel-
ative frequency of each taxon was treated as the presettlement
composition (ca. 1800) at that location. To reveal the distribution
pattern of each group or taxon, the relative frequencies were plot-
ted on basemaps using Street Atlas USA® (DeLorme Cc., Yar-
mouth, ME). Following Whitney (1994), isopleths of equal wit-
ness tree frequency (‘‘isowits’’) generalize the patterns within the
region.
Due to availability of wide ranging and detailed presettlement
surveys, the forest was arguably better documented before settle-
ment than it is today. The most detailed current data on the forest
composition is in the decennial Forest Inventory and Analysis
(FIA) project of the United States Forest Service. A comparison
of these two data sets highlights changes over the past 200 years.
256 Rhodora [Vol. 102
I calculated the ratio of the 1983 Vermont FIA (Frieswyk and
Malley 1985) relative density of trees (> 5 in. DBH) to the av-
erage witness tree frequency in equivalent species’ groups in the
state ca. 1800.
RESULTS
This study located 179 ‘ttowns” in northern New England and
New York with extensive lotting surveys (Table |). The proprie-
tory surveys date primarily from 1763 to 1810. Some non-pro-
prietory towns’ lots were surveyed as early as 1673 (Clark 1983),
while late-granted tracts in the mountains were surveyed as late
as 1850. In Vermont and New Hampshire the Proprietor’s Books
have survived in at least 185 (37%) of the towns and 105 (57%)
of these books contain numerous witness tree mentions. Although
21 other town witness tree records were uncovered, the vast ma-
jority (83%) of the New England surveys were from the propri-
etors’ records themselves. Due to the more exhaustive searches,
Vermont has a slightly higher “yield” of books or surveys than
New Hampshire; at least 33% of the Vermont towns have sur-
viving presettlkement surveys. In New York’s Adirondacks, 49
(37%) of the larger tracts and townships have equivalent surveys
available. The towns in the witness tree database come from
throughout the region. There is the greatest representation from
the heavily settled Merrimack Valley and western Vermont towns
and the least representation from east-central New Hampshire and
southeastern Vermont towns. Overall 48,260 witness trees were
tallied with a median of 179 trees in a town and a rough sample
density of 2.7 trees per km?.
Vernacular names. In 1609, Samuel de Champlain (1925)
saw “. .. fine trees of the same varieties (espéces) we have in
France” along the New York shore of Lake Champlain. Although
the early European observers were familiar with the genera in the
flora of eastern North America, the species were novel. Surveyors
invariably used colloquial names for trees, but virtually all cita-
tions can be associated with known scientific taxa. The lack of
Latin names is not surprising given the short time since the in-
troduction of the Linnaean system in 1753 and lack of useful
manuals or floras until the turn of the century. Despite their iso-
lation and lack of formal botanical education, the surveyors were
Table 1. Number of towns and witness trees represented in existing surveys from Proprietors’ Books and other archival docu-
ents.
m
Re
Number of Towns with pamper
Current a a Re SE of
Number of Prop. Prop. Book Total Survey Witness
Region “Towns” Book Lotting Lotting Dates Trees
New Hampshire 245 77 35 44 1673-1850 16,781
Vermont 251 108 70 82 1763-1820 21,150
Adirondacks 132 — — 49 1771-1831 8960
Taconics 4 — — 1790-1795 1369
4
TOTAL 628 195 105 179 1673-1850 48,260
a
L000z
SISIIOY JWSWIINasaIg JO uoNLIedaA—T]IqBo5
Table 2. Cited tree names in presettlement (1763-1820) forest surveys from 179 towns in Vermont, New Hampshire, and
northern New York. Brackets indicate possible taxonomic ambiguity. Nomenclature follows Gleason and Cronquist (1991).
Surveyor Name
Generic Names
Specific Names
Alder
Apple
Basswood
Black ash
Black birch
Black cherry
Surveyor
Synonyms
Black spruce
Wild cherry
Hawthorn
Osier, White willow
Alder birch
Brown ash, Yellow ash
Cherry birch
Spelling Variant
Burch, Berch, Birtch, Burtch
Holm, Ealm
Mapel, Maypole, Mepall
Oke, Oake, Och, Ock, Ocke
Popler, Popular, Popplr
Shadbush
Bass, Baft
Beach, Bectch
Taxa Referenced
Fraxinus sp.
Betula sp.
Quercus sp.
inus sp.
Populus sp.
Amelanchier sp.
Crataegus sp
Salix sp.
Alnus incana
Pyrus malus
Tilia americana
Fagus grandifolia
Fraxinus nigra
Betula lenta
Prunus serotina
N
Al
io)
vlopoyuy
TOA]
cOl
Table 2. Continued.
Mountain ash
orway pine
Pepperidge
S
Red birch
maple, Moosebush,
Stripped willow
Red pine
Mt. Ash
Pepraige
Pich pin
ae Popel, Poppel
sh
Read ash, Reed a
ae ch
Surveyor
Surveyor Name Synonyms Spelling Variant Taxa Referenced
Blue beech Water beech Carpinus caroliniana
Boxwood Box tree Acer negundo?
Butternut White walnut, Lemon Butnut, Buternut Juglans cinerea
walnut, Oylnut,
Oilnut, Butterwood
Buttonwood Sycamore Buttinwood Platanus occidentalis
Cedar White cedar Thuja occidentalis
Chestnut Chisnut, Chesnutt Castanea dentata
Chestnut oak Rock car es oak Quercus prinus
Fir Balsam Firr, Pur Abies balsamea
Hemlock ie pine Hamlock, Hemloc Tsuga canadensis
Leaverwood Lear wood, Liverwood Ostrya virginiana
Moosewood Moose willow, Moose Acer pensylvanicum
Sorbus americana, LS. decora]
Quercus palustris
Pinus rigida
Betula cordifolia, [B. alleghan-
iensis |
L000Z
SJSOIOf JUIWITHWIsSaI1g JO uOTRIIsa A —]]IG30D
Table 2. Continued.
Surveyor Name
Surveyor
Synonyms
Spelling Variant
Taxa Referenced
Red cedar
Red cherry
Red elm
Red oak
Sassafras
Sugar maple
Swamp maple
Wh pru
Yellow birch
Swamp white oak
ck
Reed cherry
Read oak, Reed oak, Reid
oa
Saxefax
Shag walnut
Red maple
Double spruce, spruce Sprusse
pine
Hard maple, Rock
maple, Sugar tree,
Black maple
Swamp oak
Hacematac, Larch Tamarac, Tamarisk
Silver maple
White burtch
Whight oak, Whit ocke
Juniperus virginiana
runus pensylvanica
Ulmus rubra
Sassafras albidum
arya ovata
Acer rubrum, [A. saccharinum]
Picea rubens, [P. mariana]
Acer saccharum
Acer rubrum, [A. saccharinum|]
Quercus bicolor
Larix laricina
Fraxinus nigra
Fraxinus americ
Betula iene 1B. cordifolia]
Ulmus americana
Acer saccharinum, [A. rubrum]
Quercus alba
Pinus strobus
Picea glauca
Betula aileghaniensis
O97
elopoyuy
JOA]
cOT
Table 2.
Continued.
Surveyor Name
Surveyor
Synonyms
Spelling Variant
Taxa Referenced
Yellow pine
Ambiguous Names
Balsam
Bastard maple
Hornbeam
Juniper
Moose elm
Pe
Plum
Rock birch
Wild pear
Witch elm
Whitewood
Deceptive Names
Black oak
Hacmetack
Walnut
Witch hazel
Rock white birch
Hazel, Hazelnut
Pinus rigida, [P. resinosa]
Abies balsamea, [Populus bal-
samifera]
Acer spicatum, [A. pensylvani-
cum
strya virginiana, [Carpinus
caroliniana|
Juniperus virginiana, Thuja oc-
cidentalis
Acer a
Prunt
Ulmus rubra, U. americana
not Liniodendion: Populus delto-
ides?
=i bit [Q. velutina]
pe sie [Larix laricina]
arya Sp.
Ostrya virginiana
Lo00z
S]SOIO{ JUSTUT}aS91g JO UOTIeIIB9aA—][IQ3s0D
Table 2. Continued.
Surveyor
Surveyor Name Synonyms Spelling Variant Taxa Referenced
Enigmatic names
Beattlewood ?
Bilberry tree ?
Greenwood ?
Jerwood ?
Kalmia latifolia?
Pegwood ?
Remmon Remmond, Ammon, Remon ?
Roundwood Sorbus ?
Shittum wood Sorbus ?
Spoonwood bush ?
Tobaccowood Moose(wood) Acer pensylvanicum ?
Wice not Dirca
797
vlopoyy
ZOI TOA)
2000] Cogbill— Vegetation of Presettlement Forests 205
competent naturalists. The early lotting surveys recorded 131 ver-
nacular tree names or synonyms (excluding quaint spellings) rep-
resenting 65 distinct taxa (Table 2). Interestingly, the colloquial
names were influenced by transferred English usage, and so the
most ambiguous attributions are in taxa not shared with the Brit-
ish flora (i.e., Ostrya, Carya, shrubby Acer). The surveyors were
very discerning and consistent in usage. For example, they often
made subtle species distinctions (e.g., red ash, red elm). Overall
49 recognizable species are found in the presettlement species list
for the 179 towns (Table 2). All these cited species are prominent
current members of the approximately 65 species in the region’s
tree flora. Several infrequent species were not explicitly acknowl-
edged (e.g., bur oak, big-toothed aspen, grey birch, bitternut hick-
ory), but they are certainly present, submerged in amorphous gen-
era or by misunderstood terms. There have been no apparent ex-
tirpations, but some terminology (e.g., lemon walnut, leaverwood,
pepperidge) has fallen out of use.
Although the surveyors used many explicit vernacular names,
there are still various degrees of uncertainty in some species (Ta-
ble 2). For many of the most common trees only a generic name
was cited (i.e., maple, oak, pine, birch, ash). In these genera there
is an unavoidable confusion of species, but within their range and
proper habitat many of the common species are unambiguous
(i.e., sugar maple, red oak, white pine, yellow birch, white ash).
Even in context, in some genera (i.e., cherry, poplar) the cited
species remains ambiguous. Some specific names are still equiv-
ocal (e.g., swamp maple, yellow pine, red birch) or are occasion-
ally misapplied (i.e., balsam, hornbeam, juniper). The most con-
fusing are anachronistic names that have a deceptive common
meaning today (cf. Marks and Gardescu 1992; Seischab 1992;
Siccama 1963). Thus in late 18th century vernacular usage, hac-
metack referred to any conifer, especially Picea, rather than its
current exclusive use for Larix; witch hazel was Ostrya rather
than Hamamelis; walnut meant Carya rather than Juglans; dog-
wood did not refer exclusively to Cornus; black spruce included
red spruce, whose species concept did not exist until the late-
1800s; and black oak was regularly used for red oak. A few
names remain enigmatic; those used several times might be lost
vernacular terms (i.e., remmon, shittum wood, pegwood, tobac-
cowood), but those used a single time were more likely confu-
264 Rhodora [Vol. 102
3. Composite presettlement composition of witness trees (n =
48 oo in 179 towns in northern New England and New York.
Coefficient
of
Constancy Mean Maximum Variation
Taxa (%) (%) (%) (%)
Beech 98.9 32.1 68.2 42
Spruces 87.2 14.2 52.6 96
Maples 99.4 12.1 eyo 51
Hemlock 97.2 11.6 39.3 73
Birches 99.4 8.8 37.8 67
Pines 60.6 4.8 56.8 177
Oaks 52.2 4.8 58.8 195
il 55.0 2.9 25.0 165
Ashes 78.3 2.2 11.7 94
Basswood 66. | 1.4 10.0 121
Ironwoods 57.8 1.2 9.5 149
Elms 50.0 1.0 8.2 157
Cedar 30.6 0.5 7.1 216
Poplars BG a) 0.4 4.3 198
Chestnut 10.0 0.4 12.9 406
Tamarack 16.7 0.3 24 362
Hickories 20.6 0.3 4.3 254
Moosewoods 21.7 0.2 3.9 264
Butternut 17.8 0.2 5.3 326
Cherries 22.8 0.2 2.4 263
Willow & Alders 18.9 0.1 1.6 262
Buttonwood 7.2 0.0 1.7 499
Mountain Ash 2.2 0.0 1.7 740
O 0.2
sions (i.e., wicerpee, laurel), misunderstandings (greenwood, jer-
wood), or inventions (beattlewood, bilberry tree, gumwood)
Species distributions. The composite composition over the
179 towns is an integrated view of the vegetation in the region
in 1800 (Table 3). Five taxa (beech, spruces, maples, hemlock,
birches) composed 79% of the witness trees and each occurred
in virtually every town in the region. Each of these ubiquitous
trees was abundant (mean > 8%), had relatively low variability
between towns (coefficient of variation [CV] < 100%), and could
dominate certain towns (maxima > 30%) across the region.
Beech (mean 32%) was by far the most abundant species, ex-
ceeding 60% in widely scattered towns. It constituted greater than
2000] Cogbill—Vegetation of Presettlement Forests 265
30% of the trees throughout its range, falling off only in south-
eastern New Hampshire, in the high mountains, and in the far
northeast. Spruce was second in abundance (14%), but still less
than half that of beech. It had a more restricted range than the
other dominants (constancy 87% of the towns) and was not re-
corded in a few towns of the Champlain, Hudson, or Merrimack
Valleys. Spruce abundance was variable, reaching 15% at middle
elevations across the region, 35% in the mountains, and maxima
(> 50%) in the western Adirondacks and at the Canadian border.
Maples (mean 12%) had an abundance greater than 6% through-
out the region with high pockets (> 20%) scattered across the
richer soils of Vermont. By far the majority of these maple trees
consisted of sugar maple, but its abundance was necessarily less
than the generic figures. An undetermined lesser percentage of
the trees was red or silver maple, especially in lowlands or in the
larger river valleys. Hemlock had the same mean abundance
(12%), but had a more patchy distribution than maple. There were
three large polygons (i.e., southwestern New Hampshire, central
Vermont, and especially the eastern Adirondacks) of towns with
hemlock greater than the 20% isowit. Birches were the least abun-
dant (9%) of the dominants and this figure is inflated since the
taxon is also a mixture of species. Overall birch distribution was
variable with greater than 5% everywhere and maxima of greater
than 25% in the mountains. These maxima are most likely due
to white birch, but yellow birch was apparently most important
in mid-elevations and below. Both white and yellow birch in-
creased from south to north and upslope, and both the species
were represented in all areas except the Merrimack Valley.
Pine and oak were found in slightly more that half of the towns.
Both had low overall abundance, but in a restricted part of their
range could dominate (maxima > 55%) certain towns (Table 3).
Oaks (mean 5%) were commonly found only in the Champlain
or large southern valleys. Oak was codominant (> 30%) with
pine in the Merrimack Valley and was commonly abundant (>
15%) in the Taconics and Hudson-Champlain corridor; however,
these were a mixture of oak species. In lowland valleys, areas of
maximum oak abundance, white oak dominated, with high fre-
quencies even at the northern limit of its range (13%) on Squam
Lake, New Hampshire. Red oak was most abundant (to 25%) in
the southern hills and valleys and scattered (< 5%) northward in
266 Rhodora [Vol. 102
the upper valleys, but not found on the uplands or in the moun-
tains.
Pine (mean 5%) also showed a variable and restricted distri-
bution (Figure |). The 5% isowit bounds roughly three polygons:
in the Hudson-Champlain corridor, southeastern New Hampshire,
and the Connecticut River. The nested 20% isowit defines the
high pine abundance in scattered pockets in the Champlain, Con-
necticut, Ausable, and Saco Valleys and a large extreme (maxima
> 50%) area in the Merrimack Valley. Unfortunately, the distri-
bution and composition were obscured by the lumping of pines
(i.e., pitch, white, red) into a single group. In areas of maximum
representation in the large southern valleys, the majority of the
pattern is clearly due to pitch pine, as here it was regularly cited
by name or as “‘pine plains’’. White pine co-occurred in these
valleys and was probably most common in northern ones, partic-
oy the Champlain Valley and the “‘Cohas” (Abenaki for
“white pine place’’) in the upper Connecticut Valley (Whitney
1994). Evidently white pine was the only pine on the uplands
outside of the Taconics; but remarkably, here on the hills and
mountains, white pine was consistently uncommon with very low
(< 1%) abundance. Despite its reputation and conspicuousness,
white pine was a relatively minor component of the presettlement
forest in most of northern New England (cf. Braun 1950; Clark
1983; Irland 1999; Pike 1967).
In addition to the seven most abundant trees, five other taxa
(fir, ashes, basswood, ironwoods, elms) occurred in more than half
of the towns (Table 3). All these secondary species had low av-
erage abundance (1—3%). Except for fir, which could be locally
common (> 20%) in the mountains, these species were common
associates of the dominants and had rather modest maximum ex-
pression (8—12%). Ash was the most widespread (78% of the
towns), but its component species showed contrasting distribu-
tions: black ash was more northern and in the lowlands, white
ash more in the uplands and southern, while red ash was less
common and intermediate. The remaining secondary species
(basswood, ironwood, elms) were scattered across the region, but
each reached maximum abundance in the richer lowlands such as
the Champlain Valley.
The rest of the trees in the flora were recorded in less than
33% of the towns (Table 3). All these infrequent species, includ-
ing the minor species in the grouped genera, had low average
y
10.0 \
Ae 1775
3 \
Figure 1. Relative frequencies ey of pine in 179 town-wide presettlement lotting surveys in northern New York, Vermont,
and New Hampshire. Base map is ® 1996 DeLorme Co., Yarmouth, ME.
S|soIo,J JUSWIaTNesSeIg JO UOTIeII30A—T]]IG30D [000Z
LIC
268 Rhodora [Vol. 102
Number of towns (out of 179 surveyed) and lots affected by
cited disturbances in presettlement surveys (1763-1820) in Vermont, New
Hampshire, and northern New York.
ao! Fire Windfall
Towns
Region Disturbed Towns Lots Towns Lots
New Hampshire +t 2 2 2 2
Vermont 8 2 2 6 10
Western Adirondacks 6 l | > 32
Eastern Adirondacks 14 13 71 -| 6
Lake George 5 5 22 2 =)
TOTAL af 25 98 19 50
abundance (< 0.5%) and very patchy distributions (CV > 200%).
Many of these minor species were restricted to special habitats
(i.e., swamps, dry ridges, sand plains, riparian galleries, mountain
slopes) and only three (chestnut, tamarack, cedar) with modest
maximum abundance (> 6%) were locally common in particular
habitats. Despite being distinctive indicators in the flora, all the
remaining minor species averaged less than 1.2% abundance even
when present, and were inconsequential to the prevailing com-
position of the forest
Dynamics. The presettlement surveys provide a static view
of the forest development at the time, but the surveyors also in-
dicated past disturbances in the forest. Lotting surveys commonly
included ‘‘dead”’ or “dry” trees and “‘stubs”’ or “‘stumps”’ indi-
cating a consistent low level of disturbance. The resulting forest
often had numerous “‘staddle”’ (sapling) trees cited, but trees very
rarely became large enough to merit the surveyor’s “great”? mod-
ifier. Significantly, there were 153 lots with instances of larger
“burns” or “‘windfalls’” worth recording (Table 4). Fire was the
most prevalent disturbance with some two-thirds of the highly
disturbed lots being burned. For example, in 1749 Peter Kalm
(1987) noted that on the western shore of Lake Champlain ‘“‘the
mountains are covered with trees, but in some places the forests
have been destroyed by fire.’ This is exactly the area in the
Hudson-Champlain corridor where fire was most frequent. Be-
yond this valley, or in the under-cited Merrimack Valley, which
obviously was an often burned “‘great pitch pine plain,”’ fire had
an extremely low frequency in the mountains of northern New
2000] Cogbill— Vegetation of Presettlement Forests 269
. Ratio of relative tree density in 85 towns across Vermont about
1800 to statewide FIA (Frieswyk and Malley 1985) relative tree density in
1983.
Increases Neutral Decreases
(+) —
Soft maples 25 Hard maples 1.3 Beech 02
White birches 23 Dry ashes 1.6 White oaks 0.2
Poplars 7 Sweet birches 1.0 Basswood 0.25
Cedar 4 Red oaks 0.9 Hard pines 0.4
Fir 4 Spruce 0.9 Wet ashes 0.4
Soft pines 2.3. Hemlock 0.7
Elm 0.7
England and the western Adirondacks (Table 4). Evidently, cat-
astrophic fires were restricted to sandy or rocky substrates, and
generally near the settlement frontier. In contrast to fire, windfalls
were found regularly across the region. Although commonly cov-
ering several lots at once, windfalls were smaller and more diffuse
than burns. For example, in 1816 surveyor John Richards (Field
Books, Vol. 4, NYSA) found ‘‘All the timber standing on it are
large and thrifty, with very few exceptions, the wind has made
havock [sic] among the timber in many places of [Township #
42]’’. Here in the western Adirondacks wind disturbance reached
its maximum frequency (Table 4) and the pattern has been con-
tinued with the repeated blowdowns of 1950 and 1995 at the same
site. As a result of the clumped and restricted distribution of
burns, an equal number of towns was affected by fire (13%) as
by wind (11%). Overall disturbances large enough to deserve
mention, however, affected only 21% of the towns. In the affected
towns an estimated 2.5% of the area was in burns or windthrow;
overall roughly 0.5% of the region was affected by major distur-
bances at settlement.
The presettlement forest composition is a unique baseline for
documenting the effects of land use in the region. Although all
the species of the early forest were still prominent by 1983, the
composition of the forests in Vermont have changed dramatically
since 1800 (Table 5). Species of younger forests associated with
the aftermath of human activities (i.e., soft maples, white birch,
poplars) have increased by two orders of magnitude (up to
2500%). Even white pines have more than doubled in frequency,
apparently due, in part, to the net gain between the loss due to
270 Rhodora [Vol. 102
harvesting and the regrowth in abandoned fields. Several species
that originally grew in richer lowlands (i.e., white oaks, bass-
wood, wet ashes) have also declined substantially (down to 20%
of the original). Their maximum abundance was on the most pro-
ductive land, which was intensively cleared and often remains
unforested today, such as the Champlain Valley. Several species
have remained roughly unchanged over the 200 years. Some of
this is a balance between harvesting and woodlot improvement
(maple) or a natural tendency for regeneration (ashes, sweet
birches). Spruce has had substantial decline at mid-elevations due
to climatic changes and forest harvesting, but this loss has been
nearly balanced by substantial gains in the valleys due to regen-
eration in old fields (Hamburg and Cogbill 1988). The most dra-
matic change over the past 200 years is the loss of the absolute
dominance of beech to 20% of its presettlement abundance. This
decline is apparently not due to recent bark disease, to over uti-
lization for wood, to lack of regeneration, or to land clearance.
As first pondered by Siccama (1963, 1971), the reason for the
incredible amount of beech in all northeastern presettlement sur-
veys, and its subsequent decline, remains an enigma.
DISCUSSION
Accuracy. Quantitative analyses of the survey records de-
pend on the data being an accurate estimate of tree composition
within the towns. Lotting and outline surveys of proprietory
towns are not a random sampling of the trees at the time; how-
ever, the survey design did produce samples in quasi-regular pat-
tern at locations determined a priori and covering the whole
town. As with much historical data, the methods were poorly
documented, coverage was incomplete, and the observations were
uncontrolled. For example, in 1772 in surveying the town of
Mansfield, Vermont, Ira Allen (1928) professed that ‘“‘(a) great
proportion of said lots were made on spruce or fir trees, and if I
described them as such, it would show the poorness of the town.
In my survey bills I called spruce and fir gumwood, a name not
known to the [proprietors]’’. Contrary to his claim, Allen’s own
proprietors’ survey (Mansfield Proprietors’ Book, Stowe [VT]
Town Hall) shows 18% spruce and no ‘‘gumwood” at all. Nev-
ertheless, the proprietor’s surveys were done by numerous sur-
veyors, over many years, with little incentive to skew the results.
2000] Cogbill—Vegetation of Presettlement Forests 271
In northern Vermont outline surveys, the corner-to-tree distances
were Statistically equal for all major species (Siccama 1971). Ap-
parently in these systematic surveys there was little bias in the
choice of trees (Bourdo 1956; Whitney 1994) and spatial bias, if
any, was toward the more detailed surveys (e.g., lower reaches
of the towns with the smallest lots), exactly the areas in town
later most affected by settlement. At face value, the lotting tree
tallies are a statistical sample and the relative frequencies are a
consistent and unbiased estimate of overall composition of the
forests at the turn of the 1700s.
Vegetation scale. The patterns of tree distribution exist at
three distinct, albeit nested, vegetation scales: the community or
forest type (~ 10 7 km’), the landscape or local combination of
communities (~ 10° km?), and the regional or zonal arrangement
of these landscapes (~10° km’). The town grain size (nominally
10° km?) is fixed by the mechanics of the presettlement surveys,
but conveniently preserves species variation at the landscape
scale (Delcourt and Delcourt 1996). The town-wide sample nec-
essarily averages tree abundance over multiple forest types, but
is an ideal size to reflect the local proportion of trees in those
types. Thus the town sample is appropriate for the characteriza-
tion of the landscape composition and advantageous for quanti-
fying regional patterns. The minimum of 50 trees per town is low
(Bourdo 1956) and limits the detection of infrequent species. Re-
stricted types or infrequent species are incompletely sampled, but
the analyses are accurate for the common species responsible for
gross vegetational patterns. Moreover, many of the towns had
large samples (> 400 trees) and this accounts for some estimates
of range and abundance of uncommon species.
In mountainous or hilly terrain each town captured much of
the elevational variation, so the town-wide data tend to cloud any
elevational gradients. Moreover, each town supported many of
the species in the region in a range of communities. Thus within-
town variability was high compared to between-town variability.
Therefore it is advantageous to have multiple samples within bio-
physical regions to elucidate regional patterns. The 179 towns in
the region showed major range and abundance distributions not
seen in the previous isowit maps derived from only 14 samples
(Whitney 1994). Although the gross levels of common species
abundance are similar, spatial and quantitative resolution is miss-
232 Rhodora [Vol. 102
ing. For example, the small-scale maps (Whitney 1994) misrep-
resent the actual patterns: the oak dominance in the Merrimack
Valley, the lack of pine on the uplands, the large amount of hem-
lock in the eastern Adirondacks, and the substantial presence of
spruce in southwestern New Hampshire.
Vegetation types. The vegetation of the region varied from
oak—pine in the warm southern valleys to beech—maple to spruce—
fir in the northern mountains. In 1741, Richard Hazzen (1879),
while surveying the northern boundary of Massachusetts near
Whitingham, Vermont, found the land ‘‘exceedingly good and
covered with Beach, Maple, Chestnutt &c. .. . the pigeon’s nests
were so thick that 500 might have been told on the beech [and]
Hemlocks as well.’ The beech, maple, and hemlock still domi-
nate, but the chestnut has been functionally eliminated and the
pigeons are gone completely. At the opposite extreme of the com-
positional gradient, John Richards (1816, Field Books, Vol. 4,
NYSA) while surveying Townships # 42 and 43 (now Five Ponds
Wilderness) in the western Adirondacks saw ‘‘much fine spruce,
yellow birch, beech, and maple . . . with few white pine and
black cherry trees . . . [and an] abundance of the finest spruce
and yellow birch on this land of any perhaps in the world.’ This
was and remains the archetype of a red spruce—hardwood land-
scape in the Northeast. Even in this mixed-hardwood vegetation
there was much local variation. The richer sites had more maple
and less spruce. Thus in 1773 in Norbury, New York (now Calais,
Vermont), Samuel Gale (Surveyor’s General Book, Vol. 38,
NYSA) found ‘‘choice land timbered with maple, beech, bass,
some elm, ash, birch & in patches some butternuts, with Maid-
enhair and some nettles.”
The ranges of the five dominant taxa in northern New England
and New York overlapped in a broad zone, but they did not form
a single landscape pattern. In the presettlement forest, beech was
predominant and formed a series of conifer—northern hardwood
types. Significantly, spruce was the typical conifer and neither
white pine nor hemlock typified the entire zone (cf. Braun 1950).
Although there were distinct regional variations (e.g., maple in
Vermont hills, spruce in the western Adirondacks), numerous
towns from all three states had a spruce—maple—beech composi-
tion. However common this central type, admixtures of secondary
species caused the vegetation composition to diverge from this
2000] Cogbill—Vegetation of Presettlement Forests Pa i)
hub in three primary directions. One spoke was toward colder
moosewood-fir—spruce towns of the north, the mountains, or the
western Adirondacks; the second spoke was toward drier chest-
nut—hickory—poplar—oak ridges of towns in the Taconics-Lake
George region; and the third spoke was toward the oak—pine low-
land towns of the Merrimack Valley. Within this primary pattern,
there were prominent variations, such as the abundance of hem-
lock in the eastern Adirondacks or the rich hardwoods (i.e., ashes,
butternut, buttonwoods) of the Champlain Valley.
The one major vegetation boundary was the dramatic discon-
tinuity between beech dominance on the uplands and oak—pine
dominance in the major southern valleys. This rapid transition is
akin to the “tension zone’’ between the prairie woodlands and
the northern forest in Wisconsin (Curtis 1959). The similarity
might even extend to the role of fire in maintaining the boundary.
In the lower hills of the Taconics and southwestern New Hamp-
shire there was an equivalent “‘oak—beech”’ tension zone at the
edge of the Hudson and Merrimack Valleys. This ecotone marked
a switch in dominance, as well as the coincidence of the general
range limits of spruce, yellow birch, white oak, chestnut, and
pitch pine. This major vegetation shift over a relatively short dis-
tance was even more surprising given the moderate elevational
relief. A less distinct version of this tension zone (“‘pine—spruce’’ )
extended around the Champlain Valley and weakly up the Con-
necticut Valley. Due to the condensing of the elevation gradients
and limited high elevation land, the distinct altitudinal (“‘conif-
erous—deciduous’’) ecotone was smoothed across towns in the
presettlement compositions (Cogbill and White 1991).
Historical methodology. The lotting witness tree surveys
from northern New England and New York are an empirical rep-
resentation of the natural vegetation before confounding of land
use. The presettlement dating, quantitative enumeration, unbiased
estimates, and town-wide scale, are all unique advantages of this
resource. Combined with the extensive available archival record,
this tree composition database effectively documents the regional
composition of the early forest. The summary isowits give higher
resolution and temporal control than similar “‘isopoll’? maps de-
rived from paleohistorical sampling. This summary of regional
vegetation, however, is still limited by its composite composition
and landscape scale. Utilizing exact tree locations from lotting
274 Rhodora [Vol. 102
maps within individual towns would produce a truly detailed and
spatially explicit view of the 18th century vegetation.
ACKNOWLEDGMENTS. John Burk of Harvard Forest helped find
and collate many of the New Hampshire records. I thank the
many librarians, town clerks, and repository staff, especially at
the New York State Archives, Vermont Historical Society Li-
brary, Vermont State Archives, Vermont Public Records Office,
New Hampshire State Library, New Hampshire State Archives,
and the New Hampshire Historical Society, for access to collec-
tions, retrieval of documents, and patience in helping to use the
manuscripts in their care. A special thanks to Greg Sanford, Ver-
mont State Archivist, for encouragement ever since a seminal
visit in 1983.
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Bourbo, E. A., JR. 1956. A review of the general land office survey and of
its use in quantitative studies of former forests. Ecology 37: 754-768.
BRAUN, E. L. 1950. Deciduous forests of eastern North an Hafner
ress, New York, NY.
CHAMPLAIN, SAMUEL DE. 1609 asia 1925]. Voyages du Sieur Champlain,
pp. 91-92. In: H. P. Bigger, , The Works of Samuel de Champlain,
Vol. 2. The Champlain ae Toronte. ON, Canada.
CLARK, C. E. 1983. The Eastern Frontier, the Sieg of Northern New
England, 1610-1763. University Press of New England, Hanover, NH.
CoaBILL, C. V. 1996. Black growth and soe The nature of old-growth
red spruce, pp. 113-125. In: M. B. ed., Eastern Old-Growth For-
ests, Prospects for Rediscovery and aa Island Press, Covelo, CA.
AND P. S. WuiTe. 1991. The latitude-elevation relationship for spruce-
fir forest and treeline along the Appalachian mountain chain. Vegetatio
94: 153-175.
CoGsweELL, L. W. 1880. History of the town of Henniker, Merrimack County,
w Hampshire. Republican Press Association, Concord,
CURTIS, “ T. 1959. The Vegetation of Wisconsin. Univ. Wisconcin Press,
Madison, WI.
DeLcourt, H. A. AND P. A. DELCOURT. 1996. Presettlement landscape hetero-
geneity: Evaluating grain of resolution using General Land Office Sur-
vey data. Landscape Ecol. 11: 363-381.
Foster, D. R., G. MoskIN, AND B. SLATER. 1998. Land-use history as long-
term broad scale disturbance: Regional forest dynamics in central New
England. — ems |: 96-119.
Frieswyk, T. S. AND A. M. MALLEY. 1985. Forest statistics for Vermont, 1973
and 1983. U.S.D.A. Forest Service, Resource Bull. NE-87
2000] Cogbill—Vegetation of Presettlement Forests 219
GLeASON, H. A. AND A. Cronguist. 1991. Manual of Vascular Plants of
Northeastern United States and Adjacent Canada, 2nd ed. The New York
Botanical Garden, Bronx
HAMBURG, S. P. AND C. V. CocsuL. 1988. Historical decline of red spruce
populations and climatic warming. Nature (London) 331: 428—431.
Hazzen, R. 1741 [reprinted 1879]. The boundary line on New Hampshire
and oo New England Historical & Genealogical Register 33:
323-33
IRLAND, L. a 1999. The Northeast’s Changing Forest. Yale Univ. Press, New
Haven, CT.
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northern New England. Rhodora 102: 246—247.
oe yn, J. 1675 [reprinted 1833]. An account of two voyages to New Eng-
land. Mass. Historical Soc. Coll., 3rd Ser. 3 354.
Kam, P. 1770 [edited and translated by A. B. Benson, 1987]. Peter Kalm’s
Travels in North America. Dover Publ., New York.
LONGFELLOW, H. W. 1854 [reprinted 1886]. Evangeline, a tale of Acadie. Jn:
Works. Riverside Press, Boston, MA
Lorimer, C. G. 1977. The presettlement forest and natural disturbance cycle
of northeastern Maine. Ecology 58: 139-148
Lutz, 5,-G. 1997, io aay ae settlement and present forest composition in
Kings County, New Brunswick, re M.F. thesis, Univ. of New
Brunswick, Fredericton, NB. Can
Marks, P. L. AND S. Garbescu. 1992. ia of the central Finger Lakes
Region of New York in the 1790s. New York State Mus. Bull. 484: 1-35.
McIntosu, R. P. 1962. The forest cover of the Catskill Mountain region, New
York, as oad by the land survey records. Amer. Midl. Naturalist
Moss, M. R. AND P. a. HoskING. 1983. Forest associations in extreme south-
erm Ontario ca. 1817: Biogeographical analysis of Gourlay’s “‘Statistical
cco . Canad. edna 27: 184-193
PACALA, 3.1 , C. D. CANHAM, AND J. A. SILANDE R, JR. 1993. Forest models
by 7. measurement: I. The design of a northeastern forest simulator.
anad. J. Forest Res. 23: 1980-1988.
PIKE, R. E. 1967. Tall Trees, Tough Men. W. W. Norton Co., New York.
Price, E. T. 1995, Dividing the Land. Univ. Chicago Press, Chicago, IL.
SCHWARZ, M. W. 1994. Natural distribution and abundance of forest species
nd communities in northern Florida. Ecology 75: 687—705
SEISCHAB, E K. 1990. Presettlement forests of the Phelps and Gorham Pur-
e in western New York. Bull. Torrey Bot. Club 117: 27-38
. 1992. Forests of the Holland ae oo in western New York,
ca. 1798. New York Mus. Bull. 484:
SiccaMA, T. G. 1963. Pre- a and tee forest cover in Chittenden
County, Vermont. M.S. thesis, Univ. Vermont, Burlington, VT.
mae aes o ne and al forest vegetation in northern Ver-
mont with special reference to Chittenden County. Amer. Mid]. Natu-
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SPEAR, R. W. 2000. Implications of post-glacial changes in climate and veg-
etation on the flora of the White Mountains, New Hampshire. Rhodora
102: 248-249
TORBERT, E, 1935. Evolution of land eras in Lebanon, New Hampshire.
Geogr. Rev. (New York) 25: 209-2
White, C. A. 1984. A history of the eae survey system. U.S. Dept.
Interior, Bureau of Land Management. U.S. Government Printing Office.
Washington, DC.
Wuitney, G. G. 1994, From Coastal Wilderness to Fruited Plain. Cambridge
Univ. Press, New York.
WINER, H. 1955. History of the Great fase Forest, Litchfield County,
Connecticut. Ph.D. dissertation, Yale Uni ‘ a Gb
WoopDarD, FE M. 1936. The Town Proprietors of ae The New England
Town Proprietorship in Decline. Columbia Univ. Press, New York.
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RHODORA, Vol. 102, No. 911, p. 277, 2000
FIFTY YEARS OF CHANGE IN RHODORA AND THE NEW
ENGLAND FLORA
WARREN H. WAGNER, JR.
Department of Biology, University of Michigan, Ann Arbor, MI 48109
SUMMARY. The major changes from 1950 to the present involve
destruction of natural areas, effect of invasive species as well as
species new to New England, and improved taxonomies. The pe-
riod 1900-1950 was influenced mainly by the great Harvard bot-
anist M. L. Fernald. Rhodora was one of the earliest North Amer-
ican regional journals. New techniques seem to be overtaking
classical field and herbarium studies, and many of our field and
herbarium workers feel eclipsed and no longer of value. Graduate
students are giving up ambitions of becoming systematic botanists
because they are not interested in purely laboratory or computer
research. A comparison of studies of pteridophyte taxonomy be-
tween the period 1900-1950 and the present shows profound
changes. The future of our research in New England should in-
clude careful comparison with other parts of North America, for
example the western Great Lakes. There is still much to do, and
field and herbarium studies are as important as ever. Rhodora
continues to be an inspiration.
RHODORA, Vol. 102, No. 911, pp. 278-279, 2000
LINKING THE DEEP AND RECENT PAST TO THE
MODERN NEW ENGLAND LANDSCAPE
DaviID R. FOSTER
Harvard Forest, Harvard University, Petersham, MA 01366
e-mail: drfoster @ fas.harvard.edu
SUMMARY. The very long-term record provided by paleoeco-
logical studies indicates that rates of vegetation change during the
relatively brief period since European settlement are the greatest
since the last Ice Age. Interpreting the details of these changes
and their persistent effects on modern landscapes at a range of
spatial scales provides critical information for ecologists, conser-
vation biologists, and natural resource managers. At a regional
scale (1.e., New England excluding northern Maine) the landscape
was largely deforested, farmed intensively, and, over the past 150
years, allowed to reforest naturally to produce extensive semi-
natural woodlands interspersed with urban and suburban areas.
One consequence of this history is that many open-land plants
and animals, including several that are high priorities for conser-
vation, thrived during the agricultural 18th and 19th centuries and
have declined greatly over the recent past. In contrast, as forests
have expanded and continued to mature there has occurred a re-
markable expansion of native woodland species. At a sub-region-
al scale (1.e., north-central Massachusetts) this history is associ-
ated with a broad-scale homogenization of forest canopy com-
position. Although tree species abundance varied with climate
gradients at the time of European settlement no such relationship
exists today. In contrast, many herb and shrub species do exhibit
striking variation in modern distribution with climate across the
same area. Importantly, the change in tree composition apparently
involves two processes: the very long-term decline in species
such as hemlock and beech, which actually began more than 500
years ago in response to climate change, and a regional increase
in successional and sprouting species due to land use.
On a landscape scale, vegetation structure and composition are
apparently much more homogeneous and patchy than at European
settlement as they currently vary on a fine scale with land-use
histories. Site history, along with variation in soil conditions, is
a strong determinant of modern species distributions because
plant species vary so widely in their response and ability to re-
278
2000] Foster—The New England Landscape 279
cover and re-establish after, for example, fire, forest cutting, graz-
ing, or plowing. Stand-level pollen records suggest that the can-
opy composition of most forests was completely changed by this
history and bears little resemblance to earlier forests on the same
sites (Foster and O’ Keefe 2000)
Recognition that New England is a cultural landscape shaped
in most details by its history of intensive human activity is an
essential background for understanding modern ecological pro-
cesses. Interpretation of the details of this history at geographical
and temporal scales relevant to specific concerns can afford tre-
mendous insights into land management and conservation policy
(Foster 1999).
LITERATURE CITED
Foster, D. R. 1999. Thoreau’s Country: Journey through a Transformed
eens Harvard Univ. Press, Cambridge, MA.
Dp J. O’ KEEFE. 2000. New England Forests through Time: Insights
cae the Harvard Forest Dioramas. Harvard Univ. Press, Cambridge,
MA.
RHODORA, Vol. 102, No. 911, pp. 280-298, 2000
IMMIGRATION AND EXPANSION OF THE NEW
ENGLAND FLORA
LESLIE J. MEHRHOFF
George Safford Torrey Herbarium,
Department of Ecology and Evolutionary Biology, Box U- 43,
University of Connecticut, Storrs, CT 06269-3043
e-mail: vasculum @uc uconn.edu
ABSTRACT. Nonindigenous vascular plant species have been introduced,
intentionally or unintentionally, since Europeans landed in what is now New
England some time in 1496. We know little about the native flora of New
England at that time. John Josselyn’s New England Rarities Discovered re-
corded both the native and European plants he saw duri ie his two visits to
southeastern Maine and is the earliest report on the flora ¢
England. Subsequent writers, such as Manasseh Cutler, also documented both
the native and increasing number of non-native species that became natural-
ized in this region. This paper discusses both the intentional and noninten-
tional introductions from Europe and the later introductions from eastern
a. Various naa: of unintentional introductions such as ballast plants and
eae aways” are presented. Species that are native to other re-
gions 0 cen een and that have naturalized in New England are men-
tioned. Currently, over 1000 vascular plant species that are not considered
Seen to the region exist in the New England flora. A few introductions
have become so aggressive in sane establishment around New England that
they 2 now acknowledged as invasive species. Early botanical works an
nia records are used here to document arrivals and changes in the flora.
2
Key Words: introductions, non-native plants, nonindigenous plants, New
England, flora
The flora of New England is a mosaic of native and non-native
species. The ratio of native to non-native species varies from
habitat to habitat, site to site, and time to time. Nonindigenous
species have been arriving since the earliest European explorers
set foot on New England shores. While some non-native species
arrived accidentally, many were brought here for utilitarian or
aesthetic reasons. Not surprisingly, the earliest introductions into
New England were native to Europe, later ones coming from
other regions of North America, Eurasia, Eastern Asia, or else-
where.
The current New England flora is composed of between 24 to
45 percent nonindigenous species (Table 1). These percentages
are only approximations because of different taxonomic circum-
280
2000] Mehrhoff—Immigration and Expansion 281
Table 1. Tabular summary of species by state. 'Seymour 1969; 7Dowhan
1979, Mehrhoff 1987, 1995; *Gould et al. 1998; +Sorrie and Somers 1999;
‘Campbell et al. 1995.
Non-native Percent Non-
native
State/Region Total spp. Native spp. spp
New England! 2882 1995 887 31%
Connecticut? 2625 1700 925 35%
Rhode Island? 1618 1226 392 24%
Massachusetts? 2814 1538 1276 45%
Maine 2103 1469 634 30%
scriptions, nomenclature, different appraisals of what is consid-
ered naturalized, and recent discoveries. Published works vary
depending on nomenclatural sources. Seymour’s Flora of New
England (Seymour 1969) and Dowhan’s Checklist for Connecti-
cut (Dowhan 1979) follow Fernald’s nomenclature (Fernald 1950)
for most treatments. The Vascular Flora of Rhode Island (Gould
et al. 1998) follows Cronquist (Gleason and Cronquist 1991) and
Flora of North America (Flora of North America Editorial Com-
mittee 1993+). Massachusetts’ county checklist (Sorrie and Som-
ers 1999) follows a mixture of Kartesz’s nomenclature (Kartesz
1994) and that of the Flora of North America Project (Flora of
North America Editorial Committee 1993+). Maine’s checklist
(Campbell et al. 1995) uses a variety of additional sources in-
cluding experts who are preparing taxonomic treatments for the
Flora of North America Project.
In New England, Rhode Island appears to have the lowest per-
centage of nonindigenous species, 24% (Gould et al. 1998), while
Massachusetts appears to have the highest, 45% (Sorrie and Som-
999). Published current figures are not available for Vermont
or New Hampshire.
There are two complimentary ways of evaluating the history
of the nonindigenous components of the New England flora. One
way of approaching the expansion of the flora is temporal. The
other is phytogeographic. Historical documents shed light on the
increase in non-native species over time. Concurrently, there are
elements of the introduced flora known to represent different phy-
togeographic origins. Although there were periods of introduc-
tions from different geographical regions, the temporal compo-
nent and the phytogeographic component do not exactly coincide.
Separating the two can be difficult because certain Asian taxa,
282 Rhodora [Vol. 102
such as Tree-of-Heaven, Ailanthus altissima (Mill.) Swingle, were
introduced into North America from botanic gardens in Europe
(Spongberg 1990).
TERMINOLOGY
For this paper I set the bounds of New England to be the
cumulative political boundary of the six New England states.
While this boundary is admittedly artificial, it helps clarify the
meaning of the native and non-native.
Native or indigenous are used here for those species that ex-
isted within this boundary prior to AD 1496 when the Italian
explorer John Cabot, sailing for King Henry VII of England,
landed on what are now New England shores (Newby 1982).
Native taxa are often mentioned in the early botanical literature
for New England. Herbarium collections do not exist that docu-
ment these early reports. In fact, the earliest herbarium collections
for New England that still exist (at least in North American her-
baria) appear to be from around the beginning of the 19th century.
Most native taxa are North American endemics although some
exhibit amphiatlantic or cosmopolitan distributions. Taxa that nat-
urally occurred in the region near New England and recently ar-
rived here by means of their own adaptations without the aid of
human intervention are also considered native. Eupatorium album
L., is considered native to Connecticut although it was only dis-
covered there in 1981 (Mehrhoff 1996). It had been known for
many years from Long Island (Miller and Young 1874) and is
wind dispersed. Its discovery in southeastern Connecticut was not
surprising.
Non-native or nonindigenous species as used here are taxa that
appear to have arrived in New England sometime after AD 1500.
Most of these are known io have extra North American origins.
The majority of these taxa arrived with aid, intentional or acci-
dental, from humans. As many species were intentionally intro-
duced and subsequently escaped and became established here,
there is often a known history of their introduction. In addition,
many of these are known to be native elsewhere and their oc-
currence here accepted as human-assisted. There are no herbarium
records until much later and often, as a newly discovered species
is noteworthy, there may be numerous collections attesting to its
recent discovery and novelty.
2000] Mehrhoff—Immigration and Expansion 283
Introduction is used to describe an event. By itself it implies
neither intentional introduction nor accidental introduction. These
modifiers should be used when the history of an introduction is
known or clarity of thought is necessary.
Naturalized is used to designate non-native taxa that are estab-
lished, reproducing and persisting without human intervention
and cultivation. Often, establishment can occur within natural
plant communities. Many non-native taxa occur in New England
but cannot be considered as naturalized because they must have
human intervention in order to persist. Further, naturalized im-
plies persistence over time. Some species not considered natural-
ized may exist away from cultivation for a year or two but pop-
ulations do not establish and persist for long. These should be
considered adventive (Fernald 1950).
Garden escapes are those taxa that originally were intentionally
introduced as garden plants and subsequently became naturalized
away from cultivation. The term garden escape here is used only
for taxa that are completely naturalized into the New England
flora, not for adventives. Notations on old herbarium specimens
often indicate “in garden,” “‘near garden,” “‘escaped from gar-
en,’ or “‘established.”’
Occasionally perceptions of a plant’s desirability change when
a garden plant escapes and becomes naturalized away from gar-
dens. Fernald (1940) tells how Hieracium aurantiacum L. was a
prized garden plant in the central Maine of his youth and was
then known as Venus’ Paint-brush. Once it had escaped and be-
come established away from gardens it was often seen growing
aggressively in these new sites. After a while, its colloquial name
had changed to Devil’s Paint-brush.
Invasion, invasive species, and invasives are used to imply both
an arrival event and subsequent establishment and proliferation.
These terms are only used here in reference to non-native species.
Rapid spread or aggressive growth and proliferation are implicit
with invasive species. No inference should be drawn about the
arrival event; it can either be by the biological attributes of the
species or with human assistance. The use of explosive species
or native explosive species in reference to native species that
exhibit the characteristics of invasive species might help avoid
confusion.
A weed, commonly described as a plant growing where it is
not wanted, can be native or non-native (Les and Mehrhoff 1999).
284 Rhodora [Vol. 102
For this reason, and because personal preferences and biases exist,
the term weed is not used here. Weedy, however, is a good de-
scriptive term and clearly understood by most to imply rampant
growth.
Nomenclature used here follows Cronquist (Gleason and Cron-
quist 1991) or the published volumes of the Flora of North Amer-
ica Project (Flora of North America Editorial Committee 1993+).
HISTORY OF INTRODUCTIONS
Early accounts of the pre-colonial flora are biased by interpre-
tation (Whitney 1994) and rarely go beyond generic descriptions
of the forest. The earliest written account of the flora was that of
John Josselyn, an Englishman who published two books on the
natural curiosities of the New World in the late 17th century. No
one knows the dates of Josselyn’s birth or death but we do know
he twice visited his brother who lived in the region that is now
Saco, Maine. The first visit was in 1638, when he stayed for 15
months. His second visit was in 1663, this time lasting for eight
years (Tuckerman 1865). During these two visits he recorded his
observations on the wildlife and flora he encountered. Shortly
after his return to England in 1671, he published New England
Rarities Discovered in Birds, Beasts, Fishes, Serpents, and Plants
of that Country (Josselyn 1672).
In New England Rarities Discovered, Josselyn, as the title sug-
gests, discussed five groups of organisms found in New England.
Within the plants, he further divided his listings into five subdi-
visions: ““Of such Plants as are Common with us in England,”
“Of such Plants as are proper to the Country,” ‘‘Of such Plants
as are proper to the Country, and have no Name,” “Of such
Plants as have sprung up since the English planted and kept Cattle
in New England,” and “of such Garden-Herbs amongst us as do
thrive there, and of such as do not” (Josselyn 1672). Interspersed
throughout the text are uses for the plants and animals about
which he was writing. The author, with crude line drawings, il-
lustrated nine of the plants. Josselyn’s use of vernacular names is
often confusing or difficult to decipher. He probably used John-
son’s edition of Gerard’s Herbal from 1636 as his source of in-
formation (Tuckerman 1865).
New England Rarities Discovered represents the first exposi-
tion of the New England Flora. In addition, it sets a benchmark
2000] Mehrhoff—Immigration and Expansion 285
for dates for early nonindigenous introductions. Josselyn’s fifth
section on plants is of interest as a list of garden plants that may
represent one of the earliest accounts of what plants were culti-
vated for food by early settlers.
Josselyn’s first section, ““Of such Plants as are Common with
us in England” includes native widespread species such as Typha
latifolia L. that naturally occurred here as well as in Great Britain.
Other taxa included in this first list are now considered to have
conspecific species on either side of the Atlantic. Josselyn in-
cluded here a number of species now considered nonindigenous
in New England. One can infer, as Tuckerman (1865) did, that
these nonindigenous taxa must have been introduced early in co-
lonial history because they were so well established by the time
of his visits that Josselyn mistook them for natives.
Josselyn used vernacular names known to him. Some of these
such as ‘“‘Hollow-leaved Lavender” (Sarracenia purpurea L.),
‘“Rupter-wort”” (Euphorbia sp.), or Trackle-berries” [Smilacina
racemosa (L.) Desf.] are no longer used (Tuckerman 1865). It is
often difficult to decide which taxon was meant by some of Jos-
selyn’s names. In 1865, Edward Tuckerman published an anno-
tated version of New England Rarities Discovered. In this, he
attempted to identify, using contemporary scientific names, all of
the taxa included by Josselyn. Tuckerman’s interpretations are
extremely helpful though he was not always clear about the spe-
cies to which Josselyn was referring. He attempted to interpret
Josselyn’s names in light of what was known about European and
North American floristics at that time. For example, Tuckerman
assumed that when Josselyn recorded St. John’s-wort, he probably
meant Hypericum perforatum L., now assumed by most botanists
to be introduced here. However, he commented that Josselyn
could have meant Hypericum corymbosum Muhl. (now Hyperi-
cum punctatum Lam.).
Josselyn’s fourth section, ““Of such Plants as have sprung up
since the English planted and kept Cattle in New England,”’ is
the most interesting section when considering the nonindigenous
flora. Here Josselyn listed 40 species that he felt were not native
to New England and were brought here, intentionally or uninten-
tionally, by Europeans. His section heading is interesting in that
it implies he associated the keeping of cattle with the arrival of
Puropesn ee Seeds of many species are known to have been
“stowaways” with seeds intended for agricultural uses (Fernald
286 Rhodora [Vol. 102
1905). Perhaps Josselyn had some reason to suspect that seeds
were unintentionally introduced with livestock food or bedding.
In a footnote following this section Tuckerman pointed out taxa
mentioned mostly in Josselyn’s first section that belong here.
The first and fourth parts of Josselyn’s plant lists are of interest
as lists of plants that had been introduced from the Old World by
this time. These lists help narrow the period during which the
taxa Josselyn included here were introduced. Given the state of
floristic botany in the late 17th century, it is not surprising Jos-
selyn included with his native species, taxa now thought to be
introductions. For instance, Josselyn’s Wild purcelane [sic], was
thought by Tuckerman (1865) to be Portulacca oleracea L., a
native of Europe. This report establishes this taxon as part of the
flora of New England at a very early date. Josselyn’s inclusion
of Herb Robert in his first section is interesting. Geranium rob-
ertianum L. was considered by Tuckerman (1865) to be ‘‘com-
mon to us and Europe’. Eastern North American populations
have been viewed as native here by Fernald (1950) but natural-
ized by Cronquist (Gleason and Cronquist 1991). Many field bot-
anists consider it a good indicator of rich, shaded colluvial slopes
and cool, mesic woodlands. In western North America popula-
tions of G. robertianum are viewed as non-native and invasive
(Brumback, pers. comm.). Josselyn’s inclusion of this species in
this section suggests it should be considered to be native in New
England since it is unlikely it would have become so well estab-
lished in such specific natural habitats in the short time after
Europeans arrived here.
The second written record of plants, both native and non-na-
tive, existing in New England, is that of Manasseh Cutler. Cutler
was born in Killingly, Connecticut, in 1742, educated at Yale
College, and became a pastor in Ipswich Hamlet, Massachusetts,
where he lived until his death in 1823 (Humphrey 1898). In spite
of remaining in one town for 52 years, Cutler was far from sed-
entary. We know from his correspondence and diaries (Cutler and
Cutler 1888) that he traveled widely throughout New England,
collecting as he traveled. A diary entry from July 2, 1787 re-
counts how while traveling from Middletown, Connecticut, to
New Haven he examined several plants he had collected, ‘‘for
the heat was too intense for riding” (Cutler and Cutler 1888).
Unfortunately, Cutler’s large herbarium was destroyed by fire
(Day 1901).
2000] Mehrhoff—Immigration and Expansion Zot
Cutler’s “‘“An Account of some of the vegetable Productions,
naturally growing in this Part of America, botanically arranged
[sic] was published in the first volume of the Memoirs of the
nascent American Academy of Arts and Sciences (Cutler 1785).
‘‘Botanically arranged”’ was according to the new Linnaean sex-
ual system. In his introductory paragraphs, Cutler (1785) ex-
plained that he undertook this listing of plants from “‘this part of
America”’ because he felt that while ‘‘Canada and the southern
states ... have been visited by eminent botanists from Europe”
there had been ‘‘almost total neglect of botanical enquiries [sic],
in this part of the county’. He blamed this on the fact “that
Botany has never been taught in any of our colleges, and to the
difficulties that are supposed to attend to it; but principally to the
mistaken opinion of its inutility in common life” [his italics].
Later he commented, ‘‘From the want of botanical knowledge,
the grossest mistakes have been made in the application of the
English names of European plants, to those of America.”” Cutler
was well aware of nonindigenous plants in the landscape. On this
subject he wrote, ‘“We have it, also, in our power, from the recent
settlement of the country, to determine, with great certainty, what
vegetable productions are indigenous, and present those doubts
and disputes hereafter, which have frequently taken place among
botanist in old countries. For it is very improbable that any exotic
plants are become so far naturalized as not to be distinguishable
from the natives.”
Cutler reported 66 European species established in New Eng-
land. He made no attempt to correct the confusion of using Eu-
ropean names for North American taxa. Because of this, some of
his taxa must be suspect. Under Ornithogalum, he said about
what he called Bethlemstar [sic], ““Blossoms yellow. Common in
grass lands and amongst bushes.”’ The European O. umbellatum
L., now commonly known as Star-of-Bethlehem, has white tepals.
The native Hypoxis hirsuta (L.) Coville, common in New Eng-
land grasslands and open woods, has yellow sepals and petals
and was originally published as Ornithogalum hispidum by Lin-
naeus.
Possibly the most interesting inclusion is under the genus Car-
damine. Cutler gave the common names ‘“‘Impatient”’ and then
‘Impatient Ladysmock’’. These are followed by the comments,
“Blossoms yellowish white. By springs in mountainous land.”
The European C. impatiens L. has yellowish-white petals whereas
288 Rhodora [Vol. 102
most other Cardamine that occur in New England have white or
pinkish petals. The earliest herbarium specimen seen from New
England was collected in Peterborough, New Hampshire, in 1916.
It seems unlikely C. impatiens was here and established in Cut-
ler’s time as it is not included in the seventh edition of Gray’s
Manual (Robinson and Fernald 1908) and the eighth edition of
Gray’s Manual (Fernald 1950) has it only as local from southern
New Hampshire and eastern Pennsylvania. It is possible, but un-
likely, that Cutler was seeing C. hirsuta L. but this species was
not known in New England until recently. Cardamine parviflora
L. is native to Europe and represented in New England by its var.
arenicola (Britt.) O. E. Schulz, but this is usually a taxon of dry,
sandy soils and ledges, not of “springs in mountainous land’’.
Cutler was probably reporting the native C. pensylvanica Muhl.
that occurs commonly along streams, though this species usually
has sharply white petals. The true identity of this taxon and its
historical biogeography must await further elucidation.
Some of Cutler’s other inclusions are less obscure. Many notes
about non-native species are interesting in light of current distri-
butions. Ligustrum “is not very common in the wild state.’’ He
made no mention of which species, but it must have been, given
the time, the European Common Privet L. vulgare L. Thornapple
or Jimsonweed, Datura stramonium L. “is said to be an exotic,
and that it is not found growing at any great distance from the
sea.”’ Solanum dulcamara L. was *“‘Common about fences in
moist land.” Berberis, taken by me to be B. vulgaris L. because
of his comments “‘that rye and wheat will be injured by this
shrub, .. .”’ is said to be ““Common’’.
The next account of the region’s flora was Jacob Bigelow’s
Florula Bostoniensis or Plants of Boston published in 1814. Big-
elow included 83 introduced species in the first edition. By the
third edition, published in 1840, there are 140 nonindigenous
plants enumerated (Fernald 1905). In most cases, nonindigenous
species are not distinguished in the text. Occasionally an entry
will include a comment about a possible introduction. By the time
the second edition was published in 1824, the Black Locust, Ro-
binia pseudoacacia L. had become established in New England.
Not included in the first edition of Plants of Boston, Bigelow said
of it by 1924, **The Locust tree, exceedingly valued for the hard-
ness and durability of its timber, is not, I believe, found native in
the New England states, though abundantly naturalized near hab-
2000] Mehrhoff—Immigration and Expansion 289
itations and roads.” He went no further than to explain that it is
native to North America. It is commonly taken to have occurred
as far east and north as central Pennsylvania (Elias 1987). This
is indicative however, that by this time, people had started moving
species native to other parts of North America into New England
for utilitarian purposes.
Other floras produced in the first half of the 19th century add
other species to the growing list of non-native species that had
naturalized in New England. John Brace’s flora of Litchfield,
Connecticut (Brace 1822), includes both native and non-native
species. Likewise, in 1831, Dr. Eli Ives, a professor of materia
medica at Yale College, produced a list of plants growing without
cultivation in the vicinity of New Haven, Connecticut (Ives et al.
1831). Both authors included native and non-native taxa but did
not always distinguish between them.
The Massachusetts legislature commissioned a report on the
botany of the Commonwealth that was ultimately separated into
herbaceous plants by Chester Dewey (1840) and trees and shrubs
by Emerson (1846). While Dewey’s flora includes introductory
remarks under the heading “‘Of the Useless Plants”’ that would
lead one to believe he might have provided insight into some of
the introductions, he actually provided little beyond commenting
that a species is introduced, possibly introduced, or naturalized.
Similarly, Emerson included nonindigenous species but shed no
light on how they might have been introduced. These points ap-
parently show, however, that while cognizant of the presence of
non-native species, these botanists did not view them in a nega-
tive light.
Many collections made during this same period led to the nam-
ing of species of vascular plants from New England that were
new to floristic botany. It is interesting to note that the scientific
authorities for most of the taxa included in the early works on
the New England flora were Europeans. During the first half of
the 19th century, names of New Englanders such as Bigelow,
Ives, Oakes, Robbins, Hitchcock, and Dewey appeared as au-
thorities for New England plants. The species published by these
botanists added to the numerical expansion of the regional flora.
By the last half of the 19th century lists of non-native plants
by means of introduction were appearing in the literature. There
are a number of plausible explanations for this beyond the scope
of this paper, such as better communication between Europe and
290 Rhodora [Vol. 102
North America, better training in botany, and botanists who trav-
eled abroad and knew plants in native habitats as well as in their
naturalized condition. Perhaps, too, there was an increase in the
number of North American botanists, both professional and am-
ateur, who were actively cataloging the local flora.
The effort of cataloging non-native species was perhaps pio-
neered by Lewis D. de Schweinitz, whose ‘“‘Remarks on the
Plants of Europe which have become naturalized in a more or
less degree, in the United States” was published posthumously
in 1832 (Schweinitz 1832). Schweinitz’s work appears to have
focused primarily on New York and Pennsylvania. He separated
the 137 species he enumerated into 3 categories: 1) Plants which
have become more or less generally naturalized in the United
States; 2) Plants but partially spread; and 3) Introduced only in
the vicinity in which they are or were cultivated. He further di-
vided the more or less generally naturalized species into those
introduced by cultivation, for agricultural or other purposes, and
those introduced fortuitously with agricultural seeds (Schweinitz
1832)
SOURCES OF INTRODUCED PLANTS
Introductions occurring in the latter half of the 19th century
were either intentional or unintentional. Plants were intentionally
introduced as crops for humans or livestock, for natural products
such as dyes, foods, and other intentional uses, or for esthetic
reasons. Often these escaped and became naturalized. Robinson
(1880), in the introduction to The Flora of Essex County, told of
the prevalence of gardens for purely ornamental purposes. It is
in this period that we see the rise of botanical gardens that served
the multiple functions of education, research, and recreation.
Noteworthy among these was Harvard’s Botanical Garden in
Cambridge, begun in 1806 by William Dandridge Peck and taken
over by Asa Gray in 1842 (Dupree 1959), and later, in 1872, the
Arnold Arboretum (Hay 1995; Spongberg 1990). During this pe-
riod, the polymath Jacob Bigelow and others laid out the grounds
of Mt. Auburn Cemetery in Cambridge as a kind of botanical
garden.
Seed catalogs show that many non-native plants had been in-
troduced into the trade during the first half of the 19th century
(Mack 1991). Many well-known naturalized species were first
2000] Mehrhoff—Immigration and Expansion pas) |
introduced into New England as garden plants and later escaped.
Sometimes these ‘‘escapes’’ were aided by plant-growers. Both
Trapa natans L. and Marsilea quadrifolia L. were introduced into
the wild near Boston by Louis Gauerineau, the gardener at Har-
vard’s Botanical Garden (Les and Mehrhoff 1999). Other times,
garden plants escaped. Many early labels for collections of Vin-
cetoxicum nigrum (L.) Moench mention it as escaping from gar-
dens. It is interesting to speculate that the source of the first New
England specimen of this Swallowwort (BRU!), taken on the
streets of Cambridge, Massachusetts, in 1876, was the Harvard
Botanical Garden. Fernald (1900) explained how Artemisia stel-
leriana Besser probably escaped from late 19th century private
gardens in which it was a popular bedding plant.
Ship’s ballast was an early-recognized source of non-intention-
al introductions. Ships coming to the United States in order to
bring natural resources back to Europe would arrive with rocks
and dirt as ballast to be discarded before loading the valuable
cargo for the return trip. Many port cities had “ballast grounds”’
or ballast piles to which the jettisoned ballast would be contin-
ually added. These became favorite haunts of local botanists in
search of floristic novelties. One of the earliest works on this
subject was by Aubrey H. Smith on “‘Colonies of Plants observed
near Philadelphia’ (Smith 1867). This was followed by other
reports from the Philadelphia area (Burk 1877; Martindale 1876,
1877). In 1878, Judge Addison Brown began a series of five
articles on ballast plants collected around the port of New York
City (Brown 1878a, 1878b, 1879, 1880, 1881). Many of the
plants, especially in Brown’s lists for New York, occur in New
England and it is not inconceivable that they arrived here in the
same manner, given the thriving ports and navy yards along the
coast from Connecticut to Boston and downeast to Maine. In fact,
Smith (1867) referred to some of the ballast piles in Philadelphia
near where the ‘“‘coasters’’ docked.
In the early part of the 20th century a number of small papers
were produced on plants found in the vicinity of factories where
seeds or propagules would be introduced with the products with
which the factory dealt. Some of the best known of these kinds
of introductions were the plants found with “‘wool-waste”’. From
1901 to 1932 there were a series of articles in Rhodora, mostly
by Emily F Fletcher, dealing with plants found around woolen
processing plants near Westford, Massachusetts (Collins 1901;
292 Rhodora [Vol. 102
Fletcher 1912, 1913, 1915, 1916, 1917: Weatherby, 1924, 1932).
The composting wool-waste was later used to fertilize fields (Fer-
nald 1905).
Robinson (1880) mentioned introduced plants found along the
Merrimac [sic] River down-stream from Lowell and Lawrence.
An interesting group of non-native species was collected from the
waste pile of a rubber reprocessing plant in Waterbury, Connect-
icut (Blewitt 1911, 1912). Apparently, used shoes were collected
for rubber reclamation. The nonrubber parts were thrown on the
waste piles where seeds that had hitchhiked there germinated and
grew. In an interesting postscript that may explain why some of
these species did not persist, Blewitt said, ‘‘For the past two years
many plants in a portion of this place have been killed by fumes
of an acid factory while those that survive are badly seared and
burned by the deadly gases”’ (Blewitt 1911).
One of the most interesting cases of ‘‘factory-flora’’ was the
discovery of Lepidium latifolium L. on the grounds of a glue
factory in Danvers (Morse 1924). This European species is now
abundant in parts of the southwest. In New England, it was found
in eastern Massachusetts near the coast and at one inland site in
Worcester County. It also occurs along the southwestern Con-
necticut coastline where it was thought to have been introduced
at the site of a “‘dye and licorice works’? (Eames 1935). In ad-
dition, Paulownia tomentosa (Thunb.) Steud., Lepidium draba L.,
and Tamarix pentandra Pall. were reported from the same area.
Railroads brought adaptable species, often ones with weedy
tendencies, from the developing west. The now near ubiquitous
Black-eyed Susan, Rudbeckia hirta L., is thought to have come
east in that fashion. It had reached Philadelphia by 1826 and
probably New England by 1855 (Robinson 1880). Fernald (1905)
felt that Senecio jacobea L. arrived in Portland by way of the
railroad from New Brunswick
Unintentional introductions and the escape of intentional intro-
ductions continue. Froelichia gracilis (Hook.) Mog., having
reached New England by railroads, was first collected here in
1973. Although not currently known from New England, Mile-
a-minute vine, Polygonum perfoliatum L., was first reported in
Westchester County, New York, in 1995 (R. Mitchell, pers.
comm.). At that time it was well established and within a mile
of Connecticut. It seems plausible, since a natural dispersal from
the nearest known occurrence in eastern Pennsylvania was un-
2000] Mehrhoff—Immigration and Expansion 293
likely, that it arrived here at this site in nursery stock and escaped.
Lonicera maackii (Rupr.) Maxim. was first collected in the wild
in Connecticut in 1978. This species is occasionally cultivated
and it is likely, given its history as an invasive species in the
Midwest (Luken and Theiret 1996), that its numbers will increase
in the wild in southern New England.
BIOGEOGRAPHY OF PLANT INTRODUCTIONS
An equally informative way of looking at introductions is by
considering species from different geographic origins. Most of
the early introductions were of European plants that arrived with
or after the earliest settlers (Fernald 1905). This continued until
the opening of eastern Asia for trade after 1861 (Rehder 1936).
After this, while it is still likely that some European plants were
introduced into New England, most of the new introductions were
from regions in East Asia such as Japan and China (Spongberg
1990). As they came from similar climates and geological his-
tories, species from East Asia were well adapted to exist in New
England. Often imported as ornamentals, some of these escaped
and became quickly established in the local flora.
Again, the botanical gardens often provide the earliest records
for introduced plants. In New England, the Arnold Arboretum
was actively involved in plant importations from Japan and China
by the beginning of the 20th century (Hay 1995; Rehder 1936).
Rehder (1936) reported that after 60 years, the Arnold Arboretum
had introduced at least 2500 species from around the world. Prog-
eny of many of these reached American gardens.
A catalog of plants in the Harvard Botanical Garden, thought
to have been written in 1879 (J. Warnement, pers. comm.), in-
cludes both Elaeagnus umbellata Thunb. and Berberis thunbergii
DC. Although both are now considered highly invasive, at that
point they were well-behaved members of the Garden’s holdings.
As with many invasive species, there is a variable period after
introduction before offspring begin to appear in the wild.
Recent introductions from East Asia were not seen as a prob-
lem in 1905 when M. L. Fernald delivered his address on *“‘Some
Recently Introduced Weeds” to the Massachusetts Horticultural
Society (Fernald 1905). Fernald stated that the number of non-
indigenous plants in New England was then over 600 species.
Further, he discussed only European species in spite of the fact
294 Rhodora [Vol. 102
that East Asian species had been introduced by that time (Rehder
1936). Later, in a presentation to the Franklin Society in 1939
(Fernald 1940) Fernald devoted a number of pages to the prob-
lems faced by rare plants from aggressive non-native species. One
can infer from these two papers that in 1905 most Asian species
were hardly, if at all, dispersing away from managed landscapes
into the wild but that by 1939, many of the East Asian species
were escaping and becoming well established in the wild.
CONCLUSIONS
The New England flora has steadily grown since the arrival of
Europeans in the 17th century. While many species were inten-
tionally introduced for utilitarian reasons some were also inten-
tionally introduced for aesthetics. Many arrived unintentionally.
These unintentional means of transport were often quite varied.
Many, but not all species, persisted and are part of our regional
flora today. Others that may or may not have become naturalized
did not persist until present. As recently as the middle of this
century, nonindigenous species were being imported as foods,
medicines, or ornament (Rehder 1936). In addition, still other
species, considered native to adjacent regions, have naturally ex-
panded their ranges into New England. The most recent compre-
hensive list of New England vascular flora says there are 2882
vascular plant species reported from New England (Seymour
1969). Of these, 887 are considered nonindigenous (Seymour
1969). Given recent finds and the different nomenclature, these
figures must only be accepted as approximations and it is likely
that over 1000 species should be considered naturalized here.
Introductions can be looked at both from a historic perspective
and a phytogeographic perspective. While these approaches com-
pliment each other, clear divisions in each cannot be drawn. Rosa
multiflora Thunb., a native of eastern Asia, was first introduced
into the Elgin Botanic Garden in New York by way of European
botanical gardens in 1811 (Rehder 1936).
Currently, the few non-native species that are aggressively in-
vading natural plant communities are of paramount concern for
conservationists (Brumback 1998). These invasive species are
well known and exhibit biological characteristics of species
adapted to habitat disturbance (Mehrhoff 1998). Efforts must be
taken to control their spread. Concurrently, there are other non-
2000] Mehrhoff—Immigration and Expansion 295
native species that have the potential of becoming invasive in
New England and their status must be assiduously monitored.
One final note: times have changed since Manasseh Cutler la-
mented how few botanists studied the New England flora (Cutler
1785). We know as much about our flora as we do because, for
years botanists combing the fields, woods, and other habitats
traipsed all over New England. Now there again seems to be a
paucity of field botanists. Whether you ascribe to Eames (1935)
who “had the good fortune . . . to find great quantities” of Lep-
idium latifolium or to Morse (1924) who sensed that the same
species ‘‘seems to be liable to become a hardy weed of undesir-
able character’ is not the point. What is important is that these
two individuals had the ability and interest to recognize some-
thing new, to identify it, and to document its occurrence by col-
lecting herbarium specimens. If we want to continue to monitor
changes in the New England flora we must have botanists in the
field to do so.
ACKNOWLEDGMENTS. Many botanists have discussed some of
these ideas with me. Ray Angelo (NEBC) and David Boufford (GH
and AA) more than most, have helped me with herbarium ques-
tions as well as shared their knowledge of the region’s flora and
its history. Bill Brumback has answered numerous questions on
ornamentals. P Del Tredici, R. Mitchell, and P Somers are
thanked for sharing their botanical knowledge with me. Curators
of AA, BRU, GH, MASS, NEBC, YU, and the staff of CONN are thanked
for allowing me access to their collections and helping me use
them. Judy Warnement of the Harvard University Herbaria has
been a great help and allowed me access to the libraries she
Girects.
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RHODORA, Vol. 102, No. 911, pp. 299-331, 2000
RATES OF SUCCESS IN THE REINTRODUCTION BY
FOUR METHODS OF SEVERAL PERENNIAL PLANT
SPECIES IN EASTERN MASSACHUSETTS
BRIAN DRAYTON
c/o TERC, Inc., Massachusetts Ave., Cambridge, MA 02140
ail: Brian_Drayton @terc.edu
RICHARD B. PRIMACK
Biology i said Boston pcre Boston, MA 02215
ail: primack @bio.bu.
ABSTRACT. ‘To prevent species from going extinct and to restore locally
extinct species to conservation areas, conservationists have been attempting
to create new populations of rare and endangered species. Such efforts are
still at an early stage, with the basic methodology still being developed and
many efforts resulting in failures or only modest success. The purpose of this
work was to develop some general rules about how to carry out reintroduction
efforts using four methods to create many new populations of eight perennial
species. Our results demonstrate that the chances of success were greater
when ae ‘seedli ng an ult material rather than sowing seeds on the
sites. Using larger adult aaa was more successful than using seedlings.
Adult transplants also flowered and fruited a away, in contrast to plants
derived from seeds, which rarely flowered even after several years. Digging
up the site to expose the soil and reduce omeee ice prior to sowing seed
did not result in a greater establishment of seedlings. At many sites no plants
as
of establishing new plant papul tone To increase the rate of success, at-
tempts should utilize many sites, numerous seeds or plants, and various meth-
ods in order to develop a workable methodology for the species in question.
Because of the difficulties of establishing new populations, conservation of
rare and endangered species should first protect existing populations and only
secondarily rely on reintroductions to ensure species survival.
Key Words: reintroduction methods, conservation, population re-establish-
ment, restoration ecology
It has been estimated by the Center for Plant Conservation that
perhaps 4200 of the 20,000 plant species of North America are
under threat of extinction to some degree (Center for Plant Con-
servation 1993). A recent survey of the New England flora found
576 taxa judged to be “in need of regional conservation” (Brum-
back and Mehrhoff, et al. 1996; Stevens 1998). Worldwide, per-
299
300 Rhodora [Vol. 102
haps 25% of vascular plant species may become extinct in the
coming 50 years (Raven 1987).
A primary cause of species extinctions is direct damage to the
populations, whether by destruction of habitat, over-exploitation,
or from competition from introduced plant or animal species. In
addition to these acute effects, however, there is also a mounting
chronic pressure on many species owing to a combination of hu-
man factors that alter species’ environments in ways that inhibit
or interrupt reproduction, dispersal, and colonization of new sites
and thus the establishment of new populations. Local or regional
anthropogenic effects, such as the production and dispersal of
ground-level ozone or acid precipitation, alter the chemical en-
vironment adversely for some species (witness the effects of acid
rain on Picea rubens in New England, or the contribution of
airborne sulfur compounds to Waldsterben in Germany; Schulze
et al. 1989), killing or weakening individuals, thus rendering them
more susceptible to pathogens, drought, or wind damage. Frag-
mentation of habitat can introduce changes in the biological and
physical characteristics of a location that can accumulate dra-
matically over time (Bierregaard et al. 1992; Brothers and Spin-
garn 1992; Harris and Silva-Lopez 1992; Saunders et al. 1991).
These changes can both cause the death of plants currently oc-
curring there and prevent or largely inhibit the establishment of
new populations, either by the creation of barriers to dispersal,
by the local extinction of dispersers, or by the introduction of
weedy species that compete with previously occurring species.
On a larger scale and over a longer period of time, global
climate change, especially carbon dioxide (CO,) enrichment of
the atmosphere and attendant global warming, is likely to con-
tribute as well to the cascade of plant extinctions, as the temper-
ature and precipitation regimes render areas of the current distri-
bution of many species inhospitable (Bazzaz 1996; Kutner and
Morse 1996; Peters 1992). The rate of anthropogenic climate
change currently projected (Houghton et al. 1996) would require
an adjustment of species ranges at a rate higher than any known
to have occurred during at least the past 10,000 years, and species
often will not be able to migrate naturally across the human-
fragmented landscape.
Rates of extinction of species across all five biological king-
doms are estimated by some to be as high as 0.5% per year
worldwide (Wilson 1992; Woodwell 1990). Studies of local ex-
2000] Drayton and Primack—Reintroduction of Perennials 301
tinctions in areas in which human impacts such as habitat mod-
ification and fragmentation have been sustained over a long pe-
riod are consistent with this estimate (Drayton and Primack 1996;
Newmark 1991; Robinson et al. 1994; Turner et al. 1994). As
much as one third or more of the native species have been elim-
inated from some small and high-impact conservation areas. In
the face of the local and global threats to biological diversity, the
basic conservation response has been site protection: setting aside
habitat that is maintained relatively undisturbed, in order to allow
threatened populations to survive with no further damage (Pri-
mack 1998)
This protection is necessary but probably not sufficient as a
conservation strategy (Buttrick 1992; Falk and Olwell 1992; Pres-
sey 1994). It can prevent further direct disturbance of a site, or
the effects of overexploitation of the site or population. It does
not, however, protect against the more subtle stressing effects of
climate change or pollution. It also does not counteract the long-
term impoverishing effects of habitat fragmentation, which inhibit
or interdict the metapopulation dynamics necessary to the contin-
ued survival of a species at the local and regional scales—spe-
cifically the colonization of fresh suitable sites at a rate sufficient
to offset the natural and human-induced extinction of local pop-
ulations (Grubb 1977; Holsinger 1993; Hughes and Fahey 1988;
Norton 1991; Peterken and Game 1984; White 1996).
Increasingly, in situ management includes the creation of new
populations of taxa or the augmentation of existing populations
(Falk et al. 1996; Primack 1996), despite some concerns about
implications of the practice and the indifferent success of many
programs. The restoration ecology and conservation biology lit-
erature now reports many projects in which plants are reintro-
duced to an area where they once occurred, or new populations
are initiated near existing stands, or species are introduced at
apparently suitable sites. This flush of reintroduction activity has
opened up many areas of research both on the basic biology of
the species under consideration (Drayton 1999; Primack 1996;
Schemske et al. , and on many aspects of technique that
must be considered in relation to the biology: whether to under-
take a reintroduction or augmentation plan (Gordon 1994), how
to define success for a reintroduction (Pavlik 1996; Sutter 1996),
how to select suitable sites (Fiedler and Laven 1996), and how
to design the actual introduced “‘population”’ (Guerrant 1996; Ha-
302 Rhodora [Vol. 102
vens 1998; Husband and Barrett 1996; Primack 1996). In addi-
tion, there is still much to be learned about which techniques are
most effective in restoration and reintroduction, including the rel-
ative value of seeds versus propagated material for introduction,
and the extent and nature of appropriate site preparation and after-
care.
Choosing material for reintroductions: Seeds or
plants? Because the germination and seedling stages of growth
are periods of high vulnerability and high mortality, and because
rare plant material must often be used with great care and econ-
omy, the majority of reintroductions of perennials have proceeded
by the propagation of plants ex situ, and then transplanting into
the target site (Guerrant 1996). Transplants of material in forms
such as seedlings, cuttings, or bulbs arrive at the target site al-
ready past the most vulnerable stage of life. Individuals translo-
cated in these forms tend to survive at a higher rate than seedlings
germinating in situ (Barkham 1992; De Mauro 1994; McEachern
et al. 1994; Ray and Brown 1995; Rochefort and Gibbons 1992;
Vora 1992) and initiate flowering or asexual reproduction faster
than individuals propagated from seed (Seliskar 1995; Vasseur
and Gagnon 1994). In cases where the site cannot be character-
ized quantitatively, transplants that survive provide evidence that
the site is suitable for the species and that its absence there may
be due to lack of dispersal (Barkham 1992; Lee 1993; Primack
and Miao 1992).
Yet even when it seems feasible from a logistical point of view,
transplanting does have inherent risks, since there can be signif-
icant trauma during the transplant. Plants grown ex situ by defi-
nition have not grown in situ, so that the change in environment
may subject the transplants to stress that affects their viability or
results in high levels of herbivory (Cavers and Harper 1967).
Poor horticulture or adverse conditions such as unanticipated
drought can result in high mortality in the field (Fahselt 1988).
Further, introduction of plant materials may inadvertantly intro-
duce pathogens as well (Given 1994).
Beyond the biological considerations, however, is the factor of
the cost of such an approach, which must be weighed against
potential higher rates of success as compared with the use of
seeds to initiate the new populations (Danielson 1996; Given
1994), For example, the cost of establishment of a single indi-
2000] Drayton and Primack—Reintroduction of Perennials 303
vidual of Texas Ebony by tranplanted seedling (raised ex situ)
was about $1.25, while the cost of establishment by seed was
around $0.39 per individual (Vora 1992).
Reintroductions by seed offer some important advantages over
transplants. In the first place, seeds can often be collected in large
numbers. Collection of seed can usually be accomplished without
damage to the individuals in existing populations, and this is es-
pecially important when there are only a few individuals of a
taxon remaining. For example, in the case of the threatened Prai-
rie Fringed Orchid (Platanthera leucophaea), populations are
scattered and declining to the point that pollination is inhibited
in some parts of its range. Little is known about the cultivation
requirements of this species, so transplanting of existing individ-
uals entails an unacceptable risk of mortality. The use of seeds
for the creation of new populations of this species is the most
useful short-term strategy for increasing the number of popula-
tions or for augmenting existing populations (Packard 1991).
It is possible that in a suitable site the individuals that germi-
nate and grow in situ have a better long-term chance of success
on that site than plants not “‘selected’”’ by the microenvironment
of the site. In some cases, seedlings from seeds sown in situ may
have a more rapid growth rate than seedlings transplanted from
elsewhere (Vora 1992), and rapid growth rate can be important
if light is the limiting medium so that the production of photo-
synthetic tissue is decisive for survival in the face of above-
ground competition or litter-fall.
Seeds can be dispersed soon after collection, thus ensuring that
the propagules used for reintroduction are arriving at the target
site in synchrony with the natural dispersal process. Seeds are
also amenable to several kinds of experimental plantings which
may provide important information about the biology of the spe-
cies under study. This may improve the effectiveness of recovery
or mitigation plans. For example, it may be important to design
an introduced population to have maximal genetic diversity (Dole
and Sun 1992; Fenster and Dudash 1994; Jacobson et al. 1994).
It is easier to introduce multiple populations and multiple geno-
types by means of seed than by means of transplanted material.
Another important concern is the density of the population, but
the optimal density and spatial arrangement of individuals in a
population is known for rather few species. Reintroduction by
seed allows for a variety of planting arrangements and densities.
304 Rhodora [Vol. 102
In the case of species for which abundant seed is available, one
can even design restoration or reintroduction plans at a landscape
level using mixtures of seeds and seeding techniques (e.g., Ja-
cobson et al. 1994), though this is perhaps most likely for grass-
land habitats.
Site preparation and post-translocation care. The concept
of “‘safe sites” for establishment (Harper 1977), or the “‘regen-
eration niche” (Grubb 1977), provides an important rationale for
careful site selection for the reintroduction of a species. The ra-
tionale includes a range of criteria, including biological criteria
(e.g., specific nutrient or water requirements), logistical criteria
(e.g., 18 the site accessible enough to the researcher to enable the
operation to proceed and to enable appropriate monitoring, with
‘after care’? or maintenance activities?), and ‘“‘defensive”’ criteria
(e.g., 18 the area vulnerable to human disturbance? Have man-
agement policies resulted in a high density of deer that might eat
the plants?; Fiedler and Laven 1996). In addition, there may be
other evidence to consider, such as the historical presence of the
species. The autecology of many species is not well understood.
If time and resources permit, one can conduct the studies needed
to ascertain the answers to critical questions. As this 1s not always
possible, some surrogate measures of site suitability may be re-
quired. A common example is the use of indicator species, species
whose occurrence is highly correlated with the occurrence of the
target species.
Initial experiments on which this study is based used little in
the way of site preparation (for a summary, see Primack 1996).
There is a strong a priori rationale for this, since most plants
disperse the bulk of their seeds onto unprepared sites. Further, for
many species it is not known what kinds of “‘preparation”’ might
favor establishment by seed or the survival of seeds once ger-
minated. Studies of germination requirements are not reliable
guides to the requirements for establishment, as the ideal condi-
tions for germination may not be ideal for the new seedling
(Grubb 1977). This is likely to be the reason that studies show
high laboratory germination rates but very low seedling survi-
vorship in the field (Vora 1992), or high seedling emergence and
also high seedling mortality (Barkham 1992; Bazzaz 1996).
For species whose establishment biology is not well under-
stood, some approximation can be attempted based on dispersal
2000] Drayton and Primack—Reintroduction of Perennials 305
mechanisms (Robinson and Handel 1993), germination require-
ments known or conjectured (Baskin and Baskin 1998), and on
what is known of the disturbance regime of the species’ habitat.
For example, desiccation is an important cause of mortality in
emergent seedlings (Larcher 1995). Sites can be prepared with
mulches (Jackson et al. 1990; Rochefort et al. 1992) or shaded
with branches, litter, or screens (McChesney et al. 1995) to min-
imize drying of the top layer of soil. Bringing seeds’ emergent
radicles close to mineral soil may require the removal of litter or
the mowing or removal of vegetation (Gordon 1996; Rochefort
and Gibbons 1992; Vasseur and Gagnon 1994; Vora 1992; Wat-
son et al. 1994). Removal of over-shadowing vegetation can im-
prove the light supply for early rapid growth of seedlings and
can impair root competition, significantly improving seedling sur-
vival (Danielson 1995; Pavlik et al. 1993). Cultivation of the soil
can also reduce below-ground competition (a decisive factor in
the mortality of seedlings in many systems; Bazzaz 1996), aerate
the soil, and facilitate root growth (Bainbridge and Virginia
1990). The site may be irrigated or enriched by fertilizers to fa-
cilitate rapid growth (Doerr and Redente 1983). A fire regime
may be instituted, which can remove above-ground competition,
remove thatch or litter that may prevent seeds’ reaching the soil,
and provide a nutrient pulse (Gordon 1996; Pavlik et al 1993).
Finally, some species may require protection against seed pred-
ators or herbivory on the emergent seedlings (Bainbridge et al.
1995; Barkham 1992; Chambers and MacMahon 1994; Primack
and Drayton 1997).
Post-reintroduction care (“‘soft release’’) may also be part of
the reintroduction plan. Techniques reported from the literature
include protection against seedling dessication with mulching,
screening, or irrigation (Bainbridge and Virginia 1990; Doerr and
Redente 1983; Jackson et al. 1990). Sites can be weeded (Jackson
et al. 1990) or clipped (Danielson 1995; Gordon 1996) to contin-
ue to prevent competition during early growth.
Criteria for success of a reintroduction. Increasingly it has
been recognized that a reintroduction effort must be evaluated
with reference to its original goals, and that these will vary con-
siderably from case to case (Pavlik 1996). These goals may spec-
ify an extension of a species’ range by the creation of new pop-
ulations or by increasing the size of existing populations in order,
306 Rhodora [Vol. 102
for example, to reach a threshold of attractiveness to pollinators.
In most cases, success will be achieved stage-wise, first by the
presence of individuals on the target site, then by their reaching
reproductive stage, then by their dispersing viable seed, and per-
haps finally by their establishing secondary populations. A lon-
ger-term goal may be a minimum viable population size, a target
developed on the basis of demographic modelling.
Long-term monitoring of new populations or reintroductions
can serve several critical purposes, yet systematic monitoring past
the initial stages of establishment is a surprisingly rare feature of
published reports on reintroductions. Measures of success are of-
ten expressed in terms of biomass (Doerr and Redente 1983;
Shaw 1996), per cent cover (Jackson et al. 1990), or presence-
absence (Packard 1991; Revel 1993).
Despite the large amount of attention that plant reintroduction
has received in recent years, it is still possible for a leading re-
searcher to state that there is no example of a taxon’s having been
conserved or brought to nonendangered status as a result of a
restoration plan (Pavlik 1996). In part this statement can be ex-
plained by the length of time often needed to assess the outcome
of a reintroduction, especially when working with perennials. In
part the statement also reflects the state of our understanding of
many aspects of the reintroduction process. In each section above,
one sees open questions that require further research. The recent
history of reintroduction work shows a swift development of un-
derstanding of the challenges facing such conservation work as
researchers have attempted various approaches, developed criteria
for assessing results, and collected results from a range of dif-
ferent studies and species.
The literature and examples of restoring populations of rare
and endangered species have grown considerably over the last 10
years, but the development of general approaches has been in-
hibited by a variety of factors. First, most attempts to restore
species are done with a single species, so it is unclear if the result
would be applicable to another species of different growth form,
family, or basic biology. Second, most attempts involve a single
approach rather than conducting experiments in which several
approaches are contrasted. Third, most attempts do not replicate
the approach, so it is unknown how consistent the reported results
are. Fourth, the results of many, 1f not most, such projects are
never published, and in particular it is quite likely that most un-
2000] Drayton and Primack—Reintroduction of Perennials 307
successful attempts to create new populations are never published
at all. This may lead to literature biased in favor of successful
and optimistic results. The purpose of the work presented here is
to develop generalizations on the most effective way to establish
new populations of rare, declining, and endangered species. We
used many species, several techniques, and many replicates to
develop generalizations that could be widely applicable. In this
research we focused on perennial wildflower species, as many
New England plant species are in this category, and our earlier
research investigated annual species (Primack 1996; Primack and
Miao 1992).
The present experiment was intended to answer the following
questions with regard to eight native perennial species:
1. How frequent is the establishment of new populations of
perennial species in relation to the number of propagules
arriving on a site?
Is transplantation of seedlings and adults more or less ef-
fective than reintroduction by seed?
. Does site preparation increase the success of reintroduction
by seed?
. Finally, is the establishment of new plant populations in the
wild a realistic goal for perennial wildflower species’
Sa
ies)
&
MATERIALS AND METHODS
Starting in 1993, we identified eight perennial species that were
not present but formerly attested, or whose distributions were
highly restricted, in two conservation areas in the Boston area.
None of these species was endangered or threatened in Massa-
chusetts, but the number and population size of most of them
appeared to have declined substantially over the last century.
Such species may be of conservation interest in themselves—and
thus the subject of reintroduction efforts—if the populations’ dis-
tributions were shrinking so that (presumed) genetic diversity was
diminishing, or if there were other biological, cultural, or aes-
thetic values to the species’ continued presence in a particular
locale (Hunter and Hutchinson 1994). In addition, such species
can serve as model systems for the purpose of exploring the val-
ues and limits of conservation techniques before attempts are
made to apply such techniques to endangered species.
308 Rhodora [Vol. 102
The species used for this study were as follows (nomenclature
follows Gleason and Cronquist 1991; geographic information
from Seymour 1993): Marsh Marigold (Caltha palustris); Col-
umbine (Aquilegia canadensis); Bloodroot (Sanguinaria cana-
densis); Early Saxifrage (Saxifraga virginiensis),; Spikenard (Ara-
lia racemosa), Cardinal Flower (Lobelia cardinalis), Sweet Cic-
ely (Osmorhiza claytonti); Bluets (Hedyotis caerulea).
These species are well-known, even “characteristic,” elements
in the New England flora. All species were present in the Mid-
dlesex Fells, and all were uncommon except Bloodroot, Bluets,
and Sweet Cicely. Only Marsh Marigold was present in the Ham-
mond Woods, where it existed as a single large population. While
each species has its distinct requirements, there are a few features
that should be noted. Columbine and Cardinal Flower are hum-
mingbird-pollinated, whereas the other species are insect-polli-
nated. Marsh Marigold and Cardinal Flower are wetland species,
while the others grow in forests, fields, and disturbed areas.
Sources of plant material. In the summer and fall of 1994
seeds of all species were collected from populations in eastern
Massachusetts, in most cases within 2 km of the experimental
sites. Seeds to be sown on quadrats were collected at the time of
natural dispersal, cleaned, counted, and placed on quadrats within
a week of collecting; they were stored to ensure viability in the
meantime (Baskin and Baskin 1998). In the winter of 1994, sam-
ples of the seeds of all species were sown in flats, cold-stratified
at 4°C for 10 weeks, and germinated in growth chambers to test
for viability and if necessary to provide material for transplan-
tation. All species showed germination rates in the laboratory >
50%, except for Saxifraga, for which seeds germinated at a rate
of approximately 10%.
Seedlings and adults for transplantation (see below) were ob-
tained in the spring of 1995, when possible from wild populations
in the area that were of sufficient size to allow removal of plants
for transplanting (Sanguinaria, Osmorhiza, Caltha, Saxifraga,
Hedyotis seedlings). In cases where this was not possible (Lo-
belia, Hedyotis adults, Aquilegia, Aralia), seeds were collected
from naturally occurring sites in eastern Massachusetts and prop-
agated first in the laboratory, then in suitable sheltered areas out-
side for hardening until transplantation.
2000] Drayton and Primack—Reintroduction of Perennials 309
Study sites. Experimental sites were established in the Ham-
mond Woods (Newton, MA) and the Middlesex Fells (Medford,
MA). The Hammond Woods is a conservation area approximately
80 ha in area. It comprises a mixture of deciduous woods,
swamps, parking areas, meadows, ledges, and roads. The Mid-
dlesex Fells is approximately 800 ha in area, in two roughly equal
sections isolated from each other by major highways; the reserve
overlaps five municipalities. The park is dominated by mixed
deciduous woods, but includes large and small bodies of water,
stream courses, maintained and abandoned fields, gravel carriage
roads, and hiking trails. It is used heavily for hiking, mountain
biking, picnicking, and similar recreational purposes.
Sites within each area were selected on the basis of general
topographical aspect by comparison with sites in which the spe-
cies occurred naturally in their nearest populations. Criteria in-
cluded degree of canopy closure, soil moisture, and co-occurring
indicator species. For each species, apparently suitable habitat
existed in these conservation areas, so that reasons for the absence
or decline of populations are not known. A first hypothesis is that
dispersal has limited the extent of occurrence. Further, human use
of the areas may well have contributed to reduced dispersal
(Drayton and Primack 1996 and references therein). Therefore,
the design provided several useful kinds of information about the
sites being explored: transplants that survived and seemed to es-
tablish well provided evidence that the site was suitable for the
species, at least within the time frame of the study to date. Es-
tablishment of seedlings from seed provided evidence that dis-
persal may have been limiting. Relative success of individuals of
different ages may also provide evidence about life-stages that
are particularly vulnerable in these species, information that
should be taken into account in designing a reintroduction plan
(Schemske et al. 1994).
Experimental design. At each site, four quadrats were
mapped and each marked with a numbered wooden stake in the
summer and fall of 1994. Four treatments were used; one quadrat
at each site was assigned randomly to each treatment; the number
of quadrats (replicates) for each treatment for each species is
shown in Table 1. The treatments were as follows:
Treatment 1: Seeds. A known number of seeds was sown di-
OES
Table 1. Number of replicates (quadrats) of experimental design, number of seeds sown for treatments | and 2, and number of
individuals transplanted for treatments 3 and 4. Treatments are described in Materials and Methods.
# Seedings
# Seeds Sown and Older
per Quadrat Plants per
# Replicates for Total # Seeds Quadrat for
per Treatments | Sown per Treatments 3 Total # of
Treatment Species and 4 Transplants =
Species S
Aquilegia 24 100 4800 4 192 .
Sanguinaria 12 50 1200 4 96 a
Hedyotis 16 100 3200 5 160
24 100 4800 6 288
Caltha 24 100 4800 4 192
Saxifraga 6 50 600 4 48
Lobelia 19 100 3600 4 144
Osmorhiza 24 100 4800 4 192
Total for all 149 27,800 1312
species
<
COI
2000] Drayton and Primack—Reintroduction of Perennials 311
rectly on the quadrat in the summer and fall of 1994 within
a 25 cm radius of the marker. Nothing was done to disturb
the site other than to introduce the marker.
Treatment 2: Dig and Seed. The quadrat was dug up within a
5 cm radius of the marker and to a depth of approximately
12 cm, removing possible competing herbaceous cover and
superficial roots and exposing bare soil; then the same num-
ber of seeds as in treatment | was sown in 1994.
Treatment 3: Seedlings. Seedlings were transplanted onto the
assigned quadrat in the spring of 1995, within a radius of
0.5 m of the marking stake, in holes prepared by trowel.
The sites were not altered in any other way (e.g., by re-
moval of overhanging vegetation). In the case of Hedyotis,
seedlings were watered once soon after transplanting be-
cause of unusually dry conditions.
Treatment 4: Adults. Adult plants were transplanted into the
assigned quadrat in the spring of 1995, within a radius of
0.5 m of the marking stake, in holes prepared by trowel.
The sites were not altered in any other way. In the case of
Hedyotis, adults were watered once soon after transplanting
because of unusually dry conditions. For treatments 3 and
4, the same number of individuals (seedlings and adults)
was used.
The number of replicates was determined by the number of
seeds or potential transplants that were available. The number of
seeds sown (for treatments 1 and 2) and of transplanted seedlings
and adults is shown for each species in Table 1
All sites were visited repeatedly during the growing seasons,
and data were taken annually on:
number of seedlings from seeds sown by researchers or dis-
persed by introduced individuals,
number of survivors from transplants,
number of plants flowering or setting seed in the summers
of 1996 and 1997,
number of fruits.
Although the seasons of 1996 and 1997 were quite dry in eastern
Massachusetts, no transplants were watered, nor was there any
other post-transplant care except as noted for the transplants of
Hedyotis upon first planting in 1995,
Siz Rhodora [Vol. 102
Statistical analyses were performed using the Statsoft Statistica™
(Release 4.1) program and Microsoft Excel™ versions 4 and 5.
RESULTS
The success of a reintroduction can be assessed with reference
to several questions. For perennials, these can be answered at
least provisionally in chronological order. First, are individuals of
the subject species present on any of the experimental sites? Sec-
ond, what percentage of the original propagules have resulted in
individuals surviving at the time of census? Third, are there any
individuals reaching reproductive condition, and if so, are they
setting seed? Fourth, is there evidence of a second generation at
any site?
In overall terms, the results of this experiment emphasize the
difficulty of successful reintroduction, the caution needed in gen-
eralization about methods, and the need for long-term monitoring.
Transplanting material was by far the most reliable way to estab-
lish new populations when comparing the results for all species,
but there was considerable variation among species in the rates
of success as measured both by occupancy versus treatment and
survivorship versus treatment.
Number of quadrats occupied. There was a total of 596
quadrats of all species, 149 per treatment (Table 1). Of these, by
the end of the period here studied, there were 105 occupied by
the subject species (Table 2), thus an overall rate of 19%. Of
these, 87 (78%) were reintroductions by transplant, and 15 (22%)
were by seed. The success rate of transplants was significantly
greater than establishment by seeds (x°, P < 0.001; Table 3).
Although the values varied among the species in the study, for
most species, transplants were clearly more successful than seeds
in terms of survivorship. In three species, Lobelia, Saxifraga, and
Aquilegia, no individuals from seed survived to 1997. By con-
trast, both Sanguinaria and Osmorhiza showed relatively large
numbers of quadrats occupied by seedlings from introduced
seeds: for Sanguinaria, 8 quadrats planted with seeds were oc-
cupied in 1997 (4 each for the two seed treatments); for Osmor-
hiza, 6 quadrats planted by seed were occupied in 1997. For Hed-
yotis, five quadrats planted by seed were occupied in 1997, which
contrasts with the 8 quadrats occupied by transplants.
2000] Drayton and Primack—Reintroduction of Perennials 313
Table 2. Number of quadrats occupied in 1997, by species and treatment.
Treatments are described in Materials and Methods
Species Treatment | Treatment 2. Treatment 3. Treatment 4
Aquilegia 0) 0) 5 10
Sanguinaria 4 4 6 7
edvyotis 2 3 7 1
Aralia l 1 7 11
utha l 2 18
Saxifraga 0 0 0 4
Lobelia 0) 0 0 0)
Osmorhiza 6 0 I 3
Total quadrats 14 9 28 54
Except for Osmorhiza, there seemed to be no significant dif-
ference in the success of seeds on prepared versus unprepared
quadrats. This result in 1997 was surprising, because in the pre-
vious two years of the study for several species (Sanguinaria,
Hedyotis, Aquilegia) the prepared quadrats showed higher num-
bers of individuals present. For example, in 1995 Osmorhiza
showed seedlings at 63% of the prepared quadrats, versus 13%
of unprepared quadrats. Although this was the largest disparity,
emergence of seedlings from the first seed input on prepared
quadrats was generally higher than on unprepared quadrats. Yet
by 1997, this difference had diminished in all species (Figures 1
and 2). For Osmorhiza, in 1997 no prepared quadrats (treatment
2) were occupied, while six of the unprepared quadrats (treatment
1) had individuals on them. In 1996, three of the Saxifraga pre-
pared quadrats (treatment 2) showed seedlings, as opposed to
none of the unprepared quadrats, but in 1997 no quadrats sown
with seeds showed any individuals present. For Sanguinaria,
there were four occupied quadrats for each of the two ‘“‘seed”’
treatments by 1997. The unprepared quadrats showed a signifi-
cantly higher number of seedlings present in 1997; this reversed
the situation of previous years. For Hedyotis, the prepared quad-
rats showed a significantly higher rate of occupancy in all years.
Only prepared Aralia quadrats showed any individuals from seed
present in any year. In general, the site preparation seemed to
facilitate germination and initial establishment but not to affect
longer-term persistence at a site.
With respect to the relative success of the two transplant meth-
ods, with mature versus younger plants, for most species more
314 Rhodora [Vol. 102
Osmorhiza: # Quadrat pied, by | i a signa sone da by |
year and treatment and treat
30.00 ; 16
a 14 =
4 25.00 L 42 ee
20.00 See) «|MTrt 1 10 Mcil [Succ eeeL trt 1
15.00 | jit 2 ae oo ee wtrt 2
; OTrt 3 6 ks ve
oe Ort 4 4 a oe saat au ‘
: r
¢ 5.00 S 2 enlist?
0.00 Sa Ee 0 - T
1995 1996 1997 1995 1996 1997
Year Year
— se _ ; _ =
ase hs #Quadrats oo by Caltha: #Quadrats Occupied, by year
and treatm and treatment
25
x 7
a B20 [| =
re GtTrt 1
BTrt 2
F S104 oT 3
f ‘ ot 4
C € 5 7 7
1996 1997
Year Year
Figure 1. Number of quadrats occupied per year, by treatment, for Os-
morhiza, Hedyotis, SaenOny, and Caltha. Treatments are described in Ma-
terials and Methods.
quadrats planted with mature plants were still occupied by 1997
than quadrats planted with seedlings (Figures | and 2). The ad-
vantage was most marked for Caltha, Aquilegia, and Aralia, with
these differences statistically significant. For Caltha, 18 quadrats
were occupied by mature transplants, while only 2 were occupied
by seedlings. For Aquilegia, 10 quadrats were occupied by adults,
5 by seedling transplants. For Aralia, 11 quadrats were occupied
by adults, 7 by seedlings. In one case, with Hedyotis, there was
the opposite result with seedlings occupying more quadrats than
mature plants in all years. For Sanguinaria, almost equal numbers
of quadrats were occupied by plants: 6 seedling quadrats and 7
adult quadrats. For Saxifraga and Lobelia, only mature plants
survived, and in the drought year of 1997, no Lobelia plants were
found.
Rates of success per propagule. Overall, 27,800 seeds and
2000] Drayton and Primack—Reintroduction of Perennials 315
Aquilegia: #Quadrats ee iad by Lobelia: ie cane sig by year
year and treatm and treatm
18 12
o 16 ho}
o © 10
a 14 a
3 12- g 8 |
9 104 f<}
“ %
= 8 s
g 6 Oo 44
3 4 3
ie] Go 2
%* 9 te
0 + + 4 0 T T a
1995 1996 1997 1995 1996 1997
Year Year
Aralia: #Quadrats occupied, by year a Saxifraga: panini ae by
and treatment Yea d Treatm
25 4.5
7 4-
; 20 1 3.5 -
3 4
15 eae aga 2.5 -
+ + 2 4
; 10 7-— 3154
ieee ees 1 Be
5 see] eee €
g L #05 7
0 A r 0 + LLL
1995 ie 1997 1995 1996 1997
Yea Yea
Figure 2. Number of quadrats occupied per year, by treatment, for Aq-
uilegia, Lobelia, Aralia, and Saxifraga. Treatments are described in Materials
and Methods.
1312 transplanted individuals (including both young and mature
plants) were introduced on the experimental quadrats—half on
prepared quadrats, half on unprepared. The rates of success per
propagule introduced varied widely (Table 3) but in general they
mirrored the results for rates of quadrat establishment. Thus the
transplanting of material had a very much larger rate of success—
that is, percentage of transplanted individuals surviving to 1997—
than did introduction by seed. For all species, introduction by
seeds (including both treatments) resulted in 131 individuals pre-
sent for a success rate of 0.47%. Transplanted individuals fared
better, with 23% of the 1312 transplants (including both seedlings
and plants) surviving to 1997. Species differed in the relative
rates of success, with Sanguinaria showing the most spread be-
tween seed treatments (about 4.5% for the two seed treatments)
and transplants (about 44%); most species showed rates of estab-
Number
values dene significantly by x test. Treatments are described in Materials and Methods.
of 1997 survivors per treatment, and rates of survival per propagule in each category. Superscripts indicate
Treatment 1
Treatment 2
Treatment 3
Treatment 4
# Present # Present # Present # Present
Species 1997 % of Input 1997 % of Input 1997 % of Input 1997 % of Input
Aquilegia 0 0 0 0 9 9.4 8 18.8
Sanguinaria 35 5.8 20 3.3 19 39.6 23 47.9
Hedyotis 7 0.75 47 Bal 128 120) 1 | PP ae)
Arali 1 0.042 | 0.042 20 138 20 13.8
Caltha 4 0.16 1 0.04 5 a2 49 51.4
Saxifraga 0) 0 0 0 4 8.3 4 8.33
see 0) 0 0 0 0 ee) 0 9.2
Osm 13 1.0 0 0 3 3.0 4 4.0
Total hi all spe- 60° 0.43 69" 0.5 188° 28.0 12” 16.6
cies
OLS
elopoyy
TOT TOA]
2000] Drayton and Primack—Reintroduction of Perennials 317
lishment from seed at less than 1%, significantly less than rates
by transplant. Aquilegia showed no individuals from seed present
in 1997 but had a survival rate of 10% for seedling transplants
and 19% for adults. Aralia showed survival rates of 0.04% for
the seed treatments, and 14% for the transplant treatments. Caltha
had a low survival rate from seed (0.16% and 0.04% for treat-
ments | and 2, respectively), but 5% survival for seedlings and
51% for adult transplants. In the case of Lobelia and Caltha, the
sites necessarily were near moving water, so it seems possible
that many seeds were washed away from the experimental quad-
rats before germination. No seedlings of these species were noted
downstream from the experimental sites, however.
Reproduction at experimental sites. The survival of intro-
duced material is only the first level of success for a reintroduc-
tion effort, and the reintroduction can only be considered suc-
cessful if some of the introduced individuals survive to reproduce
and become a source of reproducing offspring in the target area.
In the case of the present experiment, it is too early to assess this
level of success with respect to individuals introduced by seed.
In all cases except Hedyotis, which often flowers and sets seed
during its first year, individuals of the perennial species in this
study must reach a certain size, usually over several growing
seasons, before they will reproduce. As these sizes are not defined
in the literature so far as we can determine, this fact of life-history
means that monitoring introduced populations must be a long-
term effort.
In the case of introduced material, however, initial results can
be reported. We recorded all instances of reproduction in 1996—
97 (Table 4), and flowering individuals in 1996 (Table 5) and
1997 (Table 6). All but one species, Aralia, showed some repro-
ducing individuals during the experiment to date. It appears that
in the very dry conditions of 1996 and 1997 Osmorhiza was
prevented from reproducing, even in the few sites where there
were flowering transplants in 1995. However, in a few cases the
seeds produced by those transplants did yield seedlings in 1996.
Lobelia flowered in 1996 and two individuals set fruit (a total of
20 capsules between them), but no flowering individuals appeared
in 1997. For Caltha, only the adult transplants flowered, but a
high percentage did so (72% in 1996, with a total of 32 fruits on
47 flowering individuals; 70% in 1997, with a total of 42 fruits
318 Rhodora [Vol. 102
Table 4. Number of quadrats with reproducing individuals, total number
of fruits produced 1996—7, and presence/absence of second generation, 1.c.,
seedlings from seeds dispersed by introduced material.
Second
Species # Quadrats # Fruits Generation?
Aquilegia LO 54 no
Sanguinaria LO 31 yes
Hedyotis 14 800 yes
] O no
Caltha 19 263 no
Saxifraga 4 126 no
L ia | 14 no
Osmorhiza 10 310 yes
on 33 flowering individuals). Saxifraga showed a high percentage
of adult transplants flowering (89% in 1996, 100% of 2 individ-
uals in 1997), and essentially all flowers matured fruit though no
seedlings have appeared at these sites. Sanguinaria seedlings and
adult transplants showed similar proportions of flowering indi-
viduals in both years (about 16% in 1996, around 50% in 1997),
with a total of 31 fruits over those two years. Aquilegia showed
increasing proportions of flowering individuals (12% of seedling
transplants in 1996, 78% in 1997), but negligible fruit production
until 1997 (22 fruits noted).
Hedyotis showed the most vigorous reproduction in both years
although adult transplants showed only one flowering individual,
in 1997. The individuals appearing from seeds sown on the pre-
pared plots flowered starting in 1996 (83%) and continued in
1997 at a lower rate (21%). Seedling transplants flowered vig-
Table 5. Percentage of individuals per treatment flowering in 1996. Treat-
ments are described in Materials and Methods. ! Based on one individual.
Species Treatment | Treatment 2. Treatment 3. Treatment 4
Aquilegia 0 0 [25 37.2
Sanguinaria 0 O 15.8 16.1
Hedyotis 100! 83 93.8 0)
ralic 0) 0) 0 0)
Caltha 0 0 0 AAS
Saxifraga 0) 0) 33.3 88.9
Lobelia 0) 0 100 42.9
Osmorhiza 0) 0 0
2000] Drayton and Primack—Reintroduction of Perennials 319
Table 6. Percentage of individuals per treatment flowering in 1997. Treat-
ments are described in Materials and Methods. ! Based on one individual;
? Based on two individuals.
Species Treatment | Treatment 2. Treatment 3 Treatment 4
Aquilegia 0) 6) 77.8 33.3
Sanguinaria 0 0) 47.8 57.9
He j 0 21.3 13.3 100!
Ara 0 0 0 0
Caltha 0 0) 0 69.8
Saxifraga 0 0 0) 100
Lobelia 0) 0 0) 0
Osmorhiza O 0) 0) 6)
orously in 1996 (94%), but less so in 1997 (13%). However, this
lower proportion of flowering reflects the fact that there were
more individuals present on these sites (58 in 1997 versus 16 in
1996). The increase apparently was largely due to the establish-
ment of new seedlings from seeds dispersed the previous year.
These seedlings were all very small and did not flower, but per-
sisted through the growing season.
Table 4 summarizes the number of quadrats with reproducing
individuals per species for 1996-97, the estimated number of
fruits for those two years, and the presence or absence of seed-
lings from dispersed seeds (a ‘‘second generation’’). As of the
1997 growing season, only Sanguinaria and Hedyotis showed
quadrats with both mature flowering individuals and new seed-
lings present. The few Osmorhiza seedlings derived from 1995
flowering transplants did not appear to be of flowering size yet.
DISCUSSION
Plant reintroductions are considered an important too! in the
work of plant conservation, but there remain many unanswered
questions about techniques for reintroduction and the biology that
underlies them (Allen 1994).
The present experiment, still in progress, reinforces previous
work in which reintroduction by seed has shown very low rates
of success in establishment of new populations at even the most
basic definition of “‘success,”’ that is, presence of individuals of
the species. The rates reported here, ranging from 0% to about
6%, are similar to rates reported in a series of experiments by
320 Rhodora [Vol. 102
Richard Primack for many species in eastern Massachusetts (Pri-
mack 1996; Primack and Miao 1992). In one set of experiments
with annuals and perennials, out of 221 quadrats, a single pop-
ulation of an annual species and two populations of a perennial
species survived to reproduce and disperse seeds. Those experi-
ments showed short-lived appearances of seedlings, as reported
here, but the passage of time saw these “populations” extin-
guished.
Similar experiments in quite different habitats have shown
comparable results. For example, recruitment from seeds of 8
different species sown in the field in the semi-arid Rio Grande
Valley ranged from 11% to less than 1%, except for a single
species (Vora 1992), despite several steps taken to improve the
chances for success both by site preparation and after-care. Vas-
seur and Gagnon (1994) reported emergence rates in their exper-
iment with Allium tricoccum to vary widely from about 3% to
90%, but they did not provide data on the survival of recruits
from seeds after germination. Barkham (1992) reported seedling
survivorship of Narcissus sown in the field as “‘rapidly declining
to zero.”’ In the New England area, repeated attempts have been
made to establish new individuals and new populations of the
endangered perennial Potentilla robbinsiana in the White Moun-
tains of New Hampshire (unpubl. report). Some success has been
achieved using transplants of adults, but sowing seed in a variety
of locations has had no success whatever.
There can be many reasons for this kind of result. Many plants
need some kind of disturbance to establish successfully. Thus the
‘safe site’’ at which the propagule must arrive is not only a par-
ticular locale, but a place in time as well. Site suitability is not
only a function of characteristics such as soil composition and
the presence of competitors and predators, but also the interaction
of these with temperature and precipitation conditions.
The work of David Foster and others (e.g., Foster and Boose
1992; Whitney and Foster 1988) has shown how, on an ecological
time scale—from a few decades to a few centuries—an ecosystem
is likely to experience recurrent though unpredictable major dis-
turbances that may have important consequences for successional
processes, including the establishment or extermination of pop-
ulations of plant species. In New England, a prime example of
such a disturbance is hurricanes, whose effects on northern hard-
wood forest systems have been studied now for some years. In
2000] Drayton and Primack—Reintroduction of Perennials 321
light of this work, Primack (1996) extended his experiments to
an area artificially disturbed to recreate some of the features of a
hurricane disturbance. The radically altered light and temperature
regimes of such a disturbance can enhance or trigger seed ger-
mination, and the removal of competing vegetation and the ex-
posure of mineral soil might be expected to foster a flush of
germinations. In the event, no such response was seen for 15
perennial species sown on the experimental site, suggesting that
other factors besides, or in addition to, disturbance affect estab-
lishment.
The present experiment follows on from these, with a change
in the site preparation, and the addition of a comparison with
transplants of two different sizes. Seeds were sown in some quad-
rats with no preparation, this being the most common fate for the
seeds of these species. This unprepared sowing was compared,
however, with small-scale site preparation, which imitated in its
effects a very common type of disturbance, the uprooting of a
tree or sapling (Runkle 1985). A disturbance on this scale will
not materially alter the radiation regime of a microsite, but will
expose mineral soil and provide a site largely free from root com-
petition in the upper soil layers, and from shading by plants near-
This level of site preparation may have some positive effect
on the rate of emergence of seedlings, but in these experiments
it had no discernible effect on longer-term presence on a site.
Similar results are reported from a series of experiments with a
different set of species in sandhill conifer forests of South Car-
olina (Primack and Walker, unpubl.), in which in addition to dis-
turbance, site preparation included a nutrient pulse. From the
Cape Cod area as well, attempts to create new populations of the
endangered Sandplains Gerardia, Gerardia acuta, in grassland
sandplains, are enhanced by a carefully timed program of mowing
and burning (P. Somers, unpubl. data). Preliminary results suggest
that in this very different biological system as well, local distur-
bance does enhance the emergence of seedlings, while fertilizer
does not. The long-term consequences for survivorship remain to
be seen.
In fact, the point made by Grubb (1977) that the “‘regeneration
niche” is more than a good site for germination is quite apposite
here. Germination is the first and essential condition for a new
colonization event by seed, but the conditions must also be con-
322 Rhodora [Vol. 102
ducive to the survival of new seedlings, so that some reach the
next period of dormancy in good enough condition to survive the
winter. For a species that takes some years to reach reproductive
maturity, this second stage of recruitment lasts through several
growing seasons, with their attendant risks of adverse climatic
conditions, herbivory, and disease. The length of this ‘‘proba-
tionary period” will vary with conditions and with the species.
In the present study, Hedyotis was a species that flowered in its
first or second year, but seedlings of the other species still have
not reached reproductive size.
These experiments suggest that establishment of new popula-
tions of these species may be a very rare event, and thus suc-
cessful human reintroduction by seed will also be rare. There is
a need for more exploration of the biology of the particular spe-
cies involved, which may lead to the specification both of dis-
persal conditions and of horticultural practices that could protect
the seedlings that do emerge. Some species in this experiment,
with a single input of seeds, performed better than others. The
interaction between seed-colonist and the environment at the time
of arrival means that performances are likely to differ from year
to year (as seen in Vasseur and Gagnon 1994), and that both
abiotic and biotic conditions, including competition with other
species, are important factors (Berger 1993). It is clear in any
case that, given the low percentage of emergence for most species
in the field, reintroduction by seed requires the use of a large
number of seeds and probably more than one year. The number
of propagules used (assuming that the supply is plentiful) will
depend in part upon the ultimate population size deemed desirable
for viability in the reintroduction site. What size is sufficient for
“viability” is a subject of current research, though it is safe to
say that generalizations are perilous at the moment, since regard-
less of the definition of viability used, there remain major areas
of uncertainty that can only be resolved by longitudinal studies.
In any case, we can only conjecture how resilient a population
will be, given all possible disturbances over any particular stretch
of time (is the target 50 years? 500 years?; Howald 1996; Menges
1991; Pavlik 1996).
The present experiments show (over the course of three years’
data collection) rates of ‘‘establishment”’ (in a limited sense) from
seed dispersal ranging from about 6% to far less than 1%, with
an average around 1%. Using that figure, if the goal is a popu-
2000] Drayton and Primack—Reintroduction of Perennials 323
lation of 50 individuals, one would use 5000 seeds. This large
number of seeds would only grow larger if one’s target population
was, for example, 500, as suggested by some researchers, in order
to provide a population that might be resilient to disturbance and
environmental stochasticity over some length of time. In fact,
several of the species in this study were introduced in numbers
approaching this figure. In the short term, only two species might
be said to be present in the numbers desired (Hedyotis and San-
guinaria), but they are present not in one population but several.
This raises another design consideration that has entered the
design of plant reintroduction plans only recently, that of meta-
population structure (McEachern et al. 1994). Metapopulation
theory has formalized the insight that species often exist in pop-
ulations of populations, patchy concentrations in the landscape at
varying distances from each other, joined by gene flow in various
forms at a low rate. It is thought that this structuring of a species’
population provides resilience to disturbance not provided even
by a very large single population. The appropriate size and place-
ment of introduced populations or subpopulations is not only a
matter of ‘distributing the risk’” across varying habitats but also
of ensuring that there are enough individuals to support cross-
pollination when the species is not self-compatible. In the case
of the species that have shown the most flowering success in this
study (Sanguinaria, Hedyotis), the fact that they are pollinated by
generalist pollinators may have promoted fruiting success, while
Aquilegia, which showed good flowering but relatively poor fruit
set in both years, may have been pollinator-limited in the areas
in which the plants occurred, being too widely spaced to attract
hummingbirds. In the Hammond Woods, the flowering individ-
uals were widely separated, and there were no other stations of
the species present. In the Middlesex Fells, Aqguilegia did occur
naturally, and it appears that fruit set was somewhat higher there,
but further monitoring would be necessary to establish trends.
The attraction of appropriate pollinators remains a critical factor
for the success of introduced species that require animal or insect
pollination vectors.
In the design of a reintroduced population, especially when site
characterization may be approximate or missing some critical fac-
tor, a plan which disperses the reintroduced propagules in more
than one site is an attempt to build in the resilience that the
metapopulation may provide. In addition, the reintroduction does
324 Rhodora [Vol. 102
not risk all its resources on one or a few sites’ viability at the
time of reintroduction, thus “‘sampling” the landscape for a wider
range of safe sites (Harper 1977). This assumes as part of the
reintroduction plan that the multiple sites of introduction will
show varying rates of success and persistence, as in any coloni-
zation beyond a population’s area of concentration (Prince and
Carter 1985; Prince et al. 1985). Despite the best efforts of trained
ecologists, it may be difficult to identify the critical environmental
factors that allow or prevent the establishment of new popula-
tions. Selecting several or many sites for initial attempts increases
the chance that at least some will be successful. The sites that
show initial promise can then become target sites for more ex-
tensive reintroduction efforts.
This raises another point, however, which is relevant to rein-
troduction efforts: the “sampling” of the topography of time as
well as space. The strategy of very large inputs at one point in
time is convenient in the construction of emergency rescue plans
for threatened species, and for the creation of research programs
for doctoral theses, but it may be well to structure reintroductions
by seed to include the axis of time in the population structure.
Thus, a particular Hedyotis or Sanguinaria individual may dis-
perse at most two dozen seeds in a year. Perennials, however, are
iteroparous, that is, they will under most conditions disperse seeds
year after year. Thus their dispersal *“‘shadow”’ will take into ac-
count the interactions of site with climate. The plant conserva-
tionist may well wish to do the same, thus adding repeated dis-
persals to the same sites over the course of several years. In this
case, the 50 or 500 plants in the final target metapopulation would
not be the result of a single dispersal of 5000 or 50,000 seeds,
but of a smaller annual deposit continued for several years.
The experiments reported here, however, show that where
transplantable material is available for use, one is much more
likely to achieve success in a reintroduction by means of trans-
planting of individuals past the seedling stage. This is supported
by the results of experiments with Potentilla robbinsiana men-
tioned earlier. As discussed in the introduction to this paper, there
are important advantages to the use of seeds as the method of
reintroduction. Nevertheless, success rates are generally much
higher with established individuals than with seeds. The number
of individuals required is smaller than the number of seeds,
though the cost per individual is higher: to reach a population of
2000] Drayton and Primack—Reintroduction of Perennials 325
50 to 500, with a success rate of 25% (plausible, based on the
results reported here), would require an input of 200 to 2000
individuals, again probably distributed over multiple sites. The
mieneh rate of success per propagule makes it more possible to
‘‘structure’” something like a metapopulation. With even as few
as four individuals per quadrat, a series of 100 quadrats spread
across a target location could produce several populations sepa-
rated by enough distance to provide some protection against dis-
turbance, but close enough for occasional long-distance seed dis-
persal or exchange of pollen. In the present experiments, sites
were usually clustered, with three or four replicates of the ex-
perimental unit in one general area, separated by no more than
10 meters. The next experimental site was from 50 to 500 meters
distant. In cases like Caltha or Sanguinaria where there were
multiple occupied quadrats, the result in effect was a metapopu-
lation.
Yet there is still the question of the definition of success. For
these experiments, success cannot be determined as yet, because
for these perennial species, time to reproductive maturity may be
as much as five years or more. Thus individuals established from
seed, or from the transplant of young plants, will not begin to
reproduce for some time, if they survive. Even for reproducing
individuals, though, the monitoring time must be on the order of
a decade or more. This is in part because of the dormancy of
seeds and in part because of the relatively small number of seeds
dispersed per plant per year. If the locale is suitable for the species
(as may be deduced prima facie from the survival and reproduc-
tion of transplants), it may not always be suitable for seedlings,
as demonstrated by these same experiments. Thus if a Sangui-
naria is dispersing 15 seeds per year, with a success rate of per-
haps 6% it may take 2—5 years for these seeds to result in new
seedlings that persist for more than a year or two. The need for
a long time-horizon is emphasized by the attempts to create new
populations of the endangered orchid, Small-whorled Pogonia (/s-
otria medeoloides) using wild-collected adult transplants (Brum-
back and Fyler 1996). While there was a good rate of survival
for the first 5 years after the transplants, after 8 years virtually
all plants had died out and the remaining plants were no longer
in flower.
The experiment reported here suggests that a reintroduction
program should include reintroduction by more than one method
326 Rhodora [Vol. 102
since, as argued above, reintroduction by seed and by transplant
each has its advantages. Further, the reintroduction should be de-
signed when possible to provide new information about the bi-
ology of the species under consideration. Although the species
used in this study are common features of the New England flora,
there is little information available about their population biology
and demography, about the applicability of the metapopulation
model to them, or about the frequency and conditions under
which new populations arise. Finally, it is clear that given the
numerous hurdles that a reintroduction effort may encounter, pro-
tection of existing populations remains the fundamental ingredi-
ent in any conservation plan (Falk 1991; Lesica and Allendorf
1992), and “mitigation” of habitats even with species that are
not threatened should be done with caution. If attempts are made
to create new populations, these attempts should involve exam-
ining multiple sites and methods over a period of years to increase
the chances of success.
ACKNOWLEDGMENTS. This project was made possible by a
grant from the National Science Foundation (to R.B.P.) and a
Boston University Fellowship (to B.D.). Permission to use the
field sites was granted by the Newton Conservation Commission
and the Metropolitan District Commission. Numerous people as-
sisted in the fieldwork, most notably members of the Drayton and
Primack families; all these helpers we are glad to thank here.
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RHODORA, Vol. 102, No. 911, pp. 332-354, 2000
THE MYTH OF THE RESILIENT FOREST: ey STUDY
OF THE INVASIVE NORWAY MAP
(ACER PLATANOIDES)
SARA L. WeBB', MARC Dwyer, CHRISTINA K. KAUNZINGER2, AND
PETER H. WycCKoOFF?
Drew University, Biology Department, Madison, NJ 07940
e-mail: swebb@drew.edu
*Current address: aces University, Department of eee Evolution,
and Natural Resources, New Brunswick, NJ 0
‘Current ine Guilford College, Biology eee
Greensboro, NC 27410
ABSTRACT. In a New Jersey Quercus—Fagus—Acer saccharum forest
(Drew University Forest Preserve), the exotic tree Norway maple ee platan-
oides) is a major and pile ng presence in all size classes. Norw le
long one of North America’s favorite shade trees, is invading Eee preserves
in New Jersey and dicate Research in the Drew Forest Preserve shows
that the forest is not a tight, resilient unit that repels invasions. Norway maple
does not rely upon disturbance or edges. Norway maple seedlings are shade-
tolerant and abundant, far outnumbering sugar maple and beech seedlings in
the heart of the forest preserve. An analysis of population size and age struc-
ture shows that Norway maple trees date back to 1915 or earlier and are
present in all subsequent age classes. Spatial studies show clumped distri-
butions for Norway maples as for other plant species but show no restriction
to edges. In the understory, species richness is significantly lower beneath
Norway maple than under sugar maple or beech, and most stems under Nor-
way maple are additional Norway maples. A restoration experiment (with
tree removal plots and seedling removal plots) was begun in 1998 in a patch
of 75-year-old sugar and Norway maples; this unusually simple two-species
area will elucidate the competitive erection over time. Other invasive spe-
cies also penetrate relatively undisturbed forest; Lonicera japonica, Alliaria
petiolata, Berberis thunbergii, and Wisteria tia are i asi alongside
orway maple in the oie Forest Preserve is disturbance needed to
admit exotic pathogens or insect pests that ia native trees. The threat to
the eastern deciduous forest is grave. If the forest is not sufficiently resilient
to weather the onslaught of intentional and accidental introductions, then
intervention is urgently needed.
Key Words: Acer platanoides, Acer saccharum, biological invasions, forest
ecology, New Jersey, northern hardwoods forest, succession
Invasive species pose one the greatest of all threats to forest
integrity in eastern North America, especially in conjunction with
other anthropogenic changes: forest fragmentation and climate
ees
2000] Webb et al.—Invasive Norway Maple i
change (Vitousek et al. 1997). The eastern deciduous forest has
faced devastating invasions by introduced pathogens, particularly
the chestnut blight and Dutch elm disease, which virtually elim-
inated host trees from the canopy. Invasive phytophagous insects
have also exerted profound influence (Niemela and Mattson
1996), including the gypsy moth and the hemlock wooly adelgid
(Orwig and Foster 1998).
When it comes to introduced plants, many ecosystems world-
wide have been transformed by invasive plant species, for ex-
ample, Myrica fava in Hawaii (Vitousek and Walker 1989; Vi-
tousek et al. 1987; Walker and Vitousek 1991), Melaleuca quin-
quenervia and Schinus terebinthifolius in Florida (U.S. Congress
1993), several pines in South Africa (Richardson 1998; Richard-
son et al. 1994), and Hieracium lepidulum in New Zealand moun-
tain beech forests (Wiser et al. 1998). Wetlands are under siege
by invasive plants such as Lythrum salicaria (Hight and Drea
1991; Thompson 1991), Phalaris arundinacea (Apfelbaum and
Sams 1987; Barnes 1999) and Phragmites communis. Since El-
ton’s seminal book on the subject (Elton 1958), biological inva-
sions have been studied by a growing number of scientists (com-
pilations include Groves and Burdon 1986; Luken and Thieret
1997; Mooney and Drake 1986; U.S. Congress 1993).
The eastern deciduous forest has also begun to acquire a heavy,
if geographically patchy, load of invasive shrubs, vines, and
herbs: Lonicera japonica (Evans 1984), L. maackia (Luken 1988;
Pringle 1973; Williams et al. 1992), Berberis thunbergii (Ehren-
feld 1997), Rhamnus frangula (Post et al. 1989), Euonymus alatus
(Ebinger et al. 1984), Elaeagnus spp. (Randall and Marinelli
1996; Szafoni 1991), Alliaria petiolata (Baskin and Baskin 1992;
Byers and Quinn 1998; McCarthy 1997; Nuzzo 1991), and many
others (Randall and Marinelli 1996; U.S. Congress 1993).
Is the eastern deciduous forest also threatened by invasive
trees, alongside these smaller plants? Many exotic trees thrive at
forest margins (Ailanthus altissima; Paulownia tomentosa [Wil-
liams 1993]) and can be battled by minimizing forest fragmen-
tation. Is this true for all invasive trees, or might some also invade
the forest interior? When invasive plants move into the forest,
what are the consequences for the community as a whole? Here
we examine the invasion of Norway maple as a case study.
At first glance, the small forest preserve on Drew University’s
campus in New Jersey seemed a natural and intact forest ecosys-
334 Rhodora [Vol. 102
tem, a mixture of mature (though not old growth) oak, beech,
and maple. However, closer inspection revealed that a large and
growing proportion of the maples were not native sugar maples
(Acer saccharum) but were instead an introduced species, the
superficially similar Norway maple (A. platanoides; nomenclature
follows Gleason and Cronquist 1991).
Norway maple can be distinguished from its native congener
by its milky sap, glabrous lower leaf surface, early yellow-green
flowers in erect corymbs, and schizocarp wings at an angle of
180° (vs. 120° in sugar maple). This native of continental Europe
was introduced intentionally to Philadelphia around 1760 (Spong-
berg 1990). It is now among the top-selling shade trees in North
America (Nowak and Rowntree 1990; Nowak and Sydnor 1992).
While Norway maple is still planted along the streets of America,
it is simultaneously removed, often with considerable difficulty
and expense, from natural areas where it has spread, from New
York’s Central Park to Pennsylvania to Ontario. Unlike many
horticultural plants, Norway maple has the capacity to reproduce
and spread in diverse habitats. It is widely naturalized not only
in North America but also in the British Isles (Streets 1962; Webb
ibs wy)
Here we pose several questions about Norway maple as a po-
tential threat to the eastern deciduous forest. The overriding con-
ceptual issue is to what extent a forest resists invasion; the prag-
matic issue is to what extent intervention is thus necessary to
maintain or restore forest integrity.
1. Is Norway maple truly invasive or simply a modest and
transient presence in the forest? Dendrochronology and pop-
ulation sampling shed light on the age structure and size
structure of the Norway maple population.
Does Norway maple depend upon edges, as do many light-
demanding plants, or is it successful in the heart of the
forest? Spatial analysis helps test for any edge effects.
. How does Norway maple influence the richness and struc-
ture of the forest? We compare the undergrowth beneath
Norway maple with that beneath native trees.
How can Norway maple be removed from invaded forests,
and with what consequences? A controlled restoration ex-
periment will permit tracking of the effects of removals of
both trees and seedlings.
_
os)
=
2000] Webb et al.—Invasive Norway Maple 335
STUDY AREAS
The Drew Forest Preserve encompasses approximately 18 ha
in Madison, Morris County, New Jersey (40°46'N, 74°26'W),
within the Loantaka terminal moraine of Wisconsin glaciation.
Soils are a mosaic of well-drained and somewhat poorly-drained
loamy soils derived from gneissic glacial outwash (Tedrow 1986).
Outwash deposits are interspersed with steep dells of ice-block
origin. More generally the Preserve lies within the Piedmont
physiographic province, in the region classified by Braun (1950)
as the central hardwoods area of the oak-chestnut region. The
vegetation has characteristics of both Mixed Oak and Sugar Ma-
ple—Mixed Hardwoods (Collins and Anderson 1994; see Table 1).
Historical perspectives on the region’s forests are provided by
Russell (198la, 1981b) and Ehrenfeld (1982).
The three components of this research utilized different study
areas. The first two projects focused on the most naturalized for-
est zone, the Forest Preserve Study Area, a second-growth forest
dominated by oaks, beech, and the two maples; composition and
age structure are detailed in “‘results.”” The original tree-ring plot
(Tree Ring Study Area) was a 0.25 ha plot situated within the
Forest Preserve Study Area (Webb and Kaunzinger 1993). The
second project, the study of understory influence, included the
Tree Ring Plot but also much more of the forest, for a total Forest
Preserve Study Area measuring 3.52 ha; sampling covered 14%
of this study area in randomly selected plots (Wyckoff and Webb
1996). Stand attributes are reported in Table 1.
The third project took place in the 1.8 ha Maple Study Area,
a different forest community of younger, even-aged (75—80 year
old) sugar and Norway maples with few trees of other species.
The Maple Study Area was chosen for the study of spatial pat-
terns and for the removal experiment because of its floristic sim-
plicity. Seven other tree species were present but at very low
densities in this study area: Fagus grandifolia (3.17% density),
Quercus rubra (2.65%), Acer rubrum (0.38%), Q. alba (0.33%),
Q. velutina (0.19%), A. saccharinum (0.8%), and Q. prinus
(0.04%).
MATERIALS AND METHODS
Detailed methods for the tree ring and understory projects have
been reported in Webb and Kaunzinger (1993) and Wyckoff and
Table 1. Forest composition and stand attributes, Drew enemy Forest Preserve Study Area (3.52 ha). Note that forest
mposition is somewhat different in the Maple Study Area; see text. Based on random sampling in circular plots with 10 m
ae Includes trees with DBH = 2.5 cm. (From Wyckoff and fone 1996.)
Density Relative Mean DBH Basal Area Relative
Species (#/ha) Density (cm) (m/?/ha) Dominance
Acer platanoides 161 26.0% 8.5 2.14 7.3%
Acer saccharum 217 35.1% 14.2 7.38 25.5%
Fagus grandifolia 123 19.9% 22.1 6.94 24.0%
Ulmus americana 20 3.2% 24.0 1.29 4.4%
Prunus aviu 20 3.2% OF 0.33 1.1%
Robinia aN acacia 16 2.5% 40.3 1.47 5.1%
Quercus velutina 12 1.9% 47.1 2.16 T4A%
Fraxinus americana 10 1.6% 34.3 0.96 3.3%
Quercu 10 1.6% 51.5 2.35 8.1%
Acer saccharinum 8 1.3% 41.4 1.28 4.4%
Betula le 6 1.0% 50.2 1.20 4.1%
Quercus rubra 6 1.0% 41.4 0.82 2.8%
Acer rubru 4 0.6% 17.8 0.11 0.3%
Cornus florida 2 0.3% 22.2 0.77 0.2%
Prunus pensylvanica 2 0.3% 2.5 <0.01 <0.1%
Liriodendron tulipifera 2 0.3% 48.4 0.37 1.3%
vIOpoyy
ZOT IOAI
2000] Webb et al.—Invasive Norway Maple o37
Webb (1996), respectively. In brief, the tree ring study censused
and cored all sizable trees (DBH = 5 cm) in 1988, within a 0.25
ha plot. Smaller stems were sampled in subplots. Increment cores
from the diffuse-porous trees were soaked and sanded before
counting. Thirteen of 74 tree cores were excluded from analysis
because of missing centers due to heartrot. Estimated ages omit
time before each tree reached the core-extraction height of 25 cm.
Age distributions for the four major taxa were compared using
the Komogorov-Smirnov statistical test (Sokal and Rohlf 1981).
To compare the understories of Norway maple with those be-
neath native trees, and also to extend our picture of the Norway
maple invasion outside the dendrochronology plot, Wyckoff and
Webb (1996) established a random sampling network in fall 1993,
sampling a 3.5 ha area of the forest. Sixteen circular plots, each
measuring 314 m’, were centered on points at coordinates derived
from a random number table. Plot centers within 20 m of the
forest edge were rejected. All trees with DBH = 2.5 cm were
identified and measured for stand attributes.
Next, to assess the understory influence of Norway maple rel-
ative to that of native canopy trees, we sampled subplots under
Norway maple, sugar maple, and beech canopy trees within the
tree plots. In each tree plot, we examined up to six canopy trees
by choosing the two individuals closest to the plot center for each
of the three target species. Random coordinates were used to se-
lect additional trees for a total of 20 canopy trees for each species.
At the base of each randomly chosen Norway maple, sugar ma-
ple, and beech tree, we laid out eight | m =< | m subplots, for a
total of 160 subplots under each canopy species. All rooted vas-
cular plant stems, including nonwoody species, were identified
and tabulated. For small individuals of the tree species, we dis-
tinguished between saplings (height > 1 m, DBH = 2.5 m) and
seedlings (height = | m). We used nested ANOVA (subplots
nested within canopy trees) to avoid pseudoreplication (Hurlbert
1984) when testing each understory species for differences at-
tributable to canopy species. Post-hoc LSD tests were applied for
pairwise comparison of means when ANOVA indicated signifi-
cance at P < 0.05. A broader understory sampling was added to
better capture patterns among understory trees and saplings; we
tabulated those trees and saplings that were 50% or more over-
topped by each of the 60 randomly selected trees.
In fall 1997, two of us (Dwyer and Webb) began a third pro-
338 Rhodora [Vol. 102
Road
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| | Control | | Go "00 | |
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Plots
Figure 1. Sampling scheme and arrangement of permanent plots in the
ae Study Area, with Norway saa removal plot and control (nonremoval
ot). Permanent plot markers are at centers of eac m 10 m tree plot.
ae plots measuring 2 m x 2 m are nested within tree plots for four of
the five transects.
ject: an experiment in Norway maple removal and, ultimately, in
forest restoration. The 1.84 ha Maple Study Area was bisected
into a 0.914 ha removal plot and a 0.929 ha control plot (Figure
1); asymmetry resulted from old fencing being used to delineate
the area. Before trees and seedlings were removed, trees and seed-
lings were sampled in detail.
Five evenly-spaced transects were partitioned into 20 adjacent
linear plots measuring 2 m wide by 10 m long (the westernmost
plot in each transect was 7 m long; Figure 1). The plot centers
2000] Webb et al.—Invasive Norway Maple es ied
were permanently marked by rebar stakes with labeled metal
caps. In each plot, all woody stems with height = 1.5 m were
identified and measured. Thus this “‘tree’ category includes
smaller stems than the “‘tree’’ categories used in the above studies
of tree-ring and understory influence. This partial sampling of
trees, necessary to assess spatial patterns and statistical relation-
ships, was supplemented by a complete census of trees in both
removal and control areas. Norway maples were flagged during
sampling within the removal plot.
Seedlings (height < 0.5 m) were also sampled, using subplots
nested within the tree plots, for four of the five tree transects.
Seedling plots measured 2 m X 2 m and were centered on the
permanent markers within tree plots to facilitate future resam-
pling. Patterns were analyzed with regression using SPSS (No-
rusis 1993) and with tests of dispersion (Krebs 1999).
The removal component of this study included both trees and
seedlings. For trees, 428 flagged Norway maples were cut down
or girdled in January 1998. Machetes were used on small stems,
while 29 trees were too large for this method and were either
girdled or felled by chain saw. A total of 1430 Norway maple
seedlings were pulled up by the roots, as they were tabulated, in
two of the four transects, in 40 subplots with total area of 160
m?’. Seedling removal and seedling control (nonremoval) subplots
were situated in both tree removal and tree control plots, so that
the interaction between seedling and tree removal can be exam-
ined in the future.
RESULTS
Norway maple population age structure. Norway maple
was present within the Drew Forest Preserve by 1915. It has been
a major component of all age classes since then, with especially
heavy recruitment in the 1940s and subsequent decades (see
Webb and Kaunzinger 1993 for details). Figure 2 shows age dis-
tributions for trees with interpretable cores. The Norway maple
age distribution is not significantly different from those of sugar
maple or American beech (Kolmogorov-Smirnov test, P > 0.05),
though it is different from the age distribution profile for the oaks.
Note that apparent shortages of trees in younger age classes sim-
ply reflect the lower diameter limit for coring (10 cm in this
study). Size profiles for each tree population fill in the smaller
340 Rhodora [Vol. 102
8 Oak
: | Norway maple |_| |
Trees in 0.25 ha Plot
a n
7 ert | Nee
Sugar maple |
N 2 © OD “~
» oa ¢
Trees in 0.25 ha Plot
beer corer
| a TTT
> ~ > Ww
_il
4
: a
A 85 105 125 145 1
Age Class Age Class
Figure 2. Age distribution for the four major taxa in the Tree Ring Plot.
The “oak” category combines Quercus velutina, Q. alba, and Q. rubra. Ap-
parent ied of younger trees reflect lower diameter limit for Sle (10
cm), compare with Figure 3. Age distribution of Norway maple was signif-
icantly different from that of oaks but not from beech or sugar aoe (Kol-
mogorov-Smirnov test). From Webb and Kaunzinger (1993).
trees but otherwise parallel age profiles (Figure 3). American
beech was present by 1890 and sugar maple by 1920; like Nor-
way maple, both of these shade-tolerant species have been repro-
ducing ever since.
The overall picture of forest age structure suggests that Norway
maple appeared just as the forest became too closed for further
oak reproduction. The oldest and largest trees were oaks: Quercus
velutina, Q. alba, and Q. rubra, with estimated ages of 79-154
years (origins between 1835 and 1910; Webb and Kaunzinger
1993), and with open-grown crown morphology reflecting estab-
lishment when the site was still open pasture. The ongoing ability
of Norway maple trees to establish since forest closure attests to
its shade tolerance and ability to invade beyond disturbed ground
and sunny edges.
2000] Webb et al.—Invasive Norway Maple 341
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Diameter Class (cm) aR Class (cm)
igur Size distribution graphs ae the four major taxa in the Tree Rin
Plot, Bee all trees with DBH > 5 cm. From Webb and Kaunzinger
(1993):
Additional tree-age data are also available for the very different
Maple Study Area, the mixed stand of sugar and Norway maples.
Here, canopy trees were mostly even-aged at 75—80 years (in
1997). Both maple species were represented by abundant seed-
lings and saplings too small to core, demonstrating once again
their tolerance of shady, competitive environments.
Norway maple abundance, dominance, and reproduc-
tion. The abundance and size distributions for Norway maple
varied between study areas but in all localities showed an influ-
ential role for this species. Norway maple trees (DBH > 5 cm)
ranked second in abundance (17.2% relative density) behind
beech, among 13 tree species present in the Tree Ring Plot (Webb
and Kaunzinger 1993). In the larger Forest Preserve Study Area
(Wyckoff and Webb 1996), Norway maple also ranked second in
abundance (26%) but was outranked by sugar maple rather than
beech (Table 1). In the younger Maple Study Area, Norway ma-
342 Rhodora [Vol. 102
ple and sugar maple were nearly even in tree density (46.5% and
45.4% of stems); note that tabulations for this study area included
saplings as well as trees (all stems with height > 1.5 m).
Norway maple’s youth relative to oaks and beeches in the
Drew Forest gave it somewhat lower rankings for dominance, an
estimate of biomass based upon tree diameters (Table 1). In the
Tree Ring Plot, Norway maple was fourth in dominance (8.2%
of basal area, behind black oak, beech, and white oak; Webb and
Kaunzinger 1993); in the larger Forest Preserve Study Area, it
ranked fifth in dominance (7.3%, behind the same three species
and sugar maple). In the Maple Study Area, Norway maple
ranked second in dominance (20.6%) behind sugar maple
(52.4%); Quercus rubra ranked third (12.43%) with a few (2.65%
of density) very sizable trees; none of six other tree species had
more than 1.5% relative dominance.
Perhaps most revealing about the future of the forest are seed-
ling and sapling data. In these small size classes, Norway maple
showed an extremely strong presence in all study areas. In the
Tree Ring Plot, Norway maple accounted for nearly 60% of all
seedlings (DBH < 5 cm), with over 2000 stems per ha, or one
per five square meters (Webb and Kaunzinger 1993). In the youn-
ger Maple Study Area, dense carpets of very small maple seed-
lings (counting only stems with height < 1.5 m) comprised a 50:
50 mixture of Norway and sugar maple seedlings, with an aver-
age of 50 Norway maple seedlings per five square meters.
Spatial patterns and edge effects. Norway maple was not
restricted to forest margins; all size classes were abundant
throughout the forest interior. Both trees and seedlings of Norway
maple exhibited clumped dispersion patterns, but this was also
true for the native sugar maple (in Maple Study Area, for both
Morisita’s Index and the Variance-to-Mean Ratio; Krebs 1999:
Sokal and Rohlf 1981).
The Maple Study Area had two edges whose influence differed
sharply. The first edge was a major two-lane roadway toward the
west (Figure |); surprisingly, Norway maple dominance (as basal
area) decreased significantly toward this edge (regression, P =
0.038). A row of planted sugar maples along this roadway sug-
gests that the proximity of seed sources or perhaps some com-
petitive effect was at work; physical aspects of the edge itself did
not promote Norway maple growth in this case. Beyond this dom-
2000] Webb et al.—Invasive Norway Maple 343
ine) ee) & oOo
| i a il
Understory Species / Canopy Tree (in 8 m. sq.)
|
1o)
|
Norway maple Sugar maple Beech
Figure 4. Understory species richness beneath Norway maple as com-
pared with sugar maple and beech. Bars show average number of species in
subplots measuring 8 m*. Species richness was significantly lower beneath
Norway maple. Bars indicate one standard error. From Wyckoff and Webb
(1996).
inance response, distance from this edge did not influence density
of Norway maple trees or seedlings, nor density or dominance of
sugar maple trees or seedlings.
The second edge was a residential yard to the south of the
Maple Study Area (Figure 1). In contrast to the roadway, this
yard edge fostered significantly higher densities of Norway maple
seedlings (height < 1.5 m; regression, P = 0.040) without a sim-
ilar effect on sugar maple seedlings. Seed source again seemed
to play a more influential role than edge microclimate or light
levels. The residential yard had a dozen mature Norway maples
adjacent to the forest before the residents removed most over the
Table 2. Plant species in understory plots of the Drew Forest Preserve Study Area, with mean densities under each canopy type ~
(per 8 m?*). Significant differences are reported in the last column, based on Fisher LSD (P < 0.05). B = beech, SM = sugar maple =
NM = Norway maple.
Canopy Type
P Differences
Understory Species NM SM B (ANOVA) (P < 0.05
Trees
Acer platanoides (yearling) 54.95 16.5 22.30 0.0111 B = SM < NM
Acer platanoides (older seedling) 12.15 7.80 6.70 0.3365
Acer platanoides sae ing) 3.00 1.05 1.55 0.2592
Acer saccharum (yearling) 0.20 1.30 1.60 0.2729
Acer saccharum (older seedling) 1.15 3.00 3.50 0.3925
Acer saccharum (sapling) 0.00 0.95 0.10 0.0039 B = NM < SM as
Fagus grandifolia (seedling) 0.05 0.25 0.95 0.0264 NM = SM < B 4
Fagus grandifolia (sapling) 0.00 0.40 0.20 0.4075 ¢
Shrubs, Set and Forbs a
Actaea r 0.05 0.00 0.05 0.6092
Aster ei 0.45 0.95 1.60 0.7470
Se virginiana 0.15 8.75 33:15 0.0064 NM = SM <B
Fraxinus americana 0.50 1.00 1.35 0.7433
Ilex verticilla ta 0.40 0.70 0.85 0.6493
Lindera benzoin 0.60 295 0.05 0.0308 B = NM < SM
Lonicera japoni 4.25 7.80 0.05 0.1607
Maianthemum canadense 0.00 2.60 0.00 0.3742
Mitchella ns 0.00 1.90 0.00 0.3742 =
ae quinquefolia 0.05 3.70 O25 0.6505 =
Polygonatum pubescens 2.00 0.70 2.65 0.1823 a
Polygonium virginiana 0.05 0.00 0.00 0.3742 =
Table 2. Continued.
Canopy Type
P Differences
Understory Species NM SM B (ANOVA) iP =-0;03)
Robinia pseudoacacia 0.00 0.05 0.00 0.3742
Sassafras albidum 0.00 0.05 0.00 0.3742
Toxicodendron radicans 0.75 0.25 0.20 0.5465
Uvularia sessilifolia 0.00 0.00 0.05 0.3742
Vaccinium corymbosum 0.25 0.30 0.00 0.3286
Viburnum acerifolium 3.55 3.85 3.70 0.9940
Total Richness 16 18 15
[0007
Te 19 GQaMy
ajdepy APMION DAISVAUT
Sve
346 Rhodora [Vol. 102
past decade. A clump of sizable Norway maples was also found
within the forest near this edge. Even more telling: when seedling
density was regressed on both dominance (basal area of Norway
maple) and on distance from the edge, the distance factor lost its
significance (P = 0.874) while dominance, as a proxy for large
trees, more fully explained the pattern (P = 0.004).
While seed sources and competitive effects thus seemed to in-
fluence Norway maple, it is important to note the lack of a spatial
relationship between trees and seedlings, either presence or abun-
dance, on the finer spatial scale within the forest. For neither
Norway maple nor sugar maple seedlings was it necessary to
have mature conspecific trees within 10 m or more.
Norway maple influence on undergrowth. The understory
beneath Norway maple was distinctive in its low diversity and in
its preponderance of Norway maple seedlings. Understory rich-
ness was significantly lower beneath the average Norway maple
(¥ = 3.20 species/8 m*) than beneath sugar maple (* = 4.55
species/8 m*) or beneath beech (¥ = 4.95 species/8 m°; Figure 4;
NOVA, P = 0.0074). Of 21 understory species present (Table
2), most were found under all three canopy species: 16 beneath
Norway maple, 18 beneath sugar maple, and 15 beneath beech.
Sugar maple supported the most distinctive assemblage, on av-
erage, with slightly more Lindera benzoin, Parthenocissus quin-
quefolia, Maianthemum canadense, Mitchella repens, and Loni-
cera japonica; however, only differences in abundance of Lindera
benzoin were Statistically significant at the level of the individual
species (Table 2)
The Norway maple understory was typically a carpet of Nor-
way maple seedlings, which comprised 83% of stems and 98%
of all woody seedlings beneath Norway maple trees. Paradoxi-
cally, Norway maple understories had slightly higher densities of
understory stems (84.65 stems/8 m?’) than beech (80.40) or sugar
maple (63.85; Figure 5). However, when we excluded Norway
maple stems from the totals, this pattern was reversed, as shown
by the filled bars in Figure 5, in a statistical trend toward lower
densities of non-Norway maple stems (ANOVA, P = 0.0881).
While Norway maple seems likely to replace itself in future
generations, it also promises to replace most beech and sugar
maple trees as well, if seedling and sapling survivorship continues
at current rates. Norway maple accounted for 81% of all woody
2000] Webb et al.—Invasive Norway Maple 347
S| With Norway maple
100-— No Norway maple
&
oO
|
Understory Stems / Canopy Tree (in 8 m. sq.)
ie) fe)
r r
o=
Norway maple Sugar maple Beech
Figure 5. gees stem densities beneath Norway maple, sugar maple,
and _ beech. show average number of vascular plant stems within 8 m?
plots. ca stems of Norway maple are included in filled bars but
excluded from shaded bars. From Wyckoff and Webb (1996).
wo
B
seedlings beneath sugar maples and 80% beneath beech (Figure
6). Only under sugar maple was there much prospect for future
canopy trees that were not Norway maples. Amongst larger sap-
lings and understory trees, the pattern was somewhat different
than for seedlings: sugar maples sometimes equaled Norway ma-
ple saplings in abundance beneath sugar maple trees (Wyckoff
and Webb 1996). Elsewhere the numerical dominance of Norway
maple suggests an increasing role in the future and a concomitant
decline in native wildflowers and shrubs throughout the preserve.
Effects of Norway maple removal. Results of the removal
experiment initiated in 1997—98 must obviously await the passage
of time and future resampling of permanent plots. The baseline
348 Rhodora [Vol. 102
100
ae Norway maple
oy anan
® 80- Sugar maple
c :
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® 60- ’
o v4) Other tree species
i ral
~
Qa
re)
©
@ 40-
~
Y
e))
&
8
® 20-
”
el TE war
0 |
Norway maple Sugar maple Beech
Figure 6. Differences in assemblages of tree seedlings under Norway ma-
ple, sugar maple, and beech canopies. Bars show mean seedling densities per
8 m?’ plot. Note that Norway maple seedlings were dominant under all three
canopy species.
sampling of the Maple Study Area provided insights into the
questions outlined above, specifically about population structure
and edge effects on Norway maple. Early observations showed
that most Norway maples had resprouted and would require ad-
ditional treatment to control. Light levels were greatly elevated
in the 0.88 ha experimental removal plot, where 399 small Nor-
way maples and 29 large Norway maples were cut down. The
abundant seedlings of both sugar and Norway maple in this Ma-
ple Study Area were exhibiting a rapid growth response to the
new openings. For the seedling removal plots, the future replen-
ishment of the maple seedling bank is of great interest for both
maple species.
2000] Webb et al.—Invasive Norway Maple 349
DISCUSSION
This case study of the Norway maple invasion depicts a species
that is highly successful at penetrating an intact second-growth
forest. Norway maple is shade-tolerant, a good competitor for
conditions in forest interiors, as evidenced by its presence in all
age and size classes throughout the forest (Webb and Kaunzinger
1993). Its dominance will almost certainly increase in the future.
Forest edges are not essential for Norway maple. However, edges
coupled with adjacent landscape plantings help accelerate its
spread by increasing the seedling bank. Large tracts of unfrag-
mented forest might be invaded more slowly. Norway maple has
advantageous physiological mechanisms besides its tolerance of
shade, including early leaf expansion and late leaf drop for a
longer growth season than is seen for sugar maple (Kloeppel and
Abrams 1995). Seeds are produced abundantly and dispersed
widely (Matlack 1987).
The ability of Norway maple to depress species richness
(Wyckoff and Webb 1996) raises concerns about preserving forest
integrity. Negative consequences are likely, both for populations
of native wildflowers and shrubs and for the future of native trees
with which Norway maple competes in the canopy.
The vulnerability of our forests to this tree and to other new
arrivals, such as the chestnut blight (Good 1968), the hemlock
woolly adelgid (Adelges tsugae; Orwig and Foster 1998), and
Japanese barberry (Berberis thunbergii,; Ehrenfeld 1997), chal-
lenges the myth of the resilient forest. These examples call into
question the common generalizations that invasions are most like-
ly following disturbance or in insular habitats. A theoretical
framework on biological invasions continues to elude us because
efforts to predict invasiveness have not been generally successful.
An expectation that forests will eventually recover from dam-
age derives in part from outdated equilibrial views of nature and
in part from the regional experience in northeastern North Amer-
ica where most abandoned fields and clear-cuts do indeed return
to a forested state. Several lines of evidence challenge this com-
placent view. Recent research has shown that apparent recovery
from clear-cutting does not include return of the entire herbaceous
flora (Bratton 1994; Duffy 1993; Duffer and Meier 1992; Matlack
1994). Foster (2000) has shown that the second-growth forest
does not exhibit the presettlement forest’s topographic patterning
350 Rhodora [Vol. 102
of tree species. Thus forests need not follow a rapid trajectory to
some original steady state following perturbation. Indeed, such
recovery is unlikely in the face of complex interactions among
anthropogenic impacts: forest fragmentation, global warming, and
introduction of invasive species.
The Norway maple population explosion and the onslaught of
other invasive species together suggest that we are asking the
wrong question when we wonder why introduced species become
invasive. The real question is, ““Why not?’? What prevents some
species from becoming invasive, and what keeps in check those
species like Norway maple that seem capable of more widespread
colonization? What causes an invasive species to shift over time
from an occasional presence in the wild to a community-trans-
forming dominant?
The future of the New England forest, and of the eastern de-
ciduous forest as a whole, is very much at a crossroads. If exotic
species continue to enter the region and if invasive species are
not deterred, the result will be dramatic shifts in forest compo-
sition, structure, and function.
To protect and restore the eastern forest will take hard work
and creativity. Three distinct steps must be taken. First, invasive
species must be forcibly removed from natural areas and _ sur-
rounding lands (Randall and Marinelli 1996; Sauer 1998). Land
managers, conservation organizations, and volunteers already
have taken up this daunting charge, although much more is need-
ed. The second step is prevention of further invasions. Legal ac-
tion is essential to restrict intentional plantings of invasive spe-
cies. However, to achieve this aim also requires a cultural shift
by which native plants become desirable and even fashionable
choices in horticulture and landscaping. Local native plant or-
ganizations have had some success in promoting native plants.
An important element of success is availability and affordability
of diverse native plant materials. The third front in the war on
invasives is restoration: replacing the lost plants and providing
environments in which they can thrive (Drayton and Primack
2000). For highly disturbed or heavily invaded sites, research in
herbaria and in historical archives may be needed to reconstruct
in detail which species once grew in which habitats. Paleoeco-
logical records provide insights into tree assemblages of the past,
but offer little insight into the nonarboreal flora. In addition, sur-
vival of replanted wildflowers is compromised by deer, whose
2000] Webb et al.—Invasive Norway Maple 351
populations are elevated by human interference (Alverson et al.
1988, 1994). There is little hope that forests can maintain or re-
cover their diversity without active intervention from Homo sa-
piens, the species that has posed so many challenges to forest
integrit
ACKNOWLEDGMENTS. We are very grateful to members of the
Drew University grounds crew and especially grounds manager
Michael Virzi for their invaluable work on the Norway maple
removal project. We thank Raegan O’Lone for help with recent
field work, and countless other students and friends for past as-
sistance with the project. We also thank neighbors and friends of
the Drew Forest Preserve for their vigilance and concern: Greg
Leuser, Chris Hepburn, Mirelle Bessin, and Rolf Bessin.
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RHODORA, Vol. 102, No. 911, pp. 355-360, 2000
CLOSING REMARKS
W. DONALD HUDSON, JR.
The Chewonki asin ye 485 Chewonki Neck Rd.,
asset, ME 04578-4822
pene aes ae
When I spoke to the New England Botanical Club (NEBC) last
year about a naturalist’s perspective on science education, I ap-
proached the task with a bit of trepidation, for I have been work-
ing outside of a formal academic institution for nearly 20 years.
Providing the epilogue for today’s symposium is slightly more
daunting.
A word for those who may be new to the NEBC: the NEBC
has been around a long time. This is a wonderful association of
academic and amateur botanists who have been sharing their
work and experiences on the first Friday of the month since 1896.
David Barrington, a professional, academic botanist, and Les
Eastman, a self-trained amateur, introduced me to the NEBC 23
years ago this month. I clearly remember Friday evenings in the
room at the top of this building, participants packed between
glass-topped cases of Richard Evans Shultes’ ethnobotanical col-
lections to listen to a young man named Michael Donoghue talk
about Acer and Viburnum in the mountains of Mexico. Or, a
similar crowd listening to Les tell stories of searching for orchids
in cedar swamps of Aroostook County, Maine, with George New-
man and some eager student by the name of Les Mehrhoff in
tow. So when the committee asked me to wrap up this day, I
looked back there for some inspiration.
If you read through the early numbers of Rhodora, you will
find reports of expeditions, explorations, and investigations that
provided some of the academic foundations for the reports that
you listened to today. Certainly that is the case for the paleoeco-
logical investigations of George Jacobson and Ray Spear, which
elucidate dramatic changes in the structure of the forests, upper
elevational limits of species, shifts in treeline, and linkages of
these changes to fluctuations in temperature and other measures
of climate. Ray Spear’s work in the mountains, in particular, could
be a page right out of Fernald’s work, had he lived to be 130.
Notable for those of us who love to think about the mountains,
the 700 year period during the Younger Dryas, when average
355
356 Rhodora [Vol. 102
annual temperatures are estimated to have changed 5—10°C, must
have wrecked havoc on the prior (relative) stability of the post-
glacial alpine tundra vegetation and its associated mega-herbi-
vores. During the last 20,000 years our New England flora has
been transformed through the utter devastation of an ice age and
the somewhat more subtle influences of human commerce—not
only with axes and skidders in the 19th and 20th centuries, but
schooners and a thirst for riches that sent (mostly) northern Eu-
ropean people around the globe at a frantic pace in the 16th, 17th,
and 18th centuries, with microbes, pathogens, plant propagules,
rats, and pigs in tow (Crosby 1986).
Notable among their contributions, Charlie Cogbill, David Fos-
ter, and Les Mehrhoff have shown us the irrefutable value of the
written human record for elucidating the presettlement and early
colonial forests and associated vegetation of this region. I admit
a great affection for studies that combine evidence gleaned from
multiple disciplines, and today’s presentations fit the bill. Our
modern-day concerns about rapid plant migrations, and the ex-
traordinary loss of populations and whole species to both habitat
destruction and displacement by aggressive intruders, takes on a
whole new hue when we consider that it is just the most recent
wave of the biological expansion. Trifolium repens was an early
scout, now found on every continent. Sara Webb’s studies of Acer
platanoides show us just how aggressive are some of these rel-
ative newcomers to our flora.
The work presented today represents hundreds (if not thou-
sands) of years of collective inquiry, investigation, and analysis.
I know for certain that at least one of our speakers has years of
unfinished work stretched out before him—am I not right, Charlie
(Charles V. Cogbill)? I owe him notes and descriptions of vege-
tation for at least two remote mountain peaks in Maine before he
can complete his exhaustive survey of alpine vegetation in north-
ern New England and New York.
In the remaining few minutes of this celebratory symposium,
I offer a few thoughts and an announcement about future botan-
ical explorations—what you and your students ought to be think-
ing about for work in the coming century. I offer these thoughts
with that certain trepidation that comes before telling someone
with more experience what they ought to be doing with their life,
but . . . why not?
Many of you know that I have been railing for years that the
2000] Hudson—Closing Remarks aa]
science of our late 20th century is a fragmented discipline. Stu-
dents and the general public are poorly prepared to solve prob-
lems, not knowing where to begin to address landscape-scale is-
sues in botany, zoology, or ecology, let alone contribute to dis-
cussions affecting public policy and the stewardship of natural
resources.
Thought # 1: Teach. Or, inspire learning! The numbers
wouldn’t mean too much, but I'll wager that the general public
knows less of the world of (whole) plants in 1999 than they did
in 1899. There are certainly fewer botany or plant science majors
on most campuses that still support the discipline than there were
in 1900 or 1950 or even 1975. Please correct me where I may
go wrong here. We’re all smart enough to do something about
that, but it means creating a working environment that invites
participation by the sheer energy and high quality of its intellec-
tual inquiry.
Notably, Richard Primack’s efforts in restoration ecology have
provided a touchstone for the residents of Newton, and I imagine
that his particular botanical genius has already infected a gener-
ation of future botanists. Community members experience the ex-
hilaration of having ‘“‘made a difference,” and I’d wager that
Richard gets a real lift by sharing in that enthusiasm. And, as a
result, we know a lot more about the establishment of perennial
plants in new environments.
You know I love to explore, most recently on Baffin Island in
Nunavut, so the next thought should come as no surprise.
Thought # 2: Travel and explore with students and friends.
As the stationary tools and aids to learning continue to amaze
and distract us, so I think we need to redouble our effort to get
teams of collaborative learners and teachers into the field, if for
no other reason than to avoid William Morton Wheeler’s “dry
rot of academic biology.’ You may not have read Wheeler’s hu-
morous analysis of the state of biology, delivered in Boston 75
years ago, as the Presidential Address to the American Society
of Naturalists (Wheeler 1923). He said that we suffered then from
an academic form of Merulius lacrymans, or the dry-rot fungus:
“Undoubtedly the best culture medium for the academic
dry-rot fungus consists of about equal parts of narrow, un-
358 Rhodora [Vol. 102
sympathetic specialization and normal precocious senile ab-
straction.””
In his own fight against the tyranny of specialization, Wheeler
preferred ‘‘natural history” to “ecology,” then a young branch
of our life sciences. He stated:
“History shows that throughout the centuries, from Aristotle
and Pliny to the present day, natural history constitutes the
perennial root-stock or stolon of biological science and that
it retains this character because it satisfies some of our most
fundamental and vital interests in organisms as living indi-
viduals more or less like ourselves.”
Herb Wagner’s poignant review of the past 50 years of botan-
ical research, and especially the merciless tug-of-war on the psy-
che of the field and herbarium worker, is telling, particularly in
light of Wheeler’s earlier comments. Modern systematists have
had to adapt to increasingly more frequent changes in techniques
and tools, from the early population studies of the likes of Greg
Anderson and Charlie Heiser, through the era of numerical tax-
onomy, countless chemical analyses, to sizing gels and power
supplies. These days our work spaces resemble a forensic lab and
not an herbarium.
A couple of years ago, I attended a conference in St. Charles,
Illinois, sponsored by a world-wide business consulting firm, Ar-
thur Andersen, for both school people and business people. Our
presenters predicted some significant changes to the structure of
our primary and secondary schools across the country by 2050.
A few of those predictions:
* No school building with more than 150 students
* Multi-age classrooms and learning groups
* Teachers as facilitators of learning not dispensers of ‘‘the
truth”
If anecdotes are worth anything, more and more matriculating
freshmen have experienced the positive jolt to learning of well-
guided independent study years before entering our hallowed halls.
We risk losing them to more stimulating alternative experiences
unless we help them take the reins for their learning at the earliest
possible age. Many of the graduates of the Maine Coast Semester
at the Chewonki Foundation, all of whom are attending or have
2000] Hudson—Closing Remarks 359
graduated from places like this (Harvard University), still tell me
years later that their 16 weeks on the coast of Maine surpassed
everything else for an academic experience ... “I learned more,
ete.”
Back at work we are embarking on a new project with the
Smithsonian’s Arctic Studies Center, the Quebec/Labrador Foun-
dation, and the Center for Northern Studies. The ultimate goal is
to put college-aged students in the field with experienced profes-
sionals like yourselves, helping to establish local, community-
based archaeological, historical, ecological, economic, and envi-
ronmental projects in the Great Northern Peninsula of Newfound-
land, along the Labrador Straits, and north, past Nain, to the Torn-
gats.
We are making a deliberate effort to practice E. O. Wilson’s
concilience (Wilson 1998). At the risk of repeating myself, Pll
pitch that idea again this year. You will recall that Wilson la-
mented the lack of interest in “‘the big picture’’ and urged us to
find consilience, literally a ‘“‘jumping together’ of knowledge,
between our many and fragmented disciplines of science and the
humanities. He said:
“A balanced perspective cannot be acquired by studying dis-
ciplines in pieces: the concilience among them must be pur-
sued. Intellectually it rings true, and it gratifies impulses that
arise from the admirable side of human nature. To the extent
that the gaps between the branches of learning can be nar-
rowed, diversity and depth of knowledge will increase. They
will do so because of, not despite, the underlying cohesion
achieved. The enterprise is important for yet another reason:
It gives purpose to intellect. It promises that order, not chaos,
lies beyond the horizon. Inevitably, . . . we will accept the
adventure, go there, and find what we need to know.” (p.
As most of you know, the coast of Labrador looks and feels
like the top of Katahdin or Mt. Washington. Places like this will
“bring ’em in,” believe me. Ill wager that four weeks in Lab-
rador will do more to sink the hook in a larval botanist, archae-
ologist, or even a post-industrial economist than four years hang-
ing around the usual haunts with the usual suspects. But, I di-
gress! Back to the business at hand.
Many of us have more than a mild interest in questions about
360 Rhodora [Vol. 102
plant migrations in a world facing some potentially dramatic
shifts in climate during the coming century. How will our plant
and animal communities respond to predicted increases in annual
mean temperature? What might we expect to see or measure in
our New England mountains, or farther north along the coast of
Labrador? When the Baxter State Park Advisory Committee
asked me a dozen years ago what effect the reintroduction of
caribou might have on the Tableland of Katahdin, I shrugged and
said that, ““Whatever the change, we would never really get the
measure of it.” Beyond a plant list, there was precious little in-
formation in the record on the structure of plant communities on
that summit—no baseline data.
Thought # 3: Take good notes, the world may be changing
fast! If you haven’t established some long-term study plots to
re-visit in your retirement, this summer is not too late. Take great
care to collect good specimens with exquisite notes and details
for the labels.
Thought # 4: Share what you and your students are learn-
ing. Or, put another way, publish in Rhodora!
To that end, it gives me the greatest pleasure to be able to
announce this afternoon the establishment of a special award,
offered by the New England Botanical Club, named for the most
prolific contributor to our journal, the ‘“‘boss”’ himself. The Mer-
ritt Lyndon Fernald Award will be presented each year to the
author or authors of the paper published in Rhodora judged best
to exemplify the goals and objectives of the journal (and the
N ) to promote our knowledge and understanding of the
world of plants.
So, there you have it. Inspire learning. Get yourself and your
students into the field, perhaps with colleagues from neighboring,
or even distant disciplines. Improve your data collection, and es-
tablish some permanent plots for long-term study; the world is
changing fast. Make collections, pay attention to preserving the
record, and publish regularly in Rhodora.
LITERATURE CITED
CrossBy, A. W. 1986. Ecological Imperialism: The Biological Expansion of
Europe, 900-1900. Cambridge Univ. Press, Cambridge, England.
WHEELER, W. M. 1923. The dry-rot of our academic biology. Science 57:
61-71
WILSON, E. O. 1998. Back from chaos. The Atlantic Monthly 281(3): 41-62.
RHODORA, Vol. 102, No. 911, pp. 361-364, 2000
NOTE
CALLERY PEAR (PYRUS CALLERYANA—ROSACEAE)
NATURALIZED IN NORTH CAROLINA
Guy L. NESOM
Biota of North America Program—North Carolina Botanical Garden,
Coker Hall, CB 3280, University of North Carolina
Chapel Hill, NC 27599-3280
e-mail: guynesom @intrex.net
The Callery pear is noted by Kartesz (1999) to occur in North
Carolina, based on an undesignated specimen of Pyrus calleryana
Decne. in the NCU herbarium. The specimen referred to by Kartesz
(Larke 1649, cited below) was taken from a tree growing in a
natural habitat in the North Carolina Botanical Garden, but there
is a probability that it was planted there. The current report pro-
vides documentation for the North Carolina Botanical Garden
specimen as well as a clear example of incipient naturalization
of the same species in North Carolina.
VOUCHER SPECIMENS: NORTH CAROLINA: Orange Co., Chapel Hill, north side
of Morgan Creek in North Carolina Botanical Garden, near spring in Hunt’s
garden area, Grid 14, Aspect EK pine—hardwood bottomland, 23 Sep 1990,
Larke 1649 (NCU)—identified on the label as ‘‘Pyrus sp. (F3 hybrid—Brad-
ford x LeConte)’’; Chapel Hill, Heritage Hills subdivision in southeast corner
of county, numerous seedlings and saplings as volunteers in several yards
along Concord Drive, apparently from fruits from a single, large, cultivated
tree (apparently cv. ‘‘Aristocrat’?) at 106 Concord, 31 Oct 1999, Nesom
NC99-10-] (BRIT, NCU, TEX, US, to be distributed).
Each of the vouchers of Nesom NC99-10-] consists of a fruit-
ing branch of the parent tree and a full plant of one of the young,
deeply taprooted progeny. Most of the young plants range 3-12
dm tall at about 1—4 years old; one of them is 2.2 m tall and may
be older. Some of youngest individuals have lobed leaves, a fea-
ture not seen on mature trees. The parent tree, which was prob-
ably planted in the 1970s or early 1980s, judging from its size
and the history of the neighborhood, is about 12 m tall and has
a relatively loose, elongate-oblong crown with sharply upturned
branches, compared to the tighter, nearly globose crowns of the
shorter cultivars currently so commonly planted in urban land-
scapes.
361
362 Rhodora [Vol. 102
Everett (1981) noted that ‘nearly all pears are self-sterile.”’
Zielinski (1965) observed that within the genus, ‘‘Pyrus fauriei
appears to be unique in producing seeded fruits upon self-polli-
nation. All other species studied are self-incompatible.’’ The
abundant fruits on the parent tree in Orange County (NC99-/0-
7) presumably are from flowers outcrossed to a different clone
(see Ackerman and Creech 1966), although it seems remarkable
that there apparently are no other reproductive individuals of P.
calleryana within a radius of 0.4 kilometers of this one. From the
parent tree, 30 fruits ranged 11-27 mm in diameter and produced
an average of 1.6 mature seeds per fruit (range 1—4). Haserodt
and Sydnor (1983, p. 162) noted that ‘“‘fruiting of this cultivar
[cv. “Aristocrat,” as the tree is identified here] appears to be
heavier than for other cultivars.”” None of the progeny of NC99-
10-1 observed here (as seedlings and saplings) have matured
enough to become reproductive. All are growing within about a
40-meter radius of the parent tree. The parent tree also has pro-
duced numerous root sprouts within a 3-meter radius of its base—
these are similar in morphology to the young, independent seed-
lings and saplings.
Russell (1999) also notes that Pyrus calleryana is “beginning
to naturalize locally in North Carolina and the mid-Atlantic
states.” By 1983, it was observed to be naturalized in Maryland
‘on a wide variety of sites around the U.S. Plant Introduction
Station” in Glenn Dale (Santamour and McArdle 1983). Stewart
(1999) has observed ‘tremendous numbers” of young wild trees
of Callery pear in vacant lots along Route 450 in Bowie, Mary-
land, and along roadsides of the Capital Beltway around Wash-
ington, D.C. At one spot in Bowie, he counted ‘‘over one hundred
trees in a stretch of neglected ground about 100 ft. long and 50
ft. wide. They were so thick that in places the individual young
trees grew only a foot or two apart.’’ Stewart notes that ‘‘we seem
to have a new horticultural plague on our hands in Maryland, a
plague of pears.” The sources of these naturalized plants are ur-
ban landscapes in the United States, where ‘‘the tree is now ap-
proaching epidemic proportions’’ because of overplanting (Dirr
1990, p. 680; also see Anonymous 1986).
The Bradford pear was introduced in the eastern United States
through plantings about 1950, although the name was not pub-
lished until later (see Jacobson 1996). In the last 30 years, many
other cultivars of Pyrus calleryana have been selected and widely
2000] Note 363
planted (Dirr 1990; Huxley et al. 1992; Jacobson 1996; Santa-
mour and McArdle 1983)—although these apparently are all
sometimes informally referred to as ‘“‘Bradford,”’ technically they
should be described by other cultivar names or else generally
referred to as ‘“‘Callery pear.’ Trees of this species naturalizing
in North Carolina and other areas of the southeastern United
States apparently represent a range of different cultivars.
The original cultivar of Pyrus calleryana, the ‘‘Bradford pear,”
was developed at the U.S. Plant Introduction Station in Glenn
Dale, Maryland from seeds collected from northern China (Whi-
tehouse et al. 1963). The native range of P. calleryana includes
“11 provinces of eastern China, south of the 37th parallel,”
where it grows in mixed forests on slopes and in swamps (San-
tamour and McArdle 1983). A closely related entity from Korea,
P. fauriei Schneider, has sometimes been treated as a variety
within P. calleryana but is distinct in a number of respects (Zie-
linski 1965). A naturally occurring hybrid between P. calleryana
and P. betulifolia Bunge has been reported from Illinois (Wandell
1997)
ACKNOWLEDGMENTS. Jeff Beam and Bill Burk (University of
North Carolina Couch Botanical Library) and John Kartesz and
Amy Farstad (Biota of North America Program) helped in ac-
quiring literature.
LITERATURE CITED
ACKERMAN, W. L. AND J. L. CREECH. 1966. Long-term observation reveals
self-unfruitful trait and other desirable a aed of the Bradford
pear. Amer. Nurseryman 124(1): 7-8, 51-
ANONYMOUS. 1986. Is asl pear cee in landscapes? Amer. Nurs-
eryman 164(4): 9—
Dire, M. A. 1990. cate of Woody Landscape Plants: Their Identification,
Ornamental Characteristics, Culture, Propagation and Uses, 4th ed. Sti-
Publ. Co., Champaign, IL.
EVERETT, T. H. 1981. The New York Botanical Garden Illustrated as
pedia of pis Vol. 8, Par—Py. Garland Publ., rk.
HaseropT, H. anp T. D. SypNor. 1983. Growth habits of . aa of
Pyrus in ne J. Arboricult. 9: 160—163.
Hux.iey, A., M. GriFFITHS, AND M. Levy, eds. 1992. The New Royal Hor-
ticultural Society Dictionary of Gardening. Vol. 3, L to Q. Stockton
Press, New York.
JAcoBSON, A. L. 1996. North American Landscape Trees. Ten Speed Press,
Berkeley, CA.
364 Rhodora [Vol. 102
Kartesz, J. T. 1999. A Synonymized Checklist and Atlas with Biological
Attributes for the Mogens Flora of the ue States, Canada, and
Greenland, Ist ed. Jn: J. T. Kartesz and C. A. Meacham. Synthesis of
the North pen "Flora, Version 1.0. a Carolina Botanical Gar-
den, Chapel Hill, NC.
Russe__, A. B. 1999, Urban Tree Identification for North Carolina. North
Carolina State University Consumer Horticulture, NC Cooperative
tension Service, Web Site (http://www.ces.ncsu. adu/deptumhon/consanies!
Landscape/Pyrusca.htm
SANTAMOUR, FS. JR. AND A. J. MCCARDLE. 1983. Checklist of cultivars of
‘allery pear (Pyrus calleryana). J. Arboricult. 9: 114—116.
STEWART, B. 1999, The coming plague of pears. Univ. of Maryland Coop-
erative Extension Service Green Industry Web Site (http://
w.agnr.umd.edu/users/ipmnet/5-8art | .htm).
WANDELL, W. N. 1997. Pyrus calleryana x betulifolia tree named ‘Edge-
ood.’ United States ny plant. U.S. Patent and Trademark Office.
c 16, 1997. (10,151) 2
Wie W.E., J. L. pats ‘AND G. A. SEATON. 1963. Pe ornamental
pear a shade tree. Amer. Nurseryman 177(8): 7-8, 56-60.
ZIELINSKI, 0. B. 1965. Taxonomic status of Pyrus Fauriei Sihneider (Rosa-
ceae). es 13: 17-19
RHODORA, Vol. 102, No. 911, pp. 365—372, 2000
NEW ENGLAND NOTE
SNOW ALGAE IN THE NORTHEASTERN USS:.:
PHOTOMICROGRAPHS, OBSERVATIONS, AND
DISTRIBUTION OF CHLOROMONAS SPP.
(CHLOROPHYTA)
BRIAN DUVAL
Department of Microbiology, University of Massachusetts,
herst, MA 01003
Current Address: Commonwealth of Massachusetts,
Dept. of Environmental Protectio
627 Main St., 2nd floor, Worcester, MA 01608
e-mail: brian.duval @state.ma.us
RONALD W. HOHAM
Department of Biology,
Colgate University, Hamilton, NY 13346
e-mail: rhoham @ mail.colgate.edu
Each spring the waters of melting snowpacks revive a unique
consortium of microbes and small invertebrates that thrive within
this cold oligotrophic environment. While bacteria, fungi, protists,
rotifers, tartigrades, and small insects such as Collembola (spring-
tails) all inhabit springtime melting snow (Hoham et al. 1993),
snow algae are generally the most conspicuous inhabitants. Pop-
ulations of snow algae generally manifest as a red, orange, or
green color within several centimeters of the snow surface (Ho-
ham 1980; Kol 1968), and they may exceed 10° cells ml”! of
liquid meltwater (Hoham 1987). In high alpine regions such as
in the western United States and in polar regions, expansive
blooms of snow algae have been documented and studied exten-
sively with regard to their adaptive capacity to withstand extreme
environments (Bidigare et al. 1993; Hoham and Blinn 1979; Ho-
ham and Ling 2000; Thomas 1972; Thomas and Duval 1995).
While most studies from North America report snow algae from
high alpine areas in the Sierra Nevada, Cascade, and the Rocky
mountains, there are relatively few studies that have focused on
snow algae from the northeastern United States and Canada (Du-
val 1993; Dybas 1998; Hoham et al. 1989, 1993).
Snow algae have been described from the White Mountains
365
366 Rhodora [Vol. 102
(New Hampshire), Green Mountains (Vermont), and the Sunday
River ski area and Mt. Katahdin (Maine), as well as the Adiron-
dacks and Tughill Plateau of upstate New York (Duval 1993;
Hoham et al. 1993). Snow algae have also been described from
the Laurentian Mountains, Quebec (Hoham et al. 1989; Jones
1991), and from southern Ontario (Gerrath and Nicholls 1974),
and probably occur in other parts of eastern Canada but have yet
to be discovered. In Massachusetts, their occurrence is spotty,
limited to the higher elevations of the Berkshire Hills and Wa-
chusett Mountain where there are higher yearly snowfalls or man-
made snow (Hoham et al. 1993). There are no reports of snow
algae from southern New England or the middle or southern Ap-
palachian ranges. A reconnaissance to the West Virginia high-
lands found no evidence of snow algae in melting spring snow
(Duval, unpubl. data).
From our studies between 1972-1998, we plotted locations
where snow algae were and were not found in the northeastern
United States. These findings were compared to a snowfall ac-
cumulation map compiled by the Northeast Regional Climate
Center (Cember and Wilks 1993), and the results are shown in
Figure 1. The snow depth lines are for an “‘average winter,” 1.e.,
in the 50th percentile. Our records show that in the northeastern
United States, areas with greater than 200 cm (80 in.) of annual
snowfall are more likely to have snow algae than areas that re-
ceive lesser snowfall amounts. However, contrary to this gener-
alization is a lack of positive snow algal findings from West Vir-
ginia, many parts of the central plateau of New York State, and
wooded areas near the city of Syracuse, all of which receive
greater than 200 cm of snowfall annually. Thus, snow algal dis-
tribution probably involves other factors in addition to snowfall
accumulation such as vegetative habitat, the rate at which snow-
packs melt, or physio-chemical aspects that affect snow on a re-
gional level (Hoham et al. 1989; Jones 1991).
In the northeastern United States, populations of green snow
algae (Figure 2), are typically found in the shaded coniferous fir
and spruce forests at high elevations (Hoham et al. 1989). Here,
to avoid misnaming the species shown in Figure 3, earlier de-
scribed as Scotiella cryophila and later as a resting spore of Chlo-
romonas nivalis, we refer to the spindle-shaped resting spore as
Chloromonas sp.-A. We hope that a later explanation and Latin
2000] New England Note 367
Ly /
52 \\. 4 & =~With Snow Algae
if
RO A Without Snow Algae
Figure 1. Map of snow algal distribution in the northeastern United States
shown with average yearly snowfall accumulation.
description will help clarify the confusion surrounding the name
of this snow alga.
Additionally, we have observed a species of salmon-orange
colored snow alga in open areas from several New England ski
areas (Duval 1993; Hoham et al. 1993). Since we have observed
only a few biflagellate cells that appear to belong to Chloromon-
as, we designated the species as Chloromonas sp.-B until further
observations are made (Hoham et al. 1993). We have found Chlo-
romonas sp.-B to inhabit snow at four disjunct ski areas in New
England and have not observed this alga in natural alpine snow.
This has led us to propose that one of the mechanisms of this
snow alga’s dispersal might involve transport of resting spores on
the bottom of skis (Dybas 1998; Hoham et al. 1993).
Morphologically, the resting stage of Chloromonas sp.-B is
oval to rounded, ranges in size between 10—25 wm in diameter
(Figure 4), has an outer wall about | wm thick (Figure 5), and
resembles resting stages reported for other species of Chloro-
monas (Hoham 1975; Hoham and Mullet 1978; Hoham et al.
368 Rhodora [Vol. 102
igure 2. A green eee of Chloromonas snow algae from Mt.
Washington, New Hampshire. In North America, populations of green snow
algae are found near the snow surface in shaded areas such as in coniferous
forests
Figure 3. Spindle- ene asexual resting spores of Chloromonas sp.-A
yreen snow algae found in the New York Adirondacks and New England,
photographed at 400 x aan using Nomarski interference contrast
optics ( ). These algae are from a population collected from the Berkshire
Hills, Massachusetts.
2000] New England Note 369
Figure 4. The salmon-orange colored snow alga designated as Chloro-
monas sp.-B (Hoham et al. 1993) collected from the ski area at Killington
eak, Vermont. Filamentous fungi are generally in close proximity to, or in
contact with the algae (200 x).
Figure 5. Chloromonas sp.-B snow algal resting spores at 1000 * mag-
nification. Note the differences in cell size, shape, and the | ym thick wall
that envelopes each cell.
370 Rhodora [Vol. 102
1983). Chloromonas sp.-B is generally observed with filamentous
fungi and often appears to be in contact with them. However,
while fungi and snow algae are generally observed together in
snow samples, it is not clear if there is any exchange of nutrients
or other symbiosis-like relationship between the two microbial
types.
The difference in habitat between species of snow algae, i.e.,
the green Chloromonas sp.-A under forest canopy and the orange
pigmented Chloromonas sp.-B found in open areas, has previ-
ously been observed and attributed to variations in light intensity
(Fukushima 1963). Chloromonas sp.-B is orange in color due to
intracellular carotenoids and other pigments that may serve as
photoprotectants toward biologically harmful ultraviolet radiation
(Bidigare et al. 1993; Czygan 1970; Thomas and Duval 1995).
Indeed, the absorption spectra from solvent extractions of this
alga show absorption in the visible wavelength regions typical of
chlorophylls and carotenoids, as well as in the ultraviolet regions
(Duval 1993).
Chloromonas sp.-B is often observed at the snow surface near
shoots of sprouting Cornus sp. (dogwood), Acer pensylvanicum
(striped maple), and Betula alleghaniensis (yellow birch), and
occasionally it is found at a depth of 5—10 cm, coloring the snow
salmon-orange. This species and other snow algae consistently
appear at the end of the snow melt period (April-May) in areas
where deep snow tends to accumulate from year to year. It is
interesting to note that at lower elevations (Tughill Plateau and
Georgetown Hill, New York), snow algae have been found in
ravines where enough snow accumulates to allow for a deeper
snowpack. Chloromonas sp.-B generally appears near the snow
surface in late April and May, but has been collected as late as
July from Tuckerman Ravine, Mt. Washington, New Hampshire.
Algal predators such as rotifers and ciliated protists are often
observed within snow samples and are members of the snowpack
ecosystem that can support several levels of microbial diversity
(Hoham and Duval 2000; Hoham et al. 1993
It is our intention to stimulate the amateur microbial ecologist
through the photographs presented here to investigate melting
snowpacks, which provide an ephemeral microbial ecosystem
within their cold meltwaters.
ACKNOWLEDGMENTS. Our thanks to Dr. Lynn Margulis, Uni-
2000] New England Note 371
versity of Massachusetts, Amherst, for the use of her laboratory
and microscopy equipment, Colgate University for laboratory and
library facilities, and the University of Massachusetts Biological
Science Library.
LITERATURE CITED
BipiGare, R. R., M. C. Kennicutt II, R. IrurrtAGA, H. R. Harvey, R. W.
OHAM, AND S. A. Macko. 1993. A photoprotective function for sec-
ondary carotenoids of snow algae. J. Phycol. 29: 427—434.
CEMBER, R. P. AND D. S. WiLks. 1993. Climatological Atlas of Snowfall and
Snow Depth for the Northeastern United States and Canada. Northeast
Regional Climate Center Research Series, Cornell University. Publ. #
R 93-1, Ma
CAYGAN, FC. 1970, Blutregen und Blutschnee: Stickstoffmangel-Zellen von
Haematococcus pluvialis und Chlamydomonas nivalis. Arch. Mikrobiol.
DuvaL_, B. 1993. Snow algae in northern New England. Rhodora 95: 21-24.
Dyas, C. 1998. Alga rhythms. Adirondack Life 29: 21-24.
cei H. 1963. a ie oo in Japan. J. Yokohama Munic.
yoer (Nat. ocL, |—
eres J. FE AND K. H. HOLLS an A red snow in Ontario caused by
the dinoflagellate, Cunnadaiin pascheri. Canad. J. Bot. 52: 683-685.
Honam, R. W. 1975. The life history and ecology of the snow alga Chloro-
monas pichinchae (Chlorophyta, Volvocales). Phycologia 14: 213-226.
. 1980. Unicellular Chlorophytes-Snow Algae, pp. 61—84. Jn: E. R.
Cox, ed., Phytoflagellates. Elsevier/North Holland, New York.
. 1987. Snow algae from high-elevation, temperate latitudes and semi-
permanent snow: Their interaction with the environment, pp. 73-80. In:
J. Lewis, ed., Proc. 44th Annu. Eastern Snow Conf., Fredericton, NB,
Canada.
AND D. W. BLINN. 1979. Distribution of cryophilic algae in an arid
eo the American Southwest. Phycologia 18: 133-145.
Db B. DuvaL. 2000. Microbial ecology of snow and fresh-water ice
with Pactad: on snow algae, pp. 166—266. In: H. G. Jones, J. W. Pom
eroy, D. A. Walker, and R. W. Hoham, eds., Snow Ecology: An infer
disciplinary Examination hg Snow-covered Ecosystems. Cambridge
Univ. Press, ee ae
, A. E. Laursen, S. O. ee AND B. DuvAL. 1993. Snow algae and
other eee in several alpine areas in New England, pp. 165-173. In:
M. Ferrick and T. Pangburn, eds., Proc. 50th Annual Eastern Snow
ee ee See Canada.
H. U. Linc. 2000. Snow algae: The effects of chemical and
ee factors on res life cycles and populations. Jn: J. Seckbach,
ed. ae robial Diversity. Klewer Press, The Netherlands (in press).
OT E. MULLET. 1978. Chloromonas nivalis (Chod.) Hoh. & Mull.
ae nov., and additional comments on the snow alga, Scotiella. Phy-
cologia 17: 106—107.
372 Rhodora [Vol. 102
——., AND S. C. Roemer. 1983. The life history and ecology of
fhe snow alga Chloromonas polyptera comb. nov. (Chlorophyta, Vol-
vocales). Canad. J. Bot. 61: 2416-2429,
YATSKO, AND H. G. JONES. 1989. Recent discoveries of snow
algae in Upstate New York and Quebec cages and Lee agra
on related snow chemistry, pp. 196-200. In: J. Lewis, ed., Proc. 46th
Annual Eastern Snow Conf., Quebec City, c a.
Jones, H. G. 1991. Snow chemistry and biological activity: A particular per-
spective on nutrient cycling, pp. 173-228. In: T. D. Davies, M. Tranter,
nd ones, eds., NATO ASI Series G: Ecol. Sci., Vol. 28, Seasonal
Snowpacks, Processes of Compositional Changes. Springer-Verlag, Ber-
lin
KOL, E. 1968. Kryobiologie. Biologie und Limnologie des Schnees und Eises.
tape cea pp. 1-216. In: H. J. Elster and W. Ohle, eds., Die
nnengewasser. E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart.
Paeuac. W. H. 1972. Observations on snow algae in California. J. Phycol.
8: 1-9.
ND B. Duvac. 1995. Sierra Nevada, California, U.S.A., snow algae:
au albedo changes, algal-bacterial eae ame and ultraviolet
radiation effects. Arctic Alpine Res. 27: 389-399,
RHODORA, Vol. 102, No. 911, pp. 373-379, 2000
NEBC MEETING NEWS
February 2000. Michele Dionne, an aquatic ecologist and Re-
search Director at the Wells National Estuarine Research Reserve
in Wells, Maine, spoke on the topic: “‘Is the tide turning for salt-
marsh ecology and restoration in the Gulf of Maine?’’ The pre-
sentation centered on ecological research and habitat restoration
efforts at the Wells Reserve, one of 25 federally designated coast-
al research reserves, and a few other salt-marsh locations in the
Gulf of Maine. Describing salt-marshes as ‘“‘New England’s na-
tive grasslands,’ Dr. Dionne highlighted some of their functions
and values. Ecologically, they contribute to shoreline anchoring,
storm surge buffering, water quality, and habitat for wildlife, fish,
and shellfish. For our human society, she noted that there are
recreational, commercial, aesthetic, educational, and _ historical
values. In the Gulf of Maine salt-marshes, where mean tide ranges
are typically 8—10 ft., there is a close relationship between ele-
vation and vegetational zones, as one might expect. Here one
finds Spartina patens, salt-marsh hay, dominating high marshes
and the taller S. alterniflora, which has the capability of exuding
salt from specialized cells on its leaf blades, in the low marshes.
She described the niches of some of the other plants of the salt-
marshes as well, including Phragmites australis and Typha an-
gustifolia, which occupy the high edges of the salt-marshes and
often take over when the hydrologic regime is altered. Dionne
described several types of salt-marshes in Maine: (1) back barrier
marshes, the typical coastal marsh; (2) fringing marsh, the narrow
bands of salt-marsh lining miles of major rivers like the Kennebec
and Penobscot, and (3) finger marshes, those found in ‘‘drowned
valleys”’ associated with coves and bays.
The oldest salt-marshes in New England are reported to be
about 5000 years old. Despite changing sea-levels during this
time, they have persisted through accretion of new peat that
builds up in response to new sediments brought in by the tides.
Dionne emphasized that, conversely, natural events or human ac-
tivities, such as the construction of roads, beach barriers, jetties,
or stream control gates, can interfere with this accretion, causing
rapid erosion of salt-marsh habitat. Tidal restriction can lead to
such events as subsidence from oxidation of peat, restricted fish
passage, less exchange of nutrients and organic matter, water
freshening, encroachment by invasive species, and incremental
af2
374 Rhodora [Vol. 102
development. The Little River at the Wells Reserve is one of the
few rivers and salt-marsh systems that is minimally impacted in
the above ways. The Drakes Island marsh system on the southern
edge of the Wells Reserve, on the other hand, has been impacted
by a number of things including a road built on a berm across
the marsh’s north end. A water control gate was installed where
the Webhannet River went under the road, thus preventing spring
tides from entering the marsh from the estuarine lagoon to the
east. The gate fell off this structure in 1988, partially restoring
the tidal influence. In 1991, scientists began monitoring the
changes to the upper marsh (above the gate) where three feet of
elevation had been lost from years of abuse. Salt-tolerant plants
and marine fish quickly returned to the area, and by 1998, soils
were stabilized and a low marsh vegetation dominated by Spar-
tina alterniflora was well established in the area. In 1999, S.
patens was observed colonizing upper edges despite the marsh
still being lower than normal. A return to a high marsh is pre-
sumed to be many years away yet. In contrast, researchers found
a much more rapid restoration of Mill Brook Marsh, located near
the mouth of the Squamscott River which flows into Great Bay
in Stratham, New Hampshire. It had a somewhat similar history
of road impact and gating in 1970 followed by flow restoration
in fall 1993. Before restoration, purple loosestrife had become a
dominant species in the upstream marsh. Five years after removal
of the tide gate and installation of a large culvert, the purple
loosestrife was gone and a salt-marsh with three taxa of Spartina
had been restored.
‘“Have we stemmed the tide?”’ Dionne asked rhetorically at the
end. Her answer is that we are returning the tide to historical salt-
marsh communities in many cases but that many obstacles still
exist to maintaining their normal function. There is much concern
about development along beaches and upland edges, for instance.
Also, there are concerns about predation by the introduced green
crab on soft shell clams, an important indigenous species in the
marsh ecosystem. It appears that the Wells Reserve is key to
research and education on these issues for Gulf of Maine towns
where salt-marsh restoration is needed and maintenance is for-
ever.
—PAuL SOMERS, Recording Secretary.
2000] NEBC Meeting News S75
March 2000. The evening’s speaker was outgoing President
David Conant, whose talk was entitled ‘““The Biology of Tree
Ferns.”’ David traced his love of the ferns and field biology to
an afternoon foray to New Hampshire’s Bear Mountain with men-
tor Albion Hodgdon. Assaulted by yellow jackets, Hodgdon tum-
bled one hundred feet, head over heels, down a rocky slope and
into the crook of a tree. Fearing the worst, David hurried to Al-
bion’s side. Notwithstanding the bites of dozens of yellow jackets,
Albion rose to his feet to continue the plant hunt down the moun-
tain. During refreshments at the bottom, David thought, *‘This is
all right!”’ He happily cast his lot with Albion Hodgdon. A couple
of false starts with the ferns of New Hampshire and flora of
Sullivan County preceded David’s introduction to the tree ferns,
and he has been a student of the group since the mid-1970s.
David traced, through the 1980s, the emergence of the use of
tools like electrophoresis and analysis of chloroplast DNAs in
analyzing relationships among the ferns. Notably, the work of
Japanese botanist Haseke has confirmed many of the assumptions
of our narrative phylogenies for the ferns with his analysis of the
gene for the enzyme (rbcL) that plucks CO, from the air to build
glucose in the dark reactions of photosynthesis. He confirmed the
ancient lineage of primitive ferns like the Osmundaceae, and sort-
ed out the higher leptosporangiate ferns, just as do the narrative
phylogenies. As an aside, David said, if we are to recognize many
orders of the “‘younger”’ flowering plants, this modern work with
the ferns underscores abandonment of a single order, Filicales,
for all the ferns. For his part, David took his work with the tree
fern genus Alsophila, begun with Rolla Tryon in 1976, into the
modern laboratory. The days of plant collecting with the aid of
a converted mail van were followed by collaborations in bio-
chemistry with Gillian Cooper-Driver of Boston University and
Gus Dimaggio of Dartmouth College. Analyses of flavonoid pig-
ments and storage proteins were helpful, but not absolutely con-
clusive in sorting out the tree ferns. Together with Diana Stein
of Mount Holyoke College, David moved next to analysis of
chloroplast DNAs. David jetted all over the New and Old World
tropics to collect the ferns, shipping them back to Diana within
two days for grinding. Countless southern blots later, David re-
counted the horrible experience of trying to make sense of it all,
‘like trying to put Humpty Dumpty back together again.”” The
two scientists struggled with a number of molecular probes of the
376 Rhodora [Vol. 102
collected chloroplast DNAs, settling on ones derived from Christ-
mas ferns to retrieve the clearest set of data.
After years of work, David and Diana produced a fresh picture
of the tree ferns as three major groups centered on Alsophila,
Cyathea, and Sphaeropteris. The Cyathea clade is not restricted
to the New World tropics as previously believed, but linked
through geological time to ferns found in Western Queensland
and the Pacific. It appears that the Greater Antilles group of Al-
sophila is the most derived of the tree ferns.
David ended his presentation with striking images of hand-
prepared sections of fern stems, produced with the help of his
students at Lyndon State College. David declared that there is ‘‘a
lot to learn beyond who they (sic the ferns) are!’’—a refreshing
perspective on teaching and learning, indeed.
April 2000. Dr. Paul Godfrey of the University of Massachu-
setts at Amherst spoke on “Biodiversity of Medicinal Plants in
Northwestern Thailand.’’ Dr. Godfrey has spent the large part of
his career investigating aspects of coastal ecology along the At-
lantic seaboard. Richard Evans Shultes, who once surprised an
NEBC audience by firing a blowgun dart across a crowded hall,
inspired Paul’s interest in ethnobotany. At this juncture of his
introduction, Paul reached for a small bamboo bow and fired its
bamboo arrow across the crowded hall. He had our attention.
Paul was asked some years ago by Linda A. Swift of Hartwick
College to lend his ecological expertise to an ethnobotanical in-
vestigation of plant utilization by an Akha hill tribe village of
northwestern Thailand. Thai hill tribes have long used small-scale
swidden and crop rotation for maize and rice production. Such
swidden-based tropical agriculture is often linked with the loss
of biodiversity, though it is critical to the survival of the hill
tribes. The long-term studies of forest utilization and plant use
around Pakhasukjai Village were designed to measure the impacts
of wood gathering, agriculture, medicinal, and spiritual activities
on diversity. Drs. Godfrey and Swift expected to find the lowest
diversity in disturbed forests close to the village and the highest
in undisturbed forests further away. They selected a group of
native gingers (Zingiberaceae) to investigate, gathering data on
the abundance and distribution of species within three discreet
study areas near Pakhasukjai Village in order to evaluate modified
importance values and biodiversity indices.
2000] NEBC Meeting News owe
Paul was pleased to find J. EF Maxwell at Chaing Mai Univer-
sity during his first season in northern Thailand. As it turns out,
Dr. Maxwell has contributed the largest part of northwest Thai
plant specimens to the Harvard University Herbaria during the
past several decades. Dr. Maxwell offered his considerable ex-
pertise by acquainting Paul and his colleagues with the fine points
of collection and identification of the gingers in situ. The eth-
nobotanical team was supported at Pakhasukjai Village by the
Hill Area Development Foundation, which provided space and
facilities in its rustic center.
The Akha of Pakhasukjai Village are forest dwellers who are
spiritually bound to their surroundings. They find both good and
bad spirits in the forest. The forests provide both wood and med-
icines. Paul and his colleagues had great difficulty in learning the
uses of medicinal plants, in particular. Two layers of interpretation
are needed to get from the native language to Thai, then to En-
glish-mediated interviews. Add to the linguistic hurdles the cer-
tain possibility of deliberate deception on the part of the shaman,
and the team had its work cut out. Ten species of ginger in 7
genera were of particular interest. The list included Alpinia gal-
anga (for stomach ache and diarrhea); Amomum repoense (for
multiple medicinal uses from appetite stimulant to pain reliever);
Boesenbergia rotunda (similar medicinal uses as Amomum); Cos-
tus speciosus (stimulant, aphrodisiac, or for relief of back pain);
Curcuma longa (leaf poultice for cuts and bites); Kaempferia par-
viflora (poultice to stop bleeding cuts); and three species of Zin-
giber (for headache, stomach ache, diarrhea, or a stimulant for
breast milk production).
Paul devised an ecological sampling plan for three forest sites,
each of which had been substantially cleared nearly 50 years ear-
lier. On the first day out, the shaman lead the team by a tortuous
and turning path to a very sacred site, the Cemetery Forest. It
turned out to be the closest to the village. Diversity of gingers
was relatively low here, and greatest within a more heavily ex-
ploited site dubbed the Shrine Forest. Several of the gingers ap-
pear to respond to changes on the landscape as early successional
types. The light and regular disturbance by the Akha may con-
tribute to this increase in overall diversity.
Paul ended his presentation with a personal testimonial. He had
learned that a local Eupatorium was effective for stanching blood
378 Rhodora [Vol. 102
flow. One errant blow of the machete and Paul was able to put
the plant to the test. The wound healed nearly overnight.
—Don Hupson, Recording Secretary.
May 2000. Vice President Paul Somers introduced the even-
ing’s speaker, Dr. Avril de la Cretaz, of the Department of Natural
Resources Conservation at UMass Amherst. She spoke on her
doctoral thesis research topic, ““Understory Restoration in a Wa-
tershed Degraded by Deer Browsing and Fern Invasion,”’ and
won the Club’s award for ‘“‘best performance under adverse cir-
cumstances” by giving an outstanding summary of her research
despite a balky slide projector.
The Quabbin Reservoir is a 120,000-acre tract of land and
water of which 64% is administered by the Metropolitan District
Commission (MDC) to protect the Metropolitan Boston water
supply. Many of the upland areas of the Quabbin, farmland before
being incorporated in the watershed protection area, were planted
in white or red pine plantations to protect water quality. Deer
hunting was banned in the Quabbin from 1940 to 1991 (originally
because of fear of sabotage during WWII), resulting in a deer
population of 40—60 deer per square mile. This resulted in inten-
sive browsing on understory vegetation, essentially eliminating
tree seedling regeneration from large areas of the Quabbin wa-
tershed. The MDC is now interested in restoring a natural forested
community to the Quabbin. However, large areas of the water-
shed, including many of the pine plantations, have a dense mono-
culture of hay-scented fern (Dennstaedtia punctilobula) with es-
sentially no tree regeneration, even after the deer population has
been substantially reduced. Dr. de la Cretaz investigated the
mechanism by which hay-scented fern influences tree regenera-
tion and mechanical means of control that may allow forest man-
agers to restore the forest. Hay-scented fern, although a native,
behaves like many exotic invasives in the landscape by creating
mono-dominant stands and altering the natural community diver-
sity and dynamics. Although some studies alleged that hay-scent-
ed fern dominance was because of allelopathy, more recent work
has shown that these ferns are not necessarily allelopathic. The
fern’s effect on other species seems to be because of competition
for resources, particularly space in the thick root mat and light
that is blocked by the fern fronds.
2000] NEBC Meeting News a7
Avril also compared the ability of different tree species to be-
come established in hay-scented fern communities. She found that
only white pine and black birch were capable of developing into
saplings in a dense fern stand, because the leaves of these species
develop and expand before the fern fronds expand in the spring
and are therefore able to compete for light. White ash and oak
seedling leaves expand after the fern and thus are not as com-
petitive.
In the final stage of her research, Dr. de la Cretaz compared
three mechanical treatments to control fern growth and promote
tree regeneration: root mat removal (‘‘scalping”’), mixing root mat
and mineral soil (‘‘scarification’’), and clipping (mowing). Her-
bicide treatments are not allowed in the watershed forest. She
found that scarification actually increased the growth and domi-
nance of hay-scented fern. Root mat removal resulted in the great-
est germination response of tree seedlings, but also graminoid
dominance (sedges, especially Carex debilis, established in high
densities from the soil seed bank in the first two treatments).
Clipping, particularly if done repeatedly during the growing sea-
son, resulted in the highest tree seed germination and seedling
growth and the lowest fern and graminoid dominance. She hy-
pothesized that clipping is the most effective treatment for tree
seedlings because there are fewer graminoid competitors and a
higher nutrient availability. The root mat of the ferns may inter-
fere with seed germination, but this effect is outweighed by the
increased light available without the dense fern frond canopy.
In summary, Avril’s studies showed that the lack of tree re-
generation is a result of deer browsing following overstory thin-
ning. Browsing eliminates tree seedlings and depletes the seed-
bank, while increased understory light accelerates the growth
rates of existing fern colonies and increases spore production.
This results in the dominance of hay-scented fern: the fern’s root
mat inhibits germination of any remaining seeds, and the fronds
block the light and inhibit growth of any seeds that do germinate.
In dense fern stands, trees will not regenerate without interven-
tion, and mowing is a promising mechanical treatment that may
be effective for understory restoration.
—LISA STANDLEY, Recording Secretary pro tempore.
ANNOUNCEMENT
NEW ENGLAND BOTANICAL CLUB
GRADUATE STUDENT RESEARCH AWARD
The New England Botanical Club will offer $2,000 in support
of botanical research to be conducted by graduate students in
2001. This award is made annually to stimulate and encourage
botanical research on the New England flora, and to make pos-
sible visits to the New England region by those who would not
otherwise be able to do so. It is anticipated that two awards will
be given, although the actual number and amount of awards will
depend on the proposals received.
The award will be given to the graduate student submitting the
best research proposal dealing with systematic botany, biosyste-
matics, plant ecology, or plant conservation biology. Papers based
on the research funded must acknowledge the NEBC’s support.
Submission of manuscripts to the Club’s journal, Rhodora, is
strongly encouraged.
Applicants must submit three copies of each of the following:
a proposal of no more than three double-spaced pages, a budget,
and a curriculum vitae. Two letters in support of the proposed
research, one from the student’s thesis advisor, should be sent
directly to the Awards Committee by sponsors. All materials
should be sent to: Awards Committee, The New England Botan-
ical Club, 22 Divinity Avenue, Cambridge, MA 02138-2020.
Proposals and supporting letters must be received no later than
March 1, 2001. The recipient(s) will be notified by April 30,
2001.
This year the Graduate Awards Committee is pleased to an-
nounce two recipients of the Graduate Student Research Awards.
Dirk Albach of the Universitat Wien (University of Vienna, Aus-
tria) received support for his proposal entitled ‘Evolution, bio-
geography, and genetic diversity in Veronica alpina L. and re-
lated taxa.” Also chosen for an award was Michael Booth of Yale
University, for his proposal entitled ‘“‘Material flows across ec-
tomycorrhizal networks and plant diversity in New England for-
ests.”’ For more details on these research proposals and a listing
of the awards from 1985 to the present, consult the Club’s web
page (http://www.herbaria.harvard.edu/nebc/).
380
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381
382 INFORMATION FOR CONTRIBUTORS
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The New England Botanical Club is a nonprofit organization
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THE NEW ENGLAND BOTANICAL CLUB
Elected Officers and Council Members for 2000—2001:
President: Lisa A. Standley, Vanasse Hangen Brustlin, Inc., 101
Walnut St., P O. Box 9151, Watertown, MA 02272
Vice-President (and Program Chair): Paul Somers, Massachusetts
Natural Heritage and Endangered Species Program, | Rabbit
Hill Rd., Rt. 135, Westborough, MA 01581
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, Massachusetts College of Liberal Arts, North Adams,
MA 01247-4100
Recording Secretary: W. Donald Hudson, Jr.
Curator of Vascular Plants: Raymond Angelo
Assistant Curator of Vascular Plants: Pamela B. Weatherbee
Curator of Nonvascular Plants: Anna M. Reid
Librarian. Leslie J. Mehrhoff
Councillors: David S. Conant (Past President)
Karen B. Searcy 2001
David Lovejoy 2002
Arthur V. Gilman 2003
Jennifer Forman (Graduate Student Member) 2001
Appointed Councillors:
David E. Boufford, Associate Curator
Janet R. Sullivan, Editor-in-Chief, Rhodora
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RHODORA
Journal of the
New England Botanical Club
CONTENTS
A new species, a new combination, and new synonymy for South American
Jaltomata (Solanaceae). Thomas Mione, Segundo Leiva G., Neil R.
Smith, and Scott J. Hevner 385
Birds, pollination reliability, and green flowers in an endemic island shrub,
Pavonia bahamensis (Malvaceae). Beverly J. Rathcke ................ 392
Nomenclatural proposals in Atriplex (Chenopodiaceae). Stanley L. Welsh .... 415
Systematic notes on the Old World fern genus Oleandra. Rolla Tryon...... 428
Floristic inventory of the Waccasassa Bay State Preserve, Levy County, Florida.
J. Richard Abbott and Walter S. Judd 439
NEW ENGLAND NOTE
First records of a European moss, Pseudoscleropodium purum, naturalized
n New England. Norton G. Miller $14
NOTE
Low catchment area lakes: New records for rare coastal plain shrubs and
Utricularia species in Nova Scotia. Nicholas M. Hill, J. Sherman
Boates, and Mark F- Elderkin S18
BOOK REVIEW
Aquatic and Wetland Plants of Northeastern North America. Volume |.
Pteridophytes, Gymnosperms, and Angiosperms: Dicotyledons;
Volume 2. Angiosperms: Monocotyledons 523
NEW BOOKS 527
NEBC MEETING NEWS 529
Reviewers of Manuscripts 532
Information for Contributors 233
NEBC Membership Form 535
Index to Volume 102 536
NEBC Officers and Council Members inside back cover
Vol. 102 Autumn, 2000 No. 912
Issued: January 19, 2001
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,
rham, NH 03824
e-mail: janets@cisunix.unh.edu
ANTOINETTE P. HARTGERINK, Managing Editor
Department of Plant Biology, University of New Hampshire,
Durham, NH 03824
e-mail: aph2@cisunix.unh.edu
Associate Editors
HAROLD G. BROTZMAN STEVEN R. HILL
DAVID 8S, CONANT THOMAS D. LEE
GARRETT E. CROW THOMAS MIONE
K. N. GANDHI—Latin diagnoses and nomenclature
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DORA is available at http://www.herbaria.harvard.edu/nebc/
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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).
RHODORA, Vol. 102, No. 912, pp. 385-391, 2000
A NEW SPECIES, A NEW COMBINATION, AND
NEW SYNONYMY FOR SOUTH AMERICAN
JALTOMATA (SOLANACEAE)
THOMAS MIONE
Biological Sciences, Central Connecticut State University,
New Britain, CT 06050-4010
e-mail: MioneT@CCSU.edu
SEGUNDO LEIVA G.
Museo de Historia Natural, Universidad Antenor Orrego,
Avenida America Sur 3145, Trujillo, Peru
NEIL R. SMITH AND ScoTT J. HEVNER
Biological Sciences, Central Connecticut State University,
New Britain, CT 06050-4010
ABSTRACT. Jaltomata hunzikeri, a rare shrub of the coast of the department
of Lima, Peru, is described and shown in a photograph. Hebecladus sinuosus,
transferred as J. sinuosa, is a shrub that is widely distributed in the Andes.
Saracha lobata and S. sordideviolacea are placed in synonymy with J. dentata.
Key Words: edible fruit, Hebecladus, Jaltomata, Saracha, Solanaceae
In the process of taxonomic revision of the genus Jaltomata
we have found it necessary to describe a new species, make a
new combination, and place two binomials in synonymy with
another.
Jaltomata hunzikeri Mione, sp. nov. Type: PERU. Dept. Lima:
Prov. Barranca, 5 km north of Barranca, lomas of Pativilca,
300 m, sandy hillside, 18 Sep 1938, Stork, Horton, and Var-
gas C. 9228 (HOLOTYPE: GH; ISOTYPE: G, K, MO). Figure 1.
Planta fruticosa ad | m altitudine; axes juvenes, petioli, pe-
dunculi, pedicelli, facies abaxialis calycis villosa, pilis uniseriali-
bus, non ramosis, erectis, apice glandiferentibus; inflorescentia
floribus 10 ut maximum; corolla breviter tubulosa, limbo 16—17
mm diametro, quinque lobis triangularibus, alba, annulo azureo
prope extremum tubi; stamina 4.8-7 mm longitudine, filamenta
villosa secus proximales 45—60 partes per centum longitudinis;
stylus 6.0—7.7 mm longitudine.
385
Rhodorz
[Vol. 102
J Jaltomat
UNIVERSITY OF CALIFORNIA
Becond Botante
Pxxv, 1
No 9228
THE O10 STATE UNIVERSITY
& procumbens (Cav. )Gentry
n Davis I\ ?
1 Garden Expedition to the Andos,
hotavia, Cure, Unvaway, A x
1988-36
ROENTINA
| Director, T. 11. Goopars
ENR OL,
Peru. Depto. Lima, Prov. Cr an of
Pativiloa, Sion north of Be
June 1, 1978 hilleide,
Vary de » rounded bush to ] soro \ ty
whitish with purple throat,
Sept. 18, 19.
Figure |.
Jaltomata hunzikeri Mione, in
gas 9228, (HOLOTYPE: GH). Photo by S. J. H.
flower, Stork, Horton, and Var-
2000] Mione et al.—South American Jaltomata 387
Shrub to 1 m high. Young axes, petioles, peduncles, pedicels,
and abaxial face of calyx villous, the hairs uniseriate, unbranched
(finger-type), erect and gland-tipped. Young axes with raised lon-
gitudinal ridges (an artifact of drying?). Older axes to 1.5 cm in
diameter, terete and glabrate. Leaves alternate, often geminate,
the blades ovate, to 8 X 5.5 cm, with 3—4 pairs of primary veins,
the apex acute, the base somewhat truncate and often oblique, the
younger blades densely pubescent, the older sparsely pubescent,
the margin dentate or erose-dentate or repand and ciliate with
gland-tipped hairs 0.12—0.42 mm long; petioles to 3 cm long.
Inflorescences axillary or sometimes arising from branch dichot-
omies, umbellate, to 10-flowered including buds. Peduncle 4—9
mm long; pedicel 8.6-11.3 mm long. Calyx green at anthesis,
stellate, the lobes triangular and 4.0-5.9 mm from pedicel to tip,
1.9-3.2 mm from pedicel to sinus, the margin ciliate with finger
hairs 0.3—0.6 mm long, abaxially with both finger hairs 0.3—0.8
mm long and glands 55-70 wm long having multicellular heads
and unicellular stalks (illustrated in Mione and Serazo 1999); ca-
lyx 10 mm in diameter with fruit (mature?). Corolla short-tubular
(the tube not evident after pressing, but mentioned by collectors
on label), the limb crateriform or broadly infundibular or rotate,
16-17 mm in diameter, white with blue ring near end of tube,
with 5 triangular lobes, 6.2—11.0 mm from flower center to tip
of corolla lobe, 4.0—7.3 mm from center to sinus, the margin
ciliate with finger hairs 0.1—-0.5 mm long. Stamens 4.8—7.0 mm
long, the filaments villous on proximal 45—60% of the length, the
finger hairs 1.0-1.5 mm long; anthers 1.3-1.5 x 0.7-0.9 mm,
some basally sagittate. Pollen grains (stained 30 minutes in “cot-
ton blue’’) 26.25—31.25 jm in diameter (average 28.5 wm, n =
24). Style and ovary glabrous. Style 6.0—7.7 mm long, 0.1—0.2
mm wide at midlength; stigma capitate, not bilobed, 0.24—0.6 mm
wide perpendicular to style, exserted 0-1 mm beyond dehisced
anthers. Berry (mature?) 5 mm across, and very likely subspher-
ical and orange or red at maturity.
PARATYPE: Peru. Dept. LIMA: Prov. Barranca, 5 km north of Barranca, talus
slope of hill rising abruptly from low, narrow, coastal plain, 80 m, 5 Sep
1938, Morrison and Beetle 9099 (GH).
The specimens (the type and paratype) of this species were
treated as Saracha villosa (Zuccagni) G. Don by Macbride
(1962). We do not agree, based on study of: 1) photos of the type
388 Rhodora [Vol. 102
of S. villosa (G-DCc, F neg. 6880, NY), 2) description of the hairs
of the type of S. villosa (provided by G), and 3) the translation
to English (by N. R. S.) of the Latin description within the pro-
tologue of Atropa villosa Zuccagni, basionym of S. villosa.
Jaltomata hunzikeri is similar to J. cajacayensis S. Leiva &
Mione and J. propingqua (Miers) Mione & M. Nee, of the de-
partments of Ancash and Lima, Peru, respectively; all three
shrubs bear gland-tipped hairs and have a short-tubular corolla
with a much broader limb. Ja/tomata hunzikeri lacks corolla lob-
ules, the stigma is at approximately the same height as the de-
hisced anthers, and grows at 80-300 m in the fog-dependent,
coastal lomas habitat. The other two species have corolla lobules
alternating with the larger lobes, have stigmas exserted several
mm beyond the anthers, and grow above 1,800 m (Mione et al.
2000).
The specific epithet was chosen to honor Armando T. Hunziker,
eminent Solanaceae taxonomist.
Jaltomata sinuosa (Miers) Mione, comb. nov.
oe STNUOSUS ae sen Bot. 7: 352. 1848. Miers, Il.
Pl YPE: PERU. Dept. Amazonas: a.
poyas, es WS S.A. Liste BM; ISOTYPE: G two sheets, K). Sar-
acha sinuosa (Miers) Bitter, Repert. Spec. Nov. Regni Veg. 18:
101. 1922.
Saracha a Miers, Ann. Mag. Nat. Hist., ser. 2, 3: 450. 1849.
‘OLOMBIA. La Pena: Bogota, Goustot §.. (HOLOTYPE: P, F neg.
30250: ISOTYPE: c Witheringia glandulosa (Miers) Miers, Bk
za Nat. Hist., ser. 2, 11: 92. 1853. Miers, Ill. S. Amer. Pl. 2: 20.
857, t. 39a. mae glandulosa (Miers) Castillo & R. E. Schult.,
ae 88: 292. 1986.
Saracha vestita Miers, Ann. Mag. Nat. Hist., ser. 2, 3: 449. 1849. Type:
ECUADOR. ““Minasurcu prope Quito” on types (“‘Minashuaicu”’ is
the Defense Mapping Agency [1987] spelling), Hartweg 1292 (Ho-
LOTYPE: K; ISOTYPE: LD). Witheringia A is (Miers) Miers, Ann.
Mag. Nat. Se ser. 2, 11: 92. 185 Sem vestita (Miers)
Castillo & R. E. Schult., Rhodora 88: > 986.
eas whalenii . Knapp, T. Mione & ne Brittonia 43: 181.
91. Type: PERU, Dept. Ce Prov. Contumaza, lecho de Rio
ae Benito, alrededores de San Benito, 1300 m, Sagdstegui, Leiva,
and Sagadstegui 1247] (HOLOTYPE: HUT; ISOTYPES: IBE, MO, NY
Jaltomata sinuosa 1s superficially similar to J. sanctae-martae
(Bitter) Benitez of Colombia and Venezuela. Both species are
shrubs, are villous with gland-tipped finger hairs, and bear rotate
corollas. Jaltomata sinuosa has 3—5 flowers per inflorescence,
2000] Mione et al.—South American Jaltomata 389
flowers 2.5—3.8 cm in diameter, and orange berries while J. sanc-
tae-martae has up to 10 flowers per inflorescence, flowers to 1.8
cm in diameter, and according to Benitez de Rojas (1980), red
berries.
DISTRIBUTION, HABITAT, USES, LOCAL NAMES. Jaltomata sinuosa Oc-
curs in disturbed habitats in the Andes from western Venezuela
to Bolivia. The fruits are eaten (Dillon et al. 6193; Leiva et al.
2042: Mione et al. 672) and the local names include ‘*‘tomatillo”’
(Hawkes and Garcia-Barriga 100) and “‘uvilla de monte” (Mione
and McQueen 468, 469).
REPRESENTATIVE SPECIMENS: Venezuela. MERIDA: Vicinity of E] Royal, near
La Toma, 2440 m, 4 Nov 1978, Luteyn et al. 618] (Mo, NY).
Colombia. CUNDINAMARCA: Cordillera Oriental, Monserrate, Valle del Rio
San Francisco, 2700—2900 m, 18 Jun 1948, Hawkes and Garcia-Barriga 100
(K, US); Cordillera es western slopes, 20 km from Bogota, via Salto de
Tequendama-El Colegio road, 2470 m, 13 Jan 1976, Luteyn et al. 4817 (kK,
MO, NY).
Ecuador. IMBABURA: on road from Otavalo to Selva Alegre, 29.4 km from
junction of Panamerican Hwy, 2900 m, 28 May 1991, Spooner et al. 5113
(CONN). PICHINCHA: canton Quito, Parroquia Nanegalito, quebrada Santa Rosa,
steep slopes SW of Rio Pichan, 2000 m, 12 Jan 1995, Webster and Rhode
31234 (DAV). TUNGURAHUA: vicinity of Ambato, Feb 1919, Pachano 138 (GH,
NY, US). CHIMBORAZO: highway to Pallatanga from just S of Cajabamba, 32.1
km in from Panamerican Hwy, 3000 m, 10 Jan 1990, Mione and McQueen
468, 469 (CONN, NY). CANAR: outskirts of Asorgues, 2897 m, 27 Jun 1939,
Balls B7327 (kK, US). LOJA: road to La Toma on slopes of ae Villonaco,
ca. 10 km W of Loja, 2440 m, 7 Mar 1965, Knight 583 (w
Peru. AMAZONAS: 2 kms sions road W of Chachapoyas, "2195 m, 13 Jan
1983, King and Bishop 9179 (G, K, MO, US); Mendoza, 1400-1500 m, 19 Aug
1963, Woytkowski 8153 (MO, NY). PIURA: Huancabamba, alrededores de Sa-
palache, 2400 m, 9 Jun 1997, Leiva et al. 2042 (CONN, HAO); Ayabaca, al-
rededor de Yacupampa (Ayabaca—Cuyas), 2702 m, 21 Sep 1996, Leiva et al.
1867 (CONN, HAO). CAJAMARCA: Cutervo, garden in village of San Andrés de
Cutervo, 2050 m, 6 Nov 1990, Dillon et al. 6193 (F); Chota, 6° 33’ 54” S,
78° 38’ 42" W, 2300 m, 19 Jun 1999, Leiva 2374 and Mione 672 (CONN, HAO);
Santa Cruz, ruta Chorro Blanco—Monteseco, 1750 m, 21 Jan 1996, Leiva et
al. 1756 (HAO); Hualgayoc, entre Hualgayoc y Bambamarca, 2850 m, 11 Mar
1994, Sanchez 6868 (F); Celendin, desvio a Sorochuco, bajando Tahwan, 2900
m, 27 Dec 1988, Sdnchez 4943 (F); San Miguel, 7° 00' 02" S, 78° 50' 41” W,
18 Jun 1999, Leiva 2369 and Mione 668 (CONN, HAO); Contumaza, alrede-
dores de San Benito, 1300 m, 28 Mar 1985, Sagastegui and Leiva 12548 (BH,
NY). LAMBAYEQUE: Ferrefiafe, Cafiaris, 2600 m, 24 Jun 1989, Liatas 2486 (F).
cuzco: Acomayo, 2900 m, Feb 1937, Vargas 20] (F, GH, MO); Machu Picchu,
2134 m, 2 Feb 1938, Stafford 1224 (kK). APUR{MAC: Grau, Mancahuara, Oro-
peza Valley, 3000 m, 23 Jan 1939 Vargas 9798 (G, K).
390 Rhodora [Vol. 102
Bolivia. LA pAz: Prov. Bautista Saavadra, Charazani, 20 kms hacia Apolo,
2400 1 m, 5 Aug 1985, Beck 11396 (NY); Prov, Larecaja, Sorata, Challapampa,
ca. 2600 m, Jul-Aug 1863, Mandon 429 (G two sheets).
The type specimens of Saracha lobata Bitter and S. sordide-
violacea Bitter were both collected in Peru, department of Lima,
province of Huarochiri, near Matucana. Both types were de-
stroyed in Berlin; only photos are available for study. It was ev-
ident that these species belong to the genus Jal/tomata, but given
only photos of the types we were not able to decide whether these
should be placed in synonymy with Ja/tomata species, or trans-
ferred to Jaltomata. To solve this problem T. M., S. L. G., and
L. Yacher visited Matucana in 1998 and collected specimens that
match the photos. In the same region, at the type locality of J.
dentata (R. & P.) Benitez, we collected conspecific specimens we
identified as J. dentata. We conclude that S. lobata and S. sor-
dideviolacea are synonyms of J. dentata.
Jaltomata dentata (R. & P.) Benitez, Rev. Fac. Agron. (Maracay),
9(1): 91. 1976. Basionym: Saracha dentata R. & P., Fl. Pe-
ruv. 2: 43. 1797, t. 179b. Atropa dentata (R. & P.) Spreng.,
Syst. Veg. 1: 699. 1815. Bellina dentata (R. & P.) Roem. &
Schult., Syst. Veg. 4: 689. 1819. Witheringia dentata (R. &
P.) Miers, Ann. Mag. Nat. Hist., ser. 2, 11: 92. 1853. Type:
PERU. Dept. Lima: Prov. Canta, Obrajillo, 2732 m, Ruiz s.n.
(LECTOTYPE: MA, not seen; ISOTYPE: G).
Saracha lobata Bitter, Repert. Spec. Nov. Regni Veg. 18: 103. 1922.
YPE: PERU, ma—Oroya road, southwest of Matucana, 3,000 m,
Weberbauer 206 (HOLOTYPE: B, destroyed, F neg. 2553).
Saracha sordideviolacea Bitter, Re pan ee ec. Nov. Regni Veg. 18: 104.
1922. Type. PERU. Lima—Oroya road, Matucana, 2,600 m, Weber-
bauer 5258 (HOLOTYPE: B, destroyed, F neg. 2556)
ACKNOWLEDGMENTS. We thank L. Yacher for assistance with
field work, G. Bernardello and M. Nee for review, D. Spooner
for specimens and review, L. Gautier (G-pc) for photographs and
detailed description of a type, EF R. Barrie (BM) for confirming
samples of Miers’ handwriting, and the staff at MA for photo-
graphs of plates 3733 and 3734 of the Real Expedicién Botanica
del Nuevo Reino de Granada. Support was from research grants
from the Connecticut State University system and The National
Geographic Society (6008-97).
2000] Mione et al.—South American Jaltomata 391
LITERATURE CITED
BeNiTEZ DE Rojas, C. E. 1980. Solandceas nuevas para Venezuela. Phyto-
logia 47: 14-16.
DEFENSE MAPPING AGENCY. 1987. Gazetteer of Ecuador, 2nd ed. Names ap-
proved by the United States Board on Geographic Names. Published by
the Defense Mapping Agency, Washington, DC.
ee J. E 1962. Solanaceae. Field Mus. Nat. Hist., Bot. Ser. 13: Part
- a
MIONE, T, S. vA G., AND L. YACHER. 2000. Three new species of Jalto-
mata ens from Ancash, Peru. Novon 10:
AND L. A. SERAZO. 1999. Jaltomata lojae ae Description
and floral biology of a new Andean species. Rhodora 101: 136-142.
RHODORA, Vol. 102, No. 912, pp. 392—414, 2000
BIRDS, POLLINATION RELIABILITY, AND GREEN
FLOWERS IN AN ENDEMIC ISLAND SHRUB,
PAVONIA BAHAMENSIS (MALVACEAE)
BEVERLY J. RATHCKE
University of Michigan,
Department of Biology, Ann Arbor, MI 48109-1048
e-mail: brathcke @umich.edu
ABSTRACT. Pavonia bahamensis (Malvaceae) is a shrub that is endemic
to the southeastern Bahama Islands. Here I present the first detailed descrip-
tion of its pollination biology. On San Salvador Island, P. bahamensis appears
to be pollinated exclusively by two bird species, Bananaquits and Bahama
Woodstars. This pollination dependence was dramatically demonstrated in
one season when hurricanes decimated these bird populations, and pollen
deposition and fruit set of P. bahamensis was significantly reduced. However,
the lack of pollination limitation of fruit set during two other flowering sea-
sons and the relatively low pollen/ovule ratio (607) suggests that pollination
of P. bahamensis by these birds is generally reliable. Flowers show traits
typical for a bird pollination syndrome, except that the corolla is green. Flow-
ers are held horizontally on the shrub, rather than vertically, suggesting that
passerine birds (Bananaquits) rather than hummingbirds have been the most
effective pollinator and major selective agent for the floral traits in this spe-
cies. Individual Bahama Woodstars are mya pollinators, depositing few
or no pollen grains on stigmas pet it, however, they maintained pollination
one season when visits by Lenin were infrequent, and they contrib-
uted to the reliability of pollination for this endemic species.
Key Words: _ bird pollination, ages system, endemic plant pollination,
ering phenology, fruit set, green ate Hunrebitd
sonaticn, island pollination, nectar production, passerine pol-
lination, pollen/ovule ratio, pollen Aedes pollination lim-
itation, pollination syndrome
Islands typically have fewer pollinator species than mainlands
(Barrett 1996; Carlquist 1974; Elmqvist et al. 1992; Feinsinger
et al. 1982; Inoue 1993; Spears 1987; Woodell 1979). As a con-
sequence, many island plant species are generalized for pollina-
tion and have inconspicuous flowers (Carlquist 1974). Plants that
are specialized for a pollinator type, such as hummingbirds, can
be especially vulnerable to pollination limitation if only one or a
few pollinating species are present (Rathcke 1988a, 1988b, 1998:
Rathcke and Jules 1993; Wolf and Stiles 1989), unless those pol-
linators are abundant and predictable. Visitation to flowers has
role
2000] Rathcke—Pollination of Pavonia bahamensis 393
been demonstrated to be lower on islands for some species (Fein-
singer et al. 1982; Spears 1987), but pollination limitation has
not been measured. Many island plant species reduce or avoid
pollination limitation by auto-pollination and selfing (Baker 1955;
Barrett 1996; Carlquist 1974).
If plants have only a few, similar pollinating species, they
could experience stronger, directional selection for a specific pol-
lination syndrome (i.e., a suite of predictable floral traits adapted
to the most effective pollinator type, such as butterflies or birds,
Faegri and van der Pijl 1979; Stebbins 1970). For example, hum-
mingbird-pollinated flowers in western North America are typi-
cally red and tubular with large amounts of nectar and no fra-
grance (Brown and Kodric-Brown 1979; Grant and Grant 1976).
Recently, the concept of the pollination syndrome has been crit-
icized for being limited and misleading because flowering species
often have many different pollinators that vary over space and
time (Herrera 1996; Ollerton 1996; Waser et al.1996). Studies
show that flowers categorized in one pollination syndrome may
be pollinated effectively by other types of pollinators (Baker et
al. 1971; Feinsinger 1987; Schemske 1983; Schemske and Horv-
itz 1984; Waser 1983). As a consequence, selection on floral traits
may be weak or inconsistent (Ollerton 1996; Waser et al. 1996).
In contrast, an island plant with few pollinator species may be
more likely to exhibit a floral syndrome that accurately predicts
its pollinator type. Species on islands have been found to evolve
different pollination syndromes from their mainland ancestors
(Carlquist 1974; Inoue 1993), but the reliability of pollination and
pollination limitation has seldom been quantified for island spe-
cies.
In this study I present the first detailed description of the pol-
lination and reproductive biology of an endemic island shrub,
Pavonia bahamensis Hitchc. (Malvaceae; Bahama swamp-bush),
growing on San Salvador Island, Bahamas. The pollination of P.
bahamensis has only recently been described in brief reports, and
it appears to be pollinated exclusively by birds on San Salvador
(Rathcke 1998, 2000; Rathcke et al. 1996). I describe the breed-
ing system and the floral traits of P. bahamensis. 1 compare the
pollen/ovule ratio of P. bahamensis to the ratios categorized by
Cruden (1977) for plants with different breeding systems and dif-
ferent probabilities of pollination. I compare the floral traits with
those predicted for a classic bird pollination syndrome, and I dis-
394 Rhodora [Vol. 102
cuss the traits associated with passerine versus hummingbird pol-
lination.
Reliability of pollination may be especially crucial for this en-
demic island species. Pavonia bahamensis grows only on the
southeastern islands of the Bahamas in limited habitats near man-
groves (Correll and Correll 1982). Populations tend to be rela-
tively small and isolated, which may make this species especially
sensitive to changes in pollinator species or behavior (Rathcke
1998, 2000; Rathcke and Jules 1993). In general, species on is-
lands may be vulnerable to environmental changes such as global
warming, habitat destruction, or introduced species (Loope and
Mueller-Dombois 1989; but see Simberloff 1995). Species on
small islands such as San Salvador, which is only 150 km’, may
be especially vulnerable to environmental changes (Eshbaugh and
Wilson 1996). Therefore, documentation of the pollination biol-
ogy and the reliability of pollination may be valuable in providing
baseline data for future comparisons, as was found in Hawaii for
lobeliad plant species after the extinction of the Hawaiian Hon-
eycreeper (Smith et al. 1995).
STUDY SPECIES
Pavonia is in the family Malvaceae (the mallow family), which
has about 1800 species throughout the world (Fryxell 1999). Pa-
vonia is the largest genus in the tribe Malvavisceae and has an
estimated 100 to over 200 species that are mostly subtropical and
tropical (Fryxell 1999). The species are most diverse in South
America, but species are also found in Africa and other parts of
the Old World and in the West Indies, Central America, and Mex-
ico, reaching the southern United States (Fryxell 1999; Howard
1989)
Pavonia bahamensis is endemic to the Bahamas and is found
only on the southeastern islands (i.e., San Salvador, Long Cay,
Crooked Island, Acklins Islands, and the Turks and Caicos: Cor-
rell and Correll 1982). The northernmost limit is San Salvador
Island. Pavonia bahamensis was first described by Hitchcock in
1893 from a specimen collected in 1890 on Fortune Island (now
called Long Cay) in the Bahamas (Hitchcock 1893). It is a shrub
or small tree that grows in rocky coastal thickets (Correll and
Correll 1982) and inland adjacent to mangroves (Rathcke et al.
1996; Smith 1993). Its pollination and reproductive biology have
2000] Rathcke—Pollination of Pavonia bahamensis 395
been only recently briefly described (Rathcke 1998; Rathcke et
al. 1996). Little is known about the pollination biology of any
Pavonia species (Fryxell 1999).
SAN SALVADOR ISLAND
San Salvador is one of the easternmost islands in the Bahama
Archipelago (24°05'N, 74°30'W; Shaklee 1996), and it lies about
600 km east southeast of Miami, Florida and 340 km north of
Cuba (Smith 1993). San Salvador is a low, carbonate island,
about 19 km long and 10 km wide (Smith 1993). Although many
of the Bahama islands have been isolated and reconnected with
the fall ard rise of the ocean during the glacials and interglacials
of the Pleistocene (Sealey 1994), San Salvador has remained sep-
arated by a deep ocean trench (Carew and Mylroie 1997).
Total annual mean rainfall on San Salvador is 1007 mm (Shak-
lee 1996), with a rainy season from August to November (the
hurricane season) and a lesser rainy season in May and June
(Smith 1993). Annual temperature variation is 6°C (Shaklee
1996) with the coolest months averaging 22°C (January—Febru-
ary) and the warmest months averaging 28°C (July—August;
Shaklee 1996). The major vegetation of San Salvador is a scrub-
land or coppice (Smith 1993). Pavonia bahamensis grows in a
zone between the scrubland and the mangroves that line the in-
land hypersaline lakes and the tidal basin of Pigeon Creek (Smith
1993). San Salvador has about 440 species of vascular plants that
are native or naturalized, and 6-8% of these species are endemic
to the Bahamas (Smith 1993).
MATERIALS AND METHODS
I studied Pavonia bahamensis near the Bahamian Field Station
at the northeastern end of San Salvador Island. Most data were
collected on shrubs growing adjacent to mangroves on the south-
ern edge of Reckley Hill Pond about 500 m southeast of the field
station. Most of the flowering shrubs along the path are perma-
nently tagged and studied. These shrubs included most of the
local population in this area. Studies were done during three win-
ter flowering seasons (December—January) during the following
dates: December 23, 1994 to January 2, 1995; December 17, 1995
to January 4, 1996; and December 17, 1996 to January 5, 1997.
396 Rhodora [Vol. 102
This period encompasses the major flowering period on San Sal-
vador.
All animals observed visiting flowers of Pavonia bahamensis
were recorded throughout each study period over three winter
flowering seasons. I typically spent 2—6 hours a day in the local
site during most days for the entire research visit. Flower dimen-
sions, such as corolla length and stigma—anther distances, were
measured in the field using a metric ruler. Stigma—anther distanc-
es were measured from the bottom edge of the lowest lobe of a
stigma to the upper surface of the nearest anther. The age or
developmental stage of each flower was recorded to determine if
measurements changed over time. Individual flower phenology
was documented by marking and following flowers daily over
their life span, and their developmental stages were categorized.
Both unbagged, naturally-pollinated flowers and bagged, unpol-
linated flowers were observed to determine if pollination-induced
floral senescence occurred.
The flowering phenology of shrubs was quantified by counting
the numbers of open flowers per day per shrub. Total fruit set per
shrub was censused in June 1995 by counting fruit or enlarged,
dried peduncles remaining on each shrub. Seeds (mericarps) were
counted in fruits that had not dehisced. Ovaries that were devel-
oping one week after pollination typically matured fruit. There-
fore, fruit set data are based on ovary development after a min-
imum of one week. Ambiguous cases have been excluded, so
estimates of fruit set are conservative. A flower can produce a
maximum of five seeds, and most fruits sampled had five seeds
(x = 4.6, SD = 0.62, n = 25 fruits; 4 plants). Therefore, most
of the variation in seed production was due to fruit set, and those
values are reported here.
Nectar production reported here is based on the amount of
nectar in open flowers (standing crop) in 1996/97, because pol-
linator visitation was so rare that nectar was seldom removed.
These nectar volume values are similar or even higher than those
recorded previously for bagged flowers (Rathcke 1998; Rathcke
et al. 1996). Measurements were not included if nectar had over-
flowed the corolla. Nectar removals did not appear to stimulate
nectar production. To determine if nectar could be resorbed, nec-
tar was also measured in bagged flowers, which never had nectar
removed until the end of their floral life (day 3 or 4). Sugar
concentrations of nectar were measured using a Bellingham re-
2000] Rathcke—Pollination of Pavonia bahamensis 397
fractometer. Sugar concentrations are estimated as sucrose equiv-
alents and calculated from Brix values according to Bolten et al.
C1972).
To determine the breeding system, large flower buds were
tagged and given one of the following four pollination treatments:
1) bagged with no subsequent hand-pollination, 2) bagged with
self-pollen added, 3) open and augmented with cross-pollen from
at least two other individual shrubs, and 4) open and exposed for
natural pollination. The pollen-ovule ratio was calculated based
on the average number of anthers and the average number of
pollen grains per anther. Pollen numbers in upper and lower an-
thers were measured but were not significantly different.
Pollination limitation of fruit set was tested by augmenting
flowers with cross-pollen from at least two other shrubs and by
comparing this subsequent fruit set with the fruit set of naturally
pollinated flowers. Results of the pollination treatments are re-
ported in detail in Rathcke (2000). Pollination limitation (PL) was
estimated using a relative index based on fruit sets (FS—fruit per
flower) of pollen-augmented flowers (P+) and naturally pollinat-
ed flowers (NP) using the following equation:
(%FS of P+) ~ (%FS of NP)
(%FS of P+)
%PL = 100
If the percentages of fruit set of naturally pollinated flowers
and augmented flowers were equal, then PL = 0%. If fruit set
was zero for naturally-pollinated flowers and 100% for pollen-
augmented flowers, then % PL would equal 100% (Rathcke
2000).
The number of pollen grains necessary for maximum fruit set
was determined by comparing fruit set in bagged flowers that had
a known number of pollen grains deposited by hand on the stig-
mas. Cross-pollen from at least two other plants was used for
each flower. Pollen grains deposited by pollinators on exposed
stigmas by the end of floral life were counted in the field using
a 10 hand lens.
The effectiveness per flower visit of Bahama Woodstars was
measured by counting the number of pollen grains deposited per
individual visit to virgin flowers in 1995. Because overall polli-
nator effectiveness is determined by the frequency of visits as
well as by the amount of pollen transferred by a single visit,
overall effectiveness of the two bird pollinators was also based
398 Rhodora [Vol. 102
on average pollen loads on stigmas and pollination limitation of
fruit set over the three years.
Statistical analyses were done using SYSTAT ver. 5.01. Non-
parametric tests (Mann-Whitney U or Kruskal-Wallis tests) were
used due to small sample sizes and because the data were non-
normally distributed. These tests are conservative. Sample sizes
were based on the averages per plant unless otherwise noted, but
the total number of flowers is also shown for each sample. Most
flowers in the population were tagged and studied, so the data
nearly comprise the entire available flower population.
RESULTS
Pollinators. During three winter flowering periods, two bird
species were the only major pollinators seen visiting Pavonia
bahamensis flowers: Bananaquits (Coereba flaveola; Emberizi-
dae, Coerebinae), also called the Bahama Honeycreeper, and Ba-
hama Woodstars (Calliphlox evelynae;, Trochilidae; Rathcke
1998). Bananaquits are resident birds and are common flower
visitors to many plants (White 1991). The Bahama Woodstar is
the only hummingbird on San Salvador, and it is also a resident
(White 1991). I observed a single foraging bout by a Bahama
Mockingbird (Mimus gundlachii; Mimidae) in January 1997. I
saw a single visit by a wasp in 1996, but it visited between the
petals to collect nectar and did not effect pollination.
Flower visitation. Flower visitation by bird pollinators de-
creased greatly between 1994/95 and 1996/97. In 1994/95, Ban-
anaquits were the most common visitors to Pavonia bahamensis
flowers. They were in small flocks of 5—7 birds and appeared to
remain in the local area, visiting flowers continuously throughout
every day during 10 research days in 1994/95. Bahama Woodstars
were seen visiting flowers several times each day. In 1995/96,
Bananaquits were infrequently seen or heard in the site, but Ba-
hama Woodstars appeared to visit about as frequently as in 1994/
95. In 1996/97 after the severe Hurricane Lili, I never observed
either Bananaquits or Bahama Woodstars visiting flowers (see
also Rathcke 1998, 2000).
Description of flowers and fruits. Because of the apparent
specialization for bird pollination, | compared the observed floral
2000] Rathcke—Pollination of Pavonia bahamensis 399
Table 1. Floral traits of Pavonia bahamensis on San Salvador Island,
Bahamas, compared to those considered typical for a bird-pollination syn-
drome, including passerine vs. hummingbird pollinators (based on Howe and
Westley 1988 and see discussion in text); * denotes non-matching traits. Table
modified from Rathcke 2000.
“Typical” Bird Flower P. bahamensis
Corolla
Color *vivid; red *oreen; yellow anthers
Odor none
tubular corolla tubelike corolla, 18.1 mm
Orientation horizontal (passerine) horizontal
*vertical (hummingbird)
Anthesis diurnal diurnal
Phenology steady-state seasonal steady-state
Nectar ample ample (> 100 pl/flw/day)
Concentration 20% sucrose 20% sucrose
Volume > 100 pwl/flw/day >100 yI/flw/day
Secretion continuous continuous
traits of Pavonia bahamensis with those predicted for a bird pol-
lination syndrome (based on Howe and Westley 1988; Table 1;
see Figure 1). In contrast to the classic bird pollination syndrome,
the corolla and calyx of these flowers are green (see also Correll
and Correll 1982) and blend into the leaves, but the exserted
anthers with yellow pollen are highly visible (Figure 1). Other
traits appear to fit a bird pollination syndrome. Flowers have no
detectable odor. Flowers have five separate petals joined to the
staminal column. The calyx and corolla form a cup that retains
large amounts of nectar (Table 1). Although the petals are not
fused, they remain somewhat closed and form a tubelike corolla
that was 18.1 mm (SD = 1.74, n = 12 plants; 50 flowers) from
the edge of the corolla to the base for flowers measured in this
study. The average total length of the flower from the base to the
upper surface of the exserted stigma at maximum exsertion was
31.1 mm (SD = 3.15, n = 11 plants; 42 flowers). Correll and
Correll (1982) reported that petals were about 2 cm long and the
stamen column was 3 cm or more.
Flowers are perfect. The style typically had 10 stigmas on short
branches (n = 5 plants; 5 flowers). Anthers are located on the
stamen column that surrounds the style, and flowers I observed
had an average of 41 anthers (SD = 0.19, n = 14 plants; 26
flowers). On average, each anther contained 74 pollen grains (SD
400 Rhodora [Vol. 102
Figure 1. Flower of Pavonia bahamensis on San Salvador Island, Bahamas.
= 17.5, n = 7 plants; 13 flowers, 24 anthers). The number of
pollen grains per anther did not vary significantly with location
on the stamen column (upper versus lower). Although the anthers
encircle the stamen column, the filaments on the underside of the
column curve upward causing the anthers to be arranged on the
upper side of the stamen column (Figure 1). This arrangement of
the anthers probably ensures more effective transfer of pollen to
the body of a visiting bird (Figure 1).
Flowers exhibit herkogamy (spatial separation of male and fe-
male parts). On average, for the flowers I sampled, the uppermost
anther was separated from the nearest stigma lobe by 4.6 mm
(SD = 1.88, range = 1-10 mm, n = 8 plants: 85 flowers). How-
ever, occasionally flowers showed distances of 1 mm or less (2%
of flowers, n = 85). Even in this case, however, the few pollen
grains that could be transferred would not be sufficient alone to
promote fruit set where usually around 20 grains are needed (see
below). Pollen grains are large, spiny, and sticky and are not
easily moved by wind or by other movements. Typically pollen
must be transferred by a visitor.
2000] Rathcke—Pollination of Pavonia bahamensis 401
Flowers are solitary and are displayed singly on branches (see
also Correll and Correll 1982). The flowers are oriented horizon-
tally or at a slight upward angle (Figure 1).
Fruits (schizocarps) are dry, and the mericarps (each with one
seed) separate for dispersal. Each fruit has a maximum of five
mericarps. Most intact mature fruits had 4 or 5 seeds (x = 4.6,
SD = 0.62, n = 25 fruits on 4 plants). No mature fruits had | or
2 seeds and only 6% had 3 seeds. Total fruit production censused
in June 1995 ranged from 0—44 fruits per shrub (¥ = 16, SD =
16.8, n = 8 plants; 130 fruits). Based on these averages, each
shrub produced 74 seeds in June 1995. I never saw any evidence
of pre-dispersal seed predation. Fruits have spongy tissue and can
float for two weeks or more in the lab in fresh water.
Individual flower phenology. Flowers open throughout the
day, and stigmas are receptive for 2—3 days. Flowers are partially
protogynous (i.e., the stigma is receptive before the anthers de-
hisce and remains receptive until all the anthers have dehisced).
Stages of flower development are described below (based on 15
flowers on 5 plants; see also Rathcke et al. 1996). Day | (Stage
1): The stigma emerges through the closed corolla and gradually
the stigma lobes open and spread. Flowers are occasionally vis-
ited at this point and may have pollen deposited on the stigma.
Next, the corolla begins to open, the stigmas become exserted
beyond the corolla to their maximum length and the many anthers
on the upper half of the style sheath begin to emerge beyond the
corolla. Day 1-2 (Stage 2): The upper anthers begin to dehisce.
Day 2—3 (Stage 3): The lower anthers begin to dehisce. Later, the
stigma lobes begin to contract and move close together. Day 3—4
(Stage 4): All anthers are dehisced, the stigma lobes contract, the
style starts to retract into the corolla, and the corolla begins to
close. The stigma remains exserted beyond the corolla. Day 4—
5: The corolla and the stamen column fall. The style becomes
withered and brown. Subsequently the ovary either stays green
and begins to enlarge in size, or the ovary, sepals, and calyx turn
yellow and abscise, usually within about 10 days. Pollination does
not induce floral senescence.
Flowering and fruiting phenologies. The major flowering
of Pavonia bahamensis occurred in winter, November through
January, on San Salvador. Other flowering during the year ap-
402 Rhodora [Vol. 102
e 2. Nectar production of different flower stages of Pavonia baha-
mensis in December 1996. Microliters of nectar per flower per day an
sucrose-equivalents per ml are shown with means and standard deviations.
n = number of flowers from 7 tagged plants.
S Secretion Rate Sugar Concentration
(days of age) n wl/day mg/ml sucrose
Stage 1 (day 1) 19 72 + 90.9 1.30 + 1.646
Stage 2 (day 1-2) 6 ise 129.9 a 2.005
Stage 3 (day 2-3) 7 184 + 82.5 3.35 + 1.495
Stage 4 (day 3—4) 4 33 + 35.7 0.66 + 0.731
peared to be minor and I only saw a few flowers at other times.
However, Correll and Correll (1982) have reported flowering
throughout the year in the Bahamas.
Flowering showed a seasonal steady-state pattern (after Gentry
1974). Most individual shrubs had only 1-3 flowers open each
day during the major flowering season (*¥ = 2.3, SD = 2.42, n
= 3 years; 9 plants). Flowering of each shrub lasted for more
than a month, and new buds were produced as flowering contin-
ued.
Fruits developed from flowers produced in November—Febru-
ary were dispersing mericarps 5—6 months later in June.
Nectar production. Nectar was relatively dilute, with aver-
age sucrose concentration equal to 19.5% or 0.195 mg/ml (SD =
0.048, n = 7 plants; 43 flowers; Brix = 18.1 + 4.1: measured in
winter 1996/97). Nectar tasted sweet and had no other noticeable
flavor.
Nectar production was highest for Stage 3 (day 2—3) flowers
when it averaged 184 wl per flower (Table 2). Average lifetime
nectar production per flower was 458 wl. Nectar production was
continuous throughout the day and accumulated over the night to
high levels in the morning. Nectar in old flowers could be re-
sorbed. Bagged flowers in which nectar was never collected each
had no nectar or less than one microliter of nectar each (6 plants;
11 flowers) at the end of floral life. There was no evidence that
nectar removal stimulated nectar production.
Breeding system and pollen-ovule ratio. ©Pavonia bahamen-
sis plants depended upon birds for fruit set. Plants did not auto-
pollinate, and they were self-incompatible or weakly self-com-
2000] Rathcke—Pollination of Pavonia bahamensis 403
Table 3. Breeding system of Pavonia bahamensis on San Salvador Island,
Bahamas. Average fruit set is shown for bagged flowers with no hand-pol-
lination, bagged flowers augmented with self-pollen, open flowers augmented
with cross-pollen, and naturally pollinated flowers. % Fruit set equals 100
(fruits/flowers). ' Pollen was not augmented hand but pollen grains were
counted on naturally pollinated flowers. * Two of five flowers on one plant
produced fruit. Means within each season with preaches superscript letters
are significantly different, Mann-Whitney U tests, * P < 0.10.
Number % Fruit Set
Treatment Plants Flowers x + SD
1994/95
Bagged, no hand-pollination 5 , 0
Bagged, self-pollen 5 11 0)
Abundant pollen, >50 grains' 5 18 93 + 13.4
Natural pollination 6 22 82 + 30.9"
1995/96
Bagged, self-pollen 4 6 0
Augmented cross-pollen 1] 47 a ao
Natural pollination 1] 67 40 + 49.4»
1996/97
Bagged, self-pollen +t 16 10 + 20.0°*
Augmented cross-pollen 7 31 43°= 469°"
Natural pollination 7 64 L117.
patible (Table 3). Bagged flowers typically produced no fruit if
pollen was not deposited on the stigmas by hand. Flowers hand-
pollinated with self-pollen did not set fruit in 1994/95 or 1995/
96 (and see Rathcke 1998; Rathcke et al. 1996). However, in
1996/97 two flowers on one shrub produced fruit in the treatment
with added self-pollen (Table 3).
The pollen-ovule ratio for Pavonia bahamensis was estimated
to be 607. This was based on the following measurements: Flow-
ers had an average of 41 anthers (SD = 0.19, n = 14 plants; 26
flowers). Each anther contained an average of 74 pollen grains
(SD = 17.5, n = 7 plants; 13 flowers, 24 anthers). Using these
two averages, I estimated that flowers had an average of 3034
pollen grains. Flowers typically had five ovules.
Pollination limitation and pollen deposition. Fruit set was
not significantly pollination limited in either 1994/95 or in 1995/
96 (Table 3; Rathcke 2000). Fruit set of naturally pollinated flow-
404 Rhodora [Vol. 102
ers and that of pollen-augmented flowers were not statistically
different (Rathcke 2000). However, fruit set was strongly polli-
nation limited in 1996/97 after Hurricane Lili when populations
of the two bird pollinators were decimated (Murphy et al. 1998;
Rathcke 1998, 2000). Using the equation given in the methods,
percent pollination limitation = (43% — 11%) /43% = 74% (see
also Rathcke 2000). Pollen deposition on stigmas was also much
lower in 1996/97 than in the previous two years (Rathcke 2000).
Effectiveness of pollinators. Pollination effectiveness of a
flower visitor reflects both pollen transfer by an individual pol-
linator per visit and the frequency of visits. Bahama Woodstars
were not very effective as pollinators of Pavonia bahamensis,
both because individuals transferred little or no pollen to stigmas
and because they were relatively infrequent visitors. Because Ba-
hama Woodstars have long bills, and because they could probe
through the sides of the flowers between the petals, these birds
could access nectar without touching either the stigma or the an-
thers. In 1995/96, no pollen was transferred by individuals in 27%
of the visits to flowers (n = 11). For the visits that did transfer
pollen, the majority of visits (73%) transferred < 20 pollen grains
(¥ = 16, SD = 19.5,n = 11). A minimum of ca. 20 pollen grains
is needed for maximum high fruit set (Rathcke 2000). In 1995/
96 when Bananaquits were rare and Bahama Woodstars were the
most frequent flower visitors, both pollen deposition and fruit set
were lower than in 1994/95, although flowers were not signifi-
cantly pollination limited (Table 3; Rathcke 2000). Bahama
Woodstars were relatively infrequent visitors to flowers. During
a day, typically only one or two birds were observed visiting
flowers in 1994/95 and 1995/96. In 1996/97, no birds were seen
or heard in the site.
Bananaquits appeared to be effective pollinators, although the
effectiveness of single visits was not quantified. Bananaquits
probed flowers in two different ways; most often they probed
with their heads up so that the anthers contacted their breasts but
occasionally they probed with their heads upside down so the
anthers contacted their foreheads. The bright yellow Pavonia ba-
hamensis pollen was often evident on the foreheads of these birds
but was less obvious on their yellow breasts. Very rarely, birds
probed through the side of the flower between the petals and did
not transfer or collect pollen. Bananaquits tended to remain in
2000] Rathcke—Pollination of Pavonia bahamensis 405
small flocks and to visit flowers throughout the day. In 1994/95
when Bananaquits frequently visited flowers, pollen deposition
on stigmas was high and fruit set was not pollination limited
(Table 3).
Although I observed one Bahama Mockingbird visit flowers,
this occurred in 1996/97 when nectar was overflowing and drip-
ping from the flowers. It is unlikely these mockingbirds could
reach the nectar when other birds were removing it to low levels
in the flowers. The mockingbird had pollen covering its chest and
it is possible that it could have transferred some pollen. However,
pollen deposition in this winter period (1996/97) was low (51%
of the flowers had no pollen deposition by the end of flower life)
and fruit set was low and pollen-limited (Table 3; Rathcke 2000).
Therefore, Bahama Mockingbirds were not considered effective
pollinators, possibly because they rarely visited flowers and/or
were poor at transferring pollen.
DISCUSSION
As is common for many island plants, Pavonia bahamensis has
few pollinator species; its pollination appears to depend totally
on two bird species, Bananaquits and Bahama Woodstars. Perhaps
because it has only bird pollinators, the floral traits of P. baha-
mensis Closely fit those predicted by the bird pollination syn-
rome, except for corolla color (Table 1). The corolla is green
and is neither vivid nor red as is typical for hummingbird-polli-
nated flowers in western North America (Grant and Grant 1976;
Howe and Westley 1988; Raven 1972; Stiles 1976).
The red color of flowers that is typical for hummingbirds in
western North America is apparently not preferred by humming-
birds, but red is conspicuous to them and not to insect pollinators,
which may explain its selective advantage (Melendez-Ackerman
et al. 1997; Raven 1972). Because red is conspicuous, it has been
hypothesized that there is an advantage for plants to converge on
this single, distinctive flower color to attract migrating humming-
birds (Raven 1972). This color convergence would not be nec-
essary for plants on San Salvador where nectarivorous birds are
non-migratory. In fact, flowers visited by short-billed humming-
birds, like the Bahama Woodstar, in Central and South America
and the West Indies often show a diversity of colors (Feinsinger
1987) although green is highly unusual. For Pavonia bahamensis,
406 Rhodora [Vol. 102
the yellow pollen of the exserted anthers may provide the vivid
visual cue rather than the corolla. It is also possible that the flow-
ers exhibit an attractive color in the ultraviolet (Bleiweiss 1994;
Goldsmith 1980), but this was not tested for this species. Green
or greenish-yellow flowers are also found in three close relatives
of P. bahamensis (P. paludicola, P. troyana, and P. rhizophorae)
(Fryxell 1999), so green is not an unusual color in this lineage.
However, the maintenance of the green color may also reflect a
lack of selection for more vivid colors in areas where birds are
not migratory. Green corollas may also have an adaptive advan-
tage because they can contribute to photosynthesis and reduce
resource limitation of fruit set (Bazzaz et al. 1979; Jurik 1983).
Other characteristics of Pavonia bahamensis flowers are typical
of a bird pollination syndrome (Grant and Grant 1976; Howe and
Westley 1988; Table 1). Flowers have no detectable odor. The
calyx and corolla form a tube where nectar collects. Nectar per
flower is ample (> 100 microliters per day) with a sugar con-
centration of 20%, which is typical of bird-pollinated species
(Baker 1975; Bolten and Feinsinger 1978; Feinsinger 1983; Fein-
singer et al. 1985; Hainsworth and Wolf 1976; Opler 1983). In-
sects can access nectar by forcing their way between the petals,
as one wasp was observed to do. However, during three winter
flowering periods, only this single wasp individual was ever ob-
served to visit the flowers. This lack of visitation may support
the hypothesis that the dilute nectar deters bees and wasps, which
may need higher rewards (Bolten and Feinsinger 1978). Although
ants fed on the nectar when flowers were placed on the ground,
they were never seen in the flowers on the plant.
Pavonia bahamensis plants show a seasonal steady-state flow-
ering pattern, which is a common flowering pattern for plants that
support long-lived pollinators such as birds (Gentry 1974). Dif-
ferent flowers continued to open throughout the day, and nectar
was secreted throughout the day as is characteristic of many bird-
pollinated species (Howe and Westley 1988).
The pollinator specialization of Pavonia bahamensis is partly
enforced by pollinator availability: Bananaquits and Bahama
Woodstars are the only nectarivorous birds on San Salvador
(Murphy et al. 1998; White 1991). However, other bird species,
especially migratory warblers, occasionally visited the flowers of
other nearby species (see also Murphy et al. 1998). Insects, es-
pecially wasps and butterflies, can be common flower visitors to
2000] Rathcke—Pollination of Pavonia bahamensis 407
other plant species (Rathcke et al. 1996; pers. obs.). However,
these species were never seen visiting the flowers of P. baha-
mensis, with two exceptions. I saw a single wasp visit one flower
by pushing its way between the petals into the corolla tube; it
appeared to access nectar as it stayed for some time. I observed
one foraging bout by a Bahama Mockingbird feeding at flowers
overflowing with nectar during winter 1996/97 when the main
bird pollinators were scarce (Rathcke 1998, 2000). This bird had
yellow pollen on its breast and head and may have transferred
pollen. However, it is unlikely it could have reached the nectar
if nectar removal was at the levels seen in the previous two win-
ters (Rathcke 1998, 2000). Generalist pollinator species can pro-
vide compensatory pollination for plants, especially when nectar
accumulates in flowers and becomes available to more species,
and prevent or reduce pollination limitation (Wolf and Stiles
1989), but this was not the case for P. bahamensis. When pop-
ulations of two bird pollinators, Bananaquits and Bahama Wood-
stars, were decimated by the severe Hurricane Lili in October
1994, the fruit set of P. bahamensis was strongly pollination lim-
ited the following December—January (Rathcke 1998, 2000). This
species has no “‘fail-safe’> mechanism (Wolf and Stiles 1989) to
maintain pollination if these two bird species decline, and as such,
it is highly vulnerable to changes in their behavior or population
densities (Rathcke 1998, 2000).
Bird pollination is generally reliable for Pavonia bahamensis
when either Bananaquits or Bahama Woodstars are present, as
evidenced by the lack of pollination limitation in the two years
before Hurricane Lili decimated their populations in 1996 (Mur-
phy et al. 1998; Rathcke 1998; 2000). Hurricane Lili was a Cat-
egory 2 storm with winds up to 105 miles per hour (Rathcke
2000). In September 1999 an even more intense, Category 4 hur-
ricane, Hurricane Floyd, passed directly over San Salvador with
winds up to 150 miles per hour (Bahamian Field Station records),
but nectarivorous bird populations did not seem to be reduced;
both Bananaquits and Bahama Woodstars appeared to be at typ-
ical population levels (M. Murphy, pers. comm.; pers. obs.). Al-
though hurricanes affect San Salvador about every three years on
average (Shaklee 1996), few hurricanes may be severe enough to
reduce the nectarivorous bird populations. The strong pollination
limitation seen in 1996/97 may seldom occur. However, pollina-
tion limitation could also occur if birds are unreliable pollinators
408 Rhodora [Vol. 102
for other reasons. For example, in 1995/96 Bananaquits rarely
visited although they were common on the island (Murphy et al.
1998; Rathcke 2000). In that year Bahama Woodstars appeared
to be sufficiently effective to prevent pollination limitation al-
though pollen deposition and fruit set were lower. It is possible
that this island species usually has reliable pollination despite its
specialization, in contrast to some other island plants where pol-
lination is less certain with fewer pollinators (Feinsinger et al.
1982; Spears 1987).
The low pollen-ovule ratio (P/O) of 607 also suggests that pol-
lination by these two bird species is generally reliable. The value
of 607 is similar to the average ratio reported for plant species
with facultative xenogamy (X = 797) whereas the pollen-ovule
ratio for plants with obligate xenogamy (i.e., obligate outcrossers)
is much higher (P/O = 5860; Cruden 1977). Facultatively xenog-
amous species have more certainty of pollination than obligate
xenogamous species because they typically can auto-pollinate and
are self-compatible, although some species require pollinators
(Cruden 1977). Given that Pavonia bahamensis could be classified
as an obligate outcrosser, the low pollen-ovule ratio suggests that
this species may have unusually reliable pollination. Flowers are
unlikely to self-pollinate and outcrossing is usually required for
fruit set. The production of fruit by two selfed flowers in 1996/97
may represent the breakdown of the compatibility system when
cross-pollination is low, or it may reflect pollen contamination.
What is the evidence that either Bahama Woodstars or Bana-
naquits is the “most effective pollinator” and hence the stronger
selective agent molding the pollination syndrome (Stebbins 1970)?
The morphological match of bill and floral tube lengths suggests
that the Bahama Woodstar was the more effective pollinator. The
tube-like corolla was 18.1 mm and the bill length of the Bahama
Woodstar is ca. 17 mm (based on one museum male specimen
collected on New Providence in 1949 and deposited in the Mu-
seum of Zoology at the University of Michigan). In contrast, the
average length of Bananaquit bills measured from nares to tip was
10.8 mm (SD = 0.902, min = 7.95, max = 13.47, n = 221; M.
Murphy, unpub. data). Tongue lengths would also determine mor-
phological matching, but data are unavailable. A visual estimation
of tongue length in Bananaquits from a slide indicated that tongues
could extend 1.2—1.4 x beyond the bill length (ca. 13-15 mm long
or a total of 24—26 mm; Bruce Hallett, pers. comm.).
2000] Rathcke—Pollination of Pavonia bahamensis 409
Regardless of the morphological matching between bills and
tongues and corolla lengths, Bahama Woodstars were not effec-
tive as individual pollinators because they typically visited the
flowers through the side of the corolla and usually transferred
little or no pollen per visit. They were also relatively infrequent
visitors to flowers compared to Bananaquits, which foraged in
small flocks. Other evidence suggests that Bananaquits are more
effective pollinators than Bahama Woodstars. Bananaquits com-
monly had dense pollen loads on their foreheads and breasts and
they usually contacted the stigma and anthers when visiting flow-
ers. Pollen deposition was especially high when Bananaquits were
the major flower visitors in 1994/95 (Rathcke 2000) although it
is not known if this occurred because they transferred more pollen
per visit or because they were very frequent visitors. Bananaquits
usually visited the flowers so that pollen was deposited on their
chests, but they occasionally visited flowers while hanging upside
down so that pollen was deposited on their foreheads. In either
case, pollen could be easily transferred to the extended stigma if
the bird retained the same position during other floral visits. Pa-
vonia bahamensis appears to be an important floral resource for
Bananaquits on San Salvador (Murphy et al. 1998), and Bana-
naquits may be reliable pollinators over years. However, relative-
ly few were seen in 1995/96 and the reason for this is not clear,
suggesting that their foraging patterns may change and pollination
reliability over years may var
A second line of evidence also suggests that Bananaquits are
more effective pollinators than Bahama Woodstars. The horizon-
tal flower orientation in Pavonia bahamensis supports the syn-
drome for passerine pollination, rather than hummingbird polli-
nation. Flowers that are held horizontally, rather than vertically,
allow passerine birds to perch on nearby branches while feeding
(Bruneau 1997; Cruden and Toledo 1977). Another test for pas-
serine versus hummingbird pollination would be to examine sug-
ars in the nectar, but this remains to be done. Nectars of passerine-
pollinated species tend to have low sucrose/hexose ratios (<
0.499) whereas hummingbird-pollinated species tend to have high
sucrose/hexose ratios (Baker and Baker 1983; Bruneau 1997).
Bird pollination of Pavonia bahamensis may be relatively un-
usual for the genus Pavonia. Most species of Pavonia are thought
to have relatively generalized pollination (Fryxell 1999). How-
ever, hummingbirds are reported to be pollinators for several spe-
410 Rhodora [Vol. 102
cies that have tubular corollas and exserted stigma and anthers,
including P. schrankii with a yellow corolla (Gottsberger 1972),
P. viscosa (as P. montana) and P. malvaviscoides with red flow-
ers (Sazima 1981), and P. dasypetala (McDade and Davidar
1984; Roubik 1982; see also Porsch 1929). The green flowers of
P. bahamensis are unusual for a bird-pollinated flower. The three
closely related species (P. paludicola, P. troyana, and P. rhizo-
phorae) all have green or greenish-yellow flowers (Fryxell 1999),
Whether these species will also prove to be pollinated by birds,
or specifically by passerines or hummingbirds, remains to be de-
termined. Among these four species, P. bahamensis is unique in
having single flowers displayed among the leaves; the other three
species have racemose inflorescences that rise above the leaves.
If they are bird-pollinated, the more vertical, racemose inflores-
cence may reflect hummingbird pollination rather than passerine
pollination (see Cruden and Toledo 1977)
For Pavonia bahamensis, a species of passerine bird (Bana-
naquits) may be a more effective pollinator than hummingbirds,
but whether Bananaquits are more reliable over the long term
remains to be determined. Although Bahama Woodstars are in-
effective at transferring pollen, they maintained pollination one
flowering season when Bananaquits were infrequent visitors.
Having two pollinator species increased pollination reliability for
P. bahamensis, although it still incurs a risk of pollination limi-
tation if these two species decline or change their foraging pat-
terns (Rathcke 1998, 2000).
ACKNOWLEDGMENTS. I thank the Bahamian Field Station and
the staff for all their logistical support and help. I thank Lee Kass
and Bob Hunt for continued research and personal support. I also
thank the following people who made this research and paper
possible: Bob Hunt took photographs; Michael Murphy, Bruce
Hallett, and David Lahti provided critical data or information on
birds; Paul Fryxell very generously provided information from
his revision of Pavonia; Lee Kass, Paul Fryxell, Carol Landry,
Michael Murphy, and Rachel Simpson provided many helpful
editorial comments. I would like to dedicate this paper to Bob
mith who said that Pavonia bahamensis was his ‘favorite
plant”; Dr. Robert Smith published the first detailed description
of the vegetation on San Salvador Island; his untimely death was
a great tragedy.
2000] Rathcke—Pollination of Pavonia bahamensis 411
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RHODORA, Vol. 102, No. 912, pp. 415-427, 2000
NOMENCLATURAL PROPOSALS IN ATRIPLEX
(CHENOPODIACEAE)
STANLEY L. WELSH
Life Science Museum and Department of cae and Range Science,
Brigham Young University, Provo, UT 84601
e-mail: SLSLWEI eas com
ABSTRACT. This article includes a report of nomenclatural novelties in the
genus Atriplex, which were determined as appropriate following preparation
of a summary revision of the genus for the Flora of North America Project.
Subgeneric novelties include: Atriplex subgen. Obione, stat. nov. and Atriplex
pais Pterochiton, stat. nov. Sectional or subsectional taxa are: Atriplex
subgen. Obione sect. Pleianthae, sect. nov.; Atriplex sect. Obione subsect
Pee ae stat. nov.; Atriplex sect. Obione subsect. Saccariae, stat. nov.;
Atriplex sect. Obione subsect. Argenteae, Stat. ; Atriplex sect. Obione
subsect. Truncatae, comb. nov., Atriplex sect. ees subsect. Wolfianae
comb, nov.; ees sect. Obione subsect. Pusillae, pes noVv.; ee sect.
Obione subsect. Arenariae, comb. triple . Obione subsect. Let
cophyllae, pee nov., Atriplex sect. one eee Californicae, men
nov.; Atriplex sect. Phyllostegiae, comb. nov.; Gan Atriplex sect. Covilleiae,
sect. nov. New varietal combinations are: Atriplex oo var. alaskensis,
comb. nov., A. glabriuscula var. acadiensis, comb. ne . glabriuscula va
franktonii, cone. nov.; A, saccaria var. cornuta, a nov.; A. saccaria var.
asterocarpa, comb. nov.; A. argentea var. longitrichoma, comb. nov.; A. ar-
gentea var. rydbergti, comb. nov.; A. powellli var. minuticarpa, comb. nov.;
. wolfit var. tenuissima, comb. nov.; A. parishii var. minuscula, comb. nov.;
A. parishii var. depressa, comb. nov.; A. parishit var. subtilis, comb. nov.; A
parishii var. persistens, comb. nov.; A. cordulata var. erecticaulis, comb. nov.;
A. coronata var. vallicola, comb. nov.
Key Words: Atriplex, nomenclature
The proposals presented herein are preliminary to publica-
tion of a summary revision of the genus Atriplex L. for North
America. The main body of the article will be published as a part
of the Flora of North America Project, and is to be presented in
prepublication form, possibly through the internet system. Thus
it is deemed necessary to present the proposals to the scientific
community in standard form prior to that event.
Interpretations of the genus and its included taxa in North
America have undergone modifications since the first attempts at
revision of the genus by Sereno Watson (1874) and by Paul C.
Standley (1916). The work of Hall and Clements (1923) pre-
415
416 Rhodora [Vol. 102
sented a then revolutionary treatment in which species were en-
larged to include complexes of related taxa under a trinomial or
quadrinomial system. Since 1923 most workers chose to follow
the work of Hall and Clements in provincial treatments, but as
more information became available the large groupings of Hall
and Clements were dissected again (Bassett et al. 1983; Tascher-
eau 1972). Other papers have dealt with portions of the genus or
have added new taxa (Stutz and Chu 1993a, 1993b, 1997a,
1997b; Stutz et al. 1975, 1990, 1993, 1994, 1997, 1998; Stutz
and Sanderson 1983, 1998).
The present author (Welsh 1993, 1995, unpubl. ms.) has at-
tempted to strike a balance between the complex, and often un-
natural, groupings as interpreted by Hall and Clements, and the
more natural arrangement of individualized taxa of the later au-
thors. He has also attempted to examine all original descriptions
(see included list of references) and the type specimens of all
names involved in the genus in North America. Still, when taxa
were apparently closely allied and of the same general morpho-
logical conformation, they have been united in the present work
as varieties under an inclusive species. This is the case with most
of the nomenclatural novelties given below.
Many of the names treated herein were formally proposed by
Standley (1916), but with the taxonomic rank not designated.
They serve as basionyms for many of the infrageneric proposals
presented in this paper.
Atriplex gmelinii C. A. Mey. ex Bong., Mem. Acad. St. Petersb.
VI. 2: 160 (Observ. Veg. Sitcha 41). 1838
var. alaskensis (S. Watson) S. L. Welsh, Pome. nov., based on:
Atriplex alaskensis S. Watson, Proc. Amer. Acad. Arts 9: 108.
1874.
Bassett et al. (1983) distinguish var. alaskensis at the rank of
species from the closely allied Atriplex gmelinii by the sizes of
brown seeds (1.e., 1.7—2.7 mm wide in A. gmelinii and 2.8—3.7
mm wide in A. alaskensis). However, some seeds of A. gmelinit
measure as much as 3 mm wide, so the distinction in seed size
is not absolute. Also, A. gmelinii occasionally bears black shiny
seeds, which have not been observed in specimens of var. alas-
kensis. It appears that mainly juvenile plants have been collected,
those which lack mature fruiting bracteoles and seeds. The alas-
kensis phase occurs completely within the range of A. gmelinii,
2000] Welsh—Nomenclatural Proposals in Atriplex 417
and it might represent no more than a growth phase of the latter
species. Certainly there is considerable ecologically induced var-
iation within the gmelinii complex. Plants that grow within the
littoral, where they are inundated by high tide, show a completely
different series of facies than do those that are immediately above
the tidal zone. Leaves vary from linear to oblong, oval, and var-
ious other shapes within what has been traditionally regarded as
A. gmelinii in a strict sense (A. patula var. obtusa, sensu various
authors). The two entities, gmelinii and alaskensis are treated
herein as belonging to an inclusive A. gmelinii.
Atriplex glabriuscula Edmondston, Fl. Shetland 39. 1845.
var. acadiensis (Tascher.) S. L. Welsh, comb. nov., based on:
Atriplex acadiensis Tascher., Canad. J. Bot. 50: 1577-1579. 1272.
Historically, this taxon has been thought to be closely allied to
the ruderal weed, Atriplex patula L. (with which it is often sym-
patric, and with which it was synonymized by Gleason and Cron-
quist 1991:101), but from which it differs in some aspects (short-
er, stockier, rarely over 4 dm tall; bracteoles ovate-triangular and
joined only at the base, not rhombic-triangular with margins unit-
ed almost to the middle). The variety, like A. glabriuscula, sensu
lato, appears to be confined mainly to native habitats in saline
marshes, and apparently is not, or is seldom, a ruderal weed as
is the case with A. patula. It grows occasionally with the indig-
enous A. dioica Raf., which has elliptic (not round) seeds.
Plants examined from New Brunswick typically have at least
half of the nodes opposite, and with opposite branches of unequal
size. The plants still seem to be closely allied to, and perhaps not
always separable from the largely sympatric Atriplex glabrius-
cula. Bassett et al. (1983: 12) indicate that A. acadiensis formed
spontaneous hybrids with A. glabriuscula in the Botanic Garden
at Manchester England, noting further that these “presumably
sterile triploid hybrids exhibited marked heterosis.”
var. franktonii (Tascher.) S. L. Welsh, comb. nov., based on:
Atriplex franktonii Tascher., Canad. J. Bot. 50: 1586-1589. 72:
This taxon has been placed in synonymy of Atriplex hastata
L. (i.e., A. prostrata Boucher ex DC.) by Gleason and Cronquist
(1991: 102), but is clearly more nearly allied to A. glabriuscula
within whose range it is completely submersed. It is likewise
evidently confined to indigenous salt marsh habitats, unlike the
418 Rhodora [Vol. 102
clearly ruderal weedy status of the evidently introduced A. pros-
trata.
Atriplex subgen. Obione (Gaertn.) S. L. Welsh, stat. nov., based
on: Obione Gaertn., Fruct. II. 198. t. 126. 1791; subgen. Obione
[autonym created by subgen. Prerochiton (Torr. & Frem.) Ulbr.,
in Engl. & Prantl, Nat. Pflanzenfam, ed. 2. 16c: 509. 1934]; sect.
Obione (Gaertn.) Rchb., Consp. Regn. Veg. 164. 1828: C. A.
Mey. in Ledeb., Fl. Altaic. 4: 315. 1833.
Plants monoecious to subdioecious or less commonly dioecious
annuals or perennials, the leaves (typically) with or (uncommon-
ly) without Kranz anatomy. Staminate flowers with calyx lobes
crested or not. Pistillate flowers bibracteolate, lacking or rarely
with a perianth (in Atriplex covillei, A. pleiantha, and A. suckleyi).
Bracteoles cuneate to ovate or obovate united at least to the mid-
dle; the faces with tubercles or crests or smooth; the seeds erect:
the radicle typically superior (except in A. pleiantha), erect, the
tip adjacent to styles.
TYPE species: Atriplex muriculata Gaertn. (nom. illeg.) = A.
sibirica L.
The subgenus is comprised of numerous, indigenous North
American species and some Old World ones as well.
Atriplex subgen. Obione sect. Pleianthae S. L. Welsh, sect. nov.
TyPeE: Atriplex pleiantha W. Weber, Madrofio 10: 189. 1950;
Proatriplex (W. Weber) Stutz & G. L. Chu, Amer. J. Bot. 77:
366. 1990; Atriplex subgenus Proatriplex W. Weber, Madrono 10:
188. 1950.
Similis subgeno Obione secti Endolepe Torr. sed in bracteolis
multifloribus habentibus, sepalis staminata noncristatis et radicula
inferiore absimilis.
lants monoecious annuals, the leaves with normal (non-
Kranz) anatomy. Staminate flowers with calyx lobes not crested.
Pistillate flowers bibracteolate, enclosing 2—5 flowers, these t p-
ically with a 5-lobed perianth. Bracteoles triangular-ovate united
to the middle or above, the faces lacking tubercles, the seeds
erect, the radicle lateral, declined (the tip at opposite end from
the styles).
TYPE species: Atriplex pleiantha W. Weber.
The section is monotypic, with distribution as noted below. The
relationship of the solitary included taxon, Atriplex pleiantha,
2000] Welsh—Nomenclatural Proposals in Atriplex 419
with other species of Atriplex in the Colorado Plateau is illusory.
Great weight has been given to the presence of apparently prim-
itive inflorescences within the flowering bracteoles, but that fea-
ture is probably derived, and not primitive. The linear, pale, peri-
anth segments subtending the 3—5 flowers are not unlike those
within both A. phyllostegia and A. suckleyi, both of which also
lack Kranz anatomy in their leaves, but from which they are oth-
erwise grossly dissimilar. Rather than representing primitive fea-
tures, the presence of the perianth and in the case of A. pleiantha,
multiple flowers, these striking and seductively attractive features
appear to represent independent, derived occurrences within the
highly variable genus Arriplex. If any of them are to be segregated
within separate genera, then each should be so treated. The radicle
placement*in seeds of section Pleianthae is inferior (with the tip
of the radicle at a point diametrically opposed to the styles), pos-
sibly pointing to a relationship divergent from the other species
with perianth scales subtending the ovary in pistillate flowers.
Because of the radicle position, the section might well have been
placed within subgenus Afriplex. However, the radicle position
might also be a derived condition, at least in some cases. If so,
the relationship could well lie with other members currently treat-
ed within the subgenus Obione. It is anomalous wherever it is
placed.
Atriplex sect. Obione subsect. Graciliflorae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex VI. Graciliflorae Standl., N. Amer.
Fl. 21: 34, 45. 1916.
Leaves short-petiolate, the blades often subcordate, entire. Sta-
minate flowers in paniculate glomerules, the panicles soon decid-
uous. Fruiting bracteoles pedicellate, suborbicular (samara-like),
united, entire or nearly so, the faces lacking tubercles.
TYPE species: Atriplex graciliflora M. E. Jones, Proc. Calif.
Acad. Sei Se 717.3595;
The subsection is monotypic, with the solitary species endemic
to saline clays and silts of southeastern Utah.
Atriplex sect. Obione subsect. Saccariae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex VII. Saccariae Standl., N. Amer. Fl.
21: 34, 45. 1916.
The leaves short-petiolate; the blades mostly cordate or ovate,
entire. Staminate flowers in spicate or paniculate glomerules; the
420 Rhodora [Vol. 102
inflorescence soon deciduous. Fruiting bracteoles typically (but
not always) dimorphic; some large, pedicellate and the faces
mostly tuberculate, the others small, cuneate and unappendaged,
or lacking.
TYPE species: Atriplex saccaria S. Watson, Proc. Amer. Acad.
Arts 9: 112. 1874.
The subsection consists of a single species with three varieties,
all distributed from Wyoming and Utah south to Arizona, New
Mexico, and Texas.
Members of the subsection Saccariae are closely allied with,
and morphologically similar to, members of subsection Argen-
teae, and differing in the usually dimorphic fruiting bracteoles,
with the smaller sessile bracteoles mainly lacking surficial ap-
pendages, but with radiating appendages on the larger stipitate
bracteoles, or where the mainly stipitate bracteoles are mono-
morphic, by the appendages radiating from the globular surface.
The alliance of this complex with the argentea assemblage is
suggested by intermediacy of even the main diagnostic features.
It seems probable that the sessile, smooth-faced bracteole might
have been derived from some argentea type ancestor. Certainly
the two complexes are closely allied both taxonomically and geo-
graphically.
Atriplex saccaria S$. Watson, Proc. Amer. Acad. 9: 112. 1874.
var. cornuta (M. E. Jones) S. L. Welsh, comb. et stat. nov.,
based on: Atriplex cornuta M. E. Jones, Proc. Calif. Acad. Sci.
II. 5: 718. 1895.
var. asterocarpa (Stutz, G. L. Chu & S. C. Sand.) S. L. Welsh,
comb, et stat. nov., based on: Atriplex asterocarpa Stutz, G. L.
Chu & S. C. Sand., Madrofio 41: 199, 1994
The species consists of three infraspecific taxa. It is, in a broad
sense, a taxon with great variability, and is confined to the Amer-
ican West from Wyoming south to the Four-Corners portion of
Colorado, New Mexico, Arizona, and Utah.
Atriplex sect. Obione subsect. Argenteae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex VIII. Argenteae Standl., N. Amer.
Fl. 21: 34, 46. 1916. Synonym: Atriplex IX. Powellianae Standl..
N. Amer. Fl. 21: 34, 46. 1916.
Leaves petiolate or sessile, alternate or the lowermost opposite,
the blades typically broadest near the base, entire or dentate, often
2000] Welsh—Nomenclatural Proposals in Atriplex 421
hastate. Staminate flowers in axillary glomerules, or the glom-
erules paniculate. Fruiting bracteoles monomorphic, sessile or
pedicellate, usually broadest at or above the middle, the faces
tuberculate or smooth.
TYPE (LECTOTYPE: vide McNeill et al., Agric. Canada Monogr.
31: 17. 1983) species: Atriplex argentea Nutt., Gen. N. Amer. PI.
ly 198. 1s8ié.
The subsection consists of three (more or less) polymorphic
species, some of which are further subdivided into varieties, dis-
tributed from British Columbia east to Manitoba and south to
California, Arizona, New Mexico, and Texas.
Atriplex argentea Nutt., Gen. N. Amer. Pl. 1: 198. 1818.
var. longitrichoma (Stutz, G. L. Chu & S. C. Sand.) S. L.
Welsh, comb. et stat. nov., based on: Atriplex longitrichoma Stutz,
G. L. Chu & S. C. Sand., Madrofio 45: 128. 1998
var. rydbergii (Standl.) S. L. Welsh, comb. et stat. nov., based
on: Atriplex rydbergii Standl., N. Amer. Fl. 21: 47. 1916. Syno-
nym: A. pachypoda Stutz & G. L. Chu, Madrofio 44: 277. 1997.
The species is widely distributed over much of the American
West and exhibits a great variety of morphological subunits, some
of which are geographically correlated. The two combinations
proposed herein represent taxa with such correlations.
Atriplex powellii S. Watson, Proc. Amer. Acad. Arts 9: 114. 1874.
var. minuticarpa (Stutz & G. L. Chu) S. L. Welsh, stat. nov.,
based on: Atriplex minuticarpa Stutz & G. L. Chu, Madrono 40:
161. 1993.
This taxon, which occurs entirely within the geographical area
of the species proper, is a local endemic on fine-textured saline
substrates in eastern Utah.
Atriplex sect. Obione subsect. Truncatae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex X. Truncatae Standl., N. Amer. FI.
21: 34, 49. 1916.
Leaves petiolate or the uppermost sessile, alternate, the blades
typically broadest near the base, entire or nearly so. Staminate
flowers in axillary glomerules. Fruiting bracteoles monomorphic,
sessile or short-pedicellate, broadly cuneate, dentate at the trun-
cate apex, the faces typically smooth.
422 Rhodora [Vol. 102
TYPE (LECTOTYPE: vide McNeill et al., Agric. Canada Monogr.
31: 17. 1983) species: Obione truncata Torr. ex S. Watson =
Atriplex truncata (Torr. ex S. Watson) A. Gray, Proc. Amer. Acad.
Arts 8: 398. 1872.
The subsection is monotypic, with distribution rather broad in
the American West.
Atriplex sect. Obione subsect. Wolfianae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex XI. Wolfianae Standl., N. Amer. FI.
21: 34, 49. 1916.
Leaves sessile, alternate, the blades linear, entire. Staminate
flowers in axillary glomerules. Fruiting bracteoles monomorphic,
sessile or subsessile, cuneate, entire, the faces typically short-
tuberculate.
TYPE species: Atriplex wolfii S. Watson, Proc. Amer. Acad. Arts
97 112. 1874.
The subsection is monotypic, the solitary species with two geo-
graphical races separated herein as varieties; their distributions
include southern Wyoming, Colorado, and Utah.
Atriplex wolfti S. Watson, Proc. Amer. Acad. Arts 9: 112. 1874.
var. tenuissima (A. Nelson) S. L. Welsh, comb. et stat. nov.,
based on: A. tenuissima A. Nelson, Bot. Gaz. 34: 359. 1902.
This variety is known from southwest Wyoming, north-central
and western Colorado, and central to northeastern Utah.
Atriplex sect. Obione subsect. Pusillae (Standl.) S. L. Welsh, stat.
nov., based on: Atriplex XII. Pusillae Standl., N. Amer. Fl. 21:
50. 1916.
Sometimes villous as well as scurfy; the leaves opposite or
alternate, sessile, small, ovate to linear, entire. Staminate flowers
in axillary glomerules. Fruiting bracteoles monomorphic, sessile
or subsessile, typically ovate or hastate, broadest at or near the
base, entire or denticulate, the faces tuberculate or smooth.
TYPE species: Obione pusilla Torr. = Atriplex pusilla (Torr.) S.
Watson, Proc. Amer. Acad. Arts 9: 110. 1874.
The subsection is comprised of five species, some of them with
two or more constituent varieties. In large part, they are distrib-
uted in the Great Valley of California, with extensions to the
coastal region of southern California and to western Nevada and
2000] Welsh—Nomenclatural Proposals in Atriplex 423
southeastern Oregon. The members of the subsection are char-
acterized by its small leaves and tiny fruiting bracteoles.
Atriplex parishii S. Watson, Proc. Amer. Acad. Arts 17: 377.
1882
var. minuscula (Standl.) S. L. Welsh, comb. et stat. nov., based
on: Atriplex minuscula Standl., Fl. N. Amer. 21: 51. 1916.
var. depressa (Jeps.) S. L. Welsh, comb. et stat. nov., based
on: Atriplex depressa Jeps., Pittonia 2: 304. 1892.
var. subtilis (Stutz & G. L. Chu) S. L. Welsh, comb. et stat.
nov., based on: Atriplex subtilis Stutz & G. L. Chu, Madrono 44:
184. 1997.
var. persistens (Stutz & G. L. Chu) S. L. Welsh, comb. et stat.
nov., based on: Atriplex persistens Stutz & G. L. Chu, Madrono
AQ; 211.1993.
The parishii complex consists of a series of subordinate, small-
leaved taxa with distribution mainly in the Great Valley of Cal-
ifornia. They differ from other in subtle but evidently consistent
ways.
Atriplex cordulata Jeps., Pittonia 2: 304. 1892.
var. erecticaulis (Stutz, G. L. Chu & S. C. Sand.) S. L. Welsh,
comb. et stat. nov., based on: Atriplex erecticaulis Stutz, G. L.
Chu & S. C. Sand., Madrofio 44: 8&9. 1997.
This is yet another of the minor variants of species in the Cen-
tral Valley of California.
Atriplex coronata S. Watson, Proc. Amer. Acad. Arts 9: 114.
1874
var. vallicola (Hoover) S. L. Welsh, comb. et stat. nov., based
on: Atriplex vallicola Hoover, Leafl. W. Bot. 2: 130. 1938.
This is another variant of a species with distribution in the
Great Central Valley of California.
Atriplex sect. Obione subsect. Arenariae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex XIV. Arenariae Standl., N. Amer.
FI. 21: 34, 52. 1916.
Erect or decumbent-ascending, monoecious annuals or peren-
nials. Leaves with Kranz type anatomy, alternate, short-petiolate
or sessile, the blades typically widest at or above the middle,
entire or dentate. Staminate flowers in axillary glomerules or
424 Rhodora [Vol. 102
these in paniculate spikes. Fruiting bracteoles monomorphic, ses-
sile or subsessile, broadest near or above the base, dentate, the
faces smooth or tuberculate.
TYPE species: Atriplex arenaria Nutt. = A. mucronata Rat.,
Amer. Monthly Mag. & Crit. Rev. 2: 119. 1817.
The subsection is comprised of more than a dozen species,
eight of which occur in or near coastal regions from the eastern
U.S. to the south and west along the Gulf coast, and along the
coast of California.
Atriplex sect. Obione subsect. Leucophyllae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex XVI. Leucophyllae Standl., N. Amer.
FI. 21: 34, 58. 1916.
Erect or prostrate, monoecious perennials. Leaves with Kranz
anatomy, alternate, sessile, the blades typically widest at or near
the middle, entire. Staminate flowers in axillary glomerules or
these in short spikes. Fruiting bracteoles monomorphic, sessile,
rotund-ovate and spongy-thickened, entire or dentate, the faces
tuberculate.
TYPE species: Obione leucophylla Mog. = Atriplex leucophylla
(Mog.) D. Dietr., Syn. Pl. 5: 536. 1852.
The subsection is monotypic, with distribution in coastal Cal-
ifornia and Baja.
Atriplex sect. Obione subsect. Californicae (Standl.) S. L. Welsh,
stat. nov., based on: Atriplex V. Californicae Standl., N. Amer.
Fl. 21: 34, 44. 1916.
Prostrate, monoecious or dioecious perennials. Leaves typically
with Kranz anatomy, alternate or opposite, sessile, the blades wid-
est from below to above the middle, entire. Staminate flowers in
axillary glomerules. Fruiting bracteoles monomorphic, sessile or
short-stipitate, ovate, entire or dentate, the faces unappendaged.
TYPE species: Atriplex californica Moq., Prodr. 13(2): 98. 1849.
The subsection is comprised of three disparate but subtly com-
parable species, two of them from along the sea beaches and cliffs
of the California coast, the other from southwestern Texas and
adjacent Mexico.
Atriplex subgen Obione sect. Phyllostegiae (Standl.) S. L. Welsh,
Stat. nov., based on: Atriplex XXVIII. Phyllostegiae Standl., N.
Amer. Fl. 16: 34, 69. 1916.
2000] Welsh—Nomenclatural Proposals in Atriplex 425
Plants monoecious or subdioecious glabrate annuals. Leaves
with Kranz anatomy, alternate, petiolate, the blades variously
rhombic-triangular, oval or lanceolate, entire or subhastate. Sta-
minate flowers in axillary glomerules or in naked terminal spikes.
Fruiting bracteoles sessile or stipitate, sharply hastate and often
sharply cristate as well, united to above the middle, the enclosed
pistillate flower lacking a perianth.
Type species: Obione phyllostegia Torr. ex S. Watson = Atri-
plex phyllostegia (Torr. ex S. Watson) S. Watson, Proc. Amer.
Acad. Arts 9: 108. 1874.
Atriplex subgen. Obione sect. Covilleiae S. L. Welsh, sect. nov.
Folliis alternis sessilibus vel subsessilibus floribus staminatis
axillaribus vel in racemes terminales bracteolis omnibus similis.
Plants monoecious or subdioecious glabrate annuals. Leaves
without typical Kranz anatomy, alternate, petiolate, the blades
sharply triangular-hastate or less commonly some of them entire,
the overall shape ovate to lanceolate or elliptic. Staminate flowers
in axillary glomerules or in naked terminal spikes. Fruiting brac-
teoles sessile or stipitate, mostly 3-lobed, the lateral lobes round-
ed, united only at the base, the enclosed pistillate flower with a
calyx of 3 (1-5) segments.
Type species: Endolepis covillei Standl. = Atriplex covillei
(Standl.) J. E Macbr., Contr. Gray Herb. H. 53: 11. 1918.
The section is monotypic, with distribution in southeastern
Oregon, western Nevada, and California.
The pattern of venation is very similar to that of the closely
comparable Atriplex phyllostegia, even though the veins lack the
associated C-4 arrangement of chloroplast bearing cells in contact
with the veins. The plants differ otherwise as noted in the de-
scriptions. Placement of this species within the segregate genus
Endolepis by various workers is based on two morphological
characteristics considered to be of fundamental importance (i.e.,
the lack of Kranz leaf anatomy and the presence of sepals sub-
tending the ovary within the fruiting bracteoles). However, sepals
of staminate flowers in A. covillei lack the distinctive crests pre-
sent in A. [Endolepis] suckleyi, a feature on which the genus
Endelopis was based. The placement of A. covillei within Endo-
lepis, while convenient, does not take into account the overall
similarity of this species to the evidently related A. phyllostegia.
426 Rhodora [Vol. 102
Neither does it take into account the potential for recurrence of
sepals subtending the ovaries as possibly derived features.
Atriplex subgen. Pterochiton (Torr. & Frem.) S. L. Welsh, stat.
nov., based on: Pterochiton Torr. & Frem., in Frem., Rep. Exped.
Rocky Mts. 318. 1845. [Obione subgen. Prerochiton (Torr. &
Frem.) Ulbr. in Engl. & Prantl, Nat. Pflanzenfam, ed. 2 16c: 509.
1934; Obione sect. Deserticola Ulbr. in Engl. & Prantl, Nat.
Pflanzenfam, ed. 2 l6c: 508. 1934: Atriplex sect. Deserticola
(Ulbr.) McNeill, Bassett, Crompton & Tascher., Agric. Canada
Monogr. 31: 17. 1983; Atriplex XXVI. Nuttallianae Standl., N.
Amer. Fl. 21: 66. 1916; Atriplex XXVIII. Confertifoliae Standl.,
N. Amer Fl. 21: 70. 1916; Atriplex XIX. Canescentes Standl.,
N. Amer. Fl. 21: 70. 1916.]
The subgenus consists of dioecious or subdioecious shrubs of
western American distribution. The leaves possess Kranz anato-
my, and the radical position is typically superior. That they are
closely allied is indicated by the propensity of all or most of the
included taxa forming hybrids when they are in geographical con-
tact.
LITERATURE CITED
BAsseTT, I. J., C. W. CRompTon, J. MCNEILL, AND P. M. TASCHEREAU, 1983.
The Genus Atriplex (Chenopodiaceae) in Canada. Agric. Canada Mon-
ogr. 31: 1-72
GLEASON, H. A. AND A. CRONQuIST. 1991. Manual of Vascular Plants of the
Northeastern United States and Canada, 2nd ed. The New York Botanical
Garden, Bronx, N
HALL, H. M. AND E E. CLEMENTS. 1923. The phylogenetic method in tax-
onomy: The North American species of ene a? sothamnus, and
Atriplex. a Inst. Washington Publ. |-
STANDLEY, P. C. 1916. Chenopodiaceae. N. ron ai a
Stutz, H. C. AN ne L. Cuu. 1993a. Atriplex a) ie es
a new species a eastern Utah. Madrofio 40: 161-165.
AND 93b. Atriplex ee ane nee a new
species from Par aha: Madrono 40: 209—
A 997a. Atriplex subtilis ae ees a new spe-
cies from south- central California. Madrofio 44: 184—188
1997b. Atriplex pachypoda SeGe aanouinecsey, a new
species from southwestern Colorado and northwestern New Mexico. Ma-
drono 44: 277-2
s
A 5S. C. SANDERSON. 1990. Evolutionary studies of Atri-
ples: Pivlogeeti relationships of Atriplex pleiantha. Amer. J. Bot. 77:
4-369
2000] Welsh—Nomenclatural Proposals in Atriplex 427
——— —. 1993. Resurrection of the genus penne
and Saaneics of Atriplex phyllostegia (Chenopodiaceae). Amer. J.
Bot. 80: 592-597
AND . 1994. Atriplex asterocarpa (Chenopodiaceae),
a oe species from southern Utah and northern Arizona. Madrofio 41:
199—
, AN 1997. Atriplex erecticaulis (Chenopodiaceae):
i new species fom south-central California. Madrofio 44: 89-94
: 98. Atriplex De oe (Chenopodi-
ac oo a new rae from southwestern Nevada and east-central Cali-
fornia. Madrofio 45: 128-130.
_J.M. MELBy, AND G. K. LivinGston. 1975. Evolutionary studies of
Aen A relic ia diploid population of Atriplex canescens. Amer.
. Bot. 62: 236-2
AND S.C. on. 1983. Evolutionary studies in Atriplex: Chro-
mosome races of A. confertifolia (Shadscale). Amer. J. Bot. 70: 1536—
1547.
AND 1998. Taxonomic clarification of Atriplex nuttallit
(Chenopodiaceae) and its near relatives. Sida 18: 193-212.
TASCHEREAU, P. 1972. Taxonomy and distribution of Atriplex species in
Nova Scotia. Canad. J. Bot. 50: 1571-1594.
_— S. 1874. A revision of the North American Chenopodiaceae. Proc.
r. Acad. Arts 9: 82-126.
a. 'S. L. 1993. al ae pp. 130-136. In: S. L. Welsh, N. D. Atwood,
S. Goodrich, and L. C. Higgins, A Utah Flora. Life Science Museum,
ory Young Univ., Provo, UT.
995. Names and types of perennial Atriplex Linnaeus (Chenopo-
ic in North America selectively exclusive of Mexico. Great Basin
Naturalist 55: 322-334
RHODORA, Vol. 102, No. 912, pp. 428-438, 2000
SYSTEMATIC NOTES ON THE OLD WORLD FERN
GENUS OLEANDRA
ROLLA TRYON
Department of Biology, University of South Florida,
Tampa, FL 33620-5150
e-mail: Tryon@chuma.cas.usf.edu
ABSTRACT. A classification and key to Asian, Australian, and Pacific
Oleandra is presented to complement studies of Pichi-Sermolli (Africa) and
Tryon (the Americas). Six species are recognized in this large area. Man
species that have been proposed are evidently local variants and not recog-
nized here.
Key Words: Southeast Asia, Australia, Pacific, Oleandraceae, Oleandra,
em
Oleandra is one of the most distinctive genera of the Polypo-
diaceae (sensu lato). Its characters have been noted by Copeland
(1947), Kramer (1990), Nayar and Bajpai (1976), Pichi-Sermolli
(1965), Tryon (1997), and Tryon and Tryon (1982). Among the
especially noteworthy characters of the genus are the long, par-
allel and simple veins (branched, if at all, at their base), the oc-
currence of rhizophores (or unusual roots), the peltate stem scales,
and the articulated petiole. The glaucous coating on the stem of
nearly all of the specimens needs to be chemically studied.
The relationship of Oleandra is rather obscure in spite of sev-
eral reports that have considered this: Nayar and Bajpai (1976),
Ogura (1938), Sen and Sen (1973), Seong (1977), Tryon and
Lugardon (1991), Hasabe et al. (1995), and Pichi-Sermolli
(1965), who noted: “‘Oleandra differs from the other genera of
the Filicidae in many Heatutes ane this is the reason why its tax-
onomical position is debated.
There are about 35 accepted species in the region concerned,
while only six are recognized in this study. Some general com-
mentaries concerning the conservative assessment of species in
Oleandra are pertinent here. The characters of a species must be
those of a population, or of a series of populations, not of single
plants. The characters must be distinctive over a significant geo-
graphic area. Minor variations may develop in Oleandra in iso-
lated areas. Copeland (1958) notes under O. columbrina, ‘The
428
2000] Tryon—Systematic Notes on Oleandra 429
Palawan specimen is fairly typical. Otherwise, each region or
mountain has its own recognizable strain.” Plants may also have
some morphological features that relate to different environments.
Pichi-Sermolli (1965) remarked on the variable African species
O. distenta: ““This polymorphism is probably the result of great
ecological plasticity of the species which may grow in very dis-
similar habitats.”” Minor variants may deserve some kind of rec-
ognition, but not at the rank of species. Characters such as the
shape of the apex or base of the lamina and the distribution and
density of trichomes on the lamina are not sufficient to warrant
the recognition of species.
This study has involved ca. 400 collections from the wide re-
gion concerned. It is based on the collections at the Harvard Uni-
versity Herbaria (HUH) and the New York Botanical Garden (Ny).
It is not concerned with the details of distribution, or nomencla-
ture, or of taxonomy within the six species recognized. The se-
quence of species has no evolutionary implications except as the
key may illustrate affinities; synonyms are listed only where there
is relative certainty of their status.
This treatment may serve as a prodromus that may be amplified
or revised by a study of holotypes and of populations in the field.
Oleandra Cav., Anal. Hist. Nat. 1: 115. 1799. Type and sole
species: Oleandra neriformis (= O. nertiformis).
Neuronia D. Don, Prod. Fl. Nepal. 6. 1825. Type and sole species:
Neuronia asplenioides D. Don (= ere Wallichii
Ophiopteris Reinw., Syll. Pl. Nov. 2: 3. 1825. Type and <i species:
phiopterts jerieilcta Reinw. (= a neriiformis).
Aspidium subgenus Oleandra (Cav.) Splitg., Tidjs. Nat. Gesch. 7: 411.
1848.
KEY TO ASIAN, AUSTRALIAN, AND PACIFIC SPECIES
1. Leaves and phyllopodia borne on all sides of the aerial stem
and its branches, often in clusters (i.e., pseudowhorls); the
aerial stem with few or no rhizophores; a long internode
of the aerial stem a straight continuation, or nearly so, of
the lone mitemiode below «..n.csta.08etde Gheae eset (2)
2. Leaves monomorphic or nearly so; sori ca. halfway be-
tween the costa and the margin of the lamina, or closer
TOINS-COSIA a2ceneseeencecaeaeees 1. O. nertiformis
430 Rhodora [Vol. 102
i
2. Leaves strongly dimorphic; the fertile lamina much narrow-
than the sterile, 3-7 mm wide in the middle (rarely
more), and also usually much longer; sori near the mar-
gin of the lamina, the indusia often extending beyond
(Me WALI: Saeko ode ke as bhoee a akdn 2. O. Werneri
Leaves and phyllopodia borne only on the upper side of the
aerial stem, well-spaced or in clusters (pseudowhorls); ae-
rial stem with rhizophores on the lower side; the aerial
stem often arcuate beyond the leaf-bearing region, diver-
gent in orientation from the long internode below. (see
BI A ssp ett ae es 9, ak Be oles Sead no iscsi noe es (3)
Stem scales closely imbricate and usually appressed, con-
cealing the stem, bicolorous (marginate) beyond the
point of attachment .......................... (4)
4. Phyllopodia mostly less than 1 cm long (sometimes al-
most absent), often concealed by scales; lamina
more than 5 times as long as the petiole, often 10
times as long or more .....................
5. Stem with the scales mostly straight appressed
Sa ee wee ear eee . O. musifolia
5. Stem more or less squarrose- eee scales then
with a reflexed tip ........... 4. O. Wallichii
4. Phyllopodia mostly over | cm long, often 5 cm long or
more, not concealed by scales; lamina up to 5 times
as long as the petiole ............ 5. O. undulata
3. Stem with the scales irregularly patent, somewhat thinly
investing the visible stem, concolorous or nearly so and
light brown to reddish brown beyond the point of at-
PACING yudisi5:4 aig oat oS ate wae 6. O. Sibbaldii
ies)
Oleandra neriiformis Cav., originally as neriformis
Figures 1,
rs bantamense Blume, A. pistillare Swartz, Blechnum colu-
brinum Blanco, Oleandra Be Copel., O. Archbaldii Copel.
peas (Blume) Kunze, O. ciliata Kuhn, O. Clemensiae oe
el., O. colubrina (Blanco) Copel: O. cuspidata Baker, O. Herrei
Copel., O. hirtella Kunze, O. lanceolata Copel., O. maquilingensis
Copel., O. mollis C. Presl, O. nitida (Copel.) Copel., O. Parksii
Copel., O. pistillaris (Sw.) C. Chr, O. platybasis Copel., and
Ophiopteris verticillata Reinw.
The application of the earliest name remained in doubt until
2000] Tryon—Systematic Notes on Oleandra 431
Figure 1. Habit of Oleandra neriiformis, terminal portion of erect stem,
x ca. 1/4, adapted from Kalkman 4005 (A), New Guinea.
Christensen (1937) published on the original material at Madrid.
This is the most abundant and most widely distributed species
among those treated. Distinctive characters are the radially sym-
metrical aerial stem, the monomorphic leaves, and the sori that
are borne away from the margin. Merrill (1918) and Price (1973)
432 Rhodora [Vol. 102
Figures 2—4. . Portion of a ie of aoa neritformis, with many
phyllopodia, re Webster 14181 (GH), Fiji. 3. Oleandra Werneri, portion
of sterile lamina (left) and fertile a iiohi. both * 1, Brass 1284] (a),
New Guinea. 4. Arcuate internode of O. musifolia, base of petioles (above,
at left), I, Ry ree Allen 2418 (A), Sumatra.
both treated Blechnum colubrinum (Oleandra colubrina) as a syn-
onym of O. neriiformis.
Oleandra cuspidata was considered a valid species by Cope-
land (1940) who noted characters of the species as: ‘tindusium
small and fugacious” and ‘‘minute and fugative, not to be de-
2000] Tryon—Systematic Notes on Oleandra 433
tected on most specimens in the herbarium.” The material is
probably not exindusiate, the size and persistence varies, and a
small fugacious indusium may be undetected among the mature
sporangia as Copeland indicated.
The above synonymy mostly involves the species of Copeland,
mainly because he tended to recognize minor variants. This ra-
tionale may have value in evolutionary studies, but not in taxo-
nomic studies. In his Samoa study, Christensen (1943) rightly
indicated, ‘“‘After a careful examination of numerous specimens
I find the differences between the forms, briefly characterized
below, rather insignificant and inconstant, and I do not hesitate
to refer them all to Oleandra neriiformis and to refer Copeland’s
three Fijian species to the same. The forms run together and even
a grouping of the forms is difficult.”
Among the specimens of Oleandra neriiformis that I have seen,
the following are variants toward O. Werneri: Brass 5466, New
Guinea (GH, NY); Brass 31002, New Guinea (GH); Rosenstock
Exssic. 132, New Guinea (GH, NY); and Kajewski 537, New Heb-
rides (GH).
The species is usually terrestrial or epiphytic, but sometimes
lithophytic. When terrestrial, the erect aerial stems may grow to
2 m in height although they are usually 1—-1.5 m tall, and they
have a shrublike habit. The habit of the species when it grows as
an epiphyte is uncertain. It grows from 50-2200 m, mostly above
1000 m, in northern India and southwest China (Yunnan), Burma,
Vietnam, Malaysia and through Indonesia to the Philippine Is-
lands and to New Guinea, and in the Pacific to American Samoa.
2. Oleandra Werneri Rosenst. Figure 3.
Oleandra dimorpha Copel. and O. subdimorpha Copel.
The strongly dimorphic leaves and the very narrow, long and
falcate fertile leaves make this a distinctive species. The radially
symmetrical aerial stem (and presumably the creeping under-
ground stem) link this species with Oleandra nertiformis. The
fertile lamina is usually more than twice as long as the sterile
one. Sometimes the indusia project beyond the sterile tissue,
which may be reduced, especially toward the base of the fertile
lamina.
The specimen Van Royen 3644 (A), from New Guinea, has only
several fertile leaves, the sori are somewhat back of the margin,
434 Rhodora [Vol. 102
and the fertile lamina is shorter and broader than usual in this
species. In the latter two characters this resembles Oleandra sub-
dimorpha. This specimen and some others with aberrant fertile
leaves: Brass 2916, Solomon Islands (GH) and Brass 6886, 11870,
both New Guinea (GH), may represent hybrids with O. neriiformis
or, more likely represent variants of O. Werneri.
The species is epiphytic, rarely terrestrial, and it grows from
ca. 50-1800 m, usually above 1200 m, in New Guinea, New
Ireland, New Britain, and San Cristoval (Solomon Islands).
3. Oleandra musifolia (Blume) Kunze, originally as musaefolia.
Figure 4.
Aspidium Moritzii Kunze, A. musifolium Blume, Oleandra benguetensis
Copel., O. hainanensis Ching and Wang, O. Moritzii (Kunze) Kun-
ze, O. scandens Copel., and O. Wangii Ching.
The species is characterized by the long and arcuate, dorsiven-
tral aerial stems with their scales straight and fully appressed, and
by the short phyllopodia that are usually concealed by the scales.
Relations, based on the dorsiventral stems, seem to be with
Oleandra Wallichii, a species often of higher altitudes. Similarity
is also shown with O. undulata, a terrestrial species also with
dorsiventral stems. Oleandra musifolia may possibly be a pro-
genitor of these two.
The species is usually lithophytic, but sometimes terrestrial or
epiphytic. It grows from 250—1800 m in northwestern India and
Nepal, southeastern China, to southern India and Sri Lanka (Cey-
lon), through Indonesia (Sumatra to Timor), also the Philippine
Islands, Thailand (Siam), and northeastern Australia (Queens-
lan
4, Oleandra Wallichii (Hooker) C. Pres] Figure 5.
Aspidium Wallichianum Bory ex Bérlanger, Voy. Bot. 2: 56. 1833 (nom.
superfl. for A ichii and with the same type; the illustration,
PI. 5, is sieary Ole ee neritformis, not Sprengel, 1827). Aspid-
tum Wallichii Hooker, Neuronia asplenioides D. Don (nom. superfl.
for A. wallichii and with the same type
This species has few synonyms, perhaps because it was de-
scribed and figured early and the type material (Wallich, Nepal)
was widely distributed. It is found at elevations nearly 1000 m
higher than other species. The squarrose-paleaceous stems are
2000] Tryon—Systematic Notes on Oleandra 435
Figures 5-7. 5. Portion of squarrous-paleaceous stem of Oleandra Wal-
lopodium at extreme right, one at left, others with part of the petiole, x 1,
Rock 2026 (NY), Burma. 7. Portion of stem of O. Sibbaldii, * 1, Brass 29706
(A), New Guinea.
diagnostic for this species among its relations that have a dorsi-
ventral stem.
Oleandra Wallichii grows as an epiphyte or a lithophyte at
relatively high altitudes, 1100-3300 m, in the Himalayas of
436 Rhodora [Vol. 102
northern India and China (Yunnan), to Assam, in the mountains
of northern Vietnam, and in Taiwan.
5. Oleandra undulata (Willd.) Ching Figure 6.
Oleandra Cumingii J. Sm., O. intermedia Ching, O. macrocarpa C.
resl, O. pubescens Copal: and Polypodium undulatum Willd.
The application of the name follows Ching (1933), although it
seems that the holotype, consisting of two sterile leaves, is not
clearly identifiable. This taxon is tentatively kept at the rank of
species with the expectation that as more material becomes avail-
able it can be distinguished more clearly from Oleandra Walli-
chii. The most distinctive form of O. undulata has a short, cori-
aceous lamina; the sori are near the costa; the base of the lamina
is cuneate; and the phyllopodia are long. However, it is difficult
to distinguish some variants that resemble O. Wallichii. The un-
dulate margin of the leaf is not a useful character because the
wavy condition is not uniform along the margin, and the character
also occurs in other species. Holttum (1968) has clarified the
identity of Cuming 60 (PRC). This material is a mixture: a fertile
leaf with immature sori is clearly O. Cumingii, while a fertile leaf
with large mature sori is O. macrocarpa. The creeping, terrestrial
habit of this species may be derived from O. musifolia.
The species is usually terrestrial, but sometimes it is a litho-
phyte or rarely an epiphyte. It occurs from 100—1100 m in China
(Yunnan) to southern India, and eastward in Thailand (Siam),
Laos and Malaysia, to the Philippine Islands.
6. Oleandra Sibbaldii Greville Figure 7.
Oleandra crassipes Copel., O. gracilis Copel., O. vulpina C. Chr., and
O. Whitmeei Baker, originally as Whitmei.
This distinctive species is recognized by the light brown to
reddish brown stem scales that are not appressed. Many of the
scales are deciduous so that the stem surface may be exposed in
some places. The species resembles Oleandra Bradei of Costa
Rica in the deciduous character of the scales. Although O. vulpina
has been considered as a possibly exindusiate species, it is treated
here as a synonym of the indusiate O. Sibbaldii. A small, fuga-
cious indusium may be undetected among mature sporangia. The
sheet of Craven and Schodde 1123 (A) probably represents a hy-
brid with O. Werneri as shown by the sori near the margin and
the fertile leaves longer than the sterile ones.
2000] Tryon—Systematic Notes on Oleandra 437
Plants are usually epiphytic, or rarely terrestrial, from 950—
2450 m, mostly above 1200 m. It is found in the Philippine Is-
lands, Celebes (Sulawesi), Bones Island, New Guinea, and east-
ward in the Pacific to Tahiti and the Marquesas.
ACKNOWLEDGMENTS. I am most appreciative of the loan of
specimens by authorities of the Harvard University Herbaria and
the New York Botanical Garden; also for the use of the HUH
Library and the help there of Judith Warnement. I am indebted
to Betty Loraamm for the photographs illustrating details of the
plants, to Lynda Chandler for the drawing of Oleandra neriifor-
mis, and to Alice Tryon who has been helpful in many ways.
LITERATURE CITED
CHING, R. C. 1933. Notes on the herbarium Willdenow. Lingnan Sci. J. 12:
565-570.
CHRISTENSEN, C. 1937. Taxonomic fern studies, IM. Dansk Bot. Ark. 9 (3):
1-32.
. 1943. A revision of the pteridophyta of Samoa. Bull. Bishop Mus.
177: 3-138.
CoPELAND, E. B. 1940. fg ferns (Davalliaceae) of New Guinea. Phi-
lipp. J. Sci. 73: 345-357
1947. Genera Filicuni: Chronica Botanica, Waltham, MA.
. 1958. Fern Flora of the Philippines. 1: 1-191. Bureau of Printing,
Manila, Philippines
HASABE, M. ET AL. 1995. Fern phylogeny based on rbcl nucleotide sequences.
er. Fern J. 85: 134-181
Ho.Lttum, R. E. 1968. A commentary 0 on some type specimens of ferns in
the herbarium of K. B. Presl. Novit. Bot. Inst. Bot. Univ. Carol. Prag.
3-57. [Published in Ce ao lae to Holttum, Fl. Males. Ser. II, Vol
1(4): 262].
KRAMER, K. U. 1990. Oleandra, pp. 191-192. In: K. Kubitzki, gen. ed., The
Families and Genera of Vascular Plants. Vol. 1: Pteridophytes and Gym-
nosperms. Springer-Verlag, Berlin.
MERRILL, E. D. 1918. Species Blancoanae. Bureau of Printing, Manila, Phil-
ippi
NAYAR, 3B. K. AND N. Baspal. 1976. Morphology in relation to phylogeny of
= Davallioid-Oleandroid group of ferns. Phytomorphology 26: 333-
oan Y. 1938. Anatomy and morphology of Oleandra Wallichii oe 2 Pr,
with some notes on the affinities of the genus Oleandra. Jap. J. 9:
193-211.
PICHI-SERMOLLI, R. E. G. i ace florae aethiopicae, 11. Olean-
draceae. ates 20: 745—
438 Rhodora [Vol. 102
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Spores of the Pteridophyta. Springer-
RHODORA, Vol. 102, No. 912, pp. 439-513, 2000
FLORISTIC INVENTORY OF THE WACCASASSA BAY
TATE PRESERVE, LEVY COUNTY, FLORIDA
J. RICHARD ABBOTT! AND WALTER S. JUDD?
Department of Botany, 220 Bartram Hall,
University of Florida, Gainesville, FL 32611-8526
‘e-mail: galeans @yanoo, com
1]
“e-mail: wjudd ufl.edu
ABSTRACT. A floristic inventory of the Waccasassa Bay State Preserve in
southwestern Levy County, Florida was conducted from April 1996 to De-
cember 1997. The 12,488 ha (30,849 — Preserve yielded vouchers for a
total of 2 charophytes, 24 liverworts, 29 mo , 43 macrolichens, and 576
vascular plants. Of the vascular age nee 1S nL Goad 12 ferns, 1 cycad,
4 conifers, and 555 angiosper 78 of which are monocots. Sixty-nine
species are nonindigenous, and an ene are recorded for the first time from
Levy County. Seventy-two species are at or near their northern or southern
limits, 18 species have disjunct distributions or very restricted ranges in Flor-
ida, and 16 species are Florida endemics or near-endemics. Five natural plant
communities, as well as ruderal areas, were recognized based on field obser-
vations: tidal marsh, coastal hydric hammock, freshwater pools, basin swamp,
and mesic to scrubby flatwoods. Treatment of the coastal hydric hammock as
a single highly variable community, as opposed to a mosaic of intermixed
communities, was supported by a limited eae analysis.
Key Words: Waccasassa Bay, Levy County, floristics, phytogeography,
plant communities, Florida
Waccasassa Bay State Preserve is located within the Gulf Ham-
mock in southwestern Levy County, Florida (Figure 1). The Gulf
Hammock area, at the southern end of the Big Bend region of
Florida, is one of the largest, relatively undeveloped, continuous
forests remaining in the state. Gulf Hammock abuts the Gulf of
Mexico and is roughly bounded on the north by S.R. 24 and on
the east by U.S. 19, with the southern boundary running some-
what parallel to and just north of the Withlacoochee River. The
Waccasassa Bay State Preserve is a relatively thin strip that oc-
cupies most of the coast of the Gulf Hammock region, with 56
km (35 mi.) of indented shoreline (WBSPR 1997). Tidal marsh
and coastal hydric hammock dominate the 9745 ha (24,070 acres)
of the terrestrial portion of the Preserve (Figure 2). An additional
2743 ha (6775 acres) are submerged almost entirely by brackish
salt water, for a total of 12,488 ha (30,849 acres) in the Preserve
439
440 Rhodora [Vol. 102
a
aoe as
Figure 1. Map of Waccasassa Bay State Preserve and the Gulf Hammock
region in southwestern Levy County, Florida.
(WBSPR 1997). During this study, an additional 24 ha (60 acres)
at the southern end of the Preserve were purchased. Several ad-
ditional parcels of land that are slated for purchase would greatly
contribute to the extent and diversity of the more inland plant
communities (WBSPR 1997).
The Gulf Hammock Wildlife Management Area was estab-
lished in 1948 (Swindell 1949) and included the area of the more
recently established Waccasassa Bay State Preserve. The Preserve
was opened in 1972 from land purchased in 1971. Most of the
Preserve is surrounded by various hunt clubs on land leased from
Georgia-Pacific Railroad, the largest landholder in the Gulf Ham-
mock. A few private inholdings remain within the Preserve. Nu-
merous undeveloped roads transect the Gulf Hammock area, with
some of them providing access to gates along the inland boundary
of the Waccasassa Bay State Preserve. Public access into the Pre-
serve, however, is legal only by water.
The Florida Department of Environmental Protection (DEP),
Division of Recreation and Parks, District 2, manages the Pre-
serve with the principal mission of protecting natural habitat to
2000] Abbott and Judd—Waccasassa Bay State Preserve 441
Fiber Factory Road
Waccasassa Bay State Preserve
Figure 2. Map of Waccasassa Bay State Preserve, with delimited cae
communities. Thick lines within outer boundaries mark private inholdin
White areas are tidal marsh, stippled areas are coastal hydric hammock, yt
represents mesic to scrubby flatwoods, and “‘B”’ represents basin swamp.
ensure the survival of rare and endangered plants and animals.
The objectives of this floristic study were to document the current
flora with representative voucher specimens, to describe the var-
iation and distribution of the plant communities, and to provide
a baseline of botanical information for management and field use
by DEP personnel and other researchers.
Soils, geology, and physiography. Florida’s land area is the
highest portion of a plateau that is mostly submerged in the At-
lantic Ocean and the Gulf of Mexico. Past sea level fluctuations
have variously covered or exposed parts of the Floridian plateau,
which is of volcanic origin and now has a deep limestone foun-
dation underlying much of the shallow surface soil (Matter 1990).
The Gulf Hammock region has a low-energy coastal environ-
ment without adequate sand to sustain beaches or dunes (Burnson
et al. 1984). Very poorly drained, frequently flooded, strongly
saline soils of the Tidewater and Cracker series support tidal
442 Rhodora [Vol. 102
marsh throughout the Preserve. These mucky soils were formed
in loamy and clayey marine sediments underlain by limestone
(Slabaugh et al. 1996). Soils of the Wekiva, Demory, and Wac-
casassa series occur throughout the nontidal Preserve. These
poorly drained soils are shallow to moderately deep and were
formed in sandy and loamy marine sediments underlain by lime-
stone (Slabaugh et al. 1996).
Inland, the depth of the sandy soil mantle varies in thickness
over short distances, related to irregularities in the underlying
limestone (Rupert 1988). Field observations by the first author
suggest that there may be a trend for the areas of thickest sand
to support mesic to scrubby flatwoods, but there is no correlation
between this community and areas demarcated on the soil maps
of Slabaugh et al. (1996).
Geological formations underlying the uppermost surface Pam-
lico formation deposits (Pleistocene), in descending order, are the
Ocala Group, Avon Park and Lake City Limestones (all Eocene),
and Cedar Key Limestone (Paleocene; WBSPR 1997). The Pam-
lico Terrace is highly varied, due largely to depositional and later
erosional patterns, and includes irregular patches of sand or sandy
clay illuvium, brackish-water clay or sand and marl; pasty, sandy,
nonfossiliferous limestone; and sandy, coquina marl and locally
dolomitized marly sand (WBSPR 1997). Rock outcrops, primarily
of the uppermost Ocala member of the Ocala group of limestones,
are common in the Preserve.
Physiographically, the north peninsular Gulf coast of Florida
lies within the Terraced Coastal Lowlands, a broad, flat, topo-
graphical subdivision of the Coastal Plain, that comprises sandy,
Pleistocene shoreline deposits and erosional, Eocene limestone
surfaces (Vernon 1951). Alternatively, the area also is seen as
part of the Gulf Coastal Lowlands of the Mid-peninsular Phys-
iographic Zone (Rupert 1988). Following Vernon (1951), the Pre-
serve is entirely within the Pamlico Terrace, an ancient coastline
roughly demarcated by the 8 m (25 ft.) elevation line. According
to Swindell (1949), the Pamlico Terrace is not recognizable in
the Gulf Hammock, and there is no corresponding change in veg-
etation as it intergrades with the Talbot Terrace, an even older
coastline with elevations up to about 30 m. Vernon (1951) further
recognized the Preserve area as part of the Coastal Marsh Belt,
with a Limestone Shelf and Forested Hammocks zone along the
inland edge. Rupert (1988) included the Pamlico Terrace in his
2000] Abbott and Judd—Waccasassa Bay State Preserve 443
more broadly defined Limestone Shelf and Hammocks zone, al-
though he still recognized a Coastal Marsh Belt.
In the Preserve, the coast is often rocky, but marshy, with nu-
merous bays and inlets. Scattered islands dot the shoreline in and
near the Preserve. These are not true barrier islands, but were
formed as the Gulf of Mexico inundated the coastline, cutting off
relic Pleistocene sand dunes and isolating elevated areas from the
mainland.
Terrain within the Preserve grades slowly from sea level to
about 1.5 m (5 ft.) in places along the inland boundary (WBSPR
1997). Especially near the coast, there are superficial rock beds
that are much-eroded and pitted by solution. This karst topogra-
phy, derived from porous Eocene marine limestones, is an im-
portant part of the present-day continental shelf of Florida. Two
large springs in the Waccasassa River Basin and scattered small
sink-holes provide evidence of the importance of karst topogra-
phy in the hydrology of the region.
Hydrology. Salt water and coastal climate influences are
probably the most important elements that define the floristic
communities in the Waccasassa Bay State Preserve. However,
most, if not all, truly marine areas lie outside the legal boundaries
of the Preserve, which does not include coastal waters. There are
numerous inlets which may harbor pockets of marine communi-
ties interspersed with estuarine communities. The distinction be-
tween marine and estuarine communities relies on the amount of
dilution by fresh water, so there is obviously no sharp line of
differentiation. There are three sources of fresh water within the
Waccasassa Bay State Preserve: rainfall, the Floridan aquifer, and
several streams and rivers that all eventually drain into the Wac-
casassa Bay estuary or Withlacoochee Bay at the southern end of
the Preserve.
The Floridan aquifer is a regionally unconfined water table
diffused throughout pockets in porous Eocene limestones. Since
the uppermost layer of the aquifer, the Ocala Group deposits, are
locally exposed, the water table is also at or near the surface
throughout the area (Conover et al. 1984). This high water table,
in conjunction with the flat terrain, leads to quick soil saturation
and surface flooding, which can take weeks to drain after a major
storm (Suwannee River Water Management District [SRWMD]
1991). Numerous freshwater pools and wet depressions through-
444 Rhodora [Vol. 102
out the forested Preserve are maintained by rainfall. Discharge
from the aquifer and rainfall can lead to sheet flow of water across
much of the forested Preserve. In the Preserve and in the Gulf
Hammock area, the water table discharges via seepage, and it is
recharged through direct infiltration of rainwater (Rupert 1988).
As suggested by Williams et al. (1997), it is possible that this
aquifer discharge is, in part, responsible for locally reducing sa-
linity and maintaining islands of nonhalophytic species in scat-
tered areas near the inland edge of the Preserve’s tidal marsh.
All streams north of the Waccasassa River, and a few associated
tributaries to the south, are within the jurisdiction of the Suwan-
nee River Water Management District (Burnson et al. 1984).
Streams in the southernmost portion of the Preserve lie within
the Southwest Florida Water Management District (Waldron et al.
1984). There are 44 named streams and rivers that occur entirely
or partially within the Preserve (WBSPR 1997). The most im-
portant of these, because of its large drainage basin, is the Wac-
casassa River, which drains a total of 2424 km? (936 mi.:
SRWMD 1991). The Waccasassa River begins as a poorly defined
channel connecting swamps and ponds in the Waccasassa Flats
of Gilchrist and Alachua Counties and becomes a recognizable
channel west of Bronson, well into Levy County. Generally, the
Waccasassa River flows to the southwest, where it is fed by Blue
Springs and joined by Wekiva Creek, which is fed by Wekiva
Springs. Otter Creek and Cow Creek, which is joined by Ten Mile
Creek, also flow into the Waccasassa River. Numerous other
freshwater creeks and tidal channels drain the western and south-
ern portions of the Preserve.
Estuarine conditions prevail along the north peninsular Gulf
coast of Florida due to shallow coastal waters with abundant
freshwater discharge from shore. The Waccasassa River and nu-
merous small drainages are the primary sources of fresh water
within the Preserve boundary. Cedar Key and nearby islands
roughly mark the western limit of the Preserve, but actually have
the Suwannee River as the main factor controlling their estuarine
habitats (Wolfe 1990). The Withlacoochee River, although barely
south of the Preserve, has its entire drainage outside the Preserve,
and its freshwater discharge affects only the southernmost portion
of the Preserve.
The Waccasassa Bay system is an estuary at the mouth of the
Waccasassa River, the largest source of freshwater in the Gulf
2000] Abbott and Judd—Waccasassa Bay State Preserve 445
Hammock. The bay has an average depth of less than | m (3 ft.)
at mean low tide. Both the Waccasassa Bay and the Withlacoo-
chee Bay often have a depth of less than 1.5 m (5 ft.) for many
miles away from the coast, although there are deeper channels
that reflect old stream courses (Swindell 1949). There is no dis-
tinct line of separation between the Waccasassa Bay and the With-
lacoochee Bay, and both are part of the Big Bend Seagrasses
Aquatic Preserve. Tides are primarily diurnal, with a mean range
of 0.8 m (2.6 ft.; Hine and Belknap 1986). Tidal influence extends
several miles inland along creeks, which support tidal marsh spe-
cies rather than forested riverine swamp throughout the Preserve.
Coastal waters are multi-use areas and can be negatively im-
pacted by waste discharge, urban runoff, shoreline development,
and marine traffic. In the Waccasassa Bay, there are 18,949 ha.
(46,800 acres) of approved shellfish harvesting waters, with oys-
ter species offshore and in subtidal and intertidal areas (Gunter
et al. 1992). Numerous homes and small developments are pre-
sent in the Gulf Hammock area, but most are far from inland
shorelines along rivers and streams because of the expansive
swampy areas. Primary waterfront developments include Wil-
lams Camp on the Waccasassa River, Lebanon Station on Ten
Mile Creek, and Gulf Hammock on the Wekiva River and Mule
Creek. Otter Creek, Bronson, and Usher are in the Waccasassa
pee drainage basin but not near surface waters (Gunter et al.
1992). Other small towns in the region that serve as possible
sources of pollution and disturbance include Cedar Key, Ellzey,
Inglis, Rosewood, Sumner, and Yankeetown. The most probable
major source of pollution and disturbance near the Preserve
comes from extensive logging in the adjacent Gulf Hammock.
Clear-cutting could potentially alter surface water flow or lead to
contamination via surface runoff, and possible impact should be
closely monitored.
Climate. Levy County is at the southern limit of the continental
temperate zone, and has a peninsular subtropical climate in coast-
al areas (Jordan 1984). Summers are long, warm, and humid,
while winters are mostly warm but with invasions of cool air
from the north (Slabaugh et al. 1996). Relative humidity is often
high, with an annual mean of 78% (Swindell 1949). Average
relative humidity varies from about 55% in mid-afternoon to
about 90% at dawn (Slabaugh et al. 1996). During the summer
446 Rhodora [Vol. 102
season, humid breezes from the Gulf of Mexico lead to frequent
summer convection storms of high intensity, short duration, and
limited extent. Although lightning is a frequent component of
these summer storms, the hydrology and the sparse understory
contribute to the virtual nonexistence of wildfires in the area
(Swindell 1949). From November to February there are prevail-
ing northwesterly to northern winds which bring frontal systems
into the area, with precipitation of low intensity, long duration,
and wide coverage (Chen and Gerber 1990).
Levy County had an average annual rainfall of 127 cm (50 in.)
from 1841 to 1949 (Swindell 1949). During the same time period,
at Cedar Key, just west of the Preserve, the average annual rain-
fall was 119.4 cm (47 in.), but it varied from 68.6 cm (27 in.) to
210.8 cm (83 in.; Swindell 1949). These extremes are similar to
those more recently reported for the region, namely, 78.7 cm (31
in.) and 222.3 cm (87.5 in.; Jordan 1984). Within the Waccasassa
River Basin, the average annual rainfall from 1977-1989 was
158.8 cm (62.5 in.; SRWMD 1991). These data support the
broadly generalized maps of Jordan (1984) and Tanner (1996),
which showed several different patterns of rainfall in the Gulf
Hammock region. Thus, on average, the westernmost portion of
the northern Preserve may receive up to 20 cm (8 in.) less rain
annually than the easternmost inland portion of the Preserve.
Ironically, it is in this westernmost corner of the northern Pre-
serve that the most well-developed and extensive swamps occur.
This surely must reflect drainage patterns and not the direct rain-
fall patterns.
For the entire region, there is a pronounced rainy season from
June to September, during which time 50—60% of the mean an-
nual rainfall occurs. Up to a third of the average annual rainfall
often comes in September alone, in conjunction with tropical
storms and hurricanes (Jordan 1984). The national record for the
most rainfall in a 24 hour tae occurred at the southern end of
the Preserve in Yankeetown on 5—6 September 1950, with 98.3
cm (38.7 in.). Less than 25% of the yearly rainfall occurs from
December to March (Jordan 1984). Relatively severe drought oc-
curs in the spring every 8—10 years on average, infrequently last-
ing into the early summer (Burnson et al. 1984). The impact of
occasional dry spells is probably less defining for the area than
the frequent periods of inundation.
At Cedar Key, the average annual temperature is 22.2°C (72°F;
2000] Abbott and Judd—Waccasassa Bay State Preserve 447
Burnson et al. 1984). From 1841-1949, the average January tem-
perature was 14.7°C (58.4°F), with an extreme low of —9.4°C
(15°F; Swindell 1949). During the most severe cold snap in this
century, on 21—22 January 1985, temperatures dropped as low as
—12.2°C (10°F) in the area (Tanner 1996). The average July tem-
perature at Cedar Key was 27.7°C (81.8°F), with an extreme high
of 38.3°C (101°F; Swindell 1949). On average, 100-150 days a
year reach a maximum of 35.6°C (88°F) or higher (Tanner 1996).
Freezing temperatures, on average, occur 29 days per year (Sla-
baugh et al. 1996). Frost-free seasons were noted for 6 of 34
years of data from Cedar Key (Swindell 1949).
Average climatic patterns may typify an area, somewhat de-
termining the vegetation, but extremes of hydrology and climate
are probably more important for determining actual species com-
position. Major disturbances, such as hurricanes, though rare, are
also important in defining plant community structure and often
eventually lead to environmental heterogeneity and increased spe-
cies richness. Hurricanes occur mostly in the fall, from August
to October, with strong winds and often with torrential rains.
Sometimes two or three major storms hit or pass near the Preserve
in a single year, but usually there are many years between hur-
ricanes that severely impact the area (Matter 1990). Only five
hurricanes have hit the coast near Cedar Key since 1871 (Ho and
Tracey 1975). A weak tropical storm that hit Cedar Key in Oc-
tober 1941, produced 89 cm (35 in.) of rain inland in just 48
hours (Tanner 1996). Frontal systems in the winter and spring can
also lead to dramatic flooding and tidal surges. Given the flat
terrain, the high water table, and the far-reaching impact of tidal
surges, even a relatively minor storm can flood the area or carry
salt water vane far inland, thus affecting species composition.
Mexico currents reportedly moderate the coastal cli-
mate, eee to slightly warmer winters and slightly cooler sum-
mers along the coast, on average, than are found inland (e.g.,
Jordan 1984). The depth of inland penetration and the full extent
of this current-related climate moderation are questionable, and
such moderation must be highly variable locally. Several sub-
tropical plant species at their northern limit are present on coastal
islands in the area, while they are absent inland. The dynamic
interactions in the area between varying hydrological and climatic
extremes contribute to the confusing mosaic of ecotonal plant
associations that dominate much of the forested Preserve.
448 Rhodora [Vol. 102
HISTORY
The Waccasassa Bay State Preserve was created in 1972. The
Preserve is south of the Suwannee River, well north of Tampa
Bay, and just east of Cedar Key, three well-known areas with
long histories of human occupation. Historical detail can be found
elsewhere [see especially Gannon (1993, 1996), George (1989),
Jennings (1951), Milanich (1994, 1995), Swindell (1949), Tebeau
(1971), and Webb (1990)], and is summarized in Abbott (1998).
The Gulf Hammock Wildlife Management Area (ca. 40,470 ha.
or almost 100,000 acres) was created in 1948, and included all
of the land now considered part of the Waccasassa Bay State
Preserve. Prior to the Civil War, there was a sugar cane plantation
2 miles south of the community of Gulf Hammock and another
on the south bank of Ten Mile Creek. Both of these plantations
were just east of the current Waccasassa Bay State Preserve, and
both were abandoned at the end of the Civil War, although the
latter area had minor farming until around 1900. In the late 1940s,
Swindell (1949) found the farmed area to be indistinguishable
from surrounding forested areas, suggesting rapid regeneration.
Traditionally, people in the Gulf Hammock area mainly had
small gardens and free-ranging livestock. Agriculture has been
little-practiced due to the region’s poor drainage and shallow soils
underlain by limestone. Cattle and hogs were found throughout
the area, even well into salt marsh (Swindell 1949). There were
large herds of cattle in the area even before 1900. Cattle grazing
and hog disturbance, while still common in the Gulf Hammock
proper, have been somewhat controlled by fences along much of
the boundary of the Waccasassa Bay State Preserve. Signs of hog-
rooting were observed only in one part of the northern Preserve,
and cows were seen to have breached the fence only in an area
south of the Waccasassa River.
Records exist of as many as 20 different lumber companies in
the Gulf Hammock area, but specific details are poor or lacking.
Some evidence does exist of the impact, as compiled by Swindell
(1949) and Jennings (1951), based primarily on oral interviews
and old aerial photographs. Bald cypress was logged from
swamps shortly after the Civil War. Almost all of the coastal
hammock was cut over for southern red cedar and cabbage palm.
Red cedar was cut, especially for pencils, from 1875 until about
1920, with mills at Cedar Key and along the coast. Cabbage
2000] Abbott and Judd—Waccasassa Bay State Preserve 449
palms were cut for fiber made from the bud, starting around 1900
and still ongoing as late as 1949. There was a fiber mill on Cow
Creek around 1900. During the peak activity, an estimated
100,000 plants were being cut a year. Hardwood logging started
later, and probably peaked in the 1920s after most of the cypress
and cedar were gone. Numerous logging roads were established
in the early 1950s in the areas of driest, firmest soil. Mesic ham-
mocks were more severely affected by logging than were hydric
hammocks due to the larger number of suitable timber trees.
Swamp trees were rarely cut in the early 1950s since cypress,
ash, and swamp tupelo were already largely gone. Logging in
Gulf Hammock continues to the present, with logging in the area
of the current Preserve possibly into the 1940s or 1950s. Rem-
nants of old logging roads can still be seen, but there is virtually
no overland vehicular access within the Preserve now. Presently,
many pine stands within the Preserve are being cut in an effort
to control a southern pine beetle infestation (Dendroctonus fron-
talis). This disturbance will undoubtedly have a profound impact
on plant community structure in the affected areas.
Human activity is by no means the only kind of disturbance
that affects plant community composition in the area. Root sys-
tems are shallow due to the thin soil underlain by limestone, and
heavy winds and storms often blow down large numbers of trees.
As mentioned previously, hurricanes sometimes hit the area and,
occasionally, severe northern frontal systems come through dur-
ing the winter or spring. The aftermath of one such storm was
witnessed in the spring of 1996 when the first author saw nu-
merous large treefall areas and debris as high as 2 m throughout
much of the forest just inland from the salt marsh. One local
resident reported an 8 ft. water surge at his house over a mile
from the coast.
Despite the relatively recent development (in the last 3000
years) of the current mesic habitats in the Preserve area, the coast-
al hydric hammock found in the Gulf Hammock area contains
virtually all of the diversity and variation known in hydric ham-
mocks found elsewhere in Florida (Vince et al. 1989). It 1s pos-
sible that the area served as a refugium of sorts, as there are a
few regionally endemic (or near endemic) animals and plants:
Elaphe obsoleta williamsi, a type of rat snake described from
Gulf Hammock; Microtus pennsylvanicus dukecampbelli, the
Florida salt marsh vole known only from one location just west
450 Rhodora [Vol. 102
of the Preserve (SRWMD 1991): Pseudobranchus striatus lus-
tricolus, a subspecies of salamander described from Gulf Ham-
mock, in stagnant mucky water (Neill 1951); and Spigelia logan-
loides, Phaseolus smilacifolius, and Phyllanthus liebmannianus
ssp. platylepis. Additionally, at least two other plant species un-
common in Florida are present in great abundance within the
region, namely, Leitneria floridana and Ulmus crassifolia.
PLANT COMMUNITIES
“Unlike the rest of Florida, much of the north peninsular coast-
line has not been ditched, diked, graded, filled, or otherwise al-
tered by modern development, giving us a glimpse of what a
soggy place the Gulf coast of Florida used to be’’ (Milanich 1994,
p. 210),
An excellent, detailed, ecological characterization of Florida’s
northern peninsular Gulf coast region was provided by Wolfe
(1990). Thompson (1980) described the forest vegetation in the
northern Gulf Hammock. Plant communities within the Gulf
Hammock area, of which the Waccasassa Bay State Preserve is
a part, were described in detail by Jennings (1951), Pearson
(1951), and Swindell (1949). Although their studies were zoolog-
ical in nature and some of their plant identifications, unfortunately
unvouchered, are certainly questionable, our own observations
largely support their plant community descriptions, which were
mostly of areas further inland than the Preserve. Much of the
Gulf Hammock floods regularly, altering the species composition
somewhat from that typically found in standard community de-
scriptions used elsewhere in the state. Nonetheless, all of the
above Gulf Hammock workers described separate coastal, mesic,
and hydric forested communities, recognizing that they usually
occurred as a mosaic of poorly defined, intermixed small patches.
Jennings (1951) captured the essence of the first author’s early
BSED IS to understand the vegetation when she wrote that there
was “‘a multiplicity of ecotones, succession stages, and other con-
fusing plant aggregations in the area.”” As Swindell (1949) point-
ed out, the size of the area precludes complete field-reconnais-
sance, and it is almost impossible to distinguish similar com-
munities in the area using aerial photos. Thus, it is likely that
undiscovered pockets of vegetation may vary in composition
from the descriptions provided here.
2000] Abbott and Judd—Waccasassa Bay State Preserve 451
In order to make our work maximally useful to other workers
in Florida, we followed the community types outlined by the
Florida Natural Areas Inventory and Florida Department of Nat-
ural Resources (FNAI and FDNR 1990), with one exception as
noted later. Our emphasis was on the recognition of plant com-
munities that are discernable based on species composition and
not based on potential differences in hydrology related to micro-
topographical variances.
Five natural communities, in addition to ruderal areas, are here
recognized within the Waccasassa Bay State Preserve: tidal
marsh, coastal hydric hammock, freshwater pools, basin swamp,
and mesic to scrubby flatwoods (Figures 3 and 4). Tidal marsh
is the only floral-based group in the Preserve within the combined
FNAI marine-estuarine categories. Seagrass beds exist just out-
side the coastal boundary and are described here as they report-
edly occur in scattered localities within the many shore inlets
(SRWMD 1991). In the FNAI palustrine category, the basin wet-
lands group is represented by basin swamp, and the wet flatlands
group is represented by a type of hydric hammock. Here, we
deviate from the FNAI community description and recognize
coastal hydric hammock, following Vince et al. (1989). The term
hammock, used primarily on the coastal plain of the southeastern
United States, refers to an area of hardwood trees, often in an
otherwise treeless or pine-dominated area. Scattered throughout
the hydric hammock were numerous freshwater pools not assign-
able to an FNAI community, yet they were ecologically and flo-
ristically distinctive. Finally, in the FNAI terrestrial category, the
mesic flatlands group was represented by a mixture of mesic and
scrubby flatwoods. Upland mixed forest (synonym: mesic ham-
mock) was frequent just outside of the inland Preserve boundary
and is described here since small patches of vegetation transi-
tional to upland mixed forest occur along the inland margin of
the Preserve. Ruderal areas can occur within any of the com-
munities but are recognized by their ““weedy”’ aspect, often with
signs of human disturbance and nonindigenous plant species.
Even FNAI and FDNR (1990) admit that ““FNAT classification
is perhaps more often useful ... in potential natural vegetation
rather than existing vegetation” and that community lines are
often obscure in the field. As explained below, several other
FNAI communities could possibly be recognized within the Pre-
serve, but it is our judgement that the area is best described as a
&
N
Rhodora
[Vol. 102
2000} Abbott and Judd—Waccasassa Bay State Preserve 453
relatively thin coastal strip, predominantly with tidal marsh zones
in nonforested areas and highly variable coastal hydric hammock
in forested areas. The lack of several distinctive forest commu-
nities, as described by others in the Gulf Hammock area, though
at first confusing, was eventually understood as a reminder of the
arbitrariness of plant community delimitation and of the contin-
uum that often exists between different communities.
All of the FNAI marine-estuarine communities were present
along the coastal edge of the Preserve, but the mineral and faunal
based community groups were outside the scope of this study, as
were algal beds. Tidal swamp (synonym: mangrove forest) did
not exist per se, but there were several coastal island areas in the
southern portion of the Preserve where black mangrove (Avicen-
nia germinans) could be found.
The difference between terrestrial and palustrine systems is that
palustrine systems have soil that is inundated or saturated for
more than 10% of the growing season, resulting in plant com-
munities that are adapted to regular periods of anaerobic soil con-
ditions (FNAI and FDNR 1990). As mentioned earlier, much of
the Preserve and adjacent Gulf Hammock flooded regularly, and
the species composition was different from that typically found
in similar communities elsewhere in the state. Thus, in an area
like the Preserve where widespread inundation followed every
storm and heavy rain, the distinction between terrestrial and pal-
ustrine systems may not be real or meaningful. Nonetheless, flat-
woods and swamp are here recognized as they appeared to be
more or less distinct from coastal hydric hammock in a few areas
within the Waccasassa Bay State Preserve.
Following terminology of the Florida Natural Areas Inventory,
the forested Preserve could be described as an inseparable com-
plex mosaic of northern maritime hammock (synonym: coastal
ee
Figure 3. Plant communities nearest the coast. A. Tidal marsh abutting
coastal hydric hammock near Turtle Creek, dominated by Juncus roemert-
anus; low foreground with Batis maritima and Salicornia perennis. B. Tidal
marsh in southern Preserve, showing tidal channel and Avicennia germinans.
C. Waccasassa River, banks dominated by Juniperus virginiana and Saba
palmetto. D. Brackish water pool in southern Preserve; banks dominated by
Juncus roemerianus, with Claduim jamaicense;, background shrubs are Bac-
charis halmifolia, background trees are Pinus taeda.
elopoyuy
TOA]
cOl
2000] Abbott and Judd—Waccasassa Bay State Preserve 455
hammock), mesic hammock, hydric hammock, basin swamp, and
several other terrestrial and palustrine communities. In a study of
the ecology of hydric hammocks throughout Florida, Vince et al.
(1989) recognized the Gulf Hammock region as having its own
distinctive variant of hydric forest and called it coastal hydric
hammock, including much of the variation described as separate
intermixed communities by Jennings (1951), Pearson (1951), and
Swindell (1949). The first author quantitatively investigated the
variance in this plant community and concluded that the majority
of the forested Preserve is best treated as coastal hydric ham-
mock, a highly variable plant community where species typical
of many different FNAI communities can be found growing in
intermixed patches related to microtopography, hydrology, and
past disturbance.
Seagrass beds. In areas west and south of the mouth of the
Waccasassa River, which had an unconsolidated mud bottom,
sparse to dense seagrass beds occurred. No seagrass species were
documented within the Preserve boundary during this study, but
it is very likely that at least a few populations occurred within
the many coastal inlets. The SRWMD (1991) reported the inshore
presence of Halodule wrightii and Halophila engelmanni, al-
though no vouchers have been seen. A total of five species of
seagrass have been reported from the general area, with four of
them present between the Cedar Key area and the Withlacoochee
River (Iverson and Bittaker 1986). Thalassia testudinum (turtle-
grass) was reportedly the dominant bed-forming species. Syrin-
godium filiforme (manatee-grass), Halodule wrightii (shoal-
igure 4. Plant communites farther inland. A. Coastal hydric hammock
near Turtle Creek; dominated by Juniperus virginiana and Sabal palmetto,
understory very sparse, here with scattered rue eanaes spp. and Panicum
spp asin swamp in northwestern corner of Preserve; mixed hardwoods
as dominants, here with Liquidambar styraciflua, oe caroliniana, es
cus laurifolia, and Magnolia virginiana, with Taxodium distichum, note the
standing water, cypress knees, and an understory with Rhapidophyllum hys-
ix. C. Scrubby flatwoods along boundary trail off Dewey Allan Park Road;
canopy of Pinus taeda, with sparse Sabal palmetto; understory with Serenoa
repens, Ilex glabra, Quercus myrtifolia, Myrica cerifera, goricee ale a
Rhynchospora spp., and Liatris spp. D. Freshwater pool in southern Preserve;
dominated by Teoh domingensis, with Pinus taeda in the ea
456 Rhodora [Vol. 102
grass), and Halophila engelmanni were also reportedly present,
usually in intermixed beds. Ruppia maritima (widgeon-grass), a
freshwater to estuarine species, was observed by the first author
in the bay near the mouth of the Waccasassa River.
Tidal marsh. Coastal areas frequently inundated by salt wa-
ter and dominated by salt tolerant herbs are known as tidal marsh
(synonyms: salt marsh, coastal marsh, brackish marsh). Tidal
marsh here is alluvium-poor, with numerous karst features such
as creek channels, circular depression ponds, and limestone out-
crops. A mosaic of marshes and coastal hammocks has been cre-
ated by the low-energy karstic coastline, where small changes in
elevation, tidal inundation, soil characteristics, and fresh water
flow all control vegetation patterns (Wolfe 1990). In the area,
there is usually a very broad continuum from near-marine con-
ditions to near-freshwater conditions. The community was often
around 1.5 km wide, although it could be up to 5 km wide in
areas, due to the very gradually sloping continental shelf. Nu-
merous tidal creeks with oyster reefs and unvegetated intertidal
flats interlace with tidal marsh and support it inland along wa-
terways throughout the width of the Preserve. The following de-
scription of variation in tidal marsh is based on the first author’s
personal observations within the Preserve.
The seaward edge of tidal marsh often consisted of Spartina
alterniflora (saltmarsh cordgrass) stands. Extensive stands of Jun-
cus roemertanus (needle rush) dominated most of the area. Other
plants in the Juncus stands were sporadic and were most fre-
quently Aster tenuifolius and Solidago sempervirens, with Lyth-
rum lineare locally abundant in a few places. Along the edges of
needle rush stands, which sometimes occurred in relatively inland
habitats, most of the species mentioned below could also be
foun
Open flat depressions, often with exposed limestone, are scat-
tered throughout the tidal marsh. These depressions could be
completely bare or have mixed to nearly monospecific stands of
the following halophytic species: Salicornia perennis, Batis mar-
itima, Borrichia frutescens, and Distichlis spicata, or less com-
monly, Salicornia bigelovii, Sporobolus virginicus, Sesuvium por-
tulacastrum, and Blutaparon vermiculare. Similar areas, when
forming stands just seaward of coastal hammocks, are referred to
by FNAI as coastal grassland (synonyms: salt flat, overwash
2000] Abbott and Judd—Waccasassa Bay State Preserve 457
plain, coastal savannah). Reportedly, these areas were historically
burned by ranchers to promote better grazing conditions (Pearson
1951). In addition to all of the species found in the depressional
flats, these more inland areas usually had Triglochin striata, Li-
monium carolinianum, Spartina patens, and Agalinis maritima,
often in association with a more or less shrubby transition zone
frequently composed of Iva frutescens, Baccharis halimifolia, and
Lycium carolinianum, and occasionally with Baccharis glome-
ruliflora, B. angustifolia, and diminutive Forestiera segregata.
Areas that appeared to be less saline often had a mixture that
could include all of the above species, plus some of the following:
Fimbristylis spadicea, Ipomoea sagittifolia, Bacopa monnieri,
Cynanchum spp., Eleocharis spp., Cladium jamaicense, and
sometimes Rayjacksonia phyllocephala, Flaveria linearis, and
Eustoma exaltatum. All of the above areas frequently occurred
adjacent to or intermixed with stands of Sabal palmetto (cabbage
palm), Juniperus virginiana var. silicicola (southern red cedar),
and Quercus virginiana (live oak).
The least brackish areas, not necessarily always the farthest
inland (depending on topography), were often dominated by
Cladium jamaicense and, in a few areas, Typha domingensis, ot-
ten with pockets of Juncus roemerianus. Most of the above spe-
cies could still be found as associates, with the addition of Ac-
rostichum danaefolium, Crinum americanum, and Samolus spp.
The most prevalent submerged plant in inland brackish water was
Ruppia maritima. Myriophyllum pinnatum occurred in the least
brackish areas.
The FNAI beach dune community (synonym: coastal strand),
is essentially nonexistent in the area, but two small coastal islands
were seen with thin sand deposits. Both areas were bordered by
coastal hydric hammock and tidal marsh. One had /pomoea pes-
caprae, Sesuvium portulacastrum, Cyperus esculentus, Cenchrus
echinatus, and Heliotropium curassavicum. The other had Cakile
lanceolata, Atriplex pentandra, and Chenopodium berlandieri,
with Sideroxylon celastrinum nearby. A separate, thin, elongated,
raised sandy area parallel to the shore supported a dense stand of
giant Amaranthus australis, bordered by Spartina alterniflora and
Scirpus robustus on the seaward side and Juncus roemerianus on
the inland side. Perhaps this area could be considered a poorly
developed example of an FNAI coastal berm (synonym: coastal
evee):
458 Rhodora [Vol. 102
Numerous slightly elevated islands occur throughout the tidal
marsh, with vegetation varying from scrubby flatwoods (only a
very few in the southernmost portion of the Preserve) to coastal
hydric hammock to entirely herbaceous associations (as described
above). The abundant islands of coastal hydric hammock were
dominated by Sabal palmetto and Juniperus virginiana var. sili-
cicola, frequently with Quercus virginiana, Ilex vomitoria, Iva
frutescens, Baccharis halimifolia, and Opuntia stricta. Occasional
associates included Pinus taeda, Quercus laurifolia, Forestiera
segregata, Yucca aloifolia, Zamia floridana, and Stenotaphrum
secundatum. Rarely, almost any of the coastal hydric hammock
species, and most individuals of more tropical species, such as
Maytenus phyllanthoides and Eugenia axillaris, could be found
on islands. Interestingly, Persea borbonia, which is entirely re-
placed by P. palustris inland, seemed restricted in the area to
coastal islands. Islands of tall cabbage palms and live oaks in the
midst of marsh communities are considered prairie hammocks by
FNAL in other parts of Florida, but they are clearly just relict
coastal hammock outliers in the Preserve. The inland spread of
salt water inundation due to rising sea levels, studied by Williams
et al. (1997), has resulted in the die-off of terrestrial species, with
relict individuals of cabbage palm, southern red cedar, and live
oak. Cabbage palm, S$. palmetto, is usually the last species to
succumb, and scattered clumps, individuals, and standing dead
trunks could be found throughout tidal marsh.
Coastal hydric hammock. Hydric hammock is a community
virtually restricted to Florida. The most extensive stands of hydric
hammock in Florida occur along the Gulf Coast, often forming a
belt just inland of salt marsh, and were referred to as coastal
hydric hammock by Vince et al. (1989). Throughout the state,
hydric hammock varies from nearly monospecific stands of Sabal
palmetto to mixed stands of S. palmetto and Juniperus virginiana,
or dense hardwood stands with highly variable species compo-
sition. Statewide, hydric hammock often has a broadleaf ever-
green appearance and is typically dominated by S. palmetto, J.
virginiana, Quercus virginiana, and/or Q. laurifolia (laurel oak),
and often also has Liquidambar styraciflua (sweetgum) and Car-
pinus caroliniana (hornbeam). Gulf Hammock is the largest con-
tiguous stand of hydric hammock in Florida and includes almost
all of the variation in structure and composition found in hydric
2000] Abbott and Judd—Waccasassa Bay State Preserve 459
hammocks throughout the state. Along inland edges of the Gulf
Hammock, coastal hydric hammock is usually intermediate with
other community types, such as swamps and mesic hammock
(Vince et al. 1989). Almost all of the species known to occur in
hydric hammock in Florida can be found in the Gulf Hammock
(Vince et al. 1989), including relatively rare trees such as Prunus
americana and Ulmus crassifolia, and rare herbs such as Spigelia
loganioides, all of which were found within the Waccasassa Bay
State Preserve. The following descriptions of coastal hydric ham-
mock are based entirely on field observations by the first author.
Areas closest to salt marsh often contained near-pure stands of
Sabal palmetto and Juniperus virginiana, often with an understo-
ry of Iva frutescens, Lycium carolinianum, Baccharis halimifolia,
and Yucca aloifolia. Frequently, Quercus virginiana also was pre-
sent in the canopy, with an understory of //ex vomitoria, Myrica
cerifera, B. halimifolia, Viburnum obovatum, and often Foresti-
era segregata. This coastal-most forest was the only type of can-
opy cover included as coastal hammock by authors such as Pear-
son (1951). This area is reportedly very similar to hydric ham-
mock bordering marshes along the St. Johns and Myakka Rivers
(Vince et al. 1989). Occasionally, especially adjacent to inland
fingers of tidal marsh, Pinus taeda, Ulmus spp., and Fraxinus
spp. could be locally abundant in the canopy. Lianas were rare
in the most coastal areas, which mostly had only Smilax bona-
nox and Toxicodendron radicans, these species increased in abun-
dance with distance inland. Herbs were usually infrequent or lack-
ing in the most coastal areas, and were predominantly represented
by scattered salt marsh species and grasses, especially Chasman-
thium spp. and Panicum spp. With increasing distance from the
salt marsh, S. palmetto and J. virginiana decreased in abundance
while mixed hardwoods increased. However, within the Preserve,
which occupies a narrow coastal strip, there were virtually no
areas where S. palmetto and J. virginiana were not important
canopy members.
Inland, trees characteristic of FNAI mesic to hydric hammocks
and swamps (i.e., mixed hardwoods) became more common,
though rarely did more than two or three of these species occur
together. Most such species had a scattered distribution. The fol-
lowing abundance estimates are based on the first author’s ob-
servations over the entire area of coastal hydric hammock. Fre-
quent woody additions inland were Pinus taeda, Quercus laurt-
460 Rhodora [Vol. 102
folia, Ulmus alata, U. crassifolia, U. americana, Acer saccharum
ssp. floridanum, and Persea palustris. Mostly along the inland
boundaries, one occasionally found Carpinus caroliniana, Carya
glabra, Diospyros virginiana, Liquidambar styraciflua, Celtis lae-
vigata, Gleditsia triacanthos, Tilia americana, Quercus shumar-
dit, Erythrina herbacea, and Sabal minor. Infrequent woody
plants were Forestiera ligustrina, Morus rubra, Ptelea trifoliata,
Rapanea punctata, Sapindus saponaria, and Zanthoxylum clava-
hercules. The rarest of the intermixed woody mesic species were
Cercis canadensis, Crataegus aestivalis, Prunus americana, Aes-
culus pavia, Osmanthus americanus, Viburnum dentatum, Quer-
cus michauxit, Q. nigra, Sideroxylon lanuginosum, Aralia spi-
nosa, and Symphoricarpos orbiculatus. In wet depressions and in
areas transitional to swamps, Fraxinus caroliniana, F. pennsyl-
vanica, Acer rubrum, Magnolia grandiflora, and rarely Cornus
foemina, Carya aquatica, Nyssa biflora, and Ilex cassine, could
be found. Areas transitional to flatwoods had a greater abundance
of Pinus taeda, but there were also numerous isolated stands of
Pinus taeda scattered elsewhere through the forest.
Frequent inland lianas were Vitis spp., Smilax bona-nox, and
Toxtcodendron radicans. Occasional lianas were Sageretia min-
utiflora, Smilax auriculata, S. tamnoides, and Campsis radicans.
Berchemia scandens, Matelea gonocarpos, Bignonia capreolata,
Chiococca alba, Parthenocissus quinquefolia, Gelsemium sem-
pervirens, Ampelopsis arborea, and Lonicera sempervirens were
infrequent. Rare lianas and vines were Phaseolus smilacifolius,
Smilax smallii, S. laurifolia, Cocculus carolinus, Clematis crispa,
C. catesbyana, and Dioscorea floridana.
The ground layer was often bare or with sparse, patchy herbs,
such as Chasmanthium spp., Panicum spp., Elephantopus elatus,
and Ruellia caroliniensis. In the areas that flooded least, a rela-
tively thick ground cover of the same species could develop,
along with Zamia floridana. Additional woodland herbs that
could be found scattered or in patches included numerous sedges
and grasses, along with Salvia lyrata, Galium hispidulum, Rubus
spp., Elytraria caroliniensis, Dyschoriste humistrata, and Sani-
cula canadensis. In a few scattered areas grew Spigelia logan-
loides, Phyllanthus liebmannianus ssp. platylepis, Trepocarpus
aethusae, Euphorbia commutata, and Lithospermum carolinense.
Bare areas often had a scattering of wetland herbs, described be-
low under freshwater ponds. In many areas, there were thick,
2000] Abbott and Judd—Waccasassa Bay State Preserve 461
lawn-like stands, mostly of Stenotaphrum secundatum, with a few
areas of Axonopus furcatus. Both of these are native grasses, but
their growth pattern often seemed unnatural and perhaps reflects
remnant areas of old logging roads.
Freshwater pools. Included here are shallow rainwater de-
pressions, relatively deep isolated pools, and isolated inland chan-
nels. Most of the areas likely flood with salt water during extreme
tidal surges in connection with major storms, but sheet flow of
surface rainwater may serve to dilute them sufficiently to main-
tain their freshwater vegetation. The Florida Natural Areas In-
ventory does not have a community type that corresponds to the
freshwater pools in the Preserve. Since many of them are ephem-
eral and in depressions, they could be treated as variants of de-
pression marsh (synonyms: isolated wetland, ephemeral pond),
but the FNAI habitat description and species lists show nothing
in common with these areas. A couple of the deepest ponds (ca.
1.5 m deep) are similar to miniature sinkhole lakes, and could
perhaps be treated as versions of them, since they are karstic in
origin. But, rather than forcing several ill-fitting FNAI names here
that have highly overlapping species compositions, we chose just
to describe the variation, without recognizing the pools as a dis-
tinct FNAI community, since most of the pools are nothing more
than wet depressions whose species are mostly scattered in sat-
urated soil in areas throughout the coastal hydric hammock and
basin swamp communities.
Shallow rainwater depressions are common throughout the Pre-
serve. Many of the depressions are ephemeral, and presumably,
saturated soil conditions are responsible for supporting wetland
plants when there is no surface water. Many depressions were
devoid of vegetation or have only two or three species growing
in them. Leitneria floridana was frequent overall, usually forming
dense shrubby stands, often with nearby Fraxinus spp. and Salix
caroliniana in semi-permanently flooded areas. The largest of
these depressional areas with long-standing water often appeared
very similar to small patches of swamp forest. Most commonly,
Hydrocotyle umbellata, Samolus ebracteatus, S. valerandi, Ba-
copa monnieri, Sabatia calycina, Lippia nodiflora, Crinum amer-
icanum, Cardamine pensylvanica, Pluchea spp., ASclepias per-
ennis, and Iris hexagona were found. Saururus cernuus, Centella
462 Rhodora [Vol. 102
asiatica, Polygonum hydropiperoides, and Ammania_ latifolia
were also occasionally found in these areas.
Areas of deeply pooled fresh water are uncommon within the
Preserve. Only three such areas were seen by the first author. One
of them actually had the estuarine Juncus roemerianus, Spartina
patens, and Ruppia maritima in association with Cladium jamai-
cense, Leitneria floridana, Hibiscus coccineus, and Thalia geni-
culata. The other two pools were bordered by Cladium, Leitneria,
Hibiscus, and Panicum spp. One of them also had Hibiscus gran-
diflorus and was filled with submerged and floating mats of Ba-
copa monnieri with Nitella capillata. The other pool was more
diverse, with a dense stand of Typha domingensis in the middle
and with Nymphaea odorata, N. elegans, Chara zeylanica, My-
riophyllum pinnatum, Echinodorus berteroi, Sagittaria graminea,
and Polygonum hydropiperoides along the edges. This latter pool
was highly dynamic during the period of study. The above-listed
species were the dominants in the beginning. During a final visit
to the area, the first author found Lemna obscura and Cerato-
phyllum echinatum to be the dominants, with the original species
virtually absent. Closer inspection revealed that the water level
had risen by at least half a meter and that many of the original
species were still present as small plants along the bottom in
about one meter of water. Dozens of other deep pools were seen,
with a strong estuarine influence, as indicated by the salt marsh
vegetation and brackish water, with only one submerged or float-
ing aquatic, Ruppia maritima.
Numerous inland channels were also seen, mostly with an ob-
vious or seasonally intermittent continuity with tidal marsh. In
several areas, however, the inland boundary trail cuts across fin-
gers of elongate depressions that appear to be isolated portions
of predominantly freshwater creek channels. Vegetation included
species found near the depression pools, though it was usually
dominated by Leitneria floridana and Hibiscus coccineus. Addi-
tionally, Sagittaria lancifolia, Kosteletzkya virginica, Rumex ver-
ticillatus, and several graminoids were found primarily along a
few of these freshwater channels.
Basin swamp. Basin swamp (synonyms: cypress swamp,
hardwood swamp, mixed swamp) occupies low areas that are
flooded more frequently for longer periods of time and to a great-
er depth than hydric hammock (Vince et al. 1989). Only two
peed.
2000] Abbott and Judd—Waccasassa Bay State Preserve 463
distinct swamp areas were found within the Preserve, totaling
perhaps 15 ha., although several small forest patches in scattered
localities had standing water and trees such as Carya aquatica,
Acer rubrum, and Fraxinus caroliniana. Much of the largest
swamp area was actually very similar to adjacent coastal hydric
hammock and perhaps could have been considered as just another
variant form of hydric hammock. But the diminution in abun-
dance of Sabal palmetto and Juniperus virginiana was distinctive,
as was the hydrology, especially in conjunction with the presence
of Taxodium distichum, which is considered very rare in hydric
hammock (Vince et al. 1989). Swamps were also characterized
here by the abundance of Acer rubrum, Nyssa biflora, Fraxinus
pennsylvanica, F. caroliniana, and Magnolia virginiana, and by
the rare presence of orchids such as Malaxis spicata, Hexalectris
spicata, and Habenaria floribunda, of ferns and allies such as
Selaginella apoda, Thelypteris palustris, T. kunthii, abundant Ac-
rostichum danaefolium, and of other noteworthy plants such as
Lobelia cardinalis and abundant Rhapidophyllum hystrix.
Mesic to scrubby flatwoods. Mesic to scrubby flatwoods
(synonym: intermediate pine flatwoods) are typically dominated
by Pinus taeda (loblolly pine) and Sabal palmetto. Hydric ham-
mock species are intermixed, which is not characteristic for mesic
or scrubby flatwoods. These areas within the Preserve are not
typical for well-developed flatwoods elsewhere in the state. Thus,
most of the flatwoods-like areas in the Preserve could easily be
seen as just another variant form of a broadly defined hydric
hammock plant community. Patches of P. taeda occur naturally
throughout the Preserve, but along the boundaries of the southern
portion, these patches likely reflect past logging, as clear-cut areas
were widely replanted with P. taeda according to Vince et al.
(1989). Aerial photos from the 1950s indicate that many of the
areas considered here had been severely logged. Even though
these areas would, perhaps, best be treated as a transitional var-
iation of coastal hydric hammock, they are mostly very distinctive
in the field and are here recognized as pine-dominated areas with
a characteristic assembly of understory species in the southern
portion of the Preserve, totaling perhaps 11 ha.
Flatwoods in Florida have poor drainage and low topography,
which is true for most of these areas. Fires, usually every 10—20
years, are important in maintaining species composition in flat-
464 Rhodora [Vol. 102
woods. Only one area near the coast showed signs of having
burned within the last few years, and it had an extremely dense
stand of Serenoa repens (saw palmetto). Pinus taeda (loblolly) is
one of the least fire-adapted pines in Florida, but several of these
pine-dominated areas in the Preserve did have a little intermixed
P. elliottii (slash pine), a typical flatwoods species. The prepon-
derance of loblolly and the admixture of species more typical of
well-drained sandy areas probably reflect the effects of fire ex-
clusion and human disturbance in the area. The rareness of spe-
cies typical of scrub suggests that the area most likely was not
originally scrub, but an area of flatwoods that has been invaded
by scrub species, perhaps due to fire exclusion.
Within the Preserve, canopy members were frequently Pinus
taeda and Sabal palmetto, occasionally with Juniperus virgini-
ana, Quercus virginiana, Q. laurifolia, and Persea palustris.
Rarely, Magnolia grandiflora, Osmanthus americanus, Q. gemi-
nata, Prunus serotina, Carya glabra, and Tilia americana were
intermixed. Some of these areas were notably poorly drained and
had an understory of Serenoa repens and Myrica cerifera with
scattered Ilex glabra, Vaccinium arboreum, Lyonia fruticosa, and
L. lucida. Dense tangles of Smilax spp. were frequent. Other sim-
ilar areas also had Hypericum tetrapetalum, H. hypericoides, Car-
phephorus odoratissimus, Vaccinium myrsinites, Bejaria race-
mosa, and Liatris spp. Additional rare indicative herbs included
Aster tortifolius, Penstemon multiflorus, Silphium astericus, Le-
chea mucronata, Bulbostylis stenophylla, Cyperus retrorsus,
Buchnera americana, Piriqueta caroliniana, and Galactia elliot-
tii. The most scrubby-looking areas, oddly on sandy pockets ad-
jacent to tidal marsh, also contained Q. myrtifolia, Asimina lon-
gifolia, Myrica cerifera var. pumila, Xyris spp., and Polygala spp.
Upland mixed forest. There are no pure stands of upland
mixed forest (synonym: mesic hammock) large enough to con-
sider as distinct within the Preserve boundary. Tiny patches along
slightly elevated ridges within hydric hammock do approach up-
land mixed forest, as described by the FNAI and FDNR(1990)
and as differentiated by Vince et al. (1989). We include this brief
description here since much of the adjacent Gulf Hammock has
been described by past workers (e.g., Swindell 1949) as mesic
hammock and there may be a few undiscovered areas within the
Preserve that would best be treated as mesic hammock.
2000] Abbott and Judd—Waccasassa Bay State Preserve 465
Pearson (1951) found that even in the Gulf Hammock, mesic
hammocks are usually along ridges or islands of better drained
soils, without forming large continuous tracts. Although they are
a major portion of the overall hammock, an extensive network of
lower drainage areas separates the mesic hammock patches. His-
torically, the mesic hammocks had a very sparse understory with
visibility of several hundred meters. Jennings (1951) reported that
mesic hammocks can be inundated for several hours after heavy
rains to several days after major storms due to poor drainage in
the Gulf Hammock. Most of the species reported by Jennings
(1951), Pearson (1951), and Swindell (1949) as typical of mesic
hammock were found within the Preserve, but they were rarely
together in groups of more than 2 or 3 species and occurred
primarily along the inland boundaries.
As described in the adjacent Gulf Hammock area (Swindell
1949: Vince et al. 1989), characteristic canopy trees were Mag-
nolia grandiflora, Quercus michauxii, and Acer saccharum vat.
floridanum, often with Ostrya virginiana, Tilia americana, Cercis
canadensis, Ilex opaca, and Pinus taeda. Other common species
that were more widespread, and thus less indicative, were Car-
pinus caroliniana, Quercus virginiana, QO. nigra, Liquidambar
styraciflua, Sabal palmetto, Juniperus virginiana, Persea palus-
tris, Celtis laevigata, Ulmus alata, Acer negundo, and Aralia spi-
nosa. Common shrubs were Ilex vomitoria, Serenoa repens, Vac-
cinium arboreum, Viburnum dentatum, Callicarpa americana,
Sageretia minutiflora, Euonymus americana, and Myrica cerifera.
Less common shrubs were Zanthoxylum americanum, Ptelea tri-
foliata, and Rhamnus caroliniana. Common vines included Tox-
icodendron radicans, Ampelopsis arborea, Vitis rotundifolia,
Campsis radicans, Parthenocissus quinquefolia, Bignonia ca-
preolata, Gelsemium sempervirens, and Smilax bona-nox. Herbs
included Panicum commutatum, Oplismenus setarius, Elephan-
topus spp., Sanicula canadensis, Mikania scandens, Salvia lyrata,
Dioscorea floridana, Melothria pendula, and Mitchella repens.
Ruderal areas. Ruderal, human-created, open areas are very
infrequent in the Waccasassa Bay State Preserve, largely due to
the lack of roads and legal public access by land. In the Preserve,
most ruderal areas would be virtually impossible to distinguish
from areas of natural disturbance without the nearby fences and
tire ruts, because ruderal areas usually were dominated by native
466 Rhodora [Vol. 102
species characteristic of the adjacent communities, a testament to
the relatively pristine nature of the Gulf Hammock. Nonetheless,
virtually every species documented from the Preserve can be
found along or near a trail or road somewhere, so the emphasis
here is to point out areas where nonindigenous species are found.
Native species restricted to ruderal areas in the Preserve have
been marked accordingly in the species list. In accordance with
the Preserve’s policy for control of exotic species, non-native
species were eliminated, when found, in all localities except Fiber
Factory Road where sheer numbers made it unfeasible. Undoubt-
edly, propagules in the soil and re-introduction will maintain the
presence of most of the documented species.
The forested boundary has been mowed at least once in the
past twenty years for much of the Preserve, creating an open and,
in areas, somewhat disturbed trail up to a few meters wide. While
the boundary is typically dominated by species native to forest
gaps and wet depressions, a few nonindigenous plants rarely were
found (e.g., Apium leptophyllum, Conyza bonariensis, Hyptis mu-
tabilis, Medicago lupulina, Murdannia nudiflora, Paspalum no-
tatum, P. urvillei, Richardia brasiliensis, and Spermacoce pros-
trata).
There are a few areas where old access roads still exist or
where illegal entrance and use have created undeveloped roads
into the Preserve. One such road near Turtle Creek had a few
individuals of Medicago lupulina, Mitracarpus hirtus, and Secale
cereale, with occasional Plantago major and Youngia japonica.
A few illegal entrance roads at scattered localities were found
with Lindernia crustacea forming small patches in wet depres-
sions. Echinochloa crusgalli and Polypogon monspeliensis were
found in scattered wet pools. Stenotaphrum secundatum is a na-
tive species, but its robustness and dense growth along old road-
ways suggest that the triploid cultivar form may have been intro-
duced in areas.
One public gravel road, Dewey Allen Park Road, actually cuts
across tidal marsh in the southwestern-most boundary of the Pre-
serve. Exotics along this road were Crotalaria spectabilis, Hyptis
mutabilis, Lantana camara, Medicago lupulina, Melilotus alba,
M. indica, Paspalum urvillei, and Verbena brasiliensis, all of
them occasional.
An area known as the Northcut Property is a recent acquisition
in the southern end of the Preserve. The area was a homesite,
2000] Abbott and Judd—Waccasassa Bay State Preserve 467
with an undeveloped road transecting it. In this area occurred a
single Albizia julibrissin, dense Eremochloa ophiuroides, infre-
quent Medicago lupulina and Verbena brasiliensis, and rare
Sphagneticola trilobata.
Around 4 km (2.5 mi.) of Fiber Factory Road, an undeveloped
access road and its right-of-way, are owned by the state and are
included in this study, though they are not within the main body
of the Preserve. This roughly eight-meter swath cuts through a
wide variety of areas on its way to the coast, and perhaps a third
or more of the documented species could be found at some point
along this road. Just outside the right-of-way, was a large Melia
azedarach, but no seedlings were seen. Cows were abundant in
this area, and several feeders were along the road, with exotic
plants concentrated near them. It is in this area that signs of graz-
ing could also be seen inside the Preserve. Exotics here, most of
which were only in one or a few small areas, were Amaranthus
spinosus, Arenaria serpyllifolia, Cerastium glomeratum, Conyza
bonariensis, Coronopus didymus, Cynodon dactylon, Eleusine
indica, Hedyotis corymbosa, Hyptis mutabilis, Kummerowia Sstri-
ata, Kyllinga brevifolia, Lamium amplexicaule, Lindernia crus-
tacea, Medicago lupulina, Murdannia nudiflora, Paspalum dila-
tatum, P. notatum, Pavonia hastata, Phyllanthus urinaria, Poa
annua, Portulaca amilis, Raphanus raphanistrum, Senna obtusi-
folia, Sonchus asper, Sporobolus indicus, Stellaria media, Trifo-
lium campestre, Verbena brasiliensis, Veronica arvensis, Vicia
sativa, and Youngia japonica. Within the Preserve itself, the road
continued, locally with dense carpets of Eremochloa ophiuroides,
with rare intermixed Cyperus rotundus, Desmodium triflorum,
and Sisyrinchium rosulatum, and occasional Kyllinga pumila,
Medicago lupulina, Murdannia nudiflora, and Phyllanthus urt-
naria. Near this access point, and at two other localities in the
forested Preserve, the first author found Citrus aurantium.
Along the inland edge of tidal marsh, where piles of wrack
accrued after tidal surges, three small plants of Schinus terebin-
thifolius were found. This species was observed in great abun-
dance outside the Preserve on Cedar Key and Seahorse Key. Dis-
seminules are likely to continue to be brought in from numerous
other coastal locations as well. One coastal island had a diffuse
population of Cyperus esculentus. Another island had Tetragonia
tetragonioides along the salt marsh edge.
468 Rhodora [Vol. 102
Rather extensive logging, though scattered over the last cen-
tury, has undoubtedly affected species composition in the Wac-
casassa Bay State Preserve, and has left its mark in the form of
old access roads, which mostly have been revegetated by forest
species. During the course of this study, there was a southern
pine beetle (Dendroctonus frontalis) outbreak that necessitated
road construction and logging. The full impact of this disturbance
remains to be seen. Probably the single most important factor that
will shape the future plant communities in the Preserve, however,
is the rising sea level. It can be expected that more and more of
the Preserve will be inundated and that the plant communities
will shift inland.
QUANTITATIVE FLORISTICS: VARIATION WITHIN
COASTAL HYDRIC HAMMOCK
When the first author began this floristic inventory, much of
the forested Preserve appeared to be a confusing array of possibly
distinct plant communities, at least based on canopy dominants.
Eventually, he realized that there were areas with distinctive
swamp forest and mesic to scrubby flatwoods, but in most of the
forested Preserve, here treated as coastal hydric hammock, three
canopy extremes were seen, although all shared occasional to
abundant Sabal palmetto and Juniperus virginiana. Many areas,
especially those closest to the coast, were dominated by just S.
palmetto and J. virginiana with only a few scattered hardwoods,
primarily Quercus virginiana and Q. laurifolia. Previous workers
in the region have called these juniper and Saba/-dominated areas
coastal hammock (e.g., Swindell 1949). Some areas had a canopy
codominated by several mixed hardwoods, including trees such
as Acer rubrum, A. saccharum, Fraxinus caroliniana, F. penn-
sylvanica, Liquidambar styraciflua, and several Quercus species.
These areas with mixed hardwoods seemed similar to mesic ham-
mock and, in places, swamp forest. Other areas had a canopy
with abundant Pinus taeda and seemed to be a possible variant
of flatwoods. We decided to investigate whether the highly vari-
able coastal hydric hammock (according to Vince et al. 1989)
might be better treated as a mosaic of intermixed, yet distinctive,
plant communities.
2000] Abbott and Judd—Waccasassa Bay State Preserve 469
A general outline of our approach is as follows. Scattered
patches of the three extreme canopy types were located through-
out the Preserve. The Preserve was divided into roughly equal
northern and southern portions, separated by the Waccasassa Riv-
er. Using available access points along the Preserve boundaries,
aerial photos, a compass, and a hand-held global positioning unit,
the first author attempted to ensure that at least one of every three
map sections (U.S. Geological Survey topographical quadrangle
maps, 7.5 minute series) had a plot placed within it, so that half
the plots were scattered over the northern Preserve and half over
the southern Preserve. Typically, no more than two plots were
placed within an area of 2.6 km? (1 mi.*), roughly the amount of
area covered by an aerial photo map (Florida Department of
Transportation; scale 1 in. = 400 ft.). A plot was placed in each
of 26 forest patches. In each plot, the presence of all understory
species was recorded. Coefficient of community values (Whitta-
ker 1975), essentially modified percent similarity values, were
calculated, comparing all three of the canopy extremes to each
other. For comparison, coefficient of community values were cal-
culated between each pair of communities recognized within the
Preserve and also between each pair of communities recognized
in several other floristic studies in north-central Florida (Abbott
1998). By comparing coefficient of community values of our
coastal hydric hammock plots to those of our communities and
of communities recognized elsewhere in the state, quantitative
support may be found either for recognizing the plots as different
floristically-based plant communities or for treating the plots as
part of one highly variable community.
Most of the Preserve was found to be transitional, without
any one of the canopy extremes, although the juniper and Sabal-
dominated canopy type was most abundant. By the time eight
good patches of the coastal hammock had been found, only a
few of the other canopy types had been found. In the end, then,
the first author searched for pine-dominated and mixed hard-
wood-dominated areas until roughly equal numbers of each had
been found.
The 26 areas into which plots were placed had to meet specific
canopy cover criteria. Coastal hammock areas had to have only
Juniperus virginiana and Sabal palmetto forming over 90% otf
the canopy cover as determined by site inspection. Pine-domi-
nated and mixed hardwood areas had to have pines or mixed
470 Rhodora [Vol. 102
hardwoods comprising over half the canopy cover. Any forested
areas with infrequent to absent J. virginiana and S. palmetto were
excluded from this portion of the study, as these areas usually
represented swamp forest or mesic to scrubby flatwoods.
Once the plots were placed within the 26 subjectively chosen
forest patches, a random number of paces along a random heading
was used to establish the center point of a 100 m° plot, a circle
with a 5.6 m radius. Every vascular plant species present in each
plot was recorded (for raw data see Abbott 1998).
Since we already had chosen to follow FNAI community clas-
sification, we decided to just compare the similarity between our
plant communities with the similarity between communities rec-
ognized by other workers in north-central Florida (Amoroso and
Judd 1995; Easley and Judd 1993; Herring and Judd 1995; Tan
and Judd 1995). This comparison would provide an estimate of
the consistency between our community delimitations and those
of other workers.
A comparative reference of similarity was created using coef-
ficient of community values (Whittaker 1975) to quantify the flo-
ristic similarity between pairs of communities in our study, and
between communities in other studies. The coefficient of com-
munity value was derived by doubling the number of species
shared between two communities, then dividing by the following
sum: twice the number of shared species plus the number of spe-
cies present in each of the communities that is not shared with
the second community. Ignoring any abiotic factors involved, this
phenetic approach simply gave a measure of overall similarity
based on a modification of the percentage of species shared be-
tween any two plant communities.
Even though the categorization of the plots was based on
dominance of the canopy species, the plots within each canopy-
type category were hardly identical in canopy composition. Sev-
enteen Canopy species were encountered: 16 in mixed hardwood
areas, 7 in coastal hammock areas, and 6 in pine-dominated
areas (see annotated list below). In calculating coefficient of
community values for the plots, we noticed that some of the
plots did not contain the characteristic canopy species. These
small areas were just coincidental gaps in forest patches defined
by the surrounding canopy. For example, a few mixed hardwood
and pine-dominated plots actually lacked juniper and Sabal, al-
though they were present nearby. Quercus laurifolia and Q. vir-
2000] Abbott and Judd—Waccasassa Bay State Preserve 471
giniana were scattered throughout all of the canopy categories.
Their massive size often conveyed an impression of dominance,
but only along inland boundaries did they occur in stands with
more than a few individuals. A few of the mixed hardwood areas
were codominated by several individuals of only one or two
hardwood species, but usually there were a few different species,
with no single species appearing dominant. Areas dominated by
pure stands of juniper and Sabal were extensive only nearest the
coast. Inland, pine stands or scattered hardwoods could usually
be seen nearby. Pines were almost never found as an understory
member of areas not dominated by pines, although mature, relic
pines were often found as isolated clumps of emergent trees in
an otherwise non-pine-dominated canopy. Patches of pine-dom-
inated canopy were only rarely found in the northern Preserve,
and they were most common in the southernmost end of the
southern Preserve.
Many understory species were found only under certain canopy
extremes. A total of 124 understory species was documented in
the 26 plots: 89 species in mixed hardwood areas, 40 of which
were found only there; 53 species in coastal hammock areas, 10
of which were found only there; and 49 species in pine-dominated
areas, 22 of which were found only there. Areas with mixed hard-
woods, then, typically had a greater species richness than other
areas, possibly reflecting the rarity or infrequency of flooding by
salt water in these areas. Pine-dominated areas, although they had
twice as many restricted understory species as coastal hammock
areas, were roughly comparable to coastal hydric hammock in
total species richness. Fifteen understory species were found un-
der all three canopy types. Presumably, these species represent
the most versatile and adaptive species. Nine species were found
only in the mixed hardwood and pine-dominated areas, likely a
reflection of salt intolerance. Twenty-five species were shared by
mixed hardwood and coastal hammock areas, while only three
species were shared by both coastal hammock and pine-domi-
nated areas.
Species richness in an area was often related to factors not
reflected by canopy differences. That is, environmental hetero-
geneity often affected the understory in ways not detectable in
the canopy. Field observations while collecting these data indi-
cated that several of the species were actually restricted to wet
depressions, usually with standing water: 13 species in mixed
472 Rhodora [Vol. 102
hardwood areas, 11 in mixed hardwood and coastal hammock
areas, and 5 in coastal hammock areas. Thus, in the mixed hard-
wood and coastal hammock areas, species richness was enhanced
by environmental heterogeneity due to scattered freshwater pools.
Field observations also indicated that some of the coastal pine-
dominated areas had been invaded by salt marsh species. Nine
salt marsh species were present in association with what appeared
to be a relict canopy, with little or no regeneration of canopy
species. This situation is very similar to the pattern described by
Williams et al. (1997) for relict patches of Sabal palmetto and
Juniperus virginiana.
Seventy-two of the total 124 understory species were restricted
to just one canopy extreme in this study, suggesting a possible
association between various understory species and different can-
opy types. However, 97 species were present in three or fewer
plots, with 51 species present in one plot only. This reflects the
patchy nature of the Preserve’s forest and the relative scarcity of
most plant species. This high number of species restricted to only
one (to three) plot(s) resulted in an inadequate sample size for
determining whether or not there was a significant correlation
between the different understory species and the different canopy
extremes. Given the vast area of the Preserve, the amount of area
covered by the plots was extemely small. Thus, any patterns in
species distribution should not be considered as necessarily re-
flective of all of the Preserve’s coastal hydric hammock
The level of distinctiveness between plant communities rec-
ognized in this study corresponded to that of most communities
recognized by other workers (Table 1; Abbott 1998). All of t
plant communities in this study had coefficient of cine
values (X 100) less than 30, as did almost all of the plant com-
munities recognized by others.
he coastal hydric hammock plots were all more similar to
each other than our plant communities were to each other (i.e.,
they had higher coefficient of community values than were found
between any of the plant communities recognized herein; Abbott
1998). These similarity data support the decision to treat the plots
as variants of a single broadly defined community, coastal hydric
hammock. Since there were no readily discernable differences in
hydrology, microtopography, or soil depth or type between the
plot areas, there was also no reason to recognize the canopy var-
iants as nonfloristically based plant communities.
2000] Abbott and Judd—Waccasassa Bay State Preserve 473
Table 1. Coefficient of community values (< 100) calculated for the Wac-
casassa Bay State Preserve, an estimate of similarity based on shared plant
species. A. Plots in all communities. B. Plots in Coastal Hydric Hammock.
C. Plots in Coastal Hydric Hammock, factoring out the Wet Depression spe-
cies and Tidal Marsh species.
FL CH SW_ FP
A.
Mesic to Scrubby Flatwoods (FL)
Coastal Hydric Hammock (CH) 26
Basin Swamp (SW) 20 29
Freshwater Pools and Wet Depressions (FP) 7 10 19
Tidal Marsh (TM) 10 15 14 20
PC) JC
B.
Pine-dominated Canopy (PC)
Juniper & Sabal-dominated Canopy (JC) 39
Mixed Hardwood-dominated Canopy (MC) 38 ©6660
Pee ee
c.
Pine-dominated Canopy (PC)
Juniper & Sabal-dominated Canopy (JC) 49
Mixed Hardwood-dominated Canopy (MC) 49 62
FLORISTIC METHODS AND RESULTS
Field work was conducted by the first author in the Waccasassa
Bay State Preserve from April 1996 to December 1997. Topo-
graphic maps, soil maps, and aerial photos were used to ensure
adequate, representative coverage of plant community variation.
A compass and a handheld global positioning unit greatly facil-
itated field-efficiency and accuracy, especially given the lack of
trails, the large area, and the relative inaccessibility of much of
the Preserve. Since species richness was generally greatest away
from the coast, it was often convenient and most informative to
walk the inland boundaries, when marked, making occasional
transects toward the coast. On two occasions, an airboat was used
to survey the outer limits of the tidal marsh and island hammocks.
Several canoe trips were also made into the tidal marsh, in ad-
dition to numerous visits on foot.
Representative vouchers were deposited in the University of
Florida Herbarium (FLAS), and a partial duplicate set at Selby
474 Rhodora [Vol. 102
Botanical Gardens Herbarium (SEL). The primary references used
for identification of vascular plants were Wunderlin (1982) and
Clewell (1985), although Cronquist (1980), Godfrey and Wooten
(1979, 1981), Hall (1978), Isely (1990), and Long and Lakela
(1976) were also used. Current taxonomic revisions were con-
sulted whenever possible, as cited on the species list.
A total of 576 vascular species and subspecific taxa was doc-
umented from 353 genera and 116 families. There was | lycopsid,
12 ferns, | cycad, 4 conifers, and 555 angiosperms, 178 of which
were monocots. Sixty-nine nonindigenous species were docu-
mented from the Preserve, most of them from a single right-of-
way access road, Fiber Factory Road. Seventy-three plants were
Levy County records, having never been documented previously
in the county according to Wunderlin et al. (1997). Given that
larger numbers of species have been found in much smaller areas
in Florida (e.g., Herring and Judd 1995), and that many of the
Preserve’s species have actually only been found in the very lim-
ited ruderal areas, it seems rather clear that the Preserve is not
very species-rich. This is likely a reflection of its position as an
extreme coastal strip.
The families with the greatest representation, followed by num-
ber of species, are Asteraceae (77), Poaceae (75), Cyperaceae
(49), Fabaceae (36), Scrophulariaceae (14), Apiaceae (11), Po-
lypodiaceae (10), Rubiaceae (10), Malvaceae (10), and Lamiaceae
(9). The largest genera are Cyperus (15), Panicum (11), Carex
(10), Juncus (8), Paspalum (8), Eupatorium (7), Quercus (7),
Smilax (7), Ipomoea (6), Rhynchospora (6), and Solidago (6).
No vouchers were found for any of the previous studies, prin-
cipally zoological, in the Gulf Hammock area (Jennings 1951;
Pearson 1951; Swindell 1949), although most of their reported
plant species, once nomenclature is updated, are documented
herein or are to be expected in the area. There are also no vouch-
ers from most of the limited, previous botanical work in the Wac-
casassa Bay State Preserve, although an unpublished species list
exists for the Preserve. This unofficial list, on file at the office of
the Division of Recreation and Parks, District 2. in Gainesville,
Florida, was largely based on field identifications during 1986
and 1987 by Dr. David Hall. According to letters on file at the
district office, several of his determinations were actually made
in the FLAS herbarium from material sent in by Don Younker, a
district employee at that time.
2000] Abbott and Judd—Waccasassa Bay State Preserve 475
Personal communication with both D. Hall and D. Younker has
convinced us that they were always aware of being within the
Preserve boundaries and that the identifications were rarely in
doubt. We believe that the species not recollected by us were in
the Preserve and may still be there in rare tiny pockets, mostly
in scrubby flatwoods at the southern end of the Preserve. Thus,
for completeness and for future reference, any species whose
name was clearly traceable to the work of David Hall, yet was
not collected by us (41 species total), was included in the species
list but was not used in any other way in the analyses or descrip-
tions. Any species on the previous unofficial list from the Pre-
serve that was not documented by us, or listed in the Hall and
Younker correspondences on file, was excluded, since, in addition
to the lack of vouchers, there was no indication from where or
from whom the name came. The only vascular plant specimens
found from previous work in the Preserve are those of the second
author, who made a couple of casual collecting trips into the area
in 1980 and 1994. All of the species found by him were found
again by the first author during the course of this study.
Species of special concern or interest. Many species are of
interest in the area as they are either at or near the limit of their
natural ranges in Florida (72 spp.), are notably disjunct (13 spp.),
have a very restricted range in Florida (5 spp.), are endemic or
nearly endemic to Florida (16 spp.), or are listed as commercially
exploited, of special concern, rare, threatened, or endangered in
Florida (23 spp.). An on-line atlas of the vascular flora of Florida
was used for determining species ranges (Wunderlin et al. 1997).
For our purposes, species at their distributional limit reach Levy
County from the north or south but do not extend any farther.
Species near their distributional limit do not extend beyond two
counties along the Gulf coast to the north or south of Levy Coun-
ty. A listing of species at or near their distributional limits is
available in Abbott (1998). The five species with very restricted
ranges in Florida found in the Preserve are Leitneria floridana,
Phaseolus smilacifolius, Phyllanthus liebmannianus ssp. platyle-
pis, Spigelia loganioides, and Ulmus crassifolia. Sixteen Florida
endemics or near-endemics were documented in the Preserve: Ar-
istida patula, Berlandiera subacaulis, Campanula floridana, Car-
ex vexans, Coreopsis leavenworthii, Eupatorium mikanioides, Lo-
belia feayana, Pluchea longifolia, Rhynchosia michauxii, Scutel-
476 Rhodora [Vol. 102
laria arenicola, and Vicia floridana are endemic, while Ageratina
jucunda and Panicum dichotomum var. breve are nearly endemic
to Florida (Muller et al. 19
Twenty-three species were found that have been listed as either
commercially exploited, of special concern, rare, threatened, or
endangered in Florida (Table 2) by Coile (1993), Kral (1983),
and Ward (1979). Many of the listed bromeliads, ferns, and or-
chids are actually quite common in Florida. Perhaps some of
these species would be better treated as potentially commercially
exploited, if protection is indeed necessary. No federally-listed
protected species were found within the Waccasassa Bay State
Preserve (Wood 1996).
ANNOTATED LIST OF VASCULAR PLANTS
The vascular plant species inventoried for Waccasassa Bay
State Preserve are listed in Appendix 1. Some angiosperm family
names and/or circumscriptions here deviate from Wunderlin
(1982); in such cases references are provided and the traditional
family names are still included and are cross-referenced to facil-
itate use of the species list.
The species list is arranged alphabetically by family, genus,
and species, within the context of the larger monophyletic groups
of lycopsids, ferns, cycads, conifers, and angiosperms. Nomen-
clature follows Wunderlin (1982, 1998), unless otherwise indi-
cated in the species list, and Wunderlin (1998) was used for de-
termining exotic status. Forty-one species not found by the first
author but reported by David Hall during the 1980s are also listed
here. Most of Hall’s taxa were reportedly seen in the southern-
most portion of the Preserve, and, if still present, can be consid-
ered rare.
The plant communities in which the species occurred are tidal
marsh, coastal hydric hammock, basin swamp, mesic to scrubby
flatwoods, freshwater pools and wet depressions, and ruderal ar-
eas. There are obviously many transitional areas, and most spe-
cies, especially the more abundant ones, can be found along the
edges of, or in isolated patches within, adjacent communities.
Such transitional areas are not reflected in the species list. Rather,
multiple communities are listed only when a species was ob-
served to occur as a distinctive element in several communities.
For example, Juncus roemerianus, typically a salt marsh species,
2000} Abbott and Judd—Waccasassa Bay State Preserve 477
Table 2. Status classification of vascular plants in the Waccasassa Bay
State Preserve that are listed as commercially exploited (CE), of special con-
cern (S), rare (R), threatened (T), or endangered (E) in Florida (Coile 1993;
Kral 1983; Ward 1 979).
Status and Reference
Species Kral Coile Ward
Acrostichum danaefolium T
Dryopteris ludoviciana T
Shenk we T
exalectris spic E
Ilex cassin Cc
T
T
tT
Leitneria ae
Lobelia cardinalis
Malaxis spicata
Osmunda cinnamomea CE
Phyllanthus liebmannianus ssp. platylepis E
Rha aa um hystrix CE
Sabal n T
ee. re E
Selaginella apoda T
Smile ax smallii
4x
74
anio oides
=
=
a
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47
Zamia floridana Cc
could sometimes be found inland. Mostly this was in association
with other typical salt marsh species, and such sites were consid-
ered as transitional patches of tidal marsh. In a few areas, how-
ever, J. roemerianus occurred in association with more freshwater
species, without associated salt marsh species. In these areas, J.
roemerianus was treated as a distinctive element of the freshwater
pools. Finally, essentially every species within the Preserve could
be found along or near a ruderal area. Our attempt here was to
indicate only the species restricted to ruderal areas and the species
that usually occurred in clearings or forest gaps of adjacent com-
munities, as well as ruderal areas.
As pointed out by Amoroso and Judd (1995), the relative abun-
dance of a plant is subject to its reproductive status, seasonal
478 Rhodora [Vol. 102
variation, population changes from year to year, and the judge-
ment and acuity of the researcher. Thus, abundance values reflect
the first author’s subjective estimate of a plant’s frequency, es-
pecially in comparison to associated species or related species.
This is especially true for the basin swamp and mesic to scrubby
flatwoods communities, because both are rare within the Preserve,
occupying only a very few small areas, and any plant restricted
to either of these communities 1s automatically rare in the whole
Preserve. For all the other communities, a numerical scale (Ap-
pendix 1; modified from Thompson and Wade 1991) was used
as a guide for abundance values.
LIST OF CHAROPHYTES, LIVERWORTS, MOSSES, AND MACROLICHENS
Two charophytes, 24 liverworts, 29 mosses, and 43 macrolich-
ens were documented from the Waccasassa Bay State Preserve
(Appendix 2). Numerous other mosses and macrolichens have
been reported for Levy County and may eventually be found
within the Preserve (see Amoroso and Judd 1995; Breen 1963;
Moore 1968; also collections at FLAS). In the list of taxa, charo-
phytes, liverworts, mosses, and macrolichens are presented sep-
arately, and within each list, taxa are arranged alphabetically by
family and species.
Although bryophyte family relationships have been recently
investigated by Buck and Vitt (1986) we simply followed the
classification schemes used by the authors of the following keys
and floras. Charophytes were identified using Wood (1967). Breil
(1970) was used to identify liverworts, except for Frullania cob-
rensis (Griffin and Breil 1982). Breen (1963) and Crum and An-
derson (1981) were used to identify mosses, with nomenclature
following the latter, unless otherwise indicated. Macrolichens
were identified using Moore (1968), Hale (1979), and Harris
(1995), although nomenclature follows Esslinger and Egan
(1995). Crustose lichens were not included in this inventory, al-
though one or more crustose lichens are present in virtually all
of the lichen voucher specimens. Some of the more distinctive
crustose genera, Haematomma, Pertusaria, Buellia-like, and Gra-
phis-like entities, were observed to be very common in the Pre-
serve, especially on small branches in open sunny areas.
Bryophytes and macrolichens respond to microhabitat features
on a scale much smaller than that of vascular plant communities.
2000] Abbott and Judd—Waccasassa Bay State Preserve 479
Moisture and light intensity are undoubtedly the most important
factors controlling bryophyte and lichen distribution within the
landscape. Thus, a water-loving moss like Fissidens cristatus may
be most abundant in the wettest plant community, such as swamp,
but it can also be found in the drier flatwoods community, given
a suitable moisture regime, such as in a crevice at the shaded
base of a tree trunk. Lichens, such as many Cladonia spp., that
are typically soil-dwelling, can be found on tree trunks under
favorable conditions.
In general, mesic inland portions of the Preserve supported the
greatest number of bryophytes, with a reduction in species rich-
ness and abundance towards the coast. The most common and
widespread bryophytes were: Cheilolejeunea spp., Fissidens cris-
tatus, Isopterygium tenerum, Lejeunea spp., Leucobryum albi-
dum, and Syrrhopodon incompletus. The only bryophyte found
in tidal marsh was Frullania kunzei, on bare branches of Lycium
carolinianum. Soil-dwelling bryophytes were uncommon in the
Preserve and were largely restricted to open sand in flatwoods
(e.g., Bryum pseudocapillare, Ditrichum pallidum), a few raised
hummocks in swamps (e.g., Aneura pinguis, Odontoschisma
prostratum, Pallavicinia lyellii), and raised areas near the base of
trees (most taxa, at least in places). Frequent flooding in the Pre-
serve likely limits the ground diversity, as well as impacting spe-
cles composition on fallen branches, logs, and tree bases. The
only bryophytes consistently found on moist limestone rocks
were Barbula agraria, B. cancellata, and Marchantia domingen-
sis. Typically, in other areas, a succession of different suites of
species are associated with the transition from living trunks to
fallen logs to decomposing debris, but no strong successional
patterns were seen by the first author within the Preserve. Certain
species, such as Leucobryum albidum, Octoblepharum albidum,
and Syrrhopodon incompletus were almost always restricted to
erect Sabal palmetto trunks. Other species, such as Cryphaea
glomerata, Forsstroemia trichomitria, Leucodon julaceus, and
Radula australis, as well as most Frullaniaceae and Lejeuneaceae,
were found primarily on living hardwood and juniper trunks.
Most of the remaining bryophyte species seemed to grow any-
where that was moist enough: bark, fallen branches, logs, and
soil.
Of the lichens, Leptogium, Parmotrema (and other similar-
looking Parmelia segregates), and Usnea species were the most
480 Rhodora [Vol. 102
visually dominant and abundant. In shaded forests, lichens such
as Collema, Leptogium, and Pseudoparmelia, could be found on
bark and over bryophytes, but most lichens were growing on
trunks in open areas or on canopy branches.
Both of the charophytes were submerged aquatics, with the
only other true aquatic being the floating liverwort, Riccia flui-
tans.
Although no specific numerical values were assigned, largely
due to identification uncertainty in the field, subjective abundance
values were used to indicate relative frequency of each charo-
phyte and bryophyte species. Rare species (R) were not seen
more than once or twice. Infrequent species (1) were seen a few
times, while occasional species (O) were seen several times. Fre-
quent species (F) were scattered throughout the Preserve, but
were not consistently common. Abundant species (A) were very
common throughout the forested Preserve. Bryophytes that are
difficult to identify in the field were very likely more common
than indicated here (i.e., the accuracy of abundance values here
is a function of how distinctive the species were in the field).
There was definitely a pronounced bias toward bryophytes within
two meters of the ground. No abundance values are given here
for macrolichens, as accurate species determinations were only
had once detailed laboratory inspection and chemical tests were
carried out. Of the lichens, only some individuals of the Cladonia
species were occasionally found on the ground. All other lichens
were found on tree trunks and branches.
ACKNOWLEDGMENTS. Without the assistance of Dr. Dana Grif-
fin, we could not have included charophytes, bryophytes, or ma-
crolichens in this inventory. Numerous discussions with Dr. Kim-
berlyn Williams helped our understanding of ecological principles
and contributed to the development of the quantitative floristics
portion of this thesis. Dr. Francis “‘Jack’’ Putz is also thanked for
advice on ecological field techniques. We thank herbarium staff,
Kent Perkins and Trudy Lindler, for their assistance with technical
herbarium details. Gerald “‘Stinger’’ Guala 1s thanked for his as-
sistance with plant identification, especially grasses. Personnel of
the Florida Division of Recreation and Parks, District 2, are
thanked for their support. Kelly MacPherson and Dan Pearson
were especially helpful. Harry Mitchell and Nikki Makruski pro-
vided field assistance and keys to access gates. This study was
2000] Abbott and Judd—Waccasassa Bay State Preserve 481
conducted in partial fulfillment of requirements for the M.S. de-
gree of the first author.
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Campbell, and M. E. Barkworth, eds., Grass satis and Evolution.
Smithsonian Institution Press, Washington, DC.
APPENDIX 1
ANNOTATED LIST OF VASCULAR PLANTS OF
WACASASSA BAY STATE PRESERVE
For each species, codes for the communities in which it occurs are listed,
followed by an abundance code (sometimes with supplemental habitat or
abundance information), and collection number(s) of J. R. Abbott, unless
otherwise noted. Voucher specimens are housed in FLAS, with a partial du-
plicate set at SEL. Species that have ‘‘Hall NV” in place of the collection
umber were reported by D. Hall during the 1980s. The communities in
which these species are most likely to be expected are listed. Levy County
records, based on Wunderlin et al. (1997), are indicated by the word ‘‘new
after the collection number(s). An fue (*) denotes non-native species.
The ee. are: tidal marsh (TM), coastal hydric hammock (CH),
basin swamp (SW), mesic to bey eeulaes (PL); Seacrest Piety and
wet depressions (FP), and ruderal areas (RU). Abundance categories are: Rare
(R), | occurrences; ae ane nt (I), S—9 occurences; ed oO. 10-24
occurrences; Frequent (F), = 25 occurrences; and Abundant (A) for contin-
uous occurrence. Abundance values reflect subjective estimates of species’
frequencies.
See text for detailed information on this list.
LYCOPSIDA
SELAGINELLACEAE
Selaginella apoda (L.) Fern. — SW R, on raised hummock,; 998/
FILICOPSIDA
OPHIOGLOSSACEAE
Botrychium biternatum (Sav.) Underw. — TM, Hall NV
OSMUNDACEAE
Osmunda cinnamomea L. — SW R,; 1/0400
POLYPODIACEAE sensu lato (see Pryer et al. 1995)
Acrostichum danaefolium Langsd. & Fisch. —- CH I, SW & FP F; 9654
Asplenium platyneuron (L.) Britt. et al. - CH I, 9935
Dryopteris ludoviciana (Kunze) Small — SW R; 10929
488 Rhodora [Vol. 102
Phlebodium aureum (L.) J. Sm. — CH & SW O, epiphytic on Sabal palmetto:
6,
Pleopeltis i Aacaeee (L.) E. G. Andrews & Windham var. michauxiana
.) E. G. Andrews & Windham — CH, SW & FL E usually epiphytic;
11095 [= Polypodium polypodioides (L.) Watt; Andrews and Windham
1993; Windham 1993]
Preridium aquilinum re Kuhn — FL I; 9/07; new
*Preris vittata L. — RU R, on pe outcrops; 9707
Thelypteris hispidula (Decne.) C. F Reed — CH; Hall NV
T. kunthti (Desv.) C. V. Morton — a & SW I, locally A; 8955, 9/53, 9927,
10179, 10921
T. palustris Schott — SW R; 9129
Vittaria lineata (L.) Sm. —- CH & SW R, eae 9660
Woodwardia virginica (L.) Sm. — SW; Hall N
SCHIZAEACEAE
*Lygodium japonicum (Thunb.) Sw. — RU R, near old homesite; 9/55; new
CYCADOPSIDA
ZAMIACEAE
Zamia oo A. DC. H F; FL I; 8372 [but see Eckenwalder (1980),
Landry (1993), eee (1991), and Ward (1979) for different interpre-
ica of the correct name]
CONIFEROPSIDA
CUPRESSACEAE (incl. TAXODIACEAE; Eckenwalder 1976; Hart 1987; Hart and
Price 1990)
Juniperus virginiana L. var, silicicola (Small) Bailey — CH A; SW I; FL O;
8162 ms 1986)
Taxodium distichum (L.) Rich. — SW O; 9787
PINACEAE
ges elliottii Englem. — FL R; 3]
P. taeda L. - CH & FL E ee - 9181, 10018
ANGIOSPERMAE
ACANTHACEAE
Dicliptera brachiata (Pursh) Spreng. — CH R; 9440
Dyschoriste humistrata (Michx.) Kuntze — CH I; /0/6/, /0349
D. oblongifolia (Michx.) Kuntze — CH R; /0560
Elytraria caroliniensis (J. F Gmel.) Pers. — CH O; 9/68
Ruellia caroliniensis (J. F Gmel.) Steud. — CH F; SW R; 9439, 10/60
2000] Abbott and Judd—Waccasassa Bay State Preserve 489
ACERACEAE (see SAPINDACEAE)
ADOXACEAE (incl. part of CAPRIFOLIACEAE; Judd et al. 1994)
Sambucus canadensis L. — FP R; 10163
Viburnum dentatum L. — CH R; ae 10162
V. obovatum Walt. — CH E locally A; FL I; 9/80, 9438, 9449, 9936, 10401
AGAVACEAE
*Yucca aloifolia L. — CH O, especially on or near coastal islands; $4// (status
as native or introduced is problematic)
AIZOACEAE (incl. TETRAGONIACEAE)
Sesuvium portulacastrum (L.) L. — TM O; 9/30
*Tetragonia tetragonioides (Pall.) Kuntze — TM R; 10546; new
ALISMATACEAE
Echinodorus berteroi (Spreng.) Fassett — FP R; 948/, 9555 [= E. rostratus
(Nutt.) Engelm.
Sagittaria graminea Michx. — FP R; 9453
S. lancifolia L. — FP R; 9035
S. subulata (L.) Buch. — FP R; 9452, 9482; new
ALLIACEAE (Dahlgren et al. 1985; Fay and Chase 1996)
Nothoscordum bivalve (L.) Britt. - RU R, boundary trails; 8384, 10009
AMARANTHACEAE
Amaranthus australis (A. Gray) J. D. Sauer —- TM, SW & FP R 9/88
*A. spinosus L. — RU R; 9233; n
ee, pentandra (Jacq.) Stand]. - “TM R; 10539; new
Blutaparon vermiculare (L.) Mears — ie R; 9849
Chenopodium berlandieri Mog. — TM R; 10540
Tresine diffusa Humb. & Bonpl. ex ie — CH R; //096
Salicornia bigelovii Torr. — TM F; 8929
S. perennis Mill. -TM A; 8400 (= S. virginica L. of most authors; Clemants
1992
Suaeda linearis (Ell.) Mog. — TM F; 9507
AMARYLLIDACEAE (excl. ALLIACEAE and HYPOXIDACEAE)
Crinum americanum L. — SW & FP F; /056/
ANACARDIACEAE
Rhus copallinum L. —- CH & RU R; 9463
*Schinus terebinthifolius Raddi — TM R; 993
Toxicodendron radicans (L.) Kuntze — CH, — & FL F; 8398, 9099
490 Rhodora [Vol. 102
ANNONACEAE
Asimina longifolia Kral — FL R; 9/08 (= A. angustifolia Raf.; Kral 1997)
APIACEAE (= UMBELLIFERAE; incl. ARALIACEAE; Judd et al. 1994: Thorne
1983)
*ApIum cae ail ne ) oo — RU R; 9967; new
Aralia spino 3
Centella astatica a chee - an R:; CH, BS, FL, PP & RU F open wet
areas; 9022
Cicuta aneulet L. — SW R; /0931]
Eryngium baldwini Spreng. — ae R; 9/43
Hydrocotyle umbellata L. — , 9248
H. verticillata Thunb. — ms CH, BS, FL, FP & RU E open wet areas;
S951, 9247
Oxypolis filiformis (Walt.) Britt. var. filiformis — RU R; 10983
Ptilimnium capillaceum Sauk ‘ Be — FP I, 9/76, 10363
Sanicula canadensis L. — CH I
fone arpus aethusae Nutt. ex is - sn I; 8907, 8954, 10395
APOCYNACEAE (incl. ASCLEPIADACEAE; Endress et al. 1996; Judd et al. 1994)
Amsonia tabernaemontana Walt. —- CH & FP R; /0/90 (= A. rigida Shut-
Asclepias lanceolata Walt. — FP R; 9020
A. perennis Walt. — CH & FP O; 8903, 9208
Cynanc hum seca’ Pers. — TM . FL I, 8934, 10733
C. scoparium Nutt. —- TM R; CH F; FL I; 9520
Matelea gonocarpos (Walt.) Shinners — te O; 9184, 10189, 10481
AQUIFOLIACEAE
Ilex cassine L. — SW R; 9093, 9655
I. glabra (L.) A. Gray — FL I; 9697, 10737
I. vomitoria Ait. — CH A; FL O; 9433
ARACEAE (incl. LEMNACEAE, e.g., French et al. 1995; Mayo et al. 1995)
Lemna obscura (Austin) Daubs — FP R; 9554, /0098
Peltandra virginica (L.) Schott & Endl. — SW R; /0925; new
ARALIACEAE (see APIACEAE)
ARECACEAE (= PALMAE)
Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude ex Drude — SW R, lo-
; 9924
Sabal minor (Jacq.) Pers. — CH 1; 8900, 941]
S. palmetto (Walt.) Lodd. ex Schultes & Schultes f. —- TM, CH, SW, FL &
Serenoa repens (W. Bartr.) Small — CH R; FL O; 96/3
2000} Abbott and Judd—Waccasassa Bay State Preserve 491
ASCLEPIADACEAE (see APOCYNACEAE)
ASTERACEAE (= COMPOSITAE)
Acmella oppositifolia (Lam.) R. K. Jansen var. repens (Walt.) R. K. Jansen —
6 [= Spilanthes americana (Mutis ex L.f.) Hieron.; Jansen 1985]
Ageratina jucunda (Greene) Clewell & Wooten — CH R; 965/ (= Eupatorium
jucundum Greene
Ambrosia artemisiifolia L. — RU R; 9563
Aster carolinianus Walt. — CH R; song
A. dumosus L. — CH & RU R; //132
A. subulatus Michx. - TM & CH O; ioe 10987
a tenuifolius L. - TM F; 9597, 98/1
A. tortifolius Michx. — FL R; /0992
Baecharis angustifolia Michx. — TM F; 8/66, 11123
B. glomeruliflora Pers. - TM O; 8/65
B. halimifolia L. - TM, CH, SW, FL & FP E often locally A; 1/085, 11086
Berlandiera subacaule (Nutt.) Nutt. — FL R; 9990, 10563
Bidens alba (L.) DC. var. radiata (Sch. Bip.) R. E. Ballard ex Melchert —
CH R; RU O; 9077
B. bipinnata L. — CH & RU I; 9534, 10327
B. mitis (Michx.) Sherff — FP R; //093
Boltonia diffusa Ell. — RU R; 9457
Borrichia frutescens (L.) DC. — TM FE often locally A; 8909
Carphephorus odoratissimus (J. EF Gmel.) Hebert — FL I; 9688, 10995
Cirsium horridulum Michx. — RU R; 5388
C. nuttallii DC. - CH & RU I; 8933
Conoclinium coelestinum (L.) DC. — CH I, RU O; 9/67, 9415 (= Eupatorium
coelestinum L.
*Conyza bonariensis (L.) Crong. — RU R; /0738; new
C. canadensis (L.) Cronq. var. pusilla ae ) Crong. — RU O;
Coreopsis leavenworthti Torr. & A. Gray — FL O; ae ee eee
Eclipta prostrata (L.) L. — FP R; 9605 a E. alba (L.) Hassk.]
ean he carolinianus Raeusch. — CH I; 10922
elatus Bertol. - CH & FL F; 9474, 1104]
2 nudatus A. Gray — CH & FL; Hall NV
Erechtites hieracifolia (L.) Raf. ex DC. — RU R; 9151, 9236
Erigeron quercifolius Lam. — RU O; 8389, 9968, 10348
E. vernus (L.) Torr. & A. Gray — RU R; 10998
Eupatorium album L. — FL I; 9695, 11038
E. capillifolium (Lam.) Small — TM, FP & RU R; 9639
E. mikanioides Chapman — FL & RU I, /0402; new
E. mohrii Greene — FL R; //003
E. perfoliatum L. - TM & RU R; 9590
E. rotundifolium L. — FL & RU O; 10578
E. serotinum Michx. — FL & RU O; 9637
Euthamia caroliniana (L.) Greene ex Porter & Britt. — FL R; ////8 [= E.
tenuifolia (Pursh) Nutt. ]
Flaveria linearis Lag. - TM & RU F; CH I; 9634, 10/88, 11101
492 Rhodora [Vol. 102
Fleischmannia incarnata (Walt.) R. M. King & H. Rob. — CH R; 9644, 11134
(= Eupatorium incarnatum Walt.)
Gamochaeta pensylvanica (Willd.) Cabrera — RU R; 9950 (= Gnaphalium
pensylvanicum Willd., Nesom 1990)
Helianthus angustifolius L. — FL R; 11039
H. debili M R; [1/089
H. radula (Pursh) Torr. & Gray — FL R; /1037
Heterotheca subaxillaris (Lam.) Britt. & Rusby — FL & RU O; 9623
Iva frutescens L. — TM A; FP 1, 9/90, 1098]
I, microcephala Nutt. — RU R; cae 11087
Lactuca canadensis L. — CH;
L. floridana (L.) Gaertn. — a R: a new
Liatris gracilis Pursh — FL & RU R:; 9627
L. graminifolia (Walt.) Willd. — FL & RU I; 9567, 10993
L. tenuifolia Nutt. — FL & RU R; 9675
Melanthera nivea (L.) Small — CH & RU O; 9548
Mikania cordifolia (L. f.) Willd. — throughout O; 9587
M. scandens (L.) Willd. — throughout F locally A; 9448
Pluchea longifolia Nash — SW & FP O; 95
P. odorata (L.) Cass. — SW & FP O; 9468, 10396
P. rosea R. K. Godfrey — FP O; 97/5
Polymnia uvedalia (L.) L. — CH R; 9450
na cae List ee (Michx.) Ell. — FL R; 89/8
rrhopappus carolinianus (Walt.) ae — RU R; 9969
ia phyllocephala (DC.) R. L. Hartman & M. A. ™ &
CH O, island hammocks; 9566 (= cae ee DC.; Lane
and Hartman 1996)
Rudbeckia hirta L. — RU R; 8964, 9546
R. laciniata L. - RU R; 9
R. triloba L. var. pinnatiloba Torr. & A. Gray — CH R, in open wet area;
Senecio glabellus Poir. — FP 1; 8383
Siuphium astericus L. — FL & RU I, 894]
Solidago gene Mill. — CH I; 9678
S. odora Ait. var. chapmanii (Torr. & A. Gray) Crong. — FL & RU O; 1/006
S. rugosa Rak var. aspera (Ait.) Crong. — RU R; 9477, 11083
S. sempervirens L. — TM & CH F; 9664
S. stricta Ait. - TM & CH ES SW I; 9798
° Pa is El. — RU O; J
onchus asper (L.) Hill — R; 9951
sie ola trilobata (L.) Pruski — RU - soem: new [= Wedelia trilobata
(L.) Hitche.; Pruski 1996]
Verbesina virginica L. —- CH & RU I; 9593
*Vernonia cinerea (L.) Less. — RU:
V. gigantea (Walt.) Trel. — CH & RU " 9222, 9456, 9S15
Xanthium strumarium L. — RU R; 9594,
*Youngia japonica (L.) DC. — RU I; a
2000} Abbott and Judd—Waccasassa Bay State Preserve
AVICENNIACEAE
Avicennia germinans (L.) L. —- T™ I; 9197
BATACEAE
Batis maritima L. — TM E locally A; 98/0
BETULACEAE
Carpinus caroliniana Walt. — CH F; 8960
Ostrya virginiana (Mill.) K. Koch — CH R; 9423
BIGNONIACEAE
Bignonia capreolata L. — CH O; 8478
Campsis radicans (L.) Seem. ex Bureau — CH F; //094
BORAGINACEAE
Heliotropium curassavicum L. — TM R; 9205
Lithospermum tuberosum Rugel ex DC. — CH R; 8390; new
BRASSICACEAE (= CRUCIFERAE)
Cakile lanceolata (Willd.) O. E. Schulz — TM R; /01/66, 10542
Cardamine bulbosa (Schreb. ex Muhl.) Britt. et al. - SW R; 9967
C. pensylvanica Muhl. ex Willd. — FP F; RU I; 9938, 9983
*Coronopus didymus (L.) Sm. — RU R; 9957; new
Descurainia pinnata Sia ee a — oa R; 9948
Lepidium virginicum L. — , 994
*Raphanus raphanistrum 2 - oo R; or
BROMELIACEAE
Tillandsia bartramii Ell. - CH O; SWE elphyuc, 9444, 952]
T. recurvata (L.) L. — CH, SW, & FL FE epiphytic
T. usneoides (L.) L. —- TM, CH, SW & FL FE locally abundant, epiphytic;
9061
CACTACEAE
Opuntia stricta (Haw.) Haw. — TM & CH F; 8/75
CAMPANULACEAE
Campanula floridana S. Watson — RU R; 1/0/57
Lobelia cardinalis L. — SW R; 9580
L. feayana Gray — RU R; 10023
L. glandulosa Walt. — FP & RU R; 9790
Triodanis perfoliata (L.) Nieuwl. — RU R; 9962 (Bradley 1975)
CANNACEAE
Canna flaccida Salisb. - SW & RU R; 8959
493
494 Rhodora [Vol. 102
CAPRIFOLIACEAE (see also ADOXACEAE)
Lonicera sempervirens L. — CH I; 8359
Symphoricarpos orbiculatus Moench = CH R; 1/0200
CARYOPHYLLACEAE
Arenaria lanuginosa (Michx.) Rohrb. — CH R, near wet depression and clear-
ing; 10328; new
*A. serpyllifolia L. - RU R: 10007
*Cerastium glomeratum Thuill. — RU R; 9943, 9979; new
*Stellaria media (L.) Vill. — RU R; 9942: new
CELASTRACEAE
Euonymus americanus L. — CH R;
Maytenus phyllanthoides Benth. — a “| island hammocks; 8379, 9930
CELTIDACEAE (Grudzinskaja 1967; Judd et al. 1994)
Celtis laevigata Willd. - CH O; 1/0482
CERATOPHYLLACEAE
Ceratophyllum echinatum A. Gray — FP R; 11097
PODIACEAE (see AMARANTHACEAE)
CISTACEAE
Lechea mucronata Raf. — FL R; 9/38
CLUSIACEAE (= GUTTIFERAE; incl. HYPERICACEAE)
Hypericum cistifolium Lam. — FL R; 9/-
H. hypericoides (L.) Crantz — CH, FL ‘ an I; 9466, 9581, 9702
H. tetrapetalum Lam. — FL R; 9/42
COMMELINACEAE
Commelina diffusa Burm. f. — RU I; 9598; new
C. erecta L. — FL R; 10562
*Murdannia nudiflora (L.) Brenan — RU I; 9253; new
CONVOLVULACEAE
Dichondra carolinensis Michx. — CH, FL & RU, F; 8397, 9973
Evolvulus sericeus Sw. — CH & FL: Hall NV
Ipomoea cordatotriloba Dennst. — RU R; 9465 (= I. trichocarpa Ell.)
NV
riza Michx. — RU R; 9793
i. Ep aes (L.) G. Mey. — CH & RU I; /0340, 10920
2000] Abbott and Judd—Waccasassa Bay State Preserve 495
I, pes-caprae (L.) R. Br. — TM R; 985/
I. sagittata Poir. - TM F; CH & RU O; 9080
CORNACEAE (incl. NYSSACEAE; e.g., Eyde 1988)
Cornus asperifolia Michx. — CH R; SW I; 92/9, 10151, 10394
C. foemina Mill. - CH R; SW I; /0/50
Nyssa biflora Walt. - CH R; SW I; 10484 (Burkhalter 1992)
CUCURBITACEAE
Melothria pendula L. — CH, FL & RU I; 9220, 9636
CYPERACEAE
Bulbostylis stenophylla (Ell.) C. B. Clarke — FL R; 9680
Carex blanda Dewey — CH R; 8380; new
C. chapmannii Steud. —- CH & RU I; /00/4, 10027
C. cherokeensis Schwein. — CH & RU I; 8897, 10199
C. fissa Mack. — FP R; /0/45
C. godfreyi Naczi - CH & RU I; /003/ (Naczi 1993)
C. hyalinolepis Steud. — FP & RU I; 9997, 10185
ey heeds Sartwell ex 7 — SW R; 8949
C. vex FE J. Herm. — SW R; /0/48
ee sp. nov. (sect. Granulares) — CH R; 10029, 10146 (R. Naczi, pers.
comm.)
Carex sp. nov. (sect. tea — CH R; 8382 (R. Naczi, pers. comm.)
Cladium jamaicense Crantz M & FP FE locally A; 9032
Cyperus compressus L. — RU . 9235
C. croceus Vahl — RU R; 9542 (= C. globulosus Aubl.)
C. distinctus Steud. — FP & RU O; 9530
*C. esculentus L. - TM R; 9858
C. flavescens L. — FP & RU R; 9172, 9239, 9956
haspan L. — FP I; 9455
ligularis L. — FP R, brackish water; ///25
odoratus L. — SW & FP F;
planifolius Rich. - TM & FP; Hall NV
polystachyos Rottb. — FP & RU F; 8921, ae 9154, 9536, 10990
retrorsus Chapman — FL & RU O; 9/4/, 921
eC. rotundus L. — RU R; 9960; new
C. stri s L. — RU R; 9230
C. surinamensis Rottb. — RU R; 9224
C. tetragonus Ell. — CH F; 95/0, 9531
C. virens Michx. — FP R; /049]
Eleocharis albida Torr. —- TM R; FP F; 8387, 9/04, 9122, 9418, 9562, 10172
E. atropurpurea (Retz.) J. Pres] & C. Pres] — FP R; 9472
E. baldwinii (Torr.) Chapm. — FP; Hall NV
E. cellulosa Torr. —- TM R; 9596
E. geniculata (L.) Roemer & ag: — FP F; 9062, 9/05
E. montevidensis Kunth — FP R;
Fimbristylis autumnalis (L.) nies & Schultes — RU R; 9701]
SESES TSAO TS
496 Rhodora [Vol. 102
F. caroliniana (Lam.) Fern. — CH R, near salt marsh; //046
F. dichotoma (L.) ae FL & RU O; 9/36, 9250
F. spadicea (L.) Vahl - TM & CH F; &896, 9203, 9523, 9620, 10339, LOSS]
[= F. castanea a ) Vahl]
Futrena breviseta (Coville) Coville — FP R; 9789
*Kyllinga brevifolia Rottb. — RU R; 9240 a Cyperus brevifolius (Rottb.)
dl. ex Hassk.]
K. pumila Michx. — FP & RU I; 9261, 9479 [= Cyperus tenuifolius (Steud.)
Dandy |
Rhynchospora caduca Ell. — FL & RU O; 9027, 10362, 10573;
Rk. colorata (L.) H. Pfeiffer - CH & FP O; RU F; 8966, 9215, 9667, 10353
[= Dichromena colorata (L.) A. S. sete :
R. corniculata (Lam.) A. Gray — SW & F 9458
R. fascicularis (Michx.) Vahl — FL & oo : locally A; 8945, 9101, 9126,
9693, 10740, 11014
R. megalocarpa A. Gray — FL; Hall NV
R. microcarpa Baldw. ex A. Gray — FL & RU O; a 10335
R. miliacea (Lam.) A. Gray — CH & SW R; 9/25, 10487
R. mixta Britt. ex Small — SW & FP; Hall NV
Scirpus californicus (C. A. Mey.) Steud. — FP R; 9204
S. lineatus Michx. — SW R; /0/49
S. robustus Pursh — TM R; 9/97; new
S. tabernaemontani C. C. Gmel. — FP I; /0338 (= S. validus Vahl)
Scleria oligantha Michx. — CH & RU = — 9024
S. triglomerata Michx. — CH & RU F;
S. verticillata Muhl. ex Willd. — FL 1; en 979]
DIOSCOREACEAE
Dioscorea floridana Bartlett — CH R; 9441, 9642
EBENACEAE
Diospyros virginiana L. — CH & FL F:; 9045
ERICACEAE
Bejaria racemosa Vent. — FL R; 9/77
Gaylussacia nana (Gray) Small — FL R; 9698
Lyonia fruticosa (Michx.) G. 8. Torr. — FL R; 8935, 9/12
L. lucida (Lam.) K. Koch — FL R; 8938
Vaccinium arboreum Marsh. — CH & FL I; 8944
V. darrowii Camp — FL; Hall NV
V. myrsinites Lam. — FL R; 9/39
V. stamineum L. — FL R; /1122
EUPHORBIACEAE
Acalypha gracilens A. Gray — CH & RU R; /0984
hea blodgettii wee ex ee ) Small — CH, FL & RU FE often
exposed limestone; 9041,
2000] Abbott and Judd—Waccasassa Bay State Preserve 497
C. hyssopifolia (L.) Small — RU R; 10543
C. maculata (L.) Small — RU R; [0544
C. mesembrianthemifolia (Jacq.) Dugand — TM R; /0545; new
Euphorbia commutata Engelm. ex A. Gray — CH R; 9937
Phyllanthus caroliniensis Walt. — CH; Hall NV
P. liebmannianus Muell. Arg. ssp. platylepis (Small) G. L. Webster — CH I;
101 ie
sae aa) aL. — RU I; 9471;
ee eee vine) oa — RU; Hall NV
FABACEAE (= LEGUMINOSAE)
ponies nla Durazz. — ag R; 9786
Amorpha fruticosa L. — CH, FL & FP O; 89/9, 9201, 10154
Centrosema virginianum (L.) an — FL R; 8922
Cercis canadensis L. — CH R; 9992
Chamaechrista fasciculata (Michx.) Greene — RU O; 9081, 10556
C. nictitans (L.) Moench var. aspera (Muhl. ex Ell.) Irwin & Barneby — RU
R; 9679
Crotalaria rotundifolia J. EF Gmel. — RU O; 10729
*C. spectabilis Roth — RU R; 1/056;
; a
Desmanthus virgatus (L.) Willd. —- FL & RU R; 9089
Desmodium glabellum (Michx.) DC. — RU R; 10994
D. incanum DC. — RU I, 9076, 9540
D. marilandicum (L.) DC. — RU; Hall NV
D. paniculatum (L.) DC. — RU I; 951]
*D. tortuosum (Sw.) DC. — RU R; 96//; new
Galactia elliottii Nutt. - FL R; 10387
G. volubilis (L.) Britt. - CH & FL O; 8952
Gleditsia aquatica Ween — SW R; 8931
G. triacanthos L. — CH I; SW & FP R; 8480, 9060; new
*Kummerowia striata a Schindl. — RU R; 9447
Lespedeza oe lia (Pursh) Ell. — FL; Hail NV
L. hirta (L.) Hornem. — FL; Hall NV
* Medicago ee L. — RU R; 9/40
*Melilotus albus Medik. — RU R: 9097, 9952
*M. indicus (L.) All. — RU R; /0549
Neptunia pubescens Benth. — FL & RU I; 9086
Phaseolus smilacifolius Pollard — CH R; 8472, 9557, 9670, 11136 [This taxon
was considered to be a hybrid by Isely (1990), apparently based on one
sterile specimen. Field observation by the first author and 100% germi-
nation in a greenhouse of 221 seeds from 7 individual plants, with identical
progeny all like the parents, strongly support the recognition of this entity
as a distinct species.
Rhynchosia michauxii Vail — RU R, sandy roadside through salt marsh; //05/
R. minima (L.) DC. — FL & RU I; 9541
498 Rhodora [Vol. 102
Senna marilandica (L.) Link — RU R: /0928
*S. obtusifolia (L.) H. S. Irwin & Barneby — RU R; 92/4, 9427
Sesbania herbacea (Mill.) McVaugh — RU R; 9549 (= S. macroc arpa Muhl.
ex Raf.)
S. vesicaria (Jacq.) Ell. — FP I; RU O; ee
*Trifollum campestre Schreb. — RU R; 9977
Vicia acutifolia Ell. - CH & RU O; 8406, 8492
V. floridana S. Wats. — CH & RU O; 8407, 8467
*V. sativa L. — RU R; 9975
FAGACEAE
Quercus chapmanii Sarg. — i Hall NV
. geminata Small — FL R; 9/47
laurifolia Michx. — CH F: SW & FL O; 945]
michauxit Nutt. — CH R; 92/7
myrtifolia Willd. — FL R; 1/1/27
nigra L. — CH R; 92/8, 943]
pumila Walt. — FL; Hall NV
shumardit Buckl. — CH I; 905]
virginiana Mill. — CH E locally A; FL O; 9//4, 9422
pain ee nea
FUMARIACEAE (see PAPAVERACEAE)
GENTIANACEAE
Eustoma exaltatum (L.) Salisb. ex G. Don — TM, FP & RU O: 9509
Sabatia calycina (Lam.) A. Heller — FP F; RU O: 89/3, 10030
S. stellaris Pursh — FL & FP I; RU O: 89/0, 9021
GERANIACEAE
Geranium carolinianum L. — RU R: 9945
HALORAGACEAE
Myriophyllum pinnatum (Walt.) a et al. — FP O; 9539, 9996: new
Proserpinaca palustris L. — > 89
P. pectinata Lam. — FP: Hall i
HAMAMELIDACEAE
Liquidambar styraciflua L. — CH O; SW F; FL I; 9/52
HIPPOCASTANACEAE (see SAPINDACEAE)
HYDRANGEACEAE (distinct from SAXIFRAGACEAE; é€.g., Morgan and Soltis
Decumaria barbara L. — SW R; 11084
HYDROCHARITACEAE
*Hydrilla verticillata (L. t.) Royle — reportedly in Kelly Creek; Hall NV
2000] Abbott and Judd—Waccasassa Bay State Preserve 499
HYPERICACEAE (See CLUSIACEAE)
HYPOXIDACEAE
Hypoxis curtissii Rose - CH & RU O; 8479, 9429 (= H. leptocarpa Engelm.;
Herndon 1992a, 1992b)
IRIDACEAE
Tris hexagona Walt. — FP F; 8386
Sisyrinchium atlanticum E. P. Bickn. - CH & RU O; 8392, 10025 (This
species is not a laa with S. angustifolium, which does not occur in
Florida; Dan Ward, pers. co
*§. rosulatum E. P. Bickn. — oe - 10331, 10332 (incl. S. exile E. PB Bickn.)
JUGLANDACEAE
Carya aquatica (FE Michx.) Nutt. - SW R; 9659
C. glabra (Mill.) Sweet - CH O; SW & FL I; 9/56
JUNCACEAE
Juncus coriaceus Mack. — CH R; RU I; 10195
dichotomus Ell. — CH R, near wet depression; 9238
narginatus Rostk. — FP & RU O; 8936, 8963
megacephalus M. A. Curtis — FP R; 10367
polycephalus Michx. — FP R; 9/71
roemerianus Scheele — TM A; FP R:; /00/7
scirpoides Lam. — FP R; 10741
tenuis Willd. - RU R; 1/0194
SNS SNS SS
=x
JUNCAGINACEAE
Triglochin striata Ruiz & Pavon — TM F; 9477
LAMIACEAE (= LABIATAE; incl. part of VERBENACEAE; Cantino 1992; Thorne
992)
Callicarpa americana L. — CH & FL s 9048
Hyptis alata (Raf.) Shinners — RU I; 9225
*H. mutabilis (Rich.) Brig. — RU R; ae 9583
*Tamium amplexicaule L. — RU R; 9941;
Monarda punctata L. — RU R; 9621
ace coccinea Buc’hoz ex Etl. — CH; Hall NV
S. lyrata L. - CH & RU R; 1002]
Benes arenicola Small — FL R; 969/
eucrium canadense L. — CH O; 9/85
Trichostema dichotomum L. — FL & RU I; 9464, 9600
LAURACEAE
Persea borbonia (L.) Spreng. — CH R, island hammocks; 9/95
P. palustris (Raf.) Sarg. - CH, SW & FL F; 91/10, 10735
S500 Rhodora [Vol. 1
LEITNERIACEAE (See SIMAROUBACEAE)
LEMNACEAE (see ARACEAE)
LENTIBULARIACEAE
Utricularia foliosa L. — FP R; 9800; new
LINACEAE
Linum medium (Planch.) Britt. — FL & RU R; /0/38, 10352
LOGANIACEAE
| sempervirens (L.) W. T. Ait. - CH & FL I; 9030, 9972
eola petiolata (J. F Gmel.) Torr. & A. bea = a F; 9/74, 9229
uM ee (J. EK Gmel.) G. Don — FP; Hall N
02
al procumbens L. — RU R; 8943, 9246 fetal placement is still
)
n doubt; see Jensen 1992.
Spivelia loganioides (Torr. & A. Gray ex Endl. & Fenzl) A. DC. — CH R;
Judd 2660
LYTHRACEAE
Ammania latifolia L. — FP F; 8975; new
*Cuphea carthagenensis (Jacq.) J. F Macbr. — RU R; 9245
Decodon verticillatus (L.) Ell. — SW; Hall NV
Lythrum alatum Pursh var. lanceolatum (Ell.) Torr. & A. Gray ex Rothr. —
L. lineare L. — TM I, 9427: new
MAGNOLIACEAE
Magnolia hea - — CH O; SW F; FL R; 9/45
M. virginiana L. — 9128
MALVACEAE (incl. BOMBACACEAE, STERCULIACEAE, TILIACEAE; Judd and
Manchester 1997)
Abutilon sabes (Torr. & A. oe Torr. ex A. Gray — RU R; 10397
Hibiscus c s Walt. — FP O;
H. a oe Michx. — FP I; Beet new
Kosteletzkya virginica (L.) C. Presl ex A. — FP I; 1/0407
*Melochia corchorifolia L. — RU; Hall N
Modiola ga aes (L.) G. ae - fs 10403
*Pavonia hastata Cav. — RU R;
Sida homisi L.- RU O; i Tee
*S. L.— RU R; 1/057; new
ie americana L, var. caroliniana (Mill.) Castig. — CH I; 9057, 10020
MARANTACEAE
Thalia geniculata L. — FP R; 1/088; new
2000] Abbott and Judd—Waccasassa Bay State Preserve 501
MENISPERMACEAE
Cocculus carolinus (L.) DC. — RU R, boundary trail; 9043
MORACEAE
Morus rubra L. — CH I; 8463
MYRICACEAE
Myrica cerifera L. var. cerifera - CH, SW & FL E locally A; 10026
M. cerifera L. var. aie Michx. — FL R; /0732 (This entity 1s not usually
given taxonomic recognition, but we point it out here because we think it
may be distinct ve it should be studied in more detail. This entity is a
fire-adapted dwarf shrub restricted to well-drained sandy soils. The habitat
may represent a natural ecological barrier leading to reproductive isolation
from var. cerifera. The two taxa may also be isolated, in part, by different
blooming periods.)
MYRSINACEAE
Rapanea punctata (Lam.) Lundell — CH I, 8366, 9506 (= Myrsine floridana
A. DC.)
MYRTACEAE
Eugenia axillaris (Sw.) Willd. — CH R, island hammocks; 9200
NAJADACEAE
Najas marina L. — TM R, submerged aquatic; 9503; new
NYCTAGINACEAE
Boerhavia diffusa L. — RU R; 1054/
NYMPHAEACEAE
Nymphaea elegans Hook. — FP R; 9480; new
N. odorata Sol. — FP R; 9454
NYSSACEAE (see CORNACEAE)
OLACACEAE
Ximenia americana L. — CH: Hall NV
OLEACEAE
Forestiera ligustrina (Michx.) Poir. - CH O; 9505, 9529, 9796
F. segregata (Jacq.) Krug & Urban — TM O; CH F; 8371, 8925, 10547
Fraxinus caroliniana Mill. — SW I, 10489
F. pennsylvanica Marsh. — CH & SW O; 9054, 10184
502 Rhodora [Vol. 102
Osmanthus americana (L.) Benth. & Hook. f. ex A. Gray — SW R; 9674
ONAGRACEAE
Gaura angustifolia Michx. — RU I; 9096
Ludwigia maritima R. M. Harper — FL I; /0731
ic
Oenothera laciniata Hill — RU R: /0008
ORCHIDACEAE
Epidendrum conopseum R. Br. — CH & SW F; &/70
ee aes floribunda Lindl. — SW R; 9/21 (= H. odontopetala Reichenb.
f.)
Hexalectris spicata (Walt.) Saree ~ R; 9095
Malaxis spicata Sw. — SW
OXALIDACEAE
Oxalts corniculata L. — RU R; 9963
O. florida Salisb. ssp. prostrata (Haworth) Lourt. — RU I, 9243 [Perhaps this
should be treated as Oxalis dillennti ssp. fililpes as suggested by Eiten
(1963), but we await a modern revision. |
PAPAVERACEAE (incl. FUMARIACEAE; Judd et al. 1994; Kadereit et al. 1994,
1995: Loconte et al.1995)
Corydalis micrantha (Engelm. ex A. Gray) A. Gray — RU R; 9949; new
PASSIFLORACEAE
Passiflora lutea L. — CH; Hall NV
P. suberosa L. — CH 1; &46/
PHYTOLACCACEAE
— tolacca americana L. var. rigida (Small) Caulkins & Wyatt —- CH & RU
; 9603 (Caulkins and Wyatt 1990)
PLANTAGINACEAE
*Plantago major L. — RU R; 8957, new
P. virginica L. — RU R; 10/43
PLUMBAGINACEAE
Limonium carolinianum (Walt.) Britt. — TM F; 94/9
POACEAE (= GRAMINEAE)
oo glomeratus (Walt.) Britt. et al. var. glaucopsis (Ell.) C. Mohr —
FL R; ////9 (Campbell 1983)
2000] Abbott and Judd—Waccasassa Bay State Preserve 503
A. glomeratus (Walt.) Britt. et al. var. pumilus (Vasey) Vasey ex L. H. Dewey
— FL & RU F; 9625, 9654, 9797, 11060, 11121, 11124, 11128
. gyrans Ashe var. eee Va C. S. Campb. — FL & RU I; 9704
longiberbis Hackel — FL; Hall N
. virginicus L. var. virginicus — a & RU O; 9692, 9708 (Both old-field
and smooth variants are present.)
Aristida beyrichiana Trin. & Rupr. — FL; Hall NV
. patula Chapman ex Nash — FL & RU R; /0726, 11036 (Allred 1986)
. purpurascens Poir. — FL R; 96/4; new
. spiciformis Ell. — FL I, 9/16
Arundinaria gigantea (Walt.) Walt. ex Muhl. — RU R; 9965
Axonopus fissifolius (Raddi) Kuhlm. — RU I; 9/32, 9209, Wes ee (=A
affinis Chase)
A. furcatus (Fluegge) Hitche. — RU R;
*Bothriochloa pertusa (L.) A. Camus — an R; 9082, 96/6 (These specimens
are atypical, with non-pitted glumes.)
Cenchrus echinatus L. — RU R;
C. incertus M. A. Curtis — CH R; RU O; 9857, 1/052
C. myosuroides Kunth — CH R, open island hammock; 95/8; new
Chasmanthium laxum (L.) Yates — CH & RU F; 9653, 9711
C. nitidum (Baldw.) Yates —- CH & RU F; 896/, 9699
C. sessiliflorum (Poir.) Yates —- CH & oH F; 9669
*Cynodon dactylon (L.) Pers. — RU I, /0/93
Digitaria ciliaris (Retz.) Koel. — FL & nee F; 9622, 10580
*D. violascens Link — RU R; //045
Distichlis ee (L.) Greene — TM A; 9426, 9508
* Echinochloa colona (L.) Link — FP & RU - 9435, 10173; new
*F. crusgalli (L.) P. Beauv. — FP O; 89/6, 9568
E. walteri (Pursh) A. Heller — FP R; 9656; new
*Fleusine indica (L.) Gaertn. — . I, 9226
Elymus virginicus L. HI;
Eragrostis elliottti S. Wats. — ate A RU O; 9588, 11042, 11047
E. hirsuta (Michx.) Nees — FL & RU R; 96/8
E. virginica (Zucc.) Steud. — RU R; 9683, 1/013
*Eremochloa ophiuroides (Munro) Hack. — RU O; 9146, 9544
Eriochloa michauxii (Poir.) Hitchc. — CH R, near wet depression; 979
Eustachys glauca oe — RUF; /0366 [= Chloris glauca (Chapm.) a
E. petraea (Sw.) Desv. — RU F; 95/9 (= Chloris petraea
Leersia hexandra Sw. — FP; Hall NV
L. virginica Willd. — FP & RU R; 9663
Leptochloa fascicularis (Lam.) A. Gray — FP & RU R; 9635
Melica mutica Walt. - CH O, RU R; 8399
Monanthochloe littoralis Engelm. — TM; Hall NV
Muhlenbergia eee (Lam.) Trin. — RU R; 9855, //061]
Oplismenus hirtellus (L.) Beauv. ssp. setarius (Lam.) Mez ex Ekman — CH
F; 9652 (Scholz a
Panicum aciculare Desv. ex Poir. — FL & RU I; /0358, 10569 (Dichanthelium
is treated as a subgenus; Webster 1988; Zuloaga 1986.)
aaa
> > >
504 Rhodora [Vol. 102
P. anceps Michx. — CH & RU F; 9460
P. commutatum Schultes — CH F; 840/, 9719, 9929
P. dichotomiflorum Michx. — RU R; //009; new
P. dichotomum L. — FL & RU F,; 1/0488, 10933
P. ensifolium Baldw. ex Ell. — RU R; /0357
P. gymnocarpon Ell. — FP R; 1/090
P. laxiflorum Lam. — CH O; &477, 9148
P. portoricense Desv. ex Ham. — RU R; //002
*P. repens L. — FP; Hall NV
P. rigidulum Bosc ex Nees — CH & RU F; 9437, 9538, 10932
P. virgatum L. — CH & RU F; 9503, 9514, 9545, 10727, 11011
Paspalum caespitosum Saas - oS Hall NV
*P. dilatatum Poir. — RU R
P. floridanum Michx. — RU . ne 9804, 10399
P. langei (E. Fourn.) Nash — RU O; 9467; new
*P. notatum hie — RU F; /0365, 10581
P. repens Berg. — FP I, 9556, 10479 [= P. oo (Ell.) Kunth]
P. setaceum Michx. — RU O; 9501, 9649; ne
*P. urvillei Steud. — RU O; 9050; new
*Polypogon monspeliensis (L.) Desf. — FP & RU R; 9/73,
Saccharum giganteum (Walt.) Pers. — RU R, near salt oe or 11049
[= Frianthus giganteus (Walt.) Muhl.]
Schizachyrium scoparium (Michx.) Nash — FL R; //048
*Secale cereale L. — RU R; 10/7.
Setaria macrosperma (Scribn. & Merr.) K. Schum. — FL & RU I; 95/3
S. parviflora (Poir.) Kerguelen — CH I; RU F; 9084, 9560 [= S. geniculata
(Lam.) Beauv. |
Sorghastrum elliottii (C. Mohr) Nash — FL O: RU R: 9647, 9668, 9687,
1035; new
Spartina alterniflora Loisel — TM FE locally A; 9196, 9685, 9854, 11031
S. cf. bakeri Merr. — FL I, /0989; sterile
S. patens (Ait.) Muhl. — TM E locally A; 9202
S. spartinae (Trin.) Merr. ex Hitche. - TM O; 95/77; new
Sphenopholis obtusata (Michx.) Scribn. — Ase & RU R; /0028
Eaten indicus (L.) R. Br. - RU R
S. virginicus (L.) Kunth — TM E : Iocally fe aS 10734
es ae sec sedate (Walt.) Kuntze — CH, FL & RU A; 8899
Tridens flavus (L.) Hitehe. — FL & RU I; 9551, 9638, 9703, 11044
Tripsacum dactyloides (L.) L. — RU I; 9039
POLYGALACEAE
Polygala boykinii Nutt. - RU R, near wet depression; /0/55
P. grandiflora Walt. — FL & RU O; 90/8, 10564
P. incarnata L. — FL & RU R; 8926
P. nana (Michx.) DC. — FL R; 8942
2000] Abbott and Judd—Waccasassa Bay State Preserve 505
POLYGONACEAE
Polygonum hydropiperoides Michx. — FP O; 9232, 9478
P. punctatum Ell. — FP O; 10936, 1/001
Rumex verticillatus L. — FP R; 1/0924; new
PORTULACACEAE
*Portulaca amilis Speg. — RU R; 9255; new
POTAMOGETONACEAE
Potamogeton pectinatus L. — FP; Hall NV
PRIMULACEAE
Anagallis minima (L.) E. H. L. Krause — RU R, near salt marsh; 8396 (=
Centunculus minimus L.)
Samolus ebracteatus Kunth — TM, CH & FP F, 8394, 1/0/71
S. valerandi L. ssp. parviflorus (Raf.) Hulten — TM I; CH, SW & FP F;
10170
RANUNCULACEAE
Clematis catesbyana Pursh — CH R, near boundary trail; 964/
C. crispa L. — CH R, near wet depressions; 9040, 9859
RHAMNACEAE
Berchemia scandens (Hill) K. Koch — CH O; 94/2
Sageretia minutiflora (Michx.) C. Mohr — CH : 8358, 58466
ROSACEAE
Crataegus aestivalis (Walt.) pe & A. hake! - R; 9428;
Photinia pyrifolia (Lam.) K. Robertson & J. B. Phipps — FL R, edge of
wet depression; /0004 [= vee sarah ie (L.) Pers. ]
Prunus americana Marsh. — CH R; 9966, 10326
P. serotina Ehrh. — FL R; 9/18
P. umbellata Ell. — CH; Hall NV
Rosa palustris Marsh. — CH R, edge of wet depression; /0/58
Rubus argutus Link — CH R; 9971, 10/42
R. cuneifolius Pursh — CH R; 9103
R. trivialis Michx. — CH O; 10005
RUBIACEAE
Cephalanthus occidentalis L. — FP O; 9059
Chiocecca alba (L.) Hitche. — CH R; 8/73, 9504
Diodia virginiana L. — FP O; RU I; 9249, 9462, 10329
Galium hispidulum Michx. — CH O; 9088, 9502
G. pilosum Ait. — CH R; 10571
G. tinctorium L. — RU I; 10/52
506 Rhodora [Vol. 102
*Hedyotis corymbosa (L.) Lam. — RU R; 9254; new
H. procumbens (J. EF Gmel.) Ds FL; Hall NV
Mitchella repens L. — CH; Hall N
*Mitracarpus hirtus (L.) DC. — ai R; 10345; new [Perhaps the name should
e M. villosus (Sw.) Cham. “ ea Ward 1976.]
Psychotria nervosa Sw. — Hall N
*Richardia brasiliensis Gomez — sa R; 9242; new
Spermacoce assurgens Ruiz & Pavon — RU; Hall NV
*S. prostrata Aubl. - CH & RU R: 9689 [= Borreria ocimoides (Burm. f.)
DC.]
RUPPIACEAE
Ruppia maritima L. — TM F; FP I; 980/, 9989
RUTACEAE
*Citrus aurantium L. — CH R; 9/69; new
Ptelea trifoliata L. — CH R: 8360, 10198
Zanthoxylum clava-herculis L. — CH 1; 10197
Z. fagara (L.) Sarg. — CH; Hall NV
SALICACEAE
Saltx caroliniana Michx. — FP O; 9994, 10016
SAPINDACEAE (incl. ACERACEAE and HIPPOCASTANACEAE; Judd et al. 1994)
Acer rubrum L. — CH 1; SW O; 9037, 9926
A. saccharum Marsh. ssp. floridanum (Chapman) Desmarais — CH F; SW I;
S481, 9065
esculus pavia L. — CH R;
Sanindis saponaria L. — an . en 9516 (incl. S. marginatus Willd.)
SAPOTACEAE
Sideroxylon celastrinum (Kunth) T. D. Penn. — CH R, island hammock; /0548
( telia celastrina H.B.K.; Pennington 1991)
S. lanuginosum Michx. — CH R; 8373, 9553 [= Bumelia lanuginosa (Michx.)
Pers. ]
S. reclinatum Michx. — CH; Hall NV (= Bumelia reclinata Vent.)
SAURURACEAE
Saururus cernuus L. — FP O: 8904
SAXIFRAGACEAE (See HYDRANGEACEAE)
SCROPHULARIACEAE
Agalinis maritima (Raf.) Raf. — o - S920, 9170, 9608, 10554
A. tenuifolia (Vahl) Raf. — FL R;
2000] Abbott and Judd—Waccasassa Bay State Preserve 507
Bacopa monnieri ae Pennell — a FP & RU F; SW I; 9420, 10187
Buchnera americana L. — FL R
Conobea Fa. cowed a - or R; J0192; new [= Leucospora
multifida (Michx.) Nut
Gratiola hispida (Benth. - Lindl. fs eee — FL R; 8940
Linaria canadensis (L.) C 9953
*Lindernia crustacea (L.) ae a, _ ors R; 925], 9599; new
Mecardonia acuminata (Walt.) Small — FP R; 9258
Penstemon mutltiflorus (Benth.) Chapman ex Small — FL R; 10575
Scoparia dulcis L. —- RU R; 9257, 11008
Scrophularia marilandica L. — RU R, boundary trail near access gate; 10982;
new
*Veronica arvensis L. — RU R; 9946; new
V. peregrina L. — RU R; 9947; new
SIMAROUBACEAE (incl. LEITNERIACEAE; Fernando et al. 1995)
Leitneria floridana Chapman — FP F; 8486, 9047, 9445, 9934
SMILACACEAE
Smilax auriculata Walt. — CH F:; SW & FL O; 9058, 9109
S. : H A; SW & FL O; 8405, 9853
S. glauca Walt. — FL R; 9700; new
S. laurifolia L. — CH R; 10477
S. pumila Walt. — FL R; 11120
S. smallii Morong — CH R; 8402
S. tamnoides L. — CH F; 966/
SOLANACEAE
Lycium carolinianum Walt. — TM E locally A; 8/63, 9850
Physalis walteri Nutt. - FL & RU O; 8375, 9092, 10169
Solanum carolinense L. — RU R; 8962, 9234
S. chenopodioides Lam. — RU R; /0550
STERCULIACEAE (See MALVACEAE)
STYRACACEAE
Styrax americanus Lam. — CH R, edge of wet depression, /0024
TILIACEAE (See MALVACEAE)
TURNERACEAE
Piriqueta caroliniana (Walt.) Urban — RU R, boundary trail; 90/7
TYPHACEAE
Typha domingensis Pers. — FP R; 9/3/; new
508 Rhodora [Vol. 102
ULMACEAE (excl. CELTIDACEAE)
Ulmus alata Michx. — CH F; 8496, 94/4
U. americana L. — CH F; SW I; 8376
U. crassifolia Nutt. — CH F; 8484, 9413, 11132
URTICACEAE
Boehmeria cylindrica (L.) Sw. — SW & FP R; 10937
Urtica chamaedryoides Pursh — RU R; 9980; new
VERBENACEAE (see also LAMIACEAE)
ntana camara L. — RU R; 9079, 10164
ae nodiflora (L.) Michx. — TM I; CH, SW, FL, FP & RU F; 89/1] [=
Phyla nodiflora (L.) Greene]
*Verbena brasiliensis Vell. — RU R; /020/, 10552: new
V. scabra Vahl — RU F; 9260, 9467, 10183, 10346
VIOLACEAE
Viola affinis Le Conte — CH & SW O; 1/0022
V. triloba Schwein. — CH & SW I. 9970
VISCACEAE
Phoradendron leucarpum (Raf.) Reveal & M. C. Johnston — CH O; 9925
VITACEAE
Ampelopsis arborea (L.) Koehne — CH & FL O; 10739
Parthenocissus quinquefolia (L.) Planch. — CH i FL O; 10168
Vitis aestivalis Michx. var. aestivalis — CH R; 10
V. lee oo Engelm. ex Millardet var. oe dies Munson — CH I;
9182,
V. ae Michx. var. rotundifolia — CH & FL R; 9/83
V. vulpina L. — CH R, forest gaps; /0/35; new
XYRIDACEAE
Xyris brevifolia Michx. — FL R; 9676
X. caroliniana Walt. — FL R; 1/000
APPENDIX 2
LIST OF CHAROPHYTES, LIVERWORTS, MOSSES, AND MACROLICHENS
OF WACASASSA BAY STATE PRESERVE
Each name is followed by a brief comment on habitat or substrate, an
abundance value abbreviation, and collection number(s) of J. R. Abbott.
Voucher specimens are housed in FLAS. Abundance categories are: Rare (R),
Infrequent (I), Occasional (O), Frequent (F), and Abundant (A). See text for
detailed information on these collections.
2000] Abbott and Judd—Waccasassa Bay State Preserve 509
CHAROPHYTES
CHARACEAE
Chara zeylanica Kl. ex Willd. — two freshwater pools, R; 9443, 9792
Nitella capillata A. Br. — one freshwater pool, attached to floating mats of
Bacopa monnieri, R; 9999
LIVERWORTS (HEPATICAE)
ADELANTHACEAE
Odontoschisma prostratum (Sw.) Trev. — cabbage palm trunks and wet soil
in swamp, R; B-559, B-554
ANEURACEAE
Aneura pinguis (L.) Dum. — wet soil in swamp, R; B-568a, B-582
Riccardia latifrons Lindb. — wet fallen branches in swamp, R; B-56&
R. multifida (L.) S. Gray — wet fallen branches in swamp, R; B-530, B-566
DILAENACEAE
Pallavicinia lyellii (Hook.) S. Gray — wet soil in swamp, R; B-573
FRULLANIACEAE
Frullania cobrensis Gott. ex Steph. — on Taxodium branchlets, R; B-463
F. eboracensis Lehm. — corticolous, R; B-60/
F. kunzeit (Lehm. & Lindb.) Lehm. & Lindb. — corticolous and on branches,
A; B-334, B-486
F. obcordata (Lehm. & Lindb.) Lehm. & Lindb. — corticolous, I; a 621
F. squarrosa (Reinw., Blume & Nees) Nees — corticolous, I; B-
LEJEUNEACEAE
Ceratolejeunea laetefusca (Aust.) Schust. — corticolous, R; B-35/
Cololejeunea cardiocarpa (Mont.) Steph. — on corky Te danibay saplings,
; B-626
Lejeunea cladogyna Evans — corticolous, I; B-570
L. flava (Sw.) Nees — corticolous and on logs, O; B-436
L. laetivirens Nees & Mont. — corticolous, O; B-550, B-595
Leucolejeunea unciloba (Lindenb.) Evans — corticolous, I; B-620
Mastigolejeunea _ (Wils. & Hook.) Schiffn. — corticolous, O; B-433,
B-440, B-591,
Microlejeunea ee (Tayl.) Evans ssp. bullata (Tayl.) Schust. — corticolous,
on branches, and on logs, A; B-607
MARCHANTIACEAE
Marchantia domingensis Lehm. & Lindenb. — on moist limestone, R; B-504,
5
510 Rhodora [Vol. 102
PLAGIOCHILACEAE
Plagiochila dubia Lindenb. & Gott. — base of trees near water, O; B-553, B-
557, B-S69
RADULACEAE
Radula australis Aust. — corticolous, O; B-432, B-4S80, B-556, B-590
RICCIACEAE
Riccia fluitans L. — one freshwater pool, floating aquatic, R; B-6/1
MOSSES (MUSCI)
AMBLYSTEGIACEAE
Amblystegium varium (Hedw.) Lindb. — moist base of tree, R; B-493a
BRACHYTHECIACEAE
Homalotheciella subcapillata (Hedw.) Card. — corticolous, F; B-460
Rhynchostegium serrulatum (Hedw.) Jaeg. & Sauerb. — moist soil at base of
ree, I; B-49]
BRYACEAE
Bryum pseudocapillare Besch. — moist sandy soil, R; B-46/
CALYMPERACEAE
Syrrhopodon incompletus Schwaegr. — primarily on cabbage palm trunks, A;
-477, B-546, D2
S. texanus Sull. — on log near water, R; B-572
CRYPHAEACEAE
ie glomerata BSG. ex Sull — corticolous, O; B-588, B-619
orsstroemia trichomitria (Hedw.) Lindb. — corticolous, O; B-399
DITRICHACEAE
Ditrichum pallidum (Hedw.) Hampe — sandy soil, R; B-6/7
ENTODONTACEAE
Entodon macropodus (Hedw.) C. M. — corticolous and on logs, I; B-445
E. seductrix (Hedw.) C. M. — corticolous and on logs, I; B-349, B-427
FABRONIACEAE
Schwetschkeopsis fabronia (Schwaegr.) Broth. — on logs, I; B-560
2000] Abbott and Judd—Waccasassa Bay State Preserve 511
FISSIDENTACEAE
ReSaenS pee Wils. ex Mitt. — moist base of trees and on logs near water,
, B-478, B-575, B-579
F. ne Hedw. — moist log near water, R; B-527
HYPNACEAE
Isopterygium tenerum (Sw.) Mitt. — corticolous, on logs, and on moist soil,
F; B-350, B-414, B-549
LESKEACEAE
iphones attenuatus (Hedw.) Hueb. — corticolous, I; B-422, B-57/
A, edw.) Schimp. — corticolous, R; B-492
hia oan (Hedw.) Sull. — corticolous, O; B-400, B-489, B-567
LEUCOBRYACEAE
Leucobryum albidum (Brid.) Lindb. — primarily near base of cabbage palm
trunks, F; B-434
Octoblepharum albidum Hedw. — on cabbage palm trunks, R; B-597, B-600
LEUCODONTACEAE
Leucodon julaceus (Hedw.) Sull. — corticolous, O; B-3/2, B-44]
METEORIACEAE
Papillaria nigrescens (Hedw.) Jaeg. & Sauerb. — corticolous, R; B-624
ORTHOTRICACEAE
Schlothemia rugifolia (Hook.) Schwaegr. — corticolous, R; B-438
POTTIACEAE
Barbula agraria Hedw. — moist limestone rocks, I; B-622
B. cancellata C. M. — moist sandy soil and limestone rocks, O; B-420, B-574
SEMATOPHYLLACEAE
Sematophyllum adnatum (Mx.) E. G. Britt. — corticolous, on logs, moist soil,
and moist rocks, O; B-598, B-606
THUIDACEAE
Bryohaplocladium microphyllum (Hedw.) Wat. & Iwats. — moist soil, logs,
and tree bases, I; B-493 [= Haplocladium microphyllum (Hedw.) Broth.]
&C
ne hypnum naa (Hedw.) Buck rum — moist soil, logs, and tree
bases, I; B-546 [= Thuidium ne (Hedw.) BSG.; Buck and Crum
1990
Thuidium delicatulum (Hedw.) BSG. — moist soil, logs, and tree bases, I; B-
544, B-563
512 Rhodora [Vol. 102
MACROLICHENS
CLADONIACEAE
Cladina subtenuis (Abbayes) Hale & Culb. — on a bridge; B-423
piesa oie err. ex Sandst. — on soil; B-63.
. leporina Fr. — on wooden bridge; B-424
C. peziz a (With.) J. R. Laundon — on soil; B-632
C. ramulosa (With.) J. R. Laundon — on soil; B-6/3
C. ravenelii Tuck. — on soil and on bark; B-422
COLLEMATACEAE
Collema furfuraceum (Arnold) Du Rietz var. ee ety Degel.; B-328a
28
. pulchellum Ach. var. leucopeplum (Tuck.) Degel.; B-3
Leptogium austroamericanum (Malme) C. W. Dodee: B-329, B-456, B-562,
B-578
L. azureum (Sw.) Mont.; B-583
L. chloromelum (Sw. ex Ach.) Nyl.; B-448, B-455
L. cyanescens (Rabenh.) Koerber; B-437
L. marginellum (Sw.) Gray; B-395
L. phyllocarpum (Pers.) Mont.; B-398
L. stipitatum Vainio; B-353
PARMELIACEAE
Bulbothrix isidiza (Nyl.) Hale; B-469
Canoparmelia cryptochlorophaea (Hale) Elx & Hale; B-466
a ei le Sai boteriet) Hale; B-320
perforatum (Jacq.) A. Massal.; B-4/8
é ein (Lynge) Hale; owe SO
P. tinctorum (Delise ex ae ) Hale; B-327, B-415
P. ultralucens (Krog) H 3
Pseudoparmelia pe apne (Nyl.) Hale; B-326, B-333, B-457
Punctelia rudecta (Ach.) Krog; B-32
Ramalina i i (Sw.) Ach.; B-585
R. fastigiata (P. Ach.; B-331/, B-335, B-338
R. usnea (L.) - pacers B- 453
R. willeyi R. Howe, B-412, B-413, B-488
Rimelia reticulata are ie Hale & Fletcher; B-462
R. subisidiosa (Muell. Arg.) Hale & Fletcher; B-464
Usnea baileyi es Zahlbr.; B-339
U. mutabilis Stirton; B-602
U. perplectata Mot.; B-346
U. rubicunda Stirton; B-407
U. strigosa (Ach. ) or B-482
U. trichodea Ach. fowl
PHYSCIACEAE
Dirinaria applanata (Fee) D. D. Awasthi; B-470
Heterodermia speciosa (Wulfen) Trevisan; B-330
2000] Abbott and Judd—Waccasassa Bay State Preserve 513
Hyperphyscia syncolla (Tuck. ex Nyl.) Kalb; B-324
Physcia atrostriata Moberg; B-323, B-40/, B-612
P. neogaea R. C. Harris; B-630
Pyxine caesiopruinosa (Tuck.) Imshaug; B-543
STICTACEAE
Lobaria ravenelii (Tuck.) Yoshim.; B-525, B-592
ADDENDUM _ A final site visit on February 13, 2000 yielded the following
additions:
BRASSICACEAE
Rorippa teres (Michx.) Stucky — FP R; 13325
RICCIACEAE
Ricciocarpus natans (L.) Corda — on exposed mud, R; B—S&59
RHODORA, Vol. 102, No. 912, pp. 514-517, 2000
NEW ENGLAND NOTE
FIRST RECORDS OF A EUROPEAN MOSS,
PSEUDOSCLEROPODIUM PURUM,
NATURALIZED IN NEW ENGLAND
NORTON G. MILLER
Biological Survey, New York State Museum, Albany, NY 12230-0001
e-mail: nmiller2 @ mail.nysed.gov
Pseudoscleropodium purum (Hedw.) Fleisch. in Broth. MASSA-
CHUSETTS. Middlesex Co., Mount Auburn Cemetery, Cambridge,
ca. | km south of Fresh Pond: soil, mowed lawn, north-facing
slope under deciduous tree canopy, between Trefoil and Bellwort
paths, | Apr 2000, Miller 129/17 (NYS); on soil in rough lawn in
shade of deciduous trees, above Rose Path, northeast of Tower, |
Apr 2000, Miller 12912 (Nys); soil, sparse lawn near grove of
Picea abies trees, east and southeast and above Dell Pond, 1 Apr
2000, Miller 12910 (Nys); shaded lawn among grass, inside Scots’
Charitable Society Plot enclosure, tree canopy largely deciduous,
13 May 2000, Miller 12985 (NYS, FH). Newton Cemetery, ca. 1.5
km west of Newton Center: damp soil, south side of central pond,
mowed lawn behind Richards crypt under and near large P. abies
trees, 13 May 2000, Miller 12986 (NYS, FH).
Elsewhere is detailed the recent discovery of Pseudosclero-
podium purum in New York, where this moss has been found to
be widespread in the southern part of the state, from near Buffalo
eastward to the Albany area (Miller and Trigoboff, in press). A
large (to 8 cm long), pinnately branched feather-moss, native to
Europe, P. purum had been observed in other parts of North
America prior to its being recognized in New York. These include
Washington State and adjacent British Columbia (especially in
and near Seattle and Vancouver; Lawton 1960; Schofield 1965),
eastern Michigan (near Ann Arbor; Rohrer and Kirkpatrick 1985),
and St. John’s, Newfoundland (Brassard 1983). Lawns, gardens,
and fields are where this moss has been found most frequently in
North America so far, suggesting that its establishment and dis-
persal are closely linked to horticultural practices and lawn care.
The year and method of its introduction into northeastern North
America are unknown.
514
2000] New England Note pe]
My field studies in New York State revealed that Pseudoscler-
opodium purum occurred commonly in regularly mowed ceme-
tery lawns on moist, clayey soil shaded by conifers (particularly
Picea abies and Thuja occidentalis). While I have searched for
this moss at other sites that seemed promising, for example, vil-
lage commons and parks with lawns shaded by P. abies, I have
found it only twice in places other than cemeteries. Both of these
are in Cortland, New York (lawns of the Municipal Water Works
and the City water tower). Thus, in New York at least, P. purum
appears to occur in managed plant communities.
Nowhere in North America, so far, has Pseudoscleropodium
purum been found to produce spores, so it seems likely that its
dispersal takes place when plant fragments are transported by
unknown vectors, ones perhaps associated with lawn care or the
nursery and horticultural trades. Populations of male and female
plants (this moss is dioicous) have been found in New York in
different cemeteries in Rensselaer County, which was intensively
surveyed for P. purum (Miller and Trigoboff, in press), indicating
that a potential exists for its reproduction by spores in the north-
eastern United States. Female plants have been found in Massa-
chusetts in the Newton Cemetery population, but those in Mount
Auburn Cemetery were sexually undifferentiated at the time of
collection.
I evaluated how widespread Pseudoscleropodium purum was
in eastern Massachusetts by surveying the bryoflora of 18 cem-
eteries in Norfolk, Middlesex, and Suffolk counties, in an area of
about 800 km? bounded by the towns of Framingham, Maynard,
Acton, Bedford, Malden, Milton, and Natick. In addition, I ex-
amined parts of the Arnold Arboretum, Jamaica Plain (Boston),
where conifers and mowed lawn occurred together. The inventory
target areas were chosen utilizing the Boston North, Boston
South, Framingham, and Maynard 1:25,000 metric U.S. Geolog-
ical Survey topographic maps. Criteria used to select the sites
included large size and prominence within the community (in
eastern Massachusetts, these two factors often indicate that a cem-
etery was founded in the 1800s), tree cover (green overprint on
the maps), and location (sites more or less evenly scattered
throughout the study area). While turf mosses varied from abun-
dant to sparse in all cemetery lawns I examined, P. purum oc-
curred in only two of them, as indicated above. Moreover, my
search at the Arnold Arboretum was unsuccessful.
516 Rhodora [Vol. 102
Because Pseudoscleropodium purum may eventually spread to
cemeteries and other places in eastern Massachusetts where it
does not now occur, I list here those I searched without success
in May 2000: Norfolk Co.—Milton (Town of Milton), Woodlawn
(Wellesley); Middlesex Co.—Edgell Grove (Framingham), Forest
Dale (Malden), Glenwood (Maynard), Glenwood (Natick), Lake-
view (Wayland), Oak Grove (Medford), Ridgelawn (Watertown),
St. Patricks (Watertown), Shawsheen (Bedford), Sleepy Hollow
(Concord), Westview (Lexington), Wildwood (Winchester),
Woodlawn (Acton); and Suffolk Co.—Forest Hills (Boston). The
success rate, 11% (2 occurrences, 18 searched areas), was about
the same as that obtained in the survey of Rensselaer County,
New York. There, P. purum was found at 14 of 70 sites (20%),
but about twice as much area was surveyed (1800 km’).
In contrast to cemeteries in Rensselaer County, those in eastern
Massachusetts contained fewer small groves of Picea abies, and
therefore were locally less shaded. However, soil differences be-
tween the two regions are perhaps more important. Soil in the
eastern Massachusetts cemeteries I visited was generally well
drained and sandy or loamy, and wind exposure was greater ow-
ing to hilltop, ridge, or slope locations. More acid soils also char-
acterize cemeteries in eastern Massachusetts on the basis of the
absence of bryophytes associated with basic or circumneutral,
calcareous soil. While a search for more stations of Pseudoscler-
opodium purum in eastern Massachusetts could be productive and
should be pursued, the edaphic differences between eastern Mas-
sachusetts and New York habitats may prove to be significant
determinants of different patterns of lawn moss occurrence in
these two regions.
Pseudoscleropodium purum is more widely naturalized in North
America than reported in the most recent regional floras (e.g.,
Crum and Anderson 1981). It 1s uncertain whether this moss will
become invasive in the New York—New England region. It is so
in New Zealand where male and female plants grow together and
spores are produced (Lewinsky and Bartlett 1982). Additional sys-
tematic observation and field surveys are needed in northeastern
North America to track the status and abundance of P. purum.
LITERATURE CITED
BRASSARD, G. R. 1983. Pseudoscleropodium purum in Newfoundland, Can-
ada. J. Bryol. 12: 618-619.
2000] New England Note Si?
Crum, H. A. AND L. E. ANDERSON. 1981. Mosses of Eastern North America.
2 vols. Columbia University Press, York.
Lawton, E. 1960. Pseudoscleropodium purum in the Pacific Northwest. Bry-
ologist 63: 235-237.
Lewinsky, J. AND J. BARTLETT. 1982. Pseudoscleropodium purum (Hedw.)
Fleisch. in New Zealand. cai 8: 177-180.
MILLER, N. G. AND N. TricoBorr. In press. A European feather moss, Pseu-
doscleropodium purum, naturalized widely in New York State in cem-
eteries. Bryologist 104(1).
ROHRER, J. R. AND H. E. KirKPATRICK. 1985. Pseudoscleropodium discovered
the Great Lakes region. Bryologist 88: 24—25.
Sescur te W. B. 1965. Correlations between the moss floras of Japan and
British Columbia, Canada. J. Hattori Bot. Lab. 28: 17-42.
RHODORA, Vol. 102, No. 912, pp. 518-522, 2000
NOTE
LOW CATCHMENT AREA LAKES: NEW RECORDS
FOR RARE COASTAL PLAIN SHRUBS AND
UTRICULARIA SPECIES IN NOVA SCOTIA
NICHOLAS M. HILL
Biology Department, Mount Saint Vincent University,
alifax, Nova Scotia B3M 2J6, Canada
e-mail: NHILL@msvul.msvu.ca
J. SHERMAN BOATES AND MARK E ELDERKIN
Wildlife Division, Department of Natural Resources,
136 Exhibition Street, Kentville, Nova Scotia B4N 4E5, Canada
Nova Scotian wetlands have long been known for their diversity
of Atlantic Coastal Plain plants (Fernald 1921). They are prized
for their local abundances of rare Atlantic Coastal Plain species,
whose diversity is greatest in large catchment area lakes (Hill et
al. 1998). The large annual water level fluctuations (1—2 m) in
lakes of large catchment area (> 50,000 ha) create a wide, ephem-
eral lakeshore habitat where biomass is kept low by prolonged
flood stress in spring, intermittent flooding in summer, and ice
disturbance during winter (Hill et al. 1998). Six of ten species of
plants currently listed as Endangered, Threatened, or Vulnerable in
Canada by COSEWIC (Committee on the Status of Endangered
Wildlife in Canada) occur on the shores of these large catchment
area lakes. These species and sites are the primary focus of con-
servation efforts by the Atlantic Coastal Plain Recovery Team.
The temptation to concentrate conservation efforts on large CA
(catchment area) lakes is undeniable given the clear monotonic
increase in rare species diversity in lakes with increasing CA, but
such large CA chauvinism may not be completely justifiable. This
is because a few notable, rare coastal plain species occur in low
CA (< 2000 ha) lakes. Taschereau (1984) made the first find of
Clethra alnifolia L. in Canada on the shores of a low CA, Nova
Scotian lake and subsequently, naturalist Charlie Allen discovered
a second site at Louis Lake, another headwater lake (Newell
1997). Clearly, the shrub, C. alnifolia, did not obey the same CA
rules that applied to the rare coastal plain herbs and the shrub
518
2000] Note 519
discrepancy was also upheld when Toxicodendron vernix (L.)
Kuntze, new to maritime Canada (Hill 1989), was found along
peaty shorelines of two headwater lakes. With the exception of a
single C. alnifolia individual found at Canoe Lake (10,000 ha
CA) by MacKinnon and Maas (Newell 1997), all recent findings
of C. alnifolia continue to be made at low CA lakes. In this note,
we report finding two additional low CA lake sites for the taxon.
We also report new discoveries of rare floating, coastal plain
plants (at the opposite end of the growth-form spectrum) in the
same low CA sites.
Our field work in 1998 revealed large populations of Clethra
alnifolia in the low CA lakes, Pretty Mary and Mudflat, which
are immediately upstream from populations discovered at Mill
Lake by Leslie Rogers (det. Marian Zinck). Stands were healthy
and cottagers, unaware of the rarity of the shrub, remarked that
they had difficulty eradicating it from cleared areas on their prop-
erties. Growth in these cases appeared to be vegetative only, as
was found by Taschereau for the stands of C. alnifolia on Belli-
veau Lake (Taschereau 1984). However, we collected represen-
tative seedlings from under C. alnifolia stands on Belliveau, Lou-
is, and Pretty Mary Lakes on August 25, 1998, and grew them
in pots for a year, both outside in a cold frame and inside the
greenhouse. Seedlings were identified as Ilex verticillata (L.) A.
Gray, Rhododendron canadense (L.) Torr., and Chamaedaphne
calyculata (L.) Moench at Pretty Mary Lake; Nemopanthus mu-
cronatus (L.) Loes, and /lex verticillata at Belliveau Lake; and
Clethra alnifolia at Louis Lake. This is the first evidence that any
population of C. alnifolia in Nova Scotia can reproduce sexually.
Despite the discovery of seedling recruits at Louis Lake, all
young shoots under parent stems appeared to be vegetative suck-
ers, and even in the main range of C. alnifolia, in New Jersey,
seedling survival in intact woodland appears to be tenuous (Jor-
dan and Hartman 1995).
While lakeshore emergent herb diversity increases with in-
creasing disturbance along a lake CA gradient, shrub diversity is
negatively correlated with fetch, a variable linked to disturbance
through wind energy (Hill and Keddy 1992). The rare shrubs
discussed above occur on the shores of relatively small surface
area, low CA lakes and they grow in a zone essentially free of
ice scour disturbance. In similar fashion, large CA lakes may not
be priority habitats for rare floating plants. In Canada, Utricularia
520 Rhodora [Vol. 102
radiata Small occurs only in lakes in southwestern Nova Scotia,
where it typically grows in 1—3 m deep water in association with
Brasenia schreberi J. F Gmelin. There have been few known
reports of this species in Nova Scotia (Brown 1940; Roland 1976;
Zinck and Roland 1998); the taxon has a global ranking of G4
and in Canada is considered imperilled because of rarity of oc-
currence (6—20 occurrences; Zinck et al. 1994). While investi-
gating new sites for Clethra alnifolia, we found large populations
ot U. radiata in sheltered parts of the two low CA lakes. This
reinforced our realization that while rare coastal plain herb spe-
cies richness is tied to large CA lakes, coastal plain plants of
quite different functional groups (viz., shrubs and floating plants)
may be best represented at the opposite end of the disturbance
gradient, in low CA lakes. Utricularia radiata records include
old herbarium data (ACAD), findings made at Kejimkujic Park
(Roland 1976; updates by Peter Hope) and new findings of our
own over the past two summers (see Appendix). When these
records are put into catchment area classes, it is evident that the
species is more likely to occur in low CA lakes; out of a total of
twenty lake records, eleven were from low CA, two from large
CA, and six from intermediate CA lakes between these extremes.
At the large CA lake sites, the taxon was found in the most
sheltered locations.
Our last low CA lake addition to Nova Scotia’s rare Atlantic
Coastal Plain plants is a white-flowered form of Urricularia pur-
purea Walter, discovered in 1998 at Pretty Mary Lake. There
were extensive mats of this form and none of the typical, lilac-
flowered form, which suggests that regeneration at this site may
be strictly clonal. Utricularia purpurea forma alba has been re-
ported from a pond in New Hampshire where, as in the present
case, only mats of white-flowered plants were present (Hellquist
1974). Vouchers of our specimens are housed at the E. C. Smith
Herbarium at Acadia University (ACAD).
Conservation planning for Atlantic Coastal Plain plants has
benefitted from our knowledge of the relationship between hy-
drology and diversity of rare coastal plain communities in Nova
Scotia (Hill et al. 1998). This model allows us to concentrate field
efforts on lakes of large CA and to make recovery plans for a
suite of rare herbs occurring on these naturally disturbed lake-
shores. Despite the value of this approach in time saving and
habitat acquisition, our new records clearly indicate a need to
2000] Note 521
gather more information on low CA lakes. Further, there is a need
to assess and accomodate the conservation needs in Nova Scotia
for these species in addition to those of the disturbance-linked,
rare coastal plain herbs.
ACKNOWLEDGMENTS. We greatly appreciate the help given by
A. A. Reznicek, P. Hope, and S. P. VanderKloet.
LITERATURE CITED
Brown, M. S. 1940. Utricularia radiata, new to Nova Scotia. Canad. Field-
Naturalist 54: 44.
FERNALD, M. L. 1921. The Gray Herbarium ae to Nova Scotia, 1920.
Rhodora 23: 89-111; 130-171; 184-195; 223-245; 257-278; 284-300.
cae 8 C. B. 1974. A white- ee fant . Utricularia purpurea from
ap eo Rhodora 76: 19.
Hitt, N. M. 89. enti vernix added to the Flora of Nova Scotia.
Rhodora on 242-243.
Keppy. 1992. Prediction of rarities from habitat variables:
Coastal plain plants on Nova Scotian lakeshores. Ecology 73: 1852-—
1859
Keppy, AND I. C. WisHEU. 1998. A hydrological model for
predicting the effects of dams on the oe vegetation of lakes and
reservoirs. rey ronm. Managem. 22: 72:
JORDAN, R. A. ! a M. HARTMAN. 1995. ne sites and the regeneration of
Clethra seas L. eines on in wetland forests of central New Jer-
sey. Amer. Mid. Naturalist 133: 112-123.
NEWELL, R. 1997. Update status site for Sweet Pepperbush, Clethra al-
nifolia. Unpub. manuscript for COSEWIC, Ottawa, Canada.
ROLAND, A. E. 1976. The Coastal Plain Flora of aa National Park.
Report for Kejimkujic National Park, NS, Cana
TASCHEREAU, P. 1984. Status report on the Sweet bibs Clethra alni-
a L., Threatened Species in Canada. Unpubl. manuscript for C
ZINCK, M., K. Covert, R. MELANSON, P. MILLS, R. MILTON, T. POWER, AND
C. UNDERHILL. 1994. Wetland plants of Nova Scotia: Species of concern.
Nova Scotia Dept. Nat. Resources, Kentville, NS, Canada.
. E. ROLAND. 1998. Roland’s Flora of es Scotia. Nimbus
Publishing, Halifax, NS, Canada.
APPENDIX
RECORDS FOR UTRICULARIA RADIATA
The records have been annotated for lakes of three catchment area classes:
low CA (< 2000 ha), intermediate CA (2000—50,000 ha), and large CA (>
50,000 ha). Records were taken from three sources: E. C. Smith Herbarium
WN
Nw
i)
Rhodora [Vol. 102
(ACAD) sheets for the taxon, Kejimkujic National cone records (Roland 1976,
with map updates provided by Peter Hope), a e authors’ findings for
1998-1999. Sites listed on ACAD herbarium en were revisited to re-locate
the taxon in the field.
ACAD Herbarium Records
1. Halifax Co., Sawlor’s L. (low CA), 1940, S. Mason, A. Gorham, and
H. P. Bell; re-located 1998.
2: Tanennare Co., Ashland L. (low CA), 1957, E. C. Smith, A. C. Mac-
onald, and W. J. Curry; re-located 1998.
3. Lanenbrg ae Huey L. dow CA), 1957, E. ee ee A. C. MacDon-
d J. Curry; unable to re-locate in 19
4. Lanenure Co, Lawson L., syn. “‘Larsen L.’ cei CA), 1950;
mith, J. Taylor, D. H. Webster, and L. B. Shipp; re-located 1998.
5. aes he, ea een not recorded on sheet, may refer to Mill
L. (intermediate CA), , E. C. Smith, J. C. Taylor, D. H. Webster,
and L. B. Shipp; re- lop at Mill L. in 1999, see Neu Findings below,
#6
Kejimkujic National Park Records (P. Hope, Chief Interpreter, pers. comm.)
1. Annapolis Co., Kejimkujic L. (large CA).
2. Annapolis Co., Grafton L. aig ae ss stg A. E. Roland.
3. Annapolis Co., Puzzle L. (low CA), A. E. Roland.
4. Annapolis Co., North anal i ae CA).
5. Annapolis Co., Little Peskowesk L. (low CA).
6. Annapolis Co., Loon L. (large CA), R. Belliveau.
7. Annapolis Co., Big Dam L. (intermediate CA), 1976, A. E. Roland.
8. Annapolis Co., Turtle L. dow CA), 1976, T. Bowers.
New Findings
. Lunenburg Co., oe Mary L. (low CA), 1998, J. S. Boates, M. FE
Elderkin, and N Hill.
2. Lunenburg Co., are L. dow CA), 1998, J. S. Boates, M. F Elderkin,
and N. M. Hill.
3 Lunenburg Co., Horseshoe L. (low CA), 1998, P. Mills and J. S. Boates.
* Lunenburg Co., Darling L. (low CA), 1998, P. Mills and J. S. Boates.
. Shelburne Co., Gold L. (low CA), 1998, J. S. Boat
eae: Co., Mill L. (intermediate CA), 1999, N. ML. Hill, M. Myra,
and J. W. Hill.
7. Annapolis Co., Eleven Mile L. (intermediate CA), 1999, N. M. Hill and
J. W. Hill
pt
RHODORA, Vol. 102, No. 912, pp. 523-526, 2000
BOOK REVIEW
Aquatic and Wetland Plants of Northeastern North America by
G. E. Crow and C. B. Hellquist. 2000. Volume 1. Pterido-
phytes, Gymnosperms, and Angiosperms: Dicotyledons, \v +
480 pp. illus. ISBN 0-299-16330-X; Volume 2. Angiosperms:
Monocotyledons, lv + 400 pp. illus. ISBN 0-299-16280-X
$90.00 per volume (hardcover). The University of Wisconsin
Press, Madison, WI.
Fassett’s classic Manual of Aquatic Plants has been the main-
stay of aquatic and wetland plant taxonomy in northeastern North
America for over 60 years. Never revised by the author, the treat-
ment was resuscitated 1n1957 by E. C. Ogden who added a “‘Re-
vision Appendix” in an attempt to “bring the nomenclature into
agreement with present-day usage.’’ Happily, Crow and Hellquist
have accomplished a complete rewrite of the manual from cover
to cover. This thoroughly revised and comprehensive work finally
brings coverage of northeastern aquatics up to speed with other
regions such as the southeast (e.g., Godfrey and Wooten, Aquatic
and Wetland Plants of Southeastern United States) and the south-
west (e.g., Correll and Correll, Aquatic and Wetland Plants of
Southwestern United States).
Like Fassett’s original manual, the geographical coverage of
the new edition extends outside of the United States to include
portions of southeastern Canada, but does not cover extreme
northeastern portions of North America as might be misconstrued
from the title.
Although Crow and Hellquist are accomplished water-plant
taxonomists, they have consulted more than 30 other experts to
achieve current and dependable treatments of all taxonomic
groups. The end result represents more than 20 years of pains-
taking and meticulous taxonomic work by the authors.
The new revision bears the familiar UW trademark on the
spine. The enhanced coverage required publication in two vol-
umes (see above titles), in an 834” X 1114” format that is consid-
erably larger than its predecessor. Blue laminated boards replace
the textured cloth cover of the earlier edition’s most recent print-
ings. All taxon listings are presented in a new two-column format.
These large tomes are not meant to be field guides, and are best
left back at base camp for later referral. The complete set is also
ev)
524 Rhodora [Vol. 102
pricey at $180. The original manual is still available at $30, but
will probably soon be out-of-print permanently. (If you don’t al-
ready own one, you'd be advised to grab one of these to use as
a field book before they disappear.) Understandably, the size and
price of the new manual will deter its use as a textbook or a field
course manual.
Debuting in this edition are a section on “‘Nuisance Aquatic
Plants of the Northeast” and a useful glossary of 64 habitat terms.
The glossary is expanded to 582 terms and a full list of abbre-
viations is provided. The sections on “‘Use of aquatic plants by
birds and mammals”’ and “‘The relation of plants to fish” of the
earlier edition are omitted.
The new manual increases coverage from 752 to 1186 taxa.
Many new illustrations have been added, while keeping most of
the incredibly useful line drawings of the previous edition. Other
enhancements include expanded coverage of salt marsh and bog
species, pteridophytes, and gymnosperms (algae and bryophytes
have been excluded entirely). Treatments have been vastly im-
proved for many difficult genera. For example, for Carex, 76
species are treated as compared to fewer than 30 species in the
previous edition.
A key to “General Keys” is retained, but unlike the cumber-
some 17-choice menu of the previous edition, it is strictly di-
chotomous. Readers are lead efficiently to eight specialized keys
that more or less reflect the major morphological categories em-
phasized by the original book. These keys ultimately lead to fam-
ilies whose locations in the book are identified by volume and
page numbers. All keys have been reworked to avoid confusing
“jumping around” to odd couplets in other keys (a problem with
the original edition).
Keys are indented throughout as compared to the bracketed
format used in the old manual. The indented format is helpful by
enabling the visualization of similar taxa as they are grouped
together. Taxon names appear at the right margin of keys and
include sequential reference numbers for easy location in the ac-
companying text. This feature vastly simplifies the location of
species names and allows one to easily count numbers of taxa in
each family or genus. Species names are now separated from
descriptions and distribution data, which makes it much easier to
locate them in the text.
The keys I tried out were excellent and user-friendly. The par-
2000] Book Review 525
allel leads and non-overlapping character states in couplets are a
hallmark of excellence in key construction. So far, I have not run
into any major problems, although certain keys might still be
improved. For example, the key to Polygonum species still relies
on the traditional distinction between annual versus perennial
growth habit, which is often difficult to determine from specimens
collected without roots. Following the precedent of the original
manual, vegetative features are emphasized in keys whenever
possible, even in problematic groups like Salix and the Lemna-
ceae.
There is a good selection of references and the inclusion of
separate “‘References”’ sections at the back of each volume is a
welcome improvement over citations embedded within the text
as in the old manual. References are not duplicated in the two
volumes. This modification simplifies the location of citations be-
cause one does not have to wade through a single, longer list as
in many two-volume books.
Of course, there are a few typos (e.g., drupelet misspelled as
“druplet”’ in the glossary) and minor errors [e.g., the authorship
of Neobeckia aquatica (Eat.) Greene listed as Neobeckia aquatica
(Eat.) Britt.], but overall this is a well-edited text, especially for
its size and complexity.
I should also mention a few dislikes. Like the earlier edition,
many keys are interrupted by intervening plates of illustrations,
making it necessary to page back and forth when using the key.
The worst instance occurs in the key to Hydrocharitaceae where
the very first couplet is separated by 16 pages of plates (all Sag-
ittaria species, no less). However, most keys were uninterrupted.
I also disliked the redundancy of the general keys, section on
nuisance aquatic plants, list of abbreviations, glossary of plant
terms, glossary of habitat terms, and index in each volume. The
97 pages of identical information (along with 17 entirely blank
pages) comprises roughly 10% of the book. I would have pre-
ferred a shorter, possibly less expensive book. Although this fea-
ture was intended to make each volume “‘stand alone”’ if pur-
chased separately, it does not fully achieve this objective given
that the general keys refer to 53 illustrations from Volume I and
to 33 illustrations from Volume 2 that are not included among
the 18 figures reproduced along with the general keys. Conse-
quently, not all figures in the general keys will be available to
owners of only a single volume.
526 Rhodora [Vol. 102
The single most useful feature of the original Fassett manual
was its detailed line drawings. These are supplemented by many
illustrations carefully selected from appropriate sources. The re-
sult is a thoroughly illustrated work that greatly lessens the bur-
den of water plant identification. However, the quality of printing
used in reproducing the illustrations is mediocre. Many illustra-
tions are printed poorly, with broken lines and/or overly light or
dark print. The quality of several identical illustrations is superior
in the original manual. Although most of the figures are present-
able, the poor clarity of others lessens their utility as an aid to
identification. Furthermore, none of the figures include a scale,
making them less useful for size comparisons than those in the
original manual.
Despite the few printing and format problems, Aquatic and
Wetland Plants of Northeastern North America is more than just
a worthy successor of the old Fassett’s manual. Even though some
parts of the original manual live on in the new one, the many
improvements and innovations have resulted in a unique treat-
ment of North American aquatic plants that is a credit to the
authors. The taxonomic quality of this fine work earns a top rank-
ing among the aquatic floras written for North America. This
book is an essential resource and will surely become the new
standard for aquatic plant taxonomy in the region. Hats off to
Crow and Hellquist for providing us with a new classic that Fas-
sett himself would envy.
—DOoNALD H. Les, Department of Ecology and Evolutionary Bi-
ology, University of Connecticut, U-43, Storrs, CT 06269-3043.
RHODORA, Vol. 102, No. 912, pp. 527-528, 2000
NEW BOOKS
Annotated Checklist of the Vascular Plants of the Washington—
Baltimore Area. Part I. Ferns, Fern Allies, Gymnosperms, and
Dicotyledons by S. G. Shetler and S. S. Orli. 2000. xv +189 pp.
+ 2 pp. corrections. map of checklist area. (softcover, spiral
bound). [available from the authors, Department of Botany, Na-
tional Museum of Natural History, Smithsonian Institution, Wash-
ington, DC 20560-0166]
Contemporary Plant Systematics, 3rd Edition by D. W. Wood-
land. 2000. xiv + 569 pp. black and white photos, line drawings,
and the latest CD version of the University of Wisconsin’s Photo
Atlas of Vascular Plants. ISBN 1-883925-25-8 $64.99 (hardcov-
er). Andrews University Press, Berrien Springs, MI. [FAX 616-
471-6224; aupress @ andrews.edu]
The Flora of Manitoulin Island, 3rd Edition by J. K. Morton and
J. M. Venn. 2000. 376 pp. 124 color photographs and 996 dot
distribution maps. ISSN 0317-3348 $37.50 (softcover, spiral
bound, outside Canada). University of Waterloo Biology Series
No. 40. Department of Biology, University of Waterloo, Water-
loo, Ont. N2L 3G1, Canada.
Flora of the Northeast: A Manual of the Vascular Flora of New
England and Adjacent New York by D. W. Magee and H. E.
Ahles. 1999. xxxi + 1213 pp. 2433 dot distribution maps and
995 line drawings. ISBN 1-55849-189-9 $69.95 (hardcover). Uni-
versity of Massachusetts Press, Amherst, MA. [available from the
publisher, Box 429, Amherst, MA 01004]
The Vascular Plants of Massachusetts: A County Checklist by
Bruce A. Sorrie and Paul Somers. 1999. xvii + 186 pp. + map
of Massachusetts towns and counties. $10.00 (softcover). Mas-
sachusetts Division of Fisheries and Wildlife, Natural Heritage &
Endangered Species Program, Westborough, MA. [available from
the publisher, Route 135, Westborough, MA 01581; for orders of
30 or more, cost is $8.00 each]
a7
528 Rhodora [Vol. 102
Wildflowers of the Western Great Lakes Region by J. R. Wells,
E W. Case, Jr., and T. L. Mellichamp. 1999. xvi + 284 pp. color
photographs and line drawings. ISBN 0-87737-042-7 $39.00
(hardcover). Cranbrook Institute of Science Bulletin 63. Cran-
brook Institute of Science, Bloomfield Hills, MI. [available from
the publisher, P. O. Box 801, Bloomfield Hills, MI 48303]
RHODORA, Vol. 102, No. 912, pp. 529-531, 2000
NEBC MEETING NEWS
June 2000 Field Trip. On Friday afternoon, Mt. Tekoa in West-
field, Massachusetts, was climbed by 14 members and guests to
examine the effects of repeated fires on a rocky outcrop com-
munity. The mountain is a prominent rocky ridge behind the for-
mer Stratford Paper Company mills, and is a prominent landmark
viewed from the eastbound Massachusetts Turnpike. The ridge
has burned twice in the last six years. The group hiked up through
densely re-sprouting oak, hickory, red maple, chestnut, mountain
laurel, and witch hazel to the summit, where the oak-pitch pine
community was starting to regenerate. Several species of Vaccin-
ium were dominant, and promised good berry picking later in the
season. Highlights of the walk included the ant-dispersed sedge
Carex umbellata, Asclepias purpurascens, and Geranium bick-
nellii (or caroliniana). Thirteen species of Carex were observed.
Blackflies, thunder, lightning, and a brief downpour added ex-
citement to the experience, and presaged the early evening storm
that toppled sugar maples on the Mount Holyoke campus.
June 2000. On Friday evening, Dr. Elizabeth Farnsworth of
Smith College spoke on ‘‘Present and future impacts of invasive
plant species on wetland systems.”’’ She discussed the impacts that
invasive species currently have on wetland systems, and what
impacts they may have in a future of climatic change.
Based on county-level floras and checklists, Massachusetts and
Connecticut currently have the largest number of invasive plant
species (approximately 60 and 5Q) and observed invasions, with
a positive correlation between the number of invasive species and
the size of the human population.
Dr. Farnsworth described her research on Lythrum salicaria in
Connecticut, comparing species diversity and biomass in wetlands
dominated by Lythrum to wetlands that were composed of native
species. Studies showed Lythrum did not suppress diversity of
other plant species, but did reduce biomass, and that Lythrum may
capture resources more efficiently than other species, changing
the wetland’s nutrient and detrital dynamics. Phragmites australis
was studied at two sites where it had been removed by spraying
gylphosate herbicide. Removal was associated with a dramatic
increase in the abundance and diversity of other wetland species,
329
530 Rhodora [Vol. 102
resulting in a wetland species composition similar to undisturbed
freshwater tidal marshes.
Dr. Farnsworth looked at the future of invasive species dynam-
ics, in the probable scenario of continued increases in atmospheric
CO, and other greenhouse gas emissions, particularly since
marshes are currently effective as carbon sinks. She examined the
ways that photosynthesis and water use differ among invasive
(Phragmites and Typha angustifolia) and non-invasive (Leersia
oryzoides and Spartina alterniflora) species of freshwater and salt
marshes, with respect to the seasonal length of effective photo-
synthesis, photosynthetic rates, and nutrient balance. Changing
species composition in wetlands toward dominance of one or two
invasive species will likely alter carbon cycling in wetlands,
which in turn may have a climatic feedback effect.
June 2000 Workshop and Field Trip. On Saturday, five mem-
bers participated in a workshop, offered by Lisa Standley, on the
identification of Carex. Following a lecture and slide presentation,
which included discussion of the best keys and references as well
as important diagnostic features, the group hiked up Bare Moun-
tain in the Holyoke range. Twelve early-flowering woodland spe-
cies of Carex were found, including C. platyphylla, C. albursina,
C. laxiflora, C. digitalis, C. communis, C. albicans, and C. hir-
sutella. The woodland bulrush, Scirpus verecundus was abundant.
A small group journeyed to nearby Lawrence Swamp and Elf
Creek Conservation area to hunt for ferns. Led by the intrepid Don
Lubin, the group searched successfully for a nice population of Ly-
godium palmatum at Elf Creek, but failed to find the reported
Ophioglossum pusillum at Lawrence Swamp. Among highlights of
the natural areas and Caroline Arnold’s garden in North Amherst
were Dryopteris clintoniana, Botrychium, and Selaginella apoda.
July 2000 Field Trip: In the Footsteps of Fernald. George
Newman led a party of 21 Club members and family around the
Gaspé Peninsula of Quebec, from Mont Albert in the central moun-
tains to Grand Riviere on the southeast coast. Merritt Lyndon Fer-
nald botanized the Gaspé from 1902 to 1934. During this period,
143 of the 200 new taxa that he named were based on populations
on the Gaspé. Many of the taxa Fernald named are no longer
recognized as distinct species or varieties, but are thought to be
disjunct populations of variable western or circumboreal taxa. The
2000] NEBC Meeting News 531
area also provided much of the support of his “nunatak” theory
to explain the presence of western and arctic disjuncts. Fernald
was followed on the Gaspé by a series of remarkable Canadian
botanists, including Fr. Marie-Victorin, Jacques Rousseau, Ernest
Lepage, and A. E. Porsild, among others.
The group sampled most of the range of habitats of the Gaspé:
serpentine barrens, Thuja bogs and valleys, beaches, sea cliffs,
scree summits, talus slopes, limestone cliffs, coastal spruce-fir
forests, and wide gravel rivers. Unusual ferns were abundant:
Polystichum scopulinum (a western serpentine endemic), Aspi-
dotis densa, Cystopteris montana, Dryopteris fragrans, and Adi-
antum aleuticum at Mont Albert; Polystichum lonchitis at Cap
Bon Ami in Forillon National Park; Dryopteris filix-mas at many
locations; Cryptogramma stelleri on limestone; and tiny Botry-
chium lunaria. Orchids were also frequently observed. We saw
abundantly blooming Orchis rotundifolia in a Thuja bog in the
Parc Gaspésie. At other locations, particularly on Bonaventure
Island, we saw Platanthera dilatata, P. hyperborea, P. obtusata,
P. orbiculata, Listera cordata, L. convallarioides, Coralorhiza
maculata, C. striata, Goodyera repens, and amazingly, Cypripe-
dium parviflorum var. parvifiorum growing on the dry limestone
cliffs and talus at Cap Bon Ami.
The dry scree summits and ledges of Devonian sandstone and
conglomerates yielded a western montane/alpine flora that in-
cluded Arnica chionopappa, Saxifraga cernua, S. caespitosa, S.
aizoon, Anemone multifida, Erigeron compositus, Hedysarum al-
pinum, Shepherdia canadensis, Elaeagnus commutata, Senecio
multiradiata, Oxytropis, and Astragalus scrupulicola. The wet,
mossy sea cliffs along the north coast harbored Pinguicula vul-
garis, Saxifraga aizoides, Parnassia parviflora, Malaxis brachy-
poda, Polygonum viviparum, and Primula laurentiana. Shale and
cobble sea beaches contained Mertensia maritima, Senecio pseu-
do-arnica, [ris hookeri, Anemone canadensis, and Zygadenus
glauca. The serpentine barrens of Mont Albert were one of the
highlights of the trip, both visually and botanically, with Lychnis
alpina, Armeria labradorica, Artemesia borealis, various Salix
species, Eriophorum russeolum (E. chamissonis), and woodland
caribou. In total, the group recorded 415 species of vascular
plants in 65 families (including 34 species of Carex), and 47
species of ferns and fern allies.
—LISA STANDLEY, Recording Secretary pro tempore.
REVIEWERS OF MANUSCRIPTS
1999-2000
The Editor-in-Chief of Rhodora is grateful to the members of
the editorial staff and to each of the following specialists for their
participation in the review process. The conscientious and thor-
ough review of manuscripts by the reviewers and staff helps to
maintain the quality of this journal.
Lewis E. Anderson
Daniel F Austin
J. Craig Bailey
A. Linn Bogle
Jill L. Bubier
Howard A. Crum
Antont W. H. Damman
Brian Duval
Candace Galen
Arthur V. Gilman
Craig W. Greene
David W. Haines
Ronald L. Hartman
Robert R. Haynes
Kent E. Holsinger
Charles N. Horn
Leslie R. Landrum
David M. Lane
Geoffrey A. Levin
Milan Keser
Mark H. Mayfield
T. Lawrence Mellichamp
Norton G. Miller
Guy L. Nesom
Lorraine Olendzenski
Donald J. Padgett
Cathy A. Paris
Kim M. Peterson
Loy R. Phillippe
Albert B. Pittman
A. A. Reznicek
Paul E. Rothrock
Craig W. Schneider
Paul Somers
Daniel Sperduto
David M. Spooner
Anne W. Stork
William Thomas
Glen Thursby
Irwin A. Ungar
Michael A. Vincent
Alan Weakley
Kerry Woods
Richard P- Wunderlin
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 in-
stances noted below. Names of genera and species should be in italics
or underscored throughout. Do not underline punctuation. All pages
should be numbered in the upper right-hand corner. For guidance in
matters not addressed here, consult the editorial office by phone at
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TITLE, AUTHOR(S), AND ADDRESS(ES): Center title, in capital
letters. Omit authors of scientific names. Below title, include au-
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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
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TEXT: Main headings are all capital letters and centered on one line.
Examples are: MATERIALS AND METHODS, RESULTS, and DIS-
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Summary. Second level headings should be indented, bold, upper and
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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
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ences alphabetically by first author, rather than chronologically. With-
in 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
533
534 INFORMATION FOR CONTRIBUTORS
distribution. Specimen citations should include collector(s) and col-
lection number in italics or underscored, and herbarium acronym in
capital letters. Keys and synonymy for eB ee pee revisions should
be prepared in the style of ““A Monograph of the Genus Malvas-
trum,” S. R. Hill, RHODORA 84: 159-264, 1982. Designation of a
new taxon should carry a Latin diagnosis (rather than a full Latin
description), which sets forth succinctly how the new taxon differs
from its congeners
LITERATURE CITED: All bibliographic entries must be cited in the
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References by a single author precede multi-authored works of same
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TABLES: Tables must be double-spaced. Tables may be continued
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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 102nd 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 subscrip-
tion 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 een
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INDEX TO VOLUME 102
New scientific names are in bold face.
Abbott, J. Richard and Walter S.
Judd. Floristic inventory of the
Waccasassa_ Bay Preserve,
Levy County, Florida. 439-513
Acer platanoides 332-354, sacchar-
um 332-354
Adirondacks 250—276
Aegagropilous Desmarestia aculeata
from New Hampshire. 202—207
(New England Note)
Algae, marine 202—207
Andes, Verbesina 129-141
Angelo, Ray and David E. Boufford.
Atlas of the flora of New England:
Monocots except Poaceae and Cy-
peraceae, 1-119
ANNOUNCEMENT:
New England Botanical Club
Graduate Student Research
Award 380
Aquatic plants 13-14, 25—32, 61-64,
96-115
Aquilegia canadensis 308-326
Aralia racemosa 308—326
Atlantic coastal plain shrubs 518—
522
Atlas of the flora of New England:
Monocots except Poaceae and Cy-
peraceae. 1-119
Atriplex: subgeneric novelties, sec-
tional and subsectional mie ia
and new varietal combinations
Australia, Oleandra 428—438
Bahamas, pollination 392-414
Beaver (Castor canadensis) 175-197
Biological invasions 332—354
Birds, pollination reliability, and
green flowers in an endemic island
shrub, Pavonia oo (Mal-
vaceae). 392
Boates, J. eas 518-522
Bolivia, Genlisea and Utricularia
BOOK REVIEW
Aquatic ee Wetland Plants of
Northeastern North America.
Volume 1. Pteridophytes, Gym-
nosperms, and vee aes
Dicotyledons; Volume 2
eae MoncevGle tone
526
Guide to the Algae of New Eng-
land as Reported in the Litera-
ture from 1829-1984, Parts I
and II. A, 225-226
Thoreau’s Country: Journey through
a Transformed Landscape. 227—
229
Boufford, David E. 1-119
Breeding system 392-414
Burk, C. John 154—174
Callery pear 361-364
Callery pear (Pyrus peek ener iasg
saceae) cae ran ee in North Car-
olina. 361—364 (N
Caltha palustris 308— 306
Caryophyllaceae 214—216
Centennial Symposium: The dynam-
ics of the New England flora. 241—
360
Chenopodiaceae, Atriplex 415—427
Chloromonas spp. 365-372
Chlorophyta 365—372
Clethra alnifolia 518-522
Climate and vegetation 246-247,
248-249
Clonal plant,
dense 142—
Closing remarks. 355—
Cogbill, Charles V. are of the
presettlement forests of northern
England and New York.
250-276
Colu
Maianthemum = cana-
153
mbia, ae 138-140
Connecticut
Connecticut Pee 154-174
Conservation 243-245, 299-33]
Corynephorus canescens 208-209
Crow, Garrett E. 217-224
Dawes, Clinton J. 202—207
536
2000]
Dedication 241—242
Desmarestia aculeata 202—207
Distribution maps, New England
monocots 33-115; Chloromonas
367
Drayton, Brian and Richard B. Pri-
mack. R
Drew Forest ‘Preserve, Madison, NJ
332-354
Duval, Brian and Ronald W. Hoham
Snow algae in the northeastern
U.S.: Photomicrographs, observa-
tions, and distribution of Chloro-
monas spp. (Chlorophyta). 365-—
372 (New England Note)
Dwyer, Marc 332-354
Dynamics of the New England flora.
The, 241-360
Ecuador, Verbesina 129-138
Edge effects 332-354
Edible fruit, Jaltomata 385-391
Elderkin, Mark F 518-522
Endangered species 198-201, 210—
213
Endemic plant, pollination of 392—
414
Ferns, Oleandra 428—438
Fifty years of change in Rhodora and
the New England flora. 277
First records of a European moss,
Pseudoscleropodium purum, natu-
ralized in New England. 514-517
(New England Note)
Floodplain vegetation 154—174
Flora of New England, dynamics
241-354; specific discussions
250-276, 280-298
Flora, New England 1-119; swamp
forest 154-174; beaver wetlands
175-197; Florida 439-513
Florida 439-513
Floristic inventory of the Waccasassa
Bay State Preserve, Levy County,
Florida. 439-513
Forest ecology 332-354
Foster, David R. Linking the deep
Index to Volume 102
537
and recent past to the modern New
England landscape. 278-279
Ganger, Michael T. Do reproductive
history traits relate to seed matu-
ration in a clonal herb? 142—153
Genlisea guianensis 217-224
Globally imperiled species 198—201
Globally rare species 210—213
Green flowers 392—414
Haines, Arthur. Rediscovery of Sym-
pPhyotrichum anticostense in_ the
United States. 198-201 (New Eng-
land Note)
Hammond Woods, Newton, MA
308—319
Hebecladus sinuosus, transferred as
Jaltomata sinuosa 385-391
Hedyotis caerulea 308-326
Hehre, Edward J. 202—207
Heliantheae 129-141
Herbaceous flora 154—174
Hevner, Scott J. 385-391
Hill, Nicholas M., J. Sherman
Boates, and Mark EF Elderkin. Low
Utricularia species in Nova Sco-
tia. 518-522 (Note)
Historical ecology 250—276, 280—
298
Hoham, Ronald W. 365-372
vegetation dynamics of the lower
strata of a western Massachusetts
oxbow swamp forest. 154-174
Hudson, W. Donald Jr. Closing re-
marks. 355-360
Hummingbird pollination 392—414
Hlinois 214-216
Immigration and expansion of the
England flora. 280-298
Implications of post-glacial changes
in climate and vegetation on the
flora of the White Mountains, New
Hampshire. 248-249
Introduced species 280-298, 514-
ot Wd
Invasive species 332—354
Nn
eS)
oO
Island pollination 392—414
Jacobson, George L. Jr. Post-glacial
changes in vegetation and climate
in northern New England. 246—
247
nce 385-391; hunzikeri, sp.
dv. 385-388; sinuosa, comb.
nov. 388-390; dentata 390
Judd, Walter S. 439-—S132
Juncaceae 9-13, 49-60
Kaunzinger, Christina K. 332—354
Leiva G., Segundo 385-391
Lentibulariaceae 217-224
Lilies 14-19, 64-78
pare the deep and recent past to
the modern New England _ land-
sc se 278-27
Lobelia cardinalis 308-326
Bo term vegetation yas of
e lower strata of a western Mas-
ee oxbow nie forest.
154-174
Lortie, John P. 210-213
Low catchment area lakes
cords for rare coastal cee shrubs
and Utricularia species in Nova
Scotia. 518—522 (Note)
Maianthemum canadense 142—153
Maine 1-119, 198-201, 208—209
: 154-174,
Mathieson, Arthur C., Edward J.
Hehre, and Clinton J. Dawes. Ae-
gagropilous Desmarestia aculeata
New Hampshire. 202—207
(New England Note)
McLain, David 154—174
McMaster, Nancy D. 175-1
Mehrhof, Leslie J. Immigration and
— of the New England flo-
ra. 280-298
ee in Medford, MA 308—
Mil Norton, G. First records of a
uropean moss, Pseudoscleropo-
ree purum, naturalized in New
Rhodora
[Vol. 102
England. 514-517 (New England
Note)
oe es mas, Segundo Leiva G.,
R. Smith, and Scott J. Hevner.
At new species, a new combina-
tion, and new synonymy for South
American Jaltomata (Solanaceae).
385-39]
Myth of the Scoeele forest: Case
study of the invasive Norwi _
aS (Acer serie) The, 332-
354
Naturalized species, North Carolina
—364; New England 514-517
ee Meeting News 120-125,
230-236, 373-379, 529-53]
Nectar production 392—414
Nesom, Guy L. Callery pear (Pyrus
calleryana—Rosaceae) naturalized
in North Carolina. 361—364 (Note)
New Books 527-528
New England flora 1-119, dynamics
of 243-354, naturalized species
514-517
New Hampshire 1-119, 202-207,
210-213, 248-249, 250-276
New Jersey 332-354
New records for rare coastal plain
shrubs and Utricularia species in
Nova Scotia. 518—52? (Note)
New records for Scirpus ancistro-
chaetus in New Hampshire. 210—
213 (New England Note)
New records, Maine 198-201, 208-—
209; New Hampshire 210-213;
Bolivia 217-224; New England
4-517; Nova Scotia, Canada
518-522
New species of Verbesina from the
northern Andes (Heliantheae; As-
teraceae). Five, 129-141
New species, a new combination,
and new synonymy for South
capes Jaltomata (Solanaceae),
A. 385-391
New species, South America, Ver-
besina 129-141; Jaltomata 385-
391
New York 250—276
Nomenclatural ae in Atriplex
(Chenopodiaceae). 415—427
2000]
Non-indigenous plants 280—298
Non-native plants 280—298; 361-
364; 514-517
North Carolina 361-364
Northeastern U.S. 365—372
Northern hardwood forest 250-276,
332-354
Norway maple (Acer platanoides)
332-354
Notes on the Lentibulariaceae in Bo-
A new genus record (Genli-
sea) for the country, with two ad-
ditional species records in the ge-
nus Utricularia. 217—224 (Note)
Nova Scotia, Canada 518—522
Oleandra 428—438; nertiformis 430—
433; Werneri 433-434; musifolia
434: Wallichii 434-436; undulata
436; Sibbaldti 436-437; ke
Asian, Australian, and Pacific spe-
cies of Oleandra 429-430
Oleandraceae 428—438
Opening remarks: Plant conservation
globally and locally. 243-245
Orchidaceae 19-25, 80—96
Orchids 19-25, 80—96
Osmorhiza eiajion 308-326
Pacific, Oleandra 428—438
Panero, José 129-141
Pavonia bahamensis 392-414
Plant biogeography 250-276
, Florida 439-513
Plant conservation globally and lo-
cally 243-245
Pollen/ovule ratio 392-414
Pollination 392—414
Population re-establishment 299-331
Post-glacial 246-247, 248-249
Post-glacial changes in vegetation
and climate in northern New Eng-
land. 246—247
Potamogetonaceae 25-29, 97-108
Presettlement vegetation 250—276
ard B. 299-331
Pyrus calleryana 361 —364
Rare plants 518—522
Index to Volume 102
232
Rates of success in the reintroduction
by four methods of several peren-
nial plant species in eastern Mas-
sachusetts. 299-331
Rathcke, Beverly J. Birds, pollina-
tion reliability, and green cue
ndemic island shrub,
vonia bahamensis a nated
392-414
Raven, Peter H. Opening remarks:
Plant conservation globally and
locally. 243-245
Rediscovery of Symphyotrichum an-
ticostense he United State
198-201 (New England Note)
Reintroduction methods 299-331
Reproductive biology, Maianthemum
canadense 142-153
Reproductive history traits relate to
seed sepsis in a clonal herb?
Do, 142-153
Restoration ecology 299-331
Rhode Island 1-119
Ritter, Nur P. and Garrett E. Crow.
Notes on the Lentibulariaceae in
Bolivia: A record
(Genlisea) for the country, with
two additional species records 1
the genus Urricularia, 217-224
(Note
Robert T. McMaster and Nancy D.
McMaster. Vascular flora of bea-
ver wetlands in western Massachu-
setts. 175-197
Robinson, Harold and José Panero.
Five new species of Verbesina
from the northern eae eee
theae; Asteraceae).
pes Joshua L. and ne _ a
records for Scirpus ancistro-
pe in New Hampshire. 210—
213 (New England Note)
Rushes 9-13, 49-60
adeina pets atest 214-216;
cey t Illinois species 214
Sagina (Caryophyllace ae) in Illinois:
An update. 214—216 (Note)
Sanguinaria canadensis 308-326
Saracha lobata and S. sordideviola-
, synonymy with Jaltomata
dentata 385-391
Saxtfraga virginiensis 308-326
Scirpus ancistrochaetus 210-213
Seed semper ee can-
adense 142—
Senecio oF oe 209
Smith, Neil R. 385-391
Snow algae in the northeastern U.S.:
Photomicrographs, observations,
and distribution of Chloromonas
spp. ee 65-372 (New
England N
Solanaceae, ae 385-391
South America, le 129-141;
Jaltomata 385—2
Southeast Asia, pind 428-438
vegetation on the flora of the
White Mountains, New Hamp-
shire. 248—249
Statement of Nerd 128
Succession 332—
Surveyor’s Sn ne 276
Swamp forest 154-174
Symphyotrichum anticostense 198—
201
Symposium: The dynamics of the
New England flora 241—360
Systematic notes on the Old World
fern genus Oleandra. 428-438
Tryon, Rolla. Systematic notes on
the Old World fern genus Olean-
38
n C. Sagina (Cary-
ophyllaceae) in Illinois: An up-
date. 214—216 (Note)
Two more weeds in Maine. 208—209
(New England Note)
Utricularia nana 217-223: oliveri-
ana 217-224; known species in
Rhodora
[Vol. 102
Bolivia 220-221; radiata 518—
522; purpurea 518-522
Vascular flora of beaver wetlands in
western Massachusetts. 175-197
Vegetation dynamics 154—174
Vegetation of the presettlement for-
ests of northern New England and
New York. 250-276
Verbesina 129-141; biserrata, sp.
(8)
136-138; perijaensis, sp.
nov. 138-140
Vermont 1-119, 250—276
‘ Bay State Preserve 439—
cussion of com
Nn
erworts, mosses, and macrolichens
508-513
Wagner, Warren H. Jr. Fifty years of
change in Rhodora and the New
England flora. 277
Wagner, Warren Herbert Jr.,
tion 24]—242
Webb, Sara L., Mare Dwyer, Chris-
tina K. Kaunzinger, and Peter H.
Wyckoff. The are of the resilient
forest: Case study of the invasive
oe maple ee er platanoides).
354
dedica-
ees Stanley L. Nomenclatural
el eon in Atriplex (Chenopodi-
aceae). 415—427
— cena) 154-174; beaver
—197
a Mountains, NH 248—249
Witness tree 250—276
Wyckoff, Peter H. 332—354
Zika, Peter EF Two more weeds
Maine. 208-209 (New England
Note)
THE NEW ENGLAND BOTANICAL CLUB
Elected Officers and Council Members for 2000—2001:
President: Lisa A. Standley, Vanasse Hangen Brustlin, Inc., 101
Walnut St., P O. Box 9151, Watertown, MA 02272
Vice-President (and Program Chair): Paul Somers, Massachusetts
Natural Heritage and Endangered Species Program, | Rabbit
Hill Rd., Rt. 135, Westborough, MA 01581
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, Massachusetts College of Liberal Arts, North Adams,
MA 01247-4100
Recording Secretary: W. Donald Hudson, Jr.
Curator of Vascular Plants: Raymond Angelo
Assistant Curator of Vascular Plants: Pamela B. Weatherbee
Curator of Nonvascular Plants: Anna M. Reid
Librarian: Leslie J. Mehrhoff
Councillors: David S. Conant (Past President)
Karen B. Searcy 2001
David Lovejoy 2002
Arthur V. Gilman 2003
Jennifer Forman (Graduate Student Member) 2001
Appointed Councillors:
David E. Boufford, Associate Curator
Janet R. Sullivan, Editor-in-Chief, Rhodora
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