Rhodora

Hovdova

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

Vol. 88 January 1986 No. 853

The New England Botanical Club, Inc.

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Cover Illustration

An original drawing, seemingly the only one surviving and perhaps the only one
ever done for publication by Merritt Lyndon Fernald, used in part to illustrate his
article on cranberry species which appeared in RHODORA No, 48 (Fernald, M. L.
1902. The variations and distribution of American cranberries. Rhodora 4: 231-237
& Plate 40). The drawing was rescued from a wastebasket by Dr. Bernice Schubert; it
now hangs in the office of Dr. Carroll E. Wood at GH. The original Plate 40 caption
reads as follows: Fig. 1, Vaccinium Vitis-Idaea; fig. 2, V. Vitis-Idaea, var. minor; fig.
3, V. Oxyvcoccus; fig. 4, V. Oxvcoccus var. intermedium; fig. 5, V. macrocarpon.

Mhodora

(ISSN 0035-4902)
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 88 January 1986 No. 853

A SYNOPSIS OF NEW HAMPSHIRE SEAWEEDS'”

ARTHUR C. MATHIESON AND EDWARD J. HEHRE?

ABSTRACT

Species composition, phenology, longevity and distribution patterns of New
Hampshire seaweed populations from diverse coastal and estuarine habitats are
given. Two hundred sixteen taxa were recorded (58 Chlorophyceae, 66 Phaeophy-
ceae and 92 Rhodophyceae), including 8 new state and/or geographical records and
the recent introduction of the green alga Codium fragile subsp. tomentosovides to the
Isles of Shoals. Each major group of seaweeds showed similar phenological patterns,
with summer maxima and winter minima. The Rhodophyceae exhibited the greatest
dominance by perennials (67.4%), the Phaeophyceae had an intermediate pattern
(45.5%) and the green algae exhibited the greatest dominance by annuals (87.2%).
Overall, the open coastal sites were dominated by cold temperate species, while warm
temperate or “mixed floras” were more conspicuous in estuarine habitats. Varying
phenological and longevity patterns were also evident in coastal and estuarine habi-
tats. Most of the species (67%) occurred in both open coastal and estuarine habitats,
while 23% were restricted to the open coast and 7% to estuarine habitats. Several
unique distributional patterns were also noted, including contrasting patterns
between closely related taxa, parasitic species and their respective hosts, and different
life history stages of the same species. Several estuarine taxa represent disjunct
populations north of Cape Cod, Massachusetts; they may be relicts of an earlier
“hypsithermal” or warm period. The autecology of several disjunct taxa ts discussed,

Key Words: seaweeds, coastal, estuarine, phenology and distribution, New Hampshire

‘Scientific Contribution Number 1359 from the New Hampshire Agricultural Exper-
iment Station; also issued as Contribution Number 130 from the Jackson Estuarine
Laboratory.

?This paper is dedicated to Dr. Robert F. Scagel on the occasion of his academic
retirement and in recognition of his outstanding and pioneering efforts in marine
phycology, particularly of the Pacific Northwest.

‘Present address: 26 Park Street, South Berwick, Maine 03908

y) Rhodora [Vol. 88

INTRODUCTION

Until recently there have been few published accounts of New
Hampshire (Figures 1-4) seaweeds. Farlow (1882) and Collins
(1900, 1901, 1903 and 1906a) compiled the earliest records for the
state; subsequently, Croasdale (1941) listed a few additional records,
primarily from the Isles of Shoals (Figure 2). Wood and Straughan
(1953) described the penetration of the freshwater red alga Sacheria
fucina (as Lemanea fucina) within the tidal portions of the Oyster
River (Figure 3). Doty and Newhouse (1954) recorded several col-
lections from the Great Bay Estuary System (Figure 3) and noted a
conspicuous decrease in species numbers from the mouth to the
head of the estuary. Taylor (1957) summarized many of the earlier
records for New Hampshire and adjacent New England states in his
excellent account of the benthic marine flora of northeastern North
America.

Since 1965, a variety of floristic, phenological and ecological
studies of New Hampshire’s (Figures |-4) marine algal flora have
been conducted by phycologists at the University of New Hamp-
shire. Hehre and Mathieson (1970) described the species composi-
tion, seasonal occurrence and reproductive periodicity of 88 taxa of
red algae from various open coastal and estuarine environments.
Similar floristic and phenological data were recently summarized on
the Phaeophyceae (Mathieson and Hehre, 1982) and Chlorophyceae
(Mathieson and Hehre, 1983). The seasonal occurrence and vertical
distribution of 125 seaweeds at Jaffrey Point (Fort Stark), New
Castle, New Hampshire were recorded (Mathieson, Hehre and Rey-
nolds, 1981), as well as the distributional patterns of marine algae
within the Great Bay and Hampton-Seabrook (Figures 3 and 4)
estuary systems (Mathieson, 1975; Mathieson and Fralick, 1972;
Mathieson, Reynolds, and Hehre, 1981). Each of the estuarine areas
showed a “typical” reduction pattern inland, as well as the impor-
tance of tidal rapids in determining local and discontinuous distri-
butional patterns. A comparison of the species composition of
seaweeds from the Merrimack River estuary (Figure |), Massachu-
setts, (Mathieson and Fralick, 1973) with that of the Hampton-
Seabrook and the Great Bay estuary systems (see earlier citations)
indicates a paucity of total species and number of taxa/ station in the
Merrimack River estuary—one of the most polluted rivers in New
England (Jerome et al., 1965; Miller et al., 1971). In contrast, tidal

1986] Mathieson & Hehre — New Hampshire seaweeds 3
ae ry i Fhe
N 71°40! 71° 30! ih Cape Elizabeth
~ Kennebunk BH (7 miles } )
‘\ Ss Uetpehe eas.
* ¥ ““ Cape Arundel
N
'
& !
|
\
\
\
\ MAINE
\ Rs
Cape Neddick .
York River ae 43 10' _|
“Great Bay Estuary System
Isles of Shoals
a Dd a
Gulf of
Maine
Coast (Nearshore)
Seabrook: :: MS opisptonsSenbrenk Estuary System
MASS.
[Salisburys~ 7 42°50’ _|
Merrimack BE,
River oe
1 mile
, om
Figure |. The New England coastline between Cape Arundel, Maine and Cape

Ann, Massachusetts, including the four primary coastal-estuarine areas within New

Hampshire.

4 Rhodora [Vol. 88
Duck ° .
of vf
a 43 0O-
Appledore
Maine mutt ynose
i _ Malaga
NH i
% Cedar
Lunging 2
Square Rock
peavey
a . white 42 58-
70 38 70 36

Figure 2. The Isles of Shoals, New Hampshire- Maine.

1986] Mathieson & Hehre — New Hampshire seaweeds >

rapid sites such as Dover Point, which occur within the Great Bay
Estuary System (Figure 3), can exhibit a much greater diversity of
species than the entire Merrimack River (Mathieson, Neefus and
Emerich Penniman, 1983; Mathieson and Fralick, 1973: Reynolds
and Mathieson, 1975). Tidal rapids also exhibit a more diverse flora
than adjacent “back-eddy” sites, and such rapids may represent
major phytogeographic boundaries within estuaries (Mathieson,
Reynolds and Hehre, 1981).

A comparative phytogeographic evaluation of seaweed popula-
tions at the Isles of Shoals (Figure 2), an archipelago of eight major
offshore islands near southern Maine and New Hampshire, showed
that the mean similarity within the Shoals was approximately 82%
and that a significant proportion of the variance in species richness
per island was explained by the length of semi-exposed shoreline on
each island (Mathieson and Penniman, 1986a). Detailed studies of
the subtidal flora of New Hampshire were initiated by SCUBA
diving on the open coast and within the Great Bay Estuary System
(Mathieson, 1975, 1979: Mathieson and Burns, 1970; Mathieson,
Hehre and Reynolds, 1981). The species richness, longevity and
vertical distribution of the subtidal seaweed populations were
related to a variety of environmental parameters, including temper-
ature, salinity, light and water motion. Additional descriptive
accounts of New Hampshire algae have been given by Normandeau
Associates (1971-1980) for the Piscataqua River near the Schiller
Power Plant, by the New Hampshire Department of Fish and Game
(Nelson et al., 1981, 1982) for the Great Bay Estuary System, and by
Daly and Mathieson (1977) at Bound Rock in Seabrook. Several
additional biosystematic (Blair, 1983; Blair et al., 1982), floristic
(Hehre, 1972) and autecological investigations of New Hampshire
seaweeds have been conducted (Burns and Mathieson, 1972a. b;
Chock and Mathieson, 1976, 1983; Daly and Mathieson, 1977:
Hardwick-Witman and Mathieson, 1983; Josselyn and Mathieson,
1978, 1980; Kilar and Mathieson, 1978, 1981: Mathieson, 1979,
1982a; Mathieson and Burns, 1975; Mathieson, Neefus and Emerich
Penniman, 1983; Mathieson, Penniman, Busse and Tveter-
Gallagher, 1982; Mathieson and Prince, 1973; Mathieson, Shipman,
O’Shea and Hasevlat, 1976; Niemeck and Mathieson, 1976: Norall
et al., 1981; Sideman and Mathieson, 1983a, b, 1985: Tveter and
Mathieson, 1976; Tveter-Gallagher and Mathieson, 1980: Tveter-

6 Rhodora [Vol. 88

A.

Great Works River

~ \

r Salmon
Cocheco 7 * Falls River N
River Q
Selisngo" N eal
River 43 10/
Maine

Oyster River

Dover Point
Piscataqua
Little River
Bay .

A\
MY, Lamprey Adams Point

R River NH

db
<< 43°08

— lh o
:
a tae Point

( sQuamscott Winnicutt a :
| River River Atlantic Ocean
( 70° 50’ 70°40’

| | | |

Figure 3. The Great Bay Estuary System and the adjacent open coast of New
Hampshire- Maine.

Gallagher, Mathieson and Cheney, 1980; Zechman and Mathieson,
1985).

In the present account, a synopsis of the Chlorophyceae, Phaeo-
phyceae and Rhodophyceae from coastal/estuarine habitats in New
Hampshire is given, based upon a synthesis of the above-described
collections and data. The phenology, longevity and local distribu-

1986] Mathieson & Hehre — New Hampshire seaweeds 7

Great Boar's
Head

ae
each

a ta ae

Hampton River ‘ oe

‘ey s 3
Brown River a
)
AS a

= E ATLANTIC OCEAN

=

1/2 mile

New Hampshire

~ ~ Massachusetts

Figure 4. The Hampton-Seabrook Estuary System and the adjacent open coast

of New Hampshire and Massachusetts.

tional patterns of each taxon are summarized in a series of detailed
distribution maps. Norton (1978), among others, has emphasized
that distributional maps are significant tools in marine ecology,
particularly if a comprehensive set of environmental data is avail-
able. A detailed synopsis of the New Hampshire coastal zone is
given to aid in such geographical and ecological comparisons.

METHODS AND MATERIALS

As outlined previously, extensive collections and observations of
New Hampshire seaweeds have been made at a variety of open
coastal and estuarine sites (Figure 5, Table IV and Appendix) dur-
ing 1965-83 in order to prepare a detailed synopsis of the state’s
marine algal flora. Thus, collections were made at 212 study sites,

8 Rhodora [Vol. 88

Cocheco

River Salmon

\
Falls River \

Cc Neddick
York River Oe ere >

oy ae

Bellamy
River —
; i \
Oyster sete ‘ag MAINE ~ yy m
ws F
Ko .

Little
Bay

Lamprey
River

Winnicutt
River

NH Open
Coast (Nearshore]

.

S yuaiiaeck .
River we
o '
° 42557
Hampton-Seabrook Estuary System
1 mile
o A
7055

fe} ‘
70 45
!

Figure 5. Summary of 256 collecting sites in New Hampshire and southern
Maine.

1986] Mathieson & Hehre — New Hampshire seaweeds 9

including 23 locations where a minimum of 2 years of monthly
collections were made. Forty-nine stations were studied in the
Hampton-Seabrook Estuary, New Hampshire, during the summer
and fall of 1969 (Mathieson and Fralick, 1972), including collections
from the five major rivers and creeks, Hampton Harbor and the
adjacent open coast near Hampton. Thirteen sites from the near-
shore open coast between Portsmouth and Seabrook were docu-
mented, as well as the species composition at three of the nine major
islands at the Isles of Shoals (Mathieson and Penniman, 1986a). One
hundred forty-seven collection sites were studied within the Great
Bay Estuary System, including Great Bay, Little Bay, the Bellamy,
Cocheco, Lamprey, Oyster, Piscataqua, Salmon Falls, Squamscott
and Winnicut Rivers. Collections were also made at 44 adjacent
coastal and estuarine sites in Maine (Figure 5, Table IV and
Appendix).

Representative samples of all conspicuous species at each site
(Table IV and Appendix) were made in the littoral (on foot) and
sublittoral zones (by SCUBA). Methods of collection, identification
and processing of samples were similar to those outlined by Hehre
and Mathieson (1970) and Mathieson, Hehre and Reynolds (1981).
Monthly collections for at least 2 years were made at 23 sites rang-
ing from the nearshore open coast of New Hampshire through the
entire Great Bay Estuary System, including its tidal tributaries
(Table IV and Appendix). Intermittent or seasonal collections were
made at the other 233 sites (i.e., including 44 Maine stations). Her-
barium voucher specimens of each taxon/site were prepared and
deposited in NHA. The complete set of approximately 40,000 spec-
imens is deposited in order to document temporal and spatial char-
acteristics of the state’s marine algal flora. The primary source of
identification was Taylor (1957): even so, several other monographs
(see Mathieson, Hehre and Reynolds, 1981, for a partial listing) and
the recent nomenclatural changes summarized by South (1984) were
also employed.

THE NEW HAMPSHIRE COASTAL ZONE

The New Hampshire coastline is located approximately midway
between Cape Elizabeth, Maine, and Cape Ann, Massachusetts

10 Rhodora [Vol. 88

(Figure 1). Many geological-topographical characteristics are
common to this coastal region, including the general absence of
offshore islands, the presence of sandy barrier beaches in front of
extensive salt marshes, and the occurrence of large rocky headlands
or promontories.

As shown in Figures 1-4, the state’s coastal zone consists of four
primary areas:

1. Isles of Shoals

2. seventeen miles of nearshore open coast and adjacent salt
marshes

3. Hampton-Seabrook Estuary System

4. Great Bay Estuary System

The Isles of Shoals are located approximately 9 miles SSE of the
mouth of the Piscataqua River and 6.5 miles due east of Straw
Point, Rye (Figures | and 2). The islands oocupy an area 3 miles
north-south by 1.5 miles east-west and lie between the coordinates
42°59’N, 70°37’20"W and 43°00’30’N, 70°36’W. There are nine
major islands: five are under the jurisdiction of the Town of Kittery,
Maine (Appledore, Cedar, Duck, Malaga, and Smuttynose), and
four are within Rye, New Hampshire (Lunging, Seavey, Star and
White). Nine other rocks and ledges are present in the Island group
(Anderson, Eastern, Halfway, Mingo, Shag and Square Rocks, plus
Cedar and White Islands Ledges and Southwest Ledge). The Isles of
Shoals are massive granitic outcrops, the north and east sides of
which are exposed to extreme wave action, particularly during
storms. The west and south sides of the islands are more sheltered,
such as, Gosport Harbor on the leeward side of Star Island. Mathie-
son and Penniman (1986a) and Norall et al. (1981) give a variety of
other details regarding the physical-environmental characteristics
of the Shoals.

The southern boundary of New Hampshire’s nearshore open
coast (Figures | and 4) is at Seabrook (42°52’30’N, 70°49’W), while
the northern boundary is at the mouth of the Piscataqua River near
the entrance to Portsmouth Harbor (43°04’20’N, 70°42’42”W).
Extensive salt marshes occur along this coast, particularly near Rye,
Portsmouth and Hampton. Three major habitats are found on the
nearshore open coast: cobble-boulder, exposed headlands, and
sandy beaches (see Hehre and Mathieson, 1970, for further descrip-
tions). The metamorphic headlands at Rye Ledge and Great Boars

1986] Mathieson & Hehre — New Hampshire seaweeds 1]

Head are exposed to extreme wave action. The most extensive
sandy beaches are found in the Hampton and Seabrook areas.

The Hampton-Seabrook Estuary (Figure 4) is located entirely
within the State of New Hampshire, between latitudes 42°51’30”N
to 42°55’55”N and longitudes 70°49’30”W to 70°51’30’W. This
estuary is within the townships of Hampton, Hampton Falls and
Seabrook, and has a total area of about 3,800 acres. Five rivers
(Taylor, Hampton, Hampton Falls, Brown and Blackwater), as well
as a variety of smaller creeks and brooks are present within this
estuary.

The Great Bay Estuary System occurs within New Hampshire
and Maine (Figure 3). It consists of Great Bay, Little Bay, the
Piscataqua River, Portsmouth Harbor and its tributaries, as well as
seven other freshwater rivers (Bellamy, Cocheco, Lamprey, Oyster,
Salmon Falls, Squamscott and Winnicut), which drain into the
basin. The Great Bay Estuary System is one of the largest estuaries
on the eastern seaboard of the United States, with over 11,000 acres
of tidewater (Anon., 1960). The total drainage area of the estuary is
approximately 930 square miles, two-thirds of which is within New
Hampshire (Anon., 1960). The estuary contains about 100 miles of
shoreline. The substratum in the Great Bay Estuary System, as well
as the Hampton-Seabrook Estuary, is dominated by mud
(Hardwick-Witman and Mathieson, 1983); occasional rock out-
crops, cobbles, shells and artificial structures such as pier pilings are
also present. Overall, the substratum within the Hampton-Seabrook
Estuary is more sandy than the Great Bay Estuary System, particu-
larly toward Hampton Harbor.

The seasonal patterns of surface water temperatures on the near-
shore open coast of New Hampshire and within the Great Bay
Estuary System are illustrated in Figure 6. Typically, the maximum
temperatures occur during mid-summer through the fall. There-
after, temperatures decrease rapidly, particularly in the inner estu-
ary, with lowest values occurring January to March. Open coastal
site have a narrower temperature range than estuarine sites. For
example, surface water temperatures at the Isles of Shoals vary from
3.8° to 18.2°C, versus —1.0° to 19.0°C at Portsmouth Harbor,
—2.0° to 24.1°C at Dover Point, —1.8° to 26.5°C at Adams Point,
and —2.0° to 27.0°C within Great Bay proper (Emerich Penniman
et al., 1985; Norall and Mathieson, 1976; Norall et al., 1982). Even

30 ee ee ee Sa ee ae es ee a ee
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1973 1974 1975 1970 1977 1978 1979 1980 1981

Figure 6. Seasonal variation of surface water temperature on the nearshore open coast of New Hampshire (Jaffrey Point,
Fort Stark) and within the Great Bay Estuary System (Atlantic Terminal and Great Bay) during 1973-1981 (based upon
Emerich Penniman et al., 1985).

Cl

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1973 1974 1975 1976 1977 1978 1979 1980 1981

Figure 7. Seasonal variation of surface water salinities on the nearshore open coast of New Hampshire (Jaffrey Point, Fort

Stark) and within the Great Bay Estuary System (Atlantic Terminal and Great Bay) during 1973-1981 (based upon Emerich

Penniman et al., 1985).

[9861

spoomeas oilysduie py MON — Yo % uosaepy

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14 Rhodora [Vol. 88

greater variations (daily and seasonally) of temperatures are present
within riverine habitats of the Great Bay Estuary System. Daly and
Mathieson (1979, 1981), Daly et al. (1979), Emerich Penniman et al.
(1985), Glibert (1976), Loder et al. (1979), Norall and Mathieson
(1976), Norall et al. (1982), and Silver and Brown (1979) all gave
details regarding temperature and salinity variations within the
same geography. Overall, there is a pattern of greater variation as
well as increasing mean surface water temperatures from the open
coast to the inner estuary (Figure 8).

The seasonal patterns of surface water salinities on the nearshore
open coast of New Hampshire and within the Great Bay Estuary
System are illustrated in Figure 7. Typically, the maximum salinities
occur in the summer and fall, while the lowest salinities occur during
January to early spring—i.e., during winter and spring thaws. As
with temperatue, the most pronounced salinity variations occur
within inner estuarine sites, while adjacent open coastal areas are
more stable. For example, the surface water salinities at the Isles of
Shoals range from 31.0-33.0%o, while greater variations are evident
at Portsmouth Harbor (24.6-33.8% 0), Dover Point (0.9-30.3%o),
Adams Point (6.6-31.4%), and within Great Bay proper (2.7-
30.97% ) (Emerich Penniman et al., 1985; Norall and Mathieson,
1976; Norall et al., 1982). Overall, there is a pattern of increased
salinity variation (daily and seasonally), as well as a clinal decrease
in surface water salinities, from the open coast of New Hampshire to
the inner estuary (Figure 8).

A foot or more of ice is usually present from late December to
March in Great Bay and the major tidal rivers (except the Piscata-
qua) within the Great Bay Estuary System. The scouring effects of
ice are evident on rocks, pier pilings and other solid substrata. Large
sections of marshy shoreline may be torn loose (rafted) during the
spring thaw (Hardwick’Witman, 1985; Mathieson, Penniman, Busse
and Tveter-Gallagher, 1982). Floating ice rafts and icebergs can
often be seen on the adjacent open coast (Jaffrey Point) during the
spring thaw.

The water transparency at the Isles of Shoals is much greater than
in Portsmouth Harbor, which in turn is greater than that within the
Great Bay Estuary System (Daly et al., 1979). The depth of penetra-
tion of light (1%) in the sea determines the lower limits of plant
distribution along this natural gradient (Figure 8). Thus, attached

1986] Mathieson & Hehre — New Hampshire seaweeds 15
25
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Distance (Mi) Inland

Figure 8.
penetration on the nearshore open coast of New Hampshire and within the Great Bay
Estuary System during 1974-1978 (based upon Daly et al., 1979).

) ve 8.6 10 W1 12.2 13.8

Mean values of surface water temperatures, salinities and 1% light

16 Rhodora [Vol. 88

marine plants which require light to photosynthesize are found at
100-125 feet at the Shoals, 60-80 feet near the mouth of Portsmouth
Harbor, and 10-15 feet in the upper parts of Great Bay proper
(Figure 9). The differential water clarity of the estuary is primarily
related to the volume of silt and organic material (detritus) in the
latter habitat (Daly et al., 1979; Norall and Mathieson, 1976; and
Norall et al., 1982).

Differential levels of nutrients (nitrogen and phosphorus) are
evident on the open coast of New Hampshire and within the Great
Bay Estuary System (Figures 10 and 11), with lower values occurring
in the former areas (Norall and Mathieson, 1976; Norall et al.,
1982). In general, nutrients are highest during the winter months
from December into March; thereafter, a sharp decline occurs due
to the spring bloom of phytoplankton. Intermediate levels are usu-
ally found during the summer, and they begin to increase during the
fall. A detailed tabulation of seasonal and spatial variations of ni-

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rn “.. Dover Point
id is
> “Adams Point
o 204 on
x ,.Pierce Point
a | *Chapmans Landing
M.tw.* 0 T T T T T T T
10 5 0 5 10 15 20 25 30

Distance (Mi.)
Offshore ~- > Inland

Figure 9. The lower limits of subtidal plant distribution between the Isles of
Shoals and the inner reaches of the Great Bay Estuary System.

1986] Mathieson & Hehre — New Hampshire seaweeds 17

trogenous, phosphorous, and silicious nutrients within the Great
Bay Estuary System and the adjacent open coast of New Hampshire
is given by Norall and Mathieson (1976), Norall et al. (1982), and
Emerich Penniman et al. (1985). Additional nutrient data for the
same area are summarized by Burns and Mathieson (1972b), Glibert
(1976), Loder and Glibert (1977, 1980), Loder et al. (1979), Lyons,
Loder and Murray (1982), Mathieson and Burns (1975) and
Mathieson and Tveter (1975).

The average tidal amplitude at the Shoals and near Portsmouth
Harbor is about 8.1 feet, while it is about 6.8 feet at the head of
Great Bay proper (Anon., 1965). Two high and two low tides occur
each day in the coastal zone, and they are uniformly semi-diurnal.
Tides cause considerable fluctuations of water transparency, tem-
perature, salinity and current speeds, particularly in estuarine habi-
tats (Daly and Mathieson, 1979). Tidal currents are a conspicuous
feature of the Great Bay Estuary System (Figure 3), particularly in
narrow channels near Adams Point, Dover Point, Fox Point, and
the lower Piscataqua River (Brown and Arrellano, 1979; Celikkol
and Reichard, 1976; Mathieson, Neefus, and Emerich Penniman,
1983; Mathieson, Reynolds, and Hehre, 1981; Mathieson, Tveter,
Daly and Howard, 1977; Reynolds and Mathieson, 1975; Schmidt,
1980; Swenson et al., 1977; Trask and Brown, 1980). In such habi-
tats tidal currents of 4-6 knots are evident, with maximum currents
occurring during ebb tide (Figure 12). All of the tidal waters of the
Great Bay Estuary System enter and leave via the Piscataqua River,
creating strong tidal currents.

Domestic pollution is moderate within the Great Bay Estuary
System. Treated effluent (chlorinated and settled) is discharged from
the towns of Dover, Durham, Exeter, Newmarket and Rochester.
Occasionally, raw sewage may be discharged during extreme storm
periods when some sewage treatment plants (Dover) cannot handle
the volume. Industrial pollution (heavy metals and organic sludge)
is discharged from Dover, Rochester and Portsmouth (Capuzzo and
Anderson, 1973; Hines et al., 1984; Lyons, Armstrong, O'Neil and
Gaudette, 1982). Little industrial pollution occurs in the Hampton-
Seabrook Estuary. On the open coast of New Hampshire, little
pollution from domestic and industrial sources occurs, except for a
few “point sources” of domestic discharges in Rye and other areas.
The Isles of Shoals represent a relatively “pristine” coastal
environment.

NITRATE CUG-AT NOg-N/L)

~
Qo
TOT TTT TTT TTT TT

ve .
‘ .

t . ; A, KN: N yy, VSB
pitiurps ob itt bids ditt dt tit pir S ae pepe spp rp F994 ph ps pres pYSppitiit paurias Lia pp pid

°

/ LY
‘f v i | y _- " 7; A.
i ~y ‘ Cd

TTTTTTTTTTrTrTrTrTrTrTrTrTrrrrrrrrrrrrryrrrrrrr ery v rere rt rrr ttre rv rr vt) VP PVP TY PPV tte TY) PTV PY VP PP VY YT

Cr:

jwounwet

“¢

ee es a

eee eueeeeeeeeeenee

SONOJFMANJJASONOJFMANJIJASONOJFNANIJIJASONOJFRANJIJASONDUFMANJIJASONDIFMANJIASONOVENANJIJASONOVFNAMNJIASOND

1973 1974 1975 1976 1977 1978 1979 1980 1981

Figure 10. Seasonal variation of nitrogenous-nutrient levels (i.e. nitrate-N) on the nearshore open coast of New Hampshire
(Jaffrey Point, Fort Stark) and within the Great Bay Estuary System (Atlantic Terminal and Great Bay) during 1973-1981

(based upon Emerich Penniman et al., 1985).

81

elopoyy

88 10A]

-
8 TIT t
a E 1
al + 4
“N 2.5 + 4
a r 4
_ a q
oO a 4
a 2.0 4
L 4
— - 4
a r K 5
1 r 4
mo 1.56 { \ 7
—) r | } 4
G r ; i 4
e ik \ -
ive) r H \ 7
~e 1-0 IV AG -
= c Vw 4
a L
al o.s E
5 t
= c
= ot
Coe oa Gs as ss LLLtLLEL ELE LAELEL ELL Let tite t spt ttt ititttr tit r ppp tidy
SOMDIFMAMJIASONOJFNANJIJASONDJFNANJJASONOVEMANJSIJASONDJFMARJJASONDJFNAMJIASONDIFNANJIJASONDIFRANIJIASOND

1873 1874 1875 1876 1877 1978 1879 1880 1081

Figure 11. Seasonal variation of phosphorous-nutrient levels (i.e. orthophosphate-P) on the open coast of New Hampshire
(Jaffrey Point, Fort Stark) and within the Great Bay Estuary System (Atlantic Terminal and Great Bay) during 1973-1981
(based upon Emerich Penniman et al., 1985).

[9861

spoomeas oilysdwiey MON — 14d % UOsalyIepy

61

20 Rhodora [Vol. 88

6-
4 =—*
=
5 | \
] \
| :
4- ; ,
[ \
3- y °
(op) :
sag e
O is
S74 4 HW!
\ /
\ F
‘= \ Pee e /
\y Bee
—r if I T T T
6 8 10 12 14 16 18
AM PM

(HOURS)

Figure 12. Diurnal variations of tidal currents at Dover Point.

SPECIES COMPOSITION

A total of 216 taxa is recorded from the coastal and estuarine
environments of New Hampshire (Figures 1-5), including 58 Chlo-
rophyceae, 66 Phaeophyceae and 92 Rhodophyceae (Tables I-III).
Two of these 216 taxa (Hecatonema terminalis and Myrionema
magnusii) were not collected by us, but they were recorded by Col-
lins (1900) from the state with no specific dates nor collection sites.
All of the other taxa, except for Acrochaete repens, Bolbocoleon
piliferum, Prasinocladus marinus and Sphaerotrichia divaricata,
which were only obtained in culture after their grow-out in enriched
sea water media (Zechman and Mathieson, 1985), were collected in
one or more estuarine and/or open coastal habitats. Eight of the 216
taxa recorded herein are new records for the state, including one

1986] Mathieson & Hehre — New Hampshire seaweeds 21

brown (Sphaerotrichia divaricata) and seven red algae (Audouinella
violaceae, Callocolax neglectus, Ceramium elegans, Cruoriopsis
ensis, Halosacciocolax kjellmanii, ‘“Porphyrodiscus simulans” and
Turnerella pennyi). | Porphyrodiscus simulans and its other life his-
tory stages (cf. Farnham and Fletcher, 1976) are hereafter desig-
nated by quotes.] Each of the seven red algae represents either a
range extension or a new record for the northeastern coast of North
America. For example, the “fresh water” red alga Audouinella vio-
lacea 1s newly recorded from coastal waters of the northeast; it
grows as an epiphyte on the fresh water red alga Sacheria fucina,
and it may occur abundantly within riverine habitats. The specific
parasite Callocolax neglectus which grows abundantly on Callo-
Phyllis cristata, was previously recorded from Greenland (Pedersen,
1976) and Newfoundland (South and Hooper, 1980). Ceramium
elegans was earlier known from the Canadian Maritime Provinces
(South, 1984; Taylor, 1957), while Halosacciocolax kjellmanii and
“Porphyrodiscus simulans” were recorded from Newfoundland and
the Canadian Maritime Provinces (South, 1984). The geographical
distribution of Turnerella pennyi was previously recorded from the
Arctic to the Atlantic coast of Nova Scotia, while Taylor (1957) only
listed “Cruoriopsis ensis” from southern Massachusetts. The green
alga Codium fragile ssp. tomentosoides has recently (1983) been
found attached at Appledore Island, Maine, Isles of Shoals (Figure
2), and it could extend to adjacent New Hampshire sites (Carlton
and Scanlon, 1985; Mathieson and Penniman, 1986a).

PHENOLOGY AND LONGEVITY

A summary of the temporal variation of seaweed taxa within
estuarine—coastal waters of New Hampshire is given in Figure 13,
based upon the data in Tables I-III. The number of taxa/month
was highest in August (178) and lowest in January (105). Each of the
three major groups of seaweeds showed a similar seasonal pattern,
except that the Chlorophyceae had their highest number of taxa in
July (42). Similar phenological patterns with summer maxima and
winter minima have been noted in several other North Atlantic
areas (Coleman and Mathieson, 1975; Lamb and Zimmerman,
1964; MacFarlane and Bell, 1933; Reynolds and Mathieson, 1975;
Sears and Wilce, 1975). Chapman (1964) and Williams (1948, 1949)
have emphasized that seasonally dynamic floras and a wide range of

1 Rhodora [Vol. 88

Green

NUMBER OF TAXA

Figure 13. Temporal variations of seaweed taxa within estuarine-coastal waters
of New Hampshire, expressed as the number of taxa/ month.

annuals exist in areas with pronounced temperature fluctuations.
[he functional role of annuals in mediating the seasonal cycle of
New Hampshire seaweeds is shown in Tables I-III.

Of the total algal flora outlined in Tables I-III, 113 taxa (52.3%)
were designated as annuals and 100 taxa (46.3%) were interpreted as
perennials (Figure 14). Three algae (Cladophora sericea, Derbesia
marina and Ulva lactuca) or 1.4% of the flora, require further study
as they may be either aseasonal annuals or pseudoperennials (sensu
Knight and Parke, 1931). As outlined below, the longevity patterns
(ratios of annuals/ perennials and the percentage of perennial taxa)
for the three major groups of seaweeds are conspicuously different:
|. Rhodophyceae—30 annuals/62 perennials (0.48:1) or 67.4%
‘rennials

Phaeophyceae—36 annuals/30 perennials (1.2:1) or 45.5%

erennials

’. Chlorophyceae—47 annuals/8 perennials (5.9:1) or 13.8%
yerennials

1986] Mathieson & Hehre — New Hampshire seaweeds y

1107 [
100 - Green
4
90 Ty
ae , Annuals
[ Perennials
80 - _ Brown
V4
x 704
fia rT |
= 60 7
= b+
- 50-4
3
z 40-
Red

30-4

20-4

10-7
x = Annuals
a
i
Le
ro.) Perennials
t=
Zz
WW
oO
wa
uJ
a.

| 2 3 4 Total

GEOGRAPHICAL AREAS

Figure 14. The number and percentage of annual and perennial chlorophycean,
phaeophycean and rhodophycean taxa within estuarine-coastal waters of New

Hampshire.

Thus, the red algae exhibit the greatest dominance by perennials,
the brown algae have an intermediate pattern and the green algae
exhibit the greatest dominance by annuals.

24 Rhodora [Vol. 88

As outlined in Tables I-III, three distinct types of seasonal annu-
als (winter, spring and summer) plus aseasonal annuals can be dis-
tinguished, according to their season of maximum growth and
development. No fall annuals were found, although many summer
plants persist into fall and early winter. Thus, the fall season appears
to be a transition period between summer and winter floras. Most of
the seasonal annuals reproduce during their periods of maximum
abundance. On the other hand some aseasonal annuals ( Petalonia
fascia and Scytosiphon lomentaria var. lomentaria) are reproduc-
tive throughout the year while others such as Dumontia contorta
only reproduce during more restricted periods (Hehre and Mathie-
son, 1970; Kilar and Mathieson, 1978; Mathieson and Hehre, 1982,
1983). Similarly, these authors as well as Tveter-Gallagher et al.
(1980), pointed out that many perennials are reproductive through-
out the year, while others exhibit distinct reproductive periods.

Several taxa were only collected in situ a few times (Chlorochy-
trium moorei, “Halicystis ovalis,” Pringsheimiella scutata, Spiro-
gyra sp., Stichococcus marinus, Stigeoclonium sp., Cladostephus
spongiosus forma verticillatus, Eudesme virescens, Giffordia se-
cunda, Scytosiphon lomentaria var. complanatus, Sorocarpus
micromorus, Sphacelaria fusca, Audouinella polyides, Ceramium
elegans, Colaconema polvides, “Cruoriopsis ensis,” Erythropeltis
discigera var. discigera, Halosacciocolax kjellmanii, “ Porphyrodis-
cus simulans,” and Turnerella pennyi). Several of these plants plus
Acrochaete repens, Bolbocoleon piliferum, Prasinocladus marinus
and Sphaerotrichia divaricata, which were only found in culture
(Tables I-III), may have been missed due to their small stature. In
contrast to these “rare” taxa, many of the larger perennial forms
were ubiquitous at a wide variety of coastal and estuarine sites
throughout the year. Specific details on the seasonal occurrence and
longevity of each taxon are summarized in Tables I-III.

PATTERNS OF LOCAL DISTRIBUTION

A summary of the local distribution of the chlorophycean,
phaeophycean and rhodophycean taxa in the four major coastal-
estuarine areas in New Hampshire is shown in Figure 15. The high-
est number of taxa (179) was recorded from the nearshore open
coast between Portsmouth and Seabrook. The species richness at
the Isles of Shoals and within the Great Bay Estuary System was

1986] Mathieson & Hehre — New Hampshire seaweeds 25

E
@ —
2 s
220- % B Fc
0 >
d 38 G ie
” c 2
2004 3 a uw
2 Oo 5
}2 € 2
- Green
1807 ° ”
2 = ~
ei m7 5
1607] E tm
rf} o6§ ob
a >
(ép)
140- >
= : 7. g
= i me Bie
120+ a J rown
owen |. i 3
ac s
oon ©
> “”)
2 ; ‘
So
80+ =.
E
co
pS os
60 —]
40
20
x
Red

Figure 15. Local distribution of chlorophycean, phaeophycean and rhodophy-
cean taxa within the four major coastal-estuarine areas in New Hampshire.

nearly the same (161 and 164 taxa, respectively), although the spe-
cies composition of each was conspicuously different. The low spe-
cies richness (63 taxa) within the Hampton-Seabrook Estuary
System contrasts strongly with that of the Great Bay Estuary. Each

26 Rhodora [Vol. 88

of the three major groups of seaweeds exhibited their minimum
number of species within the Hampton-Seabrook Estuary System.
The Chloropyceae and Rhodophyceae had an approximate equality
of species numbers at the two open coastal areas and within the
Great Bay Estuary System, with 42-47 green algal taxa at the three
habitats versus 71-76 red algae at the same three sites. In contrast,
the Phaeophyceae exhibited a more pronounced difference between
the nearshore open coast and the Great Bay Estuary System, with 56
taxa at the former and 44 taxa at the latter.

As noted by Mathieson and Fralick (1972), the low species
richness within the Hampton-Seabrook Estuary may be associated
with its limited acreage (reduced habitats) and lack of stable sub-
strata. On the other hand, the relatively high species diversity at the
Isles of Shoals (comparable to the nearshore open coast and the
entire Great Bay Estuary System), is impressive as they are a small
albeit relatively “pristine” set of islands (Mathieson and Penniman,
1986a).

The species composition within the Great Bay Estuary System is
very different than that at the Shoals and the nearshore open coast,
because of the presence of several freshwater algae, the enhanced
number of “estuarine” taxa and the reduced number of “coastal”
species. A similar reduction of open coastal species within estuarine
habitats has been related to variable hydrographic conditions and
limited rocky substrata (Doty and Newhouse, 1954; Hardwick-
Witman and Mathieson, 1983; Mathieson and Fralick, 1972;
Mathieson, Reynolds and Hehre, 1981).

The coastal and/or estuarine distributional patterns of each chlo-
rophycean, phaeophycean and rhodophycean taxon are summar-
ized in Tables I-III, as well as within the individual distribution
maps (Figures 16-229). Overall, three basic patterns are evident as
follows:

1. coastal—restricted to the open coast

2. cosmopolitan—present in both estuarine and open coastal

environments

3. estuarine—restricted to estuarine environments

Approximately 23% of the total flora (49 of 216 taxa) were re-
stricted to the open coast. Twenty-five of these taxa occurred in
both open coastal habitats (Figures: 30. Cladophora rupestris; 33.
“Codiolum petrocelidis”; 36. Derbesia marina; 45. Entocladia flus-

1986] Mathieson & Hehre — New Hampshire seaweeds 27

trae; 47. Gomontia polyrhiza; 75. Alaria esculenta; 76. Ascocyclus
distromaticus; 79. Asperococcus fistulosus; 106. Laminariocolax
tomentosoides; 107. Leathesia difformis; 108. Mikrosyphar porphy-
rae; 115. Protectocarpus speciosus; 118. Punctaria plantaginea; 123.
Saccorhiza dermatodea; 141. Audouinella alariae; 158. Ceratocolax
harizii; 168. Devaleraea ramentaceum; 170. Erythropeltis discigera
var. discigera; 179. Harveyella mirabilis; 181. Leptophytum foecun-
dum, 189. Nemalion helminthoides; 196. Phyllophora traillii; 200.
Phymatolithon rugulosum; 210. Polysiphonia novae-angliae; 227.
Spermothamnion repens). On the other hand, nine taxa were res-
tricted to the Isles of Shoals (Figures: 25. Chaetomorpha minima;
35. Codium fragile subsp. tomentosoides,; 58. Pringsheimiella scu-
tata; 65. Stichococcus marinus; 83. Cladostephus spongiosus forma
verticillatus; 93. Fucus distichus subsp. anceps; 130. Sphacelaria
fusca; 178. Halosacciocolax kjellmanii; 191. Pantoneura baeri),
while 13 were only collected from the nearshore open coast (Figures:
18. Bolbocoleon piliferum; 27. Chlorochytrium moorei; 48. “Hali-
cystis ovalis”: 56. Prasinocladus marinus; 92. Eudesme virescens;
101. Giffordia secunda; 128. Sphacelaria arctica; 131. S. plumigera;
132. S. plumosa; 134. Sphaerotrichia divaricata; 143. Audouinella
polyides; 218. Porphyropsis coccinea; 229. Turnerella pennyi).

In contrast to the moderate number of open coastal taxa, 145
species or 67% of the total flora exhibited a cosmopolitan distribu-
tion, occurring in both open coastal and estuarine habitats. Several
of these cosmopolitan species were broadly distributed in estuarine
habitats (Figures: 17. Blidingia minima; 23. Chaetomorpha linum;
31. Cladophora sericea; 37, 38, 40, 41, 42 & 43. Enteromorpha ssp.;
51. Monostroma grevillei; 55. Percursaria percursa; 60. Rhizoclo-
nium riparium; 67. Ulothrix flacca; 69. Ulva lactuca; 70. Ulvaria
obscura; 77. Ascophyllum nodosum; 90. Ectocarpus siliculosus; 96.
Fucus distichus subsp. evanescens,; 98. F. vesiculosus; 111. Petalonia
fascia; 114. Pilayella littoralis; 122. Ralfsia verrucosa; 125. Scytosi-
phon lomentaria var. lomentaria; 138. Ahnfeltia plicata; 151. Calli-
thamnion tetragonum; 156. Ceramium rubrum; 160. Chondrus
crispus; 165. Cystoclonium purpureum; 169. Dumontia contorta;
177. Gymnogongrus crenulatus; 180. Hildenbrandia rubra; 195.
Phyllophora_ pseudoceranoides; 197. P. truncata; 202. Polyides
rotundus,; 205. Polysiphonia flexicaulis; 206. P. harveyi; 208. P.
nigra; 209. P. nigrescens; 216. Porphyra umbilicalis). Many other

28 Rhodora [Vol. 88

cosmopolitan species were restricted to outer estuarine sites, that is,
adjacent to the open coast, or they occurred as “disjunct” popula-
tions within estuarine tidal rapid sites such as Dover Point and Fox
Point (Figures: 21. Chaetomorpha aerea; 24. C. melagonium; 34.
“Codiolum pusillum”,; 52. Monostroma pulchrum; 57. Prasiola stip-
itata; 59. Pseudendoclonium submarinum; 73. Urospora wormski-
oldii; 74. Agarum cribrosum; 80. Chorda filum; 81. C. tomentosa;
85. Desmarestia aculeata; 86. D. viridis; 87. Desmotrichum undula-
tum; 88. Dictyosiphon foeniculaceus; 89. Ectocarpus fasciculatus;
91. Elachista fucicola; 94. Fucus distichus subsp. distichus; 95. F.
distichus subsp. edentatus; 103. Laminaria digitata; 104. L. longi-
cruris; 109, Myrionema corunnae; 112. Petalonia zosterifolia; 121.
Ralfsia fungiformis; 135. Spongonema tomentosum, 137. Ulonema
rhizophorum,; 144. Audouinella purpurea; 152. Callocolax neglec-
tus; 153. Callophyllis cristata; 161. Choreocolax polysiphoniae; 163.
Corallina officinalis; 167. Dermatolithon pustulatum; 173. Gigar-
tina stellata; 174. Gloiosiphonia capillaris; 182. Leptophytum laeve;
183. Lithophyllum corallinae; 184. Lithothamnion glaciale; 187.
Lomentaria orcadensis; 188. Membranoptera alata; 192. “Petrocelis
cruenta’, 194, Phycodrys rubens; 198. Phymatolithon laevigatum;
199. P. lenormandii; 201. Plumaria elegans; 207. Polysiphonia
lanosa; 212. P. urceolata; 213. Porphyra leucosticta; 220. Ptilota
serrata; 221. Rhodomela confervoides; 222. Rhodophyllis dicho-
toma; 228. “Trailliella intricata’). A few cosmopolitan taxa were
uncommon/rare on the open coast but widely distributed in estua-
rine habitats (Figures: 19. Bryopsis plumosa; 20. Capsosiphon ful-
vescens; 44. Enteromorpha torta; 49. Kornmannia leptoderma; 50.
Microspora pachyderma; 71. Ulvaria oxyspermum; 139. Antitham-
nion cruciatum; 157. Ceramium strictum; 159. Chondria baileyana;
175. Goniotrichum alsidii; 186. Lomentaria clavellosa (only col-
lected once on the open coast at Boone Island, Maine); 204. Polysi-
pPhonia elongata).

Sixteen seaweeds or 7% of the total flora were restricted to the
Great Bay and/or Hampton-Seabrook Estuary Systems (Figures:
29. Cladophora pygmaea; 39. Enteromorpha flexuosa subsp. flex-
uosa; 78. Ascophyllum nodosum ecad. scorpioides; 117. Punctaria
latifolia; 124. Scytosiphon lomentaria var. complanatus; 127. Soro-
carpus micromorus; 136. Stictyosiphon griffithsianus; 149. Calli-
thamnion byssoides; 155. Ceramium elegans; 166. Dasya baillouvi-

1986] Mathieson & Hehre — New Hampshire seaweeds 29

ana; 176. Gracilaria tikvahiae; 185. Lomentaria baileyana; 203.
Polysiphonia denudata; 211. P. subtilissima; 217. “Porphyrodiscus
simulans"; 224. Rhodophysema georgii). Six “fresh-water” taxa were
found attached in riverine habitats near the headwaters of tidal
tributaries (Figures: 53. Mougeotia sp.; 54. Oedogonium Spc 62,
Spirogyra sp.; 66. Stigeoclonium sp.; 146. Audouinella violacea;
225. Sacheria fucina). Some of the latter species, such as Sacheria
fucina, are known to be tolerant of reduced salinities (Wood and
Straughan, 1953).

As outlined by Mathieson, Reynolds and Hehre (1981), closely
related taxa often have distinct distributional patterns within estu-
aries. The different taxa of Chaetomorpha (Figures 21-26), Clado-
phora (Figures 28-31), Monostroma (Figures 51,52), Rhizoclonium
(Figures 60, 61), U/varia (Figures 70, 71), Ectocarpus (Figures 89,
90), Fucus (Figures 93-98), Laminaria (Figures 103-105), Punctaria
(Figures 117, 118), Ralfsia (Figures 119-122), Sphacelaria (Figures
128-133), Audouinella (Figures 141-146), Callithamnion (Figures
149-151), Ceramium (Figures 154-157), Lomentaria (Figures 185-
187), Phyllophora (Figures 195-197), Phymatolithon (Figures
198-200), Polysiphonia (Figures 203-212) and Porphyra (Figures
213-216) can all be cited. For example, of the six Chaetomorpha
species recorded (Table I, Figures 21-26), one (C. minima) was
restricted to the Isles of Shoals, while five (C. aerea, C. brachygona,
C. linum, C. melagonium and C. picquotiana) exhibited cosmopoli-
tan distributional patterns of varying degrees. Overall, C. Jinum was
the most ubiquitous and broadly distributed species (Figure 23).
The different taxa of Fucus exhibited a similar pattern (Tables I and
II, Figures 93-98), with one taxon (F. distichus subsp. anceps) being
restricted to the Isles of Shoals, and five (F. distichus subsp. disti-
chus, F.d. subsp. edentatus, F.d. subsp. evanescens, F. spiralis and
F. vesiculosus) exhibiting varying cosmopolitan distributional pat-
terns. Fucus vesiculosus was the most broadly distributed taxon of
this group (Figure 98). A comparison of the ten Polysiphonia spe-
cies (Table III, Figures 203-212) shows that two species (P. denu-
data and P. subtilissima) were restricted to estuarine environments,
one (P. elongata) was rare on the open coast, six (P. flexicaulis, P.
harveyi, P. lanosa, P. nigra, P. nigrescens and P. urceolata) exhi-
bited cosmopolitan distributions, and one (P. novae-angliae) was
restricted to the open coast. Similar distributional comparisons can

30 Rhodora [Vol. 88

also be made for different genera in the same families (sensu Smith,
1950; South, 1984): Ulotrichaceae (Stichococcus and Ulothrix, Fig-
ures 65, 67 and 68); Chaetophoraceae (Acrochaete, Bolbocoleon,
Entocladia, Pringsheimiella, Pseudendoclonium and Stigeoclonium,
Figures 16, 18, 45, 46, 58, 59 and 66); Acrosiphoniaceae (Spongo-
morpha and Urospora, Figures 63, 64, 72 and 73); Percursariaceae
(Blidingia, Gomontia, Kornmannia, Monostroma and Percursaria,
Figures 17, 47, 49, 51, 52 and 55); Ulvaceae (Capsosiphon, Entero-
morpha, Ulva and Ulvaria, Figures 20, 37-44, 69, 70 and 71); Cla-
dophoraceae (Chaetomorpha, Cladophora and Rhizoclonium,
Figures 21-26, 28-31, 60 and 61); Bryopsidaceae (Bryopsis and
Derbesia, Figures 19 and 36); Zygnemataceae (Mougeotia and Spi-
rogyra, Figures 53 and 62); Ectocarpaceae (Ectocarpus, Giffordia,
Laminariocolax, Mikrosyphar, Pilayella, Sorocarpus and Spongo-
nema, Figures 89, 90, 99-101, 106, 108, 114, 127 and 135); Ralfsia-
ceae (Petroderma, Pseudolithoderma, Ralfsia fungiformis, R.
verrucosa and Sorapion, Figures 113, 116, 121, 122 and 126); Myri-
onemataceae (Ascocyclus, Myrionema, Protectocarpus and Ulo-
nema, Figures 76, 109, 110, 115 and 137); Chordariaceae (Chordaria,
Eudesme and Sphaerotrichia, Figures 82, 92 and 134); Striariaceae
(Isthmoplea and Stictyosiphon, Figures 102 and 136); Punctaria-
ceae (Asperococcus, Desmotrichum and Punctaria, Figures 79, 87,
117 and 118); Scytosiphonaceae (Petalonia and Scytosiphon, Fig-
ures 111, 112, 124 and 125); Laminariaceae (Agarum, Laminaria
and Saccorhiza, Figures 74, 103-105 and 123); Fucaceae (Ascophyl-
lum and Fucus, Figures 77, 78 and 93-98); Cystocloniaceae (Cysto-
clonium and Rhodophyllis, Figures 165 and 222), Phyllophoraceae
(Ahnfeltia, Ceratocolax, Gymnogongrus and Phyllophora, Figures
138, 158, 177 and 195-197); Gigartinaceae (Chondrus and Gigartina,
Figures 160 and 173); Corallinaceae (Clathromorphum, Corallina,
Dermatolithon, Fosliella, Leptophytum, Lithophyllum, Lithotham-
nion and Phymatolithon, Figures 162, 163, 167, 172, 181-184 and
198-200); Kallymeniaceae (Callocolax and Callophyllis, Figures 152
and 153); Choreocolaceae (Choreocolax and Harveyella, Figures
161, 179); Palmariaceae (Devaleraea, Halosacciocolax, Palmaria
and Rhodophysema, Figures 168, 178, 190, 223 and 224); Ceramia-
ceae (Antithamnion, Antithamnionella, Callithamnion, Ceramium,
Plumaria, Pterothamnion, Ptilota, Scagelia and Spermothamnion,
Figures 139, 140, 149-151, 154-157, 201, 219, 220, 226 and 227);

1986] Mathieson & Hehre — New Hampshire seaweeds 31

Delesseriaceae (Membranoptera, Pantoneura and Phycodrys, Fig-
ures 188, 191 and 194); Rhodomelaceae (Chondria, Polysiphonia
and Rhodomela, Figures 159, 203-212 and 221); Erythropeltidaceae
(Erythropeltis, Erythrotrichia and Porphyropsis, Figures 170, 171
and 218); Bangiaceae (Bangia and Porphyra, Figures 147 and
213-216). The members of the Gigartinaceae can be cited as specific
examples (Figures 160 and 173); both Chondrus and Gigartina
exhibit cosmopolitan distribution patterns, although G. stellata has
the most restricted outer estuarine pattern and a conspicuous reduc-
tion of stature in estuarine habitats (Burns and Mathieson, 1972b).

Based upon the data in Tables I-III plus previous floristic studies

(Hehre and Mathieson, 1970; Mathieson and Hehre, 1982, 1983), it
is apparent that several endophytic, epiphytic and parasitic sea-
weeds and their respective “hosts” demonstrate contrasting distribu-
tional patterns. The following taxa can be cited as examples:

1. endophytic Mikrosyphar porphyrae growing in various Por-
phyra species, particularly P. umbilicalis (Figures 108, 213-216)

2. epiphytic Ascocyclus distromaticus growing on Palmaria pal-
mata (Figures 76, 190); Elachista fucicola on Ascophyllum
nodosum (Figures 77, 91); Laminariocolax tomentosoides and
Myrionema corunnae on various Laminaria species (Figures
103-106, 109); Protectocarpus speciosus on Chaetomorpha
aerea (Figures 21, 115); Vlonema rhizophorum on Dumontia
contorta (Figures 137, 169); Audouinella alariae on Alaria
esculenta (Figures 75, 141); and A. violacea on Sacheria fucina
(Figures 146, 225)

3. parasitic Callocolax neglectus growing on Callophyllis cristata
(Figures 152, 153); Ceratocolax hartzii on Phyllophora trun-
cata (Figures 158, 197); Choreocolax polysiphoniae on Polysi-
Phonia lanosa (Figures 161, 207); Halosacciocolax kjellmani
on Palmaria palmata (Figures 178, 190); Harveyella mirabilis
on Rhodomela confervoides (Figures 179, 221); and Polysi-
Phonia lanosa on Ascophyllum nodosum (Figures 77, 207).

Polysiphonia lanosa (Figures 77, 207), Laminariocolax tomento-
soides (Figures 103-106) and Choreocolax polysiphoniae (Figures
161, 207) are representative of the above-described species, except
for Audouinella alariae (Figures 75, 141) and A. violacea (Figures
146, 225). That is, the hemiparasite P. /anosa, which grows abun-

32 Rhodora [Vol. 88

dantly on A. nodosum on the open coast, is restricted to outer
estuarine sites, even though its host is abundant and widely distrib-
uted (Fralick and Mathieson, 1975). The common epiphyte L.
tomentosoides and the specific parasite C. polysiphoniae show a
similar restricted estuarine distribution versus their hosts. Audoui-
nella alariae and A. violacea were the only species with approxi-
mately the same distribution patterns as their hosts Alaria esculenta
and Sacheria fucina, respectively.

As noted by Dixon (1965), a comparison of the distributional
patterns of life history stages of individual taxa can be quite infor-
mative. Deviations from a “theoretical” life history can occur geo-
graphically due to perennation and various selective mechanisms
operating against a particular genome (Mathieson and Burns, 1975;
Mathieson and Norall, 1975a, b; Norall et al., 1981). Dixon (1965)
described the example of gametophytic Asparagopsis armata and
tetrasporic Falkenbergia rufolanosa, which may exhibit independ-
ent vegetative propagation and deviations from their “theoretical”
life histories at northern latitudes. The recently recorded differences
in geographical distribution in Europe for the two phases (Conway,
1960: Thomas, 1955) may be a reflection of independent vegetative
propagation. In comparing the coastal and/or estuarine distribution
patterns of different life history phases of New England seaweeds
(Tables I-III), several geographical contrasts are evident, perhaps
due to the strong environmental gradient within these areas (see
earlier description), the different physiological tolerances of various
phases (Mathieson and Burns, 1975; Mathieson and Norall, 1975a,
b: Norall et al., 1981), the different modes and magnitude of vegeta-
tive propagation, and other factors. For example, the “Codiolum
gregarium/pusillum” sporophytic stages of Urospora and Ulothrix
spp. (Kornmann and Sahling, 1977; Scagel, 1966; South, 1984) have
a more localized estuarine distribution than their corresponding
gametophytic phases (Figures 32, 34, 67, 72, 73). A similar trend is
evident in Figures 33 and 64 for the endophytic “Codiolum petroce-
lidis” sporophyte of Spongomorpha spinescens (Jonsson, 1958,
Scagel, 1966). Both gametophytic “Halicystis ovalis” and sporo-
phytic Derbesia marina (Sears and Wilce, 1970) are restricted to the
open coast, with the former being rare and the latter more common
(Figures 36, 48). The crustose sporophytic “ Ralfsia bornetii/clavata™
and foliose gametophytic stages of Petalonia fascia (Edelstein et al.,

1986] Mathieson & Hehre — New Hampshire seaweeds 33

1970) both exhibit cosmopolitan distributional patterns, although
the foliose stage is more widely distributed than the crustose phase
(Figures I11, 119, 120). Similarly, the crustose tetrasporophyte
“Porphyrodiscus simulans” of Ahnfeltia plicata (Farnham and
Fletcher, 1976) was only found at one estuarine site, while the game-
tophytic phase was collected at diverse open coastal and estuarine
habitats (Figures 138, 217). The sporophytic “Trai/liella intricata”
phase of Bonnemaisonia hamifera (Chihara, 1961, 1962), also ex-
hibits a more localized distribution than its gametophytic phase
(Figures 148, 228). Lastly, the crustose sporophytic “ Petrocelis
cruenta” and upright gametophytic phases of Gigartina stellata
(Fletcher and Irvine, 1982; Guiry and Coleman, 1982: West and
Polanshek, 1975; West et al., 1977) both extend from the open coast
into the outer-mid portions of the Great Bay Estuary System (Fig-
ures 173, 192).

Several unique ecological or phenotypic patterns were also evi-
dent. For example, the perennial psammophytic “sand-loving”
(Mathieson, 1982b) brown alga Sphacelaria radicans was restricted
to a few sand-abraded open coastal (Daly and Mathieson, 1977) and
sandy outer estuarine habitats (Figure 133). Further, attached popu-
lations of Ascophyllum nodosum were collected abundantly at
diverse open coastal and estuarine sites (Figure 77), while the ecad
scorpioides was restricted to sheltered estuarine sites (Figure 78),
entangled amongst Spartina alterniflora (Chock and Mathieson,
1976). Two examples of phenotypic plasticity should also be noted.
The fucoid brown alga Fucus vesiculosus primarily exhibits a spi-
raled morphology (var. spiralis in Taylor, 1957) in estuarine habi-
tats, while the typical non-spiraled plant is most abundant in open
coastal habitats, particularly exposed sites. Locally, most popula-
tions of Cystoclonium purpureum have tendril-like branches (var.
cirrhosum in Taylor, 1957); even so, some estuarine plants exhibit
radiating burr-like branches (forma ste//atum, Collins, 1906b),
which Taylor (1957) suggested are a pathological state. Although
South (1984) and others suggested that subspecific taxa of C. pur-
pureum are insufficiently distinct to warrant retention, the
“stellatum-type” morphology seems to be restricted to sheltered
estuarine sites.

In comparing the local distribution of plants based upon culture
and in situ collections (Tables I-III), several interesting contrasts

34 Rhodora [Vol. 88

can be made. As noted earlier, four taxa were only recorded in
culture (Acrochaete repens, Bolbocoleon piliferum, Prasinocladus
marinus and Sphaerotrichia divaricata, Figures 16, 18, 56, 134),
while the local distributional records of an additional four taxa
(“Codiolum pusillum,” Microspora pachyderma, Desmotrichum
undulatum and Isthmoplea sphaerophora, Figures 34, 50, 87, 102)
were supplemented by culture information. Thus, the single estua-
rine record of “C. pusillum” was based upon culture findings, while
similar statements can be made about the solitary nearshore open
coastal records for M. pachyderma, D. undulatum and I. sphaero-
Phora. Presumably these culture records are based upon the plants
being rare in nature, cryptic in size, and juvenile and adult stages
having different physiological tolerances/ optima, or other factors.
In this context, Mathieson and Hehre (1983) noted that attached
populations of freshwater algae like M. pachyderma are typically
restricted to inner estuarine/ riverine habitats. Even so, juvenile (cul-
tured) populations of M. pachyderma exhibit a wide tolerance to
salinity, which suggests that other biological factors may restrict the
plant’s growth in situ (Zechman and Mathieson, 1985). Several
unique culture records for Enteromorpha compressa, Spongo-
morpha arcta, Ulvaria oxysperma, Urospora wormskioldii, Des-
motrichum undulatum and Porphyropsis coccinea should also be
noted (Figures 38, 63, 71, 73, 87, 218). For example, the ubiquitous
“estuarine” alga U. oxysperma was only found at four open coastal
locations, one of which was based upon its presence in culture (Fig-
ure 71).

PHYSIOLOGICAL ECOLOGY AND DISTRIBUTIONAL PATTERNS
OF SELECT SEAWEEDS

Several estuarine taxa, or seaweeds that are rare on the open
coast of New Hampshire, represent disjunct populations north of
Cape Cod, Massachusetts, on the northeast coast of North America:
Bryopsis plumosa (Figure 19); Ulvaria oxysperma (Figure 71); Anti-
thamnion cruciatum (Figure 139); Ceramium strictum (Figure 157);
Chondria baileyana (Figure 159); Dasya baillouviana (Figure 166);
Gracilaria tikvahiae (Figure 176); Lomentaria baileyana (Figure
185); Polysiphonia denudata (Figure 203); and P. subtilissima (Fig-
ure 211). These “southerly” taxa are more widely distributed south
than north of Cape Cod and several of them extend to the tropics,

1986] Mathieson & Hehre — New Hampshire seaweeds 35

Bermuda, Florida, etc. (Taylor, 1957, 1960). At their northern dis-
tributional limits, each of these plants occurs primarily in shallow
embayments or protected habitats such as the Great Bay Estuary
System or Northumberland Straits near Prince Edward Island, New
Brunswick, Canada. All of these species, except for G. tikvahiae, are
summer annuals (Tables I and III) at their northern limits, and they
may have “modified” life histories (Mathieson and Burns, 1975:
Norall et al., 1981) and extensive vegetative reproduction. As noted
earlier, the hydrographic conditions within such northern latitudes
are much more variable than within the central portion of the
plant’s geographical range. Thus, the phenologies of such “south-
erly” species are often conspicuously different in northern than in
southern geographies (Hehre and Mathieson, 1970; Mathieson and
Dawes, 1975).

Bousfield and Thomas (1975) have recorded similar disjunct pat-
terns for many shallow water marine animals between Cape Cod, the
northern Gulf of Maine and/or the Gulf of St. Lawrence. Many
animal populations having similar temperature requirements are
isolated from each other by hundreds of miles of climatically “un-
favorable” marine coastlines, particularly during the reproductive
period that is critical to natural dispersal and to maintenance of
homogenous populations. The same authors speculated that the
most satisfactory explanation of this distribution is an historical
one; during a “hypsithermal” or warm period 7500-9500 years ago,
the relatively shallow shelf waters between Cape Cod and the Gulf
of St. Lawrence provided a uniform summer-warm environment
and dispersal pathway. Subsequent drowning and deeping of the
inshore coastal areas and increased upwelling in the Gulf of Maine
during the past 5000 years have depressed the summer temperatures
to present-day cool levels. In “post-hypsithermal” times the warm
water fauna gradually disappeared from the cooling open coastal
areas, and populations like those in the Gulf of St. Lawrence
became regionally restricted and effectively isolated from the main
populations in southern New England and further south. McAlice
(1981) has given a similar explanation for the post-glacial history of
the copepod Acartia tonsa in the Gulf of Maine and the Gulf of St.
Lawrence. He suggested that the northern populations of this
copepod, which occur in warm estuarine headwaters north of Cape
Cod, are relict ones, derived from a distribution that was once contin-

36 Rhodora [Vol. 88

uous from Cape Cod to the Northumberland Strait (New Brunswick-
Prince Edward Island). He further suggested that the disjunction of
A. tonsa at its present refuges may make it useful for studies on rates
of speciation.

The ideas presented by Bousfield and Thomsa (1975) and McAI-
ice (1981) with respect to disjunct animal distributions would appear
to be applicable to seaweeds as well. With the advent of cooling
coastal waters, populations of some seaweeds may have been forced
into warm estuarine habitats, while the main coastal populations
receded southward. If, as McAlice (1981) suggested, these relict
populations became reproductively isolated due to lower water
temperatures and westerly currents, then the potential was estab-
lished for genetic differentiation. The likelihood of this occurrence
would be increased by differences in environmental conditions
imposed upon the populations at their respective locations. Although
northern estuaries provide warm summer temperatures required for
the growth of “southerly” species, these estuarine populations must
tolerate reduced and/or fluctuating salinities and extremely cold
winter conditions. For example, the red alga Polysiphonia subtilis-
sima is primarily restricted in New Hampshire- Maine to inner riv-
erine habitats (Figure 211) where temperatures vary from 0-26°C,
and salinities from 0-22 %o (Norall and Mathieson, 1976). Since P.
subtilissima grows in northern locations as a pseudoperennial,
regenerating from perennating holdfast filaments (Hehre and
Mathieson, 1970; Yarish and Edwards, 1982), it must tolerate this
entire range of conditions. South of Cape Cod, the same species
may also occur on the open coast (Mathieson and Dawes, 1975)
where it is exposed to more uniform temperatures and stable, coas-
tal salinities. Consequently, broader temperature and salinity toler-
ances and lower optima would be of adaptive significance in
northern disjunct seaweed populations, while the same attributes
would be relatively unimportant to the continuous “southerly” coas-
tal populations.

Fralick and Mathieson (1975) and Mathieson and Burns (1971),
among others, have attempted to correlate the physiological ecology
and estuarine distributional patterns of several seaweeds. For
example, Fralick and Mathieson (1975) compared the photosynthe-
sis and respiration of four species of Polysiphonia under different
light, temperature and salinity conditions and found that they could
be separated into a “cold water” group (P. /anosa and P. elongata)

1986] Mathieson & Hehre — New Hampshire seaweeds 37

and a group with warm-water affinities (P. nigrescens and P. subti-
lissima). They speculated that the horizontal distribution of these
four Polysiphonia taxa within the Great Bay Estuary System
(Figure 204, 207, 209, 211) was primarily governed by their varying
tolerances to high temperatures and low salinities. Thus, P. subtilis-
sima, which had the highest temperature optimum, penetrated the
furthest into the estuary (Figure 211), while P. lJanosa, which had the
lowest temperature optimum, was restricted to the more coastal
stations (Figure 207). Mathieson and Burns ( 1971) conducted a sim-
ilar physiological study of the closely-related gigartinaleanm red
algae Chondrus crispus and Gigartina stellata. Both species exhi-
bited broad tolderances to salinity. Even so, C. crispus showed its
maximum photosynthesis and minimum respiration at 24 %o. in
agreement with the more open coastal habitat of Gigartina and the
more estuarine habitat of Chondrus (Figures 160, 173). Culture
studies of juvenile stages (sporelings from carpospores) from both
plants have demonstrated a similar restricted tolerance to reduced
salinities for G. stellata as compared to C. crispus (Burns and
Mathieson, 1972a). The above-described physiological and culture
studies, including those of Zechman and Mathieson (1985), demon-
strate the potential for experimentally evaluating the distributional
patterns of diverse seaweeds (Hoek, 1982a, b).

ACKNOWLEDGMENTS

We express our sincere gratitude to the following people: “Ned”
McIntosh, former captain of the R/V Jere A. Chase, who assisted
with the diving operations; a variety of marine phycologists (past
and present) at the University of New Hampshire who have helped
with many of the collections, including Steven Blair, Richie Burns,
Paula Busse, Donald Cheney, Jan Chock, Joan Conway-Lockhart,
Marty Costa, Maureen Daly, Richard Femino, William Flahive,
Steven Fuller, Phelps Fullerton, Barry Hutchinson, John Kilar,
Mike Josselyn, Cindy Mathieson, Ernani Menez, Chris Neefus.
Richard Niemeck, Timothy Norall, Chris Emerich Penniman, Clay-
ton Penniman, Norman Reynolds, John Shipman, Eric Sideman,
Eleanor Tveter-Gallagher, George Vagenas, and Tim Voorheis.
Chris Emerich penniman is also acknowledged for her help in the
preparation of several figures. The first author would also like to
gratefully acknowledge the encouragement, support and help of his
wife, Myla Jean Mathieson.

38 Rhodora [Vol. 88

List of Tables

I. Seasonal occurrence, longevity and local distribution of Chlorophyceae
Il. Seasonal occurrence, longevity and local distribution of Phaeophyceae
Ill. Seasonal occurrence, longevity and local distribution of Rhodophyceae
IV. Summary of collection sites

Key to Tables I-III
X = present

* = obtained in culture
** — residual basal material

Longevity: Ann. = annual
AAnn. = aseasonal annual
Per. = perennial
PPer. = pseudoperennial

= Isles of Shoals

= Nearshore open coast between Portsmouth and Seabrook
Hampton-Seabrook Estuary System

= Great Bay Estuary System

Local Distribution:

RwWwnh =
\|

|

Table I: Seasonal occurrence, longevity and local distribution of Chlorophyceae

[9861

om Local
Taxa JF MAMJS J A S ON D Longevity Distribution
Acrochaete repens N. Pringsh. X* X* Ann. 2*, 4*
Blidingia minima (Nag. ex Kiitz.) Kylin X KX X X X KX XK KX KX X X X_~— AAnn. 1-4
Bolbocoleon piliferum N. Pringsh. xX* xe Ann. 2
Bryopsis plumosa (Huds.) C. Ag. xX** X X X X X X X_— Ann. 1-4
Capsosiphon fulvescens (C. Ag.) Setch. et Gardn. X* X X X X X X X X_— Ann. 2,4
Chaetomorpha aerea (Dillw.) Kiitz. xX X X X X K XK X X X X X Per. 1,2,4
Chaetomorpha brachygona Harv. xX x xX X X X X X X X_— Ann. (?) 1,2,4
Chaetomorpha linum (O. F. Miill.) Kiitz. X X X X X KX X X X X X X Per. 1-4
Chaetomorpha melagonium (Web. et Mohr) Kiitz, X X X X X X X X X X X X Per 1, 2,4
Chaetomorpha minima Collins F. et Herv. xX X Ann. (?) |
Chaetomorpha picquotiana Mont. ex. Kiitz. xX X X X KX KX KX X X X X X Per. 1-4
Chlorochytrium moorei Gardn. xX Ann. 2
Cladophora albida (Huds.) Kiitz. X xX X X X X XxX AAnn, 1, 2,4
Cladophora pygmeaea Reinke X X » ae, aa, Ga, an, Same, © Per. 4
Cladophora rupestris (L.) Kiitz. X xX xX Xx Per. 1,2
Cladophora sericea (Hud.) Kiitz. X & X K X KX X X X X X X_ AAnn. 1-4

or PPer.

“Codiolum gregarium A. Braun” xX* X x Ann. 2,4

Spoomeas auiysdwuepy MIN — IYI * uosatyIepy

6t

Table I: (Cont.)

Month
Local

Taxa JF MA MJ J A S O N D Longevity Distribution
“Codiolum petrocelidis Kuck.” »,4 XxX Ann. (?) 1,2
“Codiolum pusillum (Lyngb.) Kjellm.” x xX xX X X X X X_— Ann. 1, 2, 4*
Codium fragile (Sur.) Hariot

subsp. tomentosoides (van Goor) Silva x x Per. 1
Derbesia marina (Lyngb.) Solier D4 », 4 Ann. 1,2

or PPer.

Enteromorpha clathrata (Roth) Grev. x x X X XK X X XX _— Ann. 1-4
Enteromorpha compressa (L.) Grev. xX X X X xX X X X XK X AAnn. 1-4
Enteromorpha flexuosa (Wulf. ex Roth) J. Ag.

subsp. flexuosa Bliding x X xX Ann. o
Enteromorpha flexuosa (Wulf. ex Roth) J. Ag.

subsp. paradoxa (Dillw.) Bliding x x X X X X X X_ Ann. 1-4
Enteromorpha intestinalis (L.) Link xX X X X xX X X X XK X XX AAnn. 1-4
Enteromorpha linza (L.) J. Ag. xX X X X xX X X X X X X - AAnn. 1-4
Enteromorpha prolifera (O. F. Miill.) J. Ag. xX X X X xX X X X X X X_ AAnn. 1-4
Enteromorpha torta (Mert. in Jiirg.) Reinb. xX xX X X X xX ~~ Ann. 2,4
Entocladia flustrae (Reinke) Batt. x x x Ann. (?) 1,2
Entocladia viridis Reinke xX xX X X xX X X X X X X _~ AAnn. 1,2,4
Gomontia polyrhiza (Lagerh.) Bornet et Flah. Xx x Ann. (?) 1,2

OV

elopoyy

88 TOA]

Table I: (Cont.)

ea Local
Taxa F M.S» J S N D Longevity Distribution
“Halicystis ovalis (Lyngb.) Aresch.” X Per. (?) 2
Kornmannia leptoderma (Kjellm.) Bliding xX X X x Ann. 2,4
Microspora pachyderma (Wille) Lagerh. 4 X X* X* xX* Ann. 2.4
Monostroma grevillei (Thur.) Wittr. x xX X X X= Ann. 1-4
Monostroma pulchrum Farl. xX XxX Ann. 1, 2,4
Mougeotia sp. Xx Ann. 4
Oedogonium sp. x Ann 4
Percursaria percursa (C. Ag.) Rosenv. X xX X X xX X XX AAnn. 1-4
Prasinocladus marinus (Cienk.) Waern xX Ann 2°
Prasiola stipitata Suhr. in Jessen xX X X X x xX X AAnn. 1,2,4
Pringsheimiella scutata (Reinke) Marchew. Ann. (?) I
Pseudendoclonium submarinum Wille X Xx AAnn. 1,4
Rhizoclonium riparium (Roth) Harv. . Ew Xx ».4 AAnn. 1-4
Rhizoclonium tortuosum (Dillw.) Kiitz. K- TK X Xx AAnn. (?) 12,4
Spirogyra sp. Xx Ann 4
Spongomorpha arcta (Dillw.) Kiitz. x X X x xX xX Ann ese
Spongomorpha spinescens Kitz. Xx X X Ann 1,2,4
Stichococcus marinus (Wille) Hazen 4 Ann. |

[9861

Spoamevas aitysdwiey MON — dI1YdH % uosaryI ep

Table I: (Cont.)

44

Month j ocal
Taxa JI FMA MJ J A S ON D Longevity Distribution
Stigeoclonium sp. Xx Ann. =
Ulothrix flacca (Dillw.) Thur. in Le Jolis xX xX X X X X xX X XxX Ann. 1, 2,4
Ulothrix speciosa (Carm. ex Harv. in Hook.) Kiitz. xX X X X Ann. 1, 2,4
Ulva lactuca L. xX X X X X KX KX X X X X X Ann. 1-4

or PPer.

Ulvaria obscura (Kiitz.) Gayral xX X X KX KX X X XK X X X X Ann. 1-4
Ulvaria oxysperma (Kiitz.) Bliding xX X X X KX X K X X X X X Ann. 1-4
Urospora penicilliformis (Roth) Aresch. xX X X X XK XK X X xX X X Ann. 1,2,4
Urospora wormskioldii (Mert. in Hornem.) Rosenv. X X X X X xX xX* X Ann. 1,2,4
Monthly total 24 32 36 34 33 36 42 40 30 37 34 27

Chlorophyta taxa

elOpoyYy

88 10A]

Table II: Seasonal occurrence, longevity and local distribution of Phaeophyceae

Month

Local

Taxa A MJ J AS O D Longevity Distribution
Agarum cribrosum (Mert.) Bory XxX X X KX X X X X xX Per. 1,2,4
Alaria esculenta (L.) Grev. Xx X X KX X X X X Per. 1,2
Ascocyclus distromaticus Tayl. xX XxX X X X Ann 1,2
Ascophyllum nodosum (L.) Le Jol. Mm MK, OOOO TX XX. UX X Per. 1-4
Ascophyllum nodosum (L.) Le Jol.

ecad scorpioides (Reinke) Hauck A. OM XO CK GX KX. xX Per 3,4
Asperococcus fistulosus (Huds.) Hook. x X X X X Ann 1,2
Chorda filum (L.) Stackh. Xx X X X X Ann 1-4
Chorda tomentosa Lyngb. xX X X Xx Ann 1, 2,4
Chordaria flagelliformis (O. F. Mill.) C. Ag. X X X K K X X X X Ann 1-4
Cladostephus spongiosus (Huds.) C. Ag.

forma verticillatus (Lightf.) Post. et Rupr. Per. (?) l
Delamara attenuata (Kjellm.) Rosenv. KX XOX Ann 1,4
Desmarestia aculeata (L.) Lamour. xX Xx X Kee XX x Per 1,2,4
Desmarestia viridis (O. F. Mill.) Lamour. xX X X xX xX Ann 1, 2,4
Desmotrichum undulatum (J. Ag.) Reinke xX XX X x Ann lL, 2*, 4
Dictyosiphon foeniculaceus (Huds.) Grev. xX X X X XK X X X Ann 1-4
Ectocarpus fasciculatus Harv. xX X X X X x Ann 1, 2,4
Ectocarpus siliculosus (Dillw.) Lyngb. X X X X X X X xX Ann 1-4

[9861

spoameas oilysdwiey MON — IY4dH 3? uosoaep]

tv

Table II: (Cont.)

Local

Taxa J oF M MJ J A S O N D Longevity Distribution
Elachista fucicola (Vell.) Aresch. xX X X X X X X X X XK X Per. 1-4
Eudesme virescens (Carm. ex Harv. in Hook.) J. Ag. 4 Ann 2
Fucus distichus L.

subsp. anceps (Harv. et Ward ex Carr.) Powell X X X Xx X X X X X X X Per. l
Fucus distichus L.

subsp. distichus L. emend. Powell xX X X X KX X KX X X XK X Per. 1, 2,4
Fucus distichus L.

subsp. edentatus (Pyl.) Powell xX X X X X X X X X XK X Per. 1-4
Fucus distichus L.

subsp. evanescens (C. Ag.) Powell xX X X X X X X X X X X Per. 1, 2,4
Fucus spiralis L. xX X xX X X X X X X X X _~. Per. 1,2,4
Fucus vesiculosus L. xX X X X X X X X X X X _. Per. 1-4
Giffordia granulosa (Sm.) Hamel X X X XK X X X X X~ Ann 1-4
Giffordia sandriana (Zanar.) Hamel xX X Ann 2,4
Giffordia secunda (Kiitz.) Batt. x Ann 2
Hecatonema terminalis (Kiitz.) Kylin Ann 2?
Isthmoplea sphaerophora (Carm. ex Harv. in

Hook.) Kjellm. xX X X* X Ann. 1, 2*,4
Laminaria digitata (Huds.) Lamour. xX xX X X X X X X X XK X Per. 1-4

a4

elopoyy

88 10A]

Table Il: (Cont.)

Taxa FMAMJS J A S ON D Longevity ee
Laminaria longicruris Pyl. X XM X X KX X XK X X Per. 124
Lamanaria saccharina (L.) Lamour. xX X& X KX K KX KX KX X X X ~~ Per 1-4
Laminariocolax tementosoides (Farl.) Kylin x xX x x X Ann. (?) 1.2
Leathesia difformis (L.) Aresch. xX X X KX KX K X X Ann ],.2
Mikrosyphar porphyrae Kuck. x » re, « Ann | (+ Cape
Neddick, ME)
Myrionema corunnae Sauvy. xX X X X X KX X X X X Ann 1, 2,4
Myrionema magnusii (Sauy.) Lois. Ann 2?
Myrionema strangulans Grey. x 4 Ann 1, 2,4
Petalonia fascia (O. F. Mill.) Kuntze MM. Mee XX. KX X KR XK Ann 1-4
Petalonia zosterifolia (Reinke) Kuntze xX X X x xX -& Ann 1,2,4
Petroderma maculiforme (Wolny) Kuck. x BM XK DOAeXP Ke: X XK Xx ~ Per: 2,4
Pilayella littoralis (.) Kjellm. Xx X& X KX KX X XK XK X X X= Ann 1-4
Protectocarpus speciosus (Bgrgesen) kuck. X X X X X XR 1X Ann. (?) 1:2
Pseudolithoderma extensum (Crouan frat.) S. Lund X X MX. Oe. CRUX. XX. EX XK Per. 1,2,4
Punctaria latifolia Grey. xX X X XK X Ann 4
Punctaria plantaginea (Roth) Grev. x X X Ann Le 2
“Ralfsia bornetii Kuck.” xX X X X X xX xX x Per. (?) 1-4

[9861

spoomras ailysdwey MON — d14YdH % uosaIpIeEp

SP

Table II: (Cont.)

4

Local
Taxa MJ J A O N D Longevity Distribution
“ Ralfsia clavata (Harv. in Hook.) Crouan frat.” X X X X X X_. Per. (?) 1-4
Ralfsia fungiformis (Gunn.) Setch. et Gardn. xX X X X x Per. 2,4
Ralfsia verrucosa (Aresch.) J. Ag. xX X X X X XK X Per. 1-4
Saccorhiza dermatodea (Pyl.) J. Ag. xX X X X Ann. 1,2
Scytosiphon lomentaria (Lyngb.) Link
var. complanatus Rosenv. Ann. =
Scytosiphon lomentaria (Lyngb.) Link e
var. /omentaria xX X X X X xX X_ Ann. 1-4 Q.
Sorapion kjellmanii (Wille) Rosenv. x Per. 2 (+ York 3
River, ME)
Sorocarpus micromorus (Bory) Silva x Ann. 4
Sphacelaria arctica Harv. X X X Per. (?) 2
Sphacelaria cirrosa (Roth) C. Ag. X X X X xX X X_ Per 1,2,4
Sphacelaria fusca (Huds.) S. F. Gray X Per. (?) l
Sphacelaria plumigera Holmes xX xX Per. 2
Sphacelaria plumosa Lyngb. xX xX Per. (?) 2
Sphacelaria radicans (Dillw.) C. Ag. xX xX X X xX xX Per. 2,3 —
Sphaerotrichia divaricata (C. Ag.) Kylin X* Ann 2 S
Spongonema tomentosum (Huds.) Kitz.) x xX X xX X Per. (?) 1,2,4 ce

Table II: (Cont.)

Taxa

Stictvosiphon griffithsianus (Le Jol.)
Holm. er Batt.

Ulonema rhizophorum Fosl.

Montly total
Phaeophyta taxa

JF M A M
xX
X X X X

25 34 36 43 46

Month

Local
J J A S O N D Longevity Distribution
xX Ann. 4
>, a, ca. 4 D4 Ann. 1, 2,4
48 50 55 42 36 38 27

[9361

Spaameas oitysdwey MAN — IY4IH % uosaryep

Lv

Table III: Seasonal occurrence, longevity and local distribution of Rhodophyceae

Month Local

Taxa F MA MJ J A S O N D Longevity Distribution
Ahnfeltia plicata (Huds.) Fries X X X X X X X X X XK X Per. 1-4
Antithamnion cruciatum (C. Ag.) Nag. X X X X X KX X XK X X X~ Ann 2,4
Antithamnionella floccosa (O. F. Miill.) Whittick X X KX X X X KX X X X X ~~ AAnn. 1,2,4
Audouinella alariae (Jonss.) Woelk. xX X X xX Ann 1,2
Audouinella membranacea (Magn.) Papenf. X X X X XK X XK X X XK X Per. (?) 1,2,4
Audouinella polyides (Rosenv.) Garbary xX Ann 2
Audouinella purpurea (Lightf.) Woelk. X XK X X X X KX X XK K X Per. 1,2,4
Audouinella secundata (Lyngb.) Dixon X X X X KX X KX XK X XK X_ AAnn. 1-4
Audouinella violaceae (Kiitz.) Hamel x X X X XK XK X X X~ Ann 4
Bangia atropurpurea (Roth) C. Ag. X X XK X X x xX X Ann 1,2,4
Bonnemaisonia hamifera Hariot X X X X KX KX X KX XK X Per. 1, 2,4
Callithamnion byssoides Arnott ex Harv. in Hook. xX xX Ann 4
Callithamnion hookeri (Dillw.) S. F. Gray xX X X Ann 1,2,4
Callithamnion tetragonum (With.) S. F. Gray X X X X XK XK XK XK XK XK X ~~ Per. 1-4
Callocolax neglectus Schm. ex Batt. X X X XK XK XK X XK XK X Per. (?) 1, 2,4
Callophyllis cristata (C. Ag.) Kiitz. X X X X XK KX X K X X X ~~ Per 1-4
Ceramium deslongchampii Chauy. in Duby

var. hooperi (Harv.) Tayl. XxX X X XK X X X XK X Per. (?) 1,2,4

8P

elopouy

88 10A]

Table III: (Cont.)

Month

Taxa JF MAMdJ J AS ON D Longevity bee ies
Ceramium elegans (Ducluz.) C. Ag. - x Ann. 4
Ceramium rubrum (Huds.) C. Ag. a B&R ROM OK. KK RM KM: Per: 1-4
Ceramium strictum Harv. x X X X X X X X X~ Ann 2-4
Ceratocolax hartzii Rosenv. X X X X X Ka Nek MN RM XK Per 1,2
Chondria baileyana (Mont.) Harv. x X X X X X Ann 2,4
Chondrus crispus Stackh. X X X X X X XK KX KX X X X~ Per. 1-4
Choreocolax polysiphoniae Reinsch xX X X X KX KX X X X X X X ~~ Per 1, 2,4
Clathromorphum circumscriptum (Stromf.) Fos X X X X X X X X X X X X_ Per 1-4
Corallina officinalis L. Xx X X X X X X X X X X X Per. 1, 2,4
“Cruoriopsis ensis Jao” X 4 Per. (?) 2,4
Cystoclonium purpureum (Huds.) Batt. am EK RN eR Xe NR ee 1-4
Dasya baillouviana (Gmel.) Mont. Xe K+ SEX Pe XK OX OE Ann 4
Dermatolithon pustulatum (Lamour.) Fosl. Mm OM Oe OX, OX XR EX OK OX. OX Per 1,2,4
Devaleraea ramentaceum (L.) Guiry Me OR OT OX, EXPE KY EX? 2X Per 1,2
Dumontia contorta (Gmel.) Rupr. Mm OX UX OX OX OR OR OKO OX Ann 1-4
Erythropeltis discigera (Berth.) Schm.

var. discigera x x Ann [2
Erythrotrichia carnea (Dillw.) J. Ag. X Xx X XM X X X X Ann 1-4

Spoamevas oitysdwiey MON — 49H % UosapeEp Log6l

60

Table III: (Cont.)

— Local

Taxa JI F MA MJ J A S O N D Longevity Distribution
Fosliella lejolisii (Rosan.) Howe x X X X KX X KX XK X X X Per. 1-4
Gigartina stellata (Stackh.) Batt. XxX X X X X X XK XK KX XK XK X ~~ Per. 1,2,4
Gloiosiphonia capillaris Carm. in Berk. xX X X X Ann. 1,2,4
Goniotrichum alsidii (Zanard.) Howe ».4 xX X X X Ann. 1,4
Gracilaria tikvahiae McLachlan X X KX X XK XK X X X XK X X ~~ Per. 4
Gymnogongrus crenulatus (Turn.) J. Ag. X XK X X KX X KX KX XK KX XK X _~ Per 1,2,4
Halosacciocolax kjellmanii S. Lund XxX Per. (?) l
Harveyella mirabilis (Reinsch.) Schm. et

Reinke in Reinke X X X X Per. 1,2
Hildenbrandia rubra (Sommerf.) Menegh. X X X X X KX KX KX X KX X X Per. 1-4
Leptophytum foecundum (Kjellm.) Adey xX X xX X X X X KX X X X Per. 1,2
Leptophytum laeve (Stromf.) Adey xX X X X X X X X KX X XK X~ Per 1,2,4
Lithophyllum corallinae (Crouan frat.) Heydr. xX X X X X X X X KX X X X ~~ Per 1,2,4
Lithothamnion glaciale Kjellm. X X X X X K X X XK XK XK X Per 1,2,4
Lomentaria baileyana (Harv.) Farl. X X X X X XK X_~— Ann. 4
Lomentaria clavellosa (Turn.) Gaillon xX xX xX KX KX X X X X X X _~ Per. (?) 4(+ Boone L,

Maine

Lomentaria orcadensis (Harv.) Coll. ex Tayl. 4 xX X KX X X Per 1,2,4
Membranoptera alata (Huds.) Stackh. xX X X X KX KX XK X XK X XK X ~~ Per. 1,2,4

0S

eIOpoyy

88 10A]

Table III: (Cont.)

ons Local
Taxa F A MJ JA S ON D Longevity Distribution
Nemalion helminthoides (Vell. in With.) Batt. xX X X Ann. 1,2
Palmaria palmata (L.) O. Kuntze xX xX XM X KX X X X X X ~~ Per. 1-4
Pantoneura baeri (Post. et Rupr.) Kylin xX X Per. l
“Petrocelis cruenta J. Ag.” x M XX SR OX OK XK OX. OX Per. 1, 2,4
Peyssonnelia rosenvingti Schm. in Rosenv. xX m: XA AX, Ok Oe AX Per. 1,2,4
Phycodrys rubens (L.) Batt. X X KX KX X X X X X X _~ Per, 1-4
Phyllophora pseudoceranoides (Gmel.) New.

et A. Tayl. X X X KX X X X X X X Per. 1,2,4
Phyllophora traillii Holm. ex Batt. xX X X xX XxX Per. 1,2
Phyllophora truncata (Pallas) A. Zin. X X X XK X X X X X X_. Per. 1, 2,4
Phymatolithon laevigatum (Fosl.) Fosl. x Xx X X KX X KX X X X Per. 1,2,4
Phymatolithon lenormandii (Aresch. in J. Ag.)

Adey X X X X X X X X X X_ Per 1,2,4
Phymatolithon rugulosum Adey X X X X XM Dey Per: ly?
Plumaria elegans (Bonnem.) Schm. x a Ne ee. SX OX Per. 1-3
Polvyides rotundus (Huds.) Grev. x DM, ON Ne oe. XR OR I EX BX Per 1,2,4
Polysiphonia denudata (Dillw.) Kiitz. Xx x AX X XK X XX Ann 3.4
Polysiphonia elongata (Huds.) Spreng. x X X KX X X X X X X _~ Per 2-4

Spsameas aitysdwey MIN — WYaH % uosaryepy [9861

I¢

Table III: (Cont.)

Taxa IF MAMJ J A S ON D Longevity Se tae
Polysiphonia flexicaulis (Harv.) Coll. X X X X X X XK XK XK X X X Per. 1-4
Polysiphonia harveyi Bailey xX X X X X X X X X~— Ann 1,2,4
Polysiphonia lanosa (L.) Tandy X X X X X X KX K X X XK X Per. 1-4
Polysiphonia nigra (Huds.) Batt. xX X X xX X X X X X X X _. Per. (?) 1-4
Polysiphonia nigrescens (Huds.) Grey. X X X X X X XK X X X X X Per. 1-4
Polysiphonia novae-angliae Tay]. x 4 X Per. (?) 1,2
Polysiphonia subtilissima Mont. X X X X X KX KX X X X Per. 3,4
Polysiphonia urceolata (Lightf. ex Dillw.) Grev. X KX XK X K X K XK XK X X Per 1-4
Porphyra leucosticta Thur. in Le Jol. X X X KX X X XK XK X X xX Ann 1, 2,4
Porphyra linearis Grev. xX X X X X xX xX Ann 1,2,4
Porphyra miniata (C. Ag.) J. Ag. X X X X X X X Ann 1,2,4
Porphyra umbilicalis (L.) J. Ag. Xx X X X X KX KX KX X X XK X~— Ann 1-4
“Porphyrodiscus simulans Batt.” xX Per. (?) 4
Porphyropsis coccinea (J. Ag. ex Arsesch.) Rosenv. > 4 x X* Ann 2
Pterothamnion plumula (Ellis) Nag. x x xX xX X X X X AAnn. 1,2,4
Ptilota serrata Kiitz. X X X X K K X K XK X XK X ~~ Per 1-4
Rhodomela confervoides (Huds.) Silva xX X X X X X X X X X X Per. 1,2,4
Rhodophyllis dichotoma (Lepesch.) Gobi X X X X XK K X XK X X Per 1,2,4

cs

elopoyy

88 104]

Table III: (Cont.)

Taxa

J F M:iA M.S J. A. S © ND

Longevity Distribution

Local

Rhodophysema elegans (Crouan frat. ex J. Ag.)
Dixon

Rhodophysema georgii Batt.

Sacheria fucina (Bory) Sirodot

Scagelia corallina (Rupr.) Hansen et Scagel
Spermothamnion repens (Dillw.) Rosenv.
“Trailliella intricata Batt.”

Turnerella pennyi (Harv.) Schm.

Monthly total
Rhodophyta taxa

Monthly grand total seaweed taxa

eC. De es GI eat RA ee
xX X X X xX X X

xX X Me KR ek RH

Bik Me RM MUR aN XK RM KN
X X

xX xX xX X X xX xX

xX XxX

56 65 60 66 67 75 77 83 74 73 76 66

105 131 132 143 146 159 169 178 146 146 148 120

I,

oN

2,4

Spoameas aitysdwiey MON — YH W uosayiepy [9861

c¢

Table IV: Summary of collection sites

Geographical
Area

No. and/or
Names of
Permanent
Stations

Sites with at Sites with at
Least 2 yrs. Least 2 yrs.
of Seasonal of Monthly
Collections Collections

Pertinent References

1. Isles of Shoals

2. Nearshore open
coast

3. Hampton-
Seabrook
Estuary
System

Appledore Island
Cedar Island
Duck Island
Lunging Island
Malaga Island
Smuttynose Island
Star Island

White Island

13 sites
Jaffrey Point, New Castle
43°03'22”N, 70° 42'49"W

Bound Rock, Seabrook
42° 53'30”N, 70° 48’45”W

49 total sites

Blackwater River—11 sites
Brown River— 13 sites

Xx

xX

x

x

x

xX

x

x
Xx
x
Xx

x

X

x

Mathieson (1979)
Mathieson & Penniman (1986a)
Norall et al. (1981)

Hehre & Mathieson

Mathieson, Hehre, & Reynolds (1981)
Mathieson & Penniman (1986a,b)
Mathieson, Reynolds & Hehre (1981)
Daly & Mathieson (1977)

Mathieson & Fralick (1972)
Mathieson & Penniman (1986a,b)

Mathieson and Fralick (1972)

vs

BIOpoyYy

88 190A]

Table IV (Continued)

Geographical
Area

No. and/or
Names of
Permanent
Stations

Sites with at Sites with at
Least 2 yrs.
of Seasonal
Collections

Collections Pertinent References

4. Great Bay Estuary
System

Hampton River— 18 sites
Hampton Harbor—7 sites

182 total sites

Great Bay— 16 sites
Crommet Creek, Durham
43°05’52”N, 70° 53’53”W
Nannies Island, Newington
43°04’08"N, 70°51'47”W
Thomas Point, Newington
43°04’53”"N, 70° 51'56”W
Weeks Point, Greenland
43°03'32”N, 70°51'42”W

Little Bay—21 sites
Adams Point, Durham
43°05’43’N, 70° 52’07”"W
Cedar Point, Durham
43°07'45”N, 70°51’08”"W

Dover Point, Dover
43°07'07’"N, 70° 49'42”"W

xX
xX

Mathieson, Reynolds,
& Hehre (1981)

Daly & Mathieson (1981)
Mathieson & Penniman (1986b)
Mathieson & Penniman (1986b)

Mathieson & Penniman (1986b)

Mathieson, Reynolds & Hehre (1981)

Mathieson, Reynolds & Hehre (1981)

Chock and Mathieson (1976, 1983)
Mathieson & Penniman (1986b)
Mathieson, Reynolds & Hehre (1981)
Mathieson et al. (1983)

Mathieson & Penniman (1986a,b)

[9861

Spoomeas ailysdwie py MAN — WYO W uosaryepy

Table IV (Continued)

Geographical
Area

No. and/or
Names of
Permanent
Stations

Sites with at Sites with at
Least 2 yrs. Least 2 yrs.
of Seasonal of Monthly
Collections Collections

Pertinent References

Durham Point, Durham

43°07'14”N, 70° 52’10”W
Bellamy River—10 sites
Cocheco River—17 sites
Lamprey River—9 sites

Oyster River— 14 sites
Headwater at route 108,
Durham

43°07'52”N, 70°55’'06”W

Piscataqua River—S9 sites

Atlantic Heights, Portsmouth

43°05'36’"N, 70° 46'08”W
Normandeau Schiller site
#16, just east of the
Schiller Power generating

station, Portsmouth
43°05’41”N, 70°46'51”W

x ~*~ K

8 years of
continuous
seasonal
collections

Mathieson, Reynolds & Hehre (1981)
Reynolds & Mathieson (1975)
Mathieson & Penniman (1986b)

Mathieson & Penniman (1986b)

Mathieson et al. (1977)

Normandeau Assoc. (1971-80)

9¢

elopoyy

88 10A]

Table IV (Continued)

No. and/or Sites with at Sites with at
Names of Least 2 yrs. Least 2 yrs.
Geographical Permanent of Seasonal of Monthly
Area Stations Collections Collections Pertinent References
Ibid., #17, at end of 8 years of Normandeau Assoc. (1971-80)
Long Reach Farm, Eliot, continuous
Maine seasonal
43°06'02”N, 70° 46’52”W collections
Ibid., #20, near Schiller 8 years of Normandeau Assoc. (1971- 80)
Power Plant, Newington continuous
43°06'15”N, 70° 47'47”"W seasonal
collections
Ibid., #40, near Simplex, 8 years of Mathieson & Penniman (1986b)
Pier, Newington continuous Normandeau Assoc. (1971-80)
43°06’15”N, 70°47'47”"W seasonal
collections
Ibid. #44, area just west 8 years of Normandeau Assoc. (1971-80)
of Simplex Pier and Union continuous
Oil Terminal, Newington seasonal
43°(06’28”N, 70° 47'58”"W collections
Salmon Falls River—16 sites xX
Squamscott River— 16 sites
Chapman’s Landing X Mathieson & Penniman (1986b)

Route 108 bridge, Newfields
43° 02'24’N, 70°55’43”W

Winnicut River—4 sites

Mathieson, Reynolds & Hehre (1981)

[9861

Spoameas ailysdwieH MON — a1YydH % UOsaIpIeEp

58 Rhodora [Vol. 88

LITERATURE CITED

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Biair, S. M. 1983. Taxonomic treatment of the Chaetomorpha and Rhizoclo-
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, A.C. MATHIESON AND D. P. CHENEY. 1982. Biosystematic investigations
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BousFIELD, E, L. AND M. L. H. THoMas. 1975. Postglacial changes in distribution
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Brown, W. S. AND E. ARRELLANO. 1979. The application of a segmented tidal
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Burns, R. L. AND A. C. MATHIESON. 1972a. Ecological studies of economic red
algae. II. Culture studies of Chondrus crispus Stackhouse and Gigartina stellata
(Stackhouse) Batters. J. Exp. Mar. Biol. Ecol. 8: 1-6.

AND . 1972b. Ecological studies of economic red algae. Il. Growth
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Capuzzo, J. M. AND F. E. ANDERSON. 1973. The use of modern chromium
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CaRLTON, J. T. AND J. A. SCANLON. 1985. Progression and dispersal of an intro-
duced alga: Codium fragile ssp. tomentosoides (Chlorophyta) on the Atlantic
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CELIKKOL, B. AND R. REICHARD. 1976. Hydrodynamic model of the Great Bay
Estuarine System. Univ. of New Hampshire Sea Grant Technical Rept., UNH-
SG-153.

CHAPMAN, V. J. 1964. The Algae. MacMillan and Company, Ltd., London.

CHIHARA, M. 1961. Life cycle of the Bonnemaisonaceous algae in Japan (1). Sci.
Rept. Tokyo Kyoiku Daigaku 10: 121-153.

1962. Life cycle of the Bonnemaisoniaceous algae in Japan (2). Sci. Rept.
Tokyo Kyoiku Daigaku 11: 27-54.

Cuock, J. S. AND A. C. MATHIESON. 1976. Ecological studies of the salt marsh
ecad scorpioides (Hornemann) Hauck of Ascophyllum nodosum (L.) Le Jolis. J.
Exp. Mar. Biol. Ecol. 23: 171-190.

AND 1983. Variations of New England estuarine seaweed bio-
mass. Bot. Mar. 26: 87-97.

CoLeMAN, D. C, AND A. C. MATHIESON. 1975. Investigations of New England
marine algae VII: Seasonal occurrence and reproduction of marine algae near
Cape Cod, Massachusetts. Rhodora 77: 76-104.

1986] Mathieson & Hehre — New Hampshire seaweeds 59

CoLiins, F.S. 1900. Preliminary lists of New England plants. V. Marine algae.
Rhodora 2: 41-52.

1901. Notes on algae III]. Rhodora 3: 132-137.

—___—.. 1903. Notes on algae V. Rhodora 5: 204-212.

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DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
JACKSON ESTUARINE LABORATORY

UNIVERSITY OF NEW HAMPSHIRE

DURHAM, NH 03824

66 Rhodora [Vol. 88

FIGURES 16-229. SpecIES DISTRIBUTION MAPS FOR 214 TAXA
FOUND WITHIN COASTAL—ESTUARINE WATERS OF NEW HAMSPHIRE
AND SOUTHERN MAINE.

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1986]

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1986]

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47'56- 4255
10 hm
7065" 7045 Fig, 63

SPONGOMORPHA ARCTA

78 Rhodora

[Vol. 88

SPONGOMORPHIA SPINESCENS

STICHOCOCCUS MARINUS

‘e
47654
10 bm 10 bmw
7065. 7045 7056 1045,
A , Fic. 64 , f Fic. 65
STIGEOCLONIUM SP, ULOTHRIX FLACCA
os’ 4706 4
ays 4 Ay
it tes
mh %
& fe
4
‘ 7 .
° i ~~
4758+ 47554
whe 10 bmw
s . ‘ 2 . . .
7056 70 46
10°68 748 Fig, 66 : : Fis, 67

1986] Mathieson & Hehre — New Hampshire seaweeds

fie

i?

ULOTHRIX SPECIOSA

4

SS

y

o_o
42664

ULVA LACTUCA

.
4305 4

47664

7048 Fig, 68 70°68" 7048 Fie. 69
i
Uininks Deseuns ULVARIA OXYSPERMA
4768-4
Ha
whe
bess wise ae
70°65 4 7068
i 7048 Fig. 70 f

80

Rhodora

[Vol. 88

UROSPORA PENICILLIFORMIS

ee I 1
47°05 43°05
eC
& ma
he: ae
~ “
47'55- 47564
etm Si 10 bm
7068 148 Fig, 72 7055 j 48 Fig 73

we

UROSPORA WORMSKIOLDII
YY
< ‘
7~S
Ye, |
“YY
Xd?

1066

AGARUM CRI BROSUM

NY
a?
Ry, 2
5 eet hh 4706 4
Dep §
e
£
eee.
n
4756+
1 he
7048 Fie, 74

ALARIA ESCULENTA

47554

70°68 1048 Fig, 75

Mathieson & Hehre

New Hampshire seaweeds

S|

1986]
Sle SY =
ae, AGORVEENS Bieta Lees acon Y ASCOPHYLLUM NODOSUM
ys] 4 y | i YJ
‘ = = - . -
oh N ™\y ( /
musth Mey. {
C ot ESSN ¢ ue J
. LAr j
\ ae Mise A dy? \
/ Oe, {. da P .
: ; Pas { ER —., f 43°05 430
Ss a aa a
= aaa’ jh Peay
é
a *
* Ps
ad es "S
J 7 ’e
Bi
Bran 4z'5 Vas ¢ 42°55
Wr oe
,- 10 bn Lay ‘p! 10 bm
jase i 748° Fic, 76 70°55 ny 748) Fig. 77
/ ASCOPHYLLUM NODOSUM Ne y
ECAD SCORPIOIDES ~~ } asPerococcus FISTULOSUS
| |
4 oy . : } : S
N Cupra }
a uA “tr : S Le Y [
7 eh ‘ z Las 4 } “4 )
c ‘as } Pm ee wragt pS
\ i =. rs x ) ff SSG ) s
P ™~ ye 45 s , | ™ > iis y, i
q ; Vea ry fy 4305 y a ee | ca 2 F
7 sf ¥ PSG ae a ap eae 43°05
y aot ly ye Re es ie
bs came - ae , 5 ‘" Bop ioe
J yy “KR it Cae Bo
a / fo
f :
ef dq
: ¢ & j
3
: ( <a
E J
4765- Be ( 4755
Mae TE
iP ar i= ag} >, paihan
70°55 70-48 Fic. 78 70°58 1045 Fre. 79

82

Rhodora

[Vol. 88

CHORDA FILUM

———————

| » Oke
7065 j 748 Fig, 80

47664

1065"
it

CHORDA TOMENTOSA

43°06 -

a)
4265

a
148° Fig, 81

4

oo
70 65
L

3

| CHORDARIA FLAGELLIFORMIS

7048 Fig. 82

4706

47664

CLADOSTEPHUS SPONGIOSUS
FORMA VERTICILLATUS

1986] Mathieson & Hehre — New Hampshire seaweeds 83
By
DELAMAREA ATTENUATA DESMARESTIA ACULEATA
/ s
‘ (
ae fi
oe
< xe |
. by v
=
vf ait Ps a Sted
¥. kay f 430
a,
yg
% 4
is |
( ai
*”
«
¢ 4265
eS; 10 bmw
7068 10-48 Fic, 84 7056) > 70°45 Fic, 85
7 a DESMARESTIA VIRIDIS DESMOTRICHUM UNDULATUM
‘y
. (
Ys. f
. Cy, )
,
acai toe he
‘S. Ay 2
os 4 ; te AY . Jo
43065 spe e J 4305
edt “
aA
:
a 2 ad
&
Bt. iF.
“~ ~
47564 4755
on 10 bn _
10°55) 7048 Fig, 86 7068) 7045 Fig, 87

[Vol. 88

Rhodora

DICTYOSIPHON FOENICULACEUS

J

M. ol
ee Xe hy

u e-

&

ik
88

84

45 Fig, 91

hs
>
os
Ie
<i
[
f

ei
ha:
e

ee
70 45 "
é Fis,

ECTOCARPUS SILICULOSU

Mathieson & Hehre | New Hampshire seaweeds 85

ae , tuesne vies ke oe ne
ie! z ~~ :
ver E Tk me XE es

—— s :
> 2 ie mh, x hy. -
yt a as os _ ft
rT Regatig tf vo oY

1986]

us. 10 bm Oe 10 bam
70'55' 7045 : | 55 1 y45 |
; i } Fic. 92 70°85 { 7045 Fig, 93 7
ah :
ry FUCUS DISTICHUS = FUCUS DISTICHUS
a ; atkery Beer SUBSP, EDENTATUS
) ee
i 2
= Cc
ee:
:

( f
“Hl ; 7

70:48 Fis, 95

[Vol. 88

Rhodora

86

1986] Mathieson & Hehre —- New Hampshire seaweeds

GIFFORDIA SANDRIANA

oh

ba gic

ve

87

NDA

i
ae |
2

ot:
¥ Z
:
year aos
Ce BES
Oe
en fe

oa oth
eat
id 4
3) &,
¥. ae
4765 7 ( 475
10 Sf rr
70-48 Fic. 100 70°55 y 7045 Fis. 101
ISTHMOPLEA SPHAEROPHORA = LAMINARIA DIGITATA
: ~yJ
. é
Le * 7
{
as:

7045 Fig, 102

af

Mia,

47°55

88

Rhodora

[Vol. 88

{

Sec
re watt

i

Ro
7 By

sae casa Ye

aay LAMINARIA LONGICRURIS
Mas .
{

o.-
4305 j

7045 Fic. 105

Na ie :

LEATHESIA DIFFORMIS
J 2
3 ob.
7
47684

New Hampshire seaweeds

89

1986] Mathieson & Hehre
MIKROSYPHAR PORPHYRAE EMA CORUNNAE
4 5
s ys \
sy (
Cy» a
x BY 3
oe yyy
OR Us Be, 43°05
ae “ Berl
/ /
A x
&
Pie c of
: y, ~
47'55- Re ( 4755
aig
10 bm st ee
70.68" 7048 Fig, 108 70°55) y 7045 Fig, 109
MYRIONEMA STRANGULANS avesinsie Baise
SJ
‘
{
Yes
> et “Ps
ras | ?
x Re ye
eA ag08" Sr Aue | 43°05
3 "ar
_
»
se
P Fs 4
F ( dt
: »
47564 4255
7066 was Fig, 110 7048 Fic. 11

90 Rhodora [Vol. 88

PETALONIA ZOSTERIFOLIA kw
PETRODERMA MACU LIFORME
“al —_ ef ;
>»
cs Xy ; a
Part

of

a OTECTOCARPUS SPECIO

*
site

9]

1986] Mathieson & Hehre — New Hampshire seaweeds

PSEUDOLITHODERMA EXTENSUM PUNCTARIA LATIFOLIA

4706 4 470s
x ‘ oe
g£ oo
8 8
ox. +e.
. >
4766 4 455-
be rn
70°66 j was Fis, 116 70°88 48 Fig, 17
PUNCTARIA PLANTAGINEA “RALFSIA BORNETII"
diet >
oe oa
ye
a oe
avoe- Bo 4306 4
P I
oo ro
* te
atss4 4664
he eke
ree 48 Fie, 118 do 748 Fig, 119

92 Rhodora

[Vol. 88

"RALFSIA CLAVATA” RALFSIA FUNGIFORMIS

J

2
~

ae
o \ > >» 0
3 4705 & ‘ 43°08
a gt rar!
x |

& 5

a 8.

» os

4765+ aves

10 hem i. =~ tia
had 70-48 Fic, 120 7065" v 7045 Fig, 12)
RALFSIA VERRUCOSA

ia ~y

C
oan ( 7

r i

; 2 47068 3 cH 43°05

%, 4.

fe ¥
47684 42554

ele
7068" 7048 Fie. 122 7068 7048 Fig, 123

1986]

Mathieson & Hehre — New Hampshire seaweeds

93

SCYTOSIPHON LOMENTARIA
VAR. COMPLANATUS

SCYTOSIPHON LOMENTARIA
VAR. LOMENTARIA

eo.
7065
fi

47654

7045
i

Fic. 126

en tis Wh a a ois Ke gh lle 4
. a e hae : \ebte x x 7
x , wa “ib £ . Y Ms y a { 4 ,
Ps ty SS. oO 4305 o {xf re ae 4305
A ‘ ®
». f |
ran a
J f =
(
5 4765 *) {;? 4755
y/| uae a
10 ben ¢ 10 be
70°55 { 7045 Fic, 124 70°58" N 74s Fic, 125 |
SORAPION KJELLMANI1 Sy } SOROCARPUS MICROMORUS
7 { ; 4 j

Age oe
5 @

7045 Fig, 127

94 Rhodora [Vol. 88

SPHACELARIA ARCTICA SPHACELARIA CIRROSA

47564 aves

_ ete 0 he
7065 v was Fic. 128 70°58) 7048 Fig 29
ty SPHACELARIA FUSCA

ou
7055
4

“ip.
“By =

SPHACELARIA PLUMIGERA

o..
42664

48 Fic. 130 7088 748 Fic. 131

Mathieson & Hehre — New Hampshire seaweeds

95

1986]
SPHACELARIA PLUMOSA
SPHACELARIA RADICANS

J

x ! ¥
i ee

. NK eng, |

ENS an.
x Q x
SS ea 43°05 , ved oe
ai sf

my ri

8 8

OF. iF,
42755 RE 47'56-

10 bm SesP 10 baw
7068. 70-48 Fic, 132 7065 1045 Fig, 133
SPHAEROTRICHIA DIVARICATA
SPONGONEMA TOMENTOSUM

~y

oe.
4306 4

) r
3)
iF. et
* ~
47584 ° av654
10 be 10 be
7086 7048 Fic. 134 7066 70-48 Fic. 135

Rhodora

[Vol. 88

STICTYOSIPHON GRIFFITHSIANUS

7065) 7045 Fig, 136

; 4755-
I
748 Fig. 137

i
a! ANNFELTIA PLICAT

St

748 Fic, 138

o
70 65
n

Ah’

ANTITHAMNION CRUCIATUM
se

“Ha

75 45 Fic. 139

1986] Mathieson & Hehre

New Hampshire seaweeds 97

ANTITHAMNIONELLA FLOCCOSA

.
4265

{

ah wat
y 748 Fic. 140

‘ .
i AUDOUINELLA ALARIAE
d. ~yJ
}

A

PR ‘ 5
XQ x & y
hes ‘ — f 4305
Sati

ag
v 70-48 Fic. 141

Ce
70 55
4

AUDOUTNELLA MEMBRANACEA

AUDOUINELLA POLYIDES
=

My ra
he Wy

47584
z 10 bm
70°68. { 7048 Fie. 142

70°66. {

7048 Fig, 143

Rhodora

[Vol. 88

AUDOUINELLA PURPUREA

e-
70 46
n

Fis, 144

4755 4

AUDOUINELLA

SECUNDATA

70-48 Fic. 145

STF,

GQ, AUDOUINELLA VIOLACEA

ba

4 p>
. eet 43°05 ie “ 4705
es panies
Hf : eS
4, 4,
3)
3 Re
> “~
A. 47554 47684
\ 10 be 10 hn
7088 7048 Fig, 146 70°58 748 Fig, 147

BANGIA ATROPURPUREA

1986]

Mathieson & Hehre — New Hampshire seaweeds

99

4765

a

10tm
70°66 i 7045 Fig, 148

aay

$s

“HJ
| 3

CALLITHAMNION BYSSOIDE
y E las |
~ (( ae

Ns x Wa

Sy; om
j 745 Fic. 149

7065
F

CALLITHAMNION HOOKERI

ua

47584
ete
7068 7048 Fig, 150

CALLITHAMNION TETRAGONUM

: a
é ft
~
if 4766
f eine oo
7056 70.48 Fic, 15)

[Vol. 88

Rhodora

100

<
=
<
=
no“
=
Vv
”
4
pe
=
a
3
=
=
<
J

7045 Fig, 153

v on

7048 Fig, 155

L ‘ “— ee | PT : "SB.
aay ele | ;
: 7 he 2 ll ge ~, 2
x inn ‘ee Zs “?
: pen bs | ; : Ee | 1
Ss la) : —d °
E 2 ~e < a X eS
(te eee ra / i, es! . yt ~~,
ee | ao
Le (ON fr ee AT aro is ae faa
yA SORE
Sf th yA f
_ - ol Res or
"R ( 7 he ( ee

1986] Mathieson & Hehre

New Hampshire seaweeds

101

CERAMIUM RUBRUM

o.
4266

ng! 10 bm
7056 { 7048 Fie. 156

eo .
70 65
4

CERAMIUM STRICTUM

Tae

7048 Fic, 157

YP 4

CERATOCOLAX HARTZI1

70°58" f 7048 Fig, 158

4308

47564

70668"
Ae

CHONDRIA BAILEYANA

10 be

ras’ Fie. 159

o5c8
4306

o-;
42664

102 Rhodora [Vol. 88

by! re hl

ay me f Ny, & ct vie ot > I ‘SS nik »;
pat aie" a
i a

1986] Mathieson & Hehre

New Hampshire seaweeds 103

_ ENSIS”
ae :

CYSTOCLONIUM PURPUREUM

fo

4 Ne { ( a ; {
m ua Me) |] rll ,
Scan: \ Cee ie x
{| Sp 7 5 " : | Ney i ay
2 ys whe Ue hs 2 ae Dey f, We, "os
we Regt wel ALY Cage
3 s 5 ott ° if vat at a
{ j ? *
; \ J
( ot f rte.
2 2 °
J
ne! 4765. 5 : he 4766
Se) er 5 fs seins
7058 y 745 Fic, 164 70°68 ‘f 7048 Fic. 165
14.7 DASYA BAILLOUVIANA Sy ae DERMATOLITHON PUSTULATUM
Le / LK, /
° r - - | ii -
V mo \ “ Mw —
\ 4. a Se, | | e en {
mk 13} . . " Be von pias! % : Bae
aoe wt ‘ by, oN oe we 7 ) -
she AGO ft wy he 4
4 yw uy Rete 47°05 7 of : i nee Sr} «S06
¥ Fal oth = . y, ty oe te -
P Va ," d.
/ ‘ .
i ; 6 - |
f 1. ? ft
f -
ag! ste ne vt
o> yale
Ae 10 bm as ohm
7058 ‘| 7045. Fic. 166 7055 2 7045 Fig, 167

104

DEVALERAEA RAMENTACEUM

Rhodora

[ Vol. 88

DUMONTIA CONTORTA

47584 aves
10 ta 10 he
7065 { 745 Fig. 168 71068. i 7048 = Fig, 169
ERYTHROPELTIS DISCIGERA ) ERYTHROTRICHIA CARNEA
VAR, DISCIGERA
| WAR &
4, 2
ye
er 4F08 -
Dap fs
~
¥
A. ste -
| Wie
70°58" 048 Fig, 170 doe was Fig, 171

Mathieson & Hehre — New Hampshire seaweeds

105

1986]
FOSLIELLA LEJOLISII GIGARTINA STELLATA
“J
ave ()
az os 4305
“9 j s oe
: . ais :
re
o %
a f ft
» f »
47664 af 4755
7058 { 7048 Fie, 172 7058 { 1045 Fic. 173
GLOIOSIPHONIA CAPILLARIS a ALSIDII
5
2
eg y
" hy?
re) = yy
sea ays 4 of eae, 4305
* ra P
- rts.
47564 4755-
768 748 Fig, 174 1088" 748 Fig, 175

Rhodora

[Vol. 88

: is
ay _ se

GRACILARIA TIKVAHIAE

j 745 Fig, 176

GYMNOGONGRUS CRENULATUS

748 Fig, 177

7055
4

HALOSACCIOCOLAX KJELLMANII

7048 Fis, 178

7058
A

HARVEYELLA MIRABILIS

y48' Fig, 179

1986] Mathieson & Hehre — New Hampshire seaweeds

107

“s E Ser (
£ .

‘st agi!

Can
4255

7065 70.45 Fic. 180
4

[ 7045 Fic, 18]

‘yy

I: YW, ~
Me ae, “Gor,

r *

f ys

3 (
Ng

—
a

ae
4266-

LEPTOPHYTUM LAEVE
Ps

7055) 1045 Fie. 182

‘7%
70 55
rn

LITHOPHYLLUM CORALLINAE

7048 Fic, 183

Rhodora [Vol. 88

z My LOMENTARIA BAILEYANA
LITHOTHAMNION GLACIALE > '
; by a
< Y

=,
a x

42765
ein “sf
7048 Fig. 184 hae ry

Fa
ae os

1986]

Mathieson & Hehre

- New Hampshire seaweeds 109

ey MEMBRANOPTERA ALATA

7048 Fig, 190

110 Rhodora [Vol. 88

Ww Ls
PETROCELIS CRUENTA PEYSSONNELIA ROSENVINGII

Ea
ety
( Sgie

i

.
426574

7048 F 16,193

¥ 10 bm
7068 { 70-48 Fic. 192 70°65
‘\

PHYCODRYS RUBENS

PHYLLOPHORA PSEUDOCERANOIDES

47684

7068 748 Fis. 195

1986] Mathieson & Hehre

New Hampshire seaweeds

111

PHYLLOPHORA TRAILLII

¥— 7045 Fig, 196

Pe
fp

ot

745 Fic. 197

| PHYMATOLITHON LAEVIGATUM
a . > a
a=

ia 4768 4
Jo 704 5 Fig, 198

e
4306 1

°

PHYMATOLITHON LENORMANDI I

112

Rhodora

[Vol. 88

oe -
70 55
"

PHYMATOLITHON RUGULOSUM

748 Fic. 200

70 55

PLUMARIA ELEGANS

7048 Fig, 201

a

7058.
A

POLYIDES ROTUNDUS

7048 Fie. 202

A 68

POLYSIPHONIA DENUDATA

was Fig, 203

Mathieson & Hehre

New Hampshire seaweeds 113

POLYSIPHONIA ELONGATA

4755

i 7045 Fic. 204

; POLYSIPHONIA FLEXICAULIS

4 ~ 5
¢

i

4755

a

10 bay

Fig. 205

POLYSIPHONIA HARVEY! ay’ ; ee LANOSA
J 2 ‘yy
a } . a
Le Ny (

ae f if Cy By,

= Fae wre u : es al }

x aie me x ? y re

WA
4yos " rn Sy ~ astos
7 i os a & ‘See i
a ef, d-
aa,
™ se %
oe
ot ¢ re.
: ~
a
47584 -e 4755
{
re = ein
had 7045 Fig, 206 7068" } 148° Fig, 207

114

Rhodora

[Vol. 88

70 46 Fic, 208

745 Fig, 209

7048 Fig, 210

POLYSIPHONIA NOVAE-ANGLIAE

1986}

Mathieson & Hehre

New Hampshire seaweeds i

} POLYSIPHONIA URCEOLATA

Sis

=

4308

“dy Pp

47'66-

PORPHYRA LEUCOSTICTA

4705

476554

Oe he

1048 Fie. 212 7058 70°45 Fic. 213
PORPHYRA LINEARIS PORPHYRA MINIATA
whe
7056 7948 Fic, 214 7688 48" Fie. 215

Mathieson & Hehre

New Hampshire seaweeds

[17

1986]
= = tc
ah i rer scm ).rioponeLa courervorpes
| ; yy | ( j
a's : .
=)
ln coll |
mak io
en at
\ | “4;
ye ’
yD
‘~
ee «Oh ue
a
i
70°58 ; 7045 = Fic, 220 Fic. 221
RHODOPHYLLIS DICHOTOMA Maspiaeurmmiek mUnees
SsJ
é i
4 Pe (
“,. te . f
\
if ty ~Ly. By )
Ney
}
E 7
wy A 4 'y, /
pegacy 5 aa 09 an 4 ‘a 43°05
Bat h &
34 di :
‘@ ad 4
a ep pe
7
42'55 ee, 4255
= seh
10 bn ys ‘[ 104
70-48 Fie. 222 7055 P 76-45 Fic, 223

118 Rhodora [Vol. 88

RHODOPHYSEMA GEORGI!
SACHERIA FUCINA

47564

eke eke
7065 yoas Fic. 224 70 65° 746) Fic. 225
i 7 i. -
>i 7 ) SPERMOTHAMNION REPENS
SCAGELIA CORALLINA

aw «766

4 he
ras Fie. 227

he

7048 Fie. 226 7068

- 7
7756
4

1986]

Mathieson & Hehre — New Hampshire seaweeds

119

Ni

es 10 bm
7055. yl 7045 = Fic, 228

q TRAILLIELLA INTRICATA”

4755

} TURNERELLA PENNY!

3

~y

eee
4255

120 Rhodora [Vol. 88

APPENDIX

NEARSHORE OPEN COAST BETWEEN SOUTHERN MAINE
AND NEW HAMPSHIRE

Station Number Latitude and Longitude — Description

Cl 43° 12’16”N, 70° 34°W Bald Head Cliff, Ogunquit, Maine

C2 43°09'56’N, 70°35’25”W Nubble Light, Cape Neddick, Maine

C3 43°05S’26"N, 70°30'32”W_ Sea Point, Maine

C4 43°(03’38"N, 70°41'42”W_ Kittery Point, Maine

C5 43°()3’22”N, 70°42'49"W_ Jaffrey Point, Fort Stark, New
Hampshire

C6 43°02’15”N, 70°43’20”W = Odiorne’s Point, Frost Point, Fort
Dearborn, New Hampshire

C7 43°01'30’N, 70°43’20”’W Seal Rocks, New Hampshire

C8 43°01'20’N, 70°42’1S”W Gunboat Shoals, New Hampshire

C9 43°01'25”N, 70°43'40”W North Wallis Sands, New Hampshire

C10 43°01'00’N, 70°43’55”W Concord Point, New Hampshire

Cll 43°00'05’”N, 70°44°30”W Ragged Neck, New Hampshire

C12 42° 58’20’"N, 70°45°33"W Rye Ledge, New Hampshire

C13 42° 57'°30’N, 70°46'30”W Little Boar’s Head, New Hampshire

Cl4 42°57'00’N, 70°46'44”"W_ Godfrey’s Ledge, North Hampton,
New Hampshire

C15 42°55’05’”N, 70°47'18”W_ Great Boar’s Head, New Hampshire

Cl6 42° 54’30’"N, 70°48'30”W Hampton Beach, New Hampshire

Cl7 42° 53’30’N, 70°48’45”W Bound Rock, area in the immediate

vicinity of Beckman’s Point, near
mouth of Hampton Harbor,
Hampton, New Hampshire

1986] Mathieson & Hehre — New Hampshire seaweeds 121
GREAT BAY ESTUARY SYSTEM
PISCATAQUA RIVER
(NEW HAMPSHIRE/ MAINE)

Miles
Station From

No. Latitude and Longitude Coast Description

Pl 43°04’00’N, 70°41'46"W 0S Gerrish Island, at Fort Foster,
northeast of Wood Island, Kittery,
Maine

P2 43°03’31’N, 70°42’5S8”W 2 Wentworth Point, Little Harbor,
New Castle, New Hampshire

P3 43°03'20’N, 70°43’18’"W I Little Harbor Estuary, point
northeast of Frost Point, New
Castle, New Hampshire

P4 43°03’01”N, 70°43’55”W_ 85 Witch Creek, Rye, New Hampshire

P5 43°03’25’N, 70°44’20”"W_ 1.3 Sagamore Creek, Portsmouth, New
Hampshire

P6 43°03'34’N, 70°44'15”W 1.25. Goose Island, near mouth of
Sagamore Creek, Portsmouth,
New Hampshire

P7 43°04'20’N, 70°42'30"W 8 Fort Constitution, Fort Point,
New Castle, New Hampshire

P8 43°04'24’N, 70°42’56”"W_ 1.4 Salamander Point, New Castle, New
Hampshire

P9 43°04’19”"”N, 70°43’47”"W 2.1 Shaw’s Hill, New Castle, New
Hampshire

P10 43°04'14’"N, 70°43’48"W 2.1 Riverside Cemetery, New Castle,
New Hampshire

Pll 43°04'12’N, 70°44’26"W 2.7 Shapleigh Island, Portsmouth,
New Hampshire

P12 43°04’29"N, 70°44’48"W 2.8 Pierce Island, Portsmouth, New
Hampshire

P13 43°04’44’"N, 70°45'12”"W 3.4 Memorial Bridge and adjacent
Fisherman’s Pier area, also Electric
Plant, Portsmouth, New Hampshire

P14 43°04’46’"N, 70°45’28”"W 3.6 Ceres Street, upstream from P13,
Portsmouth, New Hampshire

PIS 43°05’09"N, 70°45’40°W 4 Bridge at Rte. | bypass, west bank,

Portsmouth, New Hampshire

122

Rhodora [Vol. 88

Great Bay Estuary System/ Piscataqua River (NH,ME) (Cont.)

Station
No.

Latitude and Longitude

Miles
From
Coast

Description

P16

P17

PI8

P19

P20

P21

P22

P24

P25

43°05’13”N, 70° 45’S2”W

43°05'26"N, 70° 45°39" W

43°05'36"N, 70° 4608” W

43°05’43”N, 70° 46'10"W

43°05’52”N, 70°46'03”W

43°05’40"N, 70° 46'46"W

43°05’41"N, 70°46’51”W

43°05’51”N, 70°47'02”W

43°06'02”N, 70° 46'52”W

43°06'15”N, 70°47'47"°W

4.3

4.5

4.7

4.9

5.0

5.0

5.3

a2

5.6

5.6

Freeman’s Point, Normandeau
Schiller Site No. 12 (New
Hampshire side at Maine~New
Hampshire Rte. | bypass bridge)
Portsmouth, New Hampshire

Normandeau Schiller Site No. 13 (on
Maine side in creek west of Maine-
New Hampshire Rte. | bypass
bridge) end of Adams Lane,
Kittery, Maine

Atlantic Heights and Normandeau
Schiller Site No. 14 (west of “new”
bridge—New Hampshire side)
Portsmouth, New Hampshire

Normandeau Schiller Site No. 15
(east of navigation point and
high tension towers) Eliot, Maine

Spinney Creek, at south Eliot Road
Bridge, Eliot, Maine (including
Jerry’s Marina)

Dock at Sprague Terminal,
Portsmouth, New Hampshire

Normandeau Schiller Site No. 16 (in
cove east of Schiller Generating
Station) Portsmouth, New
Hampshire

Schiller Station, Portsmouth, New
Hampshire

Normandeau Schiller Site No. 17 and
17D (Maine side at end of Long
Reach Farm) Eliot, Maine

Newington Power Station and
Normandeau Schiller Sites Nos.
18-40 (between Schiller Plant and
Simplex Pier; benthic stations 300’
offshore LW marsh, and 500’ from
HW mark) and Normandeau Schiller

1986] Mathieson & Hehre — New Hampshire seaweeds 123

Transects A-C, Simplex Plant-Pier,
Newington, New Hampshire

Normandeau Schiller Site No. 42 (on
the west side of the Simplex Pier)
and Normandeau Schiller Transects
D & E, Newington, New Hampshire

P26 43°06’21”N, 70°47'49"W 6.5

P27

P28

P29

P30

P32

P36

43°06'32’'N,

43°06'38’"N,

43°06'28’N,

43°06'33’"N,

43°06'44’N,

43°06'53’N,

43°06'52’N,
43°07'07'N,

43°06'58’N,

43°07'00"N,

43°07'160’N,

70° 47'34”W

70° 47'47°W

70° 47'58”W

70° 48'13"W

70° 48'32”W

70° 47'°S7”"W

70° 48°08" W
70° 4806" W

70° 48°42” W

70° 49'24"W

70° 4822” W

6.45

6.6

6.7

6.85

7.05
7.4

7.6

8.2

7.65

Normandeau Schiller Site No. 19
(one-half mile east of Frankfort
Island) Park Street, Eliot, Maine

Public landing end of Green Acre
Road and just upstream and
opposite from Simplex Dock,
Eliot, Maine

Normandeau Schiller Site No. 44 (in
a large cove west of the Simplex
Pier) and Union Oil Terminal,
Newington, New Hampshire

Town Landing, Newington, New
Hampshire

Normandeau Schiller Site No. 46
(east of old shipyard and west of
Union Oil Terminal), Newington,
New Hampshire

Mast Cove (Searles Cove) and
Normandeau Schiller Site No. 21
(in Mast Cove behind Frankfort
Island) Eliot, Maine

Frankfort Island, Eliot, Maine

Mast Cove (Searles Cove) and
Normandeau Schiller Site No. 23
(east of Adlington Creek), Eliot,
Maine

Normandeau Schiller Site No. 48
(west of Atlantic terminal),
Newington, New Hampshire

Bloody Point, opposite Hilton Park
and Normandeau Schiller Site
No. 50 (cove on northeast side
of General Sullivan Bridge),
Newington, New Hampshire

North of Adlington Creek mouth,
Eliot, Maine

124

Rhodora

[Vol. 88

Great Bay Estuary System/ Piscataqua River (NH,ME) (Cont.)

Miles
Station From

No. Latitude and Longitude Coast Description

P38 43°07'18"N, 70°48'45”"W_ 7.8 East bank at first promontory
upstream from Adlington Creek
at a public dock, Eliot, Maine

P39 43°07'25”"N, 70°49'02”W 7.95 East bank opposite General Sullivan
Bridge and Normandeau Schiller
Site No. 25 (on Maine side directly
across from the eastern point of
the General Sullivan Bridge) Eliot,
Maine

P40 43°07°17"N, 70°49’25”"W_ 8.2 Offshore ledge upstream from Dover
Point, Dover, New Hampshire

P41 43°07'33’"N, 70°50’'0S”W_ 8.9 Pomeroy Cove, Dover, New Hampshire

P42 43°07'38"N, 70°49'16”"W_ 8.55 East bank along River Road and
approximately opposite Pomeroy
Cove (Hilton Park is opposite this),
Eliot, Maine

P43 43°07'S1”N, 70°49’'21"W 8.7 Stacey Creek mouth, Eliot, Maine

P44 43°07'55”N, 70°49'25”W 8.95 East bank, first major promontory
upstream of Stacey Creek, Eliot,
Maine

P45 43°07'45”N, 70°49’59"W 8.95 West bank at the end of Cote Drive,
Dover, New Hampshire

P46 43°08'05”"N, 70°49°38"W 9.35 East bank opposite Pineview Drive
ending at Rogers Pt. Road, Eliot,
Maine

P47 43°08'09"N, 70°49'58"W 9.45 — West bank, Pineview Drive ending,

Dover, New Hampshire

P48 43°08'29"N, 70°49'48”"W 9.8 East bank just southeast and
opposite from the end of Roberts
Road, Eliot, Maine

P49 43°08'38’N, 70°50’°04"W_ 10 West bank at the end of Roberts
Road, Dover, New Hampshire

P50 43°08’48”N, 70°49'55”"W_ 10.1 East bank, opposite and southeast
from Riverside Drive, Eliot, Maine

PSI 43°09’01"N, 70°50°07”W_ 10.4 West bank just northeast of

Riverside Drive, Dover, New
Hampshire

1986] Mathieson & Hehre — New Hampshire seaweeds 125

P52 43°08’59"N, 70° 49’53"W

P53 43°09'11"N, 70° 49’48"W

P54 43°09'25"N, 70° 49'40"W

P55 43°09°44’"N, 70° 49'S4"W

P56 43° 10’03"N, 70° 49°38" W

PS} 43° 10'16’N, 70° 49’43"°W

P58 43° 10°19"N, 70° 49°30" W

P59 43° 10°25"N, 70° 49°30°W

10.45

10.7

10.95

11.3

11.7

12.1

East bank just south of Sturgeon
Creek, opposite Riverside Drive,
Eliot, Maine

East bank just south of mouth of
Sturgeon Creek, Eliot, Maine

East bank just north of mouth of
Sturgeon Creek, at Tidy Road,
Ehot, Maine

Just southeast of the end of Dover
Neck Road, west bank upstream
from Sturgeon Creek, Dover, New
Hampshire

East bank at large tennis court facility,
approximately mid way between
Sturgeon Creek and the mouth of
the Salmon Falls River, at the end
of Houde Road, Eliot, Maine

Northeast of Dover Neck Road, near
power lines, Dover, New Hampshire

East bank opposite Dover Neck
Road, near power lines opposite
Gould Corner, Eliot, Maine

West bank just south of Cocheco
River junction, Dover, New
Hampshire

LITTLE BAY
(NEW HAMPSHIRE)

Station
No. Latitude and Longitude

Miles
from
Coast

Description

LBI 43°07'07"N, 70° 49'42"W

LB2 40°07'17"N, 70° 5004" W

LB3 43°06'57"N, 70° 49'46"W

8.6

8.95

8.7

Dover Point, including Hilton Park

and pilings at Sullivan’s Bridge,
Dover

Benn’s Marina, west bank of Dover

Point and upstream from Hilton
Park, Dover

Point between Great Bay Marina and

General Sullivan Bridge, and just

126

Little Bay (NH) (Cont.)

Rhodora

[Vol. 88

Miles
Station From
No. Latitude and Longitude Coast Description
northwest of Newington Station,
Newington
LB4 —-43°(06'56”"N, 70°50'02”W_ 8.9 Great Bay Marina, point due west
of LB3, Newington
LBS 43°06'54’N, 70°50'34"W 9.45 Broad Cove, Newington
LB6 43°07'21"N, 70° 50’'45’°W 9.5 Submarine ledge, southeast of Goat
Island, a peninsula in Little Bay,
Newington
LB? 43°07'45"N, 70°S1'08"W 10.2. Cedar Point including shoreline of
Little Bay and Royals Cove,
Durham
LB& 43°07'27"N, 70°S119"W 10 Goat Island and adjacent rock
outcrops, Newington
LB9 43°07'13"”N, 70°51'47"W OT Fox Point, Newington
LBIO — 43°07°14’N, 70°52’'10"W) so Durham Point except northwest tip
along the bank of the Oyster River
(01), Durham
LBII = 43°06'57”N, 70°S1’S7”W_ 11.4 Langley’s Island, formerly Sassafrass
Island; Seal Rocks and adjacent
offshore ledge, Durham
LBI2 — 43°06'54"N, 70°52'03"W_ 11.45 End of Colony Cove, just south of
Durham Point, Durham
LBI3 = 43°06’23”N, 70°52'14”W_ 11.55 East bank of Little Bay at junction
of power cable, Durham
LBI4 = 43°05’5S6”N, 70°52'02"W_ 11.9. Stone House, east bank and south of
LBI3, approximately 2/3 of the
distance between Adams Point to
Langley’s Island, Durham
LBIS = 43°0S’S1”N, 70°52’11"W_s1:2 In front of P. Sawyer’s old house,
Durham
LBI6 = 43°05’43”"N, 70°52'07"W_ 12.25. Adams Point, Durham
LBI7 = 43°0S’47”"N, 70°S1'l6”"W_ 12 First promontory north of Welch
Cove, Newington
LBI8& 43°05S’41"N, 70°S1'15”’W 12.15) Welch Cove, Newington

1986] Mathieson & Hehre — New Hampshire seaweeds |
LBI9 = 43°05’35”"N, 70°5130”W 12.25. Second promontory south of Welch
Cove, Newington
LB20 = 43°05’32”N, 70°51’44”W) 12.2, Furber Strait, Durham/ Newington
LB2I 43°05'24’N, 70°51°39"W 12.35. Promontory due east of Adams
Point, Newington
GREAT BAY
(NEW HAMPSHIRE)
Miles
Station From
No. Latitude and Longitude Coast Description
GBI 43°05’S2”N, 70°53’52”W 13.9. Crommet Creek at Bay Road Bridge,
Durham
GB2 —43°05’13”N, 70°52’21"W 12.9. Footman Islands, Durham
GB3 —43°05’05”N, 70°52’48”W_ 13.35 First major promontory southwest
of Footman Island, Durham
GB4 43°04'49’"N, 70° 53’25"W 15 Third major promontory southwest
of Footman Island, Newmarket
GBS5 43°04’03"N, 70°54’25"W_ 15.3 Moody’s Point, end of Smith Garrison
Road (except for L2 just upstream
from Moody’s Point), Newmarket
GB6 = 43°03’50"N, 70°54’45”W 15.65 Shackford Point, Newmarket (except
for LI, second promontory upstream
on Shackford Point)
GB7 ——43°03’39”N, 70°54’50”W_ 15.8 ~~ West bank, due south of Shackford
Point, near mouth of Squamscott
River, Newmarket
GB& 43°03'46"N, 70°54’34”W 14.7 Sandy Point, Greenland
GB9 43°03'35”N, 70°5S2’17"W_si14..5 Brackett’s Point, Greenland
GBIO — 43°03'32”N, 70°51’42”W_ 14.65 Weeks Point, Greenland
GBI} — 43°03'05"N, 70°S1'16”W_ 15.2. Point due west of Pierce Point, just
beyond mouth of Winnicut River,
Greenland
GBI2 = 43°03’14”N, 70°50’48”W_ 15.5 Pierce Point, Greenland
GBI3 — 43°04’05”N, 70°50’48”W_ 15.05 Fabyan’s Point, Newington
GBI4 —-43°04’08"N, 70°S1’47”W 13.85 Nannie Island, Newington
GBIS = 43°04’16"”N, 70°51’40"W_ 13.75. Woodman Point, Newington
GBI6 —-43°04’53’N, 70° S1'S56”’W 13 Thomas Point, Newington

128 Rhodora [Vol. 88
BELLAMY RIVER
(NEW HAMPSHIRE)
Miles
Station From
No. Latitude and Longitude Coast Description
Bl 43°07'47’"'N, 70°50’52”W_ 10.4 ~=Mouth, east bank near Scammel
Bridge pilings, Dover
B2 43°08’06’N, 70°50’36”"W 10.75 East bank, opposite Clements’ Point
and near toll plaza, Dover
B3 43°08’09’"N, 70°51’02”"W 10.75 Clements’ Point, Dover
B4 43°08’49"N, 70°50’54”W 11.6 ~~ West bank, opposite from Bellamy
Lane, Dover
BS 43°09’21’N, 70°SV1I7”’W 12.3 West and east banks at Nute Road,
Dover
B6 43°09’47’N, 70°51’23”W_ 12.8 — East bank at Cushing Road, Dover
B7 43°09°57"N, 70°S1’37”"W 13.1 East bank, end of Spur Road near
Greek cemetery, Dover
B8 43° 10°16"N, 70°51’52”W 13.5. West bank, near the end of Mast
Road, Dover
B9 43° 10’33”N, 70°52’'20"W_ 14 West bank, opposite Mill Street,
Dover
BIO 43° 10’39’N, 70°52’30”W 14.25 Headwaters, below tidal dam, near
Sawyer’s Mills, Dover
COCHECO RIVER
(NEW HAMPSHIRE)
Miles
Station From
No. Latitude and Longitude Coast Description
Cl 43° 10’44’N, 70°49’48”W 12.6 Mouth, on east bank and opposite
peninsula separating Cocheco from
Piscataqua River, Dover
C2 43° 10’58”N, 70°50’09"W 13.2. West bank, just opposite and
somewhat south of the end of
Three Rivers Road, Dover
C3 43°11'12”N, 70°50’'17”W_ 13.3. East bank at the mouth of Fresh
Creek, Dover
C4 43° 1110’N, 70°50’27”W 13.5. West bank, just opposite mouth of

Fresh Creek, Dover

1986]

C5

C6

C7

C8

C9

C10

Cll

Cl2

Cl3

Cl4

C15

Cl6

Cl?

Mathieson & Hehre

43°11'44’"N,

43°11'25’N,

43° H121’N,

43°11'30"N,

43° 11°30’N,

43° 1138"N,

43°11'44’"N,

43° 11'46'N,

43° 11'50’N,

43° 11'47’"N,

43° 11'47’°N,

43° 11'40’N,

43°11'41°N,

70° 50°13” W

70° SO0’34"W

70° 50°48” W

70° ST’OL"W

70° STI’ W

70° SILT” W

70° 51'26”"W

70° 51°48” W

70° 51°52” W

70° 51°58” W

70° 52°15” W

70° 52°15” W

70° 52°22” W

New Hampshire seaweeds

14

13.7

13.9

14.05

14.25

14.4

14.6

14.7

14.9

15

15.4

i

Fresh Creek at Rte. 101 Bridge (Gulf
Road), Dover

East bank, just upstream from Fresh
Creek, Dover

East of McKone Road Landing, just
downstream from lower narrows,
Dover

Lower narrows on west bank near
marsh from McKone Road
Landing, Dover

Upstream from lower narrows,
northeast of McKone Road
Landing, Dover

East bank just east of Cocheco
Country Club, opposite McKone’s
Marsh, Dover

East bank, west of Cocheco Country
Club and just upstream from the
mouth of Emerson Brook, Dover

East bank near Dover Sewage
Treatment Plant and red brick
factory, Dover

East of red brick factory, near
cemetery at Cocheco Street, east
bank, Dover

East of Dover sewage treatment
plant, west bank, Dover

Northeast of Old Water Street Bridge.
near George’s Marina and the
Davis School, Dover

Between the two Water Street bridges
and on west bank, Dover

Headwaters, at Central Avenue and
near the fish ladder, Dover

130 Rhodora [Vol. 88

LAMPREY RIVER
(NEW HAMPSHIRE)

Miles
Station From
No. Latitude and Longitude Coast Description

Ll 43°03'52”N, 70°54'53”W 5.8 Mouth, second promontory upstream
on Shackford Point, Newmarket

L2 43°(03'57”N, 70°54’49”"W 15.75. Mouth, just upstream from Moody’s
Point on north bank, Newmarket

L3 43°04'07”N, 70°55'12”W_ 16.25. East bank opposite fish seines by
private dock, Newmarket

L4 43°04'09"N, 70°55’20"W 16.4 Just beyond fish seine on west bank
near Birch Drive, Newmarket

LS 43°04'19"N, 70°SS'38"W 16.6 West bank in small cove between
lower narrows and fish seines,
Newmarket

L6 43°04'27"N, 70°55'39”W 16.75 East bank just south of lower
narrows and opposite overhead
power cables, and opposite the end
of Young’s Lane, Newmarket

L7 43°04'31"N, 70°55’47”W_ 16.85 West bank and southwest of overhead
power cables and opposite the end
of Young’s Lane, Newmarket

L8 43°04’38’N, 70°56'06"W_ «17.25. Upper narrows, east bank, Newmarket

L9 43°04’50’”N, 70°S56’01”W 17.5 Headwater, near dam to Sewage
Treatment Plant and at Rte. 108,

Newmarket

OYSTER RIVER
(NEW HAMPSHIRE)

Miles
Station From
No. Latitude and Longitude Coast Description

Ol 43°07'23”N, 70°52’20"W 11.5 Mouth, northwest tip of Durham
Point and just north of Langley
Road, Durham

O2 43°07'29"N, 70°52’17”"W_ 11.5. Mouth, Emerson’s Beach, pier area
and red boat house, opposite from
Durham Point, Durham

1986] Mathieson & Hehre — New Hampshire seaweeds 131

O03 43°07'32’N, 70°52'53”W_ 11.95 Large private dock, north of Mathes
Farm Road, Durham

O4 43°07'42’N, 70°52’36’W_s11.8 Smith Creek, mouth, Durham

O5 43°07'43’N, 70°52’58”"W 12.25. Midway between Smith and Bunker
Creeks on the Rte. 4 side, Durham

06 43°07’58’"N, 70° 53’10"W 12.5 Mouth, Bunker Creek, Durham

O7 43°07'52’N, 70°53’50”"W 13 Directly across from the mouth of
Johnson Creek, Durham

O%& 43°07'59”"N, 70° 53’48"W 13 Mouth, Johnson Creek, near
Riverview Court ending, Durham

O09 43°08'14”N, 70°54’00"W_ 13.45 Johnson Creek Bridge at Rte. 4,
Durham

O10 43°07’59”"N, 70°54’16”"W_ 13.35. Mouth, Horsehide Brook, Durham

Oll 43°08'12”N, 70°54’27”"W 13.6 = Just upstream of Durham Waste
Treatment Plant, Durham

O12 43°08'05’N, 70°54’40”W 13.75. Opposite Jackson’s Landing, Durham

O13 43°08'05’N, 70°54’48”W 13.90 Mouth, Beards Creek, Durham

O14 43°07'52”N, 70°55’06"W 14.2 Headwater at Rte. 108, along Old
Landing Road, Durham

SALMON FALLS RIVER
(NEW HAMPSHIRE/ MAINE)
Miles
Station From

No. Latitude and Longitude Coast Description

SFI 43° 10’'48”N, 70°49’40”W 12.5. Mouth, on east bank of peninsula
separating Piscataqua River from
Salmon Falls River, Dover, N.H.

SF2 43° 10'37”"N, 70°49'26"W_ 12.6 Mouth, east bank, South Berwick,
Maine

SF3 43° 10’52”N, 70°49'16”"W 12.7 First brook on Salmon Falls River,
mouth, South Berwick, Maine

SF4 43° 10’53”N, 70°49’41”W 12.8 West side, opposite the mouth of the
first brook (or SF3), Dover, New
Hampshire

SF5 43° 11°18"N, 70°49’38”W 13.25 East bank, just south of Rte. 101

(Eliot) bridge, South Berwick,
Maine

132

Rhodora

Salmon Falls River (NH) (Cont.)

[Vol. 88

Miles
Station From

No. Latitude and Longitude Coast Description

SF6 43° 11’23”"N, 70°49’48”W 13.3. West bank, opposite and somewhat
south of Rte. 101 (Eliot) Bridge,
Dover, New Hampshire

SF7 43°11'24”"N, 70°49°30"W 13.5 East side at Rte. 101 (Eliot) Bridge,
South Berwick, Maine

SF8 43° 11'25”N, 70°49’20"W 13.8 — Just upstream from Rte. 101 (Eliot)
Bridge, east bank at end of Water-
side Lane, South Berwick, Maine

SF9 43° 11'40’N, 70°48’S9"W O14 East bank, by cemetery, near Rte. 101
(Eliot) Bridge, South Berwick,
Maine

SFIO 43° 11’S50"N, 70°49'06"W_) 14.2. Above SF9, approximately 1/3 the
distance between Rte. 101 (Eliot)
Bridge and Hamilton House, South
Berwick, Maine

SFII — 42°59’47”N, 70°56’'20"W_ 14.8 = Mouth, Sligo Brook, Rollinsford,
New Hampshire

SFI2 = 42°59’43”N, 70°51’20”"W_ 15.2 Hamilton House near mouth of
Hamilton Brook, South Berwick,
Maine

SFI3. 43° 11’S9”N, 70°49’11"W 15.75 East bank, just below Leigh’s Mill
Pond, South Berwick, Maine

SFI4 43° 12’01"N, 70°49’23”W_ 15.75 West bank near Sligo Road and
opposite SF13, Rollinsford, New
Hampshire

SFIS 42°51’03”N, 70°57'00"W_ 16.3. East bank just above Leigh’s Mill
Pond, South Berwick, Maine

SF16 — 42°58’51”N, 70°56’43”W_ 16.5 Headwater at Portland Avenue

Bridge, east and west banks, South
Berwick, Maine

1986] Mathieson & Hehre — New Hampshire seaweeds 133
SQUAMSCOTT RIVER
(NEW HAMPSHIRE)
Miles
Station From

No. Latitude and Longitude Coast Description

Sl 43°03'09"N, 70°54’47"W 16.2 Mouth, on west bank at railroad
bridge, Newfields

$2 43°02’59”"N, 70°55’03”W_ 16.5. West bank near creek and just
upstream from railroad bridge,
Newfields

S3 43°02’S1”N, 70°55'02"W 16.7 West bank, just upstream of S2 and
midway between towers and Rte.
108 Bridge, Newfields

S4 43°02’33’N, 70° 55’09"W 16.9 Fast bank at towers for overhead
power cable, Stratham

SS 43°02'24”"N, 70°55’43”W_ 17.45 Bridge at Rte. 108, Chapman’s
Landing, Newfields

S6 43°02’01’N, 70°56’13”W_sI8 East bank, upstream of Chapman’s
Landing and | 4 of the way between
SS and S9, Newfields

S7 43°01735”"N, 70°5S6'04"W 18.55 East bank, halfway between SS and
S9, Stratham

S8 43°OV TIN, 70°55’57”W 19.2. East bank near private dock and
three quarters of the way between
S5 and S9, Stratham

S9 43°00'46"N, 70°56’23”W 19.75. West bank by railroad track, near
overhead power lines (towers),
Exeter

S10 43°00’01"N, 70°56’24”"W 20.75 West bank, just upstream of oxbow
cut and just north of Rte. 101
fixed bridge, Exeter

SI 42°59’48’"N, 70°S6'19"W 21 Opposite the mouth of Wheelwright
Creek, Exeter

$12 42° 59’43’N, 70°56’20”W 21.2 East bank, just upstream from the
mouth of Wheelwright Creek,
Exeter

S13 42°59’31"N, 70°5S6'42”W 21.5 West bank just upstream from
Powell’s Point, Exeter

S14 42°59'17’"N, 70°57'°03"W 21.9. West bank by dike and water outfall,

also near the mouth of Norris
Brook, Exeter

134 Rhodora [Vol. 88

Squamscott River (NH) (Cont.)

Miles
Station From
No. Latitude and Longitude Coast Description

S15 42°59'03’N, 70°S701°W 22.25 Just upstream and opposite Powder-
house Point, Exeter

S16 42° 58’52”N, 70°56’41"°W 22.7 Headwaters at tidal dam, Exeter

WINNICUT RIVER
(NEW HAMPSHIRE)

Miles
Station From
No. Latitude and Longitude Coast Description

Wi 43°02’47”"N, 70° 50’40"W_ 15.8 Mouth, on west bank and near the
mouth of Shaw Brook, across from
Portsmouth Country Club,
Greenland

W2 43°02’52”N, 70° 50’'16”W 16.25. Mouth, on east bank and downstream
from Packer’s Brook, a cove area,
Greenland

W3 43°02’°31"N, 70°50’'28"W 16.75 East bank just downstream from
railroad bridge and near the end
of Tide Mill Road, Greenland

W4 43°02’12”N, 70° 50’5S”W_ 17.25. Headwaters at the Rte. 101
(Portsmouth Avenue) Bridge,
Greenland

HAMPTON-SEABROOK ESTUARY
(NEW HAMPSHIRE)
HAMPTON RIVER AND ADJACENT TRIBUTARIES

Miles
Station From
No. Latitude and Longitude Coast Description
A-| 42°51'46"N, 70°47'02"W_ 1.3 At the mouth of the first major

tributary SE of Tide Mill Creek
on the Hampton River, Hampton

1986]

A-2

A-4

A-5

Mathieson & Hehre

42°54'40’'N,

42°54’48’N,

42°54’49"'N,

42°55‘04’N,

42°55’19’N,

42°55'34’N,

42°55'12’N,

42°55'24’N,

42°54’47’'N,

42°55'40’N,

42° 54'39"'N,

42° 54’58’'N,

42°55'12”"N,

70° 49'06"W

70° 49°40” W

70° 50'04”W

70° 50’32” W

70° 5010” W

70° 50°30” W

70° 5008” W

70° 49'08" W

70° S1'18”"W

70° 50’38” W

70° S1°16"W

70° S048” W

70° 5S0’42”W

-- New Hampshire seaweeds 135

1.9]

1.3

3.3

3.7

The northeast portion of an “island”
formed at the mouth of the Taylor
River, Blind Creek and the upper
part of Hampton River, Hampton
Falls, Hampton

Approximately 1500 feet NE of
mouth of Tide Mill Creek and the
Hampton River, Hampton

Approximately 1800 feet NW of
station A-2 on the Hampton River,
Hampton Falls, Hampton

Opposite a small brook, which
empties into the first tributary
above the mouth of Hampton
Falls River, Hampton

At the junction of the first oxbow
NE of Station A-S on the Taylor
River, Hampton

Hampton Landing on Taylor River,
Hampton

At the mouth of Nudds Canal and
Blink Creek, Hampton

Tide Mill Creek by the Route 101
bridge, Hampton

Hampton Falls River south of Depot
Avenue and near the Boston and
Maine Railroad bridge, Hampton
Falls

A site approximately 2000 feet SW
of the Boston and Maine sub-
station, which is between Lafayette
and Landing Roads. Adjacent to the
Boston and Maine railroad tracks;
itis on Taylor River in Hampton

End of Depot Avenue on Hampton
Falls River, Hampton Falls

Middle of the southernmost oxbow
near the mouth of Taylor River
and the Hampton town line

A bend in the first tributary above
(north) of Hampton Falls River
where the river crosses the

136 Rhodora [Vol. 88

Hampton-Seabrook Est./ Hampton R. & Adj. Trib. (NH) (Cont.)

Miles
Station From
No. Latitude and Longitude Coast Description

railroad tracks. Collections were
made on the harbor side of the
tracks, Hampton

A-IS 42°55’12”N, 70°51’04"W 3.7 Same as A-14, except the land side
of the railroad tracks, Hampton

A-16 — 42°55’13”N, 70°50’°42”W_ 3.5 A salt marsh on a point of land
made by the Hampton River and
the first tributary above the
Hampton Falls River, between
stations A-5 and A-14, Hampton

A-17 — 42°55’20’N, 70°49’54”"W 3.01 Hampton Landing, Taylor River,
Hampton

A-18 — 42°54’34”N, 70°49'24"W_ 1.6 The Willows—at the mouth of Tide

Mill Creek and Hampton River,
Hampton

BROWN RIVER AND ADJACENT TRIBUTARIES

Miles
Station From
No. Latitude and Longitude Coast Description

B-1 42°53’55”N, 70°49’06"W_ 1.0 A small “island” opposite Eastman’s
Slough and about 2500 feet west
of the Locke Point State Park area

B-2 42°53’45”"N, 70°50’14"W 1.7 Southernmost portion of Eastman
Slough, near Halftide Rock and at
the mouth of the Brown River,
Hampton Falls-Seabrook

B-3 42°53’40’N, 70°50’40’W_ 1.8 Just inside the mouth of Hunt’s
Island Creek at the junction of
Brown River (on the east side of
the channel). Approximately 600
feet SW of B-2, Seabrook

B-4 42° 53’59”N, 70°50’28”"W_ 2.0 Approximately 500 feet NW of the
first tributary past Hunt’s Island
Creek, Hampton Falls~Seabrook

1986] Mathieson & Hehre — New Hampshire seaweeds 137

B-5 42°54’11”N, 70°50’°18"W_ 2.2 Approximately 700 feet NE of the
mouth of Swain’s Creek, Hampton
Falls

B-6 42°54’17’"N, 70°S0'02”"W 2.3 Robbins Point, Hampton Flats,
Hampton Falls-Seabrook

B-7 42°54’16”"N, 70°S0’'14”"W 2.5 Brown’s River, first tributary
upstream from Swain’s Creek,
Hampton Falls

B-8 42° 53’59”N, 70°S0’°18”W 2.4 Approximately 800 feet upstream
from Robbin’s Point, Hampton
Falls-Seabrook

B-9 42°54’08"N, 70°50’'24”"W 2.53 Approximately 700 feet upstream
from Station B-8, Hampton
Falls-Seabrook

B-10 = 42°54’07”N, 70°50’42”W 2.8 End of Rock’s Road on the Brown’s
River, Hampton Falls-Seabrook

B-11 42° 54’23’”N, 70°50'46"W 3.0 Near the mouth of the first major
tributary east of the head waters
of Brown’s River, Hampton Falls-
Seabrook

B-12 — 42°54’14’”N, 70°S110"W 3.3 Approximately 1500 feet upstream
(west) from Station B-I1, just
before a major oxbow, Hampton
Falls-Seabrook

B-13 42° 54’26"N, 70°49'08"W_ 2.6 Swain’s Creek, neck of first oxbow,
Hampton Flats, Hampton Falls

BLACKWATER RIVER AND ADJACENT TRIBUTARIES

Miles
Station From
No. Latitude and Longitude Coast Description

C-1 42° 53’12”N, 70° 49’32”W_ 1.5 Mouth of the Blackwater River near
the first tributary SW of Mills
Point, Seabrook

C-2 42°52’19”N, 70° 50’28”W 2.2 Approximately 1200 feet SW of the
first tributary past Riverside,
Seabrook

C-3 42°52'12”N, 70°50’08”"W 2.3 Approximately 1200 feet south of C-2,
Seabrook

138 Rhodora [Vol. 88

Blackwater & Adjacent Tributaries (Cont.)

Miles
Station From
No. Latitude and Longitude Coast Description

C-4 42°52’30’N, 70°49'24"W 2.4 Approximately 500 feet SE of C-3,
Seabrook

C-5 42°52’55”"N, 70°49'34”"W 2.5 Approximately 800 feet SE of C-4,
Seabrook

C-6 42° 52’32”N, 70°49'18”"W 2.7 Approximately 800 feet SE of C-5,
near the first major tributary SE of
C-2, Seabrook

C-7 42° 52’28"N, 70°49'12"W 3.0 Approximately 1200 feet SE of C-6
near a large white rock

C-8 42°52’21”"N, 70°49:08"W 3.3 Approximately 1500 feet SE of C-7,
Seabrook

C-9 42° 52’12”N, 70°49'02”"W 3.6 By the route 268 bridge that crosses
the Blackwater River, Seabrook

C-10 42°53’10’N, 70°49'24"W si: Mill’s Point at the mouth of
Blackwater River, Seabrook

C-I1 42°53'02’N, 70°49'44’"W 1.7 Riverside, Seabrook

KNOWLES ISLAND AND MILL CREEK AREAS

Miles
Station From
No. Latitude and Longitude Coast Description
D-| 42° 53'42”N, 70°49'52”W 1.34 Mouth of Creek, between Knowles

Island and mainland, Seabrook

D-2 42° 53’42”N, 70°50’22”W_ 1.6 Before the first bend, near D-1,
Seabrook

D-3 42° 53'45”N, 70°50'34”"W_ 2.1 Walton Landing at the end of
Walton Road, Seabrook

1986] Mathieson & Hehre — New Hampshire seaweeds 139

HAMPTON HARBOR
(NEW HAMPSHIRE)

Miles
Station From
No. Latitude and Longitude Coast Description

H-1 42°53’40’N, 70°49'18’"’W_ 0.4 Hampton Harbor at the junction of
the middle piling of the tall bridge
and the tower at Hampton

H-2 42° 53'20’N, 70°49'24"W 1.0 Hampton Harbor, in the channel
near the mouth of the Blackwater
River and at the junction of the
imaginary line between Seabrook
Marina and Knowles Island,
Hampton

H-3 42°53’5S”N, 70°48’'58”W_ 0.8 Hampton Harbor, Smith and
Gilmore Marina, Hampton

H-4 42°54'05’N, 70°49'12”W_ 0.97 Hampton Harbor, Hampton Marina
at the mouth of Hampton River,
Hampton. The station was the
point protruding into the harbor
proper.

CYTOLOGY OF CAREX PURPURIFERA MACK.
(CT PERACEAE)

JAMES R. MANHART

ABSTRACT

Field work and morphological studies on section Laxiflorae Kunth of the genus
Carex L. revealed morphological variation in C. purpurifera Mack. is correlated with
geographical distribution. Carex purpurifera appears to consist of two taxa, one
found on soils derived from limestone in the western part of the range and the other
on non-limestone-derived soils in the eastern part of the range. Morphological differ-
ences were retained in greenhouse-grown plants. Pollen mother cell counts of the
western populations were n = 17, 18, and 19. The eastern populations were n = 14
only, the lowest recorded in section Laxiflorae. These cytological data support the
contention that C. purpurifera consists of two distinct taxa and the newer one, C.
manhartii Bryson, has been described elsewhere as a species.

Key Words: Cytology, Carex purpurifera, section Laxiflorae, Cyperaceae

Cytological studies of the genus Carex L. have long been a source
of considerable confusion and speculation. Members of the genus
Carex exhibit remarkable variation regarding chromosome numbers.
In North American Carex, haploid numbers range from 6 to 56 with
every number from 12 through 43 represented (Wahl, 1940). There
is also a great deal of intraspecific variation; Tanaka (1949) pointed
out that two to as many as six different chromosome numbers have
been found in 70 different species.

Polyploidy is thought to be unimportant in the evolution of the
genus Carex (Heilborn, 1924). Agmatoploidy, a special type of
aneuploidy resulting from chromosome fragmentation coupled with
unlocalized centromeres, is considered the major cause of speciation
in Carex (Davies, 1956). In organisms with localized centromeres,
breakage of the chromosomes usually results in the loss of the non-
centromere-containing fragment. However, when unlocalized cen-
tromeres are present, the fragments can be perpetuated and become
permanent components. Agmatoploidy then provides an explana-
tion for the fact that within an aneuploid series the taxa which have
the higher chromosome numbers have smaller chromosomes.

'Based on a dissertation submitted by the author to the Graduate School, University
of Georgia, in partial fulfillment of the requirements for the Ph.D. degree.

14]

142 Rhodora [Vol. 88

Faulkner (1972) showed that in section Acutae Fries., variation
within species is mainly aneuploid and that “there seems to be a
remarkable instability of chromosome numbers within species of
Carex.” This variation is structural, not quantitative, and probably
arises by chromosome fission and fusion due to unlocalized centro-
meres (Faulkner, 1972). The occurrence of agmatoploidy in Carex
has not been unequivocally demonstrated, as Grant (1981) pointed
out; however, the chromosome behavior of Carex closely resembles
that of organisms in which unlocalized centromeres have been more
clearly demonstrated (Faulkner, 1972).

Carex purpurifera Mack. was originally described and placed in
section Laxiflorae Kunth by Mackenzie (1931-35) in his revision of
North American Carex. Gleason (1952) treated C. purpurifera and
six other taxa of section Laxiflorae as varieties of C. /axiflora Lam.,
while Bryson (Ph.D. thesis, Mississippi State Univ., 1980), in a
revision of section Laxiflorae, retained all these taxa as distinct
species, including C. purpurifera. Bryson gave a more detailed des-
cription of the distribution of C. purpurifera since Mackenzie
(1931-35) had only seen plants from the type locality of Campbell
Co., TN and one other site in Tennessee. Carex purpurifera is
mainly found in soils which overlay limestone in dry to mesic woods
along steep slopes in the southern Appalachian and Cumberland
Plateau from West Virginia and eastern Kentucky south into Ala-
bama, Georgia, North Carolina, and South Carolina (Bryson,
Ph.D. thesis, Mississippi State Univ., 1980). Bryson pointed out
that specimens of C. purpurifera from south and east of the Appal-
achian Divide in Georgia, North Carolina, and South Carolina have
narrower leaves and less purple coloration on the leaf bases. During
the course of this investigation, it was noted that plants from west-
ern populations are more robust, have glaucous leaves and long-
pedunculate staminate spikes, whereas leaves of plants from eastern
populations (in a very limited area without limestone underlying the
soils) are not glaucous and the staminate spikes are short-
pedunculate or sessile. These morphological differences are retained
in greenhouse-grown plants.

Of further note, Radford et al. (1968) stated that Carex graci-
lescens Steud. is similar to C. purpurifera. This observation is espe-
cially true of the eastern plants, which would be the only ones
occurring in North Carolina and South Carolina. However, the

1986] Manhart — Carex 143

western plants are quite distinct from C. gracilescens, as this taxon
has narrow, non-glaucous leaves with relatively less purple colora-
tion on the leaf bases and a more crowded pistillate spike.

A cytological study of Carex purpurifera was undertaken for four
reasons: (1) to determine chromosome counts for C. purpurifera
since there are none in the literature; (2) to determine if there is a
correlation of chromosome numbers with the morphological and
habitat differences found between the western and eastern popula-
tions; (3) to determine if differences in chromosome numbers occur
in plants from morphologically similar populations and; (4) to allow
comparison of the chromosome numbers for the eastern popula-
tions of C. purpurifera with the three counts of n = 20 obtained for
C. gracilescens by Wahl (1940).

MATERIALS AND METHODS

Meiotic counts were obtained from staminate spikes taken from
plants grown in the University of Georgia Botany Department
greenhouse complex and preserved in a solution consisting of
chloroform:acetic acid:methanol (4:3:1 v/v). Anther squash prepa-
rations were stained with 1% aceto-carmine (Jones and Luchsinger,
1979) and photographs made of the chromosome complements
(Figures 1-4). Results are based on observations of 3 to 8 cells for
each preparation. Vouchers, each with a drawing and/or a photo of
a countable figure, have been placed in the University of Georgia
Herbarium (GA).

RESULTS AND DISCUSSION

Populations A, B, and C are from northwestern Georgia; popula-
tions B and C are 8 km from each other (Figure 5). Population D is
located in Alabama 100 km west of the Georgia populations. All of
these populations occur in soils with underlying limestone and the
plants have wide, glaucous leaves with deep purple leaf bases and
long-pedunculate staminate spikes. Population E from Macon Co..,
NC (Figure 5) occurs on soil with no underlying limestone as does
population F which is from northeastern Georgia. Plants from pop-
ulations E and F all have narrow non-glaucous leaves, with rela-
tively less purple coloring on the leaf bases and _ sessile or
short-pedunculate staminate spikes.

144 Rhodora [Vol. 88

Figures 1-4. Pollen mother cells (PMC’s) of Carex purpurifera populations.
Scale = 5 micrometers. 1. A. n = 19, 2. C.n = 17, 3. D. n = 18 (arrow indicates
location of chromosome out of focal plane), 4. E.n = 14

As with other taxa in the genus Carex, there is considerable
chromosome variation in C. purpurifera. Meiotic counts for the
four western populations vary from n = 17 to n = 19 (Table 1),
numbers which are within the range for other taxa in the C. laxiflora
complex (Wahl, 1940; Whitkus, 1981). The 7 = 14 count (Table 1) is
the only one recorded for the eastern populations and is the lowest
ever recorded in section Laxiflorae. Thus, the morphological and
distributional differences between the western and eastern taxa are
correlated with differences in the range of chromosome numbers
found in these taxa.

Manhart Carex

1986]
= eri een
/° NORTH
TENNESSEE ee CAROLINA
\ SE ESS & ie aes eek Fee, 5 aa nt SR eS ad =
iC are
' D \ Ph
| B ‘SOUTH
LA ‘CAROLINA
' \ oS;
; >
! 1 4
\ %
| \ =
i \ ~~
H \ \
i \
i ALABAMA \ GEORGIA
; 100 mi
: 160km

Figure 5. Locations of populations of Carex purpurifera sampled for cytological

study.
CONCLUSIONS

The cytological data obtained in this study, in conjunction with
morphological and habitat considerations, indicate that Carex pur-
purifera consists of two distinct taxa and also demonstrate that
chromosome data can be quite useful in solving taxonomic prob-
lems in the genus Carex. In terms of the taxonomic treatment of the
two taxa, Bryson (1985) has designated the eastern taxon as C.
manhartii Bryson, a new species distinct from C. purpurifera.

146 Rhodora [Vol. 88

Table 1. Meiotic Chromosome Counts (1) of Carex purpurifera
Population n Figure —_ Locality and Voucher
A 19 I Floyd Co., GA, east bank of Coosa River on
south side of Rome, Manhart 378
B 18 Walker Co., GA, Pigeon Mountain Wildlife

Management Area west of Lafayette and south
of GA 93, Manhart 375

Cc 17 2 Walker Co., GA, 8 km northwest of Lafayette,
Manhart 377

D 18 3 Madison Co., AL, Monte Sano Mountain east
of Huntsville, Bryson and Manhart 3156

E 14 4 Macon Co., NC, The Stewart Trail, Coweeta
Hydrologic Laboratory, Manhart 293

F 14 Union Co., GA, Woods along Chattahoochee

River | km north of Union Co.-White Co.
line, Manhart 496

There is not enough known about speciation in Carex to postu-
late solely on chromosome numbers which of these two taxa is the
“ancestral” or “most primitive” type. If it is accepted that speciation
via agmatoploidy does occur in Carex, it is still not clear whether
fission or fusion or both are involved in this process; Faulkner
(1972) cited instances in which both may be involved in section
Acutae. If the fission hypothesis is invoked, then the eastern taxon
would have given rise to the western taxon and/or C. gracilescens
n= 20; Wahl, 1940). Of course, the converse would be true if one
accepts fusion as the mode of chromosomal evolution. There is
simply not enough known at the present time to make a prudent
choice. Further studies which incorporate morphological, chemical,
distributional, and cytological data for all of section Laxiflorae
should help to shed some light on speciation processes in Carex.

ACKNOWLEDGMENTS

I thank Nanette M. Manhart for map preparation and darkroom
work, Gerald L. Smith for advice on chromosome figure interpreta-
tion, Charles T. Bryson for many helpful discussions on section
Laxiflorae, David E. Giannasi for critical review of the manuscript
and the University of Georgia Herbarium (GA) for technical sup-

1986] Manhart — Carex 147

port. This work was supported by a grant from the Botany Depart-
ment at the University of Georgia, a Grant-in-Aid of Research from
Sigma Xi, and NSF Pre-Doctoral Dissertation Improvement Grant
DEB-8 105474.

LITERATURE CITED

Bryson, C. T. 1985. A new species of Carex (Cyperaceae: sect. Laxiflorae) from
the southern Appalachians. Castanea 50: 15-18.

Davirs, FE. W. 1956. Cytology, evolution, and origin of the aneuploid series in the
genus Carex. Hereditas 42: 349-365.

FAULKNER, J. S. 1972. Chromosome studies on Carex section Acutae in north-
west Europe. Bot. J. Linn. Soc. 65: 271-301.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of the Nor-
theastern U.S. and Adjacent Canada. Vol. |, Hafner Press, New York.

GRANT, V. 1981. Plant Speciation, 2nd ed. Columbia University Press, New
York.

HEILBORNE, D. 1924. Chromosome numbers and dimensions. species-formation
and phylogeny in the genus Carex. Hereditas 5: 129-216.

JONEs, S. B. AND A. E. LUCHSINGER. 1979. Plant Systematics. McGraw-Hill Inc..
New York.

MACKENZIE, K. K. 1931-35. Cariceae. North American Flora 18(7): 1-478. New
York Botanical Garden, New York.

RaprorD, A. E., H. E. AHLES, AND C. R. BELL. 1968. Manual of the Vascular
Flora of the Carolinas. The University of North Carolina Press, Chapel Hill.

TANAKA, N. 1949. Chromosome studies in the genus Carex with special reference
to aneuploidy and polyploidy. Cytologia 15: 15-19.

Want, H. A. 1940. Chromosome numbers and meiosis in the genus Carex. Amer.
J. Bot. 27: 458-470.

Wuitkus, R. 1981. Chromosome numbers of some northern New Jersey Carices.
Rhodora 83: 461-464.

DEPARTMENT OF BOTANY
UNIVERSITY OF GEORGIA
ATHENS, GEORGIA 30602

BOOK REVIEW

CHARLES W. JOHNSON. Bogs of the Northeast, Pp. 269 + xiv, tab
76. University Press of New England, Hanover, N.H. (1985)
$12.95 paper; $25.00 cloth.

Bogs have always intrigued man—from earliest times as places of
mysterious spirits and happenings; in modern periods as unusually
interesting environmental curiosities.

The author, Vermont State Naturalist, is well qualified to write a
volume on New England bogs, for he has spent a lifetime in the
fields and forests of this part of the country. To the best of my
knowledge, this book is the first popular but scientifically sound
publication on these often misunderstood ecological peculiarities
that many developers, political officials and other uninformed citi-
zens frequently want to destroy.

The volume is divided into 17 chapters: 1) Perception of Bogs, 2)
Basic Terms & Definitions, 3) Genesis of Peatlands, 4) Ferns &
Bogs, 5) Geography of Northeastern Peatlands, 6) Plants & Ani-
mals, 7) Sphagnum Moss, 8) Carnivorous Plants, 9) Orchids, 10)
Sedges & Heaths, 11) Insects & other Invertebrates, 12) Fish,
Amphibians, Reptiles, 13) Birds, 14) Mammals, 15) Vaults of His-
tory, 16) Human Uses of Peatlands, and 17) Preservation or Obli-
teration? There follow a Bibliography of 117 entries, four appendices
(peatlands, amphibians and reptiles, birds, mammals) and a highly
detailed index.

This book is a definite contribution to New England botany.

RICHARD EVANS SCHULTES
BOTANICAL MUSEUM
HARVARD UNIVERSITY
OXFORD STREET
CAMBRIDGE, MA 02138

149

150 Rhodora [Vol. 88
BOOK REVIEW

D’Arcy, William G. and Mireya D. Correa A., Editors. 1985. The
Botany and Natural History of Panama: La Botanica e Histo-
ria Natural de Panama. Monographs in Systematic Botany,
Vol. 10, Missouri Botanical Garden. 455 pp. Dept. I1, Mis-
souri Botanical Garden, Box 299, St. Louis, MO 63166-0299
($48.00 plus 4% shipping charges)

This paperback book is the product of a symposium held in
Panama April 14-17, 1980, to commemorate completion of the
Flora of Panama, a project of long duration by many workers and
collaborators at MBG. Its most striking feature is interspersal of
Spanish summary paragraphs in English-language articles and Eng-
lish summary paragraphs in the Spanish-language articles. The
reader is thus able to jump easily from one language to the other on
nearly every page; the effect is that of a bilingual truly collaborative
effort by specialists in the two cultures. One of my colleagues at
Tufts checked the Spanish for content and readability, and ranked it
first-class. Extensive literature citations are found with each article.

The text first presents data on Panama and a list of 55 author’s
addresses followed by papers in three groups: Part I—The Organ-
isms, subdivided into Plant Groups, Animal Groups, Plants and
Geography, and Plant Study Techniques; Part I[—The Interactions
of Plants, Animals and Habitats; Part I1I—The Human Aspect,
subdivided into Plants that Affect Man, Indigenous Peoples, and
Modern Man and the Landscape. Under each, a distinguished gal-
axy of contemporary specialists present and summarize aspects of
their original work in Panama. The accounts are for the most part
fascinating, perhaps more so for me because I was a member of a
few of the parties doing the field work described. It is therefore
possible for this reviewer to vouch firsthand for the authenticity of
some of the work. Topics covered range from a history of plant
collecting in Panama through ecological, evolutionary and beha-
vioral studies of particularly prominent groups of plants and anim-
als, from uses and abuses by people of the Panamanian tropical
forests, to a genuine concern for conservation of the natural resour-
ces of Panama on the part of some Panamanians, and ending with
the saddening and sobering final observation that as yet really effec-
tive conservation policies have not become a priority of the central
government.

1986] Book Reviews 1S]

This volume is an intriguing history of thirty years of concen-
trated field effort which in every instance has an ongoing compo-
nent still pursued by most of the same authors who wrote these
papers. The flora is not yet completely tabulated, profound changes
in land use are daily altering its bases and its chances of survival,
and there is developing in Panamanians an even greater sense of
urgency to preserve what is left of these magnificent tropical forests.

Anyone interested in the future of Central America, or who is
working in any branch of its natural history, will find this volume a
rich source of Panamanian cultural history, present technical prob-
lems, and ongoing directions for future conservation and research
activities.

NORTON H. NICKERSON
DEPARTMENT OF BIOLOGY
TUFTS UNIVERSITY
MEDFORD, MA 02155

IN MEMORIAM

BERNARD BOIVIN 1916-1985

The following tribute to one of our former members, Bernard
Boivin, appeared in The Plant Press for September 1985 (Vol. 3.,
No. 3).

On 9 May 1985 Canada lost one of its most productive botanists,
Bernard Boivin. Born and raised in Montreal, he developed an
interest in botany through his undergraduate education at the Uni-
versity of Montreal and the influence of Quebec botanists, especially
Frere Marie-Victorin. He obtained a doctorate under Fernald at
Harvard University in 1943. Although most of his career was at the
Biosystematics Research Institute of Agriculture Canada in Ottawa,
Dr. Boivin spent a number of years as a researcher at the Gray
Herbarium of Harvard University, the National Museums of Can-
ada, and the University of Toronto. For several years he was a
researcher and lecturer at Laval University, where he continued to
work until shortly before his death.

A self-described herbarium botanist, Bernard Boivin literally
worked day and night on his many projects. He was the author of
more than 150 publications; the best known include “Enumeration
of the Plants of Canada,” “Flora of the Prairie Provinces,” and
“Survey of Canadian Herbaria.” He had completed a number of
as-yet-unpublished manuscripts, the most recent on Carex of east-
ern Canada. Bernard Boivin published more names of plants, many
of them new combinations, than any other Canadian botanist. Stu-
dents of systematics will miss Dr. Boivin’s lively discussions about
aspects of taxonomy and nomenclature. Even when he might dis-
agree on these matters, he was always good-humored and interested
in their work.

Bernard Boivin believed that learning about plants is an impor-
tant part of a good education for all Canadians. Realizing the value
of adequate herbarium material for research, he encouraged anyone
interested in plants to collect specimens. Many amateur botanists in
Ontario, and the rest of Canada, were inspired by Dr. Boivin’s
encouragement and as a result they have made significant contribu-
tions to our knowledge of field botany. Dr. Boivin maintained
extensive files for his “Index of Canadian Plant Collectors.” In
addition to corresponding with hundreds of individuals whose

152

1986] In Memoriam ioe

names appear on herbarium labels, he accumulated copious bio-
graphical information from newspapers and other documents. All of
this material, which will be of great value to historians, is to be kept
at the Hunt Institute for Botanical Documentation in Pittsburgh,
Pennsylvania.

LAURENCE EATON RICHARDSON 1893-1985

The Concord Journal for Thursday, October 3, 1985 carried a
notice of the death of one of our long-time members, Laurence E.
Richardson, which we quote in part:

Lawrence Eaton Richardson, 91, of Barrett’s Mill Road, a
Concord historian and a lifelong resident, died Thursday Sept. 26,
1985 in Acton. Born in Concord on December 10, 1893, he was the
son of the late Henry Preston Richardson and the late Maria
(Smith) Richardson. After attending Concord schools, he graduated
from Concord High School in 1912 and from Harvard College in
1916.

Mr. Richardson served as a captain with the military police of the
U.S. Army during World War I. During World War II Mr. Richard-
son served as a Second Lt. with the 10Ist Field Artillery in France.

For many years he was supervisor of the E & F Paint Co. in
Boston. He served on the Town of Concord Finance Committee and
was a member of the Concord Independent Battery, the Concord
Players, and the New England Botanical Club.

A well known authority on Concord history, he was the author of
Concord River (1964) and The Concord Independent Battery (1973)
and also other historical papers including a notable paper on the
history of Concord Junction.

HESLER VISITING PROFESSORSHIPS
OF FLORISTIC BOTANY

Under the terms of an endowment from Dr. and Mrs. L. R.
Hesler, the Botany Department of the University of Tennessee is
able to support a limited number of visiting professors of floristic
botany. Stipends will be awarded based on research proposals, and
may be used for salary and expenses of floristic, revisionary, or
monographic systematic research. All applicants must hold faculty
(or equivalent) status at a recognized botanical or educational
institution. Stipend amounts and residence periods will vary, and
concurrent support from the applicant’s institution is strongly
encouraged. Preliminary correspondence with specific departmental
faculty is invited or general inquiries may be sent to:

Hesler Endowment Fund
Department of Botany

The University of Tennessee
Knoxville, TN 37996-1100 USA

THE 1985 JESSE M. GREENMAN AWARD

The Greenman Award, a cash prize of $250, is presented each
year by the Missouri Botanical Garden. It recognizes the paper
judged best in vascular plant or bryophyte systematics based on a
doctoral dissertation that was published during the previous year.
Papers published during /985 are now being considered for the 18th
annual award, which will be presented in the summer of 1986.
Reprints of such papers should be sent to:

Greenman Award Committee
Department of Botany

Missouri Botanical Garden

P.O. Box 299

St. Louis, MO 63166-0299, U.S.A.

In order to be considered for the 1986 award, reprints must be
received by | July 1986.

154

NEBC 1986 AWARD
FOR THE SUPPORT OF BOTANICAL RESEARCH
IN NEW ENGLAND, U.S.A.

The New England Botanical Club will again offer an award of
$1,000 in support of botanical research to be conducted in the New
England region during 1986. This award is being made to stimulate
and encourage botanical research on the New England flora and to
make possible visits to the New England region by those who would
not otherwise be able to do so. The award will be given to the
graduate student submitting the best research proposal dealing with
field studies in systematic botany and plant ecology, but proposals
for research in other areas of botany will also be considered. This
award is not limited to graduate students at New England institu-
tions, nor to members of the New England Botanical Club. Papers
based on this research must acknowledge the NEBC’s support, and
it is encouraged that they be submitted to Rhodora, the Club’s
journal, for possible publication—subject to standard review
processes.

Applicants should submit a proposal of no more than three dou-
ble spaced pages, a budget (the budget will not affect the amount of
the award), and their curriculum vitae. Two letters, one from the
student’s major professor, in support of the proposed research are
also required. Proposals and supporting letters should be sent
before 28 February 1986 to:

Awards Committee
The New England Botanical Club
22 Divinity Avenue
Cambridge, MA 02138.
The recipient of the award will be notified by 30 April 1986.

155

MEETING ANNOUNCEMENT AND CALL FOR ABSTRACTS

New England Graduate Student Botany Meeting

The first New England Graduate Student Botany Meeting will be
held in the auditorium (Rm 153) of the Torrey Life Science Build-
ing, Department of Ecology and Evolutionary Biology, The Univer-
sity of Connecticut, Storrs, CT on Saturday 22 March 1986.
Attendance is open to all. Paper presentations will be largely re-
stricted to graduate student research (completed or in progress)
representing all areas of botany (systematics, ecology, reproductive
biology, anatomy, physiology, etc.). Time slots for paper presenta-
tions are limited and prior registration is required. Abstracts are due
by 15 Feb. 1986.

For additional information and abstract forms contact:

C. Thomas Philbrick or Debra A. Dunlop

Department of Ecology and Department of Botany and
Evolutionary Biology, U-43 Plant Pathology

The University of Connecticut The University of

Storrs, CT 06268 New Hampshire

(203) 486-4156, 486-4322 Durham, NH 03824

(603) 862-3531

Vol. 87, No. 852, including pages 449-616, was issued October 28, 1985

156

INSTRUCTIONS TO CONTRIBUTORS TO RHODORA

Submission of a manuscript implies it is not being considered for
publication simultaneously elsewhere, either in whole or in part.

Manuscripts should be submitted in triplicate (an original and
two xerographic copies) and must be double-spaced (at least 3/8”)
throughout including tables, figure legends, and literature citations.
Please do not use corrasable bond. The list of legends for figures
and maps should be provided on a separate page. Footnotes should
be used sparingly. Do not indicate the style of type through the use
of capitals or underscoring, particularly in the citation of specimens.
Names of genera and species may be underlined to indicate italics in
discussions. Specimen citations should be selected critically, espe-
cially for common species of broad distribution. Systematic revi-
sions and similar papers should be prepared in the format of “A
Monograph of the Genus Malvastrum”, S. R. Hill, Rhodora 84:
1-83, 159-264, 317-409, 1982, particularly with reference to inden-
tation of keys and synonyms. Designation of a new taxon should
carry a Latin diagnosis (rather than a full Latin description), which
sets forth succinctly just how the new taxon is distinguished from its
congeners. Papers of a floristic nature should follow, as far as possi-
ble, the format of “Annotated list of the ferns and fern allies of
Arkansas”, W. Carl Taylor and Delzie Demaree, Rhodora 81:
503-548, 1979. For bibliographic citations, refer to the Botanico-
Periodicum-Huntianum (B-P-H, 1968), which provides standard-
ized abbreviations for journals originating before 1966. All abbrevi-
ations in the text should be followed by a period, except those for
standard units of measure and direction (compass points). For
standard abbreviations and for guidance in other matters of biologi-
cal writing style, consult the CBE Style Manual, Sth ed. (original
title: Style Manual for Biological Journals). In preparing figures
(maps, charts, drawings, photos, etc.) please remember that the
printed plate will be 4 X 6 inches; be sure that your illustrations are
proportioned to reduce correctly, and indicate by blue pencil the
intended limits of the figures. (Some “turn-page” figures with brief
legends will be 3 1/2 X 6 in.) Magnification/ reduction values given
in text or figure legends should be calculated to reflect the actual
printed size. An Abstract and a list of Key Words should be sup-
plied at the beginning of each paper submitted, except for a very
short article or note.

RHODORA January 1986 Vol. 88, No. 853

CONTENTS

A synopsis of New Hampshire seaweeds
Arthur C. Mathieson and Edward J. Hehre
Cytology of Carex purpurifera Mack. (Cyperaceae)
James R. Manhart : . ; ;
BOOK REVIEWS
Bogs of the Northeast
Richard Evans Schultes : } ; St ae
The Botany and Natural History of Panama: La Botanica e Historia
Natural de Panama
Norton H. Nickerson
IN MEMORIAM
Bernard Boivin 3
Laurence E. Richardson
ANNOUNCEMENTS
Hesler Visiting Professorships
Jesse M. Greenman Award
NEBC 1986 Award Notice ; : ;
New England Graduate Student Botany Meeting

149

150

152
153

154
154
155
156

Instructions to Contributors : : : : : ; . inside back cover

‘Hovora

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

Vol. 88 April 1986 No. 854

Che Nef Lngland Botanical Club, Inc.

22 Divinity Avenue, Cambridge, Massachusetts 02138

Conducted and published for the Club, by
NORTON H. NICKERSON, Editor-in-Chief

Associate Editors

DAVID S. BARRINGTON RICHARD A. FRALICK
A. LINN BOGLE GERALD J. GASTONY
DAVID E. BOUFFORD C. BARRE HELLQUIST
WILLIAM D. COUNTRYMAN MICHAEL W. LEFOR
GARRETT E. CROW ROBERT T. WILCE

RHODORA.—Published four times a year, in January, April, July, and
October. A quarterly journal of botany, devoted primarily to the flora of
North America. Price $20.00 per year, net, postpaid, in funds payable at
par in the United States currency at Boston. Some back volumes and
single copies are available. Information and prices will be furnished
upon request. Subscriptions and orders for back issues (making all rem-
ittances payable to RHODORA) should be sent to RHODORA, 22
Divinity Avenue, Cambridge, Mass. 02138. In order to receive the next
number of RHODORA, changes of address must be received prior to
the first day of January, April, July or October.

Scientific papers and notes relating to the plants of North America
and floristically related areas, and articles concerned with systematic
botany and cytotaxonomy in their broader implications will be consi-
dered. Brevity is urged; please conform to the style of recent issues of the
journal. See “Instructions to Contributors to RHODORA” at the end of
each issue. Extracted reprints, if ordered in advance, will be furnished at
cost. RHODORA assesses modest page charges.

Address manuscripts and proofs to:
Joan Y. Nickerson
Managing Editor, RHODORA
Phippen-LaCroix Herbarium, Dept. of Biology
Tufts University
Medford, Mass. 02155

Second Class Postage Paid at Boston, Mass.

PRINTED BY
THE LEXINGTON PRESS, INC.
LEXINGTON, MASSACHUSETTS

Cover Illustration

An original drawing, seemingly the only one surviving and perhaps the only one
ever done for publication by Merritt Lyndon Fernald, used in part to illustrate his
article on cranberry species which appeared in RHODORA No. 48 (Fernald, M. L.
1902. The variations and distribution of American cranberries. Rhodora 4: 231-237
& Plate 40). The drawing was rescued from a wastebasket by Dr. Bernice Schubert; it
now hangs in the office of Dr. Carroll E. Wood at A. The original Plate 40 caption
reads as follows: Fig. 1, Vaccinium Vitis-Idaea; fig. 2, V. Vitis-Idaea, var. minor; fig.
3, V. Oxycoccus; fig. 4, V. Oxycoccus var. intermedium; fig. 5, V. macrocarpon.

Mbodora

(ISSN 0035 4902)
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

Vol. 88 April 1986 No. 854

THE NEW ENGLAND BOTANICAL CLUB
800TH MEETING

The Eight Hundredth meeting of the New England Botanical
Club was held on May 4, 1984 following a dinner at the Harvard
Faculty Club in Cambridge, Mass. This occasion was celebrated
with a special program entitled “A Tribute to Merritt Lyndon Fer-
nald” in which comments on Professor Fernald’s botanical accom-
plishments, teaching, humor and philosophy were made for the
benefit of the Club’s membership by those of his students and col-
leagues who knew him and were in attendance. Additional com-
ments were received in letters from many others.

The remarks delivered and selected sections of letters received are
here published so that all readers of Rhodora may share some of the
unique botanical history in which Prof. Fernald was so prominent.

Mary WALKER: I would like to open the 800th meeting of the
New England Botanical Club. This is the 573rd meeting and also the
800th meeting; Dr. Howard will explain that anomaly later. In view
of the nature of this meeing, we have decided to dispense with most
of the formalities. | hope people have been signing the guest book;
please continue to sign it during the evening. Instead of introducing
guests, | would like to introduce all the past presidents of NEBC
who are here tonight. We are honored to have nine of them. I will
ask them all to stand as their names are called, and you can save
your applause until they are all standing: Peter Ashton (I’m going
backwards), Lynn Bogle, Alice Tryon, Gordon De Wolf, Bill Coun-
tryman, Richard Schultes, Reed Rollins, Dick Howard, and Ralph
Wetmore. | hope that I haven’t missed any past presidents. Thank
you. Welcome!

ioe

158 Rhodora [Vol. 88

Before I turn the floor over to our program chairman, Garrett
Crow, I should say that this meeting, of course, is in honor of
Merritt Lyndon Fernald. (I’m sure there are as many pronuncia-
tions of Fernald as there are of Thoreau. I come from Concord so if
I’m not pronouncing it correctly, | will learn.) Since my student
days, I’ve been familiar with Gray's Manual, starting out with the
7th edition; when the 8th edition came out, | realized what a monu-
mental job it must have been for someone to produce that new
work, much enlarged from the 7th edition. While putting together
the poster exhibit for tonight, | came across a paper by Fernald
written in 1939 entitled, “How Soon Will the Manual be Done?”
(laughter) and asking botanists for time and patience.

While studying White Mountain botany as an Appalachian
Mountain Club Hut Naturalist, | found Fernald’s paper on soil
preferences of certain alpine and sub-alpine plants, written in 1907,
still to be a very useful document. In browsing through early and
middle volumes of Rhodora, I’ve always been impressed with the
number of articles by Fernald that were written during those
years—from Volume | to Volume 53—not only by the number of
articles, but more importantly, by the easy and fluent style in which
they were written. Wherever he could, he injected human interest.
That quality is badly lacking, | think, in botanical literature these
days. When I offered to do the exhibit for this program, it was with
a double purpose: partly to contribute just a little to the mighty
efforts of Dr. Garrett Crow and Dr. Richard Howard in putting the
program together, and partly to learn more about this unique man.
In the course of the evening, I hope to learn still more. I leave you
with one “wonder”: | wonder who, among the younger people
amongst us, will perhaps be honored by a meeting such as this; who
will be revising Gray's Manual, who will perhaps be honored at the
900th or the 1000th meeting of the Club?

I'd like to turn the floor over now to our Program Chairman, Dr.
Garrett Crow.

GARRETT CROW: Thank you. We’re delighted for the wonderful
turn-out here. It’s been an exciting meeting for me, so far, to get to
talk to so many people, and I hope it has been the same for you. I'd
like to comment about the menu. Dr. David Barrington designed it;
unfortunately, he was not able to come tonight. He’s grading exams,
which is a terrible excuse to stay away. I don’t think I'd let that

1986] NEBC 800th meeting — Remarks 159

happen to me! He must be dedicated, in some way, to that group.
But we have him to thank for the menu that’s been prepared.

I do want to mention that the June meeting will be at the Arnold
Arboretum and will begin with a tour of the arboretum in the after-
noon. Watch for the mailing on that as to the details and then the
evening program will be a talk by Peter Ashton on the Arboretum.
But we don’t want to take up any more time for such things. ..and I
want to introduce Dr. Richard Howard who will be our moderator
on this program, “A Tribute to Merritt Lyndon Fernald.”

RICHARD A. HowarRD: Thank you very much, Garrett, Madame
President. First of all, let me say that the 473rd meeting of the New
England Botanical Club, which was the 700th meeting since its
founding, was held in this very room. The head table was down at
that end; I was the speaker, and I was asked to talk about the history
of the Club. This speech was eventually published in Rhodora, and |
still have a few reprints if any of you haven’t read it (laughter); that
was in 1973. This is the 573rd meeting, as our President said, being
the 800th since the original establishment of the Club. The explana-
tion for this binomial system is the fact that after 227 meetings, in
1921, the Club found that it was going to be subject to taxation by
the Commonwealth of Massachusetts unless it incorporated. Hence,
we started the numbering system all over again. At this rate, the
900th meeting will be held in June, 1995; I would like the program
chairman to take special note of the fact that I shall probably not be
available as a speaker (laughter). The chances are that my tomb-
stone will contain that well-known epitaph that I’ve “gone to
another meeting” (laughter). Ten or eleven years ago, at the 700th
meeting, the senior members present were Ralph Bean, who joined
the Club in 1909; George Safford Torrey from the University of
Connecticut, who joined in 1912; and Donald White, 1913. All three
are gone. Henry Svenson was the member who joined in 1919. He
cannot be with us tonight but he sends his regards. Cap Weston,
1921; Ralph Wetmore, 1927, was at the head table; Cap is gone,
Ralph is here. Hugh Raup, 1929, was at the head table; he cannot
join us. Well, tonight our senior member is Harold St. John from
Hawaii, who joined the Club on December 1, 1911. This was at the
same meeting at which Fernald was elected President of the Club.

Now I'd like to set up some categories because | think Harold is
the only member of the 70-year club of membership in the New

160 Rhodora [Vol. 88

England Botanical Club. You'll hear from him in a few minutes.
Those who have been members for over 60 years, who have been
invited to attend and could not, but who sent their greetings, are
Frank Seymour who joined on December 4, 1914, the last meeting
in which Fernald was President of the Club; Henry Svenson, 1919;
James Poole, 1920; and Lyman Smith, in 1923.

Then we can set up a 50-year category—those who have belonged
to the Club for 50 years or more: G. Ledyard Stebbins, who we
thought was going to come, joined in 1926, as did George Church, in
1926; Ralph Wetmore in 1927; Hugh Raup in 1929; Ernst Abbe and
Dick Goodwin, 1930; Abbe is not here, Dick is. And while Gene
Ogden, who is here, joined the Club in 1934, we’re not considering
him a member of the 50-year club, because he did not get elected
until November of 1934! (laughter).

The history of the Club is very close to us. In December of 1895,
William Farlow and B. L. Robinson assembled a group of friends in
Farlow’s house and proposed the establishment of a botanical club.
Farlow’s house was right next door—the location of the Carpenter
Center, the Le Corbusier building that is right next to the Faculty
Club. After two meetings, they decided they would indeed have a
club, and the New England Botanical Club was officially christened
on February 5, 1896. They gathered at various people’s houses for
dinner, and they had to eat, so there were four meetings in March,
1896. At the fourth meeting Fernald became a member, and it may
be said that he dominated the club in his activities from that time
until his death. You’ll hear more about this as we go along.

I suggested for this program that we talk about Fernald, for those
of us who knew him personally are certainly becoming fewer. For
those of you who know him only as a name, perhaps some of the
things we say will tell you about this remarkable man and his per-
sonality. He certainly was the right man in the right place at the
right time. In preparing for this, we wrote to all of the old-timers
and asked that they either tape something of their recollections of
Fernald or write something out, give us a collection of stories or
episodes, and we invited them to be present tonight. About nine-
tenths of those we have contacted replied. Some of the stories were
very short; some of them are quite good stories. And of those who
are here, Ill ask about eight to talk for five or ten minutes. Then
we'll open the meeting to others to fill in the gaps or add to the

1986] NEBC 800th meeting — Remarks 161

reminiscences. Of these manuscripts that I have on hand, some of
them are rather significant. Hugh Raup has written a very nice
essay; | would like to see it published. Lyman Smith has added some
notes. Ernst Abbe defends the fact that he was the one who forced
Fernald to reconsider the Nunatak Hypothesis on the basis of his
work in Labrador. And Lily Perry, who knew B. L. Robinson and
saw the transition to Fernald’s administration, has given us some
notes. I would like to suggest that, along with the comments of this
evening, they be edited and considered for publication in Rhodora.
We'll leave that to someone else.

Well, let me set the stage for Fernald. Merritt Lyndon Fernald
was born in Orono, Maine, on the 5th of October in 1873. He was
the third child of M. C. Fernald, who at that time was professor of
mathematics and later became the first and the third president of the
Maine State College of Agriculture and Mechanical Arts, now the
University of Maine. At the same time, in 1873, Asa Gray was 63
years old, tired and overworked, as he said himself, in charge of an
herbarium, a botanical garden, a teaching program, and several
major projects on which he had to write. He had been president of
the American Academy of Arts and Sciences for ten years. He had a
worldwide reputation and the correspondence that went with it, and
he was involved in fund-raising! Even in those days, Harvard had
the edict, “Every tub on its own bottom” (laughter). The working
budget for the Gray Herbarium in 1873 was $800.

Charles Sprague Sargent had just been appointed Director of the
new Arnold Arboretum and had the additional duties of Professor
of Horticulture at the Bussey Institution in Jamaica Plain. He also
was the Director of Harvard’s botanical garden, Gray’s garden in
Cambridge, and in this he relieved Gray of part of his duties. Now
this was possible because Sargent had arranged it. He obtained from
his father, Ignatius Sargent, and from H. H. Hunnewell, a promise
of annual gifts of $500 each to provide Gray with a “pension” of
$1,000 a year, which was more than Gray’s salary, provided he
resign and devote himself to the Flora of North America. Gray had,
that year, made his last appointment of Sereno Watson as an assist-
ant in the Herbarium. The next year Sargent appointed Watson as
Curator in charge of the Herbarium. Gray accepted all this, Sargent
in charge of the garden and Watson in charge of the herbarium, but
he found himself on the outside, fighting Sargent’s “audacious

162 Rhodora [Vol. 88

scheme” to move all botanical activities to Jamaica Plain. Gray died
in 1888, the first year that a teenager, M. L. Fernald from Orono,
Maine, sent specimens to the Gray Herbarium for identification. In
1889, Watson, who was then Curator, along with Coulter, issued the
sixth edition of Gray's Manual, which was significant work because
it expanded the range, the coverage, to the 100th meridian. In 1890,
Fernald wrote to Watson:

Orono, Maine, January 30, 1890, Dr. Watson: Dear Sir, I
have just been looking at Juncus, #16, which I sent to you in
the fall and am more convinced that it is not described in the
Manual. [This is the Manual that Watson had just finished!] I
sent you a specimen with more mature fruit than any I sent
before. It seems to be something like Juncus tenuis, var.
secundus, but the flowers are not restricted to one side of the
panicle.

Watson wrote back to Fernald (and listen to this, if you think
college recruiting is something new):

I have been much pleased with the intelligent interest you
have shown in the plants of your region. I have no idea what
your plans or expectations for the future may be, nor even of
your age or how far advanced you may be in your education;
but, as a venture, I would like to say that if a career as a
botanist has attractions for you (laughter), there is an oppor-
tunity open here for a young man who is willing to begin at the
bottom and work his way upward (laughter). I am desirous of
securing the services of some intelligent, capable, quick-witted
young fellow to do work in the Herbarium that must be done.
The work requires and gives familiarity with the Herbarium
and the floras of the world in the way of caring for and distrib-
uting into herbaria the various collections that come in from
many quarters. Opportunity will be given for study and ad-
vancement and compensation sufficient at least for support. If
such a position has any attractions to you, I should be glad to
hear from you.

Fernald wrote back immediately:

The position you speak of does have attractions for me. I
think that one thing I was made for was a botanist, as from
early childhood my inclinations have been in that line. [The
punchline comes.] I dare not say whether I would like the

1986] NEBC 800th meeting — Remarks 163

position or not. We are giving it careful consideration, and my
father will doubtless write to you in a few days concerning it. I
am the third child of Dr. M. C. Fernald, aged 17, and am in
the freshman class at State College, located here, of which my
father is the president. I know very little about “textbook
botany,” my work having been mostly in the field, and should
I go to Cambridge, I should want to study as much as possible
and do the required work satisfactorily. Respectfully yours,
Merritt L. Fernald.

Well, Papa Fernald came down to see Watson and they talked
about classical education for young Merritt. It turned out that Wat-
son was able to arrange for Fernald to work half-time in the herba-
rium and, as a special student, take two courses at Harvard. Fernald
arrived in the end of July of 1891, entered the Lawrence Scientific
School and began an association that ended with his death in 1950,
nearly 60 years later.

In this transition period, Fernald was publishing, believe it or
not. He published notes on two carices of Maine in 1890. He pub-
lished a list of special plants collected in the vicinity of Portland in
1891 that went through two new editions later on.

Fernald came here in the summer of 1891, and Watson died in
1892. B. L. Robinson, who had been Watson’s assistant, was
appointed curator in 1892 to follow Watson. So here we have
Robinson, aged 26, with his assistant, this new student Fernald,
aged 18, to work under him. The institutional projects of the Gray
Herbarium at this period were supposedly four: revision of Gray's
Manual, which was then the work of Robinson and Fernald, Robin-
son having the experience, Fernald having the experience in Maine;
the continuation of the synoptical flora of North America which
Gray had started, which was Robinson’s responsibility; work on the
vegetation of Mexico and Central America, in particular identifying
plants from that area, the responsibility of Robinson, Greenman
and Fernald. The collections were those of Millspaugh and Garner,
C. C. Deam, Peck, Donnell-Smith, Rose and Lumholtz. The fourth
project of the Gray Herbarium in those days was the indexing of
newly described systematic entities for the western hemisphere. This
is what we know today as the Gray Index. It was started down in
Washington by Miss Day, who brought it up here to Harvard when
Washington could no longer finance it; it has been one of the classic
projects of the Gray Herbarium ever since.

164 Rhodora [Vol. 88

Let me go chronologically here for just a bit and give you some of
the highlights of this career of Fernald. In 1894, Fernald was the
junior author of a paper with Robinson on the identification of the
Hartman and Lloyd collections from Mexico. Robinson submitted
this paper to the American Academy and it was reprinted, as was
the tradition in those days, in the Contributions of the Gray Herba-
rium. In 1896, the New England Botanical Club was formed. Fer-
nald joined at the fourth meeting. In 1897, Fernald published his
own paper on the plants of Mexico, including a revision of the
species of Pectis, and another paper which dealt with new additions
to the flora of Mexico. Later in 1897, Fernald got his degree, a B.S.
from the Lawrence Scientific School at Harvard, magna cum laude.
In 1899, Volume 1, No. 1, of Rhodora was published. Robinson was
the editor, Fernald was one of the assistant editors. The first volume
started off with an editorial announcement. The first paper was by
M. L. Fernald on the rattlesnake plantains of New England.

In 1901, Fernald started a project called Plantae Exsiccatae
Grayanae. Now for those of you who haven't seen these classic
labels, this was a scheme in which the botanist collected 125 speci-
mens, or at least 125 sheets, and a special label was prepared in
which the bibliographic details were complete. The plant was identi-
fied, all of the literature pertinent to it was cited, and down at the
bottom it was noted where it was collected, and so on. These 125
were made up into centuries, and the centuries were distributed. The
extra 25 over the 100 sets were to take care of those who had
collected the centuries and wanted sets for themselves; so 100 sets
were distributed. Over a period of time, 1500 sets were sent out
between 1901 when Fernald started it and 1952, which I think is the
last one, so for 50 years this project continued. Alone, or with
Robinson, Fernald collected the first 54 sets, and you may be inter-
ested to know that the first numbered set was Veronica serpyllifolia
var. borealis. It was collected in Fort Fairfield, Maine, on June 6,
1901. Now, if any of you know where Fort Fairfield, Maine, is
located, you're better than | am because I had to go look it up. It’s
on the New Brunswick border in the very northeast corner of
Maine; in those days, it was the end of the railroad. Fernald was
exploring in 1901.

The second one that was collected has a very familiar name. It
was collected in Van Buren, Maine, on August 13, 1901. It was a

1986] NEBC 800th meeting — Remarks 165

species published by Sereno Watson and was called Pedicularis fur-
bisheae (laughter). And if you wonder why the plant became an
endangered species, perhaps it was that Fernald, by himself, went up
to Van Buren, Maine, in the type locality and made up 125 sheets of
the thing and it hasn’t been seen there since! (laughter).

In 1902, Fernald was appointed an instructor in Harvard College
and he started to teach. In 1905 he was promoted to an assistant
professor. In 1907 he was married; eventually they had two childern.
In 1908, he collaborated with B. L. Robinson in the production of
the 7th edition of Gray's Manual. Now the 7th edition of Gray's
Manualis probably the poorest of the whole batch. It was regarded
as a patch-up job. In fact, in a discussion later on, Henry Bartlett
said that this was simply a cut-and-paste job, and few annotations
were made on it. Fernald was not very proud of the whole thing, but
it was Robinson’s work —Robinson and Fernald. Fernald, in several
letters, says that he is going to start to work immediately and do a
good job on the 8th edition.

In 1911, Fernald was elected president of the New England Botan-
ical Club, as you see on your list here tonight. He continued in that
capacity until 1915, when he was appointed Fisher Professor of
Natural History. Robinson finally resigned in 1935 and Fernald was
appointed Curator of the Gray Herbarium, serving two years; in
1937, he became the first Director of the Gray Herbarium, a title
which he retained until 1947, when he retired. In 1950, Gray's Man-
wal was published. Fernald died that same year on September 22,
1950.

In the February, 1951 issue of Rhodora, there are tributes to
Fernald. There’s a biographical sketch by Arthur Stanley Pease.
There are sections entitled “Fernald as a Teacher” by Jack Fogg,
who has since died; “Fernald as the Reviser of Grav's Manual,” by
Henry Bartlett, who talks about the problems of the 7th edition;
“Fernald as a Botanist,” by Reed Rollins; and “Fernald in the
Field,” by Ludlow Griscom, who was a zoologist.

I knew Fernald in various ways from the time | came to Harvard
in 1938 until his death in 1950. I was a student of Ivan Johnston,
had been working with I. W. Bailey in the Biological Laboratories,
and wanted to do some taxonomic work associated with the anat-
omy | had been doing, which meant using the Gray Herbarium.
Only Fernald’s or Weatherby’s students used the Gray Herbarium

166 Rhodora [Vol. 88

as office space, so I created one of the first political crises of my stay
around here by saying, with the support of the administration, that I
needed table space in the Gray Herbarium. | was assigned table
space in what was known as the Radcliffe Laboratory. For those of
you who don’t remember it, there was segregation in those days. At
the foot of the stairs, going into the wing where the laboratories were,
there was a little lab off to the left with a door on it that was usually
closed. This was the Radcliffe Lab, and I was given space in there.
At the end, looking out over the garden, was the space where the
Harvard men had their quarters. This was meant to shame me, but
my companions in those days were Bernice Schubert, Shirley Gale
and Shiu-Ying Hu, and it wasn’t so bad at all; Fernald did come in
for lunch and eventually I got to know a little bit about him.

You look back on it and you say, “Fernald as a teacher.” In these
contributions that we have had sent in to us, the name “Botany 7,
The Flora of New England and the Maritime Provinces of Canada,”
is mentioned often. Botany 10 was “The Classification and Distribu-
tion of Flowering Plants: Advanced Studies on Special Topics,” and
here Fernald roamed. He could do anything he wanted. Then there
was a course-—and I haven’t been able to find the name or the
number of that—which we called “the drawing course.” This was
divided into two semesters: one on the monocots, one on the dicots.
Fred Taylor brought his bound drawings, and they were on display
earlier tonight. The work of that course and all it was, was that
Fernald pulled out specimens of various groups and put them down
and said, “Draw them.” It is also said that from these art works
Fernald selected the artists for his later contributions and
publications.

“Fernald as a collector?” —I apologize; I was not able to find his
field books to find out what his total numbers were. They are signif-
icant. He spent his summers collecting; | am sure you will hear
about the field trips he took with students to Cape Cod and else-
where. He roamed from Nova Scotia and the Gaspé to Virginia.
Frank Seymour wrote to me that the students used to call certain
lectures, when Fernald started talking, as the “gas-bay” lectures, as
Fernald was so windy about it. He started Plantae Exsiccatae
Grayanae, as I’ve indicated; there were 1500 numbers of that. The
majority of them were collected by Fernald; 1457 was the number of
Proserpinaca palustris, the last one that he collected. Of the last

1986] NEBC 800th meeting — Remarks 167

centuries, incidentally, ten were collected by R. A. Howard and
students in Cuba. And while it seemed a little strange to be adding
Cuban plants to this, we found that one of the most popular ones
that we sent out was Rhizophora mangle. It had the greatest fan
mail because this common mangrove was not well represented in
herbaria, and we had included the specimens of flowers and fruit
and photographs. In the photographs, I followed the lead of Walter
Hodge who had been collecting in Dominica exsiccatae sets for the
Gray Herbarium, and his photography was better than mine. You'll
still see some of these Charianthus and Melastomes under the Plan-
tae Grayanae. Regrettably, it stopped; it was a good series.

“Fernald as a scholar” It is said that he published 750 papers; I
don’t think I’ve read them all. Gray's Manual—a monument to
anyone—is the last of the big manuals written by one individual. His
work on the persistence of plants in unglaciated areas is outstanding
—the Nunatak Hypothesis that Abbe challenged. The monograph
of the linear-leaved species of Potamogeton is still cited as the out-
standing model monograph of all time. The plants of the “wine-
land,” with its discussion as to where the Vikings landed, is here.
The edible plants of eastern North America is another: whether it
was Kinsey’s manuscript or Fernald’s is still a point of debate.

“Fernald as an editor.” Well, he was in Volume I, No. 1, of
Rhodora. In 1929, in Volume 31, he became the editor and con-
tinued until his death. The October issue of 1950 is the last to bear
his name. The November (1950) issue has the last paper authored by
Fernald in Rhodora. Its title is, “Why So Many Careless Books on
Trees and Shrubs and Other Plants?” (laughter). This was character-
istic of Fernald’s reviews or his criticisms; he didn’t have a blunt
pen, I'll tell you that.

The last paper bearing Fernald’s name was in the January issue of
1951 in Rhodora and is “Botanizing on the Gaspé Peninsula,
1902-1904.” There is an editorial note that “until the day of his
death, Professor Fernald had been working on a journal account of
the field trips to the Gaspé, project to cover the years 1902-1934.”
He had sufficiently completed the years 02 to 04 to make this
portion of the manuscript publishable. We wish we could have had
the rest of it, because it’s good.

He edited the Contributions of the Gray Herbarium, and while
these were reprints of papers published elsewhere, with early ones

168 Rhodora [Vol. 88

from the Memoirs of the American Academy and the later ones
from Rhodora, they were significant in the distribution of publica-
tions that led to exchanges with the Gray Herbarium. An early
editorial note by Miss Day warns that these papers are to be cited on
their original dates and pagination and not by what is in the Contri-
butions. The first original article was Contributions No. CLXV,
which was a miscellany in honor of Merritt Lyndon Fernald at the
time of his retirement. Fernald’s last contribution to the Contribu-
tions was in 1949, but I think Carroll Wood has the honor of being
the last author reprinted in Contributions for his work on Tephro-
sia, which was in Rhodora.

“Fernald and the New England Botanical Club”— Well, he was
president, he was Curator of the Herbarium, he was on the council,
he was the Editor. | think my fondest recollections are of the contin-
uous competition he had with E. D. Merrill. Merrill came to Har-
vard as the Coordinator of Botanical Collections—Supervisor of
Botanical Collections and, in theory, was Fernald’s boss. They were
both short people. They were both very important people, and they
knew it. And they were both from Maine—and that’s equally impor-
tant! It is very clear in my recollections that Fernald never stood up
to say anything but what Merrill popped up afterwards to say some-
thing. And if Merrill started first, Fernald stood up and said his
piece, too. One of the minutes in 1911 probably summarizes it very
nicely: “Mr. Fernald filled in the remainder of the time by giving a
very interesting account of his excursion on October 12 to Wallum
Pond on the Rhode Island line in search of Sc/lerolepis. He secured
fine flowering specimens in both Rhode Island and in Massachu-
setts, thus adding this new plant to our state flora.” If you read
through that, the plant was known from Rhode Island, but he found
a pond that was on the line; when he got around the pond, it was in
Massachusetts, so it was an addition to the state flora.

Well, | have kidded a bit here about Fernald, and yet I have deep
regard for what he has done, and I know the others have, too. | would
like to ask other people to come up here and talk for five or ten
minutes. I pointed out to you that Harold St. John was elected at
the December | meeting in 1911, when the annual meetings were
held at 3 Joy Street, which I think is where the private environmen-
tal organizations rent their offices from the Appalachian Mountain
Club next door. Meetings have been held at various places; in fact,

1986] NEBC 800th meeting — Remarks 169

my first recollection of the Club was on Newbury Street when we
met in the quarters of the American Academy. President Deane was
in the Chair, and Merritt Lyndon Fernald was elected President for
the next year at this meeting at which Harold became a member.
There were 33 members present; the speaker was Mr. Deane. His
title was “The Mystery of the Maple Sap Flow,” and he concluded
that nothing positive is known as to why sap rises. Emile Williams
was the Recording Secretary, and there was a Treasurer’s Report. |
thought you might be interested that the Balance on hand at this
meeting on December |, 1911; it was $151.51. The income during
the year was $722.98; incidentally, the expenses were $799.83, which
meant that at the end of the year the Club had a balance of $74.60.
As to the income in that year, $645 was from dues. Now keep in
mind that these were dinner meetings, so everybody ate. Later on,
we went down to ice cream and donuts and ginger ale, and now
we're down to cheese and beer and so on. Things have gone down-
hill badly in the years, but costs have gone up, too. Dues produced
income of $645, and there was an entrance fee, which that year was
60c. The interest on the bank deposit was $17.53, and Mr. Treas-
urer, please note: Income from the sale of empty beer bottles was
4S¢!

Now I would like to ask Dr. Harold St. John, our senior member,
to give us afew comments. Harold.

HAROLD ST. JOHN: Thank you, Dick. I’m very glad to be present
here because the New England Botanical Club played a very impor-
tant part in my development as a botanist. At meetings of the Club,
I met numerous, dedicated, interesting, fine botanists, so I’m very
glad to be present at this meeting. Now, I can’t help being here
because I was born a botanist. I was collecting plants—I was well
along—at the age of six, and I’ve collected plants ever since.

My schooling was in the public schools of Brookline, Massachu-
setts, mostly, and my schooling was entirely in the classical trend, in
the classical program. I never had a course in botany until I came to
Harvard College. The first course I had to take was Botany 1, the
elementary course, and it was the most boring, stupid course I ever
had (laughter). It was taught by a physiologist (laughter), and he
spent most of his time in the lectures trying to imitate life; and as far
as I’m concerned, he never did! (laughter). The next year I was able

170 Rhodora [Vol. 88

to take Fernald’s course on systematic botany, and what a revela-
tion that was.

Fernald also was a born botanist. He loved his subject; he had
broad knowledge of it. He wrote beautiful English, he spoke elo-
quently, and his enthusiasm for his subject just bubbled out of him.
You couldn't help feeling the enthusiasm which he had for his
subject.

I became his teaching assistant, I think the next year. If not, the
second year after that. I handled the lab sections of his courses in
Harvard and in Radcliffe, carefully separated. After Fernald lec-
tured to Harvard, the next hour, normally, he gave the same lecture
to Radcliffe in the next room. “Why can’t I talk to the two of them?
Why can’t I have a single audience?” That was one complaint that
never ceased.

I think it was in my junior year that Fernald invited me to be his
field assistant on a trip to eastern Canada. On that same trip, he had
as one companion E. B. Bartram of Philadelphia, who at that time
was studying the flowering plants. Bartram was a big, rangy, husky,
outgoing friendly sort of a person. Not long before, he had come
back from the Spanish War, and he thought there was no use in
going to college. He was already a man, his own. Well, he got into
work in Philadelphia in the Moore-Field Company, which made
steel wire. And he proceeded to move up in the ranks into adminis-
tration. Apparently he was very well remunerated and, by the time
he got to be forty, every time you’d meet him he’d say, “Well, I’m
going to retire; I’m going into botany; I’ve made all the money I
could ever use; I’m going to retire.” But he took another ten years
for it to happen. The other companion was Bayard Long, also of
Philadelphia, who was a very keen, lean, nervous, rather irritable,
easily offended person. So we had a mixture of personalities there.
But Fernald, in the field, was a different person from his characteris-
tics that he showed in the laboratory. He was friendly and jolly and
cheerful, and no longer was he criticizing your English or pointing
out what you concluded in a paper you handed to him, that you
hadn’t proved what you concluded. In the field, he was jolly and
nice and friendly; he seemed like a different man. We spent the
summer on Prince Edward Island and in Nova Scotia. And the next
summer, the same quartet worked on the Magdalen Islands in the
Gulf of St. Lawrence and in western Newfoundland. Then the next

1986] NEBC 800th meeting — Remarks 17]

summer, Fernald raised the money, | think all of my expenses for
one summer was a hundred dollars. He sent me out to Sable Island,
off the coast of Nova Scotia, 200 miles out to sea, east of Halifax, a
20-mile sand-dune island, which is along the stretch of sandy coastal
country that runs from Virginia, southern New Jersey, Long Island,
Cape Cod, and out to sea finally to Newfoundland, the Avalon
Peninsula. That was a very interesting expedition. Two years after
that, I went with Dr. C. W. Townsend and cruised in a 40-foot
schooner, without auxiliary, 600 miles along the coast of southern
Labrador. Those opportunities were made possible to me by
Fernald.

I am wandering along from one subject to another, not knowing
how to prepare a talk like this because I figured I was following
Dick Howard and he would hit all the high spots, and I'd try to fill
in a few of the low spots.

Every five or seven years, | used to come back to the Gray Herba-
rium from my positions in the west or in the Pacific Ocean, and
always I enjoyed seeing Fernald. I remember, toward the end there
was a second-story balcony inside of the Gray Herbarium in the
room with most of the cases. At that time he had eyeglasses, oh, 8 or
10 mm thick. He couldn’t see you across the room, but one day he
was up on that balcony and looked down—“St. John, you're gettin’
bald!” (laughter). He also could see things closer to; that is, he had
these glasses on and a lens and the sheet with the plant specimen
was, at most, 10 cm from his eye. Nobody else could see any differ-
ence, but he could see a new species at that distance (laughter).

Well, [ thank you for you attention. (Applause).

R. A. Howarpb: May we all be as vigorous when we’ve been in
the Club for 70 years, too (applause). James Poole, who joined the
Club in 1920, felt that he might be able to make the trip down from
Hanover, but finally it was decided that he should not make the trip.
However, we have an interesting situation here because although he
was one of Fernald’s students, so also was was his daughter. So we
have two generations represented in Margery Poole Taylor, who is
representing not only herself but her father, Jim Poole, of the NEBC
class of 1920. Marge.

MARGERY POOLE TAYLOR: Jim Poole is sitting in Hanover
tonight, at 95, frustrated utterly that he isn’t here, I'll tell you that!

172 Rhodora [Vol. 88

He would just love to be with you, and he has given me a paper to
read to you, but he’d much rather do it himself. He writes:

I’m most grateful for the invitation to attend this 800th
meeting of the New England Botanical Club. I am particularly
happy to have the opportunity to honor Professor Fernald. |
had the privilege of attending his course in Taxonomic Botany
in 1915-16. In my estimation, it was outstanding among all my
graduate courses. His lectures were interesting, and occasion-
ally delivered by bits of MFL wit or humor. His field trips,
several of them, to the lake shores of Cape Cod were unforget-
table experiences for those in his class. I recall one instance
illustrating his keen faculty for recognition of differences in
characters of taxonomic importance. At that time, my father
owned an Oldsmobile touring car, equipped with jumper seats
in the tonneau. I borrowed it for one weekend. Its capacity
made it possible to take Professor Fernald and the class for a
trip to the Housatonic Valley with occasional visits to areas of
botanic interest along the way. As I recall the event that | am
referring to, we were bowling along at 35 miles an hour when
Fernald, in the front seat, said, “Whoa, whoa!” I stopped as
quickly as the brakes permitted, Fernald jumped out and ran
back a few yards to a grassy area beside the highway. He soon
returned with a specimen he had spotted as a different variety
of Bluegrass, probably a variety of Poa pratensis, and this he
had recognized from a car, at that high velocity! (laughter).
We were in the town of Charlemont on old Highway 2,
between Greenfield and North Adams, in either April or May
of 1916. It would be interesting to discover that particular
specimen in the Gray Herbarium (and Carroll Wood is going
to look into it). lam very sorry and disappointed that | am not
with you and have to remain in Hanover. Hearty greetings to
any of my old friends who may still remember me.

Dad is 95 this weekend and | do have a birthday card for those of
you who knew him; please come up and sign it, if | haven’t caught
up with you yet.

Now, about myself. I took his course, I think it was in °36. When I
got to Harvard, I discovered I had some “uncles.” One of them was
Ralph Wetmore. He had been a grad student with my Dad. And
Fernald was another one. And they were a little bit amused, I think,

1986] NEBC 800th meeting — Remarks 173

at this young “chick” coming in there trying to be a botanist. But |
think from the age of three, as Harold St. John said, I was imprinted
as a botanist. I just was one from the beginning. I have continued in
my field in many directions, but my days at Harvard with Fernald
were among the things I remembered particularly. I have some pic-
tures taken on a field trip of that time; and if any of you were on
field trips in °35-°36, I can’t identify anybody but Fernald and
myself, maybe you are in it—who knows? So come up and tell me. I
did include a magnifying glass for those of us who need it. Do come
up and talk to me if any of you know Dad. He is still at the Jesup
Herbarium as a curator. He goes up three or four days a week and
loves being up there. Thank you. (Applause).

R. A. Howarpb: Marge, the current Index Herbariorum lists
James Poole as the curator of the Jesup Herbarium in Hanover, as
you just mentioned: 95 years old, the only staff member ever listed
for the herbarium. Please give him our regards and best wishes on
his birthday. Sorry he couldn’t be here.

I chose for the next speaker a man who came to Harvard in °26
and joined the Club in ’27, and who today is one of the best-loved of
the senior staff members. Those of us who are much younger shared
his courses and his care and concern for students and have a very
high regard for Dr. Ralph Wetmore. Ralph.

RALPH WETMORE: Thank you. It’s certainly a privilege to be
here. There have been times when I thought I wouldn’t be, but I’m
very glad that I made it! Honest. I have been here since 1921 when I
came after my undergraduate training, for my graduate work; I hap-
pened to have chosen Fernald’s course the first year I was here. |
was down in the Museum; that’s where the Biological Laboratories
were at that time; the Biology Building was not built until the early
30s. But I confess that, so far as maintaining the botany was con-
cerned, Professor Fernald’s course—I’m still a rooter for him. I’m
very glad to speak here in his interest. | wish he could be here, too,
to hear some of the things that have been said. None of it was said,
that I know about, inimical in any way to his entire prestige, in
either his lectures, which were continuous, because they followed
the manuscript of the 7th edition of Gray’s Manual of Botany. But
as far as he was concerned, there was no question that he was loved
by all his students. His field trips were phenomenal. This has all

174 Rhodora [ Vol. 88

been said, but I can’t help but repeat. After I came here on the staff
in 1926, I must confess that for a period of years I went on every
field trip I could get to. One incident that I do want to refer to just a
few minutes, but particularly do I want to point out that from the
time he started on the long corridor down to the extension of the
Gray Herbarium used for his and others’ lectures, | must confess
that you knew it was Professor Fernald coming. Most people took
one step at a time, but he “clicked” on the floor twice with each step,
and I don’t know how he did it; I never did find out (laughter). It
was his walk—quite unusual—and it was always present.

His classes were taxonomy to be sure, and taxonomy of the
regions that he had covered in his lifetime, and botanized, but they
were different. There were so many incidents of his particular traits
on a field trip that were amusing and interesting that he always used
to spice his lectures, and believe me, they were spiced!

You've heard a number of things of his way of doing things, but I
want to mention one incident that took place, and think part of his
life would never have happened if it hadn’t been for the football
captain who was there that afternoon. He had been busy all the fall
captaining the football team, and winning, and late in the fall and
early the next spring, when there was no football, he used to go on
field trips, and in one particular one, we had gone up to southern
New Hampshire just at the time when the first touch of spring was
present. You could see the beginnings of green and we hoped to find
a few things in bloom and that we did. Professor Fernald kept close
to the edge of the water of the stream that we happened to be on at
that particular time. It was on the edge of a millpond, if that means
anything to you, the backing up of the water in the spring of the year
so that they could do whatever sawing they needed to do for the
summer before the brook or river dried up. In this particular
instance, he bent down close to the edge of the millpond which was
quite a number of feet deep. I have no idea; I didn’t test it, but in this
particular case, he got too close to the edge and disappeared!
(laughter).

We stood there—dumbfounded (laughter). Nobody did anything
—not a thing. Unfortunately for the rest of us, and to our shame, we
saw the football captain peeling off his clothes. He got the outer
clothes off and in he went. In just a few minutes, he bobbed up to
the top. It was clear that he was being held down by something; but

1986] NEBC 800th meeting — Remarks 73

we pulled him out to the shore and in his one hand he had hold of
Fernald’s hand. We got Fernald up on the shore and laid him out.
To our surprise, he had the digger in his hand. One end is a spade-
like thing that you can dig with, which he dug up roots of his plants
and the other end was broad and he could get the top part of the
plant out, with the entire root, whether it was an herb or had a root
close enough to the surface. On this particular instance, when he
got to the surface, when we fetched him out, he had the digger still in
his hand. He probably would never have told the details of what
happened at that particular episode; it was about mid-career, I sup-
pose, about 1927-28-—I don’t remember the dates, unfortunately
but one will always think of him, those of us that were present.
There may be one or two here who were there on that trip, I don’t
know, but I must confess, it was one of the outstanding achieve-
ments of the football captain (laughter), rather than any of the rest
of us who were somewhat ashamed, I'll admit. I’m not going to say
anything more about Professor Fernald. I’m on his side, rooting for
him. If he were still here, | would tell him so. Thank you very much.
(Applause).

R. A. Howarb: Thank you very much, Ralph. The point of
these two stories you've just heard is that Fernald did not drive a car
and depended on others for transportation. Perhaps some more
Fernaldian car stories will come out, but Ledyard Stebbins tells of
one graduate student who had a car that Fernald dubbed “Juncus
canadensis.” True to its name, the thing broke down and so Fernald
re-christened it “Juncus inutilis.” Incidentally, the other part of
Ralph’s story is that Fernald couldn’t swim, and yet he took groups
of students out in the field. This was my experience with him: on
going down to the Cape, we stopped at the first wet area that was
seen, Fernald led the party, and we crossed the wet area. He gener-
ally knew when it wasn’t over his head, so we didn’t repeat that
episode, but we got wet. For Fernald, it was up to his waist; for me,
it was up to my knees (laughter), but nevertheless we got wet. When
we got to the other side, he simply walked around the pond, we got
back into the cars and drove to where we were supposed to go. This
was Fernald’s philosophy: if you get your students wet immediately,
it doesn’t matter what they do during the rest of the afternoon, and
it operated very effectively.

176 Rhodora [Vol. 88

When Fernald retired in 1947, Lincoln Constance was asked to
come to Harvard and, as he says in part of his letter here, he was put
in charge, but a few weeks later he was “appointed Acting Director
since the Harvard bureaucracy would not honor my signature unless
| held a formal title. My principal responsibilities were actually only
two: to keep the Gray Herbarium running for the year, meanwhile
to assist in the selection and confirmation of Fernald’s replacement,
and to convince Fernald that he had indeed retired.” He talks about
Fernald’s replacement in this way: “I first visited Cambridge in 1942
just after Reed Rollins had accepted an appointment to the faculty
of Stanford University, and I asked Professor Fernald if he was not
as greatly pleased as I was. Fernald looked at me quizzically for a
moment and then snapped, “They’re pretty radical out there.’ So
much for Clausen, Keck and Hiesey and what Stafleu calls the ‘birth
of biosystematics’.”

Fernald’s successor has proven his merits; we’ve known him a
long time now, and he is retired. He joined the Club in 1937, so
we're getting closer in the generations here. Dr. Reed Rollins, incid-
entally, has made a tape which we can use as a manuscript later on
and we'd like to have him say something now. Reed.

ReEED C. ROLLINS: Thank you, Dick. I'll be fairly brief because
we're getting down now to the point where it’s “modern history,” at
least not quite so ancient. I suppose if | wanted to, as an administra-
tor following Fernald, I could spend a lot of time tearing him apart.
But I’m not going to do that because, first of all, there’s no point in
it and secondly, I don’t think he would deserve to be torn apart.
There is so much on the positive side that this more than counter-
balances the minor things that I ran into when I came to take over the
Gray Herbarium as Director. Fernald was not an administrator and
I think those of you who knew him would admit to that imme-
diately. He was a botanist; in fact, he was a botanist’s botanist. He
lived and breathed botany and he wasn’t really interested in any-
thing but that, so he had difficulty dealing with the bureaucracy that
then was minor compared to now in Harvard University (laughter).
So I must say that even Lincoln Constance, in his one year of
Herculean efforts to sort of make the way for me less difficult,
couldn’t manage to bring the Gray back into its best form by the
time I arrived in 1948.

1986] NEBC 800th meeting — Remarks 177

One of the tragedies that did occur, and which I always thought I
might have done something about, was the demise of the Botanical
Garden. I don’t know how many of you remember or know that, but
by the time I arrived on the scene here, the Botanical Garden had
gone the way of all flesh. The best plants had been taken out and
moved to the Arboretum to save them and the bulldozers were busy
tearing up the Garden and eventually housing was put there.

The development then proceeded in a rather different way. The
Gray Herbarium, as an isolated institution, became somewhat unten-
able and eventually was moved into its present location. Fernald
wouldn't have liked this and, in fact, was very much opposed to the
demise of the Garden even though he took very little interest in it. A
rather amusing incident had to do with the naming of the streets
that now go across the old Botanical Garden. Those of you who
know Cambridge know that the street on the back-side of the old
Gray Herbarium building is known as Robinson Avenue and the
one on the front-side is known as Fernald Drive. When we broached
the subject of naming these streets, Professor Bailey was then in
charge of the general area~he and | consulted on it--and we
hatched up the idea of naming them for Fernald and Robinson
because Gray was already honored by street names. There is Gray
Gardens East and Gray Gardens West and Gray Street, and it
seemed appropriate to have more botanists commemorated in this
way. We wanted to sound Fernald out a little bit before proceeding.
Some of the people had said that he is rather a vain man and he
really would like this sort of thing, but that his response would
probably be negative. And indeed it was negative, on the surface,
but after we probed a little we could see that he was pleased with the
idea and so we went ahead. I’m sorry that we don’t have projection
facilities because I do have a slide here that shows Fernald in front
of Fernald Drive and he was rather proud to pose for that picture.

Coming along late in the evening as I am on this program, I find
that the stories that are told by the people who knew Fernald in his
early days are rather of a different character than the stories that |
recall. I think this is largely due to the fact that Fernald, as he
progressed in his career, rather had a little different view of what he
should be doing. He became completely engrossed in the revision of
Gray's Manual. He did field work and he did write about it, and his
written work had all of the characteristics that have been men-

178 Rhodora [Vol. 88

tioned. He wrote very well. But he, somehow or other, around the
Herbarium itself, seemed to concentrate more truly in his work on
the Manual, so that he rather resented interferences. And the stu-
dents, in particular, were not as freewheeling as far as going to him
was concerned as they might have been, | think, at an earlier
period. Although, in talking to Lyman Smith the day before yester-
day in Washington about Fernald, Lyman Smith said, “You know,
Henry Svenson, one of his early students, I felt very sorry for,
because Svenson was scared stiff to go to Fernald and discuss his
thesis in detail. He thought that the thing to do was to prepare the
thesis completely and then go to Fernald, so he proceeded in that
fashion. Eventually, he went to Fernald and Fernald found some-
thing wrong with the way Svenson had prepared the descriptions.
Fernald made him make changes, so Svenson had to go through his
entire thesis, altering I don’t know how many species descriptions.
The same corrections had to be made over and over again all the way
through.” Lyman thought it to be a great shame that a graduate
student had to go through this kind of thing.

I won’t say much more. I have quite a bit more written down here.
I could talk about Fernald’s closest friends, for example. I believe J.
Franklin Collins from Providence, Rhode Island was one of his
best friends. When Collins died, he left Fernald his set of large glass
projection slides, a rather extensive collection of photographs taken
during their field trips together over an extended period of time.
Fernald used these considerably in class along with the slides he had
made himself. He was particularly interested in the distribution of
plants through the areas that have been mentioned, from the Pine
Barrens of New Jersey all the way up the coast, particularly Long
Island, Cape Cod, and then further north. He often spoke of the
plants of the coastal strip.

Bayard Long was another of his very close friends. Fernald, as
some of you certainly know, was a rather frugal man. He certainly
was well-off when he died. He played the stock market and Bayard
Long, who also played the stock market, was his confidant in this
matter, Fernald would take his tips from Bayard Long. Fernald
took this seriously to the point where when we would go on a field
trip, he would say, “Now we must stop at a Texaco station because
I’ve got stock in Texaco.”

We had memorable trips to the Cape. Fernald would hire a cot-
tage where the class would stay, and we would work out of there. I

1986] NEBC 800th meeting — Remarks 179

know that Walter Hodge, for example, took pictures of some of
these trips and a few of these were included in a series of articles
published in Rhodora at the time of Fernald’s death. Students liked
to play little tricks on Fernald. He was always very, very interested
in what he was doing and would get so engrossed that he didn’t
really pay any attention to anybody who might be clowning around
him. And so it was often the case that a student would pick a long
spear of grass or maybe a dried-up Solidago infructescence with a
piece of stem on it and then they'd stick this into his hat, you know,
and it would be towering up above. | remember one picture, | think,
Walter took with the hat so adorned. But, these kinds of things were
fun for the students and he, I’m sure, would have entered into it
more at an earlier period in his life but didn’t do it so much at that
particular time which was well toward the end of his teaching career.
Thank you very much. (Applause).

R. A. Howarp: Thank you, Reed. The Gray Herbarium in my
period was largely a one-man show. As I indicated to you, Fernald
was in charge certainly. But there was a second person around
there—Charles A. Weatherby, who graduated with an A.B. from
Harvard at the same time that Fernald got his B.S.; that was in
1897. Weatherby was a specialist in the pteridophytes and he was
entitled to have students. That was why I was out of place as being
Johnston’s student. One of Weatherby’s students—and perhaps
looking in from the side— was Dr. Rolla Tryon. Rolla, will you take
over for a few minutes? Maybe I should say “looking in from the
top,” because if any of you remember the atrium that was the Gray
Herbarium, you went up the side steps until you got to the top
balcony, then in the middle of the front of the building there was
another set of steps that took you into this aerie that was Weather-
by’s hang-out.

ROLLA TRYON: Well, what I wanted to speak about takes a little
different tack than some of the other people have, and that was to
mention the contribution of Fernald to the subject of Plant Bio-
geography. His field work has been mentioned, and certainly Fernald
was an avid collector and I suppose that, with the others he collected
with, he must have taken at least 100,000 sheets, probably more (not
that many of each kind because they often collected in duplicates).
And it’s been mentioned how he collected, for example, in Virginia,

180 Rhodora [ Vol. 88

Massachusetts (particularly Cape Cod), Maine, Nova Scotia, the
region of the Gaspé, upper Gulf of St. Lawrence, Newfoundland,
and rather later, northern Michigan and adjacent Ontario. This
collecting was clearly directed toward two lines of research: one,
that of understanding the species of the Gray's Manual area better
and getting their taxonomy straightened out, and the other was
toward biogeography.

I think that Fernald’s genius, in fact, was to synthesize this taxo-
nomic research, the biogeographic research and his field work essen-
tially into one whole. They were not at all separated. His
biogeographic method, which he developed to a point which still
serves as a model today, was first of all to see unusual ranges,
particularly disjunct ranges, and to see that a considerable number
of species had essentially the same kind of disjunct range. Then he
would begin to look for paleoclimates, paleogeological explanations
that might afford a clue to when and in what direction these ranges
that were now so isolated were once, in fact, connected. And so he
sought the explanation in recent geological history. If you read his
writings on biogeography closely, you’ll find that they’re full of the
ecology of the species he was dealing with, not just the distributions
themselves. And particularly in relation to his Nunatak Theory, as
well as others, he was in fact nibbling at the edges of population
genetics. To those of us who knew Professor Fernald and heard
what he had to talk about, it might be surprising that he would ever
get into that kind of subject at all. Of course, he wasn’t very deep
into it, but he did have to deal with these very local endemics that
did not spread in their distribution, and he was considering prob-
lems related to that.

His major theories I think many of you are quite well aware of.
One of the first related to eastern North America and Europe. In
many cases he was grumbling greatly because our species here were
still under the name given in Europe. He did a great deal of work in
sorting out these species and finding, in fact, that many of them
were closely related but different species. In this respect, he pro-
posed a theory of an earlier North Atlantic connection, when at
least there was much less water there, and that many of these species
could have migrated between North America and Europe. There
also has been mentioned his work on Nunataks, which related to the
Gaspé area particularly, and to the west, in which he envisioned
these regions as having escaped the last glaciation and species were

1986] NEBC 800th meeting — Remarks 181

disjunct because they were able to persist in these very small areas.
Mount Albert was one of the most famous of those. And then, also,
there was the matter of the coastal plain. This involved Cape Cod,
Nova Scotia, and Newfoundland. Almost all of the areas where he
went to collect had biogeographic significance to him, which is why
he went to these areas. These coastal plain elements are now very
disjunct from Cape Cod to the north, and he envisioned a shelf off
the present coastal plain that later had subsided. Originally, how-
ever, it could act as a migration route.

At the time Fernald was working, the geological evidence for
much of this—the evidence that he required was not, in fact, pres-
ent. The geologists were rather slack in Fernald’s opinion (laughter),
and he didn’t hesitate to say so in print. In fact, he rather goaded the
geologists to do some more work and give him the kind of informa-
tion he needed for his historical explanations. Whether in fact these
theories and these explanations are eventually accepted or not as
right or wrong is not really critical to Fernald’s contribution in
biogeography. He had two, certainly, lasting contributions. One was
his very strong influence on other botanists; for example, Hulten,
Porsild, Raup, Victorin, Fassett, Hodgdon, and the many col-
leagues and many students of these people as well. Also, he pub-
lished a vast amount of accurate data on ecology and the geography
of species and integrated this into a coherent whole which stands as
a model without respect to the precise accuracy of his conclusions.

I think, finally, that it is fair to say that Fernald lived in the
Golden Age of Plant Biogeography in America, and he was the
central figure in it. (Applause).

R. A. Howarp: Thank you very much, Rolla. I would like to
submit a written contribution I received, a charming several pages
from a foreign graduate student of Professor Fernald’s last class in
Harvard University. While it may be a little long to have all of it, is
Shiu Ying here? Yes. Dr. Hu, would you please tell us a couple of
those stories at least that I found so interesting?

SHiu YING Hu: Thank you, Dr. Howard. In the summer of
1947, Professor Fernald completed his teaching career in Harvard
University. I was fortunate enough to be a student in the last class he
taught. However, he was not my first teacher because his course was
in the spring. Before him, I had Professor Paul Mangelsdorf for

182 Rhodora [Vol. 88

Economic Botany, Professor Karl Sax for Cytotaxonomy, Profes-
sor Elso Barghoorn for Paleobotany, and Professor Elmer D.
Merrill, my major professor, for a 300 course. I found that
Mangelsdorf’s and Merrill’s courses were manageable; Sax’s and
Barghoorn’s courses were pretty hard. Both of those men spoke very
low and I couldn’t get the words—they spoke half way and
swallowed the rest! (laughter). And Fernald’s course was, to me, just
incredible. The reason was that Fernald did not teach with botanical
language, so I just couldn’t catch what he taught. I was poorly
prepared for his witty deliberation, clever statements and very
amusing stories.

Professor Fernald’s course was held in the old Gray Herbarium
building situated in the corner of Garden Street, Linnaean Street
and Robinson Street. When I went there, the first thing that
attracted me was a beautiful rock garden just in front of the right
wing of the building, and in the early spring, it was just beautiful.
The class consisted of about eight students; most of them were
American G.I.’s. There were two foreign students; John Constable
from Great Britain was an undergraduate, a senior, and of course,
myself. The subject matter of the course was botanical history and
phytogeographical principles. Fernald didn’t have an outline for his
lectures because everything he had was in his head, and for his guide
of the botanical history, he used a large file that Mrs. Asa Gray
collected. Those big books—I actually measured for this talk——were
400 cm long, 275 cm wide, and 10 cm thick! There were many
volumes.

At that time, we had classes in the Bio Lab and then we had to
walk all the way across campus to the Gray Herbarium, which was
opposite the Observatory. When we arrived, Fernald would wait for
us with one big volume, usually open, and he started from A to Z. It
was my first year in America. All the names sounded queer to me,
and I just couldn’t catch the sounds. Finally I moved my chair to his
side, so that as soon as he opened that big book, I copied the names!

The laboratory work, as Dr. Howard said, was in the right-hand
wing of the Gray Herbarium on the first floor. The large room was
for Harvard men, and the small room, I was told, was the Radcliffe
room. Dr. Howard was on one side and I was given a table, and
Professor Fernald came to have lunch there by himself every day.
The first time I saw Professor Fernald having lunch in the Radcliffe
room, I made a terrible mistake by American standards. It was done

1986] NEBC 800th meeting — Remarks 183

partially because of the things he ate, and partially because of my
ignorance of American academic life. Fernald came to the Radcliffe
room, sat there, opened a small cake-box, took out a banana, and a
sandwich, and then began to chew a big raw carrot without peeling
or cutting it. I was horrified! (laughter). In order to explain this
incident, I need to tell you something of my background. The rural
people of the lower Yellow River region in China eat raw onions,
garlic, green pepper, coriander, tender parts of Chinese cabbage and
raw carrots. But the people in cities, and especially the literate peo-
ple, don’t eat these things. I learned this difference through a very
bitter experience in a boarding school. I was sent to a boarding
school at the age of 11. Approximately 90 percent of my classmates
were 4 to 8 years older than I. One day a relative of mine brought a
bundle of presents from the village, and in it there were some car-
rots. I was so happy for my gifts and began to eat my carrots. A big
city girl ridiculed me, saying, “Look at this little barbarian girl
from the country. She is eating a raw carrot.” I quickly hid my
carrot and from that time on I had never had a raw carrot until |
became Americanized. So when | saw Professor Fernald chewing
that raw carrot, my hidden wounds suddenly surfaced. I walked to
Professor Fernald. I said, “Professor Fernald, do you eat raw car-
rots?” (laughter). As he chewed, he said, “Why not? Carrots are
good for you.” I said, “I heard people say only barbarians eat raw
carrots.” (laughter).

Well, you know that the Gray Herbarium was a small family, and
news went out very fast, so before the end of that afternoon every-
body knew something I did with Professor Fernald! There was a Miss
Marjorie Stone who helped Professor Weatherby with the Gray
Index. She was a very serious person and she came to find out if 1
really did say, “Professor Fernald, you are a barbarian!” (laughter).
Well, don’t laugh. The better part is coming. You see, at that time, it
was only a few months earlier that I came from China, which was
isolated for eight years and was bombed by the Japanese. During
those years, unlimited inflation ruined the lives of college profes-
sors. Instead of taking home money as salaries, (money didn’t mean
anything), the professors would, for each person of the family, take
home three bushels of rice. For example, a professor with a wife and
two children would take home 12 bushels of rice for salary, for food
and for exchange of other things. So during the war, the Chinese
professors really had a very hard time. When I came, I saw Profes-

184 Rhodora [Vol. 88

sor Fernald chewing a raw carrot and I thought that he was under-
going similar economic hardship (laughter). So as I walked out of
Gray Herbarium that afternoon, I decided to do something for this
“poor old man.” The next time I came from class, I brought two
large Malus Delicious apples. When I saw Professor Fernald start-
ing his carrot, I held one apple in each hand behind me and
approached him. I said, “Please do not eat that carrot (laughter); eat
these apples.” I put the apples in front of him, and he took up one
and ate it. Later in that afternoon, | met Carroll Wood in the hall.
And he said, “It won’t work; we won't allow it. If you get a good
grade in this course, we won’t allow it. It’s unfair. You are Professor
Fernald’s ‘apple-polisher’.” (laughter). Not understanding the signif-
icance of “an apple-polisher” in American schools, I replied, “No, I
didn’t polish his apples; he had no apples. I gave them to him.”

Fernald liked to alert his students before starting classes. One
day, as we all gathered around him, instead of opening his book he
put his hand on his bald head and said, “I forgot where we stopped.
What shall | talk about?” Well, immediately I picked up. I said,
“What about continuing the last chapter of the Book of Revelations,
Professor Fernald?” (laughter). I think that it must be because of
this comment that he gave me a passing mark (laughter). We all had
fun with Professor Fernald. Thank you. (Applause).

R. A. HowarD: For many years, all of Professor Fernald’s grad-
uate students were required to take on as a thesis, a topic that was
associated with the development of Gray's Manual. I think one of
the first people to violate that particular edict was Walter Hodge,
who somehow got some work in on the flora of Dominica, but he
was such a master photographer that, I think, Fernald admired the
technique of what he was doing, and Fernald certainly used many of
Walter’s pictures in the course of various publications. Dr. Walter
Hodge joined the Club in 1939. He now spends his time partly in
Florida and partly down on the Cape, and he is here this evening
with Barbara. Walter, will you talk?

WALTER H. HopGe: Thank you, Dick. Yes. I’m pleased to be
here, also, to participate in this tribute to Merritt Lyndon Fernald.
As Dick Howard just alluded to, I hadn’t realized that I was one of
the odd-balls who didn’t pick up a thesis topic dealing with the
Grays Manual range. It so happened that I had been studying or

1986] NEBC 800th meeting — Remarks 18S

working at Massachusetts State College, now the University of
Massachusetts at Amherst where I was a young instructor. My
former professor at Clark University, where I did my undergraduate
work, was David Potter, another Fernald student; and it was
through David Potter that I got to know Fernald and also the
reason I was up at Mass. State, which was Potter’s Alma Mater.
Anyway, while at the State College, | became interested in the
tropics; I was starting a course in an area where I eventually became
very much interested, Economic Botany, and I wanted to make a
trip into the New World tropics to collect economic plant specimens
for teaching purposes and so forth, and perhaps identify a doctoral
thesis problem. And so it happened, as Dick Howard has said, that
in 1937 I spent a couple of weeks on Dominica, then a British island.
Since it was little known botanically, as most of the Caribbean
islands were at that time, I had to visit the Gray Herbarium in order
to identify what I had collected. And in the course of this, | met up
with Fernald. I had been told earlier up at Amherst, “Well, young
man, you’re an instructor here, but you ought to go on and get a
doctorate.” That I well knew, so I broached the subject to Fernald,
stating that | had what I thought would be an interesting thesis
problem for me, a floristic study in the Lesser Antilles. Well, he
hemmed and he hawed and, as I recall, he snorted about the
botanists down in Washington who spent most of their time in the
tropics when they didn’t know a darn plant around the Washington,
D.C. area, where he was picking up novelties all the time. He and
Bayard Long were then collecting in eastern Virginia. Fernald had
had some physical problems and wasn’t able to continue “the more
rigorous field work,” up in the northeastern area, so he was
“graduating” to the flat coastal plain area of Virginia.

Anyway, he couldn’t understand why I couldn't find a problem in
that area, or at least in the Gray's Manual range. But | said, “Well, |
really hoped I could do something in the tropics. I have been teach-
ing, Professor Fernald, up at the State College and I also do photo-
graphic work, which I enjoy as an avocation.” This brightened his
eyes a bit. It happened that it was a rather fortuitous time because
Eugene Ogden, who is here tonight, was just finishing up his thesis
work and he had been doing the photographic lab work and assist-
ing in Fernald’s courses and so Fernald needed a replacement and |
guess he sort of said to himself, “Well, anyway, I'll get the photog-
raphy and the teaching done, even though Hodge doesn’t elect a

186 Rhodora [Vol. 88

thesis problem that focuses on the Gray’s Manual range.” So that’s
how it happened that, as Dick says, perhaps I broke what had been
a general precept to work under Fernald. Well, in a way, that was a
relief for oftentimes I didn’t get bogged down when Fernald would
come around, as he was wont to do, reading his own manuscripts on
coastal plain plants for the Contributions or for Rhodora, or what-
have-you, and if I had a lot of things to do, well, I could always say,
“Well, Professor Fernald, there are some prints washing down in the
darkroom. | really must get down to them.” So he would rush
around to find someone else more appropriate to whom he could
read the manuscript.

In those days, from °38 to "41, among student confreres at the
Gray were Reed Rollins, Rolla Tryon, Bob Godfrey (too bad he
isn’t here; he’d have some great tales to tell), Jim Soper, Shirley Gale
(now Shirley Gale Cross) and Bernice Schubert. Of course Reed
Rollins had been around a year or two before us, as had Louis
Wheeler as well. I forget who else was there; oh, yes, Carlos Mufioz,
a Guggenheim Fellow from Chile, with whom we had great
fun. Well, I recall, as others do here, that the most fun with Fernald
was when he was in the field. Although his lectures were full of all
sorts of interesting stories, he really blossomed when he went into
the field. Of course the trips in what we called Biology 17, down on
Cape Cod were the greatest time for most of our fun. Invariably
Fernald would rent, as Reed has said, a house and then we’d go out
into the field and that was a time when we learned among other
things about some of the materials that appear in Edible Plants of
Eastern North America. For example, I can recall that we tasted for
the first time the fresh shoots of orach, and the tubers of Stachys,
and once when Professor Fernald was explaining something, one of
us slipped a plant into the band of his fedora. It really wasn’t a
Solidago, Reed, as I recall, it was something more appropriate. It
was a Stalk of Lespedeza capitata; and all the while he went right on
telling us about the differences between this and that plant.

One of the interesting times on these trips were the evenings. Then
he would reminisce about former field experiences. I’m sure some of
you must have heard, this being the Cape, of one of his famous
problems with the local police during World War I. You know Pro-
fessor Fernald looked sort of like a little Prussian with a goatee and
while in the field he would be dressed as a European might be witha
suitcoat and a tie, no loose sport shirt or anything like that, and an

1986] NEBC 800th meeting — Remarks 187

old felt hat. Well, on this occasion he was looking for Orontium
aquaticum, the golden club, which had a northernmost extension
somewhere on the Cape. In order to locate this plant, he went to one
of the fire towers on the Cape and got out his fieldglasses (“spy-
glasses”). He was equipped of course with a vasculum— a “bomb-
case,” and as it turned out, he had a little “code book” as well, which
was a Gray's Manual check-list. To the police he was reported to be
a spy trying to help German submarine captains, presumably cruis-
ing offshore. Anyway, he was put in jail (laughter), and only aftera
call was made to Harvard’s President Elliot was he released. He was
a German spy! (laughter). | think the same thing happened during
World War II in a Virginia railroad yard, but I don't recall the
details.

One of the other stories that came out on one of those field trips
had to do with Fernald’s excursions on the Gaspé with Professor
Pease. I had spent some time in the Shickshocks, in Fernald’s
nunatak areas, and so was particularly taken with his stories of how
he and Pease first tried to navigate along the Gaspé coast. That
was before there was any highway, so they traveled down the St.
Lawrence by fishing boat, I believe, from village to village. On one
such occasion headed for the Shickshocks, they were unable to find
any overnight accommodations. As it turned out, they went to the
local post office, which was just a little frame shack with a door
opening directly onto the street, and asked the local postmaster
what he could do to help them. His reply was, “Well, there isn’t any
place where you can stay around here, but I'll tell you what. Why
don’t you just spend the night here in the post office. Of course, Vil
have to lock you in, but I’m here promptly in the morning.” They
said that was fine, and it was so arranged. But pretty soon after they
were locked in and it was time for putting down the bedrolls, one of
the “urges” came along and they wondered “Well, what the heck are
we going to do?” I don’t know who it was, Fernald or Pease-
probably it was Pease (laughter)—who said, “Well, Fernald, there’s
the mail slot!” (laughter). On the same trip, they were up in the
Shickshocks. Pease, Latin scholar, said one day, “Have you noticed,
Lyndon” (or Merritt, or whatever he called him), “what we’re using
here for T-P? It’s quite appropriate— Dryas.” (laughter). Fernald
enjoyed these earthy stories, and whenever he was getting a point
across, people may recall that he hit his forehead like this (with the
heel of his palm), whether he was reading manuscripts or what-
have-you, and that was to emphasize the point.

188 Rhodora [Vol. 88

Well, I think that is the last anecdote that I should tell. However,
I do recall that the war came on rather promptly after I left
Cambridge in 1941 and many became involved in the war effort. I
was abroad for several years and didn’t see Fernald until my return
from Peru. We had been using alcohol in collecting herbarium
specimens of Cinchona in the Andes simply because it was a little
easier then using presses. I hadn’t seen any published record of this,
so | wrote a short article for Rhodora to report on it. I think Dick
Schultes urged me to do this. He’d been doing something similar
utilizing formaldehyde. Anyway, I wrote this piece and I got a nice
reply, the last letter | received from Professor Fernald, written to me
at M.S.C. (Amherst) in April of 1947:

Dear Hodge: At the risk of making Rhodora seem to
some readers to be a saturated solution of preservatives, we
shall be very glad to use your fine paper later in the year. | am
glad that my work has been in temperate regions where it has
not been necessary to smell of alcohol (laughter). Kind regards
to the Mrs. and the youngster. Yours sincerely, Fernald,
Editor-in-Chief.

Thank you. (Applause).

R. A. Howarpb: We could get started on the Hodge-Godfrey
stories around here, too, but one of them involved the time when we
were in Cuba and Bob Godfrey knew that Bernice Schubert was
working on the genus Desmodium— Desmodium has adherent
loments, you know, that stick to your socks—and so after one day
in the field, when we came back with a set of socks that were just
loaded up with these fruits, Bob Godfrey decided that this was the
thing to send to Bernice Schubert, so he mailed them up to
Cambridge and the post office and the Harvard Herbarium were
never the same after that! Bernice Schubert was one of the key
people in Fernald’s production of Gray's Manual the 8th edition.
We hoped that she would be here tonight. Unfortunately, she was
not up to it today. I just want to read the last bit of the Preface of
the 8th Edition that Fernald wrote:

Finally, especially since the impairment of his eyesight, the
author has had the most loyal, conscientious and unlimited
aid of Dr. Schubert in the exacting details of coordinating
usages in the text, editing the manuscript for the printer,
checking and double-checking the citations of figures, and in

1986] NEBC 800th meeting — Remarks 189

the scores of other details necessary in the book as it goes to
press. My appreciation of all her helpfulness cannot be
adequately expressed.

If you read a lot of Fernald’s writing, you will never again find a
dedication that is as warm as that or as sincere as that, so we felt
that Bernice could contribute something to this program. She has
indeed supplied a few of her letters, and while we are going to ask Dr.
Carroll Wood to talk anyway, he is going to represent Bernice
Schubert, too, at the microphone. Carroll.

CARROLL Woop: Well, Bernice was unable to be here tonight.
She has been having some difficulty with a respiratory problem and
she said she just couldn’t make it. But she did agree to write down a
few things. She felt very warmly toward Professor Fernald and he
toward her, and I think she was reluctant to talk about him very
much 1n public, but she did look into her file of letters from him and
we've got some quotations from those. She writes:

“I’m sorry I can’t be present at the meeting this evening, but most
of you know that I worked rather closely with Professor Fernald,
especially in his last years of effort on the 8th edition of Gray's
Manual. Most of you are also aware of his botanical accomplish-
ments and are familiar too with one aspect or another of the
mythology that has grown up around him. I should like to have you
know a little of how I felt about him as a person, as a teacher, and as
the person for whom and with whom I worked at the start of my
botanical career. | wondered for some time about the best way to
present a few ideas and remembered a folder of letters from Profes-
sor Fernald to me on two European trips when I photographed type
specimens for him. Perhaps just a few excerpts will acquaint you
with him best. However, as an introduction to Merritt L. Fernald, a
few lines written in 1891 by his mother to Sereno Watson, then
Curator of the herbarium, may serve. Mrs. Fernald writes:

I write to ask you not to hold our son Merritt responsible
for the false and senseless reports which have in some manner
found their way into the daily papers, regarding him and his
going to work in the Harvard Herbarium. It is a matter of
great annoyance to us all and especially to Merritt, who is a
very modest and diffident boy. As the simple statement about
his leaving home, which we have made when necessary and
desirable, added to certain mistaken and exaggerated notions

190 Rhodora [ Vol. 88

expressed by some of his college mates, have in some way
furnished material for sensational reporters. Of course, we are
doing what we can to correct false impressions. And Merritt’s
disgusted exclamation, “What will Dr. Watson think of me?!”
made me wish to set the matter right with you at once.”

This is from a letter in the Gray Herbarium archives, and appar-
ently newspapers had picked up this sensational thing about the
young boy from Maine going to become a professor at Harvard.
Bernice continues:

“On the day that World War II ended in 1945, Professor Fernald
suggested that I look into the possibility of going abroad to study
and photograph type specimens for him. In late October 1946, I
managed to get off on my first trip outside the United States. On
arriving at the British Museum (Natural History), I found a letter
with a note from Professor Fernald:

To greet you at the herbarium at South Kensington where |
spent very happy weeks in August and September, 1903, and
again in the summer of 1930. You will find the botanists there
a very kind and helpful group. We are imagining you toasting
along the Gulf Stream today and very soon steaming into the
Channel.

“His note was followed by notes from several members of the staff.
By mid-November, many new botanical requests started to come in
the letters:

One item has just come up. Please make a note of the degree
of pubescence on the leaves of Fraxinus caroliniana Miller.
Ordinarily, F. carol. has been treated as the glabrous-leaved
tree and the pubescent one called a var., but Miller says it is
pubescent. Sargent and others, including me, seem to have
taken it for granted without going back to Miller. Foster and |
are reading the Manual transcript for an hour daily, checking
for consistency in small details. He is a great help, catching
errors of centimeters for millimeters, and so forth. Pease has
finished checking the Latin and Greek except for recent
manuscript, and I shall say in the preface, ‘if any errors are
found, they are obviously of sections we did not see!”

“Often there were snippets of news or gossip that followed
requests, as in the following letter of late December 1946:

1986] NEBC 800th meeting — Remarks 19]

Here is another type in the Linnaean Herbarium. Do find
out why the Linnean Society ‘with an E’ but Linnaean Herba-
rium ‘with an AE’. Weatherby asked, I too, that you examine
a photograph, and photograph with details of the lower leaf
surface of flowers, if possible, to determine whether the leaf-
lets are pubescent beneath, in which case, T (which is Thalic-
trum) purpuream displaces Thalictrum polygamum Muhl. or,
are they covered with waxy atoms in which case it displaces T.
revolutum D.C. Incidentally, Boivin is bringing down to the
meetings after Christmas a ‘young bride,’ Cousette Marcou,
director of the Children’s Gardens at the Montreal Botanical
Garden. Cousette sounds rather cuddly (laughter).

“The last letter | received during this trip was in January 1947 and
contained a rather detailed description of the Triple-A. S. meeting
held in Boston in December along with a plaintive note, saying that
he hoped I would be back soon to help him clear his table and clean
his ‘specs’, which no one had done in my absence.

“In the spring of 1950, I was fortunate to be able to go abroad
again to travel to more herbaria, to attend the first International
Botanical Congress after the war, to do some of my own research,
and to fill some more requests for Professor Fernald. Several
members of the staff, including Professor Fernald, saw me off, and
not long after my arrival in London, I received a letter with botani-
cal requests concerning Spiranthes and the following paragraph:

We were delighted to know from Miss Campbell that you
had a good trip. With blizzards and shipwrecks along our
coast, and the long delay in receiving your manuscript, we
pictured you as being like the heroine of my youth:

“To sail out on a broad ocean,

A sweet little maid took a notion.

But when the yacht rolled,

She said, “I'd give gold

To get rid of this horrible motion”.’
and there’s more:

“There was a young man from Ostend

Who vowed he’d hold out for the end.

But when halfway over

From Calais to Dover,

He did what he didn’t intend.’ (laughter).

192 Rhodora [Vol. 88

“He also mentioned in this letter that he had received proofs up to
page 1500 of the new Manual. Letters continued to come, but short-
er and mostly dictated, because during this period, he had a coro-
nary thrombosis and although he returned as soon as possible, by
Stages, to working, it was difficult, as his eyesight had been some-
what affected. However, he did see his Manual, he enjoyed and
appreciated the letters he received from all over commenting on it,
and I think he felt he had completed this work to the extent of his
ability. Surely this meeting is proof that he produced something of
value and that his influence still lives.”

And these are some comments by Bernice Schubert, whom | first
met when I came to Harvard as an ex-G.I. in 1946 by virtue of the
G.I. Bill. | had corresponded with Fernald earlier from Virginia
where I was collecting plants. | had gotten interested in taxonomy
through a student of Fernald’s, Jack Fogg, who at that time was a
professor at the University of Pennsylvania. I had intended to go
back there after the war, but when the opportunity came, I thought,
“Well, Professor Fernald is the one that I ought really to go to work
with,” and so, with Fogg’s blessing, | wrote to Fernald, and some-
how he got me into Harvard in the middle of the summer of 1946.
As Dr. Hu has said, we were both in that last class he taught in the
spring of 1947. I remember a couple of the stories she told, and
“The Last Chapter of Revelations” has always been one of my favor-
ite ones, because it was quite appropriate. The first half of his course
with the autograph collection was fascinating. As she said, he only
got about half-way through the alphabet, but I think he did get as
far as Marcus E. Jones; and he even pointed out the scurrilous
attack that Jones had made in print on Fernald himself, along with
a lot of other people. The second part of the course was mostly
readings from the geographical papers of Fernald, and most of us
had read these. That day when he came in, as I recall, he had a
volume of Rhodora open and he said he didn’t know quite what he
was going to use as a text for the day, and she remarked, “How
about the last chapter of Revelations?” Everyone laughed and Pro-
fessor Fernald got what my mother calls “the dry grins,” which is
when the joke is on you, you have to laugh, but you don’t really
want to. For years, | wondered whether Shiu Ying really knew what
she said then.

Fernald’s habit of reading to people was notorious. I got cor-
nered numerous times with that. You couldn’t get away once he

1986] NEBC 800th meeting — Remarks 193

was reading, and usually these were manuscripts that he later pub-
lished in Rhodora. It was a long time later that I realized that he
wasn't really reading to the person in front of him. He was reading
to himself. He was polishing his prose, and of course one of the best
ways to do that is to read the terrible stuff you’ve written out loud to
see how it sounds. A story that Professor Raup told me illustrates
this very nicely.

Dr. Raup has sent two pages of newsprint, which are typical
Fernald manuscripts (he has saved them all these years and sent
them to Dr. Howard) in Fernald’s handwriting, which never
changed at all throughout his entire life, as far as I can see, which
makes it very difficult to date his annotations in the herbarium,
which he frequently forgot to date himself. But Professor Raup told
me that one Saturday afternoon he came into the Gray Herbarium
to do some work and brought with him his little boy, only three or
four years old. Along came Professor Fernald who started to read to
them. The little boy was sitting on the table and was fascinated by
Professor Fernald’s goatee (laughter) as it went up and down as he
read. Raup stood around for a few minutes, then he went off and
did what he had to do in the herbarium; when he returned 45 min-
utes later, Professor Fernald was still reading to the little boy
(laughter) who was listening attentively, so it was evident to whom
he really was reading!

Well, the hour grows late. I’ve been reminded of all sorts of things
that I had forgotten all about, including the lunch and the carrot in
the Radcliffe room. It was there that Professor Fernald, who was
very kind, showed me about eating the flower buds of Hemerocallis
(daylilies) which he had cooked in a custard. He had a little jar of
those one day and tried it out on me. He also pointed out to me the
new shoots of Polygonum cuspidatum which he said made an
interesting “nibble.” I tried those and, after reading The Edible Wild
Plants of Eastern Northern America, | eventually decided that any-
thing Professor Fernald said was a “nibble” was inedible to anybody
else but him (laughter). He was willing to try anything. That’s where
I think I'll stop. Thank you. (Applause).

R. A. Howarb: We’ve all known about the manuscripts that
Fernald used and read to so many people. It was an unsized half-a-
sheet of newspaper, and it was read over and over again. My favor-
ite story about this concerned the time when Munroe Birdsey,

194 Rhodora [Vol. 88

whom some of you may know, came into the Gray Herbarium for
the first time. In the corner at the side of the door was a mail shelf
which was just high enough to hit Birdsey in about the back of the
neck. As he walked in (I was up in the second balcony so I heard all
this going on), Fernald, who usually shuffled his chair as he pushed
it back, cleared his throat—chmmm-—and it was time, obviously,
that somebody was going to be “read to.” Birdsey came in the front
door, just as Professor Fernald came into the atrium there. There
was no one in sight, and Fernald looked up at Birdsey and said, “I
think you would be interested in this,” and then proceeded to read.
About 20 minutes later, he looked again at Birdsey and said, “What
do you think of that?” and Birdsey said, “Well, that’s very interest-
ing, sir, but who are you?” (laughter) and Fernald said, “M. L.
Fernald.” And Birdsey said, “Not the M. L. Fernald?!” In the mean-
time, he had been backed up against the mail shelf so that his neck
was almost broken, but he tells the story on himself, too. That was
his introduction to M. L. Fernald. Those sheets are very, very rare
and we literally could not find any around the building. There may
be some in the archives, but when I mentioned this to Professor
Raup, he discovered in his old manuscript that he published on the
Peace River flora, that he still had a sheet which was Fernald’s
description of an Antennaria published in that particular Contribu-
tion of the Arnold Arboretum.

Being the last speaker in one of these series is the most difficult
because the best stories have been told and probably everything you
had to say has been said in one fashion or another, but we asked Dr.
Richard Schultes to give us some of his recollections of M. L. Fer-
nald. Dick.

RICHARD EVANS SCHULTES: Sorry, I’m just back from Borneo
with a jet lag, so I'll have to follow notes.

Whenever former students of Professor Fernald get together, the
tales inevitably flow endlessly throughout the evening. Fernald,
irritating but lovable, was truly what is generally known as a
“character,” of which Harvard in those years had an ample share. |
don't think there are any around now (laughter).

But Fernald never shrank from a fight. It did not matter whether
the opposition was a colleague at Harvard, a botanist somewhere
else who dared to disagree with him on a matter of taxonomy, or
even the University administration itself. His typically Maine

1986] NEBC 800th meeting — Remarks 195

straightforwardness and bluntness came to the fore. In the several
courses that I took with Fernald as an undergraduate and graduate
student, I came to respect “the Prof” for his incredible knowledge of
the temperate American flora, for his extraordinary understanding
of the historical development of North American botany, and for
his natural ability as a teacher, and yes, even for his occasionally
irascible reactions. In those years, the three dominant figures in
Harvard taxonomic botany were Fernald, Elmer D. Merrill, both
State of Mainers, and Oakes Ames, a Bostonian. Although he held
both Merrill and Ames in great respect as botanists, Fernald
enjoyed piquing both. Merrill always responded, which is exactly
what Fernald wanted. Ames, always aloof, never rose to the bait,
which infuriated Fernald (laughter).

It is said that Fernald sometimes made it difficult at the Gray
Herbarium for students of other professors, especially those profes-
sors with whom he was carrying on a scrap. I was doing my graduate
work under Ames, but Fernald always treated me courteously, even
in a fatherly way, as if I were one of his own students. Only once did
I personally sense his antipathy towards Ames. Whilst preparing for
my first collecting trip to southern Mexico, I asked Fernald if |
might look for any material for him. He asked, “Schultes, how
many corrugated cardboards are you taking?” When I replied that |
was not taking any cardboard but aluminum corrugates because |
would be working in the wet tropics, his eyes sparkled and he
remarked, “Oh, yes, | forgot. You work with that rich dilettante
down in the Botanical Museum” (laughter). He said, “We at the
Gray Herbarium cannot afford such a luxury as aluminum
corrugates!”

How well I recall the many interesting quips and jabs that often
served to impress upon us some important point in the mass of
details that without a single note he poured forth in lecture after
lecture. His characterizations of other botanists from Linnaeus to
his contemporaries was telling. For example, he described a taxon-
omist whose treatment of the Cyperaceae left much to be desired as
a “bombastic boob.” He consistently referred to the great botanist,
Liberty Hyde Bailey, whom he respected highly, as “free-skin Bai-
ley.” He carried on, in the pages of Rhodora which he edited, a
spirited fight with Gleason of the New York Botanical Garden on
the classification of raspberries, finally adding to one of Gleason’s
own papers in Rhodora a footnote to the effect that he would no

196 Rhodora [Vol. 88

longer try to set Gleason right because he did not have any time for
“more Bronxian sophistries.”

His lectures were unorthodox to say the least, interspersed with
constant references to piles of herbarium specimens. He had man-
nerisms which he used as punctuation for his lecturing. Rising on his
toes, for example, meant special emphasis—“italics,” if you will.
Stretching out both hands, palms upward, meant a question mark.
A vigorous slap on his bald head was an exclamation point. But
perhaps the idiosyncracy that I most vividly remember about this
outstanding teacher and botanist was his method of writing and
editing his own papers. He wrote always, seated at a desk in the
public part of the library and not in his own office, if indeed he had
one. He used a thick carpenter’s pencil and long sheets of plant
collecting paper. Insertions, corrections, arrows for additional
paragraphs, scratched-out phrases—all of this was such a mess that
only one secretary who had typed his material for years could
decipher the manuscript.

He always wore thick heavy-duty shoes. Often, when an unsus-
pecting graduate student would pass by, out from under Fernald’s
table would come a leg. The heavily shod foot would stamp on the
floor, there would be a clearing of the throat, and Fernald would
say, “Well, Schultes, what do you think of this?” He would then
read the whole manuscript in the greatest detail to the trapped
student, adding copious ad-lib explanations, stopping to write in
corrections, add or delete phrases. He thoroughly enjoyed this pas-
time and interspersed his reading with an occasional chuckle of
satisfaction. And I must say that, while at the time we all thought
that it was an imposition on Fernald’s part to trespass on what we
thought was our valuable time, I realize how much botanical mate-
rial I learned and how much I absorbed about writing manuscripts
from this unusual custom.

Now, before ending these reminiscences that could go on and on
and on, | must mention Fernald’s field trips. They were superb, but
there is one in particular that I will never forget. On one Saturday in
late October, we were to drive to the tip of Cape Cod. We would
stay Overnight in Provincetown in a beach house, and Fernald told
us not to take any food for he would cook up a dinner of clams and
beach spinach. I had offered to drive. The day dawned with torren-

1986] NEBC 800th meeting — Remarks 197

tial winds and rains, the backlash of a hurricane. Cambridge streets
were deep with water; branches of trees were falling. Elso Barg-
hoorn, who was to ride with me, was certain that the trip would be
postponed. When we arrived at the Herbarium, Elso suggestively
volunteered to Fernald that “it was raining” (laughter). But Fernald
chuckled and replied, “Well, the plants are still there, aren’t they?”
And we all went. But since we had had a frost, I felt that our chances
of collecting sea spinach were not too bright, so I bought and
secreted in the glove compartment several chocolate bars just in case
of an emergency. We did find sea spinach but all the leaves had
fallen off. Undaunted, Fernald collected an armful, proceeded to
boil it and serve it, stuffing the twigs into his mouth and trying to
convince all of us that it was a delicious repast (laughter). But,
before going to bed that evening, I sneaked out to the car and
devoured several chocolate bars. Fernald never knew how little |
appreciated his economic botany on that trip! (laughter).

Over the years, | have at so many times realized how fortunate |
was to have Merritt Fernald as a teacher and to have studied with
him. I’m glad that in this 800th meeting of the New England Botani-
cal Club, of which he was such a dynamo, some of us who were so
fortunate have shared with all of you what we remember of our days
with, and exposure to, this great giant of North American botany.

R. A. Howarb: Thank you very much, Dick. | am sure these
stories have brought to mind others that some of you have. For
those of you who have sent material in to us, add to it, if you come
to the thoughts. The hour is late so we won’t ask for more contribu-
tions now. We intended this program as a tribute to M. L. Fernald.
As I said at the beginning, | hope those of you who did not know
him, to whom he is only a name, have gathered from our comments
that he was really quite a person. Thank you very much.

Mary WALKER: Back in the Council minutes of late 1982, early
1983, there used to appear, “The 800th meeting is coming in May
1984. Are we going to do anything about it?” Thanks to the efforts
of Garrett Crow, who has done a lot of the organization, and to Dr.
Howard, who has been a wonderful Master of Ceremonies, and to
all of our speakers who’ve given us this picture of Fernald, I think

198 Rhodora [Vol. 88

we have done something about it. So, | wish to thank you all once
more.
May I have a motion to adjourn the 800th meeting?

[It was so moved, at 11:25 p.m. —Ed.]

LETTERS AND EXCERPTS ALSO RECEIVED FOR THE
“TRIBUTE TO FERNALD” EVENING

from Ernst C. Abbe:

As I look back on that golden age of graduate work at Harvard in
the °30’s, | well remember my desire to do well in Fernald’s eyes and
the strange twist of fate which led me to differ with him so pro-
foundly. Although my main concern was morphology, I was drawn
to the Gray repeatedly from 1930 to 1949.

The number of anecdotes growing out of those years is endless
and each one illuminates another facet of Fernald’s complex and
charismatic personality which tolerated no neutral onlookers: it
seems that all who were associated with him developed a love-hate
relationship. For instance, his sturdy New England egalitarianism is
so well illustrated by his response to Ms. McKelvey’s complaint that
she had to sleep under the same roof as her chauffeur on the Cape
Cod field trip in the fall of 1930. Fernald’s answer was that not only
would he not hesitate to sleep under the same roof but would even
sleep in the same bed with her chauffeur if necessary, thus leaving
Ms. McKelvey to decide to what extent she might wish to emulate
Fernald in this specific instance. Then there was the enormous scorn
he could heap on an opponent as in the case of Professor Jeffrey’s
“absent treatment hybrids” whose parent species were so far apart
that it would take several relays of devoted bees to effect cross-
pollination. Or his absolute fairness in sponsoring my Guggenheim
even though he knew that my collections represented a test of his
cherished Persistence Theory. Nor will I forget my surprise when he
upbraided me after our return from Virginia for having persuaded
him to buy a Virginia “peanut-fed” ham which had ruined his repu-
tation as a cook because it retained its mahogany-like consistency
after hours of cooking.

1986] NEBC 800th meeting — Letters 199

Fernald’s violent reaction to the manuscript of my paper on north-
eastern Labrador was the only really traumatic experience I had
with him, but it could not be avoided—and because | valued his
good opinion it was profoundly disruptive of my peace of mind and
diverted my activities for several years from my major area of
research. Also, a third-hand version of the affair has appeared
briefly in print so that perhaps a first-hand account is in order.

It all centered around Fernald’s Persistence Theory of which I
first heard in Wiegand’s taxonomy seminar at Cornell in which it
was treated as revealed truth and accepted by me as such. So when
Fernald held forth on it after I went to Harvard, | nodded my head
sagely and continued to accept it without question. At this time
Alexander Forbes of the Naushon Forbeses and the Harvard Medical
School asked Fernald to send one of his students on Forbes’ expe-
dition to Northern Labrador, provided such a student had a strong
back and could handle the ropes on the expedition’s Lunenburg
schooner; Fernald recommended me. Fernald told me he wanted me
to go because of my previous collecting experience on the North
Shore of the Gulf of St. Lawrence and because he wanted me to get
as many collections as possible from the tops of the Torngat and
Kaumajet Mountains which he was convinced were former nuna-
taks and should have many “relicts” waiting to be collected. He felt
that there was an excellent chance of getting to such nunatak areas
because the famous mountain climber and glacial geologist Noel
Odell was to be a member of Forbes’ expedition and had specifically
asked Forbes to add one of Fernald’s students to the personnel of
the expedition. Fernald pointed out to me that his graduate students
could not spare the time from their monographic work to do floris-
tic work of this sort, but that he would like to have me serve as
botanist on the expedition. I happily agreed to go and was vastly
flattered in the bargain, although I disliked heights and mountain
climbing; by following closely in the footsteps of Odell, I climbed a
number of mountains in northeastern Labrador. But on mountain-
top after mountaintop the plants turned out to be the ubiquitous
arctic species, not the “relicts” that were expected. It was at the
lower elevations where moisture and rock type were favorable that
the significant rarities occurred. To add to my disillusionment,
Odell was convinced on geological grounds that the highest peaks
had been ice-covered so that there could not have been any nuna-

200 Rhodora [Vol. 88

taks where “relicts” could have survived. In my opinion even if there
had been nunataks, the conditions would have been so inhospitable
that only the hardiest plants could have lived on them, and that it
was the ice-free forelands, if any existed, which might have sup-
ported the rarer plants. Yet here again Odell felt that such areas had
not existed. If Odell’s views were to be accepted, only one conclu-
sion was possible; every species, “depauperate relicts” included, had
to have migrated into the area in post-glacial time—all of these
views were sure to be abhorrent to Fernald, especially the last. Yet I
could not, in all honesty, avoid these conclusions. Various good
reasons prevented a confrontation at this time—a summer in north-
ern British Columbia with the Raups, intensive work on my thesis
with Wetmore, an NRC post-doc at Columbia with Sinnott. The
latter worked out such that I was able to spend the spring of 1935 at
Harvard again and concentrate on writing up the Torngat paper, the
painful typing of which I completed just before leaving for the Uni-
versity of Minnesota.

I took the finished manuscript of the Torngat paper to Fernald in
mid-afternoon the day before we were to leave town. Fernald sat
down with me at one of the tables in the library of the old Gray on
Garden Street and looked over the illustrations, which he liked, and
then began to read the manuscript about which he was enthusiastic,
—at first. In fact, he went off and found Weatherby somewhere and
read excerpts to him and said, “This has to be published as a Con-
tribution,” whereupon Weatherby beamed benignly, and went on
his way. The afternoon wore along, Miss Sanderson left, and soon
Fernald and | were alone and nearly through with the manuscript.
At this time Fernald came to the discussion summarized above and
immediately began to fulminate. He brought up all the information
and arguments that he had so carefully worked out for the “Persist-
ence” monograph and subsequent papers. The vast mass of detail,
the irrefutable logic were overwhelming, but I told him I could not
accept an extension of his conclusions to the area I was familiar
with, and furthermore Odell’s independent conclusions about the
glacial geology fortified my views. Fernald demanded an explana-
tion of what kind of an authority this man Odell might be, what did
he know about roches moutounees, etc., etc. | pointed out that I had
allowed for Odell’s interpretation being incorrect and that even if
there had been nunataks, etc., | was driven to conclude, against all
my earlier preconceptions, that the Persistence Theory was inade-

1986] NEBC 800th meeting — Letters 201

quate, that there had to be extensive post-glacial migrations. Fer-
nald stated that as Editor-in-Chief of Rhodora it was one of his
perquisites to insert footnotes wherever he considered that the text
of a paper needed correction. | agreed, but stated, perhaps a bit
coldly, that as a member of the New England Botanical Club it was
my privilege to submit the manuscript for publication (in retrospect,
a rather weak rejoinder on my part!). Fernald conceded this point
and then launched into a thorough dissection of my personality
(always had been a bumptious person ever since I first came to
Harvard, etc.). His enthusiasm mounted with this sally and he
recalled having known my father (Harvard, 1894) and proceeded to
dissect father’s character, scattered it in bits on the floor of the
library and figuratively did an Indian dance on the quivering
remains. Finally he did the same for my grandfather (Harvard,
1864) which showed a high degree of ingenuity, because my grand-
father was at Harvard roughly thirty years before Fernald came as a
student! While I was pointing out that such character analysis has
no bearing on the matter under discussion, the phone rang. It was
Fernald’s daughter reminding him that it was 7:30, dinner was get-
ting cold, and should she hold it any longer? Whereupon Fernald
took the manuscript and left for home without a fare-you-well! |
was exhausted and furious.

The subsequent events have always amazed me; in retrospect |
feel that Weatherby as usual had an ameliorating effect on Fernald’s
often drastic inclinations. For, a few weeks after I arrived in Minne-
sota, I received a very moderate letter from Fernald as Editor of
Rhodora saying he would publish the paper and recommending that
I change “northern Labrador” to “northeastern Labrador” in a
number of places in the text. Then, to my vast surprise, he published
it as a Contribution!

from George L. Church:

A man with a spring in his step and a lively manner entered the
laboratory and proceeded briskly to make the pile of herbarium
specimens before him seem to come alive. Indeed, as I listened to
Professor Fernald in his class in 1925, he was unfolding a vivid
picture of the natural community in which each specimen had been
collected. Furthermore, I soon realized that the enthusiasm in his

202 Rhodora [Vol. 88

discourse had been well nurtured by more than three decades of
meticulous studies of plant populations in a host of areas in New
England and eastern Canada.

As I delved into the early volumes of Rhodora, I continued to be
enthused with Fernald in the accounts of his many expeditions to
such alluring natural areas as the St. John river drainage system, the
Matapedia Valley, or the Gaspé Peninsula. A few years later, when
I myself had the opportunity to visit some of these choice collecting
places, I felt as though I had really been there earlier with M. L. F.

Although we were left to our own devices in laboratory study, we
were always free to interrupt Fernald for consultation in the Herba-
rium library where we found him vigorously writing. Before answer-
ing a question, he would often look up over his half-frame spectacles
and say, “Listen to this!” I recall particularly the time when the well
known volume entitled “Age and Area” by Willis appeared. With a
wealth of knowledge of the flora of unglaciated North America,
Fernald could easily refute Willis’ theory that the oldest species had
the widest geographical range. I can still hear him exclaim gleefully
as he looked up at us again after his demolition of Willis, —‘*Willis
should have called his work AGE AND ERROR!”

In 1925, the New England Botanical Club used to meet in the
imposing building of the American Academy of Arts and Sciences
on Newbury Street in Boston. It was at the monthly meetings there
that I enjoyed Fernald at his best as a colorful raconteur. With great
delight, he would relate his experience as a camper, mountain
climber, and collector in Maine or the Maritime Provinces. He took
great pride in his ability to explore an area in spite of such interfer-
ences as the ubiquitous, invisible mosquitoes of the northern wood-
lands or the wild dog packs of the serpentine barrens of Newfound-
land.

Certainly, hazards of field work in no way impeded Fernald in
pursuit of adequate data in support of his well known concepts of
distribution.

In fact, whenever I recognize herbarium specimens bearing anno-
tations in a clear, bold handwriting with the initials M. L. F., I relive
the excitement of following a trail blazed by Merritt Lyndon
Fernald.

1986] NEBC 800th meeting — Letters 203

from Lincoln Constance:

Out of the sea of recollections of Fernald that come back to me, |
shall confine myself to three.

I first visited Cambridge in 1942, just after Reed Rollins had
accepted an appointment to the faculty of Stanford University, and
asked Professor Fernald if he was not as greatly pleased as I was. He
looked at me quizzically for a moment and then snapped, “They’re
pretty radical out there!” So much for Clausen, Keck and Hiesey,
and what Stafleu calls “the Birth of Biosystematics....”

Sometime during my stay, I encountered a letter Fernald had
written to one of his cronies (perhaps Bayard Long?), complaining
that one of Fernald’s students had been required to take a course in
Physiology, “Whereas you and | know that the best training for a
taxonomist is a thorough grounding in Latin.”

At Fernald’s request, a meeting was held in the Director’s office
with officials of the America Book Company, a representative from
the Harvard Treasurer’s office, Professor I. W. Bailey and myself, to
see if we could speed up completion and publication of the Manual.
I told the group that so far as I could tell, the ms. had reached the
point where commencing to print the first part would ensure prompt
completion of the remainder, and it seemed that the company would
be agreeable to this interpretation. At this point, however, all other
considerations became secondary to the somewhat complicated
question of royalties. The company representative professed to be
overwhelmed by what he regarded as Fernald’s exorbitant demands,
and asked if he might smoke. Fernald, a vociferous anti-tobacconist,
was more than agreeable. I’m not sure whether he got up and lighted
the gentleman’s cigar, or did I only imagine it? (The “agreement”
came unstuck and Reed had to renegotiate it a few years later).

Finally, | had a story from Henry A. Gleason, who visited Cam-
bridge that year to serve on an appointment committee. He related
that when he visited Elmer D. Merrill at the Arboretum, Merrill was
chuckling and said, “You remember that row of moth-eaten,
broken-down elms around the Gray Herbarium? | had them cut
down!” When Gleason called on Fernald, he was fuming, “Do you
know what that fool Merrill did? He cut down the elms that Asa
Gray planted!”

204 Rhodora [Vol. 88

from William B. Drew:

I first met Prof. Fernald in the fall of 1930 when I enrolled in
Botany 7. At that time Botany met in the so-called Harvard Room,
a classroom in the Gray Herbarium. Prof. Fernald entered the
room, his beard neatly trimmed, and proceeded to begin his lecture
with the gymnosperms. Those of you who took his course may recall
that it was largely based upon a key to the families of the gymno-
sperms and angiosperms with, of course, frequent references to some
genera and species as he went along. He had a very lucid style of
lecturing which make for interesting listening. From time to time he
told us anecdotes of one sort or another.

One of the stories that was circulating around the Gray Herba-
rium at that time, I think perhaps told by Prof. Fernald on himself,
was that he had never been able to pass an automobile driver’s test.
He tried it a few times, but apparently went off the road when he
saw some interesting plant. He also averred that, as a younger man,
or perhaps a youth in Maine, he had tried collecting from a bicycle,
and he had a similar fate there. After two or three nasty spills, I
guess he probably decided to walk. At any rate, his daughter Kathe-
rine was providing the transportation to and from his home and the
Gray Herbarium during the °30’s when I was associated with him.

Another story, which perhaps may be apocryphal but as far as I
know it’s true, was told of Dr. Norman C. Fassett, one of Prof.
Fernald’s outstanding graduates, who was enroute to accept the
position at the University of Wisconsin, where he was an outstand-
ing taxonomist. It seems that Dr. Fassett and his family had to be
there in a hurry and therefore, with two or more children to cope
with, they cut holes in the wooden cover under the seat of the Model
T Ford to allow for an essentially nonstop trip, at least as far as the
toilet needs of the youngsters were concerned. Later, in 1944, |
shared briefly an apartment in Bogota, Colombia, with Dr. Fassett
and he indicated that the story was essentially true. Another anec-
dote about Prof. Dr. Fernald, if it can be called that, was his ten-
dency to be very careful with specimens on display or with any of his
large sheets of newsprint on which he wrote in such a beautiful,
flowing hand. He felt he had to be careful when certain botanists
visited the Gray Herbarium because allegedly at least one botanist
had learned through talking with Prof. Fernald, or reading his
material, about a new species which he was about to describe.

1986] NEBC 800th meeting — Letters 205

Apparently his species was scooped by this visiting botanist. So
whenever this particular individual, a very prominent botanist of the
northeast of North America, visited the Gray Herbarium, Prof.
Fernald was always very careful to see that the specimens on which
he was working were covered up and his most recent writing hidden
under some neighboring papers or specimens.

Another point that I might add was the perspective of Prof.
Fernald from the standpoint of a graduate student working under
his direction. One had to be very discreet about approaching him,
particularly when he was busily writing. | generally found it neces-
sary to make an appointment first thing in the morning when he
came in, and then, if | had good luck, | would approach him with
whatever material and questions I had in mind at the appointed
hour. Usually, if he was really steamed up about a topic, as he
frequently was, I listened to his latest writing for a considerable
period of time before we got down to cases on my own questions. As
a result, | heard a good bit of the linear-leaved North American
species of Potamogeton, published in 1932, and especially recent
discoveries in the Newfoundland flora, published in 1933. Those
were interesting papers, but it became a problem sometimes, when l
had pressing matters of my own on which I needed answers.

I recall Prof. Fernald in the mid-1940’s getting together with Lud-
low Griscom, the famous ornithologist, to go over collections they
had jointly made in the coastal plain or piedmont of Virginia. They
would work over the identifications on which Mr. Griscom was an
expert. He was particularly interested, as I recall, in carexes, but he
and Prof. Fernald would debate and study the material and reach
some conclusion. This usually occured late in an afternoon when
Prof. Griscom’s work at the museum was done for the day and he
came up and worked for an hour or two or more with Prof. Fernald
in the main gallery of the Gray Herbarium. The Virginia papers, as
far as the coastal plain was concerned, were one paper published in
1937.

Finally, | might add that Prof. Fernald was a marvelous person
with whom to go on a field trip, and each spring his Botany 7 class
made an expedition to some interesting botanical spot, such as
limestone areas in southern Vermont. There we spent a long week-
end learning a great deal about the flora of eastern North America
and particularly of that region. One other thought occurred to me:
Prof. Fernald stood up for his graduate students whenever they

206 Rhodora [Vol. 88

became involved with other members of the Harvard graduate
faculty. I recall a personal experience when I was invited to present
a seminar on my studies of the section Batrachium of the genus
Ranunculus before the group in the Biological Science building. |
hadn't told Prof. Fernald that | was going to do this, as I didn't
think he would want to be bothered or would have much interest in
it. When I presented my paper I discussed at some length the general
concept of Ranunculus and the subgeneric status of Batrachium.
This matter intrigued Prof. G. H. Parker, the eminent zoologist, and
he gave me a pretty hard time. I was a very callow youth at the time
anyway, so I did the best I could, and I think I came fairly close to
holding my own. Well, some time later, perhaps the next day, Prof.
Fernald accosted me in the Gray Herbarium and said why didn’t I
tell him I was going to give the seminar? He would have been there
to lend me support. Believe me, | appreciated that, so when I had
my final oral examination, the committee included Prof. Parker and
several other zoologists and botanists; | had no fears about Prof.
Parker that time, and much to my astonishment, | finally passed it.
Thank you; I hope you'll find this interesting.

from Joseph Ewan:

What I did not stress in my sketch of Professor Fernald in the
Dictionary of Scientific Biography is an impression which still per-
sists as a marcescent memory. It is of a large head atop a small man
housing a winning epicentricity. On my 1945 visit to the old “Gray”
on Garden Street I had sallied forth from a pinched second floor
room on Brattle Street to meet in a fertile field, the seat of historic
specimens, frames of reference, footnoted by a unique library. Rec-
ognizing a willing audience, Fernald led me to an alcove and
recommended that I listen to a few paragraphs. Thereupon, with the
obvious delight born of years of observing and collecting botanists,
he read a critique which was to bloom in the next anthesis of Rho-
dora. Here a Fernald verdict was unabashedly given. Fernald was a
holotype. Of quiet but unmistakable speech, he was sincerely inter-
ested in the response, especially if it happened to be a contrary view.
One was aware that this was Fernald at Harvard, clearly here
Fernald was the epicenter.

1986] NEBC 800th meeting — Letters 207

from Richard H. Goodwin:

My first encounter with Professor Fernald was when I took
Botany 7 with him in my sophomore year in the fall of 1930. The
initial lesson was one in local geography, as I arrived breathless at
the junction of Garden and Linnaean Streets with enough sweat
pouring off my right wrist to make note taking a serious problem.
Not being in shape to run a five-minute mile between classes, |
managed to borrow a friend’s car for the times when I had this tight
schedule. Those were the days when one could find a place to park
in Cambridge.

The course was filled with graduate students and upperclassmen,
who had far more background and maturity than I. Fernald’s lec-
tures would frequently begin with a verbal recitation of a section in
the “Key to the Families” such as: “e. Flowers unisexual, mono-
ecious, in dense spikes or heads; perianth of hair-like bristles... Ty-
phaceae.” Then would follow an interesting discourse on the
edibility of cattail rootstalks and flowering spikes, or, for the
appropriate taxon, a discussion of the phytogeographical implica-
tions of its distribution.

Later on, as a graduate student, I signed up for an advanced
course with Professor Fernald. | was the only Harvard registrant,
there were three from Radcliffe. So, strictly off the record —this was
about 1934—the course was given for the four of us, who may have
participated in the first “integrated” course at Harvard.

I can remember more than once being cornered in the Herbarium
by the Professor who, with sparkling eyes, would try out a draft of
his latest manuscript on a willing listener.

In retrospect, Botany 7 and my only too brief contacts with Fer-
nald were highly significant in shaping my career as a botanist. This
was surely not due to the very modest amount of information
retained from his lectures, nor to the proficiency attained in the use
of the drawing pen. Rather, it grew from an intangible attunement
to the natural world. Fernald was at his best in the field, and the
forays with him to Mt. Equinox and Fairlee, Vermont, and to Cape
Cod are still vividly remembered. They were the first of many to
follow on my own, and with students and colleagues.

208 Rhodora [Vol. 88
from Hiroshi Hara:

From August 1938 to November 1940, I was a Research Fellow at
the Gray Herbarium situated at that time in the Gray Botanic
Garden. The main subject of my research was the botanical relation-
ship between Eastern North America and Eastern Asia, which was
first pointed out by Prof. A. Gray in 1846, and in which Prof.
Fernald was also much interested. I studied mainly under Prof.
Fernald at the Gray Herbarium, and also visited the Arnold Arbore-
tum once a week to see specimens and literature there and to talk
with Prof. E. D. Merrill and Dr. A. Rehder, both of whom had
profound knowledge of Asiatic plants. In those days, Prof. Fernald
had been working hard to revise Gray’s Manual till late evening, and
also preparing additions to the flora of Virginia to be published in
Rhodora. My room was upstairs, and when I came downstairs in
order to check literature at late hours, I often saw Prof. Fernald
alone working at his large desk by the side of book-shelves of the
library, and heaping up specimens and books high on his desk. So |
had chances to talk with Prof. Fernald on various problems. Soon |]
published a short paper in Rhodora 41: 385-392 (1939). Regarding
the Asiatic plants newly found in North America, we intimately
collaborated, for example Cyperus microiria Steud. by Hara in
Rhodora 42: 196 (1940), and Aneilema Keisak Hassk. by Fernald in
Rhodora 42: 441 (1940).

When I told Prof. Fernald that I was looking for living material
of Phryma Leptostachya of North America, he at once kindly sug-
gested that he would take me to the place where the plant was
growing wild. On one Sunday morning, | dropped in Prof. Fernald’s
house, and drove my car to the suburbs with the Professor and his
daughter on the back seat. He guided me nicely to the place, and |
could successfully collect Phryma. Mainly based on this collection, I
later revised the North American and Asiatic plants of the genus
Phryma.

At the end of my stay I left a typewritten copy of a list of the
plants common to Japan and North America in the hands of Prof.
Fernald, and it is now preserved in the library of the Harvard Her-
baria. I brought back to Japan some seeds of Veroniscastrum virgi-
nicum (L.) Farwell from the Gray botanic garden, and the plants

1986] NEBC 800th meeting — Letters 209

raised from the seeds are still vigorously growing in my garden in
Tokyo. They are not only one of my best memorial plants in the
garden, but also they may possibly be the only living plant which
has hitherto been continuously grown from the old Gray garden.

My knowledge of the North American plants obtained during my
stay at Cambridge is really very useful for my further study on
critical comparison between closely allied taxa of North America
and Eastern Asia. These data were mostly arranged in order in my
later papers such as ‘Contributions to the study of variations in the
Japanese plants closely related to those of Europe and North Amer-
ica’ Part | and 2 (1952 & 56), and others.

In my memory, Prof. Fernald is still living as my dear father as
well as my respected teacher. | will never forget his mild face, but
with keen eyes.

from Quentin Jones:

In the Fall of 1950, Professor Fernald was reading page proof of
the 8th Edition. In fact, he was reading it to anyone that happened
to cross the foyer of the old Gray Herbarium at the wrong time.
Sometimes these “readings” lasted more than an hour. I was so
favored on several occasions.

One occasion stands out in my memory because Professor Fer-
nald got so much enjoyment out of the anecdote he told me. It seems
that during his work on the Gaspé Peninsula he had a French-
Canadian field assistant with the family name Assino, who kept
needling Fernald to name a new species after him. Fernald never
yielded to his pleading, “Because,” he said, “can you see in print a
specific epithet, Assinoanus?”

from Alexander Lincoln, Jr.:

As a student in Professor Fernald’s Botany 7 and Botany 10
courses in the early 1930’s, | came to depend on my hurried walks up
Garden Street to the Gray Herbarium of those days, set in the sunny
Botanic Garden, as a very different atmosphere from the general
drabness that tended to prevail elsewhere in Cambridge.

210 Rhodora [Vol. 88

Botany 7 was handled in what, by contrast to the minutiae of
microscopic peerings in other courses, achieved a grander scale:
whole plants to be not only seen, but actually enjoyed; the tides of
successive floras sweeping across the northern hemisphere, leaving
scattered pockets of relict species and endemics: frequent allusion to
local plant names and to plant uses of recent years if not still today,
with particular emphasis on French Canada. Then, in Botany 10,
Opportunity to examine handsomely illustrated old volumes, them-
selves floras in a different sense, and to come to know more about
early botanists and their travels, discoveries and conclusions.

Professor Fernald’s classroom presence conveyed animation,
deep interest and solid scholarship of the principles of systematic
botany, along with sonorous, good-natured commentary from his
wide range of field experience, all from a twinkley-eyed, somewhat
rotund Santa Claus figure. His full gray beard, in colorful contrast
with his often pink dome, underwent obvious change following a
trim every couple of weeks or so. Originally opposed as I was to
burdening our native plants with Latinized nomenclature, I came to
enjoy the very sound of their names as pronounced by Professor
Fernald, Professor Pease and others. Further, it soon grew apparent
that Latin names provided a far superior method of distinguishing
between species despite—over subsequent years —the annoyance of
frequent name changes as dictated by botanical congresses.

It was always a pleasure to accompany Professor Fernald on a
field trip, whether largely to collect plants or simply to learn more
about them in their natural habitat. Especially on trips to Cape Cod,
we were encouraged to taste them. Even in class, he sometimes
brought in for our appraisal samples such as muffins he had made
from acorns of white oak.

One mishap that occurred, as related to me later, interrupted the
usual flow of an October outing. The professor, a non-swimmer,
stepped off the end of a wall or low dam onto what appeared to bea
leaf-strewn flat. But the leaves concealed instead deep water. He
submerged to his chin before being helped out.

Professor Fernald remains a particularly helpful and influential
star in the constellation of teachers | experienced at Harvard.

1986] NEBC 800th meeting — Letters pad

from Wayne E. Manning:

I first knew about Professor Fernald in the use of the 7th edition
of Gray’s Manual, Professor Wiegand at Cornell told us about him
in taxonomy Classes.

| first met Professor Fernald at the International Botanical Con-
gress at Ithaca, NY in 1926. I recall that someone foolishly asked him
whether he knew the name of a certain plant; of course he knew.

On the field trip in taxonomy the summer of 1926 I showed
Professor Fernald the staminate flower buds of Vallisneria spiralis;
he had never seen these. I had known them from collections of water
plants for U.S. Biological Survey in Minnesota in 1923.

On a family trip through northern Maine in 1934, I collected a
small-flowered waterlily and could not name it. On a trip to Har-
vard the following year | showed the specimen to Professor Fernald.
He said that it was Nymphaea tetragona, not mentioned in the &th
edition of Gray’s Manual. Then he said that he would tell me a
story. He told me that Cyrus Pringle had collected a small-flowered
waterlily in Maine in the 1870's, but the collection was lost. My
collection was the first definite record for eastern U.S. Professor
Fernald very kindly added a footnote to my report of the collection
in the November 1936 issue of Rhodora.

My trips for research on the Juglandaceae in the library and
herbarium at the Garden Street Gray Herbarium were limited in
time, no more than 5 or 6 hours in any one day trip. It was a little bit
exasperating to be asked by Professor Fernald to listen to his read-
ing of his manuscripts, often critical comments about the publica-
tion of some botanical author. But of course it was pleasing to have
4a man of Professor Fernald’s caliber to take notice of me, and |
never left him until he was finished.

1 knew that Professor Fernald was working on the 8th edition of
Gray's Manual. On one of my trips to the Gray Herbarium in the
late 1940’s | asked Professor Fernald if I could see the manuscript of
Juglandaceae. I realized at once that there were a large number of
errors, but knowing his reaction to criticism by others, I said
nothing. But that day I brought to him a number of specimens in
“his” herbarium that meant that changes were necessary. Finally he
asked me to take the copy of the mss. home to study and check over.

212 Rhodora [Vol. 88

Of course that was just what I wanted, but now I had responsibility.
I wrote out several pages of corrections of the mss., and finally
dared to incorporate all of these in a corrected mss. to send to him. I
Was uncertain about his reaction. A short time later I received a
postcard, saying: “I have spent a pleasant afternoon looking over
the corrected mss. of the Juglandaceae you sent me. I am glad that
someone understands the hickories.” The material was published in
the manual as I had written it and he kindly published a footnote in
the manual that I had helped him with aid in the work on the family.
My friends asked me how I had managed to have my name in Gray's
Manual. | told them it was by hard work and good politics.

A short time later I sent Professor Fernald a mss. on the genus
Carya for publishing in Rhodora. He realized that this involved
changes for Carya in the manual—I had not dared to incorporate
these in the Gray's Manual mss.—and he withheld publication of
my material until after the publication of the manual.

I cherish very much my close connections with Professor Fernald.

from Eugene Ogden:

I did graduate work with Professor Fernald 1934- 1938. That was
a very pleasant time for me, spending most of my time in the Gray
Herbarium in the Harvard Botanical Garden.

I first met Prof. Fernald in July 1933 at meetings of the Josselyn
Botanical Society at Winthrop, Maine where he invited me to con-
tinue botanical studies under his direction at Harvard and serve as
Gray Herbarium photographer. During the four years there, he
introduced me to many distinguished visiting botanists offering the
Gray Herbarium photography services. During the fourth year it
was my great privilege to be his laboratory assistant in plant taxon-
omy for Harvard and Radcliffe. In those days the Harvard boys and
Radcliffe girls were in separate laboratories and on separate field
trips. It was obvious that Prof. Fernald greatly enjoyed the two class
trips to Cape Cod each October and the trips to Vermont each
spring.

One of my fond memories is a brief trip with Prof. Fernald and
his son, Henry, to the Catskill Mountains of New York in August
1935, presumably to look for nunataks, but I think Prof. Fernald
had become a bit weary of herbarium work and needed some relaxa-

1986] NEBC 800th meeting — Letters 213

tion in the field. We found no nunataks. The car broke down, so we
took rooms in an inn and that evening regaled each other with jokes.
Prof. Fernald was partial to the pun type and supplied some clever
ones. After the broken axle was replaced we made some collections,
including a Gray Exsiccatae set of Agrostis perennans. On the way
back to Cambridge we explored a small pond at Salisbury, Connec-
ticut, for Potamogeton longiligulatus. | put on my swim trunks, but
Prof. Fernald waded into the mucky water without removing coat
and vest, the lower parts of which were submerged.

We didn’t find the pondweed we were looking for. Prof. Fernald
replaced his wet trousers with a dry pair, and we returned to
Cambridge.

from Lilly M. Perry:

Dr. A. Lawrence Lowell once said, “What is there about the little
flowers that makes the botanists quarrel with each other?” It was
Professor Fernald who first mentioned to me that Dr. Robinson
suggested before all his able students were placed, that they wanted a
helper at the Gray Herbarium, and then he said if I were interested,
to go see Dr. Robinson. Thus began my years at Harvard. As I was
a special student of Professor E. C. Jeffrey, as soon as I returned
from Fernald’s taxonomic class, | have never forgotten Jeffrey’s
reply, “I have known Benjamin Robinson as a colleague for 23 years
and I have never known him when he wasn’t a gentleman, but be
wary of the little man whose forehead reaches to the back of his
neck.” Before I took his class, former students had warned me to
keep on his good side.

Professor Fernald was a brilliant writer, so much so that when he
published narratives of his expeditions, many copies of these (so the
librarian told me) were regularly bought by quite a number of peo-
ple who had no particular interest in botany but valued them as
good short story literature.

If you happened to drop in at the herbarium during the week, a
familiar sight in our corner of the building would be Prof. Fernald
reading a ms. to some person. Dr. Johnston used to say, “Don’t be
flattered, he’s only polishing his ms. When I polish mine, I take it
out the back door and read it aloud.”

214 Rhodora [Vol. 88

In all the years I put in at the Gray Herbarium, we clashed only
once and then it was over a remark | thoughtlessly made. One
Saturday afternoon I went to the herbarium to work on my
research. Who should show up as I got nicely settled but M. L. F.
He came over where I was and said, “Would you help me pack my
trunk for Newfoundland?” So I went with him to the closet in the
lab where both trunk and clothes were. Many times I had packed
my own, always putting the delicate things in the top tray. I didn’t
even stop to think how he would pack his. After his personal things
were tidily packed in the bottom he put in the tray. He had left his
collecting materials for the tray. He had a pair of hip boots, vascu-
lum, etc. I said where are you going to put your 7-league boots? Like
a flash he said, “Don’t be so personal,” got up and left. I felt terrible,
as I had always called those hip boots 7-league boots, but when I
went to the main room he was gone. When I went to work Monday,
the trunk was ready to ship.

Professor Fernald taught me the basis of herbarium work. (Dr.
Robinson taught me how to use the library.) We started with Old
World Juncus and Buchenov’s work as Fernald was working on his
Newfoundland collections and he wanted to see what was the rela-
tionship of the northern Old and New World species. He taught me
how to compare literature and plants. You remember Jeffrey called
him a little man. I was the shortest person on the staff and he was
next. So many times when I worked with him, it happened that the
plants he wanted to see (when we were working together) were in the
top pigeonholes. I had to step up on the base (four inches) of the
case, then hold on to the divider of the pigeonholes and reach up
with my other hand for the covers in the top, he could just reach
them comfortably; every time he asked me which one would get the
plants up there, but always he got them. | put them away after he
had seen what he wanted to know. One day I was working on the
second balcony. Fernald came in with Rendle from the British
Museum, who was spelling out for Fernald a scientific name ending
in zia. The English “zed” Fernald didn’t seem to understand, so |
said in a very low voice “Zed is our zee.” Fernald turned at once and
brought the specimen he wanted.

Fernald gave what we students called the Gossip Course. It was
said that Asa Gray went abroad to study Botany, but his wife was
interested in botanists. Anyhow, her results were in four big folios
all wrapped in gray flannel covers and kept under lock and key.

1986] NEBC 800th meeting — Letters Paes

They contained photographs of botanists and a full letter with salu-
tation and signature (very important). Fernald chose from these the
people he thought would be of value to us and told us what he
thought would be of interest. | may say not all were Europeans. |
remember two in particular: Britton, Director of the New York
Botanical Garden, who was an able man; and Brandegee from Cali-
fornia, although I understand he grew up in Brookline or its vicin-
ity. | think he must have been a small man from what follows: One
afternoon a large woman three or four steps ahead of a man came
through the inner door and asked if this was the Gray Herbarium.
On being told that it was, she turned around and said, “This is the
place; come on, Dolly,” to the man.

When Harvard had its 300th anniversary, he came into the library
and said to me, “They only gave me one ticket for this afternoon.
Shall I give it to you or Miss Stone?” I said, “Miss Stone, by all
means. She’s a regular staff member; I’m only here for the summer.”
| added it was probably only meant for a “Harvard man.” His
answer was, “You're as good a Harvard man as I know.”

mm

from Hugh M. Raup:

My earliest impressions of Professor Fernald came while I was in
graduate school in the mid-1920’s and read some of his famous
reviews. I did not know enough to understand and appreciate all the
things he was writing about, but 1 do remember thinking that he
really didn’t need to be quite so vitriolic in his denunciations. These
impressions made me rather timorous when I wrote to him in the
spring of 1929 asking whether I would be welcomed if I came to
study at the Gray Herbarium for a year beginning in the following
autumn. I also inquired about some part-time assistantship that
would help to pay expenses.

My wife and I had by that time accumulated a sizeable pile of
specimens from three summers’ plant collecting in northwestern
Canada. By the fall of 1929 another summer's collections would be
added to the pile. We realized that if we were ever to do anything
with this material we would have to take it and ourselves to some
large herbarium-library complex, preferably where there was some-
one with knowledge of the northern American flora. From all we
could learn, Harvard would be the best place. ] could use the Gray

216 Rhodora [Vol. 88

Herbarium and my wife, who had collected the lichens, could use
the Farlow. Hence my letter to Fernald.

To our amazement we had a reply almost by return mail, saying
that I would indeed be welcomed, and that he would put me on as
his assistant in his systematic botany course (Bot. 7). Naturally my
misgivings about him began to evaporate at that point. When we
arrived in Cambridge late in September he hadn’t yet returned from
Newfoundland where he had spent the summer. My first meeting
with him was when he “bounced” into the Herbarium. I can think of
no better word for the way he walked. He was about my height, but
much larger in diameter. The top of his head was bald, but he had a
full beard which was gray and I suppose could have been called a
Vandyke, though it probably hadn't been trimmed since spring and
had grown into a bush. His eyes were blue and twinkled at me over
his half moon glasses. He quickly got me established with a place to
work and storage space for the collections I had brought with me.

From then on he seemed to take a sort of fatherly interest in Lucy
and me and our affairs. He was kindness itself. Very soon after we
arrived he told us we should not expect to be entertained at his
house because his wife was an invalid and they could not have
guests. As we soon found out, however, this situation did not pre-
clude his going out to dinner when invited. We availed ourselves of
this proclivity on many occasions, usually when the monthly meet-
ings of the Botanical Club came around. An old crony of Fernald’s,
J. F. Collins, would come up from Brown University and I would
find both of them at the Herbarium and drive them to our house
for a slightly early dinner, after which we would go downtown to the
meeting. I suppose I should classify those occasions as “educa-
tional” for Lucy and me, which, of course, they were. We learned a
lot from (and about) Fernald—one of the main things we came to
Cambridge to do. But over and above this, they were just unalloyed
fun!

Both Collins and Fernald loved good food, and Fernald himself
was not tyro as a cook. They regaled us with their experiences,
usually hilarious, on their collecting trips to the Gaspé region. The
conversation never flagged when they were there. Both of them were
inveterate punsters.

Punning seemed to be epidemic around the Gray Herbarium in
those days. I had lunch one day with Fernald and A. S. Pease at a
small restaurant nearby. Soon after we had given our orders, one of

1986] NEBC 800th meeting — Letters gif

them said something with a pun in it. This started a flood, and
hardly a sentence was spoken between them throughout the meal
that didn’t have a pun in it. It was an exhibition of which I have
often wished I had a record.

Fernald did many things for us that were beyond the call of duty.
He piloted us on shopping expeditions to the push-cart markets at
Faneuil Hall. They were held on Saturday nights, and he insisted
that we should not get there until 9 o’clock when the prices would be
coming down. We would meet him at Harvard Square and go on the
subway, we with backpacks to carry home the plunder and he with a
large, ancient and much battered suitcase. His procedure at the
market was always the same. We first walked past all the carts,
noting those that had the best stuff at the best prices. Then on the
way back we bought what we wanted.

There was not much we could do for Fernald in return for all his
kindness, but we did what we could. We had an automobile, such as
it was—-a Model T station wagon in which we had come to Cam-
bridge. Fernald didn’t drive and didn’t own a Car, so everywhere he
went he walked, or used public transport, or got somebody to take
him. For the last, of course, I offered my services, for which he was
deeply appreciative. We used the car for class field trips and for
miscellaneous other excursions, but he never overdid this, and was
never demanding. One Sunday he asked if we would like to drive
out to the wet meadows along the Concord River where he knew
there were wild cranberries; we did, and came home with a few
quarts of splendid fruit.

On another occasion in the first autumn we were here, he asked
me to drive him down to Harwichport on the Cape. In the preceding
spring, before he went off to Newfoundland, he had rented a cottage
there in which his family would spend the summer. At the same time
he put in a garden for them. They had tended the garden faithfully,
harvested it in late summer, and had put up a huge amount of
canned goods. Our trip was to bring it all to his house in Cambridge,
which we did.

Except for one incident, I do not remember the details of that
trip. We had started early in the morning so as to get back in one
day. The weather was sunny, very cool and brilliantly clear, with a
stiff westerly wind. In the latter part of the morning we were riding
along a sandy road close beside a large cranberry bog surrounded by
a wire fence. The preceding night had been frosty, and the bog had

218 Rhodora [Vol. 88

been flooded up nearly to the level of the road. The cranberries were
ripening, and those that were already loose had floated to the sur-
face of the water where the wind had blown them through the fence
and piled them up right at the roadside. Fernald got me stopped and
then hopped out with a basket. In no time he had scooped up at
least half a bushel, and on we went. I was somewhat shocked by this
procedure and asked him, rather circumspectly, whether it was
legal. “Oh yes,” he said, “any fruit found on the ground in a public
right of way is public property, at least in Massachusetts.”

At that time I was just beginning to learn something of Fernald’s
character. I found out a little later that what I had seen at the
cranberry bog was thoroughly typical of Fernald. If we were riding
along on some upland country road he would spot an orchard
ahead, and, if it was immediately behind the bordering stonewall,
with overhanding branches, we would stop and he would pick up a
bag of apples. I had caught a glimpse of a phase of Fernald’s charac-
ter that could not even be guessed at from anything he ever
published.

James Truslow Adams wrote in one of his pieces that the New
Englanders had added an eleventh commandment to the Decalogue:
“Thou shalt scrimp.” Then there was the article I read in The New
Yorker many years ago about an imaginary institution called the
“N.E.S.P.D.I.P.” which, being translated, was the “New England
Society for the Prevention of Dipping into Principal.” I once men-
tioned this to Helen Bailey (Mrs. Irving Bailey, for those who didn't
know her), “Well,” she said, “when I was a girl I was taught that I
ought to live on the income from my income.” The town dump for
the village I live in is commonly spoken of as the “Petersham
Exchange,” which it really is.

I used to think that all such stories about these things were apoc-
ryphal. But after living in New England for over half a century | am
convinced that they are founded upon solid fact. I can document
this. I will even go so far as to Suggest that New Englanders are
already coded for scrimping and scrounging when they are born.
My education along this line was begun by Fernald beside that
cranberry bog.

Another opportunity for repaying Fernald for his kindness arose
by pure chance. Student registration for courses occurred soon after
we came. In those days there was no mixing of Radcliffe and Har-
vard students. Professor Fernald gave his Botany 7 lecture at 10 am

1986] NEBC 800th meeting — Letters 219

to the Harvard boys, and at 11 am he did it all over again for the
girls. That fall only two girls registered for the course, and there was
an ironclad rule that a course could not be given for less than three.
This precipitated a minor financial crisis! I say “minor”, but it didn’t
seem so at the time. A substantial wedge of Fernald’s salary came
from that Radcliffe class, along with a much lesser though distinctly
noticeable one in mine. When I told my wife about it that evening
and she said “Why don’t I register and take the class as a special
student?” When I told Fernald this the next morning he all but fell
on my neck. If Lucy had been there I think he would have kissed
her. Shortly before Christmas he appeared at our door one Sunday
morning with a little basket containing Jars of assorted jams and
jellies he had made from fruits he had gathered himself.

Our first child was born that year in mid-February. Term exami-
nations were to be early in January, and Fernald was much con-
cerned about how Lucy would get over to the Radcliffe campus
from our apartment on Gibson Terrace if the day was stormy or icy.
He went over to Radcliffe and arranged to have a proctor give her
the examination at home if the weather was bad, but the day was
bright, so she walked.

Fernald had a large, well furnished office on the second floor in
the east wing of the Herbarium building, but I never saw him do any
work there. He did his writing and studies of specimens and litera-
ture near one end of a long table that stood in the middle of the floor
of the large rotunda-like room which was the central feature of the
building. There he could see and be seen by anyone coming in the
front door. | used to wonder why he thus made himself so readily
accessible, but he seemed to prefer it that way.

His manuscript was all written by hand with a soft pencil on large
sheets of cheap news-print paper, and was easily legible. He had a
singular habit of reading it aloud, not to himself but to anyone who
happened to come by and would listen. He always read with his
pencil in his hand, using it to keep track of his lines, sentences,
words and punctuation. If anything didn’t sound quite right to him,
he would stop and correct it, then go on. After I had been captured
and read to a few times, it dawned on me that he wasn’t doing it
especially for my enlightenment. Rather, he was merely editing his
own manuscript. Someone there listening might catch something
that he missed.

220) Rhodora [Vol. 88

Most of the people who came regularly to the Gray soon learned
to avoid getting caught. If you could get through the front door very
quickly and quietly, you could dodge around through various doors
and passages and remain unseen until you got to where you needed
to be. I did this too, on many occasions. But I must admit that I did
a great deal of listening because I wanted to. Most of the things
Fernald was writing about in the years when I was regularly or very
frequently at the Gray were of immediate interest to me. I didn’t
need to read most of them when they were published because I had
already heard their essentials.

Fernald’s memory for the flowering plants and ferns of northeast-
ern America was truly prodigious. The eighth edition of Gray’s Man-
ual listed over 8000 species and lesser categories, and I think it safe
to say that he knew them all. His memory was not only for names, it
extended to habitat and geographic data, and to a huge literature
accumulated since the seventh edition was published in 1908. Most
of this was scattered through miscellaneous journals and a few
monographs, a great deal of it in Rhodora, but the only key to it was
in Fernald’s mind.

I was working on the flora of a part of N.W. Canada that was
outside the ranges covered by any extant manual, but it was in the
boreal forest where most of the species have wide east-west ranges,
far enough east to be well within the Gray's Manual range, | would
get stuck on a specimen, commonly a grass or sedge. I could get
close to it in the Manual or the Herbarium but not close enough. |
would show it to Fernald and he would immediately confirm or
correct my diagnosis up to that point and admit that he couldn't go
any further. Then he would get a far-away look in his eye and
suggest that I look in the Rhodora volume for 1912. After this sort
of thing had happened a dozen times or s0 it dawned on me that he
had never been wrong!

Quite by chance, one day I saw an exhibition of his memory
prowess. About 1938, Eric Hultén spent several months in this
country and Canada visiting the larger herbaria and getting data for
his great collection of spot maps of species ranges. He was several
weeks at the Gray. He didn’t think he would be likely to find much
at the Arboretum herbarium but because he was here he might as
well look. I told him I would take him over to Jamaica Plain and
that he should stay at our house. I was to pick him up at the Gray in
late afternoon. When I got there I couldn't find either him or

1986] NEBC 800th meeting — Letters Ze1

Fernald, even though I looked everywhere in the Herbarium. As a
last resort I opened the door into Fernald’s unused office, and there
they were with their heads together over some specimens and maps,
and surrounded by stacks of folders from the Herbarium. Hultén
always carried with him a supply of blank maps, made at various
scales. Fernald would turn over the sheets, glance at the labels,
reach over with his trusty pencil, put the point down on the map,
and Hultén would make another dot. If the specimen was one of
his own collections, Fernald didn’t even look at the label, for he
would recognize the specimen itself and remember exactly where he
had picked it up.

I don’t know how long they had been at this, but Hultén’s map
for the Maritime Provinces, Newfoundland and southern Labra-
dor was heavily sprinkled with dots. After I had watched for half an
hour or so, they quit for the day and Hultén and I left. On the path
out to the car, Hultén stopped in his tracks, turned to me and said,
“Tell me—can Fernald really do this?” I said, “Well, I’m afraid he
can.” Then we went on to the car.

Anyone who reads what I have written about Fernald thus far
would be thoroughly justified in considering that I became seriously
biased by his unfailing cordiality and generosity to me and my
family. I did, and still remain so. But as the years went on, the bias
was severely tempered by events. I knew there was another side to
his character, for I had seen it in those reviews I had read before I
knew him. He had no patience with sloppy work or with good work
that was poorly presented. Somehow, he contrived to take it as a
personal insult when he was asked or required to read it. He could
be just as irascible and thoughtless in his personal dealing with some
people as he could be cordial and considerate with others. He would
take instant and often firey dislikes to some people who came to him
for advice or suggestions. There was no way to predict when this
would happen.

Over the years I got the notion that Fernald really didn’t like
teaching. It took time away from his own studies, which he consi-
dered far more important. I never saw any evidence of preparation
for his lectures in Botany 7. When our son was born in February of
our first year in Cambridge, Lucy was out of the class for about six
weeks, so I took notes for her. But the lectures, though always
interesting (to me at least), were rambles in systematic botany. They
could not be made to “parse.”

222 Rhodora [Vol. 88

He always had a few graduate students around, some of whom
stayed with him through their doctorates while others migrated to
other segments of Biology. He had a habit of giving a new student a
taxonomic problem to cut his teeth on. This is a perfectly good idea,
but it has to be used with consideration for the student’s own prepa-
ration and proclivities. Also the problem has to be one that can be
worked out in a relatively short time. Fernald seemed never to sense
the need for all this. Rather, he would start a student off with a
genus or section of a genus that was notoriously complex and diffi-
cult, such as Poa or Calamagrostis with which he himself had trou-
ble. This led to early discouragement, and I think he lost some very
good students by it. I suspect that one of the reasons he accepted
me in the way he did was that I did not come to him as a student,
thus relieving him of all the responsibilities for seeing me through
course work, examinations, a thesis, and job hunting.

Fernald’s flaring temper sometimes resulted in distinct disadvan-
tage to his own work. One example of this is a case in which I had
no part, but about which I had first hand reports. Fernald’s famous
paper on “Persistence of plants in unglaciated areas, etc.,” published
in 1925, invited serious criticism from the very beginning. Glacial
geologists wrote it off immediately because they had no evidence
that any of Fernald’s unglaciated areas had actually escaped cover-
age by the Wisconsin ice. Another ready target was Fernald’s use of
negative evidence. A great many of his persisting plants had wide
ranges in the northern Rocky Mountain region, while their repre-
sentatives in the East were relatively rare and found only in small,
restricted areas. Who was to say they were actually absent in the
vast stretch of country between the lower St. Lawrence region and
the Rockies? This question was particularly pointed because most of
that country had never been seen by experienced botanical collec-
tors. Still another serious question was biological, and involved the
behavior of the plants themselves. If Fernald was correct in his
persistence theory, the relict species were on hand and ready to
move onto the newly exposed land when the ice melted away. But
they had not done so, and were soon crowded into rarity by vigor-
ous migrants from the south. Fernald had no good explanation for
this, and could only propose that the plants were “senescent,” “run-
ning down” as it were, as species were thought to do as they
approached extinction.

1986] NEBC 800th meeting — Letters 223

Edgar Anderson was at the Arboretum for several years in the
early 1930’s where, among other things, he was making cytotaxo-
nomic study of the genus /ris. He remembered that /ris setosa was
one of Fernald’s disjunct species having a wide range in the northern
Rockies and Yukon, but with a single variety in small areas from
southern Labrador to eastern Maine. He knew about the work of
Turresson in Sweden and the ongoing transplant studies in Califor-
nia, both of them showing the significance of heritable ecotypic
variation in species. He saw at once that here was a basis for expla-
nation of the local behavior of disjunct species, and went over to the
Gray to tell Fernald about it. Unfortunately it was the “wrong day,”
and Fernald exploded. He refused to have any part of it, and Andy
was crushed. Why he took this line I do not know, but I have always
considered the event as something of a tragedy. I think Fernald was
being handed the first reasonable biological support his theory ever
had, and he needed it.

I know of two cases in which Fernald apparently thought over his
tirades and eventually “came around.” In neither case did he ever
say anything to me about how or why he did this.

The first was concerned with the publication of the Fernald and
Kinsey book on “Edible wild plants, etc.” I had learned about the
manuscript of it soon after | came to Cambridge in 1929. Fernald
had found out that I had, in previous years, made a lot of drawings
of plants, and he got the notion that I could make illustrations for
this manuscript which he hoped to get published “some day.” One
morning he came to my desk with it, sat down and started reading it
to me. He spent most of the day at it! The book was of real interest
to me but I couldn’t get much excited about the drawings, for they
would take a lot of time that I knew I wasn’t going to have. Appar-
ently Fernald got the message, and I heard no more about it for five
or SiX years.

By that time I had moved my base of operations to the Arbore-
tum and Dr. E. D. Merrill had become Director there. During his
years at the New York Botanical Garden, Merrill had become well
acquainted with Dr. E. G. Stillman who had long been a generous
benefactor to Harvard. Dr. Stillman owned a large contract printing
establishment in Cornwall, NY and carried around in his coat
pocket the “office” of a tiny publishing business that he called the
“Idlewild Press” and ran by himself. He was always on the lookout

224 Rhodora [Vol. 88

for odd manuscripts that were good but were not likely to be attrac-
tive to commercial houses. Merrill apparently told him about the
Fernald-Kinsey manuscript at some time or other, and he imme-
diately told Merrill to get hold of it and he would print it. Dr.
Merrill talked it over with me and we debated at some length on
how we would approach Fernald, which we would have to do first.
Merrill was new here and had an idea that Fernald didn’t like him
very much, which I suspect was true. So the job fell to me and I tried
it, with dismal and eruptive failure. Fernald absolutely refused to
cooperate. This was my last personal involvement with “Edible wild
plants.” Two or three years later I heard by the grapevine that
Fernald was getting the manuscript ready for printing, and in 1943 it
was published by the Idlewild Press and printed at Dr. Stillman’s
Cornwall establishment. To this day I do not know how or why
Fernald reversed himself so completely.

The second of these strange cases arose one morning when I
found Fernald reading some galley proofs. He said he wanted to
read me something, which turned out to be a review of Stanley
Cain’s book on “Foundations of plant geography”, published in
1944. | had read the book and knew that it had some weak spots in
it, but in spite of these I considered it to be by far the best thing
available in the field it covered. Stanley had been at the Gray for
two or three weeks while he was writing it, and I knew that Fernald
liked him. But his review panned the book unmercifully. Fernald’s
specific complaints I could agree with, and did, for they dealt with
floristic geography in which Stanley’s treatment was not as up-to-
date as it should have been. Otherwise I managed to say very little
about the review, and promptly forgot it. A few weeks later a letter
came from Stanley asking whether he should do anything about a
letter he had just had from Fernald. In this letter Fernald told him
he had written a review and had it in galleys, but after thinking it
over had decided not to publish it. He had ordered the printer to
break down the type. I told Stanley that I didn’t know why Fernald
had done this, but that I was pretty sure the letter was evidence of an
entirely unique event and should be framed and hung in his office.

1986] NEBC 800th meeting — Letters 220

from Frank Seymour:

It is strange that I do not remember when I first met Dr. Merritt
Lyndon Fernald. When I went to the Gray Herbarium to identify
some specimens I had collected, I could not expect any attention
from him. He was pointed out to me at a respectful distance: “That’s
Dr. Fernald.”

For two or three summers before I entered Harvard I had been an
herbarium assistant, so, I conferred with him frequently regarding
my work. In my sophomore year (Sept. 1913-June 1914), I elected
his famous course, Botany 7. With about 27 other students in an
extraordinarily large class, we all enjoyed his humor. On reading
what one member of the class had written in his examination paper,
Dr. Fernald regaled all the members of the Staff with merriment
over the exploits at what this student called “Gas Bay.” One does
not have to ponder long to guess in what field that student’s father
made his fortune.

The following year I was privileged to be Dr. Fernald’s assistant
in the same course, Botany 7. | listened with delight, the second
time, to his lectures. A new feature that year was a lecture on fam-
ous botanists, especially American botanists.

The influence of the Continental Ice Sheets on the range of spe-
cies, or, his account of that, made it extra interesting to me, about to
begin my study of Marthas’s Vineyard. Half of that island was
covered by the Ice Sheet and the other half was not glaciated, at
least, not by the most recent glacier. My study was carried on under
the immediate direction of Dr. Fernald. It began with a series of
collecting trips to the island. A few years later, accompanied by
James M. Fogg, Dr. Fernald went to Martha’s Vineyard for a brief
collecting trip himself.

Dr. Fernald’s greatness as a botanist is seen by comparing him
with Linneaus. Linneaus entered the field when the science was
comparatively new; he had the whole field before him. From it he
could make multitudinous species. Fernald, in contrast, arose when
New England had been hand-picked for new species by several gen-
erations of botanists. Yet new species kept popping up under his

226 Rhodora [Vol. 88

wonderful eyes and mind. If Linneaus had lived in Fernald’s time,
he might not have found as many species as Fernald.

Interrupted by my 18 months service in the U.S. Navy in World
War I, I returned to work for Dr. Fernald only for about 4 months
in 1919. After that my contacts with him were few as I moved out of
easy reach of Harvard.

from Lyman B. Smith:

Professor Fernald taught me to make superb herbarium speci-
mens, to do bibliographic research, and to transfer personal enthu-
siasm into teaching.

First he learned from very fastidious amateurs to select not only
the very best but to leave sufficient space for the label in the lower
right corner and, if necessary, to choose smaller than average speci-
mens with notes that they were selected small, and what size the
plants in the stand would reach. A further refinement was extra
flowers for dissection and Fernald’s invention, the salivator, where a
flower was held open by a bit of wet newssheet.

In drying, he at least experienced the full sweep of history from
changing driers constantly with the necessity of drying the wet ones
with luck by sunlight but more often by toasting before an open fire.
Then came ventilators and circulating hot air which he at first
rejected because ventilators used alone would often leave cross-bars
on the specimens. Later he used driers between the ventilators and
the specimen with good results. I do not remember his using metal
ventilators but they have proved impractical because of weight and
the tendency of local populations to appropriate them for roofing.

In the Gaspé in 1923, Fernald taught me to collect dwarf arctic
types with my nose close to the ground; in the late 20’s with his
support, I began the collection of hundred sheet sets for the Plantae
Exsiccatae Grayanae which was a graduate course in making good
specimens and an introduction to bibliographic studies in making
the manuscript for the printed labels. Each number had _ the
significant references from one of Fernald’s new species to others
with a dozen references beginning with Linnaeus. Every reference
had to be verified, which was not too difficult with the richness of
the Gray Herbarium Library.

1986] NEBC 800th meeting — Letters PH

Fernald had little difficulty in transferring his enthusiasm for the
taxonomy of the Gray’s Manual Flora which he extended into Can-
ada and Newfoundland because his students were already commit-
ted. However, he enlivened what could have been a dull memorizing
of family characters by a wealth of anecdotes. He had his adventures
and misadventures and frequently told ones on himself. One student
enjoyed Botany 7 so much that he took a rerun by auditing it the
next year and over 60 years later I still remember family characters
by the stories with them. However, for the highest pitch of enthusi-
asm, one had to attend one of Fernald’s lectures to the New England
Botanical Club where he received almost revival meeting acclaim.

from G. Ledyard Stebbins:

My association with Professor Fernald began when I took his
course, Botany 7, in 1926-27, continued until I left Harvard in the
spring of 1931, and sporadically thereafter. Of the numerous memo-
ries I have of him, I select the following incident.

During my time the weekend field trips that he led in October to
Cape Cod were exciting affairs that no botany student who was
invited on them ever missed. Of the four that I attended, that of
1927 was the most memorable. On his Friday lecture, the day before
we left, he said: “I’ve asked Harlow Bishop (one of Weston’s gradu-
ate students) to help with transportation, so some of you will have
to ride in his car, that he took to Gaspé last summer, and named
‘Junkus canadensis’ because of its erratic behavior. I’ve ridden in
said car, and hope that it will hold up for the trip. However, I’m
afraid that any time it could descend to the status of another species,
‘Junkus inutilis.””

These words were prophetic. The conversation came Saturday
evening, just before dinner, in the main street of the town of Har-
wich. Fernald was as disturbed over the immobile vehicle as we
were, and gallantly joined the push-crew that was removing it from
the center of the street to an acceptable parking place. In doing so,
he failed to realize the modest speed that it would acquire with
everybody pushing it. All of a sudden, we found him on the pave-
ment, with his foot pinioned down by the front wheel. We quickly
extricated him, and carried him to the restaurant for dinner and to

228 Rhodora [Vol. 88

his lodging. Meanwhile, Bishop was able to get the car going, at
least temporarily.

We all expected that this disaster would end our trip. Not so, said
Professor Fernald. His foot had been examined and the damage was
minor. So the next day we drove over the usual tour of dunes and
sandy pond margins filled with the rare species that he knew and
loved so well. At every stop, two stout and faithful students would
make with their arms a comfortable sedan chair and carry him to
the pond margin, while other students brought him the specimens to
which he pointed. He would then give his inimitable descriptions of
the geographic distribution, relationships and other valuable infor-
mation about each species that he knew so well. The performance
appeared like a miniature version of Sir Joseph Hooker in India,
making notable botanical pronouncements from the howdah on the
back of an elephant. We were even more impressed than we would
have been without the accident.

HOLOCENE FRUIT, SEED AND LEAF FLORA
FROM RIVERINE SEDIMENTS NEAR
NEW HAVEN, CONNECTICUT

LEE S. PIERCE AND BRUCE H. TIFFNEY

ABSTRACT

Fruits, seeds and leaves of 48 gymnosperm and angiosperm taxa are reported from
2830+70 and 2680+30 year-old riverine sediments located just north of New Haven,
Connecticut. Most of the fossils are identified to species, providing refinement of
previously available pollen data; several species are not commonly found in Holo-
cene pollen floras.

Key Words: Holocene flora, riverine sediments, central Connecticut

INTRODUCTION

The post-glacial history of the flora and vegetation of the eastern
United States has been reconstructed from studies of fossil pollen,
and less commonly from fruits, seeds and leaves (Davis, 1969a,
1976: Bernabo and Webb, 1977; Delcourt and Delcourt, 1981;
Webb, 1981). We present data on a macrofossil flora collected from
riverine sands that were deposited about 2700 years before present
(YBP) just north of New Haven, Connecticut. We describe the sed-
iments and give a brief paleoecological interpretation of these data.

The New Haven flora is significant for two reasons: it is a macro-
fossil flora and it was collected from a river valley in Connecticut’s
central lowland. Macrofossils, in contrast to pollen, are generally
derived only from plants living near the site of deposition
(McQueen, 1969; Birks, 1973; Spicer, 1981) and are frequently iden-
tifiable to the species level, thus providing local ecological details
which studies of pollen cannot supply. Qualitative information from
macrofossils complements quantitative information derived from
pollen (Watts and Winter, 1966), the source flora and deposition of
which can be statistically characterized (e.g. Davis and Webb, 1975;
Webb and McAndrews, 1976; Webb et al., 1978; Webb et al., 1981).
Modern studies of Holocene macrofossils emphasize collection
techniques that permit a similar quantitative reconstruction of the
source vegetation (Watts, 1979; Baker et al. ,1980; Davis et al., 1980,
Delcourt et al., 1980; Barnosky, 1981). Unfortunately, much of the
material available for the present study was collected at an earlier

Zao

230 Rhodora [Vol. 88

time, and cannot be replicated; hence we limit our contribution to a
description of the flora.

The importance of a lowland river deposit is unique in that most
sites studied in New England have been upland lakes (D. Gaudreau,
pers. comm.), including the best studied site in Connecticut, Roger’s
Lake (Davis, 1969b and references therein). Traditionally, Quater-
nary palynologists have not studied river deposits because such
pollen floras are the product not only of the source vegetation and
wind patterns, but also of the sorting of pollen grains by flowing
water. This last taphonomic variable, which affects macrofossil flo-
ras to a lesser degree, cannot be easily characterized. Thus, this flora
presents data on a rarely sampled community.

SITE DESCRIPTION

The fossils were collected on the property of Landfill Associates
Inc., formerly the Stiles Clay Pit, on the west margin of the Quinni-
plac River (41°21’49”N, 72°53’7”W) in Hamden, Connecticut. Clay
mining ceased at this site several years ago; it is presently a sanitary
landfill. Excavations for sand and clay to cover refuse have exposed
a face that is approximately 5 meters high and about 100 meters
long. This section (Figure |) reveals a series of clays, sands and peats
(Bloom, 1968).

The bottom of the exposed face rests on the red clays of the New
Haven formation, deposited in a glacial lake approximately 12,000
Y BP (Brown, 1930). The clay is reported to contain fossils of tundra
plants (Beinecke and Siccama, 1973,), but no published evidence
exists, and we were unable to locate any such fossils in the field.

The clay is disconformably overlain by 2.6 to 2.9 meters of cross-
bedded, quartz-rich sands deposited by the Quinnipiac River follow-
ing draining of the glacial lake. This unit is gravel at the contact with
the clay, but grades rapidly into sand which becomes increasingly
fine toward the top of the unit. The basal portions contain many
transported logs of Quercus L. (two species), Tsuga (L.) Carr. and
Acer L. (Brown, 1930) together with smaller plant fragments. These
remains have been radiocarbon dated at 6810+170 YBP (Stuiver
and Deevey, 1961). The upper portions of the sandy unit contain
frequent lenses of leaves, fruits and seeds. Wood from this part of
the unit has been dated at 3560+180 YBP (Stuiver et al., 1963),

1986] Pierce & Tiffney — Holocene flora 231

meters

O

ee ee

fine sand

bias)
| Rae |
L’.°} coarse sand
lad aos
|e o
ob «2 ad gravel
~ ,
a _| varved clay
x cross beds
x
J
- 5 ( wood

& leaf mat

f fruits&seeds
(

Figure 1. Stratigraphic section, west margin, Quinnipiac River, Hamden, Conn.
Data from Brown, 1930, Siccama (Personal data, unpublished) and field observa-
tions by the authors.

232 Rhodora [Vol. 88

hickory nuts at 2830+70 and butternuts at 2680+30 YBP (Quater-
nary Isotype Laboratory, University of Washington, Seattle;
November, 1984; specimens QL-1881 and QL-1882).

The crossbedded sands are topped by 1.7 meters of Spartina peat
and 0.3 meters of Typha peat. The Spartina peat correlates with
other evidence (Sears, 1963; Bloom and Stuiver, 1963) for rising sea
levels in the last 7000 years. As the sea level rose, the boundary
between salt water and flowing fresh water retreated up the Quinni-
piac River. The arrival of this interface at the fossil locality is
marked by truncation of the riverine sands by Spartina salt marsh
peat. Human interference in the early 1900's restricted tidal flow in
the lower Quinnipiac, and consequently the fresh water Typha
marsh now occupying the site became established (Beinecke and
Siccama, 1973).

MATERIALS AND METHODS

Lenses of fruits, seeds and wood and leaf mats were collected
from the upper half of the gravel-sand unit. Specimens from the two
richest lenses of fruits and seeds were radiocarbon dated at 2870+70
YBP (specimens of Carya cordiformis (Wang) K. Koch) and
2680430 YBP (specimens of Juglans cinerea L.). (Quaternary Iso-
type Laboratory, University of Washington, Seattle; November,
1984; specimens QL-1881 and QL-1882). The species present in
these collections are listed in Table 1.

Fruits and seeds were identified by comparison to descriptions,
photographs and drawings in Ogden (1953), Brouwer and Stahlin
(1955), Martin and Barkley (1961), Katz et al. (1965), Berggren
(1969), Seymour (1969), Fernald (1970), Schopmeyer (1974), Mont-
gomery (1977), Elias (1980), and Hellquist and Crow (1980), and by
comparison to specimens in YU, A and GH. Leaves were identified
similarly from Petrides (1958), Symonds (1958, 1963), Fernald
(1970), and Elias (1980), and by comparison with YU, A and GH
specimens.

Photos of small fruits and seeds were taken on Kodak Panatomic
Film with a Nikon PMF photo attachment mounted on a Nikon
SMZ-10 binocular dissecting microscope. Photos of leaves and
large nuts were taken with a 35 mm camera with a 55 mm
macrolens.

1986] Pierce & Tiffney — Holocene flora 233

SYSTEMATICS

The nomenclature used here is that of Fernald (1970). The des-
criptions of leaf venation follow the method of Hickey (1973). Fruits
and seeds with bilateral symmetry are described by naming the
plane figures which would be seen if they were sectioned along the
plane of symmetry (sagittal section), along a plane that is perpen-
dicular to the plane of symmetry and which includes the longest axis
of the plane of symmetry (frontal section), and along a plane per-
pendicular to the two previously described planes (transverse sec-
tion). When “base” and “apex” are used to describe fruits, they refer
to the pedicellate or pedunculate and stigmatic ends, respectively.
For seeds, the chalaza occurs at the base, the micropyle at the apex.
The fruit and seed specimens cited in this study are stored in the
Paleobotanical Collections of the Peabody Museum, Yale Uni-
versity, collection numbers 4649 through 4825. Figured leaf speci-
mens, which were difficult to preserve, were not saved; leaf-bearing
bulk samples are also available in the Paleobotanical Collections.

LIST OF SPECIES
PINACEAE

Abies balsamea (L.) Mill.: Rare.

The 4-mm-long seed is oblong in sagittal section, lenticular in
transverse section and elliptic in frontal section, is angled on one
margin and has concave sides along the other margin, a common
feature in Abies seeds. Figure 2.

Pinus L. or Picea Dietr. sp.: Rare.

The 4-mm-long, oblong, lenticular, elliptic (sagit., trans. and
frontal sections, respectively) seed is Pinus or Picea, but without the
presence of a wing, it is impossible to determine which it is. Figure 3.
Tsuga canadensis (L.) Carr.: Several.

The ellipsoid cones are 1.2 to 1.9 cm long, with scales that are
almost as long as they are broad. Figure 4.

SPARGANIACEAE

Sparganium eurycarpum Engelm.: Several.
The 6 to 8 mm broad pithy shoulders of the 8 to 10 mm long

234 Rhodora [Vol. 88

Table 1. Species identified from two dated samples, and three associated but
undated samples, of fruits, seeds and leaves collected in Holocene sediments near
New Haven, Connecticut.

2830+70 2680+30 Undated
YBP YBP Lens |. Lens 2. Leaf Mat

PINACEAE
Abies balsamea +
Pinus or Picea sp. +
Tsuga canadensis + +

SPARGANIACEAE
Sparganium eurycarpum +
Sparganium cf. americanum +
Sparganium cf. fluctuans ++

+44

ZOSTERACEAE
Potamogeton nodosus +
Potamogeton pulcher +
Potamogeton vaginatus + +
Ruppia maritima +
Zannichellia palustris +

CYPERACEAE
Carex lupulina +
Cladium mariscoides +
Dulichium arundinaceum +
Scirpus, two species at

++ H+

LILIACEAE
Smilax sp. +

MYRICACEAE
Myrica pensylvanica +

JUGLANDACEAE
Carya cordiformis
Carya ovata
Juglans cinerea

++4
+
+

CORYLACEAE
Carpinus caroliniana
Ostrya virginiana +
Alnus sp. ve

+++

FAGACEAE
Fagus grandifolia
Quercus alba
Quercus coccinea

+44
3+

POLYGONACEAE
Polygonum, three species + + oP (2)

1986] Pierce & Tiffney — Holocene flora

CERATOPHYLLACEAE
Ceratophyllum echinatum

NYMPHACEAE
Brasenia schreberi

RANUNCULACEAE
Ranunculus cf. trichophyllus

MAGNOLIACEAE
Liriodendron tulipfera

PLATANACEAE
Platanus occidentalis

ROSACEAE
Prunus nigra
Rubus sp.
Crataegus sp.

STAPHYLEACEAE
Staphylea trifolia

ACERACEAE
Acer sp.
Acer saccharum

VITACEAE
Parthenocissus sp.
Vitis labrusca

TILIACEAE
Tilia americana

VIOLACEAE
Viola sp.

NYSSACEAE
Nyssa sylvatica

CORNACEAE
Cornus alternifolia
Cornus ammomum
Cornus drummondi

ERICACEAE
Genus indet.

CAPRIFOLIACEAE
Sambucus canadensis
Sambucus pubens

COMPOSITAE
Ambrosia artemisiifolia
Ambrosia trifida
Helianthus cf. decapetalus

+ +

+++

+++

236 Rhodora [Vol. 88

obtriangular achene are characteristic of this species. Several speci-
mens have two seed chambers. Figure 5.

Sparganium cf. americanum Nutt.: Rare.

The narrow obovoid achene is 5.8 mm long, 2.1 mm in diameter,
with a blunt apex and a narrowed base. This description agrees with
S. americanum, but the achene is also similar to the slightly larger
fruits of S. androcladum (Engelm.) Morong. Figure 6.

Sparganium cf. fluctuans (Morong) Robbins.: Rare.

The obovoid achene is 4.5 mm long, 2.1 mm in diameter, has an
enlarged, blunt apex tipped with a broken falcate stigma and a
narrow and elongate base. Although similar to S. angustifolium
Michx. and S. multipedunculatum (Morong) Rydb., the fossils lack
the constriction common in fruits of these two species. Figure 7.

ZOSTERACEAE

Potamogeton nodosus Poiret: Several.

The 4-mm-long endocarp is broadly elliptic in sagittal section,
narrowly elliptic in transverse section, oblong in frontal section, has
an undulating keel along the valve, an apical style base near the
margin opposite the valve, and convex sides. Figure 8.

Potamogeton pulcher Tuckerman. Several.

The 3.8-mm-long endocarp is obliquely obovate in sagittal sec-
tion, oblong in transverse and frontal sections, has a rounded keel
on the valve, a central projection on the opposite margin and three
basal projections. Figure 9.

Figures 2-25. Specimen identifications as indicated; further discussion in the
text. 2. Abies balsamea. Seed without wing. X4. 3. Pinus sp. or Picea sp. Seed
without wing. X4. 4. Tsuga canadensis. Ovulate strobilus. X1.5. 5. Sparganium
eurycarpum. Achene. X4. 6. Sparganium cf. americanum. Achene. X4. 7. Spar-
ganium cf. fluctuans. Achene. XS. 8. Potamogeton nodosus. Endocarp. X4. 9.
Potamogeton pulcher. Endocarp. <4, 10. Potamogeton vaginatus. Endocarp. X4.
11. Ruppia maritima. Achene. 6. 12. Zannichellia palustris. Achene. <6. 13.
Carex cf. lupulina. Achene. 3. 14. Dulichium arundinaceum. Achene. 4.
15. Cladium mariscoides. Achene. X5. 16. Scirpus species one. Achene. X5. 17.
Scirpus species two. Achene. X5, 18. Myrica pensylvanica. Endocarp. XS.
19, Carya ovata. Nut. X1.3. 20. Carya cordiformis. Nut. X1.3. 21. Juglans cine-
rea. Nut. X0.75. 22. Alnus sp. Ovulate catkin. X2. 23. Alnus sp. Scale of ovulate
catkin. X6.3. 24, Ostrya virginiana. Nut. X4. 25. Carpinus caroliniana. Nut. X4.

1986] Pierce & Tiffney — Holocene flora 237

238 Rhodora [Vol. 88

Potamogeton vaginatus Turczaniow. Several.

The 3.3-mm-long endocarp is roughly broad elliptic in sagittal
section with a sinusoidal dorsal margin, is ovate in transverse and
frontal section, has concave sides and no keels or projections. Fig-
ure 10.

Ruppia maritima L.: Rare.

The less-than-2 mm-long, compressed ellipsoid achene has a per-
sistent style similar to that of Potamogeton, but which extends
beyond the body of the seed to form an apical beak. Figure 11.

Zannichellia palustris L.: Rare.

The 2.5 mm-long, flattish achene is oblong-arched in sagitittal
section, has a stalk at one end, a style base at the other, and spine-
like projections along both sides. Figure 12.

CYPERACEAE

Carex lupulina Muhl.: Several.

The achenes average 3.7 X 2.3 X 2.3 mm and are elliptic in frontal
section, triangular in transverse section, with a long or contorted
style and a stipitate base. Figure 13.

Dulichium arundiaceum (L.) Britt.: Rare.

The flattish oblong achene plus style averages 5 mm in length and
possesses six to nine basally-attached bristles which are longer than
the achene and style together. Figure 14.

Cladium mariscoides (Muhl.) Torr.: Rare.

The achene is 2 mm in diameter. The soft outer covering has been
eroded, revealing a sphere with three longitudinal ridges and a dis-
tinctive basal ring. Figure 15.

Scirpus species one: Numerous.

The achene is 2.5 mm long, 1.5 mm wide, obovate to plano-
convex (Ssagit.), elliptic to plano-convex (trans.), obovate (front.),
with a few attached bristles. It is similar to S. acutus Muhl. or S.
validus Vahl. Figure 16.

Scirpus species two: Numerous.

The achene is 3.2 mm long, 2.9 mm wide, plano-convex (sagit.,
trans.), obovate (front.). It is similar to S. fluviatilis (Torr.) Gray
and S. robustus Pursh. Figure 17.

1986] Pierce & Tiffney — Holocene flora 239

LILIACEAE

Smilax L. sp.: Rare.

The obovate leaf with a cordate base has basal perfect acrodrom-
ous venation. The specimen is too poor to permit species determina-
tion. Figure 58.

MYRICACEAE

Myrica pensylvanica Loisel.: Rare.
A 3 X 2.5 mm ovoid endocarp with thick walls which thin at the
apex. Seeds of M. gale L. are narrowly ellipsoid. Figure 18.

JUGLANDACEAE

Carya ovata (Mill) K. Koch.: Several.
The ellipsoid nut is 2 cm in diameter, four-angled in transverse
section with a smooth surface. Figure 19.

Carya cordiformis (Wang) K. Koch.: Rare.

The subglobose nut is 2 cm in diameter, with an apical mucro and
a faintly ridged but otherwise smooth surface. C. glabra has a more
clearly ridged and nerved surface. Figure 20.

Juglans cinerea L.: Several.
The elongate-ellipsoid nut is 4 X 2.5 cm with a round base,
pointed apex, and strong, ragged, longitudinal ribs. Figure 21.

CORYLACEAE

Alnus Mill. sp.: Several.
The catkin is composed of many scales which are thickest at their
apices. A species determination is not possible. Figures 22, 23.

Ostrya virginiana ( Mill.) K. Koch: Rare.

The 5.5 mm-long X 3 mm in diameter nut is compressed-
lanceoloid to ovoid, with a smooth to faintly striate surface. Figure
24.

Carpinus caroliniana Walt.: Several.

The nut is 2.7 X 2.4 * 1.2 mm, compressed-ovoid with three or
four prominent ridges on both faces, and ridges along the margins.
Figure 25.

240 Rhodora [Vol. 88

FAGACEAE

Fagus grandifolia Ehrh.: Numerous.

The nut is obovate in frontal section, 10 to 15 mm long and 5 to
10 mm wide, has a distinct triangular transverse section, and is
borne in an involucre with four elliptic, tuberculate valves. The
elliptic leaf has a short to long acute apex, acute base, pinnate
primary venation, craspedodromous secondary venation and
toothed margins with one tooth per secondary vein. Figures 26-28,
59.

Quercus alba L.: Rare.

The involucre is 8 mm deep and 12 mm wide, covered by rounded
tuberculate scales which are not joined at the base of the cupule.
This suite of characters fits only Q. alba of the Quercus species in
the northeastern United States. The leaf, which has pinnate primary
venation and craspedodromous secondary venation, is obovate and
has obliquely descending sinuses; the four pairs of ascending lobes
are narrowly oblong and are not tipped with bristles. These features
also conform to Q. alba. Comparisons could be made to Q. stellata
Wang. or Q. macrocarpa Michx., but these do not match in all
characters. Figures 29, 60.

Quercus coccinea Meunch.: Rare.

The 5 mm-deep and 8 mm-wide involucre is closely covered by
tapering truncate scales. These cupules are clearly those of Q. cocci-
nea Meunch. Associated leaves with deep sinuses and bristle-tipped
lobes and pinnate craspedodromous veins agree with those of Q.
coccinea, but could possibly represent another species. Figures 30,
61.

Figures 26-42. Specimen identifications as indicated; further discussion in the
text. 26. Fagus grandifolia. Involucre. X1.5. 27. Fagus grandifolia. Scale of
involucre. X1.7. 28. Fagus grandifolia. Nut. X1.7. 29. Quercus alba. One half of
involucre, three views. X2. 30. Quercus coccinea. Involucre, three views. 2.3.
31. Polygonum species one. Achene. 5. 32. Polygonum species two. Achene.
x4. 33. Polygonum species three. Achene X7.5. 34. Ceratophyllum echinatum.
Nutlet. X4. 35. Brasenia schreberi. Seed. X4. 36. Ranunculus cf. trichophyllus.
Achene. XS. 37. Liriodendron tulipifera. Seed. X4. 38. Platanus occidentalis.
Receptacle. X2. 39. Rubus sp. Pyrene. X4. 40. Crateagus sp. Nutlet. X4. 41.
Prunus nigra. Endocarp, X2. 42. Staphylea trifolia. Seed. <4.

1986] Pierce & Tiffney — Holocene flora 24]

242 Rhodora [Vol. 88

POLYGONACEAE

Polygonum L. species one: Several.

The achene is 4.3 mm long X 2.5 mm wide, elliptic (front), sharply
three-angled with concave faces, yielding a star-like transverse sec-
tion. Similar achenes with concave faces are seen in P. douglasii
Greene and P. sagitittatum L., among other species. Figure 31.
Polygonum L. species two: Several.

The achene is 4.6 mm long X 2.5 mm in diameter, is triangular in
transverse section and elliptic in frontal section. The fossils are sim-
ilar to the achenes of P. robustius (Small) Fern., but this is a com-
mon achene shape. Figure 32.

Polygonum L. species three: Rare.

The achene is 2 mm long X 1.8 mm wide, compressed-ovoid, with
a small apical projection. The shape and small size are similar to P.
lapathifolium L., but this is acommon achene shape among species
of Polygonum. Figure 33.

CERATOPHYLLACEAE

Ceratophyllum echinatum Gray.: Rare.

The 6 mm-long nutlet is lenticular in sagittal and transverse sec-
tions, elliptic in frontal section, has three marginal spines plus the
bases of others which have been broken off. The number of spines,
plus the narrow marginal wing produced by their bases, indicate
that this is C. echinatum and not C. demersum L. Figure 34.

NYMPHAEACEAE

Brasenia schreberi Gmel.: Rare.
The 2.5 X 2 mm ellipsoid seed has a tuberculate surface and a
basal attachment scar. Figure 35.

RANUNCULACEAE

Ranunculus cf. trichophyllus Chaix.: Rare.
The 2 mm-long compressed ellipsoid achene is rugose, lacks a
stylar beak and is not sharply keeled. Figure 36.

1986] Pierce & Tiffney — Holocene flora 243

MAGNOLIACEAE

Liriodendron tulipifera L.: Rare.
The seed is 5 X 2.5 X 1.5 mm, flattened-ovoid, with a transversely
wrinkled surface and a basal heteropyle. Figure 37.

PLATANACEAE

Platanus occidentalis L.: Numerous.

The spherical receptacles are 5 to 10 mm in diameter and have a
pitted surface indicative of the insertions of the individual fruits.
Associated 10 to 20 cm-broad leaves have three to five shallowly
pointed, acuminate lobes with teeth scattered along their margins.
Venation is actinodromous or palinactinodromous and semicras-
pedodromous. Many specimens have stout, 5-10 cm long petioles.
Figures 3, 62.

ROSACEAE

Rubus sp.: Several.

The pyrene 3.5 X 2.5 mm, compressed ovoid to oblongoid. The
lateral surfaces are marked by deep pits which form a reticulum.
Identification to species on pyrene characters is not possible in
Rubus. Figure 39.

Crataegus sp.: Several.

The seeds are about 6.5 X 3.8 mm, rough-surfaced, rounded on
one face and bearing three ridges on the other. Identification to
species is not possible with seeds of Crataegus. Figure 40.

Prunus nigra Ait.: Rare.
The endocarp is 14 mm in diameter, flattened spheroidally, with a
low ridge along one side. Figure 41.

STAPHYLEACEAE

Staphylea trifolia L.: Several.

The seeds are about 5.5 X 5.2 X 4.1 mm, slightly compressed
ellipsoid to subglobose. The smooth surface and distinct hilar scar
with multiple vascular traces are characteristic of Staphylea. Figure
42.

244 Rhodora [Vol. 88

ACERACEAE

Acer L. sp.: Rare.

The wingless fruit is 8 X 6 X 2 mm, compressed ellipsoid. A
flattened area denotes the point of attachment to the other fruit. The
surface is marked by reticulate venation. Figure 43.

Acer saccharum Marsh.: Rare.

The 7 to 10 cm-long, deeply palmate leaf has a cordate base and
three to five pointed lobes. Primary venation is basal perfect actino-
dromous and secondary venation is semicraspedodromous. Figure
63.

VITACEAE

Vitis labrusca L.: Several.

The seeds are about 6 X 4 X 2 mm, oblong in frontal section and
elliptic in transverse section, not rounded-triangular. The body con-
stricts quickly from broad shoulders to a cuspidate, not acuminate,
apical beak. This suite of characters distinguishes Vitis labrusca
from other species in northeastern North America. Figure 44.

Parthenocissus Planch. sp.: Several.

The seeds are about 4.2 X 3.7 X 2.7 mm, with a dorsal chalaza
and narrow ventral infolds, globose, sometimes with a rounded-
triangular transverse section. The seed is too short and broad to be
Vitis, and the ventral infolds are too narrow for Ampelopsis. Figure
45.

TILIACEAE

Tilia americana L.: Several.
The capsule is spherical to ellipsoid, about 6 mm in diameter, with
five thin sutures and a slight apical point. Leaves are 10 to 15 cm

Figures 43-57. Specimen identifications as indicated; further discussion in the
text. 43. Acer sp. Fruit without wing. 3.4. 44. Vitis labrusca. Seed. X4. 45.
Parthenocissus sp. Seed. X4. 46. Tilia americana. Capsule. X3.5. 47. Viola sp.
Capsule. <5. 48. Nyssa sylvatica. Endocarp. X4. 49. Cornus alternifolia. Endo-
carp. X4. 50. Cornus amomum. Endocarp. <5. 51. Cornus drummondi. Endo-
carp. X3.6. 52. Ericaceae, genus indet. Capsule X3.5. 53. Sambucus pubens. Seed.
x6. 54. Sambucus canadensis. Seed. 6. 55. Ambrosia trifida. Achene. X5.
56. Ambrosia artemisiifolia. Achene. X4. 57. Helianthus cf. decapetalus. Achene.
X2.5,

1986]

Pierce & Tiffney

Holocene flora

245

246 Rhodora [Vol. 88

long; obovate in shape with a cordate base and a cuspidate apex.
The basal perfect actinodromous primary venation and craspedo-
dromous secondary venation lead to a finely toothed margin. The
fruit bracts are 7 to 10 cm long by 1.5 to 2.5 cm broad with pinnate,
reticulodromous to weakly brochidodromous venation and a
basally attached fruit stalk. Figures 47, 64, 65.

VIOLACEAE

Viola sp.: Rare.
The three-locular papery capsule conforms to that of Viola, but
has no characters by which it can be identified to species. Figure 47.

NYSSACEAE

Nyssa sylvatica Marsh.: Several.

The 7 X 5 X 3 mm-endocarp is marked by ten to twelve promi-
nent, rounded ridges with intervening grooves on each face and a
distinctive germination valve. Figure 48.

CORNACEAE

Cornus alternifolia L.: Several.

The globose endocarp is about 3.7 mm in diameter, with surficial
grooves containing veins. The species is distinguished by the large,
irregularly-shaped apical pit. Figure 49.

Cornus amomum Mill.: Several.

The obliquely ellipsoid endocarp is about 4 X 3 X 3 mm, and
marked by very prominent ridges. Figure 50.

Cornus drummondi Meyer.: Rare.

The ellipsoid endocarps are about 4 X 3 X 3 mm and are marked
by deep circumferential grooves. These are too large to be C. rugosa
Lam. which has a circumferential groove, but is globose. These are
also distinct from C. racemosa Lam., which is obliquely ellipsoid
and has a circumferential line, rather than a groove. Figure 51.

ERICACEAE

Genus indeterminable: Numerous.
The capsules are about 3.7 mm in diameter, spherical to obovoid
with flat to depressed centers, five-valved, and bear very small seeds.

1986] Pierce & Tiffney — Holocene flora 247

Unopened capsules display moderate to faint thickenings over the
valve sutures. While the last characteristic suggests Lyonia Nutt.,
the thickenings are not pronounced (perhaps due to erosion prior to
fossilization), and we cannot exclude the possibility that these cap-
sules belong to Andromeda L. or Chamaedaphne Moench. Figure
De:

CAPRIFOLIACEAE

Sambucus pubens Michx.: Several

The 2.5 X 2 X | mm seed is elliptic to oblong in frontal section
and compressed rounded triangular in transverse section, the sur-
face marked by transverse wrinkles. See note below. Figure 53.

Sambucus canadensis L.: Several.

The seeds are 2.8 X 1.5 X | mm, slightly longer and narrower than
those of S. pubens, but similar in shape and surface pattern. It is
possible that these seeds of Sambucus are from the same species, but
they appear to agree with the two species cited. Figure 54.

COMPOSITAE

Ambrosia trifida L.: Rare.

The achene is 8 to 12 mm long, otherwise similar to that of A.
artemisiifolia from which it is distinguished by its larger size. Figure
55.

Ambrosia artemisiifolia L.: Numerous.

The achene is 4 to 5 mm long, irregular to obovoid, with four to
seven spikes surrounding the central subulate apical beak. The beak
is | to 2 mm long and exceeds the spikes. Figure 56.

Helianthus cf. decapetalus L.: Rare.

The achene is 5 mm long and 2 mm wide, ovoid and flattened with
an apical notch and a striate surface characteristic of Helianthus
species. Figure 57.

DISCUSSION

All the taxa on the species lists (Table 1) are members of the flora
of lowland southern Connecticut. This finding is consistent with
previous work demonstrating that the vegetation of southern Con-
necticut has changed little during the past 2800 years, excluding the

[Vol. 8

Rhodora

248

1986] Pierce & Tiffney — Holocene flora 249

effects of European settlement in the past 350 years, and correlates
well with the pollen record (Davis, 1969b, 1976).

Noteworthy taxa include Staphylea trifolia, which has not pre-
viously been reported from the Holocene of New England, and
Liriodendron tulipfera, whose pollen is rarely preserved in lake sed-
iments (IT. Webb, pers. comm.).

The flora includes plants which grow in a variety of habitats:
aquatic plants of still or slow-moving water such as Ceratophyllum
echinatum, Brasenia schreberi, Zannichellia palustris, and Ranuncu-
lus trichophyllus; marsh plants such as species of Cyperaceae and
Sparganium; levee and flood plain dwellers such as Platanus occi-
dentalis, Nyssa sylvatica and Alnus sp.; forest trees which prefer
well drained soils, such as Acer saccharum, Fagus grandifolia and
Quercus alba; and species which live in open sunny areas such as
Helianthus and Ambrosia (Braun, 1950; Fowells, 1965; Fernald,
1970).

This floral variety is not surprising, given that macrofossils may
be transported up to a few kilometers by rivers (Burrows, 1980;
Drake and Burrows, 1980; Spicer, 1981). The deposit of fruits and
seeds probably represents several different plant associations grow-
ing along the banks or on the floodplain of the Quinnipiac River.
Some fossils show signs of wear, evidence of relatively long distance
transport: seeds of Pinaceae lack wings, fruits of Acer lack wings,
and the A/nus aments are in poor condition. The excellent condition
of many of the fossils from the same lens, however, suggests that
they were not transported long distances before being deposited.
Many fruits have relatively fragile structures intact, including bris-
tles on Scirpus species and Dulichium arundinaceum and styles on
Zannichellia palustris and Carex lupulina. Additionally, entire leaves
were very common in the leaf mats studied.

Figures 58-65. Specimen identifications as indicated; further discussion in the
text. 58. Smilax sp. Leaf (reflected light). X0.5. 59. Fagus grandifolia. Leaf (trans-
mitted light). X0.5. 60. Quercus alba. Leaf (transmitted light). X0.5. 61. Quercus
ef. coccinea. Leaf fragment (reflected light). X0.33. 62. Platanus occidentalis. Leaf
(reflected light). X0.125. 63. Acer saccharum. Leaf (transmitted light). X0.5. 64.
Tilia americana. Leaf (transmitted light). X0.5. 65. Tilia americana. Fruit bract
(transmitted light). 0.5.

250 Rhodora [Vol. 88

ACKNOWLEDGMENTS

We thank Mr. Vincent Pecoraro of Land Fill Associates,
Hamden, Connecticut, for permission to examine the Stiles Clay Pit
site, and for assistance in the field; Dr. Thomas Siccama (School of
Forestry and Environmental Sciences, Yale University) for provid-
ing his collections; Dr. Else Marie Friis (Geologisk Institut, Aarhus
University, Denmark) for assistance in identifying small fruits and
seeds; Ms.Denise Gaudreau and Dr. Thompson Webb III (Dept. of
Geology, Brown University) for helpful comments about the Holo-
cene biogeography and palynology of Connecticut; the reviewers,
Dr. Margaret Davis and Dr. Norton Miller, for insightful and
helpful suggestions; and the curators of A and GH for loan of
comparative material. Research partially supported by NSF grant
DEB79-05082 to BHT.

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BERGGREN, G, 1969. Atlas of seeds and small fruits of Northwestern-European
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BERNABO, J.C. AND T. WesB III. 1977. Changing patterns in the Holocene pollen
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Birks, H.H. 1973. Modern macrofossil assemblages in lake sediments in Minne-
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_R. W. SPEAR AND L. C. K. SHANE. 1980. Holocene climate of New Eng-

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: AND I. Rouse. 1963. Yale Natural Radiocarbon Measurements

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AND T. C. WINTER. 1966. Plant macrofossils from Kirchner Marsh,
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, R. A. LAseKI AND J. C. BERNABO. 1978. Sensing vegetational patterns
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PEABODY MUSEUM OF NATURAL HISTORY
AND DEPARTMENT OF BIOLOGY

OSBORN MEMORIAL LABORATORY

YALE UNIVERSITY

P.O. BOX 6666

NEW HAVEN, CT 06511

POLLINATION BIOLOGY OF PRIMULA LAURENTIANA
(PRIMULACEAE) IN MAINE

CHRISTOPHER S. CAMPBELL, NORMAN C. FAMOUS,
AND MICHAEL G. ZUCK

ABSTRACT

In eastern Maine the brightly colored flowers of Primula laurentiana Fern.
produce no nectar; their faint but pleasant scent is strongest at the center of the
corolla limb in a UV-reflecting, yellow ring called the “bird’s eye.” During our
observations, six species of insects visited the flowers, the most important being the
syrphid Helophilus groenlandicus (O. Fabricus) (Syrphidae). Since P. laurentiana is
self-compatible, long-homostylous, and its stigmas are receptive when the introrse
anthers dehisce, pollen-gathering by insects must lead to some self-pollination.
Pollen adhering to pollinators’ heads may effect cross-pollination during their
flower-constant forays. This breeding system of facultative autogamy is concordant
with the relatively low pollen-ovule ratio of about 130 to I.

Key words: Primula laurentiana, Helophilus groenlandicus, pollination, homostyly,
UV light, pollen-ovule ratio, eastern Maine

Plant biologists have long held a special interest in the genus
Primula because many of the species are heterostylous, i.e., individ-
uals differ reciprocally in stamen and style length (Ganders, 1979).
Heterostyly and the associated system of intramorph incompatibil-
ity have stimulated numerous studies of the floral morphology,
breeding systems, genetics and pollination of several Old World
species of the genus (see references in Ganders, 1979). Most of these
studies have also dealt with homostyly, a floral condition evolution-
arily derived from heterostyly and involving the juxtaposition of
anthers and stigmas. Homostylous plants are generally self-
compatible and of two kinds. Long homostyles combine long styles
with anthers positioned high in the corolla. Short homostyles, the
rare kind, have short styles and low anthers.

Primula laurentiana, the bird’s-eye-primrose, is a scapose, peren-
nial calciphile of ledges, cliffs and meadows in northeastern North
America (Scoggin, 1979). South of about 48°N it is relatively
uncommon (Famous and Campbell, 1984). We report the first work
on the pollination biology of this species. Through field and labora-
tory studies of natural populations on Great Wass Island in eastern
Maine, we have attempted to (1) determine the floral attractants and

230

254 Rhodora [Vol. 88

rewards for pollinators, and (2) relate these floral characteristics to
the pollinator fauna.

MATERIALS AND METHODS

We observed bird’s-eye-primrose plants and their insect visitors
throughout the flowering season in 1983 and returned later to col-
lect fruit from marked individuals. We focused on several fairly
dense populations totalling about 10,000 individuals located on the
southern headlands of Great Wass Island (Beals Township, Wash-
ington County). Voucher specimens of the three major pollinators
were deposited in the insect collection of the Department of Ento-
mology at the University of Maine at Orono.

To determine the source of the floral odors, we followed a method
of Faegri and van der Pil (1979). We divided 30 corollas into the
limb, the yellow center (the bird’s-eye) and the tube, placed these
parts into three separate small glass vials, and then three people
sniffed the vials and recorded their perceptions. To document the
appearance of Primula flowers in ultraviolet (UV) light, we photo-
graphed the same inflorescence in normal light and then through a
filter (Kodak Wratten 18A), which allows passage of UV light.

We fixed flowers in FPA (formalin 5: propionic acid 5: 50%
ethanol 90) and followed a standard dehydration in a graded alco-
hol series, embedding in paraffin, sectioning with a rotary micro-
tome at |2um and staining with safranin and fast green.

To determine pollen-stigma compatibility, we completely enclosed
I! plants in cages constructed of fine-mesh aluminum screening
from before to after anthesis. We self-pollinated the flowers of seven
plants with a camel’s-hair brush and left the remaining plants
undisturbed.

We used Dowrick’s (1956) technique to determine stigma recep-
tivity. First we emasculated caged flowers, selfed or crossed them
and stored them with moistened paper in a small, closed jar. After
24 hours we fixed them in FPA and stored them in 70% ethanol. We
then softened the gynoecia for two hours in 8N sodium hydroxide
and stained them for four hours in 0.1% aniline blue in 0.1 N potas-
sium phosphate. We observed squashed gynoecia in a fluorescence
microscope equipped with a Zeiss filter set 48 77 05 to determine if
pollen tubes were present.

1986] Campbell, Famous & Zuck — Primula 255

Pollen-ovule ratio estimates come from a count of all the pollen
from one anther (multiplied by five) and all the ovules from the
same flower for six different flowers. We measured scape lengths for
flowering plants and a different sample of fruiting plants from the
same population. For seed weight and dimensions we weighed 120
seeds collectively and measured length and width of 120 seeds lined
up end to end and side by side respectively.

RESULTS AND DISCUSSION

In eastern Maine, Primula laurentiana flowers in May and June.
The flowers (Figure 1) are about one cm broad, and the corolla,
which is pink to lilac with a bright yellow spot at its center, is
conspicuous in the full sun, wherein this species usually grows. The
flowers are also conspicuous because of the paucity of other insect-
pollinated flowers nearby. Sedum rosea (L.) Scop. and Viola septen-
trionalis Greene are the only entomophilous species that flower
during the peak of flowering of Primula and which also grow inter-
mixed with Primula on Great Wass Island.

Patterns of floral UV reflectance and absorption provide a poten-
tially important pollinator cue, although it should be interpreted
with caution (Kevan, 1983). In UV light the corolla limb of Primula
laurentiana is strongly absorbing except for a ring at the center
(Figure 2). This ring, which corresponds to a portion of the yellow
bird’s-eye of normal light, actually consists of five UV-reflecting
bands alternating with the petals. Generally, yellow petals are UV-
reflecting, but the usual pattern of UV-reflecting pigmentation sur-
rounding the UV-absorbing center of flowers (Silberglied, 1979) is
reversed in the bird’s-eye-primrose. The corolla tube is almost
always in shadow, making it dark at all wavelengths.

We do not know of any studies of the vision of Helophilus, the
principal pollinator of P. /aurentiana (see below). However, all four
genera of the Diptera listed by Goldsmith and Bernard (1974) have
either primary or secondary sensitivity maxima in the near UV.
Hence the UV patterns of Primula laurentiana may guide its polli-
nators to the corolla tube.

Frohlich (1976) documented the striking contrast between many
UV-absorbing flowers and their UV-reflecting background. The
background vegetation in Figures | and 2 consists mostly of grasses

256 Rhodora [Vol. 88

Figures 1-4. Primula laurentiana. 1. Inflorescence in normal light, approx. life
g

size; 2, Same inflorescence in near-UV light; 3. Photomicrograph of a portion of a
sectioned flower showing the anther (A), corolla tube (C), and stigma (S). Bar scale
500 um; 4. Fluorescence photomicrograph of self pollen and pollen tubes in a
squashed stigma. Bar scale = 50 um.

and sedges. The fact that this vegetation reflects UV light may facili-
tate pollinator recognition of Primula flowers.

The yellow rim of the corolla tube seemed to have the strongest
fragrance to all three people in the scent comparisons. We found

1986] Campbell, Famous & Zuck — Primula 257

uniseriate, multicellular trichomes in the epidermis of the same part
of the corolla. These structures appear to be secretory, but, accord-
ing to Fahn (1979), such structures generally are not responsible for
floral fragrances.

Our observation that the flowers of the bird’s-eye-primrose are
long homostylous—the stamens are located at the same level as or
(more usually) slightly below the stigma near the top of the corolla
tube (Figure 3)—is consistent with those of Fernald (1928) and
Dorwick (1956). Our sections also do not reveal any prominent
nectaries, nor could we detect with a 20X hand lens any nectar
sources in living flowers. This lack of nectar leaves pollen as the
major reward for pollinators.

During warm (above 12°C) dry and calm weather we noted six
species of insects visiting bird’s-eye-primrose flowers. Three of
these—a butterfly, Cynthia cardui (L.) (Nymphalidae), a large fly,
and a small syrphid fly—spent little time on Primula and did not
appear to be effective pollinators. Two species of bumblebee ( Bom-
bus) and a syrphid (Helophilus groenlandicus) on the other hand,
spent considerable time on the flowers (Table 1). Although we did
record a similar total number of flowers visited by the syrphid and
the two bumblebees, almost two-thirds of the flowers visited by the
latter were on one foray. The syrphid, a circumboreal species, was
clearly the most consistent flower visitor. These flies deliberately
and repeatedly probed the corolla tube while sitting on the limb. We
opened the gut of one individual collected at the end of a 10-flower,
four-minute foray and found an abundance of Primula pollen. With
their short proboscis, syrphids are generally limited to the consump-
tion of pollen rather than nectar (Faegri and van der Pijl, 1979).
Both bumblebees and this syrphid were flower constant: they did
not alight on non-Primula flowers during their forays on Primula.
Helophilus groenlandicus did visit Sedum rosea, but only after the
peak of Primula anthesis.

Controlled self-pollinations show bird’s-eye-primrose to be self-
compatible. Self pollen germinates freely (Figure 4) and grows to
the ovules within 24 hours. In contrast, in self-incompatible pollina-
tions in Primula obconica, the pollen tube rarely penetrated the
stigma in 24 hours (Dowrick, 1956). Foreign pollen performed sim-
ilarly to self pollen in P. /aurentiana.

The stigma is usually green when the anthers dehisce but later
turns dark red. Pollen tubes penetrate the stigma before and after it

258 Rhodora [Vol. 88

Table |. Pollinator observations of Primula laurentiana

Total days of observation 10
(12 May-12 June, 1983)

Total hours of observation 38.5

Helophilus Two Species of
Pollination Activity groenlandicus Bombus
Total number of forays* 101 13
Mean (S.D.; range) flowers/ foray 7.6 (7.2; 1-44) 51.9 (117.4; 4-439)
Mean (S.D.; range) seconds/ flower 7.4 (5.9; 2-42) 2.6 (1.1; 3-6)
Total flowers visited 768 675
Total minutes on flowers 94.7 29.2

“A foray is one uninterrupted sequence of floral visitations by one individual insect.

turns red. Protandry, and its potential for outbreeding, therefore
does not occur in Primula laurentiana. Since the flowers open just
before the anthers dehisce, protogyny seems unlikely.

Homostyly, the simultaneous maturity of anthers and stigmas,
and the fact that the stamens are introrse (Figure 3) ensure that the
syrphid and bumblebees must self-pollinate bird’s-eye-primrose dur-
ing their often protracted pollen-gathering visits. Our estimates of
pollen-ovule ratios (133+ 15) are close to Cruden’s (1977) mean
value (168.5 + 22.1) for his breeding system of facultative autogamy.
We don’t know how much pollen adheres to pollinators’ bodies
and thereby effects cross-pollination.

In general, caged Primula laurentiana sets seed when artifically
selfed or when not touched. How does fertilization occur in these
untouched plants? We could not find within any flowers small
insects such as thrips or small beetles, which might have passed
through the cage and which have been suggested as possible pollina-
tors in other species of Primula (Woodell, 1960). We suspect that
the wind blows the flowers against the cages and forces the anthers
against the stigma. We don’t know if such “wind” pollination occurs
normally.

Mean scape length increases significantly (t = 12.96, P > 0.01;
Zar, 1974) between anthesis (8.7 + 2.8 cm) and fruit maturation
(15.7 + 2.8 cm). The seeds are light (0.0607 mg) and small (0.473 <
0.653 mm). The erect capsules open apically, and the seeds are

1986] Campbell, Famous & Zuck — Primula 259

gradually released in a “salt shaker” fashion. They do not show
obvious adaptations for wind or animal dispersal, nor do they float.
Self-compatibility would facilitiate the establishment of new popu-
lations from individuals isolated after long-distance dispersal.

Primula laurentiana is morphologically very similar to P. farinosa
L., the bird’s-eye-primrose of the mountains of Europe (C.S.C.,
personal observation of plants in Austria). Primula laurentiana is
octoploid or hexaploid (Dorwick, 1956; Vogelmann, 1960) and
homostylous, as in some other polyploid Primulas (Vogelmann,
1960; Dowrick, 1956). Polyploidy may actually have provided a
greater opportunity for the recombinations necessary to produce
homostyly (Dowrick, 1956). Primula farinosa is characterized by
the canstrasting suite of ancestral character states: diploidy, heteros-
tyly and self-incompatibility.

ACKNOWLEDGMENTS

We thank C. D. Richards and D. R. Whitehead for help in the
field, E. A. Osgood of the Department of Entomology of the Uni-
versity of Maine at Orono and F. C. Thompson of the U.S.D.A.
Systematic Entomology Department for insect identifications, and
M. W. Frohlich and B. St. J. Vickery for comments on an early
draft. This work was supported by a grant from the Maine Chapter
of The Nature Conservancy.

LITERATURE CITED

Cruben, R. W. 1977. Pollen-ovule ratios: a conservative indicator of breeding
systems in flowering plants. Evolution 31: 32-46.

Dowrick, V.P. J. 1956. Heterostyly and homostyly in Primula obconica. Hered-
ity 10: 219-236.

FAEGRI, K. AND L. VAN DER Pist. 1979. The principles of pollination ecology. 3rd
ed. Pergamon Press Ltd. Oxford, England.

FAHN, A. 1979. Secretory Tissues in Plants. Academic Press. London.

Famous, N. C. AND C. S. CAMPBELL. 1984. Lomatogonium rotatum (Gentiana-
ceae) and Primula laurentiana (Primulaceae) in Maine: New localities and gen-
eral distributions. Rhodora 86: 425-429.

FERNALD, M.L. 1928. Primula section Farinosae in America. Rhodora 30: 59-77,
85-104.

FROHLICH, M. W. 1976. Appearance of vegetation in ultraviolet light: Absorbing
flowers, reflecting backgrounds. Science 194: 839-841.

GANDERS, F. R. 1979. The biology of heterostyly. New Zealand J. Bot. 17:
607-635.

260 Rhodora [Vol. 88

Go-psMITH, T. H. ANDG. D. BERNARD. 1974. The visual system of insects. Jn: M.
Rockstein, Ed., The Physiology of Insecta. Vol. 2, 2nd ed. Academic Press, New
York and London. Pp. 165-272.

Kevan, P.G. 1983. Floral colors through the insect eye: What they are and what
they mean. /n: C. FE. Jones and R. J. Little, Eds., Handbook of Experimental
pollination Biology. Science and Academic Editions, New York. Pp. 3-30.

ScoGGAN, H. J. 1979. The Flora of Canada. Part 4—Dicotyledoneae (Loasaceae
to Compositae). National Museum of Natural Sciences Publications in Botany,
No. 7 (4).

SILBERGLIED, R. E. 1979. Communication in the ultraviolet. Annu. Rev. Ecol.
Syst. 10: 373-398.

VOGELMANN, H. W. 1960. Chromosome numbers in some American farinose
Primulas with comments on their taxonomy. Rhodora 62: 31-42.

WoopvELL, S. R. J. 1960. What pollinates primulas? New Scientist 8: 568-571.

Zar, J. H. 1974. Biostatistical analysis. Prentice Hall. Englewood Cliffs, New
Jersey.

DEPARTMENT OF BOTANY AND PLANT PATHOLOGY
UNIVERSITY OF MAINE
ORONO, MAINE 04469

THE CYTOGEOGRAPHY OF CHRYSOPSIS MARIANA
(COMPOSITAE: ASTEREAE): SURVEY OVER
TREsRNGE OF. THE SPECIES

JOHN C. SEMPLE AND C. C. CHINNAPPA

ABSTRACT

Chromosome numbers determined from 49 populations of Chrysopsis mariana
(L.) Ell. ranged from diploid to octoploid: 27 = 8, 2n = 16, 2n = 24, 2n = 32
respectively. Seedlings from one 6x population had 5-9 small supernumerary chro-
mosomes. These counts plus 8 previously reported counts indicate an allopatric distri-
bution pattern of cytotypes throughout the range of the species. All cytotypes were
found in Florida, whereas only hexaploids occurred in other parts of the range.
Diploids occurred in two disjunct areas of Florida. Tetraploids and octoploids were
found only in the northeastern portion of the Florida peninsula.

Key Words: cytogeography, polyploidy, goldenaster, Chrysopsis, Astereae

INTRODUCTION

Chrysopsis mariana (L.) Ell. is the most widely distributed species
in this recently revised genus (Semple, 1981). The first chromosome
number reported for this species was 2n = 24 (Smith, 1966). Harms
(1974) reported a meiotic count of m = 12 and postulated that the
species was a tetraploid with base number of x = 6. Semple (1977)
reported counts of 2” = 8, n = 12, and 2m = 24 and concluded that
the base number was x = 4. Semple and Chinnappa (1980) reported
the occurrence of an octoploid plant with 2n = 32. The diploid
karyotype was illustrated and described in Semple and Chinnappa
(1980) along with karyotypes of the other species in the genus. The
existence of the tetraploid race was noted by Semple (1981), but no
locations or vouchers were cited. Field work during the last several
years has resulted in an additional 49 populations being sampled.
The chromosome number determinations and their geographic dis-
tributions are reported for the first time in this paper.

MATERIALS AND METHODS

Meiosis was observed in pollen mother cells (PMC’s) prepared as
follows. Capitulescence buds (heads) were fixed in the field in 3:1
absolute alcohol: glacial acetic acid, then stored in 70% EtOH under
refrigeration until examined. Anthers were dissected out of florets

261

262 Rhodora [Vol. 88

removed from the heads and squashed in 1% acetic orcein stain
under a coverslip coated with egg albumin in glycerin. Observations
were made on fresh preparations. Permanent slides were made by
floating off the coverslip with adhering cells in 10% acetic acid, then
dehydrating in a 70%-95%-100% EtOH series before mounting on a
clean slide with Euparol. Permanent slides remain in the possession
of the first author.

Mitotic counts were determined from root tip cells prepared as
follows. Achenes collected in the field were germinated in vermicu-
lite in the greenhouse facilities at the University of Waterloo.
Actively growing root tips were removed from seedlings, pretreated
in 0.01% colchicine for 2 hrs., then fixed in Modified Carnoy’s
Fixative (4:3:1 / chloroform: absolute EtOH: glacial acetic acid, by
volume) and stored without solution change at -4° C. Before
squashing, root tips were hydrolized in IN HCI for 30 min at 60° C.
The meristematic tips were then squashed as above. Permanent
slides were made as above.

RESULTS

Chromosome number determinations from seedlings and wild
individuals from 49 populations are listed in Table I. A dozen
diploid populations were found in Florida; four tetraploid popula-
tions were sampled in northeastern Florida; one octoploid popula-
tion was sampled near the coast in northeastern Florida. Twenty-nine
hexaploid populations were sampled from the southern and central
portions of the range excluding Florida. Seedlings grown from
achenes collected at one population in eastern Virginia were found
to have varying numbers of small euchromatic supernumerary
chromosomes (2n = 24 + 5-9 supernumeraries).

DISCUSSION

In total, 57 populations of Chrysopsis mariana have been
sampled cytologically, including those reported previously. The
geographical distribution of the cytotypes is illustrated in Figure 1.
The range of the species is indicated by broken lines (Semple, 1981).
Although some areas of the range are sparsely sampled, the overall
distribution pattern of cytotypes is clear. Based on a much smaller
sample of published and unpublished counts, Semple (1981) noted

1986]

Table 1.

Semple & Chinnappa — Chrysopsis 263

Chromosome number determinations and voucher data for CArysopsis

mariana from the United States. All collections by Semple & Chmielewski unless
otherwise indicated: Bt = L. Brouillet, C = J. Canne,S = J. Semple, S&S = J. & B.
Semple. Vouchers in WAT.

COUNT
2n =

COLLECTION LOCATION AND VOUCHER DATA (WAT)

Florida. Osceola Co.: N of Davenport, S 5344.

Florida. Calhoun Co.: W of Clarksville, S & Godfrey 3/05. Lafayette Co.:
S of Cooks Hammock, S & S 7444. Liberty Co.: W of Bloxham, § & S
7424. Madison Co.; US-221, 4.9 mi N of county line, S, Br & C 39/4,
Pasco Co.: US-301, just N of county line, S & S 54/7. Sumter Co.: FL-44,
3.8 mi E of county line, S, Br & C 3976. Suwanee Co.: SE of Wellborn,
S & S 7451. Taylor Co.: W of Hampton Springs, S & S 744/. Wakulla
Co.: N of Ivan, S & S 7426; N of Panacea, S & S 7439. Washington Co.: E
of Bonifay, S & Godfrey 3181.

Florida. Levy Co.: N of Inglis, S, Br & C3958. Putnam Co.: FL-19, 0.5 mi
N of Oklawaha River, S, Br & C 3958.

Florida. Duval Co.: SE of Callahan, S & S 7546. St. Johns Co.: W of St.
Augustine, S & S 7544.

South Carolina. Chesterfield Co.: S of Pageland, S, Brammall & Hart
3036. Williamsburg Co.: N of Rhems, 6/24.

Alabama. Etowah Co.: N of Attalla, 6302. Randolph Co.: N of Wedowee,
63/3. Tuscaloosa Co.: Northport City, 6369. Wilcox Co.: S of Millers
Ferry, 6360. Florida. Gadsden Co.: E of Chattahoochee, S & Godfrey
3284; US-90 6.3 mi W of county line, S & Godfrey 3186. Jackson Co.:
Marianna, S & Godfrey 3/83. Georgia. Coffee Co.: SE of Broxton, S & S
7413. Evans Co.: S of Claxton, S & S 7560. Lowndes Co.: W of Valdosta,
S & S 7418. White Co.: N of Robertstown, 62/4. Wilcox Co.: NW of
Abbeville, S & S 74//. Kentucky. Whitley Co.: KY-92 2.8 mi E of county
line, S, Brammall & Hart 2994. Mississippi. Alcorn Co.: E of Strictland,
6348. Rankin Co.: US-49 3 mi N of road to Star, S, Br & C 3808. New
Jersey. Burlington Co.: N of Chatworth, 6258. Cape May Co.: NE of
Dennisville, 6253. North Carolina. Catawba Co.: S of Millerville, S, Br &
C 4076. Henderson Co.: W of Chimney Rock, S, Brammall & Hart 3029.
Transylvania Co.: E of Connestee, 6/75. Wayne Co.: NW of Cliffs of
Neuse State Park, 6044. Wilkes Co.: McGrady, S, Br & C 408]. Ohio.
Scioto Co.: Shawnee State Forest, S, Brammall & Hart 2983. South
Carolina. Chester Co.: SE of Chester, 6097. Richland Co.: NW of Colum-
bia, S, BT & C 4067. Tennessee. Morgan Co.: W of Rugby, S, Brammall &
Hart 3006. Virginia. Culpepper Co.: NW of Lignum, 5957. Madison Co.:
SW of Culpepper, 5947. Middlesex Co.: NE of Harmony Village, 5975.

24 + 5-9 supernumeraries

32

Virginia. Essex Co.: SE of Chance, 5967.
Florida. Flagler Co.: N of Bunnell, S 7535.

264 Rhodora [Vol. 88

that hexaploids were by far the most widely distributed cytotype
and that there were a few counts of other cytotypes from popula-
tions in Florida. The new data reported in this paper indicate that
within Florida, hexaploids are in fact restricted to a small area
northwest of Tallahassee. This area is not floristically like areas to
the south (R. K. Godfrey, pers. comm.). Based on the present sam-
ple, we interpret the hexaploid count reported from the Merritt
Island area of Florida (Semple, 1977; indicated in Figure | by an
open circle) to be a possible F; hybrid between a tetraploid and an
octoploid, both of which are now known to occur in that area.
Otherwise, the count indicates that there is a disjunct population of
hexaploids in that part of Florida.

The new data also indicate that within Florida, the 2x, 4x and 8x
cytotypes are allopatric. Furthermore, the distribution of these cyto-
types fits the general pattern of distribution in Florida of the x = 5
species for the genus Chrysopsis as a whole. The x = 5 diploids are
distributed in three patterns (Semple, 1981): 1) only in the panhan-
dle area (e.g., C. Januginosa Small); 2) only in the peninsula area
(e.g., C. latisquamea Pollard); or 3) as disjunct panhandle and
peninsula populations (e.g., C. /inearifolia Semple). Several of these
general patterns are represented in C. mariana. The diploids show
disjunction, the last type of distribution pattern. Nine 2x popula-
tions occur in the Florida panhandle area and three were found in
the central peninsula area. Tetraploids were found only in the
northern peninsula area between the two groups of diploids, and
thus have the second distribution type. The two 8x samples came
from near the Atlantic coast on, or north of, Merritt Island. The 8x
plants are unusual in that they have very large heads for the species.
Such plants occur only in eastern Florida and thus also have the
second type of distribution pattern.

In conclusion, it appears that each cytotype is endemic to a par-
ticular part of Florida. One of these areas, the region northwest of
Tallahassee, is the southern end of a broadly distributed floristic
region and thus by chance the hexaploid race is much more broadly
distributed than the other cytotype races of the species. The sample
size within Florida, however, is relatively small, and a much larger
sample may reveal some sympatry that is not indicated by the cur-
rent data base. Geographic isolation and subsequent chance diver-
gence were hypothesized to be major factors in the evolution of the

1986] Semple & Chinnappa — Chrysopsis 265

Cytogeography of

Chrysopsis mariana DS 60" |
a ———s|
Figure |. The cytogeography of Chrysopsis mariana. Superimposed on a state

outline map of eastern United States are the range of the species (broken lines) and
the locatons of all known populations from which chromosome numbers have been
determined. A key to the symbols is included.

genus Chrysopsis (Semple, 1981). These same phenomena also
appear to account for the pattern of cytotype distribution within C.
mariana.

ACKNOWLEDGMENTS

This work was supported by National Research Council and Nat-
ural Sciences and Engineering Research Council of Canada Operat-
ing Grants to the first author. The following people are thanked for
their assistance in the field: R. A. Brammall, L. Brouillet, J. Canne,

266 Rhodora [Vol. 88

J. Chmielewski, R. K. Godfrey, C. Hart and B. Semple. Jerry
Chmielewski is thanked for reading and commenting on a draft of
the manuscript.

LITERATURE CITED

Harms, V.L. 1974. Chromosome numbers in Heterotheca, including Chrysopsis
(Compositae: Astereae), with phylogenetic interpretations. Brittonia 26: 61-69.

SeMPLE, J.C. 1977. Cytotaxonomy of Chrysopsis and Heterotheca (Compositae:
Astereae): a new interpretation of phylogeny. Canad. J. Bot. 55: 2503-2513.

1981. A revision of the goldenaster genus ChArysopsis (Nutt.) Ell. nom.

cons. (Compositae: Astereae). Rhodora 83: 323-384.

AND C. C. CHINNAPPA. 1980. Karyotype evolution and chromosome
numbers in Chrysopsis (Nutt.) Ell. sensu Semple (Compositae: Astereae).
Canad. J. Bot. 58: 164-171.

SMITH, E. B. 1966. /n: IOPB Chromosome number reports. Taxon 15: 155-163.

J.C.S.

DEPARTMENT OF BIOLOGY
UNIVERSITY OF WATERLOO
WATERLOO, ONTARIO N2L 3G] CANADA

C.C.C.

DEPARTMENT OF BIOLOGY
UNIVERSITY OF CALGARY

CALGARY, ALBERTA T2N IN4 CANADA

THE PRAIRIE FRINGED ORCHIDS:
A POLLINATOR-ISOLATED
SPECIES PAIR!

CHARLES J. SHEVIAK AND MARLIN L. BOWLES2

ABSTRACT

Plants referred in the past to Platanthera leucophaea (Nutt.) Lindl. have been
found to comprise two species differing in pollination mechanisms, morphology, and
geographic distribution. The more eastern species, P. leucophaea, is lectotypified,
and the more western species is formally described as P. praeciara. Relationships of
the species are discussed.

Key Words: Orchidaceae, Platanthera leucophaea, Platanthera praeclara, lectotype,
pollination, evolution, biogeography

INTRODUCTION

The prairie fringed orchids, Platanthera leucophaea (Nutt.) Lindl.
auct., are some of the showiest plants in the genus. They are among
only a few orchid species characteristic of the prairies of the central
United States and are largely limited to this region. Although origi-
nally often frequent and occasionally occurring in vast numbers,
they have become increasingly rare due to the wholesale destruction
of the prairies. These circumstances have contributed to a great
interest in these plants, yet they have been little studied and less
understood. Their present rarity and characteristic drastic popula-
tion fluctuations make them difficult subjects for observation.

Our familiarity with these plants is based on studies of their ecol-
ogy (Sheviak, 1974; Bowles, 1983) and pollination biology (Bowles,
unpub.). Examination of herbarium specimens suggested that plants
from areas west of the Mississippi River were dramatically larger-
flowered than more eastern plants, and the present study was
initiated in order to assess the significance of this difference.

'Published as New York State Museum Journal Series No. 438
?Present address: Natural Land Institute, 320 South Third Street, Rockford,
Ilinois 61108 and The Morton Arboretum, Lisle, Illinois 60532.

267

268 Rhodora [Vol. 88

GARDEN AND FIELD STUDIES

A few plants from Illinois and Wisconsin had been cultivated by
the authors for a number of years; others from North Dakota were
collected for this study. In 1982 a plant from North Dakota and two
from Wisconsin bloomed in cultivation at Albany and allowed
comparison and experimentation. Forty-five populations also were
studied in the field over a broad geographic area, including stations
in North Dakota, Nebraska, Kansas, Minnesota, lowa, Wisconsin,
Illinois, Michigan, Ohio, and Ontario. These studies aided the
assessment of the morphological differences noted in the cultivated
plants.

Floral Characters

Under uniform light and temperature regimes, the first flowers on
both the eastern and the western plants opened on the same day,
suggesting no difference in phenology. The size differences apparent
in herbarium specimens were evident also in live material (Figure 1),
and floral characters including color, fragrance, shape of the lateral
lobes of the lip, petal shape, and column structure also differed. The
significance of most of these characters could not be judged from
the small sample, but the differences suggested features for subse-
quent analysis. Differences in column structure, however, were of
obvious significance, for the importance of column form in Platan-
thera speciation is well established (Stoutamire, 1974; Inoue, 1983).

Columns of the eastern plants were comparatively small and
rounded. Pollinaria [hemipollinaria sensu Dressler, 1981] were
closely spaced, the caudicles parallel, the viscidia directly below the
pollinia, facing each other, and only |.2-3.2 mm apart in the culti-
vated plants. In contrast, the larger columns of the western plant
were noticeably angular, the caudicles widely diverging and directed
somewhat forward, with the viscidia 6.2-7.5 mm apart. Field studies
confirmed these differences in column structure and indicated that
they are constant across broad geographic areas. It is apparent that
the two column morphologies result in different pollinaria place-
ment on pollinators and serve to isolate the plants through mechan-
ics and pollinator specificity. Hence, the differences in column
structure suggest that the eastern and western plants comprise dif-
ferent species.

1986] Sheviak & Bowles — Platanthera 269

Figure |. Inflorescences and flowers of Platanthera praeclara and P. leucophaea.
a-d: Platanthera praeclara. a-c: Sheviak 2222a, Richland Co., North Dakota. d:
Bowles 501, Ransom Co., North Dakota (isotype). e-h: Platanthera leucophaea. e-g:
Sheviak & Bowles 1828a, Kenosha Co., Wisconsin. h: Bowles s.n., Kenosha Co.,
Wisconsin.

270 Rhodora [Vol. 88

Cytology

Chromosome numbers were determined for two cultivated plants
each from North Dakota (Sheviak 2222a, 2222b: NYS) and Wiscon-
sin (Sheviak & Bowles 1828a, 1828b: NYS). Mitotic figures were
obtained from ovules and root tip meristems as in Sheviak and
Catling (1980) and Sheviak (1982). All numbers were found to be
2n = 42, and although this meager sample does not provide evidence
on cytological conditions throughout the range of the plants, it does
indicate that cytology is of no significance to the taxonomic prob-
lem at hand.

Hybridization

Cultivated plants blooming in 1982 were artificially hybridized
and selfed to determine compatibility of eastern and western plants.
Although limited, the results suggest compatibility. Copious seed
resulted from all pollinations, and capsules appeared fully deve-
loped. Microscopic examination suggested that seed was compara-
ble and highly fertile irrespective of parentage.

HERBARIUM STUDIES

Specimens were borrowed from herbaria located throughout the
range of the plants. Virtually all specimens could be easily sorted on
gross morphological differences into two groups of plants of com-
paratively eastern or western origin. Statistical analyses were then
employed to determine which characters contributed to this ease of
determination and to assess the variability and significance of the
distinctive features noted in the live plants. Thirteen characters were
measured on each of 29 eastern and 27 western specimens chosen to
represent the evident range of variation in gross morphology and
selected from across the plants’ geographic ranges. Specimens
chosen for measurement and their geographic origins are cited in
Table |. Characters measured and a summary of the measurements
obtained is presented in Table 2. Most of these characters are self-
explanatory, but a few require some discussion.

Spur maximum diameter measures the broadest part of the distal
enlargement in the flattened state. Petal apex length is the length of
the petal beyond its broadest point.

Column structure was severely distorted by pressing in all speci-
mens examined, and after moistening with warm water, concen-

1986] Sheviak & Bowles Platanthera 271

trated ammonia, strong saline, or other solutions, no clear
indication of the original position of the viscidia was evident.
Another character which might provide data related to viscidia
orientation was consequently sought. In addition to a pronounced
trend toward larger column size in the western plants, the shape of
the columns when viewed from the side differs markedly (Figure 1).
The rostellum lobes, which bear the caudicles and viscidia, are pro-
nounced in the western plants, thrusting downward and forward in
pressed material and providing the column with a strongly angular
appearance. In eastern plants, the lobes are much smaller and give
the column a more compact and rounded appearance. These differ-
ences-between columns of the two groups of plants are directly
related to viscidia placement. Accordingly, a measurement was
made of the height of the column from the apex of the lobe to the
crown Of the pollinium; this measurement most nearly assesses the
key character of position of the viscidia.

Results of morphological analyses

Data derived from the 56 measured specimens were subjected to
principal components analysis (SPSS: Factor). Two distinct groups
were recognized by PCA (Figure 2), and these groups correspond to
the eastern and western groups which had been established by sub-
jective consideration of gross morphology. This correlation extends
even to a few atypical or seemingly out-of-range specimens which
had been determined by consideration of a combination of charac-
ters stressing column morphology. The PCA results support these
determinations and further indicate the validity of the eastern and
western concepts.

Utilizing the groupings established by subjective study and PCA,
a stepwise discriminant function analysis (SPSS: Discriminant) was
performed in an effort to determine the significance of individual
characters and to aid in determination of herbarium material (Fig-
ure 3 and Table 2). Again, in DFA two groups are clearly estab-
lished, and these groups correspond to those obtained by subjective
determination and by PCA. Significant characters for determina-
tion were, in order of decreasing importance: lateral sepal width,
column height, inflorescence length, spur length, lateral sepal
length, and flower number. These results establish the utility of
column height as a character and also identify several other charac-
ters that clearly contribute to subjective determination.

272 Rhodora [Vol. 88

Table 1. Specimens of Platanthera leucophaea and P. praeclara employed in
statistical analyses by subjectively determined group.

Locality [County] Specimen

Eastern (P. leucophaea)

ONTARIO
Huron Morton s.n. (DAO 328245b)
Middlesex Burgess 1769 (DAO)
Simcoe Walshe 143 (DAO)
NEW YORK
Oswego Wibbe s.n. (NYS)
Wayne Hankenson s.n. “a” (NYS)
NEW JERSEY
Sussex Rushby s.n. (F349473)
OHIO
Stark Reihl s.n. 7/1835 (MO)
MICHIGAN
Cheboygan Erlanson 481c (MICH)
Cheboygan Ehlers 2507a (NYS)
Eaton w/o collector (NYS)
Tuscola Case s.n. 7/14/61 (MICH)
INDIANA
Elkhart Deam 57927 (IND)
Hamilton Shipman 2765 (MICH)
Lagrange Deam 20669 (IND)
Laporte Deam 55231b (IND)
WISCONSIN
Jefferson Wadmond 1504 (MINN)
ILLINOIS
Cook A. Chase 138la (ALL)
Cook McDonald s.n. 6/1886 (ILL)
Cook Hill 58, 1875b (ALL)
Cook Babcock s.n. (F33398)
Fayette Benjamin s.n. 6/14/1948 (ILL)
Kankakee Hill 231, 1873 ALL)
Madison (?) Eggert s.n. [ex Denslow 142] (NYS)
McDonough Young s.n. (IND)
McDonough Myers 632 (MWI)
Stark V. Chase 665 (ALL)
Winnebago Bebb s.n. (F 62188)
IOWA

Henry

Mills 1843 (MO)

1986] Sheviak & Bowles — Platanthera

NEBRASKA
Dodge Engberg s.n. (NEBI69756)

Western (P. praeclara)

MINNESOTA

Clay Ottoson 98 (MINN)

Faribault Shimek s.n. (NEB5S9440)

Fillmore Hill s.n. “a 7/1861 (ILL)

Freeborn Rosendahl et al. 7277 (MINN)

Houston Wheeler 299 (MINN)

Kandiyohi Frost 316 (MINN)

Nicollet Ballard 1067 (MINN)

Polk Ownbey 4954a (MINN)
IOWA

Black Hawk Burk 601 (ALL)

Decatur Fitzpatrick & Fitzpatrick 18 (MQ)

Dickinson Shimek s.n. (F 1564537)

Emmet Cratty s.n. (NYS)

Pottawattamie Shimek s.n. (MO1245647b)

Union Fay 3156 (TEX)

Webster Churchill 1110a (NEB)

Webster Churchill 1110b (NEB)

Winneshiek Tolstead s.n. (NEB236617)
NORTH DAKOTA

Ransom Seiler 3460 (MO)

Richland Seiler 3344 (KANU)
SOUTH DAKOTA

Minnehaha Over 7030 (SDU)
NEBRASKA

Cass Fricke s.n. (NEBI69784)

Chemung Devorak s.n. (NEB169781)

Cherry Smith & Pound 219a (NEB)

Dodge Eastman s.n. (NEB16975Sa)

Grant Bates s.n. (NEB169786)

Otoe Turrell s.n. (NEB242215)
KANSAS

Douglas Snow 8061 (KANU)

274 Rhodora [Vol. 88

Table 2. Means, standard deviations, and discriminant function character
weights for the 56 specimens of Platanthera cited in Table I.

Discriminant

Means (0) Function
East West Character
Character [/euwcophaea] [praeclara] Weights*
Column height 3.9 (0.4) 6.5 (0.8) 1.04734
Lateral sepal
length 8.1 (1.2) 12.0 (1.1) .24089
width 5.0 (0.6) 8.3 (1.0) 51681
Lip
length 18.0 (2.3) 25.6 (3.4)
width 20.5 (2.9) 30.0 (5.0)
Spur
length 35.6 (4.8) 45.7 (5.9) 08228
maximum diameter 1.8 (0.3) Zt (0.5)
Petal
length 9.6 (1.3) 13.1 (1.8)
width 5.8 (1.2) 9.5 (1.9)
apex length 2.6 (0.6) 2.7 (0.6)
Flower number 19.4 (7.9) 12.6 (4.5) .07686
Inflorescence length 122.7 (48.2) 92.1 (32.3) 02894
Plant height 676.4 (191.0) 566.3 (138.2)

*From the sum of the products subtract the constant 12.61585.

The significance of the correlation between morphological groups
and column structure is clear: the eastern and western plants repres-
ent distinct species.

TYPIFICATION OF Platanthera leucophaea

Orchis leucophaea Nutt., a product of Nuttall’s 1819 collecting in
the Arkansas Territory, was described from “Kiamesha, Red River”
(Nuttall, 1834). For several days in June of that year, he collected on
calcareous prairies in the vicinity of the confluence of the Kiamichi
and Red Rivers, in the present state of Oklahoma (Nuttall, 1821).
This site is south of the known range of the western species and is far
disjunct from that of the eastern. The apparent type collection com-
prises two sheets, one each at PH and BM. The sheet at PH bears
Nuttall’s original field label with the characteristically cryptic nota-
tion ““Fl. O * (Nutt.) Ark. Red riv.”” This specimen is an inflores-

1986] Sheviak & Bowles — Platanthera ya pe

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Principal Component 1 (58.3%)

Figure 2. Positions of 56 specimens on the first and second principal components
derived from measurements of 13 morphological characters in moistened herbarium
specimens. Solid dots = P. praeclara, open circles = P. leucophaea.

cence which, after pressing, was nearly destroyed by insects. The
sheet at BM bears a more carefully executed label, on which is
written, apparently in the same hand, “Orchis *leucophaea Red
River Prairies,” with “leucophaea” crossed out. On this sheet are
mounted two specimens, one a leafy stem with an inflorescence very
similar to that of the PH specimen and also heavily damaged by

276 Rhodora [Vol. 88

P. leucophaea P. praeclara
L Ix xr
1 M 1 |
4-7
3 =
2 4
1 +
0-

-6 -4 -2 0 2 4 6
Discriminant Score

Figure 3. Histogram of discriminant scores of 27 specimens of Platanthera prae-
clara and 29 of P. leucophaea. Also plotted are group centroids (X), the holotype of P.
praeclara (P), a topotype of P. leucophaea [Leavenworth s.n., NY] (1), and the
lectotype of P. leucophaea (L). Measurements of the lectotype were made on dry
flowers and then adjusted for the projected effects of moistening, based on the actual
changes in each measurement in the topotype. An estimate of two additional flowers
was included to compensate for a missing segment of the rachis. Failure to adjust for
dry material and for missing flowers each shifts the score to more negative values.

insects, and the other an inflorescence of P. grandiflora (Bigel.)
Lind]. with a number of flowers in excellent condition. The identity
of this second inflorescence and lack of insect damage to it indicate
a mixture of collections in the herbarium; this mixture may have led
to the name having been crossed off the label. The more complete
BM specimen and the specimen at PH, despite their poor condition,
are readily identifiable and are clearly of the eastern species. This
identity and the inclusion of the inflorescence of P. grandiflora raise
the question of authenticity of the specimens. Are these the speci-
mens Nuttall collected at the type locality, or might they be others
with which he replaced a missing type? Nuttall is known to have lost
and replaced other specimens (Stuckey, 1967). A few years after
Nuttall’s publication, the eastern plant was collected in Ohio by
Sullivant; his plants were subsequently cultivated at the Cambridge
(Massachusetts) Botanic Garden, which Nuttall curated until 1834.
Perhaps Nuttall collected the specimens on a later visit to the
garden. In order to assess this possibility, an effort was made to

1986] Sheviak & Bowles — Platanthera Zit

locate any duplicate of the type which Nuttall might have sent to
Hooker, de Candolle, or other correspondents. Nuttall apparently
did not distribute duplicates of his new Arkansas species, however
(Graustein, 1967), and, indeed, none has been located. Less direct
means of establishing the authenticity of the specimens were conse-
quently necessary.

Documentation of the presence of the eastern species in southern
Oklahoma is provided by two collections at PH and NY. One spec-
imen, from the Short herbarium at PH, is labelled “Flowering in
June, vicinity of Fort Towson” (written) and “Dr. Leavenworth,
Fort-Towson, Arkansas” (printed). Unfortunately, a note on the
sheet indicates that the label may have been interchanged with a
Sullivant specimen from Ohio. Both of the specimens involved are
clearly of the eastern species, but the possibility of confusion pre-
vents the unequivocal establishment of the identity of the plant at
the collection site. The second specimen is then useful for verifica-
tion. A part of the Torrey Herbarium at NY, it is labeled simply
“Arkansas, Dr. Leavenworth, 1837.” An army surgeon, M. C. Lea-
venworth was stationed for five years in western Louisiana and
southeastern Oklahoma. During the growing seasons of 1834 and
1835, he was stationed along the Red River, Arkansas Territory,
first in the Cross Timbers region and later at Fort Towson. Late in
1835 he was transferred to Florida and was forced to leave extensive
botanical collections at the fort. In 1836, he was transferred to Fort
Jessup, Louisiana (where he had been stationed prior to his assign-
ment in Arkansas Territory) and later to Camp Sabine. During his
years in Louisiana he collected in that state and in Texas, and he is
not known to have collected again in the Arkansas Territory
(McVaugh, 1947). He began in 1836 to ship specimens to Torrey,
and the date on the specimen evidently records the year in which
Torrey received it. Leavenworth received a shipment of his Arkan-
sas specimens while at Camp Sabine (letter to Torrey, 22 Jan 1837)
and later that year expressed his pleasure at Torrey’s receipt of
unspecified Arkansas specimens (letter to Torrey, | Jul 1837). From
what is known of Leavenworth’s travels, it is evident that the speci-
men was collected along the Red River in southeastern Oklahoma,
and it is reasonable to conclude that both specimens are from the
same locality. Fort Towson was located near the mouth of the Kia-
michi River, and hence Leavenworth’s specimens are evidently top-

278 Rhodora [Vol. 88

otypes; they clearly establish the presence of the eastern plant along
the Red River.

The Leavenworth specimens indicate that Nuttall could indeed
have collected the eastern species in Oklahoma. The specimen at PH
and the more nearly complete of the two at BM are parts of the type
collection, and the name Orchis leucophaea is properly applied to
the eastern species. The flowers of the PH specimen collectively
display all floral features; this is not the case on the specimen at BM.
This situation, together with slightly better data on the original field
label at PH and the mixture at BM, leads us to designate the speci-
men at PH as lectotype. The western species is described here:

Platanthera praeclara Sheviak & Bowles, sp. nov. (Figures la-d; 4)

Flores maximi, eburnei, albescentes. Sepala ovata vel suborbicu-
lata, 9.0-14.1 mm longa, 6.8-10.0 mm lata. Petala flabellata, trun-
cata, saepe emarginata, 9.0-16.5 mm longa, 6.5-13.5 mm lata.
Labellum 17-32 mm longum, 20-39 mm latum, penitus trilobum,
lobis flabellatis, incisis, fimbriatis. Calcar gracile, clavulatum, 36-55
mm longum. Columna lata, rostello bilobo, lobis triangularibus,
patentibus, viscidiis 6-7 mm disjunctis.

Herb erect, stout, 38-85 cm tall, glabrous throughout. Leaves
lanceolate to ovate-lanceolate, ascending, the bases sheathing the
stem, up to 26 cm long, 5 cm wide. Raceme large, showy, 4.8-11.6
cm long, 5.5-9.0 cm wide. Flowers creamy white, very large. Ovaries
slender, up to 27 mm long. Sepals ovate to suborbicular, the lateral
obliquely asymmetrical, 9.0-14.1 mm long, 6.8-10.0 mm wide. Petals
flabelliform, truncate, often emarginate, the apical margin lacerate,
9.0-16.5 mm long, 6.5-13.5 mm wide. Lip deeply three-lobed, the
lobes narrowly to broadly flabellate, the lateral sometimes very
broad and overlapping the median, deeply incised and fringed,
17-32 mm long, 20-39 mm wide. Spur slender, curving, clavellate,
36-55 mm long. Column broad, the rostellum two-lobed, the lobes
triangular, wide-spreading, the viscidia 6-7 mm apart. Chromosome
number 2n = 42.

Type: U.S.A. North Dakota. Ransom Co.: Swales in sand prairie
haymeadows being mowed for hay. Sheyenne National Grasslands,
TI34N, RS3W, Fifth Principal Meridian. 14 July 1982. M. L.
Bowles 501 (HOLOTYPE: NYS; ISOTYPE: AMES, F, ILL, NY).

1986] Sheviak & Bowles — Platanthera 279

Etymology: The epithet praeclara was chosen for its singularly
appropriate series of translations: very bright, beautiful, splendid,
glorious, distinguished, noble.

COMPARATIVE MORPHOLOGY

Platanthera praeclara is very similar to P. leucophaea in gross
morphology, but the two species are readily separated by a number
of floral characters. Most of these directly reflect the larger size of
the flowers of the former species, and these differences are apparent
in Figure | and Table 2. Other characters include flower color,
fragrance, lip and petal shape, sepal shape, flower number, and the
length and density of the inflorescence. The salient character,
column structure, has been treated in the discussions of garden and
herbarium studies and is well depicted in Figure 1.

Floral color tends to differ slightly. The lip and petals of Platan-
thera praeclara are predominantly a creamy white, and the sepals
are similar although suffused with a faint greenish cast. In contrast,
flowers of P. leucophaea appear markedly whiter due to the lip and
petals being pure white and contrasting strongly with their green
claws and wholly green sepals. Flowers of P. praec/ara often fade to
white as they age, however, and the sepals of P. /eucophaea may be
somewhat whitish.

Floral fragrance in both Platanthera praeclara and P. leucophaea
is light and very sweet, and intensifies after sunset. In the cultivated
plants, however, the fragrance of P. leucophaea is somewhat more
spicy than the very delicate scent of P. praeclara. This difference is
quite apparent and was noted by other observers.

The obvious difference in the shape of the lateral lobes of the lip
in the cultivated plants (Figure 1) is not taxonomically significant.
The broad, downward-spreading lobes of the Platanthera leuco-
Phaea flower and the much narrower lobes in P. praeclara are repre-
sentative of the species, but variation is common.

Petal shape tends to differ markedly between the species. The
petals of Platanthera praeciara are nearly triangular in outline, very
broad, truncate, and often somewhat emarginate. Those of P. leu-
cophaea, in contrast, are typically merely obovate, but occasionally
vary toward those of P. praeclara.

280 Rhodora [Vol. 88

The sepals of Platanthera praeclara are comparatively broader
than those of P. /leucophaea. The importance of lateral sepal size (as
determined by DFA), in contrast to various corolla characters is
perhaps surprising, but is easily understood. Sepal enlargement is
negligible after the flower opens, whereas the petals and lip greatly
increase in size. Hence corolla measurements will vary depending on
age of the flower, and their maximum dimensions might be envir-
onmentally influenced. The differences in bud size (reflected in sepal
measurements) contribute significantly to the different aspects of
the plants.

Flower number, inflorescence length, and apparent density of the
inflorescence are related characters which contribute to the distinc-
tive aspects of the two species. Flowers are generally fewer and the
inflorescence shorter in Platanthera praeclara than in P. leuco-
phaea. Because of the larger flower size in P. praeclara, spaces
between flowers are much reduced and the inflorescence typically
appears much more dense than in P. leucophaea. In general, P.
praeclara produces shorter, denser inflorescences of fewer, larger
buds and flowers than P. /eucophaea.

ECOLOGY

The ecology of these plants has been discussed by Bowles (1983).
West of the Mississippi River and in much of the prairie peninsula,
Platanthera leucophaea and P. praeclara occupy mesic to wet cal-
careous prairie. Further eastward, however, P. lewcophaea occurs in
a region supporting a much more diverse assortment of habitats,
and it also occurs in marshes, fens, and bogs. These plants exhibit
adaptations to periodic drought and may exist in a subterranean,
dormant or mycotrophic state for one or more years; they are noted
for dramatic, periodic mass flowerings following several-year peri-
ods of apparent absence. This behavior appears to be linked to
fire-stimulated growth and flowering, although other factors, such
as rainfall and soil moisture levels, are likely involved as well.

POLLINATION BIOLOGY

These plants are classic examples of phalaenophily (Faegri and
van der Pijl, 1971) and, more specifically, sphingophily (van der Pijl
and Dodson, 1966). The horizontally displayed flowers are noctur-
nally fragrant, white, without colored nectar guides, deeply fringed,

1986] Sheviak & Bowles Platanthera 281

bilaterally symmetrical, with extruded columns, and with nectarif-
erous spurs that are the longest of any north temperate member of
the genus. Within this general syndrome, column structures of Pla-
tanthera leucophaea and P. praecilara dictate different pollination
mechanisms. The restricted spur entrance controls pollinator posi-
tion in the “key-hole” sense of Dressler (1981), and pollinaria
placement is limited to either the proboscis or eyes, which are the
only organs to which the viscidia will adhere properly.

Sphingophily in Platanthera leucophaea is supported by Robert-
son (1893), Case (1964), and our observations. [Moth specimens to
be deposited at Entomology Laboratory, University of Wisconsin,
Madison.] Robertson is the only worker to have identified pollen
vectors [Xylophanes tersa (L.) and Eumorpha achemon (Drury):
Sphingidae] and to have described pollination mechanics. He
reported that pollinaria were deposited on the proboscis and usually
were removed one at a time, as the viscidia were only 2 mm apart
and the moth directed its proboscis to one side or the other. From
this information it is clear that his observations pertain to P. leuco-
Phaea. Pollination in P. praeclara has not been reported.

Pollination of Platanthera leucophaea and P. praeclara was stu-
died in the field from 1979 to 1983. Even when working with ultra-
violet lights and red-filtered lanterns, observation of swift-flying
nocturnal pollinators is difficult, and only three visitations by
sphinx moths have been witnessed. At P. leucophaea, Sphinx eremi-
tis (Hiibner) was collected while feeding at flowers and was found
to carry a pollinarium at the base of its proboscis; Manduca sexta
(L.) was observed and photographed while feeding but was not
collected. In both cases abundant seed set occurred, and these insects
probably were pollinators. Manduca quinquemaculata (Haw.) was
collected while visiting P. praec/ara, but pollinaria removal was not
evident, and the insect probably was not a pollinator.

In order to gain more information, pollination was studied under
controlled conditions in Rockford using potted Platanthera leuco-
phaea and P. praeclara and laboratory-reared Manduca sexta.
These long-tongued moths (see below) served as pollinators of P.
leucophaea, contacting viscidia and removing pollinaria with the
proboscis. Most moths were able to reach nectar of P. praeclara
without effecting pollination, but one individual with an abnormally
short proboscis contacted viscidia and removed pollinaria with its
eyes.

282 Rhodora [Vol. 88

Following removal of pollinaria, the caudicle taxis necessary to
orient the pollinia for stigmatic contact on subsequent visits is
markedly different in the two species. Pollinaria of Platanthera leu-
cophaea, initially born vertically, bend forward, assuming a position
nearly parallel to the proboscis, and with the pollinia only slightly
elevated. This position is necessary in order for them to be inserted
between the closely-spaced rostellum lobes and to contact the
recessed stigma. In contrast, pollinaria of P. praeclara rotate later-
ally, either toward the left or right, assuming a central position
above and slightly forward of the moth’s head. This position permits
contact with the stigma without interference from the wide-spread
rostellum lobes.

These different movements and column structures provide a phys-
ical barrier to hybridization between Platanthera leucophaea and P.
praeclara. A \aterally oriented pollinarium of P. praeclara borne ona
moth’s eye will be deflected by the rostellum lobes of P. leucophaea
and will not contact the deeply recessed stigma. Forward-directed
pollinia of P. leucophaea will be unaffected by the wide-spread
rostellum lobes of P. praeclara, but the larger flower dimensions of
the latter species suggest that the position of the pollinia close to the
proboscis may prevent their contacting the stigma elevated above
the entrance to the spur. Hence, hybridization between P. leuco-
pPhaea and P. praeclara would appear to be unlikely, even in mixed
populations.

The different pollination mechanics of Platanthera leucophaea
and P. praeclara reflect selection for different groups of pollinators.
Placement of pollinaria on eyes dictates a certain pollinator specific-
ity, for spur length and the distance between the viscidia together
limit pollinator dimensions. Proboscis length must be sufficient to
reach nectar, but not so great as to prevent the moth’s eyes from
contacting the viscidia, and the distance across the eyes must
approximate that between the viscidia. Longer-tongued moths will
be able to reach nectar without contacting the viscidia, and moths
with a small distance across the eyes will not remove pollen regard-
less of proboscis length. Measurement of spurs in herbarium speci-
mens of P. praeclara (Table 2) disclosed a mean length of 45.7 mm
(o = 5.9). The volume of nectar in unvisited flowers appears to vary
greatly, but data could not be obtained from unbagged specimens in
the field. Field experience and study of the plant in cultivation
suggest that the distal 10-15 mm of the spur are commonly filled

1986] Sheviak & Bowles — Platanthera 283

with nectar. Assuming 10 mm as an average minimum, maximum
proboscis lengths of primary pollen vectors would need to occur
within an extreme range of 30-50 mm, and more likely 35-45 mm.
Moths with proboscis lengths greater than spur length, and moths
with the distance across the eyes much less than the distance
between the viscidia will not serve as pollinators and will function as
nectar thieves.

Proboscis lengths in prairie region sphinx species vary greatly.
Gregory (1964) and Fleming (1970) listed values for several species.
During the present study, measurements were made of one or two
specimens [NYSM] each of nineteen common prairie region species.
Of these, five had reduced, non-functioning mouth parts [ Pachy-
sphinx modesta (Harr.), Paonias excaecatus (J. E. Smith), P. myops
(J. E. Smith), Smerinthus cerisyi Kby., S. jamaicensis (Drury) }; in
nine, the proboscis was too short to reach the nectar in most flowers
[10-20 mm: Amphion floridensis B. P. Clark, Ceratomia amyntor
(Geyer), C. undulosa (Wlk.), Darapsa myron (Cram.), D. pholus
(Cram.), Deidamia inscripta (Harr.), Hemaris diffinis (Bdv.); 21-25
mm: Hemaris thysbe (F.), Sphinx gordius Cram.]; and five had
proboscis lengths sufficient to reach common nectar levels [34-43
mm: Eumorpha achemon (Drury), Hyles lineata (F.), Sphinx dru-
piferarum J. E. Smith, S. kalmiae J. E. Smith, S. vashti Stkr.].
Significantly, in this last group the measurements across the eyes of
all but S. vashti were highly consistent, 5.8-6.4 mm, figures closely
corresponding to viscidia separation in Platanthera_ praeclara.
Hence, these species could serve as pollinators. In contrast, this
measurement varied greatly (3.7-6.2 mm) in the shorter-tongued
species and was not correlated with proboscis length. In §. vashti,
with a proboscis length of 34 mm, the distance across the eyes was
only 4.7 mm, suggesting that the moth might often fail to contact
viscidia, despite its moderate proboscis length. Consequently, this
species may serve as a nectar thief. In Manduca quinquemaculata
and M. sexta, proboscis lengths average 80 to 130 mm (Gregory,
1964; Fleming, 1970), and these species are very likely nectar
thieves, a conclusion supported by our pollination experiments.
Gregory’s report of a mean of 60 mm for Sphinx drupiferarum
suggests that this species may also at times function as a nectar thief,
although his more western populations may not be representative of
moths from the prairies.

284 Rhodora [Vol. 88

Flowers of Platanthera leucophaea, in contrast, deposit pollinaria
on the proboscis at precisely the correct point for later stigmatic
contact. Hence a wide range of proboscis lengths is acceptable, and
eye relationships are irrelevant. Nectar production in these plants is
highly variable (Bowles, unpubl.). The distal 10-15 mm of the spur
is dilated and accumulates a larger quantity of nectar than the more
slender tube. Assuming 15 mm as a common maximum, the mean
spur length of 35.6 mm (o = 4.8) in these plants (Table 2) suggests
that a minimum pollinator proboscis length of 15.8 mm has been
selected for in this species. Some spurs of unvisited flowers, how-
ever, Over time may become entirely nectar-filled, and shorter-
tongued moths may then obtain nectar. It may be significant that
flowers of P. leucophaea are occasionally visited by noctuids and
other moths. These species may remove pollinaria, and although
short-tongued and inefficient at nectar removal, they routinely visit
a number of flowers and may function at least as secondary vectors.

Our field observations of Manduca in natural populations of Pla-
tanthera leucophaea and P. praeclara support an hypothesis of nec-
tar thievery. In one population of P. praeclara (Adair Co., lowa),
most inflorescences showed signs of visitation (low nectar levels),
yet few pollinaria were removed and a specimen of Manduca quin-
quemaculata captured while visiting the plants bore no pollinaria. A
population of P. leucophaea (Grundy Co., Illinois) at which Man-
duca sexta was observed and photographed (but not collected) had
a high frequency of pollinaria removal.

RELATIONSHIPS

Platanthera praeclara and P. leucophaea form a species pair
closely related to another evident pair, P. grandiflora (Bigel.)
Lindl.-P. psycodes (L.) Lindl., and to two other species, P.
peramoena (A. Gray) A. Gray, and P. lacera (Michx.) G. Don.
These six species form a natural group. Their relationships have
been interpreted by Stoutamire (1974), who stressed column struc-
ture and resulting differences in placement of pollinaria on pollina-
tors. He proposed two evolutionary lines, one marked by eye-
deposited pollinaria (P. grandiflora, P. peramoena) and the other by
deposition on the proboscis (P. psycodes, P. lacera). He suggested
that the great similarity in gross morphology between P. grandiflora
and P. psycodes was not due to a close relationship but rather to
convergent specialization for diurnal lepidopteran pollination. This

1986] Sheviak & Bowles — Platanthera 285

conclusion was supported by hybridization data which suggested
that P. grandiflora was somewhat isolated from P. psycodes and P.
lacera but that these latter two species hybridized successfully and
were apparently isolated primarily by pollinator differences. His
understanding of the prairie species was incomplete, but from his
description of the column it is clear that his comments refer to P.
leucophaea. He considered its column to be somewhat intermediate
between the two primary types, and he suggested that P. /leucophaea
arose from the grandiflora-peramoena line through specialization
for sphinx pollination.

The recognition of a sphinx-pollinated species pair comparable to
the grandiflora-psycodes pair suggests that pollination mechanism
and attendant column structure may not indicate evolutionary lines
in the group, but rather may have been repeatedly selected in
response to similar pressures. This interpretation is based on the
evidently close relationship between Platanthera leucophaea and P.
praeclara, a relationship apparent from similarities in morphology
and, perhaps, habitat preferences. Although the species are strik-
ingly similar and, together, dramatically different from their rela-
tives, most of the obvious differences reflect specialization for
sphingophily and could be the result of convergence. Other floral
features not obviously part of the syndrome, however, also suggest a
relationship. These include the triangular petals of P. praeclara and
some P. leucophaea; the broad, downward-spreading lateral lobes
of the lip in P. leucophaea and some P. praeclara; and the orienta-
tion of the pollinia, which in these species are directed forward,
forming a hood at the apex of the column. These features represent
extreme developments of tendencies present in the other species of
the group; their coincident expression in these very similar species
indicates a close relationship. This relationship is also reflected in
vegetative habit, a character which also discriminates the psycodes-
grandiflora pair. Each of these pairs represents opposite extremes in
the pattern of vegetative variation in the group. Leaves of P. leuco-
Phaea and P. praeclara are comparatively short, broadest near the
base and tapering to acute apices, rigidly ascending with the broad
bases prominently infolding and sheathing the stem. In P. psycodes
and P. grandiflora, leaves are longer, broadest near or above the
middle and rounded to a blunt apex, and wide-spreading to droop-
ing. Hence, the species pairs are evident also from vegetative
features.

286 Rhodora [Vol. 88

Viscidia provide additional evidence of significance to evolution-
ary interpretation. In Platanthera, viscidia highly specialized for
proboscis deposition are elongated, whereas those deposited on eyes
are round (Inoue, 1983). Viscidia in P. /acera are linear-oblong, and
those in P. grandiflora and P. praeclara are round. Those of P.
leucophaea, however, are very similar to those of P. praeclara, and
in P. psycodes they are narrowed only slightly and are rather varia-
bly shaped, but definitely not elongated. This situation suggests that
P. leucophaea and P. psycodes are less completely adapted to pro-
boscis deposition than is P. /acera, and that these three species do
not form an evolutionary line. Stoutamire’s hybridization data do
not provide unequivocal evidence on relationships, because the
indicated isolation may either initiate or result from speciation,
furthermore, our data suggest compatibility between P. praeclara
and P. leucophaea. Consequently, it appears unlikely that the coin-
cident patterns of floral and vegetative characteristics evident in the
two species pairs are the results of repeated selective events. It is
more reasonable to view the two white-flowered species and the two
purple-flowered species as two pairs of closely related species, with
the members of each pair isolated by differences in pollination
mechanics. In each case it seems most likely that the proboscis-
depositing column type arose from the other type as an adaptation
reducing nectar thievery and increasing reproductive efficiency. This
hypothesis is supported by the biogeography of the species.

BIOGEOGRAPHY

Platanthera leucophaea and P. praeciara are essentially allopatric
(Figure 4). The western P. praeclara occupies an area largely corres-
ponding to the Missouri River drainage, the center of the tallgrass
prairie formation. In contrast, P. leucophaea occurs primarily in the
upper Mississippi River drainage and the Great Lakes region in an
area corresponding to the prairie peninsula. From this region, dis-
junct stations are scattered eastward through a zone rich in prairie
species. The Red River occurrences probably represent relictual
populations.

The prairie peninsula is a comparatively young feature which
appears to have developed in a northeastward direction following
the close of the Wisconsinan glaciation. Since then, its eastern
margins have been retreating westward, leaving behind numerous

1986] Sheviak & Bowles — Platanthera 287

Figure 4. Distributions of Platanthera praeclara (triangles) and P. leucophaea
(dots).

disjunct communities and isolated populations of prairie species.
Populations of Platanthera leucophaea in the East probably were
established during this period of pronounced prairie influence and
are consequently relictual. Insofar as the prairie peninsula is a recent
extension of the western prairies, the present distribution of P. /eu-
cophaea may have developed more recently than that of P. prae-
clara. This situation is significant and may provide evidence of the
origin of P. leucophaea.

The column of Platanthera leucophaea appears to represent an
advancement over that of P. praeclara. Proboscis deposition should
lead to greater reproductive success through utilization of a greater
diversity of pollinators. This situation may be related to the distri-
bution of P. /eucophaea in the prairie peninsula. This species may
have arisen from P. praeclara and colonized northeastward as the
prairie peninsula developed. Its ability to utilize a variety of pollina-
tors would have been of great significance as it colonized new types

288 Rhodora [Vol. 88

of habitat and encountered different faunas. Regardless of the time
of its origin, these adaptively significant features enabled P. leuco-
phaea to expand its range across an ecologically diverse and un-
stable region and to occupy a variety of habitats. In contrast, P.
praeclara remains restricted to essentially a single habitat in a region
that is biologically and climatically uniform and stable.

It is significant that Platanthera grandiflora and P. psycodes dis-
play a similar pattern. The eye-depositing P. grandiflora is relatively
restricted in range, essentially limited to the Appalachian region,
whereas the proboscis-depositing P. psycodes is widespread through-
out the glaciated Northeast. It appears that this species pair mirrors
the pattern in the prairie species. The reproductively more efficient
P. psycodes has been able to colonize a wide, diverse geographic
region, whereas its more specialized relative has been comparatively
less successful. Hence, an increase in reproductive efficiency
through evolution of less specific pollination mechanisms may bea
potential in the group which has been repeatedly exploited as a
strategy in response to climatic and biotic fluctuations.

A NOTE ON COMMON NAMES

The great popular appeal of the prairie fringed orchids and the
consideration given these plants under federal, state, and provincial
endangered species programs will doubtless result in the appearance
of new common names now that two species are recognized.
Accordingly, we appeal for adoption of the names “Eastern Prairie
Fringed Orchid” and “Western Prairie Fringed Orchid” for these
two species. These names are simple, descriptive, and based on past
usage.

ACKNOWLEDGMENTS

We thank the curators and staffs of the herbaria that lent speci-
mens for this study, including AMES, BKL, BM, DAO, DEK, F,
ILL, IND, KANU, KNOX, KSC, MICH, MIN, MO, MWI, NEB,
NY, NYS, OCLA, OKL, OKLA, OS, PH, SDU, TEX, UARK,
UMO, US, WIS. We also appreciate the efforts made to locate
Nuttall, Hale, and Leavenworth specimens and other critical collec-
tions at CGE, DUKE, DWC, E, G, K, LIV, LSU, MANCH, NLU,
NO, OXF, VDB. A number of persons provided information on

1986] Sheviak & Bowles — Platanthera 289

plants in their areas, assisted in the field, and contributed specimens
and live plants: F. W. Case, Jr., Saginaw, Michigan; P. M. Catling,
Agriculture Canada; P. Currier, Platte River Whooping Crane Hab-
itat Maintenance Trust; L. K. Magrath, College of Science and Arts
of Oklahoma; T. L. McCabe, New York State Museum; D. Roosa,
Iowa State Preserves Advisory Board; W. Smith, Minnesota Natu-
ral Heritage Program; R. Warner, North Dakota Natural Heritage
Program; T. Wieboldt, Virginia Polytechnic Institute. Location of
the Wyoming station of Platanthera praeclara was established and
brought to our attention by W. F. Jennings, Boulder, Colorado.
Live Manduca pupae were generously provided by J. Keesy, Uni-
versity of Wisconsin, Madison. We are indebted to E. A. Gossen,
New York State Museum Library, for digging out much critical
biographical and archival material, and to the New York Botanical
Garden Library for providing the Torrey correspondence for study.
Drawings in Figure | are by Linda Lobik. We wish to thank P. M.
Catling for his thoughtful review of the manuscript.

LITERATURE CITED

Bow tes,M.L. 1983. The tallgrass prairie orchids Platanthera leucophaea (Nutt.)
Lindl. and Cypripedium candidum Muhl. ex Willd.: Some aspects of their sta-
tus, biology, and ecology, and implications toward management. Natural Areas
J. 3: 14-37.

Case, F. W., Jk. 1964. Orchids of the Western Great Lakes Region. Cranbrook
Institute of Science, Bloomfield Hills.

Dresser, R. L. 1981. The Orchids: Natural History and Classification. Harvard
University Press, Cambridge.

FAEGRI, K. AND L. VAN DER PuL. 1971. The Principles of Pollination Ecology.
2nd. ed. Pergamon Press, Oxford.

FLEMING, R. C. 1970. Food plants of some adult sphinx moths (Lepidoptera:
Sphingidae). Michigan Ent. 3: 17-23.

GRAUSTEIN, J. E. 1967. Thomas Nuttall, Naturalist: Explorations in America
1808-1841. Harvard University Press, Cambridge.

GreGory, D. P. 1964. Hawkmoth pollination in the genus Oenothera. Aliso 5:
385-419.

INouE, K. 1983. Systematics of the genus Platanthera in Japan and adjacent
regions with special reference to pollination. J. Fac. Sci. Univ. Tokyo III 13:
285-374.

McVauGH, R. 1947. The travels and botanical collections of Dr. Melines Conklin
Leavenworth. Field and Lab. 15: 57-70.

NuTTAaLi, T. 1821. A Journal of Travels into the Arkansa Territory, During the
Year 1819. University Microfilms facsimile series 63. 1968.

1834. Collections towards a flora of the Arkansas Territory. Read before

the American Philosophical Society April 4, 1834. Trans. Am. Phil. Soc. 5: 161.

290 Rhodora [Vol. 88

Put, L. VAN DER AND C. H. Dopson. 1966. Orchid Flowers: Their Pollination
and Evolution. University of Miami Press, Coral Gables.

RoBeRTSON, C. 1893. Flowers and insects. Bot. Gaz. (Crawfordsville) 18: 47-54.

SHEVIAK, C. J. 1974. An introduction to the ecology of the Illinois Orchidaceae.
Ill. St. Mus. Sci. Papers XIV.

1982. Biosystematic study of the Spiranthes cernua complex. New York
St. Mus. Bull. 448.

_ AND P.M. CaTLinG. 1980. The identity and status of Spiranthes ochro-
leuca (Rydb.) Rydb. Rhodora 82: 525-562.

STOUTAMIRE, W. P. 1974. Relationships of the purple-fringed orchids Platanthera
psycodes and P. grandiflora. Brittonia 26: 42-58.

Sruckey, R.L. 1967. The “lost” plants of Thomas Nuttall’s 1810 expedition into
the old Northwest. Michigan Bot. 6: 81-94.

Coda S.

BIOLOGICAL SURVEY

NEW YORK STATE MUSEUM

THE STATE EDUCATION DEPARTMENT
ALBANY, NEW YORK 12230

M. L. B.

DEPARTMENT OF PLANT BIOLOGY
UNIVERSITY OF ILLINOIS
URBANA, ILLINOIS 61801

NEW COMBINATIONS
IN THE SOLANACEAE

RAFAEL CASTILLO AND
RICHARD EVANS SCHULTES

1. In 1977, A. T. Hunziker reestablished the generic name Merin-
thopodium Donnell Smith, distinguishing the genus from Markea
L. C. Richard by its dorsifixed anthers exceeding the corolla tube
and by the corolla which is valvate, not quincuncial in aestivation
(Hunziker: Kurtziana 10 (1977) 30-31). He cited four species from
Central America and one from Colombia. He furthermore cited
Markea vogelii Cuatrecasas from Colombia, noting that “...1 do
not find any difference between this species and Merinthopodium
uniflorum (Lundell) A. T. Hunziker, except that the anthers are
somewhat longer.”

An examination of Merinthopodium uniflorum indicates that the
lower surface of the leaves is covered with sharp-pointed rigid tri-
chomes that are basally swollen and almost a millimeter long. When
these hairs are broken, they leave very conspicuous pointed scars.
The rigidly coriaceous, subobtuse leaves also have an abundance of
glandular papillae, especially beneath.

In our work of identifying the solanaceous plates of the Real
Expedicion Botanica del Nuevo Reino de Granada, we have dis-
covered an illustration (Plate No. 7367) which we believe represents
a species hitherto included in Markea: M. vogelii Cuatrecasas. In
the United States National Herbarium there is a specimen represent-
ing this species concept—the type collection, Vogel 159 from Mon-
teredonda, Departmento de Cundinamarca, Colombia, 1800 m.,
October 13, 1956.

This collection has thin, flexible leaves that are attenuately acute.
The leaves and twigs are devoid of hairs or glandular scars. Addi-
tional collections undoubtedly will provide further differences
between these two concepts, but we believe that these two differ-
ences are sufficient to indicate that Merinthopodium uniflorum of
Mexico and Markea vogeliti of Colombia must be considered dis-
tinct species in Merinthopodium.

ral

292 Rhodora [Vol. 88

We, therefore, are making the necessary new combination:
Merinthopodium vogelii (Cuatrecasas) Castillo et R. E. Schultes,
comb. nov.

Markea vogelii Cuatrecasas, Journ. Wash. Acad. Sci. 49: 269, fig.
Ic. 1959: TypE: Vogel 159, Us.

We are pleased to express our gratitude to Dr. José Cuatrecasas
for his graciously offered help.

2. Research in identifying the plates of the Expedicion Bota-
nica has revealed the need for another new combination. Plate Nos.
3733 and 3734 represent a species hitherto known as Saracha vestita
Miers.

In 1973, J. L. Gentry reestablished the generic name Jaltomata
Schlechtendahl (Phytologia 27: 286. 1973). He made new combina-
tions for two Central American species but none for any from South
America.

Jaltomata vestita (Miers) Castillo et R. E. Schultes, comb. nov.

Saracha vestita Miers, Ann. and Mag. Nat. Hist., ser. 2, 3: 449.
1849.

During our research into the Solanaceae, we have discovered that
another species which has been included in Saracha should now be
transferred to Ja/tomata. Consequently, the following new combi-
nation is proposed:

Jaltomata glandulosa (Miers) Castillo et R. E. Schultes, comb. nov.

Saracha glandulosa Miers in Ann. and Mag. Nat. Hist., ser. 2, 3:
450. 1849.

RC.
UNIVERSIDAD PEDAGOGICA

Y TECHNOLOGICA DE COLOMBIA
TUNJA, COLOMBIA

R.E.S.
BOTANICAL MUSEUM OF HARVARD UNIVERSITY
CAMBRIDGE, MA 02138

BOOK REVIEW

Davip B. LELLINGER with photographs by A. Murray Evans. 1985.
A Field Manual of the Ferns and Fern-Allies of the United
States and Canada. ix + 389 pp. Smithsonian Institution
Press, Washington, D. C. ($45.00 cloth; $29.95 paper)

This book is a valuable addition to the manuals on ferns for both
professional and amateur botanists. The introductory material is
presented in a way that can be understood by those not familar with
the biology of ferns. The glossary at the back of the book is very
clear and has illustrations of characters difficult to describe. Once to
the keys there are a number of characters used to facilitate choices,
rather than dependence on one character. The keys are bracketed
keys and each dichotomy indicates the previous choice made, thus
steps can be retraced. The descriptions are precise; thus the book
should be useful to botanists at all levels of expertise. The innova-
tion of the hybrid charts at the end of the book should prove to be a
useful addition to fern manuals.

There are two pairs of photographs which have been misplaced:
386 and 388 (Nephrolepis exaltata and N. biserrata) are reversed as
are 399 and 400, (Marsilea macropoda and M. vestita). The photo-
graphs are excellent overall. One wishes that the penny used for
scale in 75 and 96 could have come out of the photographer’s pocket
more often since for someone unfamilar with the plants it would be
helpful to know how large or small they really are. Many of the
photographs are not very useful for diagnostic characters between
species, especially in the Lycopodium species: e.g. alopecuroides
and prostratum, and the tristachyum complex. A beginner will be
hard pressed to distinguish these species. Perhaps some diagnostic
sketches or a statement in the descriptions as to species easily con-
fused and the particular characters to check would be useful in
future editions. For the serious pteridologist there are some as yet
undescribed species in the book under Botrychium and Polypodium
but that does not decrease its usefulness for identification.

The checklist might be confusing at first since the generic names
are indented but the specific epithets are not. This makes some of
the genera difficult to find quickly and I would prefer the genera to
be in boldface or capitals.

293

294 Rhodora [Vol. 88

Overall I find the book extremely useful. It lists all the fern species
in the area covered and includes much pertinent information. An
extensive bibliography leads one on to more detailed studies. The
photographs are very attractive and each one is worth much more
space than it has been given. Indices of both common and scientific
names are provided. Much work and care has gone into the produc-
tion of this manual and it will be referred to again and again by
botanists interested in the lower vascular plants of North America.

JUDITH E. SKOG

BIOLOGY DEPARTMENT
GEORGE MASON UNIVERSITY
FAIRFAX, VA 22030

JOINT FIELD MEETING: June 15-18, 1986

The Annual Joint Field Meeting of
The Northeastern Section of the Botanical Society of America
The Philadelphia Botanical Club, and
The Torrey Botanical Club

will be held June 15 to 18, 1986 at Paul Smith’s College. Paul
Smith’s College is located in the upper Adirondacks, on the shore of
Lower St. Regis Lake. Accommodations are in dorms built for the
1982 Olympics, with private baths. Additional standard dorm
accommodations are available.

Trips to sites of interest will be led by authorities on the Adiron-
dack flora. Evening lectures will provide local background and his-
tory. Many historical and recreational facilities are located in the
vicinity.

Prior registration is advised. Information is available from:

Dr. Charles Burch

Wells College
Aurora, NY 13026

Vol. 88, No. 853, including pages 1-156, was issued January 30, 1986

295

INSTRUCTIONS TO CONTRIBUTORS TO RHODORA

Submission of a manuscript implies it is not being considered for
publication simultaneously elsewhere, either in whole or in part.

Manuscripts should be submitted in triplicate (an original and
two xerographic copies) and must be double-spaced (at least 3/8”)
throughout including tables, figure legends, and literature citations.
Please do not use corrasable bond. The list of legends for figures
and maps should be provided on a separate page. Footnotes should
be used sparingly. Do not indicate the style of type through the use
of capitals or underscoring, particularly in the citation of specimens.
Names of genera and species may be underlined to indicate italics in
discussions. Specimen citations should be selected critically, espe-
cially for common species of broad distribution. Systematic revi-
sions and similar papers should be prepared in the format of “A
Monograph of the Genus Malvastrum”, S. R. Hill, Rhodora 84:
1-83, 159-264, 317-409, 1982, particularly with reference to inden-
tation of keys and synonyms. Designation of a new taxon should
carry a Latin diagnosis (rather than a full Latin description), which
sets forth succinctly just how the new taxon is distinguished from its
congeners. Papers of a floristic nature should follow, as far as possi-
ble, the format of “Annotated list of the ferns and fern allies of
Arkansas”, W. Carl Taylor and Delzie Demaree, Rhodora 81:
503-548, 1979. For bibliographic citations, refer to the Botanico-
Periodicum-Huntianum (B-P-H, 1968), which provides standard-
ized abbreviations for journals originating before 1966. All abbrevi-
ations in the text should be followed by a period, except those for
standard units of measure and direction (compass points). For
standard abbreviations and for guidance in other matters of biologi-
cal writing style, consult the CBE Style Manual, Sth ed. (original
title: Style Manual for Biological Journals). In preparing figures
(maps, charts, drawings, photos, etc.) please remember that the
printed plate will be 4 X 6 inches; be sure that your illustrations are
proportioned to reduce correctly, and indicate by blue pencil the
intended limits of the figures. (Some “turn-page” figures with brief
legends will be 3 1/2 X 6 in.) Magnification/ reduction values given
in text or figure legends should be calculated to reflect the actual
printed size. An Abstract and a list of Key Words should be sup-
plied at the beginning of each paper submitted, except for a very
short article or note.

RHODORA ; April 1986 Vol. 88, No. 854

CONTENTS

The New England Botanical Club 800th meeting
Remarks
Letters
Holocene fruit, seed nis leaf fore ne riverine bedaninds near New Hives,
Connecticut
Lee §. Pierce and Bruce H. Tiffney
Pollination biology of Primula laurentiana (Primulacese) in Maine
Christopher S. Campbell, Norman C. Famous, and
Michael G. Zuck :
The cytogeography of Chrysopsis mariana Commedia ita Shrvey
over the range of the species
John C. Semple and C. C. Chinnappa
The prairie fringed orchids: A pollinator-isolated species pair
Charles J. Sheviak and Marlin L. Bowles
New combinations in the Solanaceae
Rafael Castillo and Richard Evans Schultes
BOOK REVIEW
A Field Manual of the Ferns and Fern-Allies of the United States and
Canada
Judith E. Skog
ANNOUNCEMENT
Annual Joint Field Meeting

157
198

229

253

261

267

291

293

<a

Instructions to Contributors : : j < ‘ ; . inside back cover

Hovora

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

Vol. 88 July 1986 No. 855

Che New England Botanical Club, Inc.

22 Divinity Avenue, Cambridge, Massachusetts 02138

Conducted and published for the Club, by
NORTON H. NICKERSON, Editor-in-Chief

Associate Editors

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WILLIAM D. COUNTRYMAN MICHAEL W. LEFOR
GARRETT E. CROW ROBERT T. WILCE

RHODORA.—Published four times a year, in January, April, July, and
October. A quarterly journal of botany, devoted primarily to the flora of
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Scientific papers and notes relating to the plants of North America
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dered. Brevity is urged; please conform to the style of recent issues of the
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Cover Illustration

An original drawing, seemingly the only one surviving and perhaps the only one
ever done for publication by Merritt Lyndon Fernald, used in part to illustrate his
article on cranberry species which appeared in RHODORA No. 48 (Fernald, M. L.
1902. The variations and distribution of American cranberries. Rhodora 4: 231-237
& Plate 40). The drawing was rescued from a wastebasket by Dr. Bernice Schubert; it
now hangs in the office of Dr. Carroll E. Wood at A. The original Plate 40 caption
reads as follows: Fig. 1, Vaccinium Vitis-Idaea; fig. 2, V. Vitis-Idaea, var. minor; fig.
3, V. Oxycoccus; fig. 4, V. Oxycoccus var. intermedium; fig. 5, V. macrocarpon.

Mbhodora

(ISSN 0035 4902)
JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 88 July 1986 No. 855

THE MORPHOLOGY AND CYTOLOGY OF
POLYSTICHUM X POTTERI HYBR. NOV.
(= P. ACROSTICHOIDES X P. BRAUNID

DAVID S. BARRINGTON

ABSTRACT

The hybrid between Polystichum acrostichoides and P. braunii is common but
overlooked in Vermont. Based on a qualitative analysis, the hybrid is structurally
intermediate for most traits, but character states are not predominantly medial or
nearer to those of tetraploid P. braunii. Indusia of the hybrid are much larger than
those of either progenitor, suggesting a summation of development programs for
indusial development. Stomates of the triploid hybrids are intermediate and nearer to
those of P. braunii in size, but those of diploid P. acrostichoides are unexpectedly
larger than those of tetraploid P. braunii. Chromosome counts confirm that the
hybrid is commonly triploid, and that non-homologous pairing is high, as reported
for other Polystichum hybrids (mean number of bivalents is 21.7). The hybrid is
described and named as P. X potteri.

Key Words: Polystichum, hybrid, fern morphology, Vermont

INTRODUCTION

During the past few years botanists have frequently encountered
the hybrid between Polystichum acrostichoides and P. braunii at
Vermont stations where both parents are common. Review of her-
barium materials has yielded a set of these hybrids from the north-
ern Appalachians, all determined as P. braunii. This hybrid now
appears to be common, at least in the Green Mountains, but over-
looked because it shares several features, notably a similar lamina
dissection, with its tetraploid progenitor, Braun’s Holly Fern (P.
braunii). In this paper, I provide basic structural, cytological, eco-
logical, and geographical documentation for the hybrid. Since I
argue that commonly occurring entities in nature need names, I am

297

298 Rhodora [Vol. 88

also providing an epithet, type designation, and Latin description.

The first and only previous report of the native hybrid between
Polystichum acrostichoides and P. braunii was by Thompson &
Coffin (1940). It was based on a single small sporophyte that they
encountered in Smuggler’s Notch, Vermont during the summer of
1937. Thompson and Coffin used 18 characters to evaluate the
hybrid. They reported that the hybrid shared seven character states
with P. acrostichoides and three with P. braunii. Of the seven char-
acter states reported as intermediate, three were nearer to P. acrosti-
choides, three were medial, and one was nearer to P. braunii. One
character state was not intermediate (leaf length was shorter than
that of either progenitor). This relatively early report provided no
information on cytology or variation in spores or sporangia.

Morzenti (1962), in developing her ideas on pseudomeiotic spo-
rogenesis, described the cytology of a Polystichum acrostichoides X
P. braunii hybrid from the garden of Harold Rugg (at Hanover,
New Hampshire), thought to be a derivative of the plant discovered
by Thompson and Coffin. At meiosis the Rugg plant had 41 bival-
ents and 82 univalents (two of four sets of chromosomes pairing).
Assuming that the P. acrostichoides parent was diploid (x = 41) and
the P. braunii parent was tetraploid (x = 82) (Léve et al., 1977), the
origin of the Rugg plant must have been via an anomalous cytologi-
cal pathway. Morzenti suggested that the Rugg plant was tetraploid
(rather than triploid as expected) because P. acrostichoides had
contributed an extra set of chromosomes, either from a tetraploid
sporophyte or, according to Morzenti (1962), from “duplication of
the normal diploid genome of that species in the hybrid” (for
instance, via an unreduced gamete). The tetraploid plant from
Rugg’s garden had some sporangia that yielded 16 giant, viable
spores. These giant spores yielded gametophytes with spermatozoa
and archegonia, but no sporophytes.

SYSTEMATIC TREATMENT

Polystichum X potteri Barrington, hybr. nov. = Polystichum acros-
tichoides (L.) Roth X P. braunii (Spenner) Fée.

D1aGNnosis: Hybrida sterilis e Polysticho acrostichoide atque P.

braunii prodiens, ab illo differt foliis pinnatisectis vix supra medium

1986] Barrington — Polystichum 299

latissimis apicem versus non contractis, carenti foliis laxis sine spo-
rangiis; ab hoc petiolis et pinnis basalibus longioribus. Indusia
hybridae indusiis parentium grandiora.

HoLotyPe: Vermont, Orange Co., Strafford, Beacon Hill; low
ledges in shady maple woods above Chadsey residence, Barrington
939: VT.

PARATYPES: The Appendix provides citations of the numerous
paratypes.

Stem prostrate, short-creeping, branching at irregular intervals,
bearing a single whorl of rigidly ascendent leaves. Leaves long-
petiolate, chartaceous to coriaceous; lamina long-lanceolate, nar-
rowed to a truncate base, apically attenuate, but not abruptly
narrowed into the fertile apical portion (Figure 1); lamina dissection
twice-pinnate, but the distal half of most pinnae merely pinnatifid to
pinnatisect (Figure 2); basal acroscopic pinnules of most pinnae
crenate to pinnatifid, one-third to twice again as long as the nearest
more distal pinnules; pinnules crenate and long-spinulose, ascend-
ent along the pinna-rachis (costa axis attached at 50-60 degrees
from the pinna-rachis). The distal half or two-thirds of each lamina
with approximate sori, (lax trophophylls not seen). Indusia irregu-
larly crenate-margined, ca. 1.0 mm in diameter. Sporangia com-
monly but not always indehiscent; tapetal remnants clinging to
sporangium interior. Spores irregular, pale.

Indument of petioles and rhizome apex a dense cover of broad-
lanceolate, amber to stramineous scales with a few short marginal
setae and scurf like that of the rachis. Rachis scurf abaxially lanceo-
late to long-lanceolate, amber to stramineous, weakly to heavily
short-setate, more or less descendent along the rachis; adaxially
filiform and stramineous. Leaf-surface indument abaxially well
developed, of long (ca. 0.7 mm) very narrow-lanceolate scurf.

EtyMoLoGy: Polystichum X potteri is named for the late Henry
Potter of West Rutland, Vermont, lifelong student of the Vermont
fern flora and one of the state’s finest naturalists, who was still
active last field season at 94.

I have found it useful to develop a set of categories for the qualita-
tive character states encountered in the hybrids. Relative to those of
its progenitors, hybrid character states are necessarily of three sorts:

300

Rhodora [Vol. 88

FLL
Then

ubité

Figures | and 2: Figure 1. Leaf silhouettes of Polystichum species: from left to
right, P. acrostichoides (Barrington #1123, original leaf length 63 cm). P. X potteri
(Barrington #939, original leaf length 77 cm), and P. braunii (Barrington #1138,
original leaf length 66 cm).

Figure 2. Pinna silhouettes of Polystichum species (left pinna of each pair from
three-quarters up lamina, right pinna from center): from left to right, P. acrosti-
choides ( Barrington #1123), P. X potteri (Barrington #939), and P. braunii ( Barring-

ton #1138). Bars = 2cm.

1986] Barrington — Polystichum 301

shared with one of the progenitors, intermediate, or anomalous
(that is, neither shared nor intermediate). Intermediate character
states are more or less variable between the two progenitors. The
less variable intermediate states are either medial (clustered at the
mean between the progenitors) or nearer one or the other of the two
progenitors.

Twenty-two qualitative characters were scored for the two species
and their hybrid (Table |). Comparison of the hybrid with its pro-
genitors revealed that the hybrid shared five character states with
Polystichum braunii, but only two with P. acrostichoides. Thirteen
character states were intermediate between the two progenitors:
three of these were nearer P. acrostichoides, eight were medial, two
were nearer P. braunii, and one was variable. A single character
state, indusium diameter, was anomalous —it was larger than that of
either progenitor. In sum, the characters assessed were variable in
their expression relative to the two progenitors.

An Anderson hybrid index (Anderson 1936, 1949) was computed
for Polystichum X potteri, using the qualitative character states
(Table 1). Contrary to Anderson’s usual approach, the index was
computed for the composite characters diagnostic of the group of
hybrids as a whole and not for the individual character states of
each hybrid. Based on the hybrid index, the hybrid is intermediate
and slightly closer to P. braunii than to P. acrostichoides (57th
percentile, given P. braunii is the 100th percentile).

An Anderson hybrid-index score was computed for the sporo-
phyte from Rugg’s garden that was presumably the source of Mor-
zenti’s tetraploid count. (Only characters available from the
herbarium sheets were possible to score.) The MICH collection was
at the 42nd percentile, and the GH collection was at the 41st percen-
tile; that is, both were slightly closer to P. acrostichoides than were
the triploid plants of the hybrid. The Thompson & Coffin specimen
at GH was scored at the 6Ist percentile, that is similar to the compo-
site index for the hybrids.

Guard-cell measurements, which were done to corroborate
ploidy-level information derived from cytology, yielded unexpected
results. Guard-cell pair length and width of the diploid species were
a bit larger than those of the tetraploid species, rather than a bit
smaller (Table 2). Guard-cell measurements of the triploid hybrids

Table 1. Structural features of Polystichum X potteri and its progenitors compared (Anderson Hybrid Index scores in
parentheses: 0 = shared with acrostichoides, | = intermediate and nearer acrostichoides, 2 = medial; 3 = intermediate and

nearer braunii; 4 = shared with braunii.)

acrostichoides (a) braunii (b)
Character (all 0) hybrid (all 4)
Petiole length long intermediate, nearer a (1) short
Lax trophophylls present shared with b (4) absent

Attitude of fertile leaves
Lamina texture

Evergreenness

Lamina base width

Point of maximum lamina
width

Lamina dissection medial

Angle of costae
Stomate length
Rachis-scale setae
Rachis-scale texture

Adaxial rachis scales

stiffly ascendent
coriaceous

evergreen

nearly = medial width

near lamina base

l-pinnate to |-pinnate,
pinnatifid

45°
long
present
dull

narrow-lanceolate to
lanceolate

shared with a (0)
medial (2)

intermediate, nearer a (1)

medial (2)
medial (2)

intermediate, nearer b (3)

medial (2)

intermediate, nearer b (3)
intermediate, nearer a (1)
intermediate, various (2)

shared with b (4)

laxly ascendent
herbaceous to chartaceous

deciduous to unevenly
evergreen

one third the medial width

above the middle of the
lamina

l-pinnate, pinnatisect

80°
short
rare
lustrous

filiform

COE

vIopoyuYy

88 ‘1OA]

Table 1. Continued

acrostichoides (a)

braunii (b)

Character (all 0) hybrid (all 4)
Abaxial lamina scurf size small medial (2) large
Abaxial lamina scurf shape filiform intermediate, nearer a (1) narrow-lanceolate
Lamina scurf frequency rare medial (2) abundant
Fertile lamina portion contracted shared with b (4) not contracted
Receptacle shape round to elongate shared with b (4) round
Sorus proximity confluent medial (2) remote
Receptacle sclerenchyma abundant shared with b (4) weak
True indusium diameter small not intermediate (not small

indexed)
True indusium shape entire medial (2) crenate
Anderson Hybrid 0 48 84

Index Totals

[9861

wnyousdjog — uo\duleg

t0€

304 Rhodora [Vol. 88

Table 2. Guard-cell pair measurements in um for Polystichum X potteri and its
progenitors (n = 30 guard cell pairs measured for each sporophyte)

PARENTS AND PUTATIVE TRIPLOIDS

hybrid
(putative
Locality acrostichoides _ triploids) braunii
Wheelock
DSB # 1172 946 1175
mean length (s. d.) 54.1 (3.48) 51.3 (3.45) 49.0 (2.67)
mean width (s. d.) 40.6 (1.93) 34.0 (2.36) 31.8 (2.81)
Strafford
DSB # 1185 939 (holotype) 1183
mean length (s. d.) 52.6 (3.20) 51.1 (3.20) 47.7 (3.79)
mean width (s. d.) 37.9 (2.57) 32.3 (2.13) 31.2 (1.98)
Barnard
DSB # 1144 1097 1119
mean length (s. d.) 54.4 (4.09) 48.0 (2.82) 46.7 (3.10)
mean width (s. d.) 37.1 (2.54) 31.0 (2.16) 31.0 (1.85)
Mean for all sites
mean length (s. d.) 53.6 (0.97) 50.0 (1.83) 47.8 (1.14)
mean width (s. d.) 38.6 (1.88) 32.5 (1.52) 31.2 (0.38)
PUTATIVE TETRAPLOID
Rugg s. n. (GH) Rugg s. n. (MICH)
mean length (s.d.) 59.8 (5.36) mean length (s.d.) 64.0 (4.43)
mean width (s. d.) 12.5 (0.78) mean width (s.d.) 42.0 (2.26)

were nevertheless intermediate and nearer to those of Polystichum
braunii. Guard-cell dimensions of the Rugg collections at GH and
MICH (both presumed to be vouchers for Morzenti’s tetraploid
hybrid) were larger than those of any other plants in the sample
(Table 2).

One unusual structural variant was seen among the hybrids col-
lected in Vermont. The basal pinnae in most leaves of Barrington
1155 (VT) are the same length as the widest pinnae in the blade, but
the next few pinnae acropetally are irregular in length: some are
short like those at the base of Polystichum braunii leaves; others are
long like those of P. acrostichoides.

1986] Barrington — Polystichum 305

GEOGRAPHY & ECOLOGY

Polystichum X potteri is known from Quebec, Nova Scotia, New
Brunswick, New Hampshire, Vermont, New York, and Pennsylvania.
In Vermont the hybrid is found in cool, wet forests from 1000 to
3000 feet. The hybrids are often found in ice-carved notches and on
wet, steep, rocky slopes—both habitats characterized by unstable,
thin-soiled substrates. Both parents are present at all of the Vermont
stations known to the author.

Polystichum braunii has a circumboreal distribution (Hultén,
1962). Compared with other circumboreal ferns, it is relatively
northern in distribution. For instance, in Europe it is confined to the
maritime parts of Scandinavia and montane regions, and in western
North America it is entirely coastal and northern. In Vermont P.
braunii is limited to wet, disturbed substrates in forests at altitudes
above 1000 feet. In contrast, P. acrostichoides is endemic to North
America. It is a common species of woodlands in eastern North
America where it is best developed on wooded, thin-soiled slopes,
which are frequently disturbed. The hybrid between the two was
encountered in Vermont at fairly high altitudes where P. braunii
(near its lower limits) and P. acrostichoides (near its upper limits)
occupy the same habitat.

Eight of the 14 Vermont sites yielded only a single hybrid sporo-
phyte despite careful searching, whereas two sites had two sporo-
phytes each, one had five, one had ten, and one had twenty-four.
Some of the sporophytes at these sites are clustered and appear to be
part of a single clone, suggesting that vegetative reproduction
increases numbers of plants in populations. However, the large
populations are for the most part well spread out over the terrain,
from which | infer that they are separate genets, not ramets derived
from branching of stems.

CYTOLOGY & SPORE VARIATION

Seventeen sporophytes yielded cytological preparations with uni-
valents. The three sporophytes yielding counts were triploids with
univalents and various numbers of bivalents ranging from 19 to 26
(Table 3; Figure 3-6). During early anaphase the bivalents are slow
to separate, resulting in the characteristic early anaphase figures

306 Rhodora [Vol. 88

Table 3. Chromosome number and pairing behavior in Polystichum X potteri

Barrington Collection

Number & Sporocyte Letter Univalents Bivalents Trivalents
939A 71 26 0
939B 81 21 0
942A 74 23 l
942B 77 23 0
942C 87 18 0
942D 79 22 0
1094A 85 19 0
Mean 79.1 21.7 0.14
(s. d.) (5.73) (2.69) (0.37)

also illustrated in European work on Polystichum hybrids (Manton
and Reichstein, 1961; Sleep and Reichstein, 1967). Chromosome
segregation is irregular in meiosis, so that micronuclei are common
in the resultant spores.

The spores resulting from meiosis are for the most part wrinkled
and irregular in shape. Giant spores resulting from inclusion of all
the meiotic nuclei in a single cell, like those observed by Morzenti
(1962), were common in some of these triploid hybrids (Barrington
909 VT). Indehiscent, pale, collapsed sporangia are useful in field
identification of the hybrids, but indusia only rarely fail to evert as
in Dryopteris hybrids (Wagner and Chen, 1965). Some of the spo-
rangia of the hybrids open as the summer progresses; the proportion
of sporangia which open varies with the sporophyte. Pale bits of
tapetal material which cling to the interior of the opened sporangia
of the hybrid, but which are absent from those of the parents, are
diagnostic. These tapetal remnants were seen in all herbarium spec-
imens of hybrids at VT.

DISCUSSION

Polystichum X potteri has been largely overlooked because it
superficially looks so much like its tetraploid progenitor, P. braunii.
Lamina dissection is probably the single most important character
used in determining northeastern North American specimens of
Polystichum and, in this character, the hybrid is intermediate and
nearer P. braunii.

Intermediacy of sterile hybrids, assumed to be Fis only, often

1986] Barrington — Polystichum 307

serves as the basis for hypothesizing hybrid ancestry of plant line-
ages (Wagner, 1983), but scoring of hybrids is often based on intui-
tion rather than on character analysis. Hybrids between tetraploids
and diploids are commonly expected to approach the tetraploid in
structure because the tetraploid is genetically represented twice, but
the diploid only once. However, all of the various possible character
states are represented among the characters surveyed in the present
hybrid. Although medial states are most common, they only
account for about one-third of the characters, and only two charac-
ters are intermediate and nearer Polystichum braunii. Since structu-
ral features are elaborated by various genetic systems, most of them
polygenic, a complex array of character states is to be expected in
hybrids. A quantitative analysis of the character states of progeni-
tors and hybrids would serve as the basis for a better understanding
of hybrid character states in P. X potteri.

The Anderson-hybrid-index scores for the Rugg specimens (GH,
MICH) were closer to Polystichum acrostichoides than was the
composite hybrid-index score. Though these data are qualitative,
(and the tie of the tetraploid count to these Rugg specimens is
circumstantial) they suggest that an extra set of chromosomes from
P. acrostichoides resulted in leaf characteristics more like those of
the diploid species. The hybrid-index score for the Thompson &
Coffin specimen suggests that it was a triploid, and consequently
probably not from the same plant as the Rugg collections (contrary
to Morzenti, 1962).

The original qualitative scoring of the hybrid (Thompson and
Coffin, 1940) differs from the present one in showing more charac-
ters shared with the progenitors and fewer medial character states.
Their emphasis was on contrasting the contribution of the progeni-
tors and not on the expression of each of the characters they used.

The unusually large indusium size of the hybrid is particularly
interesting because it is a novel character state not found in the
progenitors. In this case the larger size of the hybrid indusia suggests
that the parental genomes have been summed to program for a
longer developmental period, generating a larger indusial flange
(that is, the round top, not the stalk). The precise developmental
sequences in both parents could be compared to investigate this
idea. Indusium size is important in characterizing species of Polysti-
chum in Mexico (Smith, 1981), so that transformations in this par-

308 Rhodora [Vol. 88

Figures 3 and 4: Figure 3. Late prophase of meiosis I (791 and 2211), Barrington
#942, 1OSOX. Figure 4. Camera lucida drawing of sporocyte in figure 3.

1986] Barrington — Polystichum 309

re)
Oo &
a
on ar
o °° tb
a t aS Bad i) a
‘ =
0 9

?
Seg BN %
Soe &, 40

oo. @ ‘ant

Figures 5 and 6: Figure 5. Metaphase of meiosis I (741, 2311, 111] indicated with
arrow), Barrington #942, 800X. Figure 6. Camera lucida drawing of sporocyte in
figure 5.

310 Rhodora [Vol. 88

ticular character are possibly important in considering the evolution
of the genus as a whole.

The irregular pinnae of Barrington 1155 can be interpreted as a
result of mixed developmental programs from the two parents:
broad basal pinnae determined by the Polystichum acrostichoides
program, followed by pinnae of various lengths resulting from com-
promises betwen the two parental programs. In contrast, normal
hybrids evidence an orderly progression of compromises between
the two parental programs, resulting in leaves with regular pinna
development intermediate between that of the two parents. Random
prevalence of one parental developmental program over another has
been invoked to explain a class of irregular features found in
hybrids (Barrington, 1985).

The most intriguing character is the stomate size of the three
entities. The reasonably good inverse correlation of stomate length
with ploidy level known for the species and the triploid hybrid is
counter to that demonstrated in classical studies (e.g., Wagner,
1954). Perhaps the diploid progenitor or progenitors of Polystichum
braunii are both characterized by stomates that are much smaller
than those of the native eastern North American P. acrostichoides.
The large stomate size of the Rugg collections, presumably vouchers
for the tetraploid, suggest that the presence of an extra acrosti-
choides genome has an unusual effect on the determination of sto-
mate size.

The contrast between the present triploid chromosome counts
and the tetraploid count first reported for this hybrid is remarkable.
The hybrid apparently arises most often as a result of fusion of
normal gametes of the two parents. A fertile hexaploid race of
Polystichum X potteri may possibly be found, as was a fertile tetra-
ploid population for Asplenium ebenoides (Wagner, 1954). Such a
plant, backcrossing to P. acrostichoides, could have generated the
tetraploid documented by Morzenti. No fertile hexaploid hybrids
have yet been encountered; all of the large sample of sporophytes
recently collected in Vermont are presumed to be sterile triploids,
since they all show pale tapetal remnants inside the sporangia. Thus,
Morzenti’s original hypothesis for the origin of the tetraploid is the
simplest, given our present understanding of the hybrid.

Many Polystichum hybrids form more bivalents than would be
expected as a result of pairing of homologous chromosomes (by

1986] Barrington — Polystichum 311

definition the result of homoeologous pairing), ranging from an
average of I1 (range 6-24) in European P. braunii X lonchitis
(Lovis, 1977) to 26 (range 19-33) in Californian P. dudleyi X muni-
tum (Wagner, 1973). In Cystopteris, Vida (1974) has reported a
similar high number of bivalents in the apomictically generated
diploid cytological race of C. fragilis. Lovis inferred from these data
that Polystichum and Cystopteris polyploids are segmental allopoly-
ploids, that is they share “valuable complex linkages” common to
both ancestors, but are otherwise chromosomally divergent. The
present hybrid adds to the list of hybrids with unusual levels of
pairing. This high level of pairing is of potential interest, since the
other polystichums hybridizing with P. braunii are much less diver-
gent morphologically. Lovis (1977) reported that Sleep saw similar
high levels (overall range nine to 25) of bivalents in synthesized
hybrids of P. acrostichoides with P. lonchitis, P. munitum, and P.
setiferum.

The origin of Polystichum braunii remains an interesting and
unsolved problem. None of the five hybrids involving P. braunii in
Europe and North America shows a full set of bivalents, so that no
diploid from these regions is implicated as a progenitor. Though no
conclusive information is available from eastern Asia, the close
structural similarity to P. makinoi, a diploid, suggests that the
ancestry of P. braunii is Asian.

ACKNOWLEDGMENTS

I thank Lisa Andrews, David Conant, Philip Cook, Christopher
Haufler, Steven R. Hill, R. James Hickey, Peter Hope, Cathy Paris,
Frank and Libby Thorne, David Wagner, and Peter Zika for their
contributions to the completion of this work.

LITERATURE CITED

ANDERSON, E. 1936. Hybridization in American tradescantias. Ann. Mo. Bot.
Gard. 23: 511-525.

1949. Introgressive Hybridization. John Wiley & Sons, NY.

BARRINGTON, D. S. 1985. The morphology and origin of a new Polystichum
hybrid from Costa Rica. Syst. Bot. 10: 199-204.

HuLTEéN E. 1962. The circumpolar plants I. Vascular cryptogams, conifers,
monocotyledons. Kongl. Svenska Vetenskapsakad. Handl. 5: 1-275.

Live, A., D. Love, AND R. E. G. PICHI-SERMOLLI. 1977. Cytotaxonomical
Atlas of Pteridophyta. J. Cramer, Vaduz, Germany.

312 Rhodora [Vol. 88

Lovis, J. D. 1977. Evolutionary patterns and processes in ferns. Advances Bot.
Res. 4: 229-415.

MANTON, I. AND T. REICHSTEIN. 1961. Zur Cytologie von Polystichum braunii
(Spenner) Fée und seiner Hybriden. Ber. Schweiz. Bot. Ges. 71: 370-383.

Morzenti, V. M. 1962. A first report of pseudomeiotic sporogenesis, a type of
spore reproduction by which “sterile” ferns produce gametophytes. Amer. Fern
J. 52: 69-78.

SLEEP, A. AND T. REICHSTEIN. 1967. Der Farnbastard Polystichum X meyeri
hybr. nov. = Polystichum braunii (Spenner) Fée X P. lonchitis (L.) Roth und
seine Cytologie. Bauhinia 3: 299-374.

Smith, A. R. 1981. Part 2, Pteridophytes. /n: Breedlove, D. E., Ed., Flora of
Chiapas. California Academy of Sciences. San Francisco, California.

THompson, R. H. AND R. L. Corrin. 1940. A natural hybrid between Polysti-
chum braunii (Spenner) Fée and P. acrostichoides (Michx.) Schott. Amer.
Fern J. 30: 81-88.

Vipa, G. 1974. Genome analysis of the European Cystopteris fragilis complex. I.
Tetraploid taxa. Acta Bot. Acad. Sci. Hung. 20: 181-192.

Wacner, W. H., JR. 1954, Reticulate evolution in the Appalachian aspleniums.
Evolution 8: 103-118.

1973. Reticulation of Holly Ferns (Polystichum) in the Western United
States and adjacent Canada. Amer. Fern J. 63: 99-115.

1983. Reticulistics: the recognition of hybrids and their role in cladistics

and classification. Jn: Platnick, N. I., and V. A. Funk, Eds., Advances in Cladis-

tics Vol. 2. Columbia University Press, New York, pp. 63-79.

, AND K.L. CHEN. 1965. Abortion of spores and sporangia as a tool in the

detection of Dryopteris hybrids. Amer. Fern J. 55: 9-29.

PRINGLE HERBARIUM

DEPARTMENT OF BOTANY
UNIVERSITY OF VERMONT
BURLINGTON, VT 05405-0086

APPENDIX
Exsiccatae for Polystichum X potteri Barr.

(Specimens determined by D. H. Wagner, Univ. of Oregon, which I
have not seen are indicated as det. DW. All others—those at GH,
NHN, VT, and YU—are paratypes except for the holotype, Barring-
ton 939.)

CANADA: (all det. DW) Nova Scotia. Inverness Co.: near Harvard Lakes, A.
Prince & C. Atwood 1442 (DAO). Co. indet.: Folleigh, M. Malte s. n. (CAN 216749).
New Brunswick. York Co.: Keswick, J. Brittam 26, (GH). Quebec. Brome Co.: Water-

1986] Barrington — Polystichum 313

loo, Br. Marie-Anselm s. n. (DAO). Montmorency Co.: Montmorency Falls, Macoun
69235 p. p. (CAN).

UNITED STATES: New Hampshire. Coos Co.: White Mountains, quite high, E.
Tuckerman s. n. in 1843 (GH). Vermont. Caledonia Co.: Sutton, W slope of Mt. Hor,
Barrington 777, Zika 3386, 3390 (vt); Wheelock, E face of Mt. Ide, near summit,
Barrington 944, 945, 946, 947, 948, 949, 950, 1084, 1085, 1167 (vr). Chittenden Co.:
Underhill, Nebraska Notch, Barrington 904, 912, 928 (vt); Underhill, base of the chin
of Mansfield, Pursh s. n. in 1838 (GH). Essex Co.: Brighton, W side of Rt. 114 near
Morgan town line, Barrington 834, Zika 4130 (vt). Lamoille Co.: Eden, Access Road
to Belvidere Serpentine Mine, Hickey 906 (personal herbarium R. James Hickey); E
side of Rt. 100 near Lowell town line, Barrington 909, 959 (vt); Stowe, F. Bumstead
s. n. (YU); Cambridge, Smuggler’s Notch, Barrington 918 (vt), garden specimen from
Smuggler’s Notch, R. H. Thompson & R. L. Coffin s. n. in 1940 (Gu), J. Churchill
s. n. (MO 1055822, det. DW). Orange Co.: Braintree, J. Bates 496 (YU); Brookfield, E
side of Brookfield Gulf, 1200 feet, Barrington 1081, 1082, Zika 7196, 7199 (v7);
Strafford, Beacon Hill, Barrington 939 (holotype), 940, 941, 942, 943 (vT). Orleans
Co.: Westmore, E side of Mt. Hor, 1500 feet, Zika 7348 (vT); Town indet., Wil-
loughby Mt., J. Churchill s. n. (Mo 1055821, det. DW). Rutland Co.: Sherburne,
Shaw Hill Brook ravine, 1550 feet, Barrington 750 (vt); W slope of Wolf Hill,
Barrington 749a (vt). Windsor Co.: Barnard, Barnard Gulf, Barrington 1091, 1092,
1093, 1094, 1096, 1097, 1098, 1099, 1100, 1103, 1105, 1106, 1116, 1125, 1131, 1134,
1135, 1140, 1143, 1148, 1151, 1153, 1155, 1157 (vt); same locality, E. Kittredge s. n.
(Mo 993604, det. DW); Cavendish, W side of Proctorsville Gulf, Barrington 898 (VT);
Sharon, Honey Brook Ravine, Barrington 990 (vt); Woodstock, Miss Strong s. n.
(HNH). New York. (all det. DW) Delaware Co.: Arkville, P. Wilson s. n. (NY). Co.
indet.: Ox Clove, Catskill Mts., C. Lown s. n. (NY). Pennsylvania. Sullivan Co.:
Ganoga Glen, H. Pretz 3922, 3923a, (MICH, det. DW). Wayne Co.: Scott Twp., NE
side of Schrawder Mt., | mi N of Island Lake, W. Dix s. n. (us, det. DW); NE slope
of Shrawder Mt., E. Wherry s. n. (GH; Us, det. DW). Starrucca, Lake Shehawken, W.
Dix s. n. (GH); Sterling, R. C. Harlow s. n. in July, 1939 (Gu), in July 1941 (GH), Gas
Hollow, 2 mi. from Jericho (GH).

PLANTAGO MARITIMA AND CAREX MACKENZIEI
NEW FOR SASKATCHEWAN:
ADDITIONAL RARE INLAND STATIONS FOR
TWO SEACOAST SALT MARSH SPECIES

VERNON L. HARMS, DONALD F. Hooper, AND LES BAKER

ABSTRACT

Plantago maritima L. and Carex mackenziei Krecz., two seacoast salt marsh
species rare at inland stations, are reported from the lower Carrot River valley of
east-central Saskatchewan, Canada as new to the flora of the province. Taxonomic
considerations for Plantago maritima and phytogeographical observations and
comments for both species are included.

Key Words: Plantago maritima, Carex mackenziei, inland stations, salt-marshes,
Saskatchewan

During the 1984 and 1985 summer seasons, we found and col-
lected plants of the Seaside Plantain, Plantago maritima L., at two
separate locations in the lower Carrot River valley of east-central
Saskatchewan. This species is reported here as new to the flora of
Saskatchewan. Our first collection was on the broad, saline, marshy
flats about 0.7 km south of the Nitenai River, about 2.5 km south of
the Carrot River, and north of Highway no. 55 (The Kelsey Trail)
about 7 km west of its junction with Highway no. 9, about 85 km
north of Hudson Bay Junction (c Sec. 22, T 53 N, R 02 W 2nd M;
53° 35’ 25”N, 102° 13’W; 17 July 1984, no. 32767). The plants were
locally abundant on the marshy borders of salt-spring pools where
they were.closely associated with Triglochin maritima L. Our
second collection was on the extensive saline, marshy flats just south-
east of Shoal Lake, about 0.5 km west of Pakwaw Lake village, on
the Shoal Lake Indian Reserve, about 100 km east of Nipawin (ne
Sec. 10, T 52 N, R 05 W 2nd M; 53° 28’ 45”N, 102° 39’ W; 18 July
1984, no. 32837). The plants occurred in a similar habitat to those at
the previous location, being limited to the very edges of salt-spring
pools. They were locally frequent with Triglochin maritima, the
only immediate associate, with the exception of some scattered
Scirpus validus plants that were more or less emergent just off the
pond edges.

Plantago maritima sensu lat. (including P. juncoides Lam., P.
decipiens Barneoud, P. borealis Lange, P. oliganthos Roemer &

315

316 Rhodora [Vol. 88

Schultes, and P. glauca Hornem.) is a circumboreal and bipolar,
primarily sea-coast species which in North America reportedly
occurs along the Pacific Coast (Aleutian Islands and southern
Alaska, south to southern California), the Atlantic Coast (Green-
land, southern Baffin Island and Labrador, south to New Jersey),
and along the shores of the Hudson, James, and Ungava Bays (Hul-
tén, 1949, 1968: Fernald, 1950; Gleason, 1952; Bassett, 1973;
Hitchcock et al., 1959; Scoggan, 1979; Porsild & Cody, 1980). It is
apparently absent on the North American Arctic Coast except for a
single reported station on the Mackenzie River delta (Hultén,
1968; Scoggan, 1979) which, however, was omitted by Cody (1979)
and Porsild & Cody (1980). (Figure 1). The characteristic seacoast
habitats for the species are moist, saline sand-beaches, rock-ledges
and tidal marshes.

In addition to its maritime coastal distribution indicated above,
Plantago maritima has also been recorded at the following, quite
widely separated stations in the continental interior: CANADA:
Yukon Territory: Kluane National Park, in “alkaline pools” (Doug-

?

GA ae , yoy & PLANTAGO MARITIMA L.

FIG. 1

Figure |. Recorded distribution of Plantago maritima in North America. Hatch-
ing indicates the maritime coastal distribution; closed dots include the isolated and
continental interior stations; open circle and arrow indicate the newly reported Sas-
katchewan sites.

1986] Harms, Hooper & Baker — Plantago & Carex 317

las, 1974; Douglas et al., 1981). District of MacKenzie, Northwest
Territories: north shore of McTavish Bay of Great Bear Lake, at
“saline springs” (Cody, 1979; Scoggan, 1979; Porsild & Cody, 1980).
Alberta: Wood Buffalo National Park, on “saline soil” (Argus &
White, 1978; Scoggan, 1979) or in “saline marshes” (Packer, 1983;
Packer & Bradley, 1984). Boivin (1972) listed it also from “Heart
Lake,” presumably this being in east-central Alberta, northeast of
Lac la Biche, at 55° 02’ N, 111° 30’ W. Packer & Bradley (1984)
further mapped the species at a lower Clearwater River site (56° 41’
N, 110° 46’ W) in northeastern Alberta, based on a collection by
Peter Lee on 28 June 1983 from “saline marshes” (J. Packer, pers.
corr. 26 Oct. 1984). Scoggan (1979) also lists Plantago maritima for
Alberta from “saline soil along the Red Deer River, Alberta, where
taken by John Macoun in 1881.” But the latter appears erroneously
referred by Scoggan to the Red Deer River in Alberta rather than to
the river of the same name in west-central Manitoba, as seems
clearly indicated by Macoun’s (1884, p. 393) own citation: “in great
profusion in and around salt marshes on Red Deer River about one
mile from Lake Winnipegosis, Lat. 53° (Macoun).” Manitoba:
Dawson Bay of Lake Winnipegosis and Red Deer River above
mouth, at “salt springs,” in “salt marshes,” or on “saline” or “alka-
line soils” (Macoun, 1884; Scoggan, 1957; Boivin, 1972; Scoggan,
1979; White & Johnson, 1980). A report by Lowe (1943) of “P.
decipiens” as rare on “sea beaches and alkaline areas” on the “Old
Souris channel s. of Melita” in Manitoba, was apparently dis-
counted by Scoggan (1953, 1957) as presumably based on misidenti-
fied P. elongata. U.S.A.: Utah: Great Salt Lake (Hitchcock et al.,
1959; Scoggan, 1979).

Our newly reported Carrot River valley populations of Plantago
maritima from along saline spring pool edges in the lower Carrot
River valley add to the foregoing list of inland stations, and are the
first records for Saskatchewan. They are located approximately 110
and 125 km, respectively, northwest of the interior Manitoba station
at Dawson Bay—Lower Red Deer River, where the species was first
collected over a century ago by John Macoun.

In the Canadian Prairie Provinces, the species most similar to
Plantago maritima, at least in general appearance, is probably P.
elongata Pursh (including P. pusilla auct., non Nutt.). But P. elon-
gata consists of smaller (mostly <10 cm high), more delicate,

318 Rhodora [Vol. 88

slender-stemmed, annual plants with slender taproots, in contrast to
our mostly larger, coarser, perennial P. maritima plants with thick-
ish roots and usually branched caudices. Also our P. maritima
plants, in contrast to those of P. elongata, have (1) leaves fleshy
(rather than non-fleshy), (2) corolla tubes short-villous (rather than
glabrous), (3) corolla lobes spreading and over | mm long (rather
than often erect and less than | mm long), (4) anthers 4 (rather than
2), (5) spike axis distinctly short-pubescent (rather than glabrous),
and (6) flowering and fruiting spikes thicker and denser (rather than
slender and loosely flowered).

The New World populations of Plantago maritima appear best
accepted as subsp. juncoides (Lam.) Hultén, separable from the
Eurasian subsp. maritima on the basis of their broader bracts and
more numerous ovules (Hitchcock et al., 1959; Bassett, 1973).
Further varietal segregates have often been distinguished within
North American populations (Fernald, 1925, 1950), but also fre-
quently questioned as being perhaps merely environmental forms
(Rousseau, 1942; Gleason, 1952; Hitchcock, 1959). Our collected
sample of 38 Saskatchewan plant specimens can be characterized as
follows: (1) leaves entire, stiffishly erect, (I-) 1.5-3 (-4) mm wide,
(7-) 10-20 cm long, mostly somewhat shorter than the spike-bearing
scapes; (2) spike-bearing scapes 10-23 cm high, strictly erect or often
somewhat curved above; (3) spikes 1-10 cm long, densely compact
or flowers somewhat separated below; (4) bracts mostly distinctly
keeled, the tips not prolonged, and the margins glabrous to usually
slightly or sometimes distinctly ciliolate; (5) calyx-segment margins
glabrous or often remotely ciliolate; (6) seeds oblongish, 1.3-2.5 mm
long; and (7) anthers I-1.5 mm long, with their “subulate-tips”
about 0.25 mm long. Upon application of the taxonomic criteria
given by Fernald (1925, 1950), most of our material would be identi-
fied as his P. juncoides var. decipiens (Barneoud) Fern., although
some characters of his P. oliganthos R. & S. are frequently present
(e.g., bracts distinctly keeled, they and the calyx-segments often
glabrous or only remotely ciliolate). A few of the smaller plants with
shorter spikes might well be individually identifiable as P. juncoides
var. glauca (Horneman) Fern., although they clearly belong within
the same local populations from which our sample was collected.
While the Saskatchewan collections represent a basis too limited
for taxonomic conclusions, their populational variation at least

1986] Harms, Hooper & Baker — Plantago & Carex 319

suggests the probable presence of only weak distinctions, if any,
between the “forms” represented by the epithets: decipiens, glauca,
and oliganthos. But even if these three “taxa” were to be merged
under a broader P. maritima var. glauca Horneman (the epithet
among these three with priority at the varietal rank), taxonomic
questions would still remain regarding its separation from the more
western P. maritima var. juncoides (Lam.) Gray and var. californica
(Fern.) Piper, and the more eastern taxon that has often been
recognized as P. juncoides var. laurentiana Fern. These taxonomic
questions are beyond the scope of this paper.

During the 1985 summer, within the broad Nitenai River salt
marsh, our first collecting site for Plantago maritima, we also discov-
ered and collected plants of Carex mackenziei Krecz. as new to the
flora of Saskatchewan (se Sec. 22, T 53 N, R 02 W 2nd Wa os" 35’
25” N, 102° 13’ W; 24 June 1985, no. 33,803). The plants occurred
on wet, somewhat raised, hummocky, salt-spring mounds (in con-
trast to salt-spring pool edges where Plantago maritima was found).
Despite a relatively extensive although hardly thorough search of
the broad, over 150-hectare marsh, only a single relatively limited
local population consisting of about 100 plants was found, con-
tained within an area of less than 100 m2. The other plant species
most closely associated with this sedge were Triglochin maritima L.
Scirpus rufus (Huds.) Schrad., and Glaux maritima L.

Carex mackenziei is a circumpolar maritime species, characteris-
tic of tidal and other saline marshes. In North America it reportedly
occurs in Alaska along the Arctic, Bering and Pacific coasts from
Kotzebue Sound south to Sitka (Hultén, 1968), at the Mackenzie
River delta in the Northwest Territories on the Arctic coast
(Hulten, 1968; Porsild & Cody, 1980), along the Hudson Bay and
James Bay coasts from southern Keewatin District to the Great
Whale River in Quebec, and along the Atlantic and Gulf of St.
Lawrence coasts from Labrador, south to Nova Scotia and Maine
(Fernald, 1950; Gleason, 1952; Hultén, 1968; Scoggan, 1978; Por-
sild & Cody, 1980). Our present Saskatchewan report of Carex
mackenziei is particularly significant since it apparently represents
the first known record for the species from the continental interior
of North America. (Figure 2).

With regard to the newly reported Saskatchewan occurrences of
two characteristically maritime coastal salt marsh species, it may be

320 Rhodora [Vol. 88

7 , a
fo, / J ae , ele 3
/ ~ scale os = oe
/ S a ee - to km ~~ va ai
pS size a faa
FIG. 2 CAREX MACKENZIEI

Figure 2. Recorded distribution of Carex mackenziei in Western Canada. Closed

dots indicate the previously reported localities; the open circle and arrow indicate the
newly reported Saskatchewan site.

significant that the salinity of the pertinent Carrot River Valley salt
springs and marshes in which these species were found is mainly due
to their sodium chloride content rather than the sodium sulfate that
is characteristic of most Saskatchewan saline lakes and marshes.
The occurrences of maritime coastal salt marsh species such as
Plantago maritima and Carex mackenziei in the continental interior
of Western Canada as rarities at widely spaced isolated outposts
pose an interesting, albeit difficult, phytogeographical question with
respect to how such distributional patterns were originally attained.
A post-Pleistocene marine incursion extensive enough to account

1986] Harms, Hooper & Baker — Plantago & Carex ot

for such distributions is hardly a viable postulate. We would
hypothesize that such seacoast species likely spread along semi-open
receding shores of the large and perhaps at times estuarian-like
post-glacial Lakes Agassiz and McConnell, especially during their
latter stages (about 5000-4000 years ago) when connections or near-
connections existed to Hudson Bay and the Arctic Ocean, respec-
tively. Subsequent survival would then have occurred only at a few
persistent, favorable interior sites, which in the cases of P. maritima
and C. mackenziei appear to have been mainly sodium chloride
salt-springs within saline marshes.

Plantago maritima was accorded rarity status for the Yukon
(Douglas et al., 1981), the continental Northwest Territories (Cody,
1979), Alberta (Argus & White, 1978; Packer & Bradley, 1984),
Manitoba (White & Johnson, 1980), and Ontario (Argus & White,
1977). Carex mackenziei was accorded rarity status for the continen-
tal Northwest Territories (Cody, 1979) and Manitoba (White &
Johnson, 1980). Both species now deserve a similarly rare status for
Saskatchewan as well.

The collections cited in this paper are those of the authors, with
voucher specimens deposited at SASK and USAS. Duplicates will
also be filed in DAO and CAN.

ACKNOWLEDGMENTS

We gratefully acknowledge the field assistance of John Lathlin at
the Shoal Lake Reserve, as well as the subsequent contribution of
John H. Hudson (of the Saskatchewan Research Council, Saska-
toon) in verifying our Carex identifications. We also thank August
J. Breitung for suggesting our careful 1985 search for sedge species
in these salt marsh species

LITERATURE CITED

Arcus, G. W., AND D. J. Wuite. 1977. The Rare Vascular Plants of Ontario.
Syllogeus no. 14, Botany Div., Nat. Mus. Can., Ottawa.

, AND 1978. The Rare Vascular Plants of Alberta. Syllogeus no.
17, Botany Div., Nat. Mus. Can., Ottawa.

BasseTT, I. J. 1973. The Plantains of Canada. Can. Dept. Agric. Res. Br., Mono-
graph 7, Ottawa, Ontario.

Bolvin, B. 1972. Flora of the Prairie Provinces, Part [1I—Connatae. Provanche-
ria 4, Mem. |’Herbier Louis-Marie, Univ. Laval, Que.

322 Rhodora [Vol. 88

Copy, W. J. 1979. Vascular Plants of Restricted Range in the Continental
Northwest Territories, Canada. Syllogeus no. 23. Nat. Mus. Can., Ottawa.
DouG.ias, G. W. 1974. Montane zone vegetation of the Alsek River region,

southwestern Yukon. Canad. Journ. Bot. 52: 2505-2532.

, G. W. Arcus, H. L. Dickson, AND D. J. BRUNTON. 1981. The
Rare Vascular Plants of the Yukon. Syllogeus no. 28, Nat. Mus. Can., Ottawa.

FERNALD, M. L. 1925. The maritime plantains of North America. Rhodora 27:
93-104.

1950. Gray’s Manual of Botany, 8th ed. American Book Co., N. Y.

GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of the North-
eastern United States and Adjacent Canada, Vol. 1—The Pteridophyta,
Gymnospermae and Monocotyledoneae, and Vol. 3—The Sympetalous Dicot-
yledoneae. The N.Y. Botanical Garden, N.Y.

Hitcucock, C. L., A. CRONQUIST, M. OWNBEY, AND J. W. THOMPSON. 1959.
Vascular Plants of the Pacific Northwest. Part 4: Ericaceae through Campanu-
laceae. Univ. Wash. Press, Seattle.

HuLtén, E. 1949. Flora of Alaska and Yukon, IX. C. W. K. Gleerup, Lund,
Sweden.

1968. Flora of Alaska and Neighboring Territories. Stanford Univ.
Press, Stanford, Calif.

Lowe, C. W. 1943. List of the Flowering Plants, Ferns, Clubmosses, Mosses and
Liverworts of Manitoba. Natur. Hist. Soc. Manitoba.

Macoun, J. 1884. Catalogue of Canadian Plants. Part II—Gamopetalae. Daw-
son Bros., Montreal.

Packer, J.G. 1983. Flora of Alberta, 2nd ed., Univ. of Toronto Press, Toronto.

, AND C. E. BRADLEY. 1984. Checklist of the Rare Vascular Plants
of Alberta with Maps. Prov. Mus. of Alberta, Natural Hist. Occ. Paper, No. 6,
Edmonton.

PorsI_p, A. E. AND W. J. Copy. 1980. Vascular Plants of Continental Northwest
Territories, Canada. Nat. Mus. Can., Ottawa.

Rousseau, J. 1942. La forme naine du Plantago juncoides et d’autres espéces.
Contr. Inst. Bot., Univ. Montréal 44: 59-64.

ScoGGAN, H. J. 1953. Botanical Investigations in the Glacial Lakes Agassiz-
Souris Basins. Nat. Mus. Can. Bull. no. 128. Ottawa: 103-107.

1957. Flora of Manitoba. Nat. Mus. Can., Bull. no. 140, Ottawa.

1978. The Flora of Canada, Part 2—Pteridophyta, Gymnospermae,

Morocotyledonae. Nat. Mus. Can., Ottawa.

1979. The Flora of Canada, Part 4—Dicotyledoneae (Loasaceae to
Compositae). Nat. Mus. Can., Ottawa.

WuiteE, D. J., AND K. L. JOHNSON. 1980. The Rare Vascular Plants of Manitoba.
Syllogeus no. 27. Nat. Mus. Can., Ottawa.

1986] Harms, Hooper & Baker — Plantago & Carex

Me Oa) 3 B

THE W. P. FRASER HERBARIUM
UNIVERSITY OF SASKATCHEWAN
SASKATOON, SASKATCHEWAN
CANADA S7N 0WO

DoF cH

BOX 40

SOMME, SASKATCHEWAN
CANADA SOE INO

LB.

BOX 90

CARRAGANA, SASKATCHEWAN
CANADA SOE 0K0

323

DISPERSION PATTERN OF AERIAL SHOOTS
OF THE COMMON MARSH REED
PHRAGMITES AUSTRALIS (POACEAE)

ANDREW N. DAVIS AND TIMOTHY L. BRIGGS

ABSTRACT

The dispersion pattern of aerial shoots of the common marsh reed, Phragmites
australis (Cav.) Trin. ex Steud., was determined by density counts from a marsh
population in southeastern Massachusetts using quadrat methods. Statistical ana-
lyses of 0.0625 m? quadrat data indicated a random dispersion pattern. An aggre-
gated dispersion pattern is indicated by analyses of 0.25 m? quadrat data. The
aggregated dispersion pattern appears to be a reproductive pattern based on the
mode of population development of P. australis in which extensive vegetative propa-
gation forms aggregations of aerial shoots. In addition, the number of shoots per
aggregate exhibited a logarithmic distribution that conforms to the concept of a
viable unit.

Key Words: Phragmites australis, dispersion pattern, marsh reed, aerial shoots,
ecology, southeastern Massachusetts

INTRODUCTION

The common marsh reed, Phragmites australis (Cav.) Trin. ex
Steud. [= P. communis Trin. (Clayton, 1968)] may be the most
widely distributed angiosperm in the world, growing in fresh or
brackish waters in pure or nearly pure stands. This species has been
found in strata that date back to the Miocene (Tertiary Period) (de
la Cruz, 1978), and is believed to have evolved in subtropical regions
from where it subsequently spread to temperate areas in both hem-
ispheres. Its present distribution ranges from Norway and Canada
in the north to southern temperate areas in Australia, Africa, and
South America (Haslam, 1972; Holm et al., 1977).

Phragmites australis is a perennial grass with annual aerial shoots
that terminate in a distinctive feather-like flower cluster. Although
seed set is annual, most are inviable, making establishment from
seed rare (Harris and Marshall, 1960; Haslam, 1971a, 1971b, 1972).
Meiotic disturbance, flower infertility, pollen lethality, self-incom-
patibility, inflorescence density, and unfavorable weather during
flowering and seed setting, may all lower the percentage of viable
fruits produced (Haslam, 1972), as also may clonal and habitat
differences. Even with viable seeds, field germination is often poor,

325

326 Rhodora [Vol. 88

and seedlings have rarely been reported (Harris and Marshall, 1960;
Haslam, 1972). In addition, young seedlings may be killed by frost,
high salinity, flooding, desiccation, or competition (Haslam, 1972).

Phragmites australis also reproduces asexually via vegetative
propagules and long-lived rhizomes. The possible range of suitable
habitats for seed germination is significantly narrower than that
which allows vegetative propagation; e.g., seeds will not germinate
in, 5 cm of water, yet rhizomes grow and spread in | m of water
(Haslam, 1972; Holm et al., 1977). The reported infrequent occur-
rence of seedlings in nature and the extensive vegetative propaga-
tion exhibited by P. australis led us to investigate the dispersion
pattern of this species in a single marsh population. Biological and
physical factors that may influence the spatial distribution and den-
sity of aerial shoots are considered.

PROCEDURE

Field Methods

Density counts of aerial shoots of Phragmites australis in
Demarest-Lloyd Memorial State Park, South Dartmouth, Massa-
chusetts (approx. 70°59’17”W, 41°31’40”N) were made March 20,
1984, prior to the emergence of new growth. The P. australis stand is
in a brackish marsh adjacent to Georges Pond, approximately 175
m along the south side of the state park access road. In order to
minimize the effects of environmental heterogeneity on our data,
sampling was confined to a 5 m-wide band parallel to and ca. 10 m
from a freshwater runoff culvert. Twenty-five 0.0625 m2 and twenty-
five 0.25 m? quadrat counts were taken randomly. Only shoots at-
tached to rhizomes were counted in order to minimize the possibility
of sampling shoots formed prior to the 1983 growing season. Possi-
ble implications of this are addressed below.

Statistical Analyses

The Index of Dispersion was used to test the null hypothesis that
Phragmites australis shoots are distributed randomly (Greig-Smith,
1964). Data that agreed with this null hypothesis were analyzed by
the Chi-square goodness of fit test to verify the fit to a standard
Poisson (random) distribution (Greig-Smith, 1964).

Data from the quadrat size that deviated significantly from a
Poisson distribution and indicated aggregation were tested for

1986] Davis & Briggs — Phragmites 327

agreement with the negative binomial distribution. Both a Chi-
square goodness of fit test (Bliss and Fisher, 1953) and Anscombe’s
third moments test (Anscombe, 1950) were used.

RESULTS

Field density counts of Phragmites australis shoots are presented
in Table |. The results from the Index of Dispersion test indicate
that the dispersion pattern of 0.0625 m? quadrat data did not deviate
significantly from a Poisson distribution (Table 2). The Chi-square
goodness of fit test confirms a Poisson distribution for the 0.0625 m2
quadrat data (.70 > p > .50). The Index of Dispersion test shows
that the deviation from the random model is highly significant for
the 0.25 m? quadrat.data (p < .001), indicating an aggregated distri-
bution (Table 2). The Chi-square goodness of fit test for 0.25 m2
quadrat data shows no significant difference between observed and
expected values in a negative binomial distribution (.75 > p > .50).
Anscombe’s third moments test also indicates that the 0.25 m2 quad-
rat data fit a negative binomial distribution (T = -262.6 and 2S.E.
=439.2).

Calculations for statistical analyses are available by request.

Table |. Field density counts of Phragmites australis aerial shoots.

0.0625 m? quadrats: 5,5, 4,5,6,5, 1,5, 5, 6,9, 7,4, 9, 13, 6, 10, 8, 13, 6, 8, 6, 4, 7, 11

0.25 m? quadrats: 28, 23, 29, 23, 13, 9, 24, 19, 21, 34, 26, 22, 18, 20, 11, 14, 20, 28,
26, 32, 33, 22, 26, 39, 29

Table 2. Index of Dispersion test for Phragmites australis aerial shoot data.

Index of Dispersion

Quadrat = ~

Size (M?) N X S? S?/X x?(df) P

0.0625 25 6.72 8.21 1.22 29.30(24) .30 > p > .20(R)
0.25 25 23.56 53.59 2.27 $4.60(24) p<.001 (A)**

(R) = random dispersion pattern (.95 > p > .05)
(A) = aggregated dispersion pattern (p < .05)
** = highly significant:

DISCUSSION

An understanding of dispersion patterns of organisms, i.e. the
spacing of individuals with respect to their neighbors, can lead us to

328 Rhodora [Vol. 88

causes of their distributions and abundances. The occurrence of any
non-random dispersion pattern (aggregated or even) requires out-
side force or expenditure of energy by the organism. For the latter
to occur, the organism must gain some advantage from the pattern.
The importance of recognizing the dispersion pattern lies in the
correlation of that pattern with biological and physical factors in
order to understand the ecology of the species. Causes of distribution
patterns within populations may be vectorial, reproductive, social,
coactive, or stochastic (Hutchinson, 1953), but are not necessarily
-mutually exclusive, as several may operate together or succeed one
another in time.

In our study, it was determined that aerial shoots of Phragmites
australis are distributed in an aggregated pattern at the 0.25 m2
quadrat size. Gorham and Pearsall (1956) noted “a good deal of
aggregation in Phragmites,” but fail to document this observation
or suggest possible mechanisms leading to the aggregation.

The mode of vegetative reproduction of Phragmites australis can
be correlated with the aggregated distribution of the species. Popu-
lation development within an established area occurs primarily by
proliferation of underground rhizomes leading to extensive clones,
some of which may persist for hundreds of years (Haslam, 1971a,
1972). Horizontal rhizomes that reach 2-3 m in length (Holm et al.,
1977), branch to form other horizontal rhizomes and initiate vertical
rhizomes (Haslam, 1970, 1972). Aerial shoots are produced from
buds near the growing tips of the vertical rhizomes. The shoots
emerge in a “bunched arrangement” (Haslam, 1970) that can
account for the aggregated dispersion pattern observed. Factors
influencing underground portions of P. australis plants are un-
known, but may also be important in determing the aggregated
dispersion pattern of shoots.

The aggregated dispersion of Phragmites australis shoots fits the
negative binomial distribution; the clumps are randomly spaced but
the number of individual shoots per clump exhibit a logarithmic
distribution. Thus, the presence of one aerial shoot increases the
probability that others will be there. Haslam (1971b) suggested a
viable unit of ca. 10 shoots per seedling for British populations of P.
australis. It is not until this stage that a sedling develops a stress-

1986] Davis & Briggs — Phragmites 329

resistant horizontal rhizome (Haslam, 1971b). When a number of
shoots occur together, there is greater possibility for survival under
stressful conditions by reorientatron of food reserves. The concept
of a viable unit for a seedling can be extrapolated to the degree of
aggregation of aerial shoots produced annually from a vertical rhi-
zome, and may explain the negative binomial distribution of the
aggregated shoots in our study. The mean number of shoots at
aggregation was 23.56, indicating a larger viable unit for our popu-
lation in comparison with British populations as noted above (Has-
lam, 1971b). Physical factors [floods, drought, nutrients, light,
temperature, and salinity (Haslam, 1970, 1971la, 1971b, 1972)] and
biological factors [competition and herbivory (Haslam, 1970,
1971b)] may influence the density of aerial shoots and size of
clumps. The concept of a viable unit warrants further research and,
along with the mode of vegetative reproduction, may explain the
aggregated distribution of aerial shoots and the degree to which it
occurs.

We recognize that the observed aggregated dispersion pattern for
Phragmites australis shoots may be a transient pattern at this stage
in the development of the population. Maturation of the P. australis
stand may result in changes in the dispersion pattern. Also, another
dispersion pattern may have been masked if more than one year
class of shoots were sampled. In dense populations in Britain, some
shoots may remain standing for up to 2 seasons, with older shoots
becoming brittle (Haslam, 1972). However, we counted only shoots
attached to rhizomes, and the winter conditions at our site may
preclude older, brittle shoots staying attached to their rhizomes
through two winters.

In summary, aerial shoots of Phragmites australis are distributed
in an aggregated manner and according to the negative binomial.
This distribution appears to be a reproductive pattern based on the
mode of vegetative propagation. Biological and physical factors
may be involved in determining density of aerial shoots and size of
the clumps. Degree of aggregation may be a response to environ-
mental stresses, and conforms to the concept of a viable unit. Bio-
logical implications of the vegetative mode of development for
stands of this species are evident in the widespread distribution and
success of P. australis.

330 Rhodora [Vol. 88

ACKNOWLEDGMENTS

Our gratitude is expressed to Craig Edwards for his invaluable
help with the statistics and constructive criticisms of this work.
Thanks go to Bob Wilce, Steve Miller, Lars Carlson, Melinda God-
frey, and Paul Godfrey for their comments on the manuscript. We
thank Alan Hankin, Director of the Lloyd Center for Environmen-
tal Studies, for use of the facilities and access to Demarest-Lloyd
Memorial State Park.

LITERATURE CITED

ANSCOMBE, F. J. 1950. Sampling theory of the negative binomial and logarithmre
series distributions. Biometrika 37: 358-382.

Buiss, C. 1. AND R. A. FISHER. 1953. Fitting the negative binomial distribution to
biological data. Biometrics 9: 176-200.

CLAYTON, W. D. 1968. The correct name of the common reed. Taxon 17:
168-169.

De La Cruz, A. A. 1978. The production of pulp from marsh grass. Econ. Bot.
32: 46-S0.

GorHAM, E. AND W.H. PEARSALL. 1956. Production ecology. III. Shoot produc-
tion in Phragmites in relation to habitat. Oikos 7: 206-214.

GREIG-SMITH, P. 1964. Quantitative Plant Ecology, 2nd ed. Butterworths,
London.

Harris, S. W. AND W. H. MARSHALL. 1960. Experimental germination of seed
and establishment of seedlings of Phragmites communis. Ecology 41: 395.
HasLaM, S.M. 1969. The development of shoots in Phragmites communis Trin.

Ann. Bot. 33: 695-709.
1970. The development of the annual population in Phragmites commu-
nis Trin. Ann. Bot. 34: 571-591.
1971la. Community regulation in Phragmites communis Trin. I. Mono-

dominant stands. J. Ecol. 59: 65-73.

1971b. The development and establishment of young plants of Phrag-

mites communis Trin. Ann. Bot. 35: 1059-1072.

1972. Biological flora of the British Isles. Phragmites communis Trin. J.
Ecol. 60: 585-610.

Hout, L. C., D. C. PLUCKNETT, J. V. PANCHO AND J. HERBERGER. 1977. The
World’s Worst Weeds— Distribution and Biology. Univ. Press of Hawaii, Hono-
lulu. Ch. 59, pp. 373-378.

HUTCHINSON, C. E. 1953. The concept of pattern in ecology. Proc. Acad. Nat.
Sci., Phila. 105: 1-12.

DEPARTMENT OF BOTANY
UNIVERSITY OF MASSACHUSETTS
AMHERST, MASSACHUSETTS 01003

SEASONAL SUCCESSION AND VERTICAL DISTRIBUTION
OF PHYTOPLANKTON IN CANDLEWOOD LAKE, CT

STANLEY J. FREEDA! AND PETER A. SIVER

ABSTRACT

Preliminary to development of a management plan for Candlewood Lake (Fair-
field and Litchfield Counties, CT), phytoplankton populations were recorded at
several sites from April, 1983 through January, 1984. Vertical phytoplankton profiles
were identified and counted to derive seasonal succession patterns and population
concentrations. Vertical profiles of nitrate, phosphorus, chlorophyll a, phaeophytin
a, temperature, conductivity, dissolved oxygen, and light were also recorded at the
sampling sites. Distributions of phytoplankton were seasonal; approximately 104
cells/ml occurred in the epilimnion during summer stratification. The lake was dom-
inated (89%) by blue-green algae (Oscillatoriaceae and Nostocaceae). Diatoms and
green algae were important during spring and winter, respectively. Horizontal phy-
toplankton distributions throughout the lake were similar and correlated well with
chlorophyll a concentrations. Nitrate concentrations were generally low (< 0.3 mg
NO3/1) to non-detectable during the summer. Total phosphorus was found to be
high, averaging 35 ywg/| on the surface. Candlewood Lake is thus in an early eutro-
phic state.

Key Words: phytoplankton, eutrophication, seasonal succession, nitrate, phos-
phate, western Connecticut

INTRODUCTION

Candlewood Lake (Fairfield and Litchfield Counties), the largest
lake in Connecticut, covers 2194 hectares and has 105 km of shore-
line. Over the past decade, this lake experienced accelerated eutro-
phication. In the early 1970’s, it was classified as early mesotrophic
(Norvell and Frink, 1975). By the late 1970’s, phosphorus levels had
doubled and the lake was classified as mid to late mesotrophic
(DEP, 1983). In 1983, phosphorus concentrations were sufficient to
classify sections of the lake as eutrophic (Siver, 1984).

In addition to increased phosphorus levels, major changes in
aquatic vascular plant populations have occurred since 1979. Myri-
ophyllum spicatum L. has become the major dominant aquatic vas-
cular plant (Siver, 1984), and its heavy growth has greatly impeded

‘Present Address: Department of Botany and Plant Pathology, University of New
Hampshire, Durham, NH 03824.

a3!

332 Rhodora [Vol. 88

recreational activities on Candlewood Lake, a major economic and
recreational resource in western Connecticut.

In general, algal populations are not a major problem, although
dense blooms have been reported in isolated bays. Lake manage-
ment programs aimed ai controlling weed growth such as aquatic
plant population controls by herbicides and lake drawdowns are
under consideration. Baseline information is relevant prior to the
proposed management practices, to answer the question to what
extent such management practices would affect lake phytoplankton.
The present study was conducted in order to determine distribution
of phytoplankton in Candlewood Lake in relation to its present
physical and chemical parameters.

MATERIALS AND METHODS

Phytoplankton Enumerations

Vertical quantifications of phytoplankton populations were made
in Danbury Bay (Site 1) at 1.0 m intervals from April through
November, 1983, and in January, 1984. During April, May and
August, similar vertical profiles were taken from New Fairfield Bay
(Site 2) and Lattin’s Cove (Site 3) (Figure 1). In May and August,
epilimnetic samples were taken from three additional sites for com-
parative purposes: Pocono Point (Site 4), Orchard Point (Site 5)
and Chatterton Point (Site 6).

All samples were collected using a horizontally positioned van
Dorn bottle and were fixed with Lugol’s Solution (Franson, 1980);
phytoplankton were identified and counted at 600 with an
inverted microscope using settling chambers constructed with the
slight modifications of Coulon and Alexander (1972). Time periods
required to settle all cells completely in the sample volume ranged
from 24 to 48 hours (Furet and Benson-Evans, 1982). Counts were
made along horizontal transects across the diameter of the settling
chamber. Because the top ten taxa accounted for 95% to 99% of the
total cell concentration in each sample, they were used to calculate
the composition of phytoplankton in each algal class.

Chemical and Physical Parameter Enumerations

Water samples for chlorophyll analysis were taken bimonthly
during the summer and monthly during autumn and winter at verti-
cal meter intervals from sites | to 3. Samples at sites 4 to 6 were
taken at | m and from an equal mix of the top seven meters of

1986] Freeda & Siver — Phytoplankton 333

Town of
NEW MILFORD

z
Town of a
SHERMAN =

=

°

a Sf” ie

ag se

‘< 4

¢ Town of

Squantz Pend

Chatterton
Point

BROOKFIELD

Orchard Point

Turtle Bay

Town of

NEW FAIRFIELD NORTH

’
Byars Pocono Point
New Fairfield Bay :

\
Lattin's Cove
,

City of
DANBURY

CANDLEWOOD LAKE
FAIRFIELD & LITCHFIELD COUNTIES
CONNECTICUT NOT TO SCALE

Figure |. Map of Candlewood Lake (Fairfield and Litchfield Counties, CT)
showing locations of the towns, major bays, and the six study sites. Drawing not to
scale.

epilimnetic water. From 200 to 1000 ml of sample were passed
through glass fibre filters buffered with 0.2 ml of saturated mag-
nesium carbonate (MgCO;) suspension. Chlorophylls were extracted
with 90% acetone using a power tissue grinder, the suspension was

334 Rhodora [Vol. 88

centrifuged at 2000 rpm for 10 min, and concentrations of chloro-
phylls a, b, and c were determined using the trichromatic method
(Franson, 1980). Because chlorophyll a may be overestimated by
breakdown products such as phaeophytin a, another method was
used to estimate the chlorophyll a and phaeophytin a levels
(Lorenzen, 1967).

Nitrate concentrations were determined by reducing the nitrate to
nitrite using cadmium reduction. The nitrite level was measured
spectrophotometrically by diazotizing with sulfanilamide and cou-
pling with N-(1-napthyl)-ethylenediamine (NED) to form a violet
azo dye (Franson, 1980). Each 75 ml sample was first passed
through a glass fibre filter, mixed with 25 ml of NHsCl-EDTA, and
allowed to flow through the cadmium reduction column (modified
from Eppley, 1978). The first 50 ml were discarded and the re-
mainder used for nitrate determinations; columns were rinsed
between samples with 40 ml of dilute NHsCI-EDTA.

Total phosphorus was determined after acid persulfate digestion
of non-filtered water samples using the stannous chloride method
(Franson, 1980); pH was determined using a Fisher model 750
pH/ion meter.

At each site, vertical profiles of conductivity, dissolved oxygen,
and light levels were determined; the Secchi disk depth was also
recorded. Temperature and conductivity were measured using a YSI
model 33 temperature/conductivity meter; light was measured with
a Licor Li-185 B Quantum Sensor Photometer, and dissolved oxy-
gen was measured using a YSI model 58 Dissolved Oxygen meter
that was standardized by the Winkler titration method (Franson,
1980).

RESULTS

Phytoplankton

During the study, phytoplankton concentrations ranged from a
low of 2.9 X 103 cells/ml in January to a high of 1.3 X 105 cells/ml in
August. In Danbury Bay, mean concentrations increased from 1.5 X
104 cells/ml in April to the seasonal maximum of 5.43 X 104 cells/ml
in August, and then gradually decreased to a seasonal minimum of
8.27 X 103 cells/ml during January (Figure 6).

Vertical distributions of phytoplankton numbers in Danbury Bay
were uniform from April through June, ranging from 0.5 to 3.0 X

TOTAL CONCENTRATION

wn
w

Ht

1934

JAN 28

so PHYTOPLANKTON (x 10! cetis/m.)

a

NOV 20

PHYTOPLANKTON
(x 10° ceus/m)

OCT 23

1983 SEP 10

») JUN 10
APR 23 bee

a
10.0

DEPTH (meTERS)

a Bie PHYTOPLANKTON VS DEPTH

Figure 6. Phytoplankton concentration versus depth and time in Danbury Bay (Site 1), Candlewood Lake, CT. The sum of
the concentrations at each depth is represented by the bars above each profile.

[9861

uo}yueldoyyd — IdAlS ® epsal4

Ste

336 Rhodora [Vol. 88

104 cells/ml (Figure 6). As the summer progressed, large epilimnetic
populations developed, with a maximum of 8.5 X 104 cells/ml at 6
m on August 19. Vertical distributions were again uniform during
October, November and Janaury.

The phytoplankton taxa in Danbury Bay were similar throughout
the study, but varied in concentration and, therefore, dominance.
Overall, Cyanophyceae represented 89% of the total cells in the bay
during the study. On a monthly basis, Cyanophyceae accounted for
over 68% of the total flora each month, except during January, at
which time they comprised 49% of the phytoplankton community
(Figure 7). The Bacillariophyceae were most abundant in the spring
(31% in April), and decreased to insignificant levels by June. The
Chlorophyceae and Cryptophyceae were most important during
autumn and winter (Figure 7).

Phytoplankton concentrations were similar at all sites (Figure 8).
For example, in May the mean phytoplankton numbers were low at
all sites, ranging from 5.28 X 103 to 1.43 X 104 cells/ml. During
August, all sites had mean epilimnetic phytoplankton levels between
3.86 X 104 to 1.18 X 105 cells/ml.

Phytoplankton community structure was similar at all six sites
during May and August (Table 1). On May 21, at all but the Chat-
terton Point site, the Cyanophyceae accounted for greater than 88%
of the phytoplankton. The dominant taxa were Oscillatoria Agard-
hii Gomont, O. amphigranulata van Goor, and Anabaena laxa
Rabenhorst. During August, the blue-green algae accounted for
over 80% of the flora at all six sites, with O. amphigranulata and A.
laxa being the most abundant taxa.

Chemical and Physical Parameters

Temperature profiles at the Candlewood Lake sites denoted a
typical, dimictic, temperate lake with a temperature range of
0.9-25.6°C. Conductivity ranged from a winter minimum of 60
umhos/cm to a summer maximum of 133 wzmhos/cm. During most
of the study period, pH values were between 7 and 8, except in the
hypolimnion during summer anoxic conditions. Saturated levels of
dissolved oxygen occurred throughout the lake, except in the hypo-
limnion and portions of the metalimnion during summer stratifica-
tion. The light extinction coefficient, n, ranged from 0.67-1.33,
increasing as the summer progressed. Light did not penetrate below
7 m on sunny days (when incident light levels were greater than 1500

1986] Freeda & Siver — Phytoplankton a7

60 J

PERCENT CELLS (2)

CO cvanopayceae Wi
BACILLARIOPHYCEAE f
50} (0) CRYPTOPHYCEAE
CHRYSOPHYCEAE
} cHLOROPHYCEAE

40 T T T T T ay y '
APR 23. MAY 21 JUN 10 JUL 23 AUG S AUG 19 SEP 10 OCT 23 NOV 20 JAN 28
1983 DATE 1984

Figure 7, Percentage of phytoplankton per algal class in Danbury Bay (Site 1),
Candlewood Lake, CT, April, 1983-January, 1984.

120

110 ~
100 -
90 =
BO +
7o -
60 +
50 +
40 +
30 =

20 +
10 4
_ IY

SITE
72 MAY 21 SS AG 19
Figure 8. Phytoplankton concentrations at | m in Danbury Bay (Site 1); New
Fairfield Bay (Site 2); Lattin’s Cove (Site 3); Pocono Point (Site 4); Orchard Point
(Site 5); and Chatterton Point (Site 6) in Candlewood Lake, CT on May 21 and
August 19, 1983.

PHYTOPLANKTON (x 10° ceuis/M)

WLLLLLLLLLLLL

" LLL

338 Rhodora [Vol. 88

Table 1. Percentage of total cells belonging to each class found on the sampling
dates May 21 and August 19, 1983 at | m for the six sampling sites, Candlewood
Lake, CT.

Percentage of Total Cells (%)

Site Class 21 May 1983 19 August 1983

l Cyanophyceae 94 100

2 Cyanophyceae 90.3 92.4
Bacillariophyceae 5.9
Chrysophyceae 4.3

3 Cyanophyceae 97 79.7
Chlorophyceae 18.7

4 Cyanophyceae 88 97
Bacillariophyceae 6.9

5 Cyanophyceae 91.9 99.3
Bacillariophyceae 3.9

6 Cyanophyceae 65 89
Bacillarigphyceae 21 4.4
Chrysophyceae 4.4
Chlorophyceae 6.2

wE/cm2), or below 4-5 m on cloudy days (with incident light levels
of 40-400 »E/cm?). The Secchi depth rarely extended below 3 m.

Nitrate concentrations were low at all sites during the study
period, ranging from non-detectable levels to 0.262 mg NO3;/1. In
Danbury Bay, nitrate concentrations of 0.086 mg NO;/1 were
detected during spring overturn (April); however, levels decreased
to non-detectable by mid-summer (Figure 2). Nitrate concentrations
increased during autumn overturn and reached a seasonal maxi-
mum in January.

Total phosphorus in Danbury Bay ranged from 10 to 125 ywg/1
during the study period (Figure 3). Phosphorus concentrations were
often higher in surface waters (0.5 m), decreased slightly with depth
to 8 m, and then increased substantially below 8 m. Mean phospho-
rus levels of surface water and water between | and 8 m were 35 and
25 ug/l, respectively. Below 8 m, the mean phosphorus concentra-
tion was 55 ug/l.

Chlorophyll a concentrations throughout Candlewood Lake
ranged from 0 to 44 mg/m, with the greatest concentration of
chlorophyll a in the epilimnion during the summer. Lower, more

1986] Freeda & Siver — Phytoplankton 339

uniform levels were recorded during spring and autumn overturns
(e.g., Danbury Bay, Figure 4). The chlorophyll levels corresponded
well with phytoplankton concentrations throughout the year (Fig-
ure 5). Phaeophytin a levels were generally greatest on the lake
bottom and were rarely detected in the epilimnion (Figure 4). Chlo-
rophyll b and c concentrations were non-detectable throughout
most of the year, especially in the summer. Peaks of 2.74 and 0.54
mg/m} for chlorophyll c and b, respectively, occurred during Jan-
uary.

DISCUSSION

Over the past five years, Candlewood Lake has experienced
change, probably the result of cultural eutrophication. Phosphorus
levels in Candlewood Lake during the early 1970’s were indicative of
an early mesotrophic condition (Frink, 1971; Norvell and Frink,
1975). By 1980, average phosphorus concentrations had increased to
the point where the lake was considered mid-mesotrophic (DEP,
1983). During 1983, mean phosphorus levels in Danbury and New
Fairfield Bays were 75% and 35% greater, respectively, than in
1980-81 (Siver, 1984). The rapid rise in the lake’s phosphorus con-
centration was probably due to a combination of factors associated
with cultural eutrophication (e.g., failing septic systems, increased
runoff, fertilizers) and with loading from the sediments. Further
research will be necessary to determine the relative importance of
each contributing factor.

Major changes in both species and densities of aquatic vascular
plants have occurred since 1979 within the lake (DEP, 1983; Siver,
1984). Ceratophyllum sp., Vallisneria americana Michx., and
Potamogeton spp. were dominant forms in many areas of the lake
during 1979 (DEP, 1983); however, by 1983 Myriophyllum spica-
tum L. was by far the most abundant aquatic macrophyte through-
out the lake.

Concurrent with enhanced nutrients and aquatic weeds has been a
further shift toward domination of the phytoplankton community
by the Cyanophyceae. Although blue-green algae dominated the
lake’s phytoplankton community during the late 1970’s and early
1980’s, Chlorophyceae and Bacillariophyceae were also common
(DEP, 1983). Phytoplankton analysis prior to the present study
included only surface samples. In the present study, including verti-
cal analyses of phytoplankton concentrations throughout the year,

NITRATE (me/L)

Nitrate concentrations versus depth and time in Danbury Bay (Site |), Candlewoed Lake, CT.

Ore

vlopoyYy

88 10A]

DEPTH (meters)

Figure 3. Total phosphorus versus depth and time in Danbury Bay (Site 1), Candlewood Lake, CT.

[9861

uo}yueldoiyg — JdaAlg 2 epra14

Ipe

---- PREOPHYTIN A (m/w?)
CHLOROPHYLL A (me/?)

DEPTH = (meters)

Figure 4. Chlorophyll a and phaeophytin a concentrations versus depth and time in Danbury Bay (Site 1), Candlewood
Lake, CT. Concentrations estimated by the Lorenzen Method. chlorophyll a — WW phaeophytin a

cht

vIOpouYy

88 1A]

\o
oO
n
=

Freeda & Siver — Phytoplankton 343

8,05 ky METER ria m= 20

PHYTOPLANKTON (x 104 cents /mL)
~-- = CH/9M) WY TIAHAOYOTD

tT T t tT T T
APR 23 JUN 10 JUL 23 AUG 5S SEP 10 OCT 23 NOV 20 JAN 28
1983 DATE 1984

Figure 5. Seasonal levels of phytoplankton and chlorophyll a at 0.5 in Danbury
Bay (Site 1), Candlewood Lake, CT. ___ phytoplankton a _ — _ chlorophyll a

the Cyanophyceae dominated the lake’s phytoplankton community,
accounting for 89% of all cells. Concentrations of phytoplankton in
surface waters ranged between 104 and 105 cells/ml, which was
comparable to previous studies. However, we found that moderately
high surface phytoplankton concentrations were present throughout
the epilimnion (6-8 m) from June to September. Thus, following a
calm period, the epilimnetic blue-green population could migrate to
the surface after the formation of pseudovacuoles, to form an ex-
tensive bloom. Such bloom formation was observed during the
study in several bays following calm periods. However, because of
its long fetch, surface waters in the lake are generally well mixed,
preventing surface migrations of the blue-greens.

In 1980 and 1981, the Connecticut Department of Environmental
Protection reported mean ratios of total nitrogen to phosphorus
greater than 15:1 and concluded that phosphorus was the limiting
nutrient (DEP, 1983). The higher phosphorus concentrations
recorded in 1983 could have led to periods of nitrogen deficiency. If

344 Rhodora [Vol. 88

so, a shift toward nitrogen-fixing taxa might be expected. In
summer 1981, the phytoplankton were at least 61% Oscillatoriaceae,
non-nitrogen-fixing blue-green algae (DEP, 1983), but in 1983, 67%
of the algae were Nostocaceae, a blue-green family characterized by
heterocystic taxa and presumably nitrogen-fixing. Thus, while the
lake remained primarily a blue-green-dominated lake, a shift toward
nitrogen-fixing taxa was evident, which enabled the blue-green
algae present in 1983 to compensate for nitrogen-poor conditions.
This finding could be an indication that nitrogen rather than phos-
phorus was, at times, the limiting nutrient in the lake.

Death and decomposition of macrophytes result in the release of
nutrients, especially phosphorus, into the water column (Nichols
and Keeney, 1973; Strange, 1976). Based on tissue phosphorus levels
and the determination of macrophyte population numbers, it was
estimated that the phosphorus concentrations in Danbury Bay
could double if all tissue phosphorus were released (Simpkins, Cas-
sia, and Siver, 1984). If the macrophytes were killed with the chemi-
cal herbicide 2,4-D as has been proposed, phosphorus could be
released in large amounts. This death, in turn, could lead to an
immediate increase in Nostocacean algae, especially Anabaena flos-
aquae and Aphanizomenon flos-aquae.

Aquatic vascular plants may obtain their phosphorus in two
ways, either by removing it from the sediments via root systems, or
by absorbing it from the water column through the leaves (Wetzel,
1983). The effect that control of these macrophytes would have on
the phytoplankton would depend on which system the macrophytes
are using to obtain phosphorus. If they are acting as pumps, remov-
ing phosphorus from the sediments, their removal would negate a
source of phosphorus to the phytoplankton and phytoplankton
concentrations might decrease. Conversely, if the weeds are absorb-
ing phosphorus from the water column, they represent competition
for phosphorus to the algae. Thus, their control would remove
competitive pressure from the algae and phytoplankton concentra-
tions might increase.

Chlorophyll a concentrations have often been used as estimates of
productivity and biomass (Kalff and Knoechel, 1978; Wetzel, 1983)
as well as a parameter to indicate lake trophic status (Vollenweider,
1979; Wetzel, 1983). The range of chlorophyll a found in the present
study (0-44 mg/ m3) is most closely associated with a eutrophic lake
(Vollenweider, 1979). The correlation between phytoplankton and

1986] Freeda & Siver — Phytoplankton 345

chlorophyll a concentrations indicated a seasonal succession pattern
with a major summer peak (Figure 5). Thus, chlorophyll data could
be used to estimate the phytoplankton concentrations in Candle-
wood Lake which were not enumerated in the present study. Based
on these data, large seasonal algal maxima were observed at all six
sites in the lake during the summer, with concentrations between 104
and 105 cells/ml. These concentrations were similar to those found
by the Connecticut DEP in 1979-1981 (DEP, 1983). Since winter
concentrations were also similar, it appears that the seasonality of
phytoplankton cell concentrations has remained the same over the
last 5 years, suggesting eutrophication of the lake (Wetzel, 1983).

The similar phytoplankton concentrations found at all six sites
(Table 1) was unexpected since the sites were spread over 6 miles,
and because of the highly dissected morphological nature of the
lake. The horizontal distributions of phytoplankton may, in fact, be
similar throughout the lake.

Based on data from this study, Candlewood Lake is a eutrophic
lake. Because the change in trophic status of the lake occurred over
a relatively short period of time (10 years), it is probable that rever-
sion to an early mesotrophic condition could also occur rapidly if
the nutrient load into the lake were reduced, much as it was in Lake
Washington (Edmundson, 1970).

LITERATURE CITED

ANON. 1983. Phase I Diagnostic/ Feasibility Study, Candlewood Lake, Brook-
field, Danbury, New Fairfield, New Milford, Sherman, CT. Water Compliance
Unit, Department of Environmental Protection, State of Connecticut.

CouLon, C. AND V. ALEXANDER. 1972. A sliding chambered phytoplankton set-
tling technique for making permanent quantitative slides with applications in
flourescent microscopy and audioradiography. Limnol. Oceanogr. 17: 149-152.

EDMUNDSON, T. V. 1970. Phosphorus, nitrogen, and the algae in Lake Washing-
ton after diversion of sewage. Science 169: 690-691.

Epptey, R. W. 1978. Nitrogen uptake. /m: Helleburst and Craigie, Eds., Phyco-
logical Methods Vol. 2, Cambridge University Press, New York. p. 401-409.
FRANSON, M.A. H., Ed. 1980. Standard Methods for the Examination of Water
and Wastewater. I5th ed. American Public Health Association, American
Waterworks Association, and Water Pollution Control Federation, New York.

Frink,C.R. 1971. Candlewood Lake: A tentative plant nutrient budget. CT Agr.
Exp. Sta. Circular 238.

FuretT, J. E. AND K. BENSON-EVANS. 1982. An evaluation of the time required to
obtain complete sedimentation of fixed algal particles prior to enumeration. Br.
Phycol. J. 17: 253-258.

346 Rhodora [Vol. 88

KALFF, J. AND R. KNOECHEL. 1978. Phytoplankton and their dynamics in oligo-
trophic and eutrophic lakes. Ann. Rev. Ecol. Syst. 9: 475-495.

LoRENZEN, C. J. 1967. Determination of chlorophyll and pheo-pigments: spec-
trophotometric equations. Limnol. Oceanogr. 12: 343-346.

NICHOLS, D. AND D. KEENEY. 1973. Nitrogen and phosphorus release from decay-
ing water milfoil. Hydrobiologia 42: 509-525.

NorveLL, W. A. ANDC.R. FRINK. 1975. Water chemistry and fertility of twenty-
three Connecticut lakes. CT Agr. Exp. Sta. Bull. 759.

SIMPKINS, T., D. Cassia, AND P. Siver. 1986. The potential effect of herbicidal
control of macrophytes on phosphorus levels in Candlewood Lake, CT. Pro-
ceedings from the Aquatic Environment: Problems and Perspectives. Fairfield
University Press, Fairfield, CT. p. 117-125.

Siver, P. A. 1984. A preliminary report on the phosphorus dynamics, macro-
phyte growth, and algal flora of Candlewood Lake, 1983. Completion report to
the City of Danbury, Danbury, CT.

STRANGE, R. 1976. Nutrient release and community metabolism following appli-
cation of herbicide to macrophytes in microcosms. J. Appl. Ecol. 13: 889-897.

VOLLENWEIDER, R. A. 1979. Das Naehrstoffbelastungskonzept als Grundlage
fuer den externen Eingriff in den Eutrophierungsprozess stehender Gewaesser
und Talsperren. Z. Wasser- u. Abwasser-Forschung 12: 46-56.

WeTzeL, R.G. 1983. Limnology, 2nd ed. Saunders College Publishing Co., New
York, NY.

DEPARTMENT OF BIOLOGICAL

AND ENVIRONMENTAL SCIENCES
WESTERN CONNECTICUT STATE UNIVERSITY
DANBURY, CONNECTICUT 06810

NEW WOOL-ALIEN CRUCIFERAE (BRASSICACEAE)
IN EASTERN NORTH AMERICA:
LEPIDIUM AND SIS YMBRIUM

IHSAN A. AL-SHEHBAZ

ABSTRACT

The African Sisymbrium turczaninowii, Lepidium africanum, and L. schinzii and
the South American L. bonariense are reported for the first time for North America.
The Eurasian S. /oeselii and the North American L. austrinum, L. lasiocarpum, and
L. oblongum are new to the southeastern United States. All eight species were
collected from waste grounds around wool-combing mills in South Carolina. A key
to the eastern North American species of Lepidium is given.

Key Words: Cruciferae, Brassicaceae, Lepidium, Sisymbrium, wool-alien weeds,
North America, South Carolina

INTRODUCTION

Many noxious weeds became naturalized in eastern North Amer-
ica shortly after their initial introductions in the seventeenth century
by early European settlers. Ballast, wool, imported seed stocks, and
agricultural products were important carriers of small fruits and
seeds of numerous exotic plants. Fruits with hooks or spines are
easily dispersed by clinging to animal wool or fur. Although a very
few genera of the Cruciferae possess such fruits, the majority have
relatively small seeds often capable of exuding mucilage when mois-
tened. Such seeds can effectively be transported by adhering to
uncleaned fleeces. It is not surprising, therefore, that many alien
crucifers are common inhabitants of waste grounds around wool-
combing mills.

This study is based on specimens collected in South Carolina by
Harry E. Ahles and John G. Haesloop from waste grounds around
the Santee Wool-Combing Mill (near Jamestown on route 45, Ber-
keley County) and around the Wellman Wool-Combing Mill (north
of Johnsonville on route 41, Florence County). All specimens cited
are deposited in NCU. This paper deals with eight species, of which
three of Lepidium and one of Sisymbrium have not been previously
recorded for North America.

347

348 Rhodora [Vol. 88

LEPIDIUM

Species of Lepidium are probably the most common and success-
ful crucifers of wool-alien flora. Their success may be attributed to
their small mucilaginous seeds, their abundance on all continents
but Antarctica, their weedy tendencies, and most importantly their
predominantly autogamous breeding system. Seed characters and
abundance of the species are important for adherence of seeds to the
wool, while the breeding system and weediness are essential for the
establishment of plants in their new habitats. As described below,
some of the species have been collected from the same locality for
three or four consecutive years, but the present fate of most of them is
unknown.

Despite significant morphological differences between Lepidium
virginicum L. and L. densiflorum Schrader, a few authors (e.g.
Radford et al., 1968; Smith, 1978) have not recognized the latter or
have reduced it to a synonym of the former. Lepidium virginicum
usually has conspicuous petals and accumbent cotyledons; the
petals, however, may abort in flowers which develop toward the end
of the season, and the cotyledons are incumbent in some of its
western North American varieties. In L. densiflorum the petals are
either absent or aborted and the cotyledons are always incumbent.
It is possible, therefore, to confuse the two species. Lepidium densi-
florum tends to have fruits smaller than those of L. virginicum, but
fruit size is unreliable for distinguishing the western North Ameri-
can plants of both species. The type of pubescence on the infructes-
cence axis and, to a lesser degree, the shape of the fruit (see couplet
N of the key below) are very useful in separating them.

The following key is provided to aid the identification of 14 native
and exotic eastern North American lepidiums, including three
recorded here as new for the continent. The European L. /atifolium
L. and L. heterophyllum (DC.) Bentham are sporadically distrib-
uted in New England, New York, and New Jersey, but neither has
been reported from the Southeast. The native L. austrinum Small,
L. lasiocarpum Nutt., and L. oblongum Small are very rare in the
eastern limits of their natural ranges in Arkansas and Louisiana, but
are widespread in California, the southwestern states, and northern
and central Mexico. Undoubtedly, these last three have been intro-
duced into South Carolina on imported wool, and are recorded here

1986] Al-Shehbaz — Cruciferae 349

as well. Each of the six numbered species is further described follow-
ing the key.

Key to the eastern North American species of Lepidium

A. Upper cauline leaves perfoliate or sagittate.
B. Upper leaves perfoliate, ovate to orbicular; basal leaves bi-
pinnatisect; fruiting pedicels glabrous; flowers yellow;
SOCUS WINKED 5 cecilia en esuee sees anes L. perfoliatum L.
B. Upper leaves sagittate, oblong to linear or lanceolate; basal
leaves undivided to rarely lobed; fruiting pedicels densely
pubescent; flowers white; seeds wingless.

C. Annuals or biennials; stems usually solitary at the base;
fruits densely covered with vesicles; style included or
slightly exserted from the apical notch of fruit; anthers
VOUOW. 2 45cekoes ie ries see es L. campestre (L.) R. Br.

C. Perennials; stems several from the base; fruits without or
with a few vesicles; style usually exserted | mm or more
beyond the apical notch of fruit; anthers purple ........

eee ee err et L. heterophyllum (DC.) Bentham
A. Upper cauline leaves tapering to the petiole or the base.
D. Fruits 5-6 mm long; stamens 6; cotyledons 3-lobed ........
SPAS SCRE RONSS EWA TY Eek eee aes L. sativum L.
D. Fruits 1.5-3.5 (-4) mm long; stamens 2 (6 in L. latifolium);
cotyledons entire.

E. Fruiting pedicels erect-appressed; nectar glands filiform,
0.3-0.6 mm long; stems with retrorse, flattened, scalelike
(IGROMES 6 dia ceae Sees cause 6. L. schinzii Thell.

E. Fruiting pedicels ascending to spreading; nectar glands tri-
angular or round, less than 0.2 mm long; stems glabrous or
pubescent, without scalelike trichomes.

F. Stoloniferous perennials, (SO-) 70-130 cm high; fruits
wingless and not notched at apex; stamens 6; stigma 2-3
times broader than the style; petals longer than the sepals

[2 cssr whee aan ee aces eyes wees L. latifolium L.

F. Annuals or biennials, 10-45 cm high, rarely perennials to
75 cm high (L. africanum); fruits winged and notched at
apex; stamens 2; stigma about as wide as the style; petals
much shorter than the sepals (longer in L. virginicum).

350 Rhodora [Vol. 88

G. Fruits pubescent or only ciliate at margin.
H. Fruits with subappressed trichomes, lower surfaces
of fruiting pedicels glabrous ................0.
(eRAER MONEE REE eee ee 2. L. austrinum Small
H. Fruits with spreading trichomes, lower surfaces of
fruiting pedicels pubescent.................000.

G. Fruits glabrous.

I. Petals conspicuous, equalling to twice as long as the
SOURIS 4s Gaweveds eepisds wens L. virginicum L,
I. Petals inconspicuous, rudimentary, or absent, always

shorter than the sepals.
J. Upper cauline leaves pinnatifid to laciniate; axis of
the infructescence with trichomes to 0.8 mm long.
K. Fruiting pedicels 1.5-2 (-2.5) mm long, stems
usually prostrate .... 5. L. oblongum Small
K. Fruiting pedicels (2-) 3-4.5 mm long, stems
usually erect or ascending .................

J. Upper cauline leaves entire, serrate, or dentate;
axis of the infructescence glabrous, minutely
papillose, or with trichomes usually less than 0.1
mm long.

L. Basal leaves bi- or tripinnatisect, seeds wingless,
lower surfaces of young fruiting pedicels
sparsely papillose, apical wing of fruit sub-
acute, plants fetid .......... L. ruderale L.

L. Basal leaves once pinnatifid to undivided, seeds
narrowly winged or margined (wingless in L.
africanum), lower surfaces of young fruiting
pedicels glabrous, apical wing of fruit obtuse,
plants not fetid.

M. Fruits oblong or rarely oblong-ovate, seeds
wingless....1. L. africanum (Burm. f.) DC.

M. Fruits orbicular to broadly ovate or obovate,
seeds with a marginal or distal narrow wing.

N. Axis of the infructescence with straight,
usually subclavate, minute papillae; fruits

1986] Al-Shehbaz — Cruciferae 351

obovate to oblong-obovate or rarely sub-
orbicular, usually widest above the mid-
dle; cotyledons always incumbent.......
Se eer er L. densiflorum Schrader
N. Axis of the infructescence with curved,
usually subappressed, minute trichomes,
rarely glabrous; fruits orbicular to broadly
ovate, widest at or below the middle,
cotyledons accumbent or obliquely so,
very rarely incumbent... L. virginicum L.

1. Lepidium africanum (Burm. /) DC.

Specimens examined: South Carolina. Jamestown, Ahles & Haes-
loop 22599; Ahles & Haesloop 25814; Ahles & Haesloop 40523;
Johnsonville, Ahles & Haesloop 53524.

This species isa native of tropical and southern Africa. It is
naturalized in Australia and is a wool-alien in Britain (Hewson,
1982; Ryves, 1977). It is reported here from North America for the
first time. Lepidium africanum closely resembles L. ruderale but
differs from it in its larger size, undivided basal leaves, and arcuate
fruiting pedicels. Lepidium ruderale has bi- or tripinnatisect basal
leaves and straight fruiting pedicels. The origin of the material cited
here is very likely Africa because of the presence of other African
wool-alien species in the same localities.

2. Lepidium austrinum Small
Specimens examined: South Carolina. Jamestown, Ahles & Haes-
loop 52818; Ahles & Haesloop 53361.

This North American species is distributed in Texas, New Mex-
ico, Oklahoma, Kansas, and northern and central Mexico. It has
not been recorded from the southeastern United States. Hitchcock
(1936) recognized Lepidium austrinum as a distinct species, but later
(Hitchcock, 1945a) reduced it to L. lasiocarpum Nutt. var. orbicu-
lare (Thell.) C. L. Hitchc. In my opinion, L. austrinum is sufficiently
distinct to merit specific rank.

3. Lepidium bonariense L.
Specimen examined: South Carolina. Jamestown, Ahles & Haes-
loop 40522.

352 Rhodora [Vol. 88

It was not possible to trace the origin of the seed material from
which the above collection was derived. Lepidium bonariense,
recorded here for the first time for North America, is indigenous to
South America and is a widely naturalized weed in Australia, west-
ern Europe, and tropical and southern Africa (Hewson, 1982; Car-
valho e Vasconcellos, 1964; Jonsell, 1975; Marais, 1970). I follow
Hitchcock (1945b) and Thellung (1906) in treating this polymorphic
species without infraspecific taxa. Thellung (1912, 1914) recognized
eight varieties in L. bonariense, but | am unable to find adequate
morphological grounds for maintaining most of them.

4. Lepidium lasiocarpum Nutt.
Specimens examined: South Carolina. Jamestown, Ahles & Haes-
loop 22598; Ahles & Haesloop 38301.

The native range of this species extends from Colorado westward
into California and southward into the southwestern United States
and adjacent northern Mexico. Lepidium lasiocarpum has been
reported from Louisiana by Thomas & Allen (1982), but it is ap-
parently a rare plant in that state. Of the eight varieties recognized
by Hitchcock (1936, 1945a) in Lepidium lasiocarpum, the collec-
tions cited here belong to the type variety.

5. Lepidium oblongum Small
Specimens examined: South Carolina. Jamestown, Ahles & Haes-
loop 22559; Ahles & Haesloop 38302; Ahles & Haesloop 53476.

Lepidium oblongum is distributed from California eastward
through the southwestern states, Kansas, and central Arkansas and
southward into northern and central Mexico and Guatemala.
Hitchcock (1945a) recognized two varieties in L. oblongum. All of
the three collections cited here belong to the type variety.

6. Lepidium schinzii Thell.

Specimens examined: South Carolina. Johnsonville, Ahles 4045]
(2).

This is the first record of this South African species in the New
World, and the collection cited must have been derived from seeds
introduced from its native range. Lepidium schinzii is also a wool-
alien in Europe. It is easily distinguished from the other lepidiums in

1986] Al-Shehbaz — Cruciferae 353

North America by its appressed-erect fruiting pedicels and by its
stems with retrorse, flattened, scalelike trichomes.

SISYMBRIUM

The genus Sisymbrium is represented in North America by an
indigenous species, S. auriculatum A. Gray (Texas, New Mexico,
and northern Mexico), and by six Eurasian weeds (Rollins, 1981).
Almost all of the North American species assigned previously to
Sisymbrium by Payson (1922) are now believed to belong to Thely-
podiopsis Rydb. and Schoenocrambe Greene (Rollins, 1982). The
following discussion deals with two species, of which one is a new
addition to the New World’s weed flora.

1. Sisymbrium turczaninowii Sond.
Specimens examined: South Carolina. Jamestown, Ahles & Haes-
loop 42907; Johnsonville, Ahles 40434 (2):

This South African species has not been previously recorded from
the New World. It is a densely pubescent perennial in its native
habitat, but the specimens above are annuals with a sparse indu-
mentum. The characteristic features of Sisymbrium turczaninowii
are: pinnatifid, sessile upper cauline leaves with dentate lobes;
ebracteate inflorescences; stout, short (1-2.3 cm), terete, erect sil-
iques with antrorsely strigose valves; and small 0.7-0.8 X 0.3-0.5
mm), biseriate seeds.

2. Sisymbrium loeselii L.
Specimens examined: South Carolina, Jamestown, Ahles & Haes-
loop 53808; Johnsonville, Ahles 53754.

This Eurasian weed is abundant in the Canadian prairie provinces
and in some of the mountain states (Rollins, 1981). It is sporadic in
widely separated localities in New England, Pennsylvania, Michi-
gan, and North Dakota, but has not been recorded from the south-
eastern states. Sisymbrium loeselii is eaily distinguished from the
other sisymbriums in North America by the combination of its div-
aricate fruiting pedicels narrower than the siliques, the terete, short
1-4.5 cm) siliques not overtopping the floral buds, and its 4-7 mm
long petals.

354 Rhodora [Vol. 88

ACKNOWLEDGMENTS

I thank Reed C. Rollins for his critical review of the manuscript
and for making available the specimens on which this study is based.
I am indebted to Barbara Nimblett for typing the manuscript. I
thank an anonymous reviewer for his careful review of the
manuscript.

LITERATURE CITED

CARVALHO E VASCONCELLOS, J. 1964. Lepidium. In: Tutin, T. G. et al., Eds. Flora
Europaea 1: 330-333.

Hewson, H. J. 1982. The genus Lepidium L. (Brassicaceae) in Australia. Bruno-
nia 4: 217-308.

Hitchcock, C. L. 1936. The genus Lepidium in the United States. Madrofio 3:
265-320.

1945a. The Mexican, Central American, and West Indian Lepidia. /bid.
8: 118-143.

1945b. The South American species of Lepidium. Lilloa 11: 75-134.

JONSELL, B. 1975. Lepidium L. (Cruciferae) in Tropical Africa. A morphological,
taxonomical and phytogeographic study. Bot. Notiser 128: 20-46.

Marais, W. 1970. Cruciferae. Jn: Codd, L. E. et al., Eds. Flora of Southern
Africa 13: 1-118.

Payson, E. B. 1922. Species of Sisymbrium native to North America north of
Mexico. Univ. Wyoming Publ. Sci. I: 1-27.

Raprorpb, A. H., H. E. AHLES, AND C. R. BELL. 1968. Manual of the Vascular
Flora of the Carolinas. The University of North Carolina Press, Chapel Hill,
North Carolina.

Roiuins, R.C. 1981. Weeds of the Cruciferae (Brassicaceae) in North America.
Jour. Arnold Arb. 62: 517-540.

1982. Thelypodiopsis and Schoenocrambe (Cruciferae). Contrib. Gray
Herb. 212: 71-102.

Ryves, T. B. 1977. Notes on wool-alien species of Lepidium in the British Isles.
Watsonia | 1: 367-372.

SmiTH, E. B. 1978. An Atlas and Annotated List of Vascular Plants of Arkansas.
University of Fayetteville, Fayetteville, Arkansas.

THELLUNG, A. 1906. Die Gattung Lepidium (L.) R. Br. Eine monographische
Studie. Mitt. Bot. Mus. Ziirich 28: 1-340.

1912. Lepidii generis formae novae ex Museo botanico Berolinensi,
Repert. Sp. Nov. [1: 309-310.

1914. Lepidium bonariense L. novis varietatibus ex herbario Stuckerti-
ano auctum. /bid. 13: 301-303.

THOMAS, R. D. AND C. M. ALLEN. 1982. A preliminary checklist of the dicotyle-
dons of Louisiana. Contrib. Herb. Northeast Louisiana Univ. 3: 1-130.

1986] Al-Shehbaz — Cruciferae x5)

ARNOLD ARBORETUM
HARVARD UNIVERSITY
22 DIVINITY AVE.

CAMBRIDGE, MA 02138

CURRENT STATUS OF MAGNOLIA VIRGINIANA
IN MASSACHUSETTS

RICHARD B. PRIMACK, EDWARD HENDRY AND PETER DEL TREDICI

ABSTRACT

The sweet bay magnolia (Magnolia virginiana L.) from Gloucester, Massachusetts,
was almost eliminated by flower collectors in the early 20th century from its one
locality in New England. The population appears to have recovered since that time
and now consists of approximately 129 multi-stemmed plants which have a total of
1800 stems. Approximately 33% of the plants produced fruits in 1984. Plants have
shown a positive growth in height and number of stems in the two-year period,
1982-1984. No naturally occurring seedlings were evident. The population appears to
be persisting through the vegetative vigor of existing plants, some of which may be
long-lived. Seedlings grown from seed collected in the Gloucester stand in 1980 and
transplanted back into the area in 1982 showed excellent survival (48%) and moderate
growth as of 1984.

Key Words: Magnolia, rare plants, ecology, Massachusetts

INTRODUCTION

The sweet bay magnolia (Magnolia virginiana L.) has only one
known population in New England, with the next population occur-
ring 150 miles to the south on the eastern shore of Long Island
(Little, 1971). The New England population of this small, freely-
sprouting tree occurs in a swamp near the coast in Gloucester, Mas-
sachusetts. Since its discovery, this plant population has received
the attention of both botanists and the general public because of
status as a “southern” species growing far from its normal range.
The residents of the Kettle Cove section of Gloucester, where the
magnolia grows, were so enthusiastic over the discovery of the plant
that they changed the name of the community to Magnolia in the
19th century. Del Tredici (1981) has reviewed in detail the history of
the population, horticultural varieties of the species, and seed ger-
mination behavior. The purpose of this article is to present a statis-
tical summary of the current status of the population, particularly in
relation to the past condition of the population. These data will be
used to determine whether current management of the site is
appropriate.

357

358

1623.
1806.

1814.

1858.

1875.

1889.

Rhodora [Vol. 88

HISTORICAL BACKGROUND

The town of Gloucester was founded.

Theophilus Parsons discovered the magnolia population
while riding in a carriage along woodland road between
Kettle Cove and Fresh Water Cove in Gloucester. The loca-
tion described by Parsons (from Kennedy, 1916) is the
“swamp on the western side of the road as you go from
Manchester to Gloucester, and before you come to a large
hill over which the road formerly passed. It grows so near
the road as to be visible even to the careless eye of the
traveler.”

Jacob Bigelow (1814) in the Plants of Boston stated that the
magnolia “attains a height of a dozen feet, but is sometimes
killed down to the roots by severe winters. ...It grows
plentifully in a sheltered swamp at Gloucester, Cape Ann.”

Henry David Thoreau (1962) visited the swamp on Sep-
tember 22. He described the magnolia bushes as being 2 to 7
feet in height. The susceptibility to frost-damage was again
noted: “I saw last year’s shoots, which had died down sev-
eral feet, and probably will be the fate of most which has
grown this year.”

People had begun collecting flowers from the magnolia
plants to sell in Boston and Salem. So widespread had this
practice become that George B. Emerson (1875) writing in A
Report on the Trees and Shrubs Growing Naturally in the
Forests of Massachusetts said that scores of trees had been
broken down by people collecting flowers to sell.

J. G. Jack (1889) confirmed how desperate the situation had
become for this population: “So eagerly have the flowers
been sought for by collectors and especially by those who
wished to make money out of the sale of both plants and
flowers, that there has been some apprehension that the day
would soon come when the Magnolia could only be classed
in New England floras as one of the indigenous plants of the
past.” In that same year, some hope for the protection of the
population developed. Mr. Samuel E. Sawyer, the owner of
the swamp, set aside the land to be managed as “Ravens-
wood Park”, for the enjoyment of the public.

1986] Primack, Hendry & Del Tredici — Magnolia 359

1913. Unfortunately, the condition of the population continued to
deteriorate. C. E. Faxon (quoted in Kennedy, 1916) stated,
“When I first visited the swamp some 45 years ago (roughly
1868) there were plenty of good specimens all about, some-
times 15 feet tall or more.” However, by 1913, “only two
little plants a few feet high had escaped the Magnolia
hunters—such had been the destruction!”

1970. The trustees of Ravenswood Park recognized that the mag-
nolias were being shaded out by the growth of large canopy
trees of hemlock, white pine, and red maple. Consequently,
a program of thinning out the large trees was initiated.

1980. Peter Del Tredici (1981) of the Arnold Arboretum observed
that the magnolias appeared to be growing well and fruiting.
He estimated that there were 40 to 50 multi-stemmed mag-
nolia clumps, most being 6 to 12 feet in height, with one
plant 18 feet in height. Del Tredici commented that the
swamp at present must be more shaded than the past, as
shown by the abundance of the shade-tolerant inkberry (Ilex
glabra) which was not mentioned as being present by Jack
(1889). Also, Jack (1889) listed the light-demanding cran-
berry (Vaccinium macrocarpon) as growing densely in the
swamp, but this plant no longer grows there.

These scattered observations provide a picture of a magnolia
population that was probably extensive in the early 19th century,
declined greatly in the late 19th and early 20th century as a result of
stems being broken down during collection of flowers, and then
recovered to its present status. However, the exact status of the
population is difficult to determine without a quantitative survey of
the population in which individuals are tagged, mapped and meas-
ured through time.

MATERIALS AND METHODS

A study was initiated on November 14, 1982 by two of the authors
(RBP and PDT), accompanied by two Boston University students.
During this survey, every multi-stemmed plant clump was tagged
with a metal tag, typically at shoulder height. For a few plants, there
was some difficulty estimating what constituted one genetic or phy-
siological individual, since stems are sometimes bent to the ground

360 Rhodora [Vol. 88

and root, forming new clumps of shoots at distances of up to 3 m
from the original clump. After tagging, a rough map was made of
the population and plants were evaluated for the following three
characters:

|. HEIGHT was visually estimated to the nearest 0.3 m for the
tallest stem in aclump. Estimates were improved by practicing
on several plants measured with a tape-measure. Some error is
probably associated with this estimate.

2. NUMBER OF STEMS for the plant was counted, with every shoot
emerging from the ground counted as a separate stem. Often
many new stems would develop from an existing stem which
had been bent down and partially covered with leaves.

3. NUMBER OF FRUITS present on the plant. This number is a signifi-
cant underestimate of fruit production since many fruits had
already fallen off the plants.

The site was revisited on November 3, 1984, the field party con-
sisting of the three authors and eleven Boston University students.
Previously tagged plants were located, and measurements taken
again on the three plant characters. The larger field party made it
possible to survey the area more intensively and locate plants missed
in the previous survey. This effort provided a confirmation of Wu’s
law of plant population biology, “The more you look, the more you

”

see.

RESULTS

1982. 119 clumps were located and tagged, many more than had
been anticipated. Heights of the clumps varied from 0.3 m to 8.4 m,
with a mean of 3.6 m. Number of stems per clump varied from | to
64, with a mean of 15.2 stems. Many of these stems were relatively
small and often less than 0.6 m in height. The total number of stems
in the population is about 1800. Using Spearman Rank Correla-
tions, there is no correlation between plant height and the number
of stems (r = 0.139), indicating that these two measures of plant size
are essentially independent of each other. The distribution of height
in the population appears to be normal (Figure 1), indicative ofa
mature population in which adult trees are persisting and growing,
but with relatively little regeneration by seed. The same pattern
holds when number of stems per plant is examined (Figure 2): 32

1986] Primack, Hendry & Del Tredici — Magnolia 361

40
we 30
a ]
Cc
.e)
are
=
oO
. oT
re) 0)
0
0-12 13-24 25-36 37-48 49-60 61-72 > 72
Height (dm)
30
—”
= a
Cc
5S 2
a.
—
oO 10
Oo
ie)
1-4 5-8 9=+12 13-16 17-20 21-24 >24
Number of Stems
Figure 1-2. Distribution of plant height and number of stems per plant in the

magnolia population in 1982. 1. Plant height. 2. Number of stems per plant.

plants had fruits, with an average of 6.7 fruits each; one plant had 37
fruits.

1984. Of the original 119 plants, 114 were re-located, 3 plants
could not be found, and 2 plants were apparently dead; in addition,
15 new plants were located. While one of these plants may have been
established from seed since 1982, the other 14 plants are clearly

362 Rhodora [Vol. 88

older plants that were overlooked in the 1982 census or were beyond
the fringes of the area then surveyed. The larger size of the 1984
survey team made possible our locating these additional plants. 129
plants were tagged, mapped, and measured in 1984.

In 1984, mean plant height of the original plants was 3.9 m, a
significantly greater height than in 1982, (t-test; p < 0.05). These
results suggest that the population is increasing in height. Another
indication of vegetative growth was that 72 out of 112 plants showed
an increase in height, compared with 1982 measurements, while only
22 plants showed a decline in height; 18 plants measured no change.

Mean number of stems per plant of the original plants was 17.5,
again a significant (p < 0.05) increase over 1982. The largest plant
had 104 stems in 1984, while the same plant had only 64 stems in
1982. A total of 74 plants showed an increase in the number of stems
over 1982, 24 plants showed a decrease, and 15 plants showed no
change.

The fact that the new plants found in 1984 were probably just
overlooked in 1982 is emphasized by the fact that they had a mean
height (3.4 m) and a mean number of stems (12.8) which were
smaller but reasonably similar to the values of the tagged plants.

A total of 42 plants produced fruit in 1984, representing 33% of
the population. Of these same plants, 24 had also produced fruit in
1982, indicating some stability in the reproductive structure of the
population. Fruiting plants produced an average of 5.1 fruits with a
total of 214 fruits in the population. One plant produced 37 fruits.

In the fall of 1980, seeds were collected from the site and grown in
pots at the Arnold Arboretum. In May 1982, 500 of these 18 month-
old seedlings were transplanted back into the swamp at two loca-
tions, one about 50 meters from the northern edge of the wild
population and the other at the southern edge. Prior to planting, the
canopy was opened up in these two areas by removal of several large
trees. As of November 1984, 239 of the seedlings (48%) were still
alive. The tallest plants were about 75 cm high with most plants in
the 15-25 cm range. When planted, the seedlings were all under 10
cm. While this increase in height is not particularly dramatic, it is
not much less than that shown by ten seedlings from the same seed
lot grown under light shade in the outdoor nursery at the Arnold
Arboretum in Jamaica Plain, Mass. As of February, 1985, these
nursery seedlings planted at the same time as the Ravenswood
plants, ranged in height from 20 to 54 cm, with an average height of
31 cm.

1986] Primack, Hendry & Del Tredici — Magnolia 363

DISCUSSION

The population of sweet bay magnolias in Gloucester has clearly
recovered from those dismal days of the late 19th and early 20th
century when all of the stems were broken down. The population is
healthy in many ways. Roughly 35% of the plants have been flower-
ing and fruiting, suggesting that the plants are in good physiological
condition. There is an abundance of plants (129 clumps), a great
increase over the 2 plants observed in 1913. The number of stems
observed in 1982 (approximately 1800) is impressive, and argues for
the health of the population, even though many of the stems are
small. Measurements of height and number of stems from 1982 and
1984 indicate that the average size of individuals is increasing.
Further, the human pressure on the population has obviously
decreased; the area is now in a well-managed park and people no
longer collect the flowers to sell and probably rarely visit the swamp
where the magnolias grow. These factors all indicate that the popu-
lation is in good condition.

The fact that no seedlings or young plants were noted in the
Ravenswood population, despite good seed production, indicates
that the population is under some limitation. Even with the tree
thinning of 1970, the swamp is still too shaded to allow seedling
establishment and consequently a self-maintaining magnolia popu-
lation. The continued presence of shade-tolerant inkberry and the
absence of light-demanding cranberries further emphasizes that the
swamp is probably too shaded for magnolia establishment.

The main mechanism maintaining the population at present is
clearly vegetative reproduction, involving abundant sprouting from
the root systems and rooting of stems that have been bent down by
large branches falling from the trees above. This rooting of prostrate
stems provides a mechanism for limited lateral movement of a
clone. It is a reasonable speculation that most of the magnolia
plants present in the swamp today were there in the early 19th
century when the population was first discovered, and have main-
tained themselves until the present time by sending up new stems
from the persistent root system, much like the chestnut tree (Casta-
nea dentata) does today. Magnolia plants in southern savannas
sprout readily after fires, and these Gloucester magnolias are
adapted to sprout following a severe frost which kills the stems. This
ability to sprout pre-~adapted the species for recovery when stems
were broken during collection of flowers. Further, we speculate that
several of the larger magnolia clumps might have been present in the

364 Rhodora [Vol. 88

swamp for several thousand years, or ever since the population was
first established following retreat of the glacier. It is not possible to
prove or disprove this speculation at present. However, detailed
studies of isozymic variation on this population, as has been done
for salt-marsh cordgrass (Silander, 1979) and alder (Huenneke,
1985), might reveal the extent to which individual genotypes have
spread through the swamp as a result of vegetative reproduction.
An alternative hypothesis would be that Magnolia virginiana
invaded the swamp via bird dispersal at some point in the eighteenth
century following logging and/or burning of the swamp. Following
this initial invasion, the magnolias did not become a prominent
feature of the landscape until 1804, when first noticed by Theophi-
lus Parsons. This hypothesis would explain why the plant went
undiscovered for so long in an area that was settled in 1623.

ACKNOWLEDGMENTS

Funds to support this project came from the Boston University
Biology Department and the Arnold Arboretum of Harvard Uni-
versity. Help of our many field assistants is appreciated, particularly
on those cold, damp November days. Statistical assistance was pro-
vided by Pamela Hall. Helpful comments on the manuscript were
provided by Bruce Sorrie, Kamal Bawa, and Pamela Hall.

LITERATURE CITED

BiGELow, J. 1814. Florula Bostoniensis: A Collection of Plants of Boston and Its
Environs. Cummings & Hillard, Boston.

Det Trepici, P. 1981. Magnolia virginiana in Massachusetts. Arnoldia 41:
36-49.

Emerson, G. B. 1875. A Report on the Trees and Shrubs Growing Naturally in
the Forests of Massachusetts, 2d ed. Little, Brown, and Co., Boston.

FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Co.,
N.Y.

HueNNeKE, L. F. 1985. Spatial distribution of genetic individuals in thickets of
Alnus incana spp. rugosa, a clonal shrub. American Journal of Botany 72:
152-158.

Jack, J.G. 1889. Magnolia glauca in its most northern home. Garden and Forest
2: 363-364.

KENNEDY, G. G. 1916. Some historical data regarding the sweet bay and its sta-
tion on Cape Ann. Rhodora 18: 205-212.

Littte, E.L. Jr. 1971. Atlas of United States Trees, Vol. 1: Conifers and Impor-
tant Hardwoods. Misc. Publ. 1146 U.S.D.A. Forest Service.

1986] Primack, Hendry & Del Tredici — Magnolia 365

SILANDER, J. A. JR. 1979. Microevolution and clone structure in Spartina patens.
Science 203: 658-660.

THOREAU, H. D. 1962. The Journal of Henry D. Thoreau. Reprint of 1906 ed.
Dover Publ., New York.

R. B. P. AND E. H.
BIOLOGY DEPARTMENT
BOSTON UNIVERSITY
BOSTON, MA 02215

P. DT.

ARNOLD ARBORETUM OF
HARVARD UNIVERSITY

JAMAICA PLAIN, MA 02130

STUDIES IN THE ARISTIDA (GRAMINEAE)
OF THE SOUTHEASTERN UNITED STATES.
IV. KEY AND CONSPECTUS.!

KELLY W. ALLRED

ABSTRACT

The grass genus Aristida in the southeastern United States is reviewed. Twenty
species are present in the region, all indigenous to the area. New combinations
proposed are Aristida purpurescens Poir. var. tenuispica (Hitche.) Allred and var.
virgata (Trin.) Allred.

Key Words: Aristida, floristics, southeastern United States

This paper concludes a series of investigations of the Aristida of
the southeastern United States. The first paper dealt with the
purpurescens-tenuispica-virgata complex, the second with A.
intermedia and A. longespica, and the third compared the /anosa
and palustris entities (Allred, 1984a, 1985a, b). All were phenetic in
nature and based solely on herbarium collections. These studies
were preparatory to a treatment of this genus for the Vascular Flora
of the Southeastern United States. Several questions have arisen
and remain unanswered by the studies. For example, little is known
about chromosome numbers, breeding systems, or extent of eco-
typic variation of these grasses. It is hoped that such information
can be acquired from future systematic studies of this troublesome
but interesting genus.

Examination of Aristida collections from 26 herbaria (see
Acknowledgments) resulted in the documentation of 20 species for
the region covered by the Vascular Flora of the Southeastern United
States project (Alabama, Arkansas, Delaware, Florida, Georgia,
Kentucky, Louisiana, Maryland, Mississippi, North Carolina,
South Carolina, Tennessee, Virginia, and West Virginia). For all
taxa recognized herein, one of the types or a photo of the type was
seen. Although all taxa were indigenous to the southeastern United
States, there are three records of adventives worthy of note. Gleason
and Cronquist (1963) reported A. adscensionis L. as being “rarely

‘Journal Article No. 1111 of the New Mexico Agricultural Experiment Station, Las
Cruces, NM 88003.

367

368 Rhodora [Vol. 88

adventive” in the northeastern United States, and Hitchcock and
Chase (1951) reported it as far east as Missouri, but no specimens
were seen of this species from the southeast flora region. In addi-
tion, there were several collections of Australian species in James-
town, South Carolina, from around a wool-combing mill. These
grasses combined features of Aristida armata Henrard, A. bentha-
mii Henrard, and A. inaequiglumis Domin; they are easily distin-
guished from any North American Aristida by a groove (sulcus) on
the ventral side of the floret, formed by the involute margins of the
lemma that do not completely enclose the palea and flower. Also,
Aristida divaricata Willd. was collected from the Soil Conservation
Service Nursery at Chapel Hill, North Carolina in 1941. This species
is found on the dry southwestern plains and will probably not
become naturalized as part of the southeastern flora.

A few terms should be defined. The awn column is formed by the
connivent or coalescent bases of the awns above the lemma, and is
usually twisted (for example, Aristida tuberculosa). This structure
differs from the beak of the lemma, which is the slender, narrowed
terminal portion of the lemma body just below the awns; it is also
often twisted (for example, A. spiciformis). The awns themselves are
free from one another, and are measured from the summit of the
beak or awn column to the tip of the awn.

At this point in our understanding of these Aristida, it would be
unwise to speculate on the phylogeny of the taxa, but it may be
helpful to give some indication of the apparent morphologic rela-
tionships among them. The groupings that follow are based only on
external morphology and the intuitive insights gained after examin-
ing several thousand herbarium specimens. The taxa included
within a group may be considered to be generally similar to one
another, but no biologic relationships are implied. The groups are
informal, with no taxonomic standing or rank, and in some cases
are based on similar such groups found in Hitchcock and Chase
(1951).

Group Dichotomae. All of these taxa have a tendency toward
coiling of the central awn; this coiling is most pronounced in Aris-
tida dichotoma and A. basiramea, and only slightly developed in A.
longespica. Aristida oligantha and A. ramosissima intergrade, as do
A. dichotoma and A. basiramea.

1986] Allred — Aristida 369

. oligantha

. ramosissima

. dichotoma vars. dichotoma and curtissii

. basiramea

. longespica vars. longespica and geniculata

mR RR»

Group Tuberculosae. These taxa are members of the generic
section Arthratherum, in which the lemma is jointed at the summit
and breaks from the awn column. Our species differ mainly in size
of the spikelet parts.

A. tuberculosa
A. desmantha

Group Purpureae. This group enters our region only in a few
scattered localities in Arkansas and Louisiana. The nearest relatives
are all plains and desert taxa of the southwestern United States,
Mexico, and South America.

A. purpurea vars. purpurea and longiseta

Group Purpurescentes. More heterogenous than the others, this
group contains taxa that are very similar to Aristida purpurescens
(tenuispica, virgata, condensata, gyrans) as well as taxa that seem to
fit nowhere else but have a general similarity of panicle or spikelet.
Aristida stricta, A. rhizomophora, and A. lanosa have peculiar
vegetative features that distinguish them.

. purpurescens vars. purpurescens, tenuispica, and virgata
. condensata

. gyrans

. palustris

. mohrii

. simpliciflora

. Stricta

. lanosa

. rhizomophora

Group Divaricatae. Used by Hitchcock and Chase (1951) for
species with abruptly spreading panicle branches, this group is
expanded somewhat to include otherwise similar species with
reduced lateral awns. Aristida floridana is tentatively placed here
because of its similarity to A. ternipes, a species of the western

RoR RAA AAA DA

370 Rhodora [Vol. 88

deserts, which in turn is very close to A. patula and other members
of the group Divaricatae.

A. patula

A. floridana

Group Spiciformes. This Aristida group shows a parallel devel-
opment to the group 7uberculosae in the long, twisted lemma beak.
It differs, however, in that the beak appears to be an elongation of
the lemma body rather than a twisting together of the awns. Also,
the dense panicle and unequal, awned glumes are distinctive in this
group.

A. spiciformis

Following the key is a conspectus for each numbered species
which gives the correct name and author, pertinent synonyms, iden-
tifying features, general habitat and distribution, and miscellaneous
notes. The distribution data are provided only for those states
covered by the “Southeastern Flora,” and not the entire range of
the species. A representative spikelet of each taxon is illustrated
with figures taken from Henrard (1929, 1932).

KEY TO THE SPECIES

1, Firat GIMME 3-7 NOVO 5 oc 4-45ouedtiewnebbawnsesaadeanvees (2)
2. Central awn of lemma 0.9-3 cm long; awn of second glume 3-7
Ti ON a oon hee se esas ds weer ceeaes 15. A. ramosissima

2. Central awn of lemma 3-7 cm long; awn of second glume
VOnd7 WOM IONE «ci esisss ve eas cenvnses 10. A. oligantha

D5. SUSE RI aE ROVE oo gp ooo ns oe noses Heeen Oda eers (3)
3. Central awn spirally coiled at the base like a corkscrew. .. .(4)
4. Lateral awns 4 mm or less long, erect ..... 4. A. dichotoma

4. Lateral awns 5-12 mm long, spreading.... 1. A. basiramea

3. Central awn not spirally coiled, straight to curving or con-
torted, but not corkscrew-like ............ 0.0.0 ee eee (5)

5. Lateral awns markedly reduced, 1/3 or less the length of the
COGCENE AW 6 oho 5 6 ee eds (6)

6. Plants annual; panicle narrow, rarely wider than 5 cm, the

DERNC BCE BPOCE sa verccesaicaddasase 8. A. longespica

6. Plants perennial; panicle open, almost always wider than

1986] Allred — Aristida 371

5 cm, the branches spreading ................... (7)

7. Lateral awns 0-1 mm long; central awn sickle-shaped
Se ee ee ST re Er Deere tet 5. A. floridana

7. Lateral awns 3-10 mm long; central awn straight to
somewhat flexuous but not sickle-shaped..........
bPogshes SEeGh ewe ney ah ceuyiemer es 12. A. patula

5. Lateral awns well developed, at least 1/2 the length of the
CO OIG ito eos waving mwas ny emaodtenecs (8)

8. Leaf blades tightly involute, stiff and wiry, the young
blades pilose on the lower (outer) surface near the mid-

rib especially toward the base, also often densely pilose

at the throat and collar region ......... 19. A. stricta
8. Leaf blades, flat, or if involute then glabrous at the throat
BNO COMAL TEMION: <4 nw s0 20s Rei das'wainnkeeeundka (9)

9. Nodes of panicle axis with a tuft of lanose or floccose
hairs: sheaths lanose or floccose, the hairs cobwebby,
kinked and intertwined .............. 7. A. lanosa

9. Nodes of panicle axis glabrous or sparsely pilose;
sheaths glabrous to pilose, the hairs straight, not
cobwebby, and usually appressed ............ (10)

10. Lemma body separated from the awns or awn
column by a joint (zone of articulation), the body
and awns eventually disarticulating (check mature
BPS 4 oss eso oo ad danke ecineunees (11)

11. Panicle spiciform, dense, compact, the spikelets
very congested, the branches appressed; plants
perennial, the culms simple or sparingly
DIONCNGO a3. 70043 sore ate 18. A. spiciformis

11. Panicle open, loose, somewhat fan-shaped, the
spikelets only slightly congested, the branches
ascending to spreading; plants annual (may be
robust), the culms usually much branched ....
(eCtseea ena he Deedee ees ah k es os (12)

12. Glumes, at least the second, longer than 2 cm;
awn column (formed by a twisting together

of the awn bases) 8-15 mm long; callus

mostly 3-4 mm long .... 20. A. tuberculosa

372 Rhodora [Vol. 88

12. Glumes both shorter than 2 cm; awn column

2-5, rarely 7, mm long; callus mostly 2-2.5

MM. JONG. cavews ceexaces 3. A. desmantha

10. Lemma body passing directly into the awns, without

a joint of articulation, the body and awns not
CiSATUCINAUNG. coisas eves neeasseeewes (13)

13. Summit of lemma with an obvious, elongated beak mostly
10-30 mm long (rarely only 7 mm long) ............+2+06-
eT ToT eT Tee TT Te eee Te 18. A. spiciformis

13. Summit of lemma without a beak, or if slightly beaked then
obscure and less than 7 mm long... .......0eccessees (14)

14. Panicle open at maturity, the branches widely spreading to
reflexed from axillary pulvini; first glume longer than the
SACOG ac aa n4s54cdhanen cease ese caueee es 12. A. patula

14. Panicle narrow, with mostly erect-appressed or ascending
branches without axillary pulvini, if the branches somewhat
open and drooping (in A. purpurea and A. rhizomophora)

then the first glume shorter than the second ......... (15)

15. Plants with vigorous thick rhizomes; basal sheaths becom-
ing markedly fibrous in age and shredding into thread-

MG RERINGNIS i cee sca esac noes 16. A. rhizomophora

15. Plants tufted, lacking rhizomes; basal sheaths not becom-
ing fibrous and not shredding into thread-like segments

Seer ee eee ete Corre rer erry re (16)
16. Spikelets borne singly at each node of the main axis, the
inflorescence a spike or raceme ........ 9. A. mohrii

16. Spikelets at least two per node of the main axis, or borne
on branches at least below, the inflorescence racemose

OF DANI ised heen esesaeswnnecesen es (17)

17. First glume 1/3 to 3/4 the length of the second glume
Ee eTE CREE NEN CLAERTR ENR CA AE RA ERA RSE (18)

18. Lemma awns 0.8-—1.5 cm long; lemma 5-6 mm long
ee Se rere Pe ree ee ere ee er 6. A. gyrans

18. Lemma awns 2-10 cm long; lemma 6-16 mm long
Te ee Te ee ee ee 13. A. purpurea

17. First glume more than 3/4 the length of the second
SMG 6s nda ARE ARTO RASS OMAST RAE EES (19)

19. PIQHts AUNUG) soo css aseassncnes 8. A. longespica

B95. Plants Perennial o.566655654 6 ees ek davon es (20)

1986] Allred — Aristida 373

20. Central awn 1.5-4 cm long; first glume promi-
nently 2-keeled and 9-13 mm long in mature
SPECINENS «656. csc ines hoods 11. A. palustris

20. Central awn 0.8-2.5 cm long; first glume mostly
I-keeled (2-keeled in some A. purpurescens)

and/or 6-9 mm long .............ee0. (21)
21. Central awn about twice as thick as the lateral awns, divergent
NG UCHGIe TONNES 6. 6 ooo blew bewadsaaaesseu en es (22)

22. All three awns divergent to reflexed, somewhat contorted at
the base; spikelets mostly two per node (occasionally 3),
one pedicelled and one subsessile..... 17. A. simpliciflora

22. Lateral awns (sometimes also the central) mostly erect to
ascending, usually not contorted at the base; spikelets
mostly many per node, but sometimes only two, pedicelled
OF SUDSESSUC. 5 55s isin earch knw 4% 14. A. purpurescens

21. Central awn about the same thickness as the laterals, erect to
IVGUION eyo siad dnt ncdenhhu sec anederee wed ewaxs (23)

23. First glume 1-4 mm longer than the second...............

a ee ee a ee ee eer eee ee 14. A. purpurescens

23. First glume no more than | mm longer than the second, or

1107 6) Se Nave oe aa A ao (24)

24. Culms usually robust, the lower shoots 3-6 mm broad;
panicle branches 4 cm or more long; first glume I-nerved

Te CTT EES eT OTe CCT TOC CC 2. A. condensata

24. Culms usually slender, the lower shoots 1-4 mm broad;
panicle branches 1-4 cm long; first glume 1-2 nerved

CeEee ee uedtee de casei sheeeda tee ees (25)
25. Callus 0.4-0.8 mm long.......... 14. A. purpurescens
ode Calis 1-2 nia TONG. 5 cs 5oaa Gee eee es 6. A. gyrans

1. Aristida basiramea Engelm. ex Vasey, Bot. Gaz. 9:76. 1884.
Branching annuals to 45 cm tall; panicles narrow and loose, (2) 4-10
cm long; glumes subequal, the first 8-10 mm long, the second 10-12
mm long; lemma about 10 mm long; central awn spirally coiled at
base in 2-3 spirals, 10-12 mm long; lateral awns usually spreading,
5-10 mm long. Distribution: Sandy waste places, fields; AL, FL.
(Figure 1).

Aristida basiramea grades morphologically into A. dichotoma
var. curtissil; see note under A. dichotoma.

374 Rhodora [Vol. 88

Figures 1-9. Spikelets and calluses of Aristida of the southeastern United States
(from Henrard, 1929, 1932). Approximate scales in parentheses are for spikelet first
and callus second. 1. A. basiramea (X 2,6). 2. A. condensata (X 1.3, 2.6). 3. A.
desmantha (X 0.8). 4. A. dichotoma var. curtissii (X 1.6, 5.6). 5, A. dichotoma
var. dichotoma (X 3.2). 6. A. floridana (X 2.4). 7. A. gyrans (X 1.6, 3.2).
8. A. lanosa (X 1, 2.4). 9. A. longespica var. geniculata (X 1.6, 4).

Allred — Aristida a75

|
\ 17 18

Figures 10-18. Spikelets and calluses of Aristida of the southeastern United
States (from Henrard, 1929, 1932). Approximate scales in parentheses are for spi-
kelet first and callus second. 10. A. longespica var. longespica (X 2.8, 4). 11. A.
mohrii (X 1, 2.6). 12. A. oligantha (X 0.8, 4). 13. A. palustris (X 1, 2.6). 14.
A. patula (X 1, 2.6). 15. A. purpurea var. longiseta (X 0.5, 3.2). 16. A. purpurea
var. purpurea (X |, 3.2). 17. A. purpurescens var. purpurescens (X 1.3, 3.2). 18.

A. purpurescens var. tenuispica (X 1.3, 3.2).

376 Rhodora [Vol. 88

Figures 19-25. Spikelets and calluses of Aristida of the southeastern United
States (from Henrard, 1929, 1932). Approximate scales in parentheses are for spi-
kelet first and callus second. 19. A. purpurescens var. virgata (X 1.6, 4). 20. A.
ramosissima (X 1.2, 4). 21. A. rhizomophora (X 1.2, 4). 22. A. simpliciflora (X
1.6, 5.2). 23. A. spiciformis (X 2,4). 24, A. stricta (X 1.6, 4.8). 25. A. tubercu-
losa (X 0.5, 1.6).

1986] Allred — Aristida Sti

2. Aristida condensata Chapm., Bot. Gaz. 3:19. 1878. [A. conden-
sata var. combsii (Scribn. & Ball) Henr.] Erect, robust perennials to
1.5 m tall; sheaths glabrous to pilose, but not lanose; panicles nar-
row, (15) 20-55 cm long, the branches mostly longer than 4 cm,
erect to ascending or slightly spreading; glumes subequal, 6-10 (12)
mm long; lemma 6-8 mm long; awns usually contorted at the base,
the central 10-15 mm long, the laterals 8-13 mm long. Distribution:
Pine savannahs, oak-pine uplands, sandy hills; AL, FL, GA, MS,
NC; SC.-(Figure 2):

Specimens with glabrous lower sheaths have been recognized as
Aristida condensata var. combsii. There appears to be no geogra-
phic or morphologic basis other than pubescence for this variety,
and it is submerged within A. condensata without formal recogni-
tion. Aristida condensata is part of the A. purpurescens complex
and needs to be studied relative to the other entities of this group:
purpurescens, virgata, tenuispica, and perhaps also stricta, gyrans,
simpliciflora, and mohrii.

3. Aristida desmantha Trin. & Rupr., Acad. St. Petersb. Mem. VI.
Sci. NAt. 5:109. 1842. Branching annuals to 80 cm tall; sheaths
glabrous to pilose-floccose; panicles loose and open, 10-20 cm long,
the branches ascending and the spikelets erect; glumes subequal, the
body 10-12 mm long with an awn 2-5 mm long, the first slightly
longer than the second, lemma 7-10 mm long; awn column 2-5 (7)
mm long, articulated with the lemma summit; awns subequal, 2-3
cm long. Distribution: Fields, waste places, dry pine woods; AR,
LA. (Figure 3).

Aristida desmantha and A. tuberculosa are members of the
mostly African section Arthratherum, wherein the lemma is termi-
nated by an awn column that breaks from the lemma at maturity.
The point of articulation is not always evident, especially in young
plants.

4. Aristida dichotoma Michx., Fl. Bor. Amer. 1:41. 1803.
First glume as long as or nearly equal to second glume; lemma
sparsely appressed-pubescent, 3-8 mm long ................
Serer ee ee es eee eee ee ee ee ee 4a. var. dichotoma
First glume 1/2 to 2/3 as long as second glume; lemma glabrous
to scaberulous, 6-11 mm long.............. 4b. var. curtissii

378 Rhodora [Vol. 88

4a. A. dichotoma var. dichotoma. Plants annual, culms usually
much-branched at base; panicles narrow, spikelike 2-11 cm long;
glumes subequal, the first 3-6 mm long, the second 4-7 mm long;
lemma 3-8 mm long; central awn spirally coiled, 3-6 mm long;
lateral awns erect, about | mm long. Distribution: Disturbed areas,
fields, pine clearings; throughout the region. (Figure 5).

4b. A. dichotoma var. curtissii A. Gray, Man. Bot. North. U.S. ed.
6. 640. 1890. [A. curtissii (A. Gray) Nash; A. basiramea Vasey var.
curtissii (A. Gray) Shinners.] Plants annual, the culms branched at
base but less so than in var. dichotoma; panicles spikelike 2-11 cm
long; glumes unequal, the first 5-8 (10) mm long, the second 7-13 mm
long; lemma 6-11 mm long; central awn spirally coiled, 4-8 mm
long; lateral awns erect, 2-4 mm long. Distribution: Pine woods,
fields, disturbed areas; AL, AR, FL, GA, MD, MS, NC, SC, TN,
VA, WV. (Figure 4).

I follow here the treatment of Vaughn (M.S. thesis, Univ. of
Oklahoma, Norman, 1981) in relegating Aristida curtissii to varietal
status under A. dichotoma. Vaughn demonstrated the phenetic
intermediacy of A. curtissii relative to A. dichotama and A.
basiramea, but with greater similarity to A. dichotoma. Aristida
basiramea seems to be no more distinct from A. dichotoma than is
var. curtissii, and could likewise be treated as a variety of A. dicho-
toma.

5. Aristida floridana (Chapm.) Vasey, Grasses U.S. Descr. Cat. 35.
1885. [A. ternipes sensu Small, not Cav.] Erect perennials to about |
m tall, only sparingly branched below; panicles loose, 30-45 cm
long, the branches erect to ascending and narrow, or spreading and
open; glumes unequal to subequal, the first 10-14 mm long, the
second 8-9 mm long; lemma gradually narrowed to a single awn, the
midrib prominently thickened, 30-35 mm long from callus to awn
tip; central awn gradually curving, sickle-shaped; lateral awns
totally lacking or minute points only. Distribution: Roadsides, rail-
road embankments; FL. (Figure 6).

Aristida floridana is known in the United States only from Key
West and Ramrod Key of the Florida Keys, Monroe County. Beetle

1986] Allred — Aristida 379

(1983) also reported this species from Yucatan, Campeche, and
Quintana Roo, Mexico. It is apparently most closely related to A.
ternipes Cav., a species of the southwestern deserts to central
America.

6. Aristida gyrans Chapm., Bot. Gaz. 3:18. 1878. Tufted, erect per-
ennials, 20-65 cm tall; sheaths mostly shorter than the internodes,
but occasionally longer; blades filiform, involute; panicles narrow,
lax, 10-30 cm long; glumes mostly strongly unequal, the first 6-9
(11) mm long, the second 9-12 mm long, some plants with subequal
glumes; lemma 6-7 mm long, with an acute prominent callus 0.8-2
mm long; awns subequal, I-1.5 cm long, somewhat contorted at the
base. Distribution: Sandy pinelands, oak scrub; FL, GA. (Figure 7).

The relationships of Aristida gyrans are unclear. Even though the
glumes are usually strongly unequal, it does not appear to be related
to the group Purpureae, which are mostly plants of the arid south-
west. The callus, awns, and habit suggest an affinity to the A. purpu-
rescens complex.

7. Aristida lanosa Muhl. ex Ell., Bot. S.C. and Ga. 1:143. 1816. [A.
lanosa var. macera Fern.] Robust, tufted perennials to 1.5 m tall;
sheaths longer than the internodes, lanose or floccose, the pubes-
cence occasionally patchy and sparse; panicles narrow, elongate, (25)
35-70 (82) cm long, with tufts of woolly hair at the nodes of the
main axis; glumes subequal, the first 8.7-18 mm long, the second
8.4—-15 mm long; lemma 6.5-10 mm long; awns unequal, the central
12-28 mm long, the laterals 7-17 mm long. Distribution: Dry fields,
pine-oak woods, uplands; throughout the region except KY. (Figure
8).

Although several recent authors have suggested a close similarity
of Aristida lanosa and A. palustris, with some even merging the two
taxa, these two species are easily distinguished by both vegetative and
spikelet features (Allred, 1985b). Aristida palustris lacks lanose
sheaths and panicle nodes, and can be distinguished further from A.
lanosa by having sheaths shorter than the internodes, 2-nerved
glumes, longer calluses, and longer awns.

8. Aristida longespica Poir. in Lam., Encycl. Sup. 1:452. 1810.
Central awn mostly I-10 mm long; lateral awns mostly 0-5 mm

380 Rhodora [Vol. 88

PONS scuaky case wene eines epee des tenes 8a. var. longespica
Central awn mostly 12-27 mm long; lateral awns mostly 6-18 mm
PONS os eae a ok ey ena ee aweneeees 8b. var. geniculata

8a. A. longespica var. longespica. Much-branched annuals; pani-
cles narrow and spikelike; glumes mostly subequal, 2-8 mm long;
lemma 2.5-7 mm long; central awn erect to reflexed, 1-10 (14) mm
long; lateral awns erect to ascending, 0-5 (8) mm long. Distribution:
Disturbed ground, pine and oak woods; throughout the region.
(Figure 10).

8b. A. longespica var. geniculata (Raf.) Fernald. [A. intermedia
Scribn.] Much-branched annuals, frequently robust and appearing
perennial late in the growing season; panicles narrow and spikelike,
usually with many spikelets; glumes subequal, 4-11 mm long;
lemma 3.5—10 mm long; central awn erect to reflexed, (8) 12-27 mm
long; lateral awns erect to horizontal, (1) 6-18 mm long. Distribu-
tion: Disturbed ground, pine and oak woods; throughout the region
except KY and DE. (Figure 9).

The morphologic similarity of Aristida longespica and A. inter-
media was documented in an earlier paper (Allred, 1985a), justifying
the reduction of A. intermedia to varietal status. The variety genicu-
lata tends to be a more robust plant with larger spikelets and longer
awns. Specimens that are intermediate betwen the two varieties are
not uncommon; identification of these is difficult and often arbi-
trary. Nevertheless, because the two extremes are so strikingly dif-
ferent, it seems appropriate to maintain a distinction at the varietal
level. Both taxa frequently possess a peculiar banding pattern on the
lemma (as does A. purpurescens and rarely A. oligantha and A.
ramosissima), but it appears that the expression of this feature is
arbitrary and not correlated with any other feature.

9. Aristida mohrii Nash, N.Y. Bot. Gard. Bull. 1:436. 1900. Erect,
tufted perennials to 110 cm tall; blades of flowering shoots flat,
lightly pilose near the ligule; inflorescence 20-45 cm long, un-
branched, spicate, the spikelets on very short, stout pedicels on the
main axis; glumes subequal, 9-11 mm long; lemma 7-10 mm long;

1986] Allred — Aristida 381

awns subequal, |.3-2 cm long, contorted, spreading. Distribution:
Sandy pinelands and oak barrens; AL, FL, SC. (Figure 11).

The spicate inflorescences in Aristida mohrii and A. simpliciflora
have led to their frequent confusion with one another. The awns of
A. simpliciflora are distinctive, with the central about twice as
thick as the laterals, whereas in A. mohrii the awns are of equal
thickness. The two species may also be distinguished by the shorter
glumes, lemma, and callus, and less robust habit of A. simpliciflora.

10. Aristida oligantha Michx., Fl. Bor. Amer. 1:41. 1803. Much
branched annuals to 55 cm tall, similar to Aristida ramosissima;
spikelets often sessile or nearly so; first glume 3-7 nerved, 14-25 mm
long; second glume 18-31 mm long, with an awn 7-17 mm long
between two setae; lemma 14-20 mm long; central awn 3-7 cm long;
lateral awns 2.5-6.0 cm long. Distribution: Waste places, old fields;
throughout the region. (Figure 12).
See comments under Aristida ramosissima.

11. Aristida palustris (Chapm.) Vasey, Grasses U.S. Descr. Cat. 35.
1885. [A. affinis of American authors] Stout, tufted perennials to
1.5 m tall; sheaths mostly glabrous, occasionally pilose, mostly shor-
ter than the internodes; panicle narrow, stiff, to 55 cm long; glumes
subequal, (7.5) 9-13.5 mm long, the first prominently 2-nerved and
2-keeled; lemma 6-9 mm long; awns unequal, the central 1.5-4 cm
long, the laterals 0.8-3.5 cm long. Distribution: Moist ground,
marsh and pond shores, lowlands; AL, FL, GA, LA, MS, NC, SC.
(Figure 13).

The correct name for this species is not Aristida affinis (Schultes)
Kunth, as in commonly used; A. affinis is the same as A. purpures-
cens and placed in synonymy there (Allred, 1985b). See comments
under A. /anosa regarding the relationship of A. palustris and A.
lanosa.

12. Aristida patula Chapm. ex Nash, Bull. Torr. Bot. Club 23:98.
1896. Erect perennials with short rhizomes, to | m tall; panicles
30-50 cm long, loose, open, the branches stiffly spreading to droop-
ing at maturity; spikelets appressed at the ends of the branches;
glumes slightly unequal, 10-13 mm long, the first slightly longer
than the second; lemma 10-12 (15) mm long to base of awns; central

382 Rhodora [Vol. 88

awn straight, 20-25 mm long; lateral awns 3-10 mm long. Distribu-
tion: Roadsides, sandy fields, low pinelands; FL. (Figure 14).

13. Aristida purpurea Nutt., Trans. Amer. Phil. Soc. 5:145. 1837.
Summit of lemma 0.3-0.8 mm broad, not gradually narrowed;
awns usually stout, 4-10 cm long; second glume 14-25 mm long
Pee ere rey reer rere ere er ere er 13b. var. longiseta
Summit of lemma 0.1-0.3 mm broad, gradually narrowed; awns
slender, mostly 2-6 mm long; second glume mostly shorter
Lee 10 WE dye saeco caaeneraaennes 13a. var. purpurea

13a. A. purpurea var. purpurea. Tufted perennials 25-60 cm tall;
blades involute; panicles loose, the branches usually somewhat
drooping; glumes unequal, the first about 1/2 as long as the second;
lemma 6-12 mm long, narrow at the summit; awns subequal, (1.5)
2-6 cm long. Distribution: Waste places, roadsides, sandy prairies;
AR, LA. (Figure 16).

13b. A. purpurea var. longiseta (Steud.) Vasey, in Rothr., Rpt. U.S.
Survey W. 100th Merid. 6:286. 1878. [A. longiseta Steud.] Tufted
perennials 12~50 cm tall; blades involute; panicle branches erect and
stout to delicate and drooping; glumes unequal, the first about 1/2
as long as the second; lemma 10-16 mm long, broad at the summit;
awns subequal, 4-10 cm long. Distribution: Waste places, roadsides;
LA. (Figure 15).

A common grass of the plains and deserts of the western United
States, Aristida purpurea is known only from a few localities in
Arkansas and Louisiana. Our plants are best treated at the varietal,
rather than the specific level (Allred, 1984b). The variety /ongiseta
currently is known only from one collection in St. Tammany Parish,
Louisiana, and one from Kershaw County, South Carolina. The
record from South Carolina appears to be either an introduction or
perhaps an error on the label. Variety /ongiseta was also collected
from the Soil Conservation Service Nursery in Chapel Hill, North
Carolina.

14. Aristida purpurescens Poir. in Lam., Encycl. Sup. 1:452. 1810.
Central awn divaricate to reflexed, about twice as thick at the base

1986] Allred — Aristida 383

as the erect lateral awns................... l4c. var. virgata
Central and lateral awns spreading, straight to spirally contorted
but not reflexed, and all about the same thickness at the base.
First glume mostly longer than the second; awns straight or
slightly contorted at the base; blades 1-3 mm wide, often
CUPS $3655 chasis acetates 14a. var. purpurescens
First glume shorter than or equal to the second; awns spirally
contorted at the base; blades about | mm wide, usually not
COME 6oc ik soacene Hearne cianees 14b. var. tenuispica

14a. A. purpurescens var. purpurescens. [A. affinis (Schulte)
Kunth] Tufted perennials to | m tall; lower sheaths longer than the
internodes, mostly glabrous, occasionally pilose with appressed
hairs; blades mostly flat, often curling; panicles narrow, densely to
loosely flowered; glumes usually unequal, the first longer than or
equal to the second, 6-10 mm long, the second 5-8 mm long; lemma
6-8 mm long; awns 1.5-2.5 cm long, about equal in thickness and
angle, usually ascending, the laterals sometimes shorter than the
central. Distribution: Waste places, pine savannahs, fields; through-
out the region. (Figure 17).

14b. A. purpurescens var. tenuispica (Hitchc.) Allred, comb. nov.
Basionym: Aristida tenuispica Hitchcock, Contr. U.S. Natl. Herb.
22:581. 1924. Tufted perennial to | m tall; sheaths longer or shorter
than the internodes, glabrous; blades flat to loosely rolled, mostly
not curling; panicles narrow, loosely flowered; glumes subequal, 8-9
mm long, the first equal to or slightly shorter than the second and
1-2 nerved; lemma 6-7 mm long; awns 1.2-1.8 cm long, subequal in
thickness, angle, and length, usually spirally contorted at the base.
Distribution: Low pine and oak woods, prairies, roadsides; AL, FL,
GA, MS, NC, SC. (Figure 18).

l4c. A. purpurescens var. virgata (Trin.) Allred, comb. nov. Basio-
nym: Aristida virgata Trinius in Sprengel, Neu. Entd. 2:60. 1821.
Tufted perennial to | m tall; sheaths glabrous, longer or shorter
than the internodes; blades flat, curling or not; panicles narrow,
densely to loosely flowered; glumes subequal or the first slightly
longer, 6-7 mm long; lemma 4-7 mm long; central awn reflexed and
about twice as thick at the erect lateral awns, the central 1.3-2 cm

384 Rhodora [Vol. 88

long, the laterals 0.8-1.3 cm long. Distribution: Pinelands, oak
woods, bogs, waste places, prairies; AL, AR, FL, GA, LA, MS, NC,
SC, TN, VA. (Figure 19).

There is significant morphological intergradation among the
three entities purpurescens, tenuispica, and virgata, with spikelets
corresponding to more than one entity sometimes present within the
same panicle (Allred, 1984a). The lack of a strongly correlated set of
features that distinguishes the three taxa prompts their relegation to
varieties of a single species. Further studies of chromosome number,
breeding behavior, and habitat are needed.

15. Aristida ramosissima Engelm. ex Gray, Man. Bot. North. U.S.
ed. 2. 550. 1856. [A. ramosissima var. uniaristata A. Gray; A. ramo-
sissima var. chaseana Henr.] Much-branched annuals to 60 cm tall,
similar to Aristida oligantha; spikelets sessile or nearly so; first
glume 3-7 nerved, 11-20 mm long; second glume 16-25 mm long,
with a short awn 3-7 mm long between two seta; lemmas (8) 14-22
mm long; central awn 9-28 mm long; lateral awns 0-12 (25) mm
long. Distribution: Pine savannahs, roadsides, fields; AR, KY, LA,
TN. (Figure 20).

The variety chaseana, with nearly equal awns, is transitional
between this species and Aristida oligantha, and perhaps deserves
formal recognition. It is usually identified as A. oligantha. The var-
iety uniaristata, with obsolete lateral awns, is somewhat anomalous
in the section Chaetaria of Aristida, this feature being typical of the
section Streptachne. Further biosystematic studies of A. oligantha
and A. ramosissima are needed.

16. Aristida rhizomophora Swallen, J. Wash. Acad. Sci. 19:196.
1929. Perennials to 100 cm tall, from vigorous rhizomes; old basal
sheaths fibrous and shredding into threadlike segments; panicles
20-45 cm long with erect to somewhat flexuous branches; glumes
unequal, the first 6-12 mm long including the awn, the second 13-18
mm long including the awn; lemmas 9-13 mm long; awns often
arcuate-recurved, the central 15-30 mm long, the laterals 13-20 mm
long. Distribution: Sandy pinelands and prairies; FL. (Figure 21).

1986] Allred — Aristida 385

17. Aristida simpliciflora Chapm., Bot. Gaz. 3:18. 1878. Erect,
tufted perennials to 80 cm tall; sheaths shorter than the internodes;
inflorescence slender, racemose, usually with 2-3 spikelets per node,
one short-pedicelled, one long-pedicelled; glumes subequal, 6-8 mm
long; lemma 5-6 mm long; central awn strongly reflexed, about
twice as thick as the laterals, 1-1.5 cm long; iateral awns spreading
to divaricate, 1-1.5 cm long. Distribution: Low dense pine barrens
and woods; AL, FL, MS. (Figure 22).

Aristida simpliciflora shows strongest morphologic affinities with
A. mohrii and A. purpurescens var. virgata. Awn and panicle fea-
tures successfully distinguish this species.

18. Aristida spiciformis Ell., Bot. S.C. and Ga. 1:141. 1816. Erect
perennials to | m tall, the culms little-branched; panicles (5) 8-24 cm
long, dense and compact, spiciform, with numerous congested spike-
lets; glumes unequal, the first with a body about 4 mm long and
abruptly awned with a 4-12 mm long awn, the second 7-10 mm long
and narrowed to an awn 10-12 mm long; lemma 5-6 mm long and
terminated by a twisted beak 7-30 mm long; awns mostly subequal,
(1) 2-3 cm long, the lateral awns thinner than the central. Distribu-
tion: Low pine savannahs, oak woods; AL, FL, GA, MS, NC, SC.
(Figure 23).

The dense panicle, awned glumes, and twisted lemma beak are
characteristic of the species. Although not developing a zone of
articulation and not considered a member of section Arthratherum,
the lemma beak frequently breaks from the lemma body, presuma-
bly from a weakening of the thin lemma tissue above the caryopsis.

19. Aristida stricta Michx., Fl. Bor. Amer. 1:41. 1803. Tufted, erect
perennials, occasionally with short rhizomes, 45-100 cm tall; blades
tightly involute, firm, pilose on the lower (outer) surface near the
midrib especially toward the base, also at the collar and throat
regions; panicles narrow, 18-35 cm long; glumes about equal, 7-12
mm long; lemma 6-8 mm long; central awn 10-15 mm long; lateral
awns 10-13 mm long. Distribution: Pinelands, sandy fields; AL, FL,
GA, MS, NC, SC. (Figure 24).

Herbarium specimens of this species almost always show charred
plant bases.

386 Rhodora [Vol. 88

20. Aristida tuberculosa Nutt., Gen. Pl. 1:57. 1818. Branching
annuals to | m tall; panicles 10-20 cm long, loose with somewhat
stiff, ascending branches; spikelets erect, yellowish-brown; glumes
subequal, 20-30 mm long, the second slightly longer than the first,
gradually acuminate and awned; lemma 10-14 mm long from callus
to point of articulation with the column; awn column 8-15 mm
long, breaking at maturity; awns subequal, 3-4 cm long. Distribu-
tion: Sandy fields, pinelands, disturbed areas; AL, FL, GA, MD,
MS, NC, SC, VA. (Figure 25).

The spikelets (glumes, lemma, awns) in Aristida tuberculosa are
generally larger than in A. desmantha. In both, the awn column is
formed by the connivent awn bases, frequently twisted in A. tuber-
culosa, and not formed by the narrowing of the lemma apex.

ACKNOWLEDGMENTS

I] thank the curators of the following herbaria for their generous
loans of specimens: APSC, ARK, AUA, DUKE, FLAS, FSU, KY,
LAF, LSU, MARY, MO, NCSC, NLU, NO, NSU, PH, SMU,
TENN, UNA, URV, US, USCH, USF, VDB, VPI, WVA. I also
thank an anonymous reviewer for several helpful suggestions.

LITERATURE CITED

ALLRED, K. W. 1984a. Studies in the Aristida (Gramineae) of the southeastern
United States. 1. Spikelet variation in A. purpurescens, A. tenuispica, and A.
virgata. Rhodora 86: 73-77.

1984b. Morphologic variation and classification of the North American

Aristida purpurea complex (Gramineae). Brittonia 36(4): 382-395.

1985a. Studies in the Aristida (Gramineae) of the southeastern United
States. Il. Morphometric analysis of A. intermedia and A. longespica. Rhodora
87: 137-145.

1985b. Studies in the Aristida (Gramineae) of the southeastern United
States. III]. Nomenclature and a taxonomic comparison of A. /anosa and A.
palustris. Rhodora 87: 147-155.

BEETLE, A. A. 1983. Las Gramineas de México. Vol. I. Editorial Calypso, Mex-
ico City.

GLeAsON, H. A. AND A. Cronguist. 1963. Manual of the Vascular Plants of
Northeastern United States and Canada. Van Nostrand Reinhold Co., New
York.

1986] Allred — Aristida 387

HENRARD, J.T. 1929. A monograph of the genus Aristida. 1. Meded. Rijks-Herb.
Leinden No. 58.

1932. A monograph of the genus Aristida. Il. Meded. Rijks-Herb.
Leiden No. 58A.

Hircucock, A. S. 1924. The North American species of Aristida. Contr. U.S.
Natl. Herb. 22: 517-586.

AND A. CHASE. 1951. Manual of the Grasses of the United States. Ed. 2.

USDA Misc. Publ. 200.

DEPARTMENT OF ANIMAL AND RANGE SCIENCES
BOX 3 - 1, NEW MEXICO STATE UNIVERSITY
LAS CRUCES, NM 88003

NOMENCLATURAL NOTES ON HEDYOTIS ROSEA
RAFINESQUE AND A NEW COMBINATION
IN HOUSTONIA!

EDWARD E. TERRELL

ABSTRACT

The nomenclatural application of the name Hedyotis rosea Rafinesque (1817) is
discussed. It is concluded that the name applies to the species also known as Housto-
nia pygmaea C. H. & M. T. Muller (1936). The new combination, Houstonia rosea
(Rafinesque) Terrell, is proposed and a neotype is designated.

Key Words: Hedyotis, Houstonia, Rubiaceae, nomenclature

Hedyotis rosea Rafjnesque (1817) and Houstonia pygmaea C. H.
and M. T. Muller (1936) are two competing names for a diminutive
early spring-flowering rubiaceous annual that occurs in the south-
eastern and southcentral United States. The protologue of Hous-
tonia pygmaea describes the species quite adequately, as it includes
a detailed description in English with a rather full Latin diagnosis,
comments on habitat, distinguishing characters, and citations of
type specimens. The name Hedyotis rosea Raf. was first adopted by
Waterfall (1953) and later used by Lewis (1970) in his treatment of
Hedyotis for the “Manual of Vascular Plants of Texas.” The proto-
logue is very brief and application of the name tenuous. The follow-
ing discussion takes up in detail the question of whether this name
referred to the same species as Houstonia pygmaea.

Rafinesque (1817) described Hedyotis crassifolia (Houstonia
pusilla) and Hedyotis rosea as follows:

“243. Hedyotis crassifolia Raf. Ramis tenuis divaricatis sulcatis
unifloris, foliis sessilibus oblongis acutis glabris integris carnosis,
floribus longe pedunculatis. Raf.—Anonyme I. Rob. p. 454. This
plant although resembling very much the Houstonia, is a real Hedy-
otis having a capsul two celled and polysperme. It blossoms in
February, it varies with white, deep violet and pale violet flowers,
stem only two inches high, tube of the corolla filiform, four stamina

'Scientific Article No. A-4339, Contribution No. 7328 of the Maryland Agricultural
Experiment Station.

389

390 Rhodora [Vol. 88

in the tube nearly sessile, style short, stigma oblong, capsul heart
shaped, with many minute seeds.

“244. Hedyotis? rosea Raf. Repens, floribus roseis—Anonyme
2. Rob. p. 454. This may be the Houstonia tenella of Lyon and
Pursh; Robin does not describe it, but he merely says it is still
smaller than the foregoing, creeping, and with flowers of a pale rose
colour.”

Rafinesque’s “Florula Ludoviciana” (1817) was a translation
from the French of “Flore Louisianaise,” the third volume of an
account by Claude C. Robin (1807) of a journey in western Florida,
Louisiana, and parts of the West Indies. Rafinesque’s florula was
more than a translation, however, but a new work, providing
binomials for the taxa lacking binomials in Robin’s book (Merrill,
1949: Ewan, 1967).

It is desirable to find out whether Robin’s original description,
from which Rafinesque’s protologue was taken, provides additional
information about the identity of Hedyotis rosea. Robin described
the plants in French, adding a Latin genus name or approximation
to a genus name. For the plants in question Robin described three
unnamed species under the general heading “Garance (rubia).” The
first species was in 1817 named Rubia techensis Raf., possibly a
species of Galium (Merrill, 1949). The description of the second
species was faithfully translated by Rafinesque, who named it
Hedyotis crassifolia. The third species, H. rosea Raf., was described
as follows:

“Une autre plus petite, se trainant et tragant davantage, porte des
fleurs rose tendre: elles pourraient, au printemps, émailler agréa-
blement des compartimens de parterres ou de gazons.

“Ces espéces paraissent se rapprocher de l’o/den/andia, si ce n’est
que le tube de la corolle est trés-alongé, tandis que l’ol/denlandia
est a peine tubulé.”

In their descriptions, Robin and Rafinesque provided four char-
acteristics for Hedyotis rosea: (1) plants smaller than those of H.
crassifolia, (2) flowers pale rose, (3) corolla tubes longer than those
of Oldenlandia species, and (4) plants creeping. It is necessary not
only to determine what species best fits this brief description, but
also to know the identity and characteristics of Hedyotis crassifolia,
as Hedyotis rosea was directly compared to it.

1986] Terrell — Hedyotis & Houstonia 39]

Hedyotis crassifolia Raf. was adequately described and has been
generally accepted (e.g., Lewis, 1970) as referring to the species also
known as Houstonia pusilla Schoepf or H. patens Elliott, the com-
mon “small bluets” of the southeastern states, ranging from north-
ern Maryland (where adventive) to northwest Florida, west to
Texas, and north to Illinois. It has the following characteristics: (1)
plants 1.5-20 (usually 5-15) cm. tall, (2) corollas usually purple with
reddish centers, but varying to violet, lavender, pink, or white with
yellow centers (various color forms sometimes occur in a single
population), (3) corolla tubes 2~5 mm long, and (4) plants erect or
spreading.

The characteristics of Houstonia pygmaea (=Hedyotis rosea?)
are: (1) plants 1-4 (—5) cm. tall), (2) corollas varying from white to
pale pink or pale rose, (3) corolla tubes 3.5-6 mm long, and, (4)
plants erect, spreading, or sprawling. It is apparent that three of the
four described characteristics of Hedyotis rosea apply to Houstonia
pygmaea: the latter is smaller than Hedyotis crassifolia; it usually
has pale rose or pale pink flowers (along with some white flowers);
and it and H. crassifolia have elongate corolla tubes compared to
the rotate or short-tubular corollas of Oldenlandia boscii (DC.)
Chapman, O. uniflora L., and O. corymbosa L., the three common-
est Oldenlandia species present in the Gulf Coastal Plain. The
fourth characteristic, however, does not agree. In reference to the
creeping habit, Rafinesque speculated that Hedyotis rosea could be
the same as Houstonia tenella Pursh, but the latter is now known to
be a synonym of the very different species, H. serpyilifolia, centered
in the southern Appalachians. To find out exactly what Robin may
have meant by “se trainant et tragant davantage,” I requested a
translation from Smithsonian botanist Dr. Marie-Héléne Sachet.
She translated it literally as “‘dragging itself? and creeping,” and
commented that “What is meant is that the plant is completely
prostrate, with stems running along the ground, like stolons.”

Are there any creeping taxa in the southeastern states that have
the characteristics of Hedyotis rosea? Two creeping rubiaceous spe-
cies exist there. Houstonia procumbens is creeping, but almost in-
variably has white flowers and is not necessarily smaller than H.
pusilla ( Hedyotis crassifolia). \t usually has broadly ovate or subor-
bicular leaves, in considerable contrast to Hedyotis crassifolia. It

392 Rhodora [Vol. 88

occurs in southeastern Louisiana east to Florida, Georgia, and
South Carolina. Another species, Oldenlandia salzmannii, is an
accidental introduction from South America; it was recently discov-
ered near Pensacola, Florida, and in adjacent Alabama (Fosberg
and Terrell, 1985). It is creeping and has pink or light purple flow-
ers. Its corolla tubes are longer than those of most Oldenlandia
species, but relatively shorter and distinctly wider than those of the
houstonias. It is known that Robin visited Pensacola, but unlikely
that this supposedly recent introduction was present there at that
tyme.

Could Hedyotis rosea have referred to a pink variant of H. crassi-
folia? This conclusion is doubtful as pink flowers are scarce in H.
crassifolia and when present would often be accompanied by other
color variants in the same population. Furthermore, plants of H.
crassifolia are commonly larger, as noted previously.

That Hedyotis rosea was based on a species not in the Rubiaceae
seems unlikely because of Robin’s supplemental description (omit-
ted by Rafinesque) regarding the occurrence of elongated corolla
tubes in both H. crassifolia and H. rosea. This omission seems to
point to a close relationship between these two species and excludes
from consideration many non-rubiaceous herbs in the Louisiana
flora. | asked Dr. R. Dale Thomas, who has collected extensively in
Louisiana, whether he knew of any other herb besides Houstonia
pygmaea that fits the description of Hedyotis rosea, and his reply
was negative. To this I would add that the combination of pale rose
tubular corollas on consistently very small plants is unusual.

Neither Robin nor Rafinesque cited specimens or locations for
Hedyotis crassifolia and H. rosea. For Rubia techensis Raf. the
name Teche was mentioned, referring to the bayou Teche in south-
ern Louisiana. Mentioned by Robin and Rafinesque are the Oua-
chita region, the Red River (flows north to south through
Louisiana), and New Orleans. It is also known that Robin visited
the Opelousas area in southern Louisiana, and Natchez, Mississippi
(J. Ewan, pers. comm.). At present, Houstonia pusilla (Hedyotis
crassifolia) is very common and widespread in Louisiana. Housto-
nia pygmaea is less common, but occurs in at least 24 parishes
throughout the state except the extreme south. It occurs in St.
Landry Parish, which includes the city of Opelousas. Accordingly, it

1986] Terrell — Hedyotis & Houstonia 393

seems likely that Robin could have come across plants of this
species.

Waterfall (1953), in taking up Hedyotis rosea, discussed in some
detail Rafinesque’s description of Hedyotis rosea, but he did not
refer back to Robin’s work. He emphasized that the rose color of the
flowers is striking and distinctive, and also stated that the species “‘is
not repent with us (it appears so in some well developed speci-
mens).” Waterfall went on to state his opinion that despite the brief
protologue, Rafinesque “did furnish it with a description that seems
to characterize it. Can it, then, be rejected as a nomen dubium?”

In summary, three out of four characteristics of Hedyotis rosea
agree with those of Houstonia pygmaea. Furthermore, the latter
species occurs over most of Louisiana and certainly in some of the
areas traversed by Robin. The only discordant item in the descrip-
tion of Hedyotis rosea is the creeping characteristic which translated
literally suggests a stoloniferous or trailing species. Should we reject
the name Hedyotis rosea as a nomen dubium on the grounds that
Robin meant quite literally creeping and was, therefore, referring to
some other species than Houstonia pygmaea? Is it easier to make a
mistake regarding flower color than to err as to creeping?

The weight of the evidence given above leads me to accept the
name Hedyotis rosea as the same as Houstonia pygmaea. It is possi-
ble that Robin saw massed plants of the tiny annual and supposed
without close observation that they were creeping, or that he made
some kind of observational error. Also, his brief description in con-
trast to that of Hedyotis crassifolia suggests that he may not have
been very familiar with a species so casually described. The accep-
tance of the name also agrees with usage established by Lewis (1970)
in the manual of the Texas flora.

Rafinesque (1817) based his descriptions solely on those of Robin
(1807), as there are no extant specimens of Robin’s (see Stafleu and
Cowan, 1983); therefore, it is advisable to designate a neotype. A
neotype was selected by me from a collection widely distributed to a
number of herbaria: H. E. Moore, Jr. 1075 from Texas, distributed
as Plantae Exsiccatae Grayanae 1479 under the name Houstonia
Pygmaea (Moore originally labelled it H. croftiae Britton & Rusby).
It so happens that the neotype (GH) shows sprawling plants (Figure
1); could such plants as these have led Robin to use the word
creeping?

394 Rhodora [Vol. 88

AG
Past . PLANTAE EXSICCATAE GRAYANAE

1479. Houstonia pygmaea C,H. & M. T. Moeiien

In Bull. Torr, Bot, Cl. lxiti, $3 (1986); Reeves & Bain,
Fi. 8. Centr. Texas, 231 (1946).

Syn, Houstonia patens var. pusilla Gray, Syn. Fi. (2).
25 (1884), Howsionia minor var. pusilla (Gray) Small, Fl.
Se. U.S. 1107, 1888 (1908), Hedyotis Taylorae Fosberg
in Shinners in Field & Lab, xvii (4), 109 (1969),

Low winter annual; very infrequent, usually occarring with
Houstonia patens Eli., but readil Toenthas ty fe lower,
spreading habit, short- late, delicate pink flowers, the flow-
ers slightly fragrant and with « ring of yellow hairs at top of
corolla-tube; capsule laterally compressed, the length nearly four
times the width,

Neglected field, Amigo, Smith County, Texas.

Coll. H. E. Moon, Ja. (no, 1075) March 3-6, 1946

PLANTS OF
jtene

PE Soa Leas Butt Qual

Figure |. Neotype of Houstonia rosea (Raf.) Terrell.

Generic delimitations among Hedyotis, Houstonia, and Olden-
landia were discussed by Terrell (1975) and are under further study.
It is questionable whether any of the North American species
referred to Houstonia or Hedyotis are closely related to Hedyotis

1986] Terrell — Hedvyotis & Houstonia 395

fruticosa L., the type species, a native of Sri Lanka. Although taxo-
nomic relationships of Houstonia rosea within Houstonia are
ambiguous, it seems best to relate the species tentatively to other
early spring-flowering bluets with similar habits. These true bluets
are in the H. caerulea group, including H. caerulea L. (the type
species); H. serpyllifolia Michaux [ Hedyotis michauxiana Fosberg];
Houstonia pusilla Schoepf [H. patens Elliott; Hedyotis crassifolia
Raf.; Houstonia minima Beck]; H. micrantha (Shinners) Terrell
[Hedyotis australis W. H. Lewis & D. M. Moore]; and H. pro-
cumbens (Walter ex J. F. Gmelin) Standley [ Hedyotis p. (Walter ex
J. F. Gmelin) Fosberg]. The base chromosome number for this
group is x = 8, except fot H. procumbens with n = 7 (Lewis, 1962).
Houstonia rosea has n = 7 (Lewis, 1962), but is not very closely
related to H. procumbens, judging by reproductive morphology.
Pollen of H. rosea is similar to but smaller than that of other
members of the H. caerulea group (Lewis, 1965). Seeds of H. rosea
are more or less cymbiform with an elongate hilar ridge in a ventral
depression, thus they resemble seeds of the H. rubra Cav. and H.
purpurea L. groups rather than the subglobose seeds of H. caerulea
and its close relatives. Although seed morphology is very important
in Houstonia and allied genera, it is.difficult in this instance to
ignore the chromosome number, which agrees with the H. caerulea
group rather than with the H. rubra group (x = 11) and H. purpurea
group (x = 6).

The new combination and synonymy are given below. In the
absence of a type specimen for Houstonia rosea, a neotype is desig-
nated, following ICBN, Art. 7.4, 7.8, and T5 (Voss et al., 1983).

Houstonia rosea (Rafinesque) Terrell, comb. nov.

Hedyotis rosea Rafinesque, Florula Ludoviciana 77. 1817. NEo-
TYPE, here designated: Texas. Smith Co.: neglected field, Amigo,
3-6 March 1946, Moore 1075, distributed as Plantae Exsiccatae
Grayanae 1479 (as Houstonia pygmaea),; NEOTYPE GH!; ISONEO-
TYPES DUKE!, MO!, NCSC!, NO!, NY!, US!, and others.

Houstonia patens Elliot var. pusilla A. Gray, Syn. Fl. N. Am. 1, 2: 25. 1884.
LectotyPe: Louisiana. Rapides Parish: Alexandria, Hale s.n. (GH!) (lectotype
designated by W. H. Lewis, Ann. Missouri Bot. Gard. 55: 31-33. 1968).

Houstonia pygmaea C. H. & M. T. Muller, Bull. Torrey Bot. Club 63: 33-34. 1936,
non Hedyotis pygmaea Roemer & Schultes, Syst. Veg. 3: 526. 1818. (= Houstonia
wrightii A. Gray). SYNTYPES: Texas. Dewitt Co.: 16 Feb and 6 Mar 1934, Muller 3

396 Rhodora [Vol. 88

and 4, specimens originally deposited in Muller herb. at Cuero, Texas, and “cotypes”
at NY, but present locations unknown, as NY now has no types of this species.
Another specimen by same collectors, but not cited as a type: Texas. Dewitt Co.:
Mission Valley, 11 Feb 1934, Muller / (GH!).

Hedyotis taylorae Fosberg, in Shinners, Field and Lab. 17: 169. 1949.

Hedyotis minima (Beck) Torrey & Gray f. albiflora Lathrop, Rhodora 59: 95.
1957. HoLotype: Kansas. Woodson Co.: Sec 32, T25S, RISE, Lathrop and McGre-
gor 35 (KANU) (fide W. H. Lewis, Ann. Missouri Bot. Gard. 55: 31-33. 1968). (Note:
Hedyotis minima f. minima (Houstonia minima Beck) = Houstonia pusilla Schoepf
sens. lat.).

ACKNOWLEDGMENTS

I thank Drs. F. R. Fosberg, J. H. Kirkbride, Jr., W. H. Lewis,
Harold Robinson, and Marie-Héléne Sachet for manuscript
reviews and valuable nomenclatural advice; they are not to be held
responsible for the opinions expressed here. Dr. Sachet also pro-
vided a translation as cited in the text. Helpful data were contrib-
uted by Michael Canoso, and Drs. Joseph Ewan, Patricia Holmgren,
and R. Dale Thomas.

LITERATURE CITED

Ewan J. 1967. Introduction, pp. i-xl, for Rafinesque, C. S. 1817. Florula Ludo-
viciana. Reprint by Hafner Publ. Co., New York.

FosBerG, F. R. AND E. E. TERRELL. 1985. A recently established exotic in west
Florida and Alabama (Hedyotis salzmannii or Oldenlandia salzmannii;, Rubia-
ceae). Castanea 50: 49-51.

Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North Amer-
ica. Amer. J. Bot. 49: 855-865.

1965. Pollen morphology and evolution in Hedyotis subgenus Edrisia
(Rubiaceae). Amer. J. Bot. 52: 257-264.

1970. Hedyotis, pp. 1487-1490. Jn: Correll, D. S. and M. C. Johnston,
Eds. Manual of the vascular plants of Texas. Texas Research Foundation,
Renner, Texas.

MERRILL, E. D. 1949. Index Rafinesquianus. Arnold Arboretum of Harvard
University, Jamaica Plain, Mass.

MuLter, C. H. AND M. T. 1936. A new Houstonia in southcentral Texas. Bull.
Torrey Bot. Club 63: 33-34.

RAFINESQUE-SCHMALTZ, C. S. 1817. Florula Ludoviciana. C. Wiley & Co., New
York. 178 pages.

Rosin, C.C. 1807. Voyages dans l’interieur de la Louisiane, de la Floride occid-
entale, et dans les isles de la Martinique et de Saint Domingue, pendant les
années 1802-1806. 3 vols. Vol. 3. Flore Louisianaise. F. Buisson, Paris. Vol. 3,
551 pages.

1986] Terrell — Hedyotis & Houstonia 397

STAFLEU, F. A. AND R. S. Cowan. 1983. Taxonomic literature. Vol. 4. Second
edition. Regnum Veg. 110.

TERRELL, E.E. 1975. Relationships of Hedyotis fruticosa L. to Houstonia L. and
Oldenlandia L. Phytologia 31: 418-424.

Voss, E.G. etal. 1983. International Code of Botanical Nomenclature. Regnum
Veg. III.

WATERFALL, U. T. 1953. The identity of Hedyotis rosea Raf. Rhodora 55:
201-203.

DEPARTMENT OF BOTANY
UNIVERSITY OF MARYLAND
COLLEGE PARK, MARYLAND 20742

TAXONOMY OF CAREX SECTION FOLLICULATAE
USING ACHENE MORPHOLOGY

DANIEL E. WUJEK AND FRANCIS J. MENAPACE

ABSTRACT

Scanning electron microscopy was used to examine the surfaces of achenes of
Carex section Folliculatae and the allied section Collinsiae for systematic relation-
ships. Epidermal patterns were distinctive and consistent for each taxon. Variation of
epidermal cells was most evident with respect to size of cell, the number, thickness,
and sinuosity of anticlinal walls per cell, and shape of the apex of silica bodies present
in two of the taxa. Recognition of Carex lonchocarpa is supported.

Key Words: Carex, systematics, achene, Cyperaceae, SEM, EDS

INTRODUCTION

Carex folliculata is one of three taxa traditionally placed within
the section Folliculatae. Mackenzie (1935) recognized three distinct
species (C. folliculata L., C. michauxiana Boeckl., and C. loncho-
carpa Willd.). Other authors (e.g., Small, 1933; Fernald, 1950; Glea-
son, 1952; Gleason & Cronquist, 1963; Radford, Ahles & Bell, 1968;
Correll & Correll, 1975; Godfrey & Wooten, 1979) also treat this
group in their floras as being composed of three taxa (Table 1), but
recognize C. folliculata var. australis Bailey in synonymy of C.
lonchocarpa.

The members of this group have not been critically reviewed since
their original descriptions. This study was initiated to investigate the
Carex folliculata aggregate and the closely allied section Collinsiae
in eastern North America. Scanning electron microscopy (SEM) of
achene characteristics was utilized in the study. Our findings (Tal-
lent and Wujek, 1983a, 1983b; Menapace and Wujek, 1985) confirm
those of earlier studies by Walter (1975), Toivonen and Timonen
(1976) and Standley (Ph.D. thesis, Univ. of Washington, Seattle,
1981; 1985) that achene epidermal characteristics provide criteria
useful for delimitation of taxa at various levels.

MATERIALS AND METHODS

Perigynia containing mature achenes of all species and varieties of
Carex sections Folliculatae and Collinsiae were taken from herba-
rium specimens deposited at the University of Michigan. Collection

399

400 Rhodora [Vol. 88

Table |. Collection data for Carex species used for SEM observations.

Section Folliculatae
C. folliculata L.
Ohio: Bissell 228; Indiana: Lyon (s.n.); Mass.: Fogg 443; Canada (Ont.):
Brunton 4289
C. lonchocarpa Mackenzie
North Carolina: Ahles 45925; Mississippi: Tracy 4830
C. michauxiana Boeck].
Vermont: Grout 662, Ball 1426; Canada (Que.): Roy 535; (Ont.): Grassel
4374, Meyer 1/321

Section Collinsiae
C. collinsti Nutt.
New Jersey: Hermann 3213, Canby (s.n.); Georgia: Ashe 287

data for SEM studies are given in Table 1. Achenes were dissected
out of the perigynia and subjected to a cleaning procedure of sonifi-
cation, followed by soaking for 18-36 hrs. in a 1:9 concentrated
sulfuric acid: acetic anhydride solution. This latter step, modified
from Walter (1975), is essential to remove the outer walls of epider-
mal cells for examination of silicon ornamentations. Achenes were
then rinsed in distilled water and desiccated prior to sputter coating
with 30 nm of gold. Two or three achenes of each plant were exam-
ined by an AMR 1200 SEM and photographed using Polaroid P/N
665 film.

Qualitative x-ray spectra of the epidermal cells were obtained
with an energy-dispersive spectrometer (EDS; Ortec 5000) attached
to a Hitachi S 520 SEM. Beam accelerating voltage was maintained
at IS kV.

OBSERVATIONS AND DISCUSSION

Features of the achene surfaces are constant within species, even
in herbarium specimens up to 50 years old. Among the species,
achene epidermal cells ranged from apparent lack of silica bodies
[ Carex folliculata (Figure 1), C. michauxiana (Figure 2)] through
the presence of central silica bodies [C. /onchocarpa (Figure 3), C.
collinsii (Figure 4) ].

A typical achene for these sections has epidermal cells with four
to eight anticlinal (radial) walls that are generally sinuous, but may
be straight in some taxa (Carex folliculata, C. collinsii). A single or
double hemispherical body (Figures 3, 4) arising from the inner

1986] Wujek & Menapace — Carex 401

fe!
| ia,
-

Figures 1-4. SEM photographs of Carex achenes. 1. C. folliculata. 2. C.
michauxiana. 3. C. lonchocarpa. 4. C. collinsii. Scale bar = 40 um

periclinal wall of the epidermal cells is smooth apically and not
nodulose as has been reported in other Carex taxa (Menapace, M.S.
thesis, Central Mich. U., Mt. Pleasant, 1985). X-ray spectrometry
confirmed that the major element in the hemisphere body is silicon,
of which the chemical form in these plants is silica.

402 Rhodora [Vol. 88

In Carex folliculata (Figure 1), each achene epidermal cell is
nearly as wide as long, and generally has straight anticlinal walls
and a smooth platform that lacks a central body.

The epidermal pattern of Carex michauxiana (Figure 2) is most
similar to that of C. folliculata in terms of overall size and morphol-
ogy of cells. The silica platform does not, however, extend to the
anticlinal wall but possesses a rounded margin producing a narrow
wing toward the base of the achene.

Each epidermal cell of Carex lonchocarpa (Figure 3) has sinuous
anticlinal walls and one, occasionally two, central bodies. Platforms
are flat except for being concave at the base of each central body.

The achene epidermal cells of Carex collinsii (Figure 4) usually
have five straight anticlinal walls. Each platform possesses a single
central body with a buttress which is less concave than that observed
in C. lonchocarpa.

The epidermal patterns of the achene surfaces can be used to
differentiate the taxa in the sections Folliculatae and Collinsiae.
Studies of achene surfaces in other groups of Cyperaceae, particu-
larly in Scirpus and Eriophorum (Schuyler, 1971), Cyperus (Den-
ton, 1983), and in Carex (Walter, 1975; Toivonen & Timonen, 1976;
Standley, Ph.D. thesis, U. of Washington, Seattle, 1981, 1985; Tal-
lent & Wujek, 1983a, 1983b; Menapace, M.S. thesis, Central Mich.
U., Mt. Pleasant, 1985; Menapace & Wujek, 1985) have shown the
usefulness of achene epidermal studies in the elucidation of
species relationships.

In section Folliculatae and Collinsiae, the most distinctive epi-
dermal patterns are those of Carex lonchocarpa and C. collinsii, both
of which have prominent silica central bodies with conspicuous but-
tresses. The nature of the central body in these two taxa appears to
reflect convergence. They differ, however, in many other aspects of
their overall morphology (Mackenzie, 1935).

Apart from showing exclusive or distinctive achene micro-mor-
phological characters, all four taxa are distinguishable from each
other. The gross morphology of Carex folliculata and C. loncho-
carpa, substantiated by achene micromorphology, supports Mac-
kenzie’s (1935) recognition of the two taxa at the species level.

ACKNOWLEDGMENTS

This study was supported by a CMU Summer Fellowship to
D.E.W. We are grateful to A. A. Reznicek, MICH for allowing us

1986] Wujek & Menapace — Carex 403

to examine and remove perigynia from selected specimens. We
thank Dow Chemical Co., Midland for conducting the EDS
analysis.

LITERATURE CITED

CorreLL, D. S., AND H. B. Corrett. 1975. Aquatic and Wetland Plants of the
Southwestern United States. Stanford Press, Stanford, CA.

Denton, M.L. 1983. Anatomical studies of the Luzulae group of Cyperus (Cype-
raceae). Syst. Bot. 8: 250-262.

FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Co.,
Boston, MA.

GLeAsON, H. A. 1952. The New Britton and Brown Illustrated Flora of the North-
eastern United States and Adjacent Canada. Van Nostrand and Reinhold Co.,
New York, N.Y.

GLEASON, H. A., AND A, CRONQUIST. 1963. Manual of Vascular Plants of North-
eastern United States and adjacent Canada. Van Nostrand Co., Princeton,
N.J.

Goprrey, R. K., ANDJ. W. Wooten. 1979. Aquatic and Wetland Plants of South-
eastern United States. U. of Georgia Press, Athens, GA.

MACKENZIE, K. K. 1935. Cyperaceae-Cariceae. North American Flora 18: 169-
478.

MENAPACE, F. J. AND D. E. WuseK. 1985. Scanning electron microscopy as an aid
to sectional placement within the genus Carex (Cyperaceae): sections Lupulinae
and Vesicariae. Micron and Microscopia Acta 16: 213-214.

RADFORD, A. E., H. E. AHLES AND C. R. BELL. 1968. Manual of the Vascular
Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill.

SCHUYLER, A. E. 1971. Scanning electron microscopy of achene epidermis in spe-
cies of Scirpus (Cyperaceae) and related genera. Proc. Acad. Nat. Sci. Philadel-
phia 123: 29-52.

SMALL, J. K. 1933. Manual of the Southeastern Flora. Univ. of North Carolina
Press, Chapel Hill.

STANDLEY, L. A. 1985. Systematics of the Acutae group of Carex (Cyperaceae) in
the Pacific Northwest. Syst. Bot. Mongr. 7: |-106.

TALLENT, R. C., AND D. E. Wusek. 1983a. Taxonomy of several Carex species
using micromorphological characters. Amer. J. Bot. 70: 120 (abstr.).

AND . 1983b. Scanning electron microscopy as an aid to taxonomy
of sedges (Cyperaceae: Carex). Micron and Microscopica Acta 14: 271-272.

TOIVONEN, H., AND T. TIMONEN. 1976. Perigynium and achene epidermis in some
species of Carex subg. Vignea (Cyperaceae), studied by scanning electron
microscopy. Ann. Bot. Fennici 13: 49-59.

WALTER, K. S. 1975. A preliminary study of the achene epidermis of certain
Carex (Cyperaceae) using scanning electron microscopy. Mich. Bot. 14: 67-72.

DEPARTMENT OF BIOLOGY
CENTRAL MICHIGAN UNIVERSITY
MT. PLEASANT, MI 48859

CAREX STRIATA, THE CORRECT NAME FOR
C. WALTERIA NA (CYPERACEAE)

A. A. REZNICEK AND P. M. CATLING

In 1893, Liberty Hyde Bailey proposed the name Carex walteri-
ana to replace the then familiar C. striata Michaux for a widespread
coastal plain species ranging from Massachusetts to Florida and
Alabama. This name was a replacement because of the existence of
a prior C. striata Gilibert (1792). Bailey’s name has gained universal
acceptance in North American floras (Small, 1933; Mackenzie,
1935; Fernald, 1950; Gleason, 1952; Gleason & Cronquist, 1963;
Radford, Ahles, and Bell, 1964).

As pointed out by Hylander (1945) and McVaugh (1949), and
noted again by Crins (1985), Gilibert (1792) did not consistently use
the Linnaean binary system of nomenclature. The binomials in Gili-
bert’s work are thus not validly published under art. 32.1(b), for his
“epithets” are not to be treated as epithets (see art. 23 ex. 11) (Voss,
et al., 1983). Later use of such binomials does not create homonyms.
Michaux’s Carex striata, therefore, is not a later homonym, but
rather the earliest correct name for the species now known as C.
walteriana.

Carex striata Michaux, Fl. Bor.-Amer. 2: 174. 1803—Holotype:
“Hab. in Carolina,” Michaux (P, IDC microfiche 6211:110
MICH!)

Carex walteriana L. Bailey, Bull. Torrey Bot. Club 20: 429. 1893. Based on C.
striata Michaux.

Plants of Carex striata from South Carolina southward normally
have more or less pubescent perigynia. Plants occurring from South
Carolina northward supposedly have glabrous perigynia. Michaux’s
type has pubescent perigynia and the glabrous, northern plants have
been segregated as C. striata var. brevis L. Bailey [C. walteriana var.
brevis (L. Bailey) L. Bailey] (Bailey, 1889, 1893).

LITERATURE CITED
BarLey, L. H. 1889. Studies of the types of various species of the genus Carex.

Mem. Torrey Bot. Club I: 1-85.
—____. _ 1893. Notes on Carex-XVII. Bull. Torrey Bot. Club 20: 417-429.

405

406 Rhodora [Vol. 88

Crins, B. 1985. Gilibert’s plant names. Plant Press 3: 27-28.

FERNALD, M. L. 1950. Gray’s Manual of Botany, Ed. 8. Amer. Book Co., New
York.

Giipert, J. E. 1792. Exercitia Phytologica. [J. B. Delamolliére], Lyon. 2 vols.

GLEASON, H. A. 1952. Illustrated Flora of the Northeastern United States and
Adjacent Canada. New York Bot. Gard., New York. 3 vols.

AND A. CRONQUIST. 1963. Manual of Vascular Plants of Northeastern

United States and Adjacent Canada. Van Nostrand Reinhold Co., New York.

HyLANperR, N. 1945. Nomenklatorische und systematische studien tiber Nor-
dische gefasspflanzen. Uppsala Univ. Arsskr. 7: 1-337.

MACKENZIE, K. K. 1935. Cyperaceae-Cariceae. N. Amer. Fl. 18: 169-478.

McVauGu, R. 1949. Questionable validity of names published in Gilibert’s floras
of Lithuania. Gentes Herb. 8: 83-90.

Raprorp, A. E., H. E. AHLES, AND C. R. Bett. 1964. Manual of the Vascular
Flora of the Carolinas. University of North Carolina Press, Chapel Hill.

SMALL, J. K. 1933. Manual of the Southeastern Flora. Author, New York.

Voss, E. G., et al. 1983. International Code of Botanical Nomenclature Adopted
by the Thirteenth International Botanical Cogress, Sidney, August 1981. Reg-
num Vegetabile Vol. I 11.

A. A. R.

UNIVERSITY OF MICHIGAN HERBARIUM
NORTH UNIVERSITY BLDG.

ANN ARBOR, MICHIGAN 48109

P.M.C.

BIOSYSTEMATICS RESEARCH INSTITUTE
AGRICULTURE CANADA

WM. SAUNDERS BLDG., C. E. F.

OTTAWA, ONTARIO

CANADA KIA 0C6

NEW ENGLAND NOTES

REDISCOVERY OF /SOTRIA VERTICILLATA (WILLD.)
RAF. (ORCHIDACEAE) IN VERMONT

MICHAEL EFFRON AND ERROL C, BRIGGS

The large whorled pogonia (/sotria verticillata) is at the northern
limits of its range in northern New England, and is considered an
endangered species in Maine, New Hampshire, and Vermont (Crow
et al., 1980). In these three states, this orchid is less frequently found
than the other member of the genus, the federally listed /sotria
medeoloides (Pursh) Raf.

Isotria verticillata has been reported from two stations in Maine in
this century. The most recent report was from Waterford in 1974
(Campbell and Eastman, 1980). Brackley (1985) reported only one
extant population in New Hampshire.

Documented stations for this orchid in Vermont include a collec-
tion by Benedict in the 1840’s “at the High Bridge” in Colchester,
collections in Pownal (1897), in Bristol (1898), and a number of
collections in Burlington and Colchester in 1902, 1903, and 1904.
During the last 81 years, the species: has not been reported from
Vermont, leading the committee currently revising the state’s
endangered species list to assume it was extirpated in the state.

A vigorous population of /sotria verticillata was discovered on
June 4, 1985 in the town of Colchester (Chittenden County), several
miles from Benedict’s station. The orchids grow in a level forest of
middle-aged Acer rubrum, Quercus rubra and Q. alba, with occa-
sional Pinus Strobus. Understory trees and shrubs include Acer
pensylvanicum, Ostrya virginiana, Hamamelis virginiana, Kalmia
angustifolia, Vaccinium vacillans, Viburnum acerifolium, and Cory-
lus cornuta. The herbaceous community contains Maianthemum
canadense, Medeola virginiana, Trientalis borealis, Aralia nudicau-
lis, and Pteridium aquilinum. Names follow Fernald, 1950.

The population of /sotria verticillata at this station can be conser-
vatively estimated at over 1000 stems, and the species is the predom-
inant herbaceous plant at many sites in the area. The number of
plants and the apparent vigor of the colony is in keeping with the
occurrence of the species in more southerly parts of its range. With
the exception of an unvouchered account of /. verticillata from

407

408 Rhodora [Vol. 88

Aroostook County, Maine (Goodale, 1861), however, these plants
represent the northernmost station for either species of /sotria.
Specimens (Effron 0051) have been deposited at VT and NEBC.

LITERATURE CITED

FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Co.,
N.Y.

BRACKLEY, F. 1985. The Orchids of New Hampshire. Rhodora 87: 1-117.

CAMPBELL, C., ANDL. M. EASTMAN. 1980. Flora of Oxford County, Maine. Tech.
Bull. 99 Life Sciences and Agriculture Experiment Station, Univ. of Maine.
Orono.

Crow, G. E., W. D. CountRYMAN, G. L. CHURCH, L. M. EASTMAN, C. B. HELL-
guist, L. J. MEHRHOFF, AND I. M. Srorks. 1981. Rare and endangered vas-
cular plant species in New England. Rhodora 83: 259-299.

Goopate, G. L. 1861. Botanical notes on the new lands, pp. 361-372 Jn: Sixth
Annual Report, Agriculture and Geology of Maine. Stevens and Hayward,
Augusta.

AQUATEC, INC.
ENVIRONMENTAL SERVICES
75 GREEN MOUNTAIN DRIVE
SOUTH BURLINGTON, VT 05401

NEW STATIONS FOR PLATANTHERA FLAVA
AND TRIPHORA TRIANTHOPHORA
AND OTHER OBSERVATIONS

PHILIP E. KEENAN

ABSTRACT

New stations for Platanthera flava (L.) Lindl. and Triphora trianthophora (Sw.)
Rydb. were discovered through field studies in New Hampshire and Maine. A short
discussion of habits and ecology is also included.

Key Words: Platanthera flava, Triphora trianthophora, ecology, New Hampshire

PLATANTHERA FLAVA

Platanthera flava (L.) Lindl., the well-named tubercled orchid, is
one of our more infrequently seen orchids. Many botanists and
amateur orchid lovers have never seen it. Brackley (1984) considered
it uncommon in New Hampshire, listing it for three widely divergent
counties: Coos, Cheshire and Strafford. Eastman et al. (1981) listed
only one or two records for Maine in the last few decades, while
Crow et al. (1981) considered it rare in all of New England with the
exception of Connecticut. Once under review for Federal listing, the
Government dropped it from further consideration as being more
abundant than warranted for listing (Federal Register, 15 December
1980, vol. 45 no. 242).

In the past few years it has been my good fortune to discover three
new stations in York County, Maine, three in Rockingham County,
N.H. and three in Strafford County, N.H. The most recent discov-
ery in the town of Madbury, N.H. contained more than two thou-
sand plants, many of them bunched together, most of them in
bloom, and all within an area approximately fifteen by thirty
meters. According to Brackley (personal communication) this is the
largest station in New Hampshire. The plants were very robust, a
few exceeding the usual heights given—50 cm by Luer (1975) and
7 dm by Fernald (1950)—by several millimeters. The plants are
growing in a wet meadow environment, surrounded by forest, and
maintained as a meadow by periodic, but not annual, mowing.
While scattered in other parts of the meadow, the main site featured
a rank growth of sensitive fern (Onoclea sensibilis) which grew

409

410 Rhodora [Vol. 88

along the border of the woods and provided a uniform backdrop for
the orchids. One could conjecture as to the long-term effect of this
invasive weedy fern on the orchid niche. Typical companion plants
included J/ris versicolor (blue flag), Lilium canadense (meadow lily),
Lysimachia terrestris (swamp candles), Geum rivale (water avens),
Thalictrum polygamum (tall meadow-rue), Spiraea latifolia
(meadowsweet) Spiraea tomentosa (steeplebush), Geranium macu-
latum (wild geranium), and Eupatorium maculatum (Joe-Pye-weed)
among others. An interesting side-light involves the Joe-Pye-weed
here: long after all the other Joe-Pye-weed has reached peak bloom
and gone beyond, the patch growing beside the tubercled orchid is
just beginning to bud, and in some years fails to open because of
frost arrival. | have never seen this phenomenon anywhere else.

One of the peculiar characteristics of Platanthera flava is the
persistence of its flowers. The yellowish-green flower is in prime
condition for only about two weeks, but the calyx remains green in
many specimens well into September, sometimes until the first frost
kills it. After the first two weeks of peak color, the corolla begins to
blacken rapidly; the spur, lateral petals and lip become black about
the same time. The ovary, however, retains its green color. This
differential aging facilitates the off-season chance of identifying the
species all summer long. Many of the stems persist through the
winter (as does Goodyera pubescens).

Unfortunately, many orchid stations (including other orchids of
grassy habitats) are at least temporarily obliterated when the lan-
downer mows the meadow at the “wrong” time. Mowing undoubt-
edly accounts for some of the difficulty in finding this inconspicuous
orchid and makes it a little more uncommon.

Color photographs—slides, prints and enlargements—are in my
collection, including the closeup of one specimen with a handsome
deep yellow crab spider—with two maroon thorax marks—on a
spike of 40-50 flowers. I have seen many of the white crab spiders
with the pink thorax marks but never the yellow. In addition to the
photographic documentation, vouchers have been deposited in
NHA.

1986] New England Notes 411

TRIPHORA TRIANTHOPHORA

Triphora trianthophora (Sw.) Rydb., the nodding pogonia or
three-birds orchid, is a most fascinating plant which I have been
studying for a number of years. During this time I have also discov-
ered three new stations in Holderness, N.H. (Grafton County). This
orchid is saprophytic, subterranean and erratic in nature. It is also
tiny, pretty and ephemeral. Each flower blooms only for a day—in
prime condition; some flowers will remain unfresh into the second
day. Usually only one or two flowers open at once, sometimes three.
Since there are only one to six buds produced on each plant, the
effective bloom period for an individual plant is only about three
days. Each one of these three blooming days is usually separated by
an interval of several days. The flowering period, therefore, can be
as long as two or three weeks, but with only three days of actual
open flowers, due to the simultaneous mass flowering of a popula-
tion on each of these approximately three days. This situation is
unique among our wild flowers, and explains why no flowers are
detected on several field trips during the anticipated blooming
period. In 1974, I made five trips to a Triphora station in ten days’
time and found it in bloom, in mass, only on the last day. The day of
bloom occurred forty-eight hours after a colder than usual night, a
phenomenon known as thermoperiodicity. This response has been
shown experimentally under artificial conditions also (Luer, 1975).
Sometimes an isolated plant in a population will bloom alone, “off
schedule” or be a second-day flower. The order of opening is also
erratic. The top and bottom bud may open together while the mid-
dle one remains closed, or sometimes, when the plant has only two
buds, the top one will open first.

In 1984 I observed another interesting occurrence. On August 25
the station was in mass bloom. Eight days later the ovaries of all the
plants had been nipped off, while more than half of all the plants
had been eaten in their entirety. On the previous August 25, after
completing a photographic session with an especially highly-colored
clump, I noticed a chipmunk come out of his hole beside a decayed
tree trunk which was lying on the ground in close proximity to the
prized group of Triphora. There was no sign of this clump eight
days later. The attractiveness of the succulent stems to rodents has
been noted for plants under cultivation and no doubt is also a

412 Rhodora [Vol. 88

problem in the wild, as I have observed for several years. It must
also account for some of the erratic appearances of this beautiful
orchid—here today, gone tomorrow.

Vouchers have been deposited in NHA.

LITERATURE CITED

BRACKLEY, F. E. 1984. The Orchids of New Hampshire. Rhodora 85: 1-119.

Crow, G. E., W. D. CountrYMAN, G. L. CHurcH, L. M. EASTMAN, C. B. HELL-
Quist, L. J. MEHRHOFF AND I. M. Storks. 1981. Rare and endangered vascu-
lar plants in New England. Rhodora 83: 259-299.

EASTMAN, L. M. ANDS.C. GAWLER. 1981. Rare Vascular Plants of Maine. Criti-
cal Areas Program. 1981. State Planning Office, Augusta, Maine 04333.

FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Co.,
New York.

Luer,C. A. 1975. The Native Orchids of the United States and Canada Excluding
Florida. N.Y. Botanical Garden, Bronx, N.Y.

31 HILLCREST DRIVE
DOVER, NH 03820

FOUR RARE EURASIAN ADVENTIVES IN
THE FLORA OF WESTERN MASSACHUSETTS

MICHAEL H. GRAYUM AND MARIAN F. ROHMAN

ABSTRACT

Recent collections of four weedy species of angiosperms native to Eurasia are
reported from Hampshire County, Massachusetts. Elsholtzia ciliata and Lepidium
perfoliatum are reported for the first time from western Massachusetts. Three collec-
tions of Poa bulbosa from the Amherst area mark the first records of this species
from New England. Apera (Agrostis) interrupta is reported for the firt time from the
eastern seaboard of the United States.

Key Words: Labiatae, Cruciferae, Gramineae, Eurasian adventives, new records,
Massachusetts

INTRODUCTION

Botanical collecting by the authors in the Pioneer Valley area of
Hampshire County, Massachusetts over the past six years has
revealed the presence of four rare angiosperm weeds native to Eura-
sia. Since the distribution of these species is of general relevance to
the New England flora, the collections are detailed below. The order
is that of Gray’s Manual (Fernald, 1950).

Poa bulbosa L.

Fernald (1950) reported Poa bulbosa only as far north as Long
Island and Yonkers, New York. It can now be added to the New
England flora on the basis of three collections, all from the Pioneer
Valley area of Hampshire County where it appears to be becoming
rather well established.

The first collection of Poa bulbosa from New England was appar-
ently that of the late Harry E. Ahles, as a lawn weed on North
Maple Street, Hadley: 21 May, 1979, Ahles 86583 (MASS, NEBC).
The species was later collected in a similar situation along North
Pleasant Street, Amherst: 20 May, 1981, Grayum 3080 (MASS). It
has recently been discovered in relative abundance on the campus of
the University of Massachusetts, Amherst: 3 May, 1984, Rohman &
Grayum 2335 (MASS).

Apera interrupta (L.) Beauv.
This species has generally been treated in North American floras
as Agrostis interrupta L. Most recent European authors, however,

413

414 Rhodora [Vol. 88

prefer to segregate the oligotypic genus Apera, which (though rather
disturbingly polythetic) may be distinguished from Agrostis sensu
stricto on the basis of the following characteristics [key adapted
from Hitchcock (1950) and Tutin (1980) ]:

Lemma firm at maturity, shortly bearded at the base (in ours),
awned from just below the tip; awn straight, usually at least 4
mm long; upper glume 3-veined; palea strongly 2-nerved, about
as long as the lemma; rachilla prolonged back of the palea as a
WGK IU hae cee coe ca cieo uk ners skReRewers Apera

Lemma thin, glabrous basally or variously hairy or bearded, awn-
less or awned from near or below the middle; awn, if present,
straight or geniculate, often less than 3 mm long; upper glume
usually l-veined; palea usually nerveless and shorter than the
lemma, or obsolete; rachilla usually not prolonged or, if so,
minutely pubescent to vilose peck ce de staedsuwine ss Agrostis

Although Apera spica-venti (L.) Beauv. (= Agrostis spica-venti
L.) is a rather widespread weed in North America, most records of
A. interrupta are from the Pacific coast. Gleason & Cronquist
(1963) reported a collection from the vicinity of St. Louis, Missouri
(see also Miihlenbach, 1979), and there have been several recent
collections from the midwestern United States and adjacent Canada
(see Solheim & Judziewicz, 1984). The species seems to be spreading
rapidly eastward, having been collected in Missisquoi County,
Quebec by 1979. The following specimen, from the campus of the
University of Massachusetts, Amherst, Hampshire County, is
believed to represent the first collection from the eastern seaboard
of the United States: 7 June, 1982, Rohman & Grayum 2030
(MASS, NEBC).

The following herbaria were checked, without success, for eastern
specimens of Apera interrupta that might have been overlooked:
MASS, NEBC, GH, NY, and MO. It was obviously not possible to
check all of the hundreds of herbaria in the eastern United States, so
it is conceivable that one or more of these may harbor additional
specimens of this rare weed.

Apera interrupta may be distinguished from the more common A.
spica-venti by several characteristics not alluded to in most North
American floras. The following key, which also includes a third

1986] New England Notes 415

Eurasian species not yet reported from North America, has been
adapted from Bor (1970) and Tutin (1980):

Panicle branches long, spreading; lemma shortly bearded at the

base; anthers 1-2 mm long ..... A. spica-venti (L.) Beauv.
Panicle branches short, erect

Upper glume awned; lemma shortly bearded at the base; anth-

ers 0.3-0.5 mm long ........... A. interrupta (L.) Beauv.
Upper glume shortly awned; lemma glabrous at the base; anth-
Ors Ca: 1,5 Tm 1008. 6 ees cine eee A. intermedia Hackel

Additional features that may aid in identifying Apera interrupta
are its narrow (1-4 mm broad), more or less involute leaves and
generally shorter ligules (2-5 mm, as opposed to 3-12 mm in A.
spica-venti). In general, any small, annual, Agrostis-like grass with a
strict inflorescence and conspicuously awned lemmas should be
checked against the description of Apera interrupta, which is cer-
tainly more widespread than records indicate.

The station for Apera interrupta reported above was mowed just
three days following the collection, and subsequently ravaged by a
construction project. No sign of the species has been seen since.

Lepidium perfoliatum L.

Collected on the campus of the University of Massachusetts,
Amherst, Hampshire County: 27 April, 1984, Grayum & Sleeper
3272 (MASS). This record is apparently the first for Hampshire
County, and for western Massachusetts in general. According to
unpublished recoreds of H. E. Ahles, the most recent New England
collection is from Essex County, Massachusetts, in 1954. Prior to
that date, L. perfoliatum had been collected in Worcester County,
Massachusetts, in 1938, but never any further west in New England.

On the University of Massachusetts campus, Lepidium perfolia-
tum grows alongside Poa bulbosa at the same site—the south side of
the Student Health Center—formerly occupied by Apera interrupta.
This commonality of habitat suggests that these three adventive
species may owe their introduction to an exotic and rather impure
source of soil or, more likely, grass seed. The area was indeed
reseeded following a recent construction project, but the collection
of Apera predated these events. Poa bulbosa, as has been noted, is
also known from two other disparate locales in the Amherst area. It

416 Rhodora [Vol. 88

thus seems improbable that the presence of three of the four adven-
tives reported herein is traceable to a single introduction.

Elsholtzia ciliata (Thunb.) Hylander

Although this rare labiate was collected in Chittenden County,
Vermont as recently as 1975 (Ahles 81660, MASS), it has not been
collected in Massachusetts since a 1958 collection from Essex
County (according to unpublished data of H. E. Ahles). The western-
most Massachusetts collection is from Worcester County (1952).
The following specimen thus represents the first record from Hamp-
shire County, as well as from western Massachusetts in general:
along railroad tracks at east side of Arcadia Lake, Belchertown, 13
September, 1978, Grayum 1533 (MASS). The site is at an elevation
of about 315 feet.

ACKNOWLEDGMENT

We express our sincere appreciation to the late Harry E. Ahles,
whose enthusiasm and expertise were an inspiration to legions of
botany students (ourselves included) at several different universities,
and whose meticulous and voluminous data books contributed sub-
stantially to the present report.

LITERATURE CITED

Bor, N. L. 1970. Gramineae. 1-573. Jn: K. H. Rechinger, Ed. Flora Iranica No.
70.

FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th ed. American Book Co.,
New York.

GLEASON, H. A. AND A. Cronquist. 1963. Manual of Vascular Plants of North-
eastern United States and Adjacent Canada. D. Van Nostrand Co., Princeton,
N. J.

Hitcucock, A. S. 1950. Manual of the Grasses of the United States. U.S. Gov't
Printing Office, Washington.

MUHLENBACH, V. 1979. Contributions to the synanthropic (adventive) flora of
the railroads in St. Louis, Missouri, U.S.A. Ann. Missouri Bot. Gard. 66: 1-108.

SOLHEIM, S. L. AND E. J. Jupziewicz. 1984. Four noteworthy Wisconsin plants.
Phytologia 54: 490-492.

Tutin, T.G. 1980. Gramineae. 118-267. In: T. G. Tutin, V. H. Heywood, N. A.
Burges, D. M. Moore, D. H. Valentine, S. M. Walters and D. A. Webb, Eds.
Flora Europaea. Vol. 5. Cambridge Univ. Press, London.

1986] New England Notes
MiH..G.

MISSOURI BOTANICAL GARDEN

P.O. BOX 299

ST. LOUIS, MO 63166

M.F.R.

DEPARTMENT OF BOTANY
UNIVERSITY OF MASSACHUSETTS
AMHERST, MA 01003

417

THE FIRST RECORD OF POA BULBOSA L. (POACEAE)
FOR VERMONT

PETER F. ZIKA

In the past 20 years, several adventive weeds reported for the first
time from the campus of the University of Vermont have become
pests in the state. These weeds include Geum urbanum L., Polygo-
num caespitosum Blume var. longisetum (DeBruyn) Stewart (Zika
et al., 1983) and Puccinellia distans (L.) Parl. (Seymour, 1967).
Therefore it seems useful to note the occurrence of a new adventive.
Vermont’s first station for Poa bulbosa L. came from a weed patch
near a dumpster by Cook Science Building on the University of
Vermont campus, in Burlington, Chittenden County, on 6 May
1985. The species was recognizable only because a few days of rainy
spring weather had delayed the mowing schedule of the ground crew
and allowed some inflorescences to form. For five years the writer
had walked past this very spot several times a week and not noticed
the species.

The specimens collected (Zika 8974 NEBC, VT) showed no nor-
mal spikelets, only viviparous inflorescences, and aggressively
spreading bulbous bases connected by short runners. The species
may have been introduced as an impurity in lawn seed or hay bales,
or by the muddy construction machinery commonly found on
campus.

A check of the Pringle Herbarium showed no Poa bulbosa speci-
mens from Vermont or any other New England state. The range
map in Hitchcock and Chase (1950) indicated no records of this
European weed north or east of New York State. Thus P. bulbosa is
recorded here as new to Vermont, and as a recent northeastern
extension of range. Although Seymour (1982) did not include the
species in his New England flora, Marian Rohman (pers. comm.)
reports four records from two counties in western Massachusetts,
where the species was first collected in the mid-1970’s by Harry E.
Ahles.

LITERATURE CITED

Hitcucock, A. S. (Revised by A. Chase) 1950. Manual of the Grasses of the
United States. 2nd ed. U.S.D.A. Misc. Pub. No. 200, U.S. Gov. Printing Office.

419

420 Rhodora [Vol. 88

SEYMouR, F. C. 1967. Notes from the Pringle Herbarium, I]. Rhodora 69:
377-380.
1982. The Flora of New England. 2nd. ed. Phytologia Memoirs V. Mol-
denke & Moldenke, Plainfield, NJ.
ZikA, P. F., R. J. STERN AND H. E. AHLES. 1983. Contributions to the flora of the
Lake Champlain Valley, New York and Vermont. Bull. Torrey Bot. Club 110:
366-369.

PRINGLE HERBARIUM
BOTANY DEPARTMENT
UNIVERSITY OF VERMONT
BURLINGTON, VT 05405

BOOK REVIEW

A. C. JERMY AND T. G. WALKER. 1985. Cytotaxonomic Studies
of the Ferns of Trinidad. Bull. Brit. Mus. Nat. Hist. Bot 13:
133-276. (Price £22.00)

During the course of an earlier study on the ferns of Jamaica,
which focused on the cytological features of a tropical island fern
flora, it was determined that a broader scope was needed to place
the evidence within a larger framework. Accordingly, Trinidad was
selected for a similar study and the results are now reported here.

The Bulletin consists of three separate papers. The first by A. C.
Jermy is a general review of the geology, soils, climate and vegeta-
tion of Trinidad, and the last by Jermy and Walker describes nine
sterile hybrids and five new species, three of them diploid and two
tetraploid. The main portion of the Bulletin consists of the second
paper on “The cytology and taxonomic implications” by T. G.
Walker. This is a major contribution to fern cytology and especially
to the cytology of tropical American ferns. It contains 155 records
of taxa and their cytotypes.

Surveys were made of the island ferns during trips from 1963
through 1974. A list of the species and their chromosome numbers
and ploidy levels contains many new reports for American ferns,
although some species were previously reported, especially from
Jamaica, but also from Mexico, Central America, Puerto Rico, and
Brazil.

The results are discussed under the treatments of families and
genera and previous reports are integrated with the Trinidad find-
ings. Morphological characters are often included in these accounts
as well as details of species complexes and hybrids. The studies are
especially critical in difficult groups as Lygodium, Gleicheniaceae,
Adiantum, Polypodium, Goniopteris, Asplenium, and Blechnum.

Although the author states that (except for Polypodium sens. lat.)
the classification of Crabbe et al. (Brit. Fern Gaz. 11: 141-162) is
followed, there are in fact departures from that system. For exam-
ple, Nephrolepis and Oleandra in a subfamily of the Aspleniaceae
rather than the Davalliaceae. The treatment of the Cyatheaceae
includes Cnemidaria under Cyathea, rather than a distinct genus,
and Sticherus is placed under Gleichenia.

421

422 Rhodora [Vol. 88

Much work is devoted to karyotyping, including a lengthy expla-
nation of a new methodology. Nineteen karyograms are illustrated,
some of large ones as Polypodium phyllitidis with 148 chromo-
somes. This impressive data is used to distinguish alloploid taxa as
Lygodium venustum and 4X Polypodium aureum from an interpre-
tation as possible autoploids. However, many problems still persist
with this method that place constraints on general evolutionary
conclusions.

The cytology has valuable implications for the species classifica-
tion and the recognition of polyploid complexes. The results
emphasize the importance of field study followed by rigorous cyto-
logical analysis for an understanding of the dynamics of tropical
fern floras. Much more needs to be done, for as Walker comments
under the genus Blechnum “... it is clear that as far as the B. occi-
dentale complex is concerned only the tip of the iceberg has been so
far exposed.” The same could be said for Adiantum and other gen-
era where cytological knowledge can be expected to provide critical
data on problems evident from taxonomic studies.

ROLLA TRYON

ALICE TRYON

HARVARD UNIVERSITY HERBARIA

22 DIVINITY AVE., CAMBRIDGE, MASS. 02138

BOOK REVIEW

EDWARD G. Voss. 1985. Michigan Flora, Part II Dicots
(Saururaceae-Cornaceae). xix + 724 pp. Cranbrook Institute
of Science, Bulletin 39 and the University of Michigan Herba-
rium. (Price $12.50)

Part II of the Michigan Flora continues the outstanding coverage
of the plants from Michigan growing outside of cultivation. The
coverage includes nearly 1000 species in the apetalous and polype-
talous families. The parts of the Michigan Flora correspond to the
divisions of Gleason’s New Britton and Brown Illustrated Flora.
The style is identical to Part / published in 1972: each family treat-
ment includes a key to genera and species, descriptions of taxa,
references, county dot maps, and representative illustrations. A key
to genera has been purposely omitted until Part /// is published.

This volume, like Part /, is a comprehensive study of Michigan
plants. The authors consulted the herbaria in Michigan and selected
herbaria throughout the United States for localities and measure-
ments. All measurements stated are from Michigan specimens with
the exception of rare plants.

The keys are excellent and easy to use. Nomenclature is up-to-
date. If nomenclatural problems occur, Voss discusses his treatment
decision. Varieties and subspecies are not included in the keys, but
are treated in the writeup. Habitat data were compiled from herba-
rium sheets, Voss’s personal field observations, or from original
local information.

The format of the book is superior. The print is clear and use is
made of various print sizes and types. The maps and illustrations are
clear. Not all species are illustratéd, but there is at least one repre-
sentative species included per genus. A glossary and index to botan-
ical and common names are also included.

Part IT as wellas Part I, of the Michigan Flora is a must for every
botanist, botanical student, conservationist, or person interested in
plant identification in Michigan. For those of us outside of Michi-
gan, this is a valuable reference. This volume may be used by anyone
in the northeastern portion of the United States and southeastern

423

424 Rhodora [Vol. 88

Canada. This book is a “must” purchase and is also a great bargain
for only $12.50. I highly recommend Part // and eagerly await Part
III.

C. BARRE HELLQUIST

DEPARTMENT OF BIOLOGY

NORTH ADAMS STATE COLLEGE
NORTH ADAMS, MASSACHUSETTS 02147

IN MEMORIAM
FRANK C. SEYMOUR 1896-1985

The Boston Globe for January 20, 1985, carried a notice that Dr.
Frank C. Seymour, 89, who joined NEBC in 1915 and who served as
President in 1963 and 1964, died in Dartmouth, MA on January 15.
Dr. Seymour was a renowned botanist, author and Congregational
minister. Many of his plant specimens are in local herbaria, includ-
ing the Knowlton Herbarium of the University of Massachusetts.
He served as Curator of the Pringle Herbarium at UVM, as Assist-
ant Professor at the University of Florida in Gainesville, and as a
research assistant at the Missouri Botanical Garden. He wrote a
comprehensive “Flora of New England,” which has gone into its
second printing. Burial was in Granville, MA.

425

426 Rhodora [Vol. 88

IN MEMORIAM
HENRY K. SVENSON'- 1897-1986

Dr. Henry K. Svenson, 89, who joined NEBC in 1919, died on
March 4. The following tribute appeared in the Boston Globe for
Friday, March 7, 1986.

Henry K. Svenson, a botanist who coauthored a guide to the flora
of Cape Cod, died Tuesday in Cape Cod Hospital in Hyannis after a
long illness. He was 89.

Born in Hinnaryd, Sweden, Mr. Svenson moved to Malden in
1898 and worked in his family’s musical instrument manufacturing
business in Boston.

In 1921, he graduated from Harvard University. After serving in
the US Army, he returned to Harvard, receiving his doctorate in
botany in 1929.

From 1930 to 1946, he was curator of the Brooklyn Botanical
Gardens in New York. He was chief botanist for the Vincent Astor
Expedition to the Galapagos Islands during his first year there.

In 1946, he was appointed curator in forestry and botany at the
American Museum of Natural History in New York City. Six years
later, he joined the US Geological Survey’s military branch, and
worked on vegetation maps of Europe and North America.

A year after his retirement in 1966, Mr. Svenson moved to Oster-
ville. In 1979, he coauthored “The Flora of Cape Cod” with Robert
W. Pyle.

Mr. Svenson was a member of the Cape Cod Museum of Natural
History, the Osterville Men’s Club, the Explorers Club of America,
the New England Botanical Club and the Joselyn Botanical Society.

Vol 88, No. 854, including pages 157-295, was issued April 29, 1986

INSTRUCTIONS TO CONTRIBUTORS TO RHODORA

Submission of a manuscript implies it is not being considered for
publication simultaneously elsewhere, either in whole or in part.

Manuscripts should be submitted in triplicate (an original and
two xerographic copies) and must be double-spaced (at least 3/8”)
throughout including tables, figure legends, and literature citations.
Please do not use corrasable bond. The list of legends for figures
and maps should be provided on a separate page. Footnotes should
be used sparingly. Do not indicate the style of type through the use
of capitals or underscoring, particularly in the citation of specimens.
Names of genera and species may be underlined to indicate italics in
discussions. Specimen citations should be selected critically, espe-
cially for common species of broad distribution. Systematic revi-
sions and similar papers should be prepared in the format of “A
Monograph of the Genus Malvastrum”, S. R. Hill, Rhodora 84:
1-83, 159-264, 317-409, 1982, particularly with reference to inden-
tation of keys and synonyms. Designation of a new taxon should
carry a Latin diagnosis (rather than a full Latin description), which
sets forth succinctly just how the new taxon is distinguished from its
congeners. Papers of a floristic nature should follow, as far as possi-
ble, the format of “Annotated list of the ferns and fern allies of
Arkansas”, W. Carl Taylor and Delzie Demaree, Rhodora 81:
503-548, 1979. For bibliographic citations, refer to the Botanico-
Periodicum-Huntianum (B-P-H, 1968), which provides standard-
ized abbreviations for journals originating before 1966. All abbrevi-
ations in the text should be followed by a period, except those for
standard units of measure and direction (compass points). For
standard abbreviations and for guidance in other matters of biologi-
cal writing style, consult the CBE Style Manual, Sth ed. (original
title: Style Manual for Biological Journals). In preparing figures
(maps, charts, drawings, photos, etc.) please remember that the
printed plate will be 4 X 6 inches; be sure that your illustrations are
proportioned to reduce correctly, and indicate by blue pencil the
intended limits of the figures. (Some “turn-page” figures with brief
legends will be 3 1/2 X 6 in.) Magnification/ reduction values given
in text or figure legends should be calculated to reflect the actual
printed size. An Abstract and a list of Key Words should be sup-
plied at the beginning of each paper submitted, except for a very
short article or note.

RHODORA July 1986 Vol. 88, No. 855

CONTENTS

The morphology and cytology of Polystichum X potteri hybr. nov. (= P
acrostichoides < P. braunii)
David S. Barrington
Plantago maritima and Carex piackinbatel new er Saskatchewan Additional
rare inland stations for two seacoast salt marsh species
Vernon L. Harms, Donald F. Hooper, and Les Baker
Dispersion pattern of aerial shoots of the common marsh reed Phragmites
australis (Poaceae)
Andrew N. Davis and Timothy L. Briggs
Seasonal succession and vertical distribution of phytogiaainos in Candle-
wood Lake, CT
Stanley J. Freeda and Peter A. Siver
New wool-alien Cruciferae (Brassicaceae) in canna North hnachlss Lepi-
dium and Sisymbrium
Ihsan A, Al-Shehbaz
Current status of Magnolia virginiana in Massachusetts
Richard B. Primack, Edward Hendry, and Peter Del Tredici
Studies in the Aristida (Gramineae) of the southeastern United States. IV.
Key and conspectus
Kelly W. Allred
Nomenclatural notes on Hedyotis rosea Rafi inesque aid anew sigidiaiien in
Houstonia
Edward E. Terrell ‘
Taxonomy of Carex section Folliculatae oes jdhaes morpuuiens
Daniel E. Wujek and Francis J. Menapace
Carex striata, the correct name for C. walteriana iCrmernceaal
A. A. Reznicek and P. M. Catling ,
NEW ENGLAND NOTES
Rediscovery of Isotria verticillata (Willd.) Raf. (Orchidaceae) in Vermont
Michael Effron and Errol C. Briggs
New stations for Platanthera flava and Triphora trianthaghors snd other
observations
Philip E. Keenan
Four Eurasian adventives in the Sie of walters Massachapeitts
Michael H. Grayum and Marian F. Rohman
The first record of Poa bulbosa L. (Poaceae) for Vermont
Peter F. Zika
BOOK REVIEWS
Cytotaxonomic Studies of the Ferns of Trinidad
Rolla Tryon and Alice Tryon
Michigan Flora, Part II, Dicots ‘enarnbiiad-Cabeaines
C. Barre Hellquist
IN MEMORIAM
Frank Seymour
Henry Svenson

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Roovdova

JOURNAL OF THE NEW ENGLAND BOTANICAL CLUB

Vol. 88 October 1986 No. 856

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Cover Illustration

An original drawing, seemingly the only one surviving and perhaps the only one
ever done for publication by Merritt Lyndon Fernald, used in part to illustrate his
article on cranberry species which appeared in RHODORA No. 48 (Fernald, M. L.
1902. The variations and distribution of American cranberries. Rhodora 4: 231-237
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Tbhodora

(ISSN 0035-4902)
JOURNAL OF THE

NEW ENGLAND BOTANICAL CLUB

Vol. 88 October 1986 No. 856

CAREX OLIGOCARPA (CYPERACEAE), A RARE SEDGE IN
CANADA NEWLY DISCOVERED IN QUEBEC

STUART G. HAY AND DANIEL GAGNON

ABSTRACT

The first authentic record in Québec of Carex oligocarpa is reported in the
Ottawa Valley. This discovery, in the Eardley Escarpment, represents a significant
outlying northerly station for this rare sedge and is only the seventh known location
in Canada. Aspects of the site and forest community are described. The phytogeo-
graphic importance of the new record is examined in the context of other unique and
rare elements of the Outaouais flora.

Key Words: Carex oligocarpa, rare, new record, phytogeography, Ottawa Valley
Québec, Canada

Carex oligocarpa Schk. ex Willd., a rare Canadian sedge, was
recently discovered at the Eardley Escarpment of Gatineau Park in
the Ottawa Valley, Québec. This constitutes the first authentic
Québec record. The new, outlying station represents a significant
range extension and is of considerable interest in view of current
work on rare and endangered vascular plants in Canada.

Carex oligocarpa and C. Hitchcockiana Dewey are the sole
members of the section Oligocarpae Carey. These two sedges are
rich woodland species of the deciduous forest region of eastern
North America. They are often rare in peripheral parts of their
range and both are at the northern limit of their distribution in
southern Ontario or southwestern Québec.

Floras covering northeastern North America, including the most
recent by Scoggan (1978), usually signal the presence of Carex oli-
gocarpa in southwestern Québec. The revised edition of Flore Lau-
rentienne (Marie-Victorin, 1964) also includes it, as an addendum.
Fr. Louis-Alphonse (1956, M.Sc. thesis, Univ. of Montréal)

427

428 Rhodora [Vol. 88

mapped the species from four locations southeast of Montréal,
based on a supposed collection from Philipsburg and other relevant
collections from Montréal Island, Chambly and Mt. Johnson con-
served at CAN, DAO, MT, and QFA. However, Ball & White
(1982) and Boivin (unpublished data) discount the species from the
province’s flora. A thorough herbarium search for The Rare Vascu-
lar Plants of Québec (Bouchard et al., 1983), revealed no substan-
tiating specimens. Erroneously identified specimens of remotely
similar species in the section Laxiflorae or in the closely allied sec-
tion Griseae evidently account for the mistaken references to C.
oligocarpa prior to this paper. Although somewhat similar to C.
Hitchcockiana, C. oligocarpa is readily distinguished by its glabrous
leaf sheaths and straight-beaked achenes.

Authentic Carex oligocarpa was first discovered in the course of
an ecological study of the Eardley Escarpment (fault-scarp of the
Precambian Shield) in the Ottawa Valley (Gagnon, 1980, M.Sc.
thesis, Univ. of Montréal; Gagnon & Bouchard, 1981). A single
collection of the sedge was made but the specimen was misidentified
as C. ormostachya Wieg. and its proper identity was only revealed
by a recent revision of the sheet. The species appears to be very rare
on the escarpment. The field data (Gagnon, 1980, M.Sc. thesis, Univ.
of Montréal, sample stand #69) indicate that C. oligocarpa (sub C.
ormostachya) was recorded as a single clone from a | m? quadrat
within a 50 m X 20 m sampling plot. The site has not been reexam-
ined since the discovery to assess the species frequency more accu-
rately. The label information on the voucher specimen is as follows:
Cté. de Gatineau; Parc de la Gatineau, escarpement d’Eardley.
Forét séche dominée par Ostrya virginiana, Tilia americana,
Carya cordiformis (alt. 420’, approx. 45°30’N, 75°54’W). 16 juin
1978. Gagnon & Bergeron S4M-12 (MT)

The new station for this sedge was found in a young stand of
maple-hickory forest community type as described by Gagnon and
Bouchard (1981). Ostrya virginiana (Mill.) K. Koch was the domi-
nant tree species due to past selective logging of the forest. Sub-
dominants were Tilia americana L., Carya cordiformis (Wang.) K.
Koch and Juglans cinerea L. The sapling layer was dominated by
Ostrya virginiana, Tilia americana, and Carya cordiformis. Her-
baceous species characteristic of these maple-hickory communities
such as Amphicarpa bracteata (L.) Fern., Desmodium glutinosum
(Muhl.) Wood, Oryzopsis racemosa (J. E. Smith) Richer, Phryma

1986] Hay and Gagnon — Carex oligocarpa 429

leptostachya L., Sanguinaria canadensis L. and Solidago caesia L.
were also recorded. The maple-hickory forests of the escarpment are
situated in the lower areas of south-facing slopes where they benefit
from increased moisture conditions because of seepage as well as
from a warmer microclimate (Gagnon & Bouchard, 1981). The site
was situated on a steep (40%) south-facing slope at the foot of the
escarpment. The soil was a well-drained Orthic Melanic Brunisol
developing on colluvial debris of diopsidic gneiss. The maple-
hickory communities of the escarpment generally have soils derived
from tills low in calcium, but seepage due to topographical position
increases soil nutrient levels, including calcium. This is similar to the
situation described in southern Minnesota (Wheeler & Ownbey,
1984) where the preferred habitats of Carex oligocarpa are steep,
hardwood-covered slopes and ravines on calcareous soils. Macken-
zie (1931-35) described this sedge as a calciphile or near-calciphile
of rich woodlands in calcareous districts. Other rare or uncommon
sedges such as Carex backii Boott, C. cephaloidea Dewey, C. cepha-

/ LAKE HURON

Pp

A aie

ré
t ee
| \ EB Toronto
| | : LAKE ONTARIO
i f ae) ; \- Buffalo
LAS A =
/ ay ~*~ _ e
ae
fs o LAKE ERIE we CAREX OLIGOCARPA

Figure 1. Distribution of Carex oligocarpa in southwestern Québec (solid dot,
reported in this paper) and southern Ontario (triangles). Open triangles indicate
pre-1910 collections; solid triangles represent recent collections at Point Pelee and
Pelee Island (map after Ball & White, 1982).

430 Rhodora [Vol. 88

lophora Muhl., C. Hitchcockiana Dewey, C. platyphylla Carey and
C. sprengelii Dewey were also recorded in this community and are
also usually associated with calcareous woodlands in the northern
part of their range.

Carex oligocarpa is essentially a southern and eastern species
whose range coincides largely with the eastern deciduous forest. In
the Great Lakes region, it reaches its northern limit in southeastern
Minnesota, where it has only recently been reported (Wheeler, 1981;
Wheeler & Ownbey, 1984), and in southernmost Wisconsin, Michi-
gan and Ontario. Further to the east, it is found in New York State,
and isolated populations occur in Vermont, Massachusetts and
Connecticut. The species is rare throughout most of its northern
range. It is on the lists of rare and endangered vascular plants of
Michigan (Wagner et al., 1977), Ontario (Ball & White, 1982), Ver-
mont (Countryman, 1978), Massachusetts (Coddington & Field,
1978) and Connecticut (Dowhan & Craig, 1976; Mehrhoff, 1978).

In Ontario, where its distribution has recently been mapped (Ball
& White, 1982), Carex oligocarpa is known from only six locations
along Lake Erie and Lake Ontario (Figure 1). Four of these records
are pre-1910, from Durham Co. (Macoun s.n., DAO #172252),
Prince Edward Co. (Macoun 2031, 31551, CAN), Elgin Co.
(Macoun 16569, CAN) and Welland Co. (Klugh s.n., TRT #189605).
In 1969, C. oligocarpa was discovered in Essex Co. at Point Pelee
(Maycock 13678, in herb. propr.). The status and habitat of the
colonies at Point Pelee National Park have recently been examined
by S. Varga (1984). Carex oligocarpa has also been discovered in
1981-84 on Pelee Island where A. A. Reznicek and M. J. Oldham
have made several noteworthy collections (Reznicek 6313, MICH;
Oldham 2571, CAN, MICH, TRTE; 2664, CAN, TRTE; 375/,
CAN, MICH, TRTE; 3767, TRTE; 4284, TRTE). At the Pelee
location, which was too recently reported to be included on the
Ontario map by Ball and White (1982), the species appears to be
relatively widespread on calcareous soils in deciduous woods and
open savannah.

Carex oligocarpa is an important addition to the Québec list of
rare vascular plants since in Canada its only other known occur-
rence is in Ontario, where it is already classed as a rare species. It is
the latest addition to an already important contingent of rare vascu-
lar plants known from the Eardley Escarpment area of the Ottawa
Valley (Brunton & Lafontaine, 1974; Gagnon, 1980, M.Sc. thesis,

1986] Hay and Gagnon — Carex oligocarpa 43]

Univ. of Montréal, 1985; Gagnon & Bouchard, 1981). Bouchard et
al. (1983, 1985) have shown that the Outaouais region of Québec
constitutes the richeSt floristic sector for rare species in the province.
Moreover, a recent study of the ecological distribution of rare spe-
cies in different plant communities of the central sector of the
Ottawa Valley (Gagnon, 1985) shows that the Eardley Escarpment
harbors the highest concentration of rare plants, particularly in the
oak forests and the maple-hickory communities. The recent addi-
tion to the area’s flora (Gillett & White, 1978) of such species as
Carex muhlenbergii Schk. (Gagnon, 1985), Claytonia virginica L.
(Gagnon, 1980) and now Carex oligocarpa is perhaps an indication
that still other rare southern elements may be found.

Various authors (Brunton & Lafontaine, 1974; Gagnon, (1980,
M.Sc. thesis, Univ. of Montréal); Marie-Victorin, 1934; Raymond,
1950, 1951; Rousseau, 1974) have suggested that post-glacial con-
nections with the Great Lakes may have played a critical role in the
establishment of numerous, often rare elements of southern and
mid-western affinity in the Ottawa Valley flora. Cody (1982) has
examined the distribution of a number of plants in southern Ontario
which show disrupted patterns from the extreme southern part of
the province to north of Lake Ontario. A number of these species
display a fragmented pattern similar to that of Carex oligocarpa.
They are known from few, scattered populations limited essentially
to the Lake Erie-Lake Ontario basins, with outlying colonies occur-
ring northward in the Ottawa Valley. Cody interpreted their distri-
bution in terms of present-day climatic conditions as well as
post-glacial migration corridors. He also suggested possible regres-
sion from more widespread distributions during the more climati-
cally favorable post-Champlain Sea period.

ACKNOWLEDGMENTS

We are grateful to the curators of CAN, DAO and QFA who
made available label information or herbarium specimens originally
identified as Carex oligocarpa from Québec. B. Boivin (QFA) con-
firmed our identification and furnished information regarding prior
misidentifications of C. oligocarpa in Québec. P. W. Ball (TRTE)
kindly provided collection data for the Ontario locations and M. J.
Oldham (Essex Region Conservation Authority) permitted us to
map his recent collections from Pelee Island.

432 Rhodora [Vol. 88

LITERATURE CITED

BALL, P. W. AND D. J. Wuite. 1982. Carex oligocarpa Willd. In: G. W. Argus
and D. J. White, Eds. Atlas of the Rare Vascular Plants of Ontario. Part |. Natl.
Mus. Can.

BoucHarD, A., D. BARABE, Y. BERGERON, M. DuMAIS ETS. Hay. 1985. La phy-
togéographie des plantes vasculaires rares du Québec. Naturaliste can. 112:
283-300.

, ,M. Dumais AND S. Hay. 1983. The rare vascular plants of Qué-
bec. Natl. Mus. Can., Syllogeus 48.

BRUNTON, D. F. AND J. D. LAFONTAINE. 1974. An unusual escarpment flora in
western Québec. Can. Field-Nat. 88: 337-344.

CoppINGTOoN, J. AND K. G. Fie_p. 1978. Rare and endangered plant species in
Massachusetts. U.S. Fish & Wild. Serv., Newton Corner, MA.

Copy, W. J. 1982. A comparison of the northern limits of distribution of some
vascular plant species found in southern Ontario. Naturaliste can. 109: 63-90.

COUNTRYMAN, W. D. 1978. Rare and endangered vascular plant species in Ver-
mont. U.S. Fish & Wildl. Serv., Newton Corner, MA.

Downan, J.J. ANDR. J. CRAIG. 1976. Rare and endangered species of Connecti-
cut and their habitats. State Geological and Natural History Survey of Conn.,
Dept. of Environmental Protection. Report No. 6.

GaGnon, D. 1980. Addition de Claytonia virginica L. a la flore de l’'Outaouais
au Québec. Naturaliste can. 107: 301-302.

1985. Synécologie des plantes vasculaires rares des milieux forestiers de

V’Outaouais central, Québec. Naturaliste can. 112: 333-341.

, AND A. BOUCHARD. 1981. La végétation de l’escarpement d’Eardley,
parc de la Gatineau, Québec. Canad. J. Bot. 59: 2667-2691.

GitteTt, J. M. AND D. J. Wuire. 1978. Checklist of Vascular Plants of the
Ottawa-Hull Region, Canada. Natl. Mus. Can.

MACKENZIE, K. K. 1931-1935. Cyperaceae, Cariceae. North American Flora.
New York Botanical Garden, New York. 18: 1-478.

MARIE-VICTORIN, Fr. 1934. Le fleuve Saint-Laurent, milieu biologique pour les
plantes vasculaires. Proc. & Trans. Roy. Soc. Can. Ser. 3, 28: 1-17.

1964. Flore laurentienne, 2° éd. par E. Rouleau. Les Presses de l’Uni-
versité de Montréal, Montréal.

MenRHOFF, L. J. 1978. Rare and endangered vascular plant species in Connecti-
cut. U.S. Fish & Wildl. Serv., Newton Corner, MA.

RayMonD, M. 1950. Esquisse phytogéographique du Québec. Mém. Jard.
Bot. Montréal 5: 1-147.

1951. Sedges as material for phytogeographical studies. Mém. Jard.
Bot. Montréal 20: 1-23.

Rousseau, C. 1974. Géographie floristique du Québec-Labrador. Distribution
des principales espéces vasculaires. Les Presses de l'Université Laval,
Québec.

ScoGGAN, H. J. 1978. The Flora of Canada, part 2. Pteridophyta, Gymnosper-
mae, Monocotyledoneae. Natl. Mus. Can. Publ. Bot. 7(2): 93-545.

1986] Hay and Gagnon — Carex oligocarpa 433

VARGA, S. 1984. Status and species/habitat management analysis for 16 plant
species; Point Pelee National Park (Carex oligocarpa, pp. 197-209). Parks Can-
ada, Ontario Region.

WaGner, W. H., E. G. Voss, J. H. BEAMAN, E. A. Bourbo, F. W. Case, J. A.
CHURCHILL AND P. W. THOMPSON. 1977. Endangered, threatened and rare
vascular plants in Michigan. Michigan Bot. 16: 99-110.

WHEELER, G. A. 1981. New Records of Carex in Minnesota. Rhodora 83:
119-124.

AND G. B. OwnBey. 1984. Annotated list of Minnesota Carices, with phy-

togeographical and ecological notes. Rhodora 86: 151-231.

S.G. H.

HERBIER MARIE-VICTORIN

INSTITUT BOTANIQUE DE L’UNIVERSITE DE MONTREAL
4101 EST, RUE SHERBROOKE

MONTREAL (QUEBEC)

CANADA HIX 2B2

DG,
DEPARTEMENT DES SCIENCES BIOLOGIQUES
UNIVERSITE DU QUEBEC A MONTREAL, C.P. 8888
MONTREAL (QUEBEC)

CANADA H3C 3P8

A NEW SPECIES OF /LEX (AQUIFOLIACEAE)
FROM HAITI

WALTER S. JUDD

ABSTRACT

A new species, //ex blancheana, is described from a high elevation cloud forest
along the Riviere Blanche, south of Morne la Visite, Massif de la Selle, southern
Haiti. This species is distinguished from other tropical American species of /lex by
the combination of small, conspicuously veined, and entire margined leaves with
obtuse to slightly emarginate apices, and fasciculate inflorescences of three to nine,
four-merous flowers, each with an ovoid and very prominent stigma. The hardwood
communities of the ravines and the steep, more or less north-facing slopes at higher
elevations in the vicinity of Morne La Visite support a diverse and highly endemic
flora, which is known mainly through the field work of Erik L. Ekman.

Key Words: /lex, Aquifoliaceae, flora of Haiti

The species described in this paper was discovered in the Massif
de la Selle of southern Haiti near Morne la Visite. Field work in this
region formed part of an inventory of the flora and fauna of the
recently established Morne la Visite National Park. Discovery of
this species brings to sixteen the number of species of //ex recorded
for Hispaniola (Liogier, 1981). Three other hollies occur in this
region of the Massif de la Selle: //ex cubana Loes., I. fuertesiana
(Loes.) Loes., and /. macfadyenii (Walp.) Rehder. In addition /lex
tuerckheimii Loes. occurs just to the east in the vicinity of Pic la
Selle. A fifth species, //lex obcordata Sw. (= I. formonica Loes.),
occurs to the west in the high elevation cloud forests of the Massif
de la Hotte.

The high elevation cloud forests and pinelands of the Morne la
Visite National Park and adjacent areas have been extensively
explored by Erik L. Ekman (Ekman, 1926; xerographic copy of
unpublished field notes of E. L. Ekman at FLAS), and also were
botanized earlier by W. Buch, Padre E. Christ, and Padre L.
Picarda (Moscoso, 1943). The diverse and highly endemic flora of
the region is in need of additional botanical exploration, however,
as evidenced by the discovery of the species described herein.

'This paper is Florida Agricultural Experiment Station Journal Series No. 6859.

435

436 Rhodora [Vol. 88

Ilex blancheana W. Judd, sp. nov. (Figure 1).

Frutex vel arbor parva ad ca. 4 m alta. Folia elliptica vel obovata,
(0.9-) 1.5-2.9 cm longa, (0.7-) 1.1-2 cm lata, coriacea, glabra, ad
apicem plerumque obtusa, rotundata, vel emarginata, ad basin
cuneata; margo integer, planiusculus vel subrevolutus; nervatura
brochidodroma; petioli 4-6 mm longi. Inflorescentiae axillares, fas-
ciculatae, cum 3 ad 9 floribus. Flores 4-meri. Pedicelli 3.5-6.5 mm
longi, leviter pubescentes. Calyx lobis 0.5-1 mm longis, 0.7-1 mm
latis, abaxialiter pubescentibus. Petala oblonga/ ovata, 2-2.3 mm
longa, 1-1.5 mm lata, alba, leviter pubescentia. Staminodia ca.
1.5-1.7 mm longa. Ovarium globosum, glabrum; stigma conspicuum
et ovoideum. Fructus non visus.

Evergreen shrub or small tree to ca. 4 mm tall with smooth bark.
Indumentum of unicellular nonglandular hairs to 0.1 mm long.
Twigs ridged, sparsely pubescent. Buds triangular, ca. 0.7-1.5 mm
long, with two external pubescent scales. Leaves alternate with min-
ute (ca. 0.5-0.7 mm long) triangular stipules; petiole 4-6 mm long,
adaxially grooved, slightly pubescent, especially on adaxial surface;
blade elliptic to slightly obovate, (0.9-) 1.5-2.9 cm long, (0.7-) 1.1-2
cm wide, flat, coriaceous, the apex obtuse to slightly retuse, rarely
acute, with a short blunt mucro, the base cuneate to broadly so, the
margin entire, plane to slightly revolute along proximal portion of
blade, the adaxial surface lustrous and glabrous, slightly pubescent
on impressed midvein, the abaxial surface glabrous, but very spar-
sely pubescent on extreme proximal portion of midvein, the vena-
tion brochidodromous, adaxially obscure, abaxially conspicuous.
Inflorescences axillary fascicles, the axis 1-2 mm long, glabrous to
sparsely pubescent, bearing 3 to 9 flowers. Flowers imperfect (plants
dioecious), 4-merous, each subtended by a pubescent triangular
bract to | mm long, and paired minute stipules. Pedicels 3.5-6.5 mm
long, very sparsely pubescent; bracteoles 2, alternate to opposite,
triangular, 0.2-0.4 mm long, pubescent, borne from midpoint to
within lower 1/3 of pedicel. Sepals 4, triangular with acute to obtuse
apices, 0.5-1 mm long, 0.7-1 mm wide, slightly connate at base,
adaxial surface sparsely to moderately pubescent, especially near
apex. Petals 4, oblong/ ovate with rounded apex, slighlty connate at
base, 2-2.3 mm long, I|-1.5 mm wide, white, adaxially slightly papil-
lose/ pubescent, abaxially sparsely pubescent. Fertile stamens not
seen; staminodia of carpellate flowers 4, filament ca. | mm long,

1986] Judd — Ilex blancheana 437

Figure |. /lex blancheana: A, habit; B, leaf; C, flower. Drawn from the holotype.

slightly adnate to corolla, anther 0.5—0.7 mm long. Ovary 4-loculed,
globose, glabrous, terminated by a conspicuous, ovoid, slightly 4-
lobed stigma. Drupes not seen.

TYPE: HAITI. Department du L’Ouest: Massif de la Selle, Morne
la Visite National Park, cloud forest on upper slope of ravine of
Riviere Blanche near large waterfall, ca. 2.5 km southwest of “Park
Center”, south of Morne la Visite, ca. 1750-1800 m alt. Uncommon.
Walter S. Judd 4403 (HOLOTYPE, FLAS!; ISOTYPES, A! EHH!, S!).

438 Rhodora [Vol. 88

EtTyMOLoGy: The specific epithet refers to the Riviere Blanche, a
beautiful river flowing through the high plateau of the Massif de la
Selle in the Morne la Visite region, which supports a high diversity
of angiosperm species along its ravine slopes.

Ilex blancheana is easily distinguished from other tropical Ameri-
can species of //lex by the combination of small (i.e., less than 3 cm
long), conspicuously veined, and entire margined leaves with more
or less obtuse to slightly emarginate apices, and fasciculate inflores-
cences of three to nine flowers. In addition the flowers are 4-merous,
and the stigma is ovoid and very prominent. The species is perhaps
most similar to /. berteroi Loes. (incl. var. buchiana Loes.), which
has larger leaves, fewer-flowered inflorescences, and flowers with a
conical stigma. Similarity in stigma form is also evident with /.
fuertesiana. In Liogier’s (1981) key to the hollies of Hispaniola //ex
blancheana would key to the vicinity of /. guianensis (Aubl.)
Kuntze, a species of lower elevations with obovate to oblong leaves
ca. 5-13 cm long, and /. obcordata, a species with only one-flowered
pistillate inflorescences.

The cloud forest (i.e., moist forests of mixed hardwoods) and
forests of Pinus occidentalis Sw. of higher elevations of the Massif
de la Selle are diverse and contain numerous endemics (ca. 34%
endemism among the flowering plants of Morne la Visite National
Park; Judd, unpublished data). Hardwood communities similar to
the one in which //ex blancheana was collected occur along the steep
more or less north facing slopes of the Massif de la Selle from Tete
Opaque, Belle Fontaine, and Pic Cabaio west to Morne la Visite.
This forest extends westward to Morne d’Enfer and eastward
toward Pic la Selle, and shows much local geographic variation.
These cloud forests also extend into the larger ravines (and sink-
holes) in the high elevation pinelands of the Massif de la Selle. The
Riviere Blanche contains the best developed hardwood community
of any ravine in the Morne la Visite region. Woody taxa occurring
with the newly described holly include: Ardisia fuertesii Urb.*', A.
picardae Urb.*, Brunellia comocladiifolia H. & B. subsp. domin-
gensis Cuatr., Buddleja domingensis Urb.*, Bumelia subintegra
Urb. & Ekm.*, Bunchosa ekmanii Urb. & Ekm.*, Calyptranthes cf.
pitoniana Urb. & Ekm.* Cestrum coelophlebium O. E. Schultz, C.
monoeurum Urb. & Ekm.*, C. violaceum Urb.*, Citharexylum

'Species endemic to Hispaniola are indicated by an asterisk.

1986] Judd — Ilex blancheana 439

caudatum L., Coccoloba picardae Urb.*, Daphnopsis crassifolia
(Poir.) Meissn.*, Dendropanax arboreus (L.) Decn. & Planche,
Dendropemon bistriatus Urb.*, Dendrophthora cupressoides
(Macf.) Eichl., Didmyopanax tremulun Krug & Urb.*, Eremolepis
wrightii Griseb., Garrya fadyenii Hook., Gesneria spp., Malpighia
macracantha Urb. & Ndz.*, Meliosma impressa Krug & Urb.*,
Miconia favosa (Desr.) Naud.*, M. ferruginea (Desr.) DC.*, M.
selleana Urb. & Ekm.*, Myrsine coriacea (Sw.) R. Br. ex Roem. &
Schult., Ocotea acarina C. K. Allen*, O. wrightii (Meissn.) Mez,
Oreopanax capitatum (Jacq.) Dene. & Planch., Persea anomala
Britt. & Wils., Picrasma selleana Urb.*, Pinus occidentalis Sw.*,
Rhytidophyllum spp., Solanum crotonoides Lam., Trema micran-
tha (L.) Blume, and Weinmannia pinnata L. There is hope that these
interesting cloud forests can be preserved through establishment of
the Morne la Visite National Park, although at this time clearing
and burning are still occurring in the area.

ACKNOWLEDGMENTS

I thank Dr. Charles Woods (of the Florida State Museum), coor-
dinator and principal investigator of the U.S.A.I.D.-sponsored Bio-
geophysical Inventory of the National Parks of Haiti, who
organized the field trip to the Massif de la Selle, Haiti. Thanks are
also given to Mr. Paul Paryski, Assistant to the Director,
I.S.P.A.N., Port-au-Prince, Haiti, who took me to the locality at
which /lex blancheana was collected. I am grateful to Dr. David W.
Hall for his helpful suggestions concerning the manuscript, Wendy
Zomlefer for preparing the fine illustration, and the curator of the
herbarium of the Swedish Museum of Natural History (S) for loan
of comparative material of /lex.

LITERATURE CITED

EKMAN, E. L. 1926. Botanizing in Haiti. U.S. Naval Med. Bull. 24(3): 483-497.

LioGiger, A. H. 1981. Aquifoliaceae, pp. 7-13. in: Flora of Hispaniola: Part I.
Phytologia Mem. 3: 1-218.

Moscoso, R. M. 1943. Catalogus florae domingensis. Parte I. Spermatophyta. L.
& S. Printing, New York.

DEPARTMENT OF BOTANY
220 BARTRAM HALL EAST

UNIVERSITY OF FLORIDA

GAINESVILLE, FL 32611

ADDITIONS TO THE VASCULAR FLORA
OF KENT ISLAND, NEW BRUNSWICK

ALEXANDER L. MCILRAITH!

ABSTRACT

Ten previously unrecorded species of vascular plants are listed for Kent Island,
New Brunswick.

Key Words: Vascular plants, new records, Kent Island, New Brunswick

Kent Island (45°35’N, 66°46’E) is the largest island of the Three
Islands group located approximately nine kilometers south of
Grand Manan Island, New Brunswick in the Bay of Fundy. Vegeta-
tion patterns on the island were described by McCain (1975) and a
list of the vascular plant species was also published (McCain et al.,
1973).

Documenting the species composition of a small island permits
assessment of changes which may occur over long periods of time.
Hodgdon and Pike (1969) indicated that such records may shed
light on plant dispersal mechanisms and species turnover in island
communities.

During the period from May 7 to August 14, 1984, ten previously
unrecorded species were found on Kent Island. I do not know
whether they were overlooked previously or represent new immi-
grants; it is reasonable to suggest that at least some have become
established in the 15 years since the last plant survey. All the species
have been recorded on nearby Grand Manan Island (Weatherby
and Adams, 1945). Some species may no longer be present on Kent
Island; for example, I did not observe Rosa rugosa or Larix laricina.
Identifications of eight specimens collected were verified by G.
Keleher; these are deposited at WIN. Two species were identified in
the field but were not collected because of their apparent rarity.
Nomenclature follows Kartesz and Kartesz (1980).

SPECIES LIST

Lycopodium clavatum L. (Lycopodiaceae): Collected on a hum-
mock of Pleurozium schreberi under Picea glauca near center of the
island (WIN-41220).

!This paper is Contribution No. 55 of the Bowdoin Scientific Station.

44]

442 Rhodora [Vol. 88

Lycopodium complanatum L. (Lycopodiaceae): Collected in an
old field (South Field) located in center of the island (WIN-41214).

Botrychium matricariifolium (A. Braun ex Doll) A. Braun ex
Koch (Ophioglossaceae): Observed at the southeastern edge of
South Field, close enough to the sea to be exposed to salt spray and
high winds. Roland and Smith (1969) indicated that this species has
been recorded in exposed habitats in Nova Scotia.

Rosa nitida Willd. (Rosaceae): Collected on east side of sphag-
num bog on west side of the island (WIN-41215).

Lathyrus palustris L. var. pilosus (Cham.) Ledeb. (Leguminosae):
Collected on edge of rocky east shore near middle of the island
(WIN-41225).

Acer spicatum Lam. (Aceraceae): Collected in dense woods at
north end of the island (WIN—41219).

Euphrasia randii B. L. Robins. (Scrophulariaceae): Collected at
north-east end of South Field (WIN-41216). McCain et al. (1973)
previously recorded this species only on nearby Sheep Island.

Sambucus racemosa L. var. pubens (Michx.) Koehne (Caprifolia-
ceae): Collected below a bald eagle nest near east shore in open
Picea glauca woods at north end of the island (WIN-41224).

Carex stipata Muhl. ex Willd. (Cyperaceae): Collected in bog on
west shore of the island (WIN-41218). McCain et al. (1973) pre-
viously recorded this species only on nearby Hay Island.

Platanthera lacera (Michx.) G. Don (Orchidaceae). A single indi-
vidual of this species was observed in bloom north of the gull hills at
south end of the island.

ACKNOWLEDGMENTS

I thank Dr. C. E. Huntington for logistic support and J. A. Ger-
vais for field assistance while I was undertaking research on Kent
Island. Financial support was received from an Undergraduate
Research Award of the National Science and Engineering Research
Council of Canada and Dr. R. M. R. Barclay.

1986] Mcllraith — Kent Island, NB 443

LITERATURE CITED

Hopcpon, A. R. ANDR. B. PIKE. 1969. Floristic comparison of three bird islands
in the gulf of Maine. Rhodora 71: 510-523.

KarRTESZ, J.T. ANDR. KARTESZ. 1980. A Synonymized Checklist of the Vascular
Flora of the United States, Canada and Greenland. Vol. II. The University of
North Carolina Press, Chapel Hill.

McCain, J. W. 1975. A vegetational survey of the vascular plants of the Kent
Island group, Grand Manan, New Brunswick. Rhodora 77: 196-209.

, R. B. PikE, AND A. R. HopGpon. 1973. The vascular flora of Kent
Island, Grand Manan, New Brunswick. Rhodora 75: 311-322.

ROLAND, A. E., AND E. C. SmitH. 1969. The Flora of Nova Scotia. The Nova
Scotia Museum, Halifax.

WEATHERBY, C. A. AND J. ADAMS. 1945. A list of the vascular plants of Grand
Manan, Charlotte County, New Brunswick. Contributions No. 158 from the
Gray Herbarium of Harvard University, Cambridge, MA.

DEPARTMENT OF BOTANY
UNIVERSITY OF MANITOBA
WINNIPEG, CANADA. R3T 2N2

THE POLLINATION BIOLOGY OF
CYPRIPEDIUM ACAULE (ORCHIDACEAE)

RICHARD W. Davis!

ABSTRACT

The pollination biology of Cypripedium acaule, the pink lady’s slipper, was studied
by hand-pollinating flowers in the field and by examining pollination and fertiliza-
tion patterns in naturally occurring populations near Amherst, Massachusetts. Fruit
formation in C. acaule appears to be limited by pollinators rather than by other
resources. Pollination intensity appears to be independent of floral density.

Key Words: Cypripedium acaule, orchid, pollination, Western Massachusetts

INTRODUCTION

The stemless pink lady’s slipper, Cypripedium acaule Ait., typi-
cally grows in acidic soil under a fairly open canopy. The plant
consists of two large, oval, basal leaves growing from a rhizome.
The solitary flower is borne on a scape arising between the basal
leaves, and is subtended by a large, leaf-like bract.

Many observers have noted that Cypripedium acaule, even in
dense populations, rarely forms capsules (Correll, 1950; Case, 1964;
Kurfess, 1965). Brackley (1985) states that fruit set in New Hamp-
shire is consistently low, usually less than 10%. Fruit set may be
affected by a lack of pollinators, or by accidents occurring between
pollination and fruit maturation (Correll, 1950). Late frosts could
also lower fruit set (Case, 1964), as could a lack of food resources.
However, the causes of low fruit set have not been studied. The
purpose of this paper is to determine why fruits infrequently occur.

The nature of pollination in Cypripedium acaule promotes out-
crossing (Darwin, 1877). The pollinator enters the flower through a
slit in the labellum, but is unable to exit through this slit because of
its infolded margins. The pollinator exits by first passing under-
neath the stigma and then one of the two lateral anthers. Each
anther contains a sticky mass of pollen, which is attached to the
thorax of the pollinator as a unit by pressure the pollinator exerts
against the anther. Absence of a pollen mass is a reliable indicator of
a visit by a pollinator to a flower (Plowright et al., 1980).

'Present address: 145 Winsor Avenue, Watertown, MA 02172

445

446 Rhodora [Vol. 88

After the pollinator crawls beneath and past the anther it passes
through a small opening at the base of the flower and flies away.
Effective pollinators are too large to enter the flower through this
opening, and thus a visit to a flower is a one-way trip. The pollinator
may then visit and cross-pollinate a second flower.

The pollinators of Cypripedium acaule are bumblebee queens
(genera Bombus and Psithyrus). Although there seem to be no
reports of pollinator visits to C. acaule, two Bombus species have
been identified as pollinators. Stoutamire (1967) identified Bombus
vagans and B. borealis queens as probable pollinators, and Plow-
right et al. (1980) found several B. vagans queens carrying pollen
masses.

Bumblebees are probably attracted to the lady’s slipper flower by
its color and by a sweet, sugary smell concentrated in the sepals and
lateral petals (Stoutamire, 1967). There do not seem to be any
rewards for pollinators. Nectar is not secreted, and bumblebees are
unable to collect the pollen because of the anther’s position on the
flower (R. Davis, personal observation).

The lack of pollinator rewards in a bumblebee-pollinated flower
is of importance in understanding its pollination biology. Bumble-
bees have been shown to avoid flowers lacking rewards after sam-
pling them, in favor of flowers which do offer rewards (Heinrich,
1979). This obviously adaptive behavior on the part of bumblebees
should lead to lower fruit set in deceptive, non-rewarding flowers.
This condition has been shown to be true for other orchid species
not offering food rewards, such as Calopogon tuberosus and
Calypso bulbosa (Ackerman, 1981; Boyden, 1982; Thien and
Marcks, 1972). However, lower fruit set in orchids is offset by the
greater number of seeds produced in a single capsule, which in
Cypripedium acaule ranges from 14,000 to 54,000 seeds per capsule
(Withner, 1959).

It is hypothesized that the formation of large flowering popula-
tions might enhance the reproductive success of Cypripedium
acaule; there might be a positive correlation between floral density
and attractiveness to pollinators, with more pollinators visiting the
denser populations. The denser populations, by massing both visual
and olfactory cues, could be more successful at attracting inexpe-
rienced bumblebees, and increase the number of pollinated flowers
within a population.

1986] Davis — Cypripedium acaule 447

METHODS

On a site near Mt. Holyoke, in Hadley, Hampshire County, Mas-
sachusetts, an unpollinated population of 85 flowers was selected
for experimental hand-pollination with cross- and self-pollen. On 2
June 1983, 75 flowers were cross-pollinated with the pollen obtained
from neighboring flowers, which appeared to be separate individu-
als; single pollen masses were removed with a finger tip and applied
to each stigma. The ten remaining flowers were pollinated with
self-pollen. In April 1984, the number of mature capsules from 1983
was counted; mature capsules successfully overwinter and may per-
sist for at least two years because of the lignification of both the
scape and capsule wall.

Three belt transects were established at this same site in 1983 to
observe pollination and fertilization patterns in natural populations.
Each transect was ten meters wide, beginning and ending at the
borders of each subpopulation. Two of the transects were twenty
meters long, the third ten meters long. In 1984 these same three
transects were used again, along with six new transects at Mt.
Holyoke. Six transects were also established at Rattlesnake Gutter
Road in Leverett, Massachusetts, and three at Teewaddle Hill
Road, also in Leverett, for a total of 18 transects. These transects
ranged from 5 to 22 meters in length.

The number of flowers was counted in each belt transect. Each
flower was inspected for removal of pollen masses; pollen masses
were judged removed if the anther was empty. Evidence of pollina-
tion such as pollen on the stigma or an enlarged ovary and collapsed
labellum was also recorded.

The number of pollen masses removed was counted in 1983 on
9-10 June, when the flowers were rapidly senescing. In 1984 the
counts were made on 9-11 June. In 1983 the number of pollinated
flowers was recorded on 22 June, after all of the flowers had
senesced. In 1984 pollinated flowers were recorded simultaneously
with the counting of the removed pollen masses. Pollinators landing
on or entering flowers were also recorded.

RESULTS

Cypripedium acaule appears to be highly fertile. Out of 75 cross-
pollinated flowers, 56 (75%) set fruit. Seven of the 10 self-pollinated

448 Rhodora [Vol. 88

flowers (70%) also formed capsules. These results are similar to
those obtained by Newhouse (M.S. thesis, Michigan State, 1976), in
which she had 100% fruit set with 4 cross-pollinated flowers and 2
selfed flowers. Newhouse enclosed 8 unpollinated flowers, none of
which formed capsules; autogamy does not seem to occur in the
absence of pollinators.

The results (Table 1) suggest that Cypripedium acaule is infre-
quently visited by pollinators. Out of a total of 278 flowers in 1983,
only 13.3% of the pollen masses were removed, and 9.4% of the
flowers pollinated. In 1984, out of 986 flowers, 6.5% of the pollen
masses were removed and 4.3% of the flowers pollinated. Other
observations supporting infrequent pollinator visits to C. acaule
come from Plowright et al. (1980), who found that only 3%-6% of
the pollen masses were removed from two populations in New
Brunswick.

Table |. Summary of Field Observations, 9-10 June 1983 and 9-11 June 1984.
(1983 transects = 3; 1984 transects = 18.)

Data 7 7
Category 1983 X SD Range _ 1984 xX SD _ Range
Flowers 278 92.7 69.7 26-165 986 54.8 42.1 12-162
Pollen Masses

Removed 74 24.7 11.0 14-36 128 7.1 7.9 0-34

Pollinated Flowers 26 8.7 4.9 3-12 42 2.3 2.6 0-11

Several pollinator visits and pollinators were observed in 1983
and 1984. On 10 June 1983 two pollinators were seen at Mt.
Holyoke, both of them bumblebee queens. One was observed enter-
ing a flower and exiting with a pollen mass on her thorax. The
other, a Psithyrus ashtoni queen, a species not previously known to
visit Cypripedium acaule, was found inside a flower with a pollen
mass on her thorax.

Nine bumblebee queens were seen landing on or entering flowers
on 2 June 1984, all of them at the same site near Mt. Holyoke. Five
entrances and exits by bumblebees were observed. Two of these bees
carried pollen masses into flowers, and deposited them on the stig-
mas of the visited flowers.

Pollination in Cypripedium acaule seems to be density-
independent. No significant relationships were found between C.
acaule flower density and either the percentages of pollen masses

1986] Davis — Cypripedium acaule 449

removed or the percentages of pollinated flowers, using the Spear-
man Rank Correlation Test (R, = —0.1560 and —0.3476, repec-
tively; N = 18).

DISCUSSION

In light of the relatively high fertility of Cypripedium acaule, and
the low frequency of pollen mass removal, capsule formation is
overwhelmingly pollinator-limited. C. acaule thus follows the same
pattern as other deceptive, non-rewarding orchids, which set rela-
tively few fruits but offset this limitation by producing a large
number of seeds within a mature fruit.

Because bumblebees quickly learn to avoid non-rewarding flow-
ers, it is not surprising, in retrospect, that pollination intensity and
floral density should be randomly associated. When low densities of
flowers are as effective as greater densities in attracting pollinators,
what may be of more importance is the density of pollinators in the
foraging range of Cypripedium acaule. If C. acaule flowers are typi-
cally pollinated by “naive” bumblebees unfamiliar with C. acaule,
the number of pollinator visits will depend upon the number of
naive bees in the vicinity; these bees may visit the same number of
flowers in a given area irrespective of floral density.

Othe plants co-flowering in association with C 'ypripedium acaule
could also affect C. acaule fruit set if they attract bumblebees into
the area. Co-flowering plants located in the belt transects included
Maianthemum canadense, Trientalis borealis, Gaylussacia baccata,
Vaccinium angustifolium and V. stamineum. The most wide-spread
associate was M. canadense, which was located in all transects,
sometimes in great density. However, M. canadense is pollinated by
solitary bees and Dipterans (Thaler and Plowright, 1980). One
bumblebee, a worker, was seen foraging on an M. canadense inflo-
rescence on 2 June 1984, but this bee was too small to have been an
effective pollinator of C. acaule. There was some casual evidence,
however, that a positive relationship might exist between the pres-
ence of the ericaceous shrubs, which are bumblebee-pollinated, and
an increased number of pollinator visits to the associated C. acaule
populations; this possible association might be worth further study.

ACKNOWLEDGMENTS

This paper is based in part on an M.S. thesis submitted to the
Department of Botany, University of Massachusetts at Amherst, in

450 Rhodora [Vol. 88

partial fulfillment of degree requirements. I thank J. Ramstetter and
L. Jinishian for field assistance, R. Miller for insect identification,
and E. L. Davis, P. Godfrey, P. Alpert, J. Ramstetter, and two
anonymous reviewers for their comments on this paper.

LITERATURE CITED

ACKERMAN, J.D. 1981. Pollination biology of Calypso bulbosa var. occidentalis
(Orchidaceae): a food deception system. Madrofio 28: 101-110.

BoyDEN, T.C. 1982. The pollination biology of Calypso bulbosa var. americana
(Orchidaceae): initial deception of bumblebee visitors. Oecologia 55: 178-184.

BRACKLEY, F. E. 1985. The orchids of New Hampshire. Rhodora 87: 1-117.

Case, F. W. 1964. Orchids of the western Great Lakes region. Cranbook Inst.
Sci., Bloomfield Hills, MI.

CorreELL, D.S. 1950. Native orchids of North America. Chronica Botanica Co.,
Waltham, MA.

Darwin, C. 1877. On the Various Contrivances by Which Orchids Are Fertilised
by Insects. 2nd ed. D. Appleton & Co., New York.

HEINRICH, B. 1979. Bumblebee economics. Harvard University Press, Cam-
bridge, MA.

Kurress, J. F. 1965. Through the letter slot. Amer. Orchid Soc. Bull. 34(8):
710.

PLOWRIGHT, R. C., THOMSON, J. D. AND THALER, G. R. 1980. Pollen removal in
Cypripedium acaule (Orchidaceae) in relation to aerial fenithrothion spraying in
New Brunswick. Can. Entomologist 112(8): 765-770.

STOUTAMIRE, W. P. 1967. Flower biology of lady’s slippers. Michigan Bot. 6(4):
159-175.

THALER, G. R. AND PLowriGHT, R.C. 1980. The effect of aerial insecticide spray-
ing for spruce budworm control on the fecundity of entompohilous plants in
New Brunswick, Canad. J. Bot. 58(18): 2022-2027.

THIEN, L. B. AND MARCKS, B. G. 1972. The floral biology of Arethusa bulbosa,
Calopogon tuberosus, and Pogonia ophioglossoides (Orchidaceae). Canad. J.
Bot. 50: 2319-2325.

WitHNeR, C. L. 1959. Orchid physiology. Jn: The Orchids: A Scientific Survey,
C. L. Withner, Ed. The Ronald Press Co., New York, 1959.

DEPARTMENT OF BOTANY
UNIV. OF MASS.
AMHERST, MA 01003

RE-EVALUATION AND LECTOTYPIFICATION
OF SCIRPUS RETROFRACTUS L.

RICHARD CARTER! AND CHARLES E. JARVIS

ABSTRACT

The nomenclatural history of Cyperus retrofractus (L.) Torr. is discussed. An
authentic Linnaean specimen is reinterpreted as the lectotype for the basionym Scir-
pus retrofractus L. and arguments are made for the reapplication of this name to a
species which, since 1906, has been called Cyperus dipsaciformis Fernald.

Key Words: Cyperaceae, Cyperus retrofractus, C. dipsaciformis, C. hystricinus,
C. plukenetii, nomenclature, typification

The Cyperus retrofractus complex consists of three North Ameri-
can species for which there are four names available. These species
are widely distributed in xeric habitats throughout much of south-
eastern United States. Kiikenthal (1936) in his comprehensive
treatment of Cyperus placed this complex in section Umbellati
which is composed of perennials with umbellate inflorescences of
simple spikes and few-flowered subterete spikelets. Members of the
Cyperus retrofractus complex are distinguished from other Umbel-
fati by their uniformly retroflexed spikelets and relatively larger
akenes and spikelet scales. This group has had a turbulent nomen-
clatural history (Summarized in Table 1) that began with Linnaeus’
description of Scirpus retrofractus in Species Plantarum (1753) p.
aU:

retrofradus, 17, SCIRPUS culmo triquetro,umbella fimplici:{picarum
flofculis retrotractis.
Cyperi genus indianum, panicula fpeciofa, fpiculis pro-
pendentibus atvis. Pluk. phyt. 41s. f. 4.
Habitat in Virginia. &

In 1805 (p. 375) Martin Vahl treated Scirpus retrofractus under
Mariscus with Scirpus retrofractus L. as a synonym. This treatment
was followed by Elliott in 1821 and again by Torrey in 1836. How-
ever, Torrey (in Gray, 1848) later transferred the species to Cyperus,
thus making the currently accepted combination Cyperus retrofrac-
tus (L.) Torr. There were apparently few problems in applying this

'Present address and that to which reprint requests should be sent: Department of
Biology, Valdosta State College, Valdosta, Ga 31698

451

452 Rhodora [Vol. 88

name until 1906, when Fernald segregated and described two addi-
tional species, thus making it necessary to determine exactly what
Linnaeus meant by Scirpus retrofractus. The original diagnosis is
brief and on its own is of little help. As was usual for his time,
Linnaeus cited no specimens but did cite an illustration by Leonard
Plukenet (tab. 415, fig. 4: 1742) in the synonymy of the species.
Fernald (1906) used this illustration as his basis for application of
the name C. retrofractus (L.) Torr. This interpretation was followed
by Small (1933), by Kiikenthal in 1936 (although he treated the
species as varieties), and by Horvat (1941). In the meantime, how-
ever, Fernald learned that the Linnaean Herbarium (LINN) pos-
sessed a specimen labeled by Linnaeus “17 retrofractus” (no. 71.36;
see Savage, 1945). From a photograph of the specimen, Fernald
could tell it was not the same plant depicted by Plukenet but was
instead, he thought, the related glabrous species he had described in
1906 as C. hystricinus. This left Plukenet’s plant, which had been
called C. retrofractus, without a name; so Fernald in 1945 rear-
ranged his 1906 nomenclature and reduced C. Aystricinus to a syn-
onym of the newly interpreted C. retrofractus (L.) Torr. and
described the plant illustrated by Plukenet as C. plukenetii. In doing
so, Fernald named and described all three species in the complex.

Recently, the Linnaean specimen has been re-examined by us and
determined to be neither the glabrous species that Fernald (1945)
thought it was (previously described as Cyperus hystricinus in 1906)
nor the plant in Plukenet’s illustration cited by Linnaeus (1753).
Instead, it is what since 1906 has been called C. dipsaciformis Fer-
nald, and a member of the only species in the group to which the
name “retrofractus” has not been applied. In light of this unsettling
information, arguments can be made for taking either of two
courses.

First, one could argue that it would be preferable to adopt the
pre-1945 view of Fernald and others, and accept as the type Plu-
kenet’s illustration. This solution has two advantages: (1) since this
approach has already been taken and the species have already been
treated as varieties in this manner by Kiikenthal (1936), it would
prove nomenclaturally more conservative if they are treated as vari-
eties again; (2) the plant in Plukenet’s figure has been unequivocally
attributed by Fernald (1906) to the most morphologically distinc-
tive, central, and most abundant and widespread of the three spe-
cies. By taking this course, the taxonomy would be in greater
concordance with the nomenclature and hence more logical.

1986] Carter and Jarvis — Scirpus retrofractus 453

The other solution is to accept a Linnaean specimen as the lecto-
type of Scirpus retrofractus L. Clarke (1895) argued convincingly
for the use of marked specimens of Cyperaceae at LINN instead of
the sometimes vaguely rendered and often confusing illustrations
cited by Linnaeus. However, each case must be judged on its merits
and the sources of information used in the protologue carefully
weighed. Further, Recommendation T4b of the ICBN (Voss, 1983)
requires that when there is a choice, a specimen should be selected as
lectotype over an illustration. Finally, Article 8.1 states that the
author who first designates a lectotype must be followed. Fernald
did not designate a type in 1906; he wrongly assumed the plate to be
the basis of the name. However, in 1945 Fernald was aware of both
elements and clearly referred to the specimen at LINN as “the type.”

In addition to the previously cited authentic specimen at LINN,
there is at the Linnaean Herbarium in Stockholm (S) a specimen
(IDC Microfiche no. 21.1) bearing the number “17” and the annota-
tion “retrofractus.” However, these annotations are not in Linnaeus’
hand and we therefore do not regard the specimen as a syntype.
Moreover, a photograph of this specimen has been examined by us
and determined definitely not to be in this complex.

Certain other information in Linnaeus’ hand found on the reverse
side of specimen 71.36 (LINN) and pertaining to its identity indicates
that it was sent to Linnaeus by Gronovius (“Gron.”) and is evidently
number “457” of John Clayton. It was therefore collected in Virgi-
nia (Savage, 1945; Stearn, 1957; Reveal, 1983). It is interesting to
note that Linnaeus did not cite this specimen indirectly in the proto-
logue as he did in other instances of Clayton material acquired by
Gronovius, presumably because it seems not to have been cited in
Gronovius’ Flora Virginica (1739).

Plukenet’s illustration is rather questionably matched by a speci-
men in his herbarium, now found in the Sloane Herbarium at BM
(HS 92: 79). However, if that specimen was the basis of the illustra-
tion, significant modifications were made in the form and arrange-
ment of the heads. Linnaeus would not in any case have seen this
specimen and he referred Plukenet’s polynomial to Scirpus retro-

_fractus on the basis of the information available in the illustration.

In light of the information presented here, we believe it best to
follow Fernald (1945) in accepting the specimen no. 71.36 (LINN) as
the lectotype of Scirpus retrofractus L. and to make the necessary
nomenclatural adjustments (Table 1); that is, we apply the name

Table 1. Historical application by various authors of names in the Cyperus retrofractus complex.
Fernald Fernald (1906) Present
Names! Pre-1906 Horvat (1941) Kiikenthal (1936) Fernald (1945) Treatment
“plukenetii” retrofractus retrofractus var. retrofractus plukenetii plukenetii
“hystricinus” ” hystricinus var. hystricinus retrofractus Aystricinus
“dipsaciformis” . dipsaciformis var. dipsaciformis dipsaciformis retrofractus

'Because Fernald used modern type-methods and his names can be unambiguously applied, they are used here as convenient

reference points.

vSP

elopoyy

88 1A]

1986] Carter and Jarvis — Scirpus retrofractus 455

Scirpus retrofractus L. to what has been passing since 1906 as C.
dipsaciformis Fernald, and relegate C. dipsaciformis to the synon-
omy of C. retrofractus (L.) Torr., and re-establish C. hystricinus
Fernald. Below are currently accepted names with their synonyms:

1. Cyperus retrofractus (L.) Torr. in Gray, Man. Bot. North. U.S.
519. 1848.

Scirpus retrofractus L., Sp. Pl. 1:50. 1753. Type: u.s.A. Virginia, Clayton 457.
(LECTOTYPE: no. 71.36 LINN!).

Mariscus retrofractus (L.) Vahl, Enum. Pl. 2:37. 1806.

C. dipsaciformis Fern., Rhodora 8:127. 1906. Type: u.s.A. District of Columbia,
Washington, 22 July 1896, Steele s.n. (HOLOTYPE: GH!; ISOTYPE: US!).

C. retrofractus (L.) Torr. var. dipsaciformis (Fern.) Kiikenthal in Engler, Pflan-
zenreich IV(20)101: 509. 1936.

2. Cyperus hystricinus Fern., Rhodora 8: 127. 1906. TyPE: U.S.A.
New Jersey, near Haddenfield, 13 October 1867, C. F. Parker
5.n. (HOLOTYPE: GH!).

C. retrofractus (L.) Torr. var. Aystricinus (Fern.) Kiikenthal in Engler, Pflanzen-
reich [V(20)101: 509. 1936.

3. Cyperus plukenetii Fern., Rhodora 47:110. 1945. Type: u.s.A.
Virginia, Princess Anne County, Cape Henry, 28 and 29 July
1934, M. L. Fernald and B. Long (HOLOTYPE: GH!)

ACKNOWLEDGMENTS

Much of this paper is taken from a dissertation submitted by
Carter to the Graduate School of Vanderbilt University. The
authors extend thanks to Dr. Robert Kral for his counsel and advice
while Carter was his student at Vanderbilt University, and to Dr.
James Reveal for helping Jarvis in dealing with the Plukenet mate-
rial. Also we wish to thank an anonymous reviewer for helpful
comments and Miss M. J. Keene for carefully typing the manuscript.
Publication costs were met by the Faculty Research Fund of
Valdosta State College.

456 Rhodora [Vol. 88

LITERATURE CITED

CLarKE, C. B. 1895. On certain authentic Cyperaceae of Linnaeus. Journ. Linn.
Soc., Bot. 30: 299-315.

Evxtiott, S.A. 1821. A Sketch of the Botany of South Carolina and Georgia. Vol.
1. Charleston, SC.

FERNALD, M. L. 1906. Some new or little known Cyperaceae of eastern North
America. Rhodora 8: 126-130.

1945. Botanical specialities of Virginia. Rhodora 47: 93-142.

Gray, A. 1848. Manual of Botany of Northern United States. J. Munroe and Co.
Boston & Cambridge, MA.

Gronovius, J. F. 1739. [repr. 1946]. Flora Virginica. Murray Printing Co.
Cambridge.

Horvat, M. L. 1941. A revision of the subgenus Mariscus found in the United
States. Catholic Univ. Amer. Biol. Ser. No. 33. The Catholic Press. Washington,
D.C.

KUKENTHAL, G. 1936. Cyperaceae—Scirpoideae—Cypereae. /n: Engler, A.,
Pflanzenreich IV(20)101; 1-671.1.

LINNAEUS, C. 1753. Species plantarum, Vol. 1. Salvius. Stockholm.

PLUKENET, L. 1742. Amaltheum Botanicum. Vol. IV. London.

REVEAL, J. L. 1983. Significance of pre-1753 botanical explorations in temperate
North America on Linnaeus’ first edition of Species Plantarum. Phytologia 53:
1-96.

SAVAGE, S. 1945. A Catalogue of the Linnaean Herbarium. Linnaean Society of
London. London.

SMALL, J.K. 1933. [repr. 1972]. Manual of the Southeastern Flora. Hafner. New
York.

STEARN, W. T. 1957. An introduction to the Species Plantarum and cognate
botanical works of Carl Linnaeus (introduction to the Species Plantarum fac-
simile of the Ray Society): 1-176. London.

Torrey, J. 1836. Monograph of North American Cyperaceae. Ann. Lyceum Nat.
Hist. New York 3: 249-288.

VAHL, M. 1805. Enumeratio plantarum. Vol. 2. Molleri et Filii. Copenhagen.

Voss, E. G., Ed. 1983. International Code of Botanical Nomenclature. W. Junk.
Boston, MA.

R.C.

DEPARTMENT OF GENERAL BIOLOGY
VANDERBILT UNIVERSITY
NASHVILLE, TN 37235

O59) Se 8

BRITISH MUSEUM (NATURAL HISTORY)
CROMWELL ROAD

LONDON SW7 SBD

UNITED KINGDOM

SYNOPSIS AND ACHENE MORPHOLOGY OF
POLYGONUM SECTION POLYGONUM
(POLYGONACEAE) IN CANADA

STEVEN J. WOLF! AND JOHN MCNEILL

ABSTRACT

Thirteen species and two subspecies are recognized in Polygonum section
Polygonum in Canada. Four achene types are recognized in the section: smooth,
roughened, papillose and striate-papillose. Descriptions, a discussion of comparative
features, distributions, chromosome numbers and a key to the species are presented.
Two new taxa are described: P. caurianum ssp. hudsonianum and P. franktonii.

Key Words: Polygonum section Polygonum, Polygonaceae, synopsis, achene mor-
phology, Canada

In Canada, Polygonum section Polygonum comprises an ex-
tremely variable assemblage of both native and introduced taxa
which occur in a variety of habitats from coastal shores, beaches
and saline marshes to dry, disturbed, weedy habitats along road-
sides, footpaths and wasteplaces. The section is characterized by its
flowers borne in axillary clusters, its jointed petioles and a basic
chromosome number of x = 10. Existing treatments of the section in
Canada are extremely divergent in the number of species that they
include and in criteria used for species delimitation. Fernald (1950)
recognized 12 Canadian species in Polygonum section Polygonum,
one of which, P. douglasii Greene, is no longer inlcuded in the
section (Léve and Live, 1956; Mertens and Raven, 1965). In
addition, Fernald (1950) noted that the name P. aviculare L.
encompassed a vast array of forms, and recognized three varieties in
this taxon. Gleason (1952) recognized six species in the section in
eastern Canada and treated the P. aviculare complex as a single
polymorphic species. In an account of chromosome numbers, Love
and Love (1956) recognized 21 species and a number of infraspe-
cific taxa in eastern Canada, and attempted to clarify the applica-
tion of the name P. aviculare.

In more recent treatments of Polygonum section Polygonum,
particular attention has been focused on defining the limits of the P.

'Present address: Department of Environmental, Population and Organismic
Biology, University of Colorado, Boulder, Colorado 80309.

457

458 Rhodora [Vol. 88

aviculare complex and establishing taxonomic criteria for the delim-
itation of its segregates (Styles, 1962; Mertens and Raven, 1965;
Savage and Mertens, 1968; Jones and Mertens, 1970; McNeill,
1981). Styles (1962) recognized four species in the P. aviculare com-
plex in the British Isles and established the utility of such characters
as heterophylly, fruit size and shape, perianth features and chromo-
some number in distinguishing members of the section. Styles (1962)
concluded that P. aviculare s. str. is a heterophyllous hexaploid
while P. arenastrum Boreau is a homophyllous tetraploid. Mertens
and Raven (1965) extended Styles’ (1962) work to North America
and recognized nine species, which included seven in Canada and
three in the P. aviculare complex: P. aviculare s. str., P. arenastrum,
and P. boreale (Lange) Small. Shortly thereafter, Savage and
Mertens (1968) recognized a fourth species in the P. aviculare com-
plex, P. buxiforme Small. Scoggan (1978) recognized eight species
in Polygonum section Polygonum, suggested that P. aviculare s. str.
is very rare in Canada, and included a large number of synonyms
under P. arenastrum. Recently, McNeill (1981) demonstrated that
both the homophyllous tetraploid P. arenastrum and the hetero-
phyllous hexaploid P. aviculare are very common in eastern Canada
and are readily distinguishable by a number of additional features.
These features include achene size and shape, length of perianth
segments, pollen type, and leaf size and shape.

In the course of McNeill’s (1981) investigation of the Polygonum
aviculare complex, it became apparent that it is comprised of a
number of both native and introduced taxa, some of which have
never received formal taxonomic recognition. It also soon became
apparent that the entire section Polygonum in Canada needed to be
re-examined with respect to the taxonomic criteria employed in the
group, especially in light of more recent cytological data (Moore,
Mertens and Highwood, 1970; Wolf and McNeill, 1986). The inves-
tigation reported here was undertaken in an effort to delimit the
taxa of Polygonum section Polygonum in Canada and to re-
evaluate taxonomic criteria, particularly achene features, that have
been traditionally employed in the group. This study is based on
more than 200 field collections as well as specimens from the follow-
ing herbaria: ACAD, ALTA, BSUH, C, CAN, CU, DAO, GH,
MO, NYS, QUC, TRT, US, VT, WAT and WIN.

1986] Wolf and McNeill — Polygonum 459

MORPHOLOGY AND TAXONOMIC CRITERIA

Although many species of Polygonum section Polygonum appear
superficially similar, they are readily distinguished by a combination
of features of the general habit, leaves, ochreae, fruiting perianth,
and achenes. The following discussions detail morphological varia-
tion and characters of taxonomic significance within the section.

HasitT. All species of the section are herbaceous annuals. Habit
varies from dense, prostrate mats to loosely ascending or erect.
Most species are glabrous; however, Polygonum buxiforme is fre-
quently gray-glaucescent and P. raii Bab. is glaucous. Stem height
varies from 2 cm in P. caurianum Robins. ssp. caurianum up to 2 m
in P. ramosissimum Michx., with such species as P. buxiforme and
P. ramosissimum having somewhat woody and thickened bases.
Other diagnostic features include elongated internodes as in P. ne-
glectum Besser and P. erectum L., branching from the base in P.
caurianum and P. raii, and upcurved branching in P. achoreum and
P. prolificum (Small) Robins.

LEAVES. Several features of the leaves are diagnostic for the var-
ious species. These include leaf length, shape, apex, and petiole
length. Leaf shape varies from linear (P. neglectum) to oblong or
obovate in several species and apices vary from acuminate to acute,
obtuse or rounded. Leaf length varies from 3 mm in P. caurianum
ssp. caurianum up to 60 mm in P. erectum and P. ramossissimum.
Also characteristic of some species is a condition, termed hetero-
phylly, in which leaves of two markedly different sizes occur on the
same plant. In the heterophyllous P. aviculare, P. erectum and P.
ramosissimum cauline leaves are more than three times as long as
leaves on the lateral branches.

OCHREAE. Ochrea length in Polygonum section Polygonum var-
ies from 1-10 mm with most ochreae becoming brown and lacerate
with age. However, in such species as P. buxiforme and P. erectum
they remain hyaline-silvery and intact, and are therefore usually
quite conspicuous and diagnostic for these two species. Ochrea
venation is diagnostic for some species, particularly P. raii with 3-5
nerves. It is sometimes confused with the more southerly P. glaucum
Nutt., which has 8-16 nerves.

460 Rhodora [Vol. 88

FRUITING PERIANTH. Several features of the fruiting perianth
are important in the delimitation of the species of Polygonum sec-
tion Polygonum. Among these are total length, the degree to which
the segments are divided and whether they overlap, the shape of the
perianth tips, and color of the margins. Perianth length varies from
1.6 mm in P. caurianum ssp. caurianum to 5.5 mm in P. raii, with
the length of the longest segment being important in delimiting the
subspecies of P. caurianum. The degree to which the perianth seg-
ments are divided relative to their total length is very useful in the
delimitation of species. In P. achoreum Blake the perianth is divided
one third or less, in P. arenastrum it is divided half way and in the
other sepcies it is divided two-thirds to three-fourths or more. Also,
in such taxa as P. arenastrum, P. neglectum and P. caurianum ssp.
caurianum the segments do not overlap in fruit, while in the remain-
ing taxa they do overlap. Perianth tips in most species are flat and
flared; however, in such species as P. buxiforme, P. erectum and P.
fowleri Robins. they are conspicuously cucullate. Perianth margins
are mostly white to pink within species, with the exception of
yellow-green in P. achoreum, P. erectum and P. ramosissimum,
and purple in P. caurianum ssp. caurianum.

ACHENES. ‘Traditionally, achene characters such as size, color,
shape, and surface texture have been heavily relied upon in the
delimitation of species in Polygonum section Polygonum; however,
some of these characters have never been satisfactorily described or
defined. Additionally, the fact that many species produce elongated,
exserted achenes in late season, in addition to normal achenes, has
led to a proliferation of names in the section. Achene size is diagnos-
tic for many species, particularly those of the P. aviculare complex,
and varies from 1.5-2.2 mm in P. caurianum ssp. caurianum to
3.5-5 mm in P. raii. Most species have brown to dark brown
achenes, with the exception of P. achoreum (yellow-green to tan)
and P. caurianum ssp. caurianum (dark brown to purple). Achene
shape in all species is trigonous; however, the relative width and
shape of the individual sides are diagnostic for most species. Species
such as P. aviculare and P. erectum have three subequal convex
sides; P. buxiforme has three convex sides, one of which is broader
than the other two; such species as P. arenastrum and P. franktonii
S. J. Wolf & McNeill have one narrow concave side and two broad
convex sides. The other species have various combinations of con-
vex and concave sides.

1986] Wolf and McNeill — Polygonum 461

Although achene features have been consistently used in the
identification of the taxa of Polygonum section Polygonum, the
descriptive terminology has heretofore been vague, ill-defined and
sometimes technically incorrect. Such terms as smooth, glossy, shin-
ing, lustrous, sub-lustrous, dull, striate, granular and punctate are
most frequently used in keys and descriptions of the species (Fer-
nald, 1950; Léve and Léve, 1956; Styles, 1962; Mertens and
Raven, 1965; Savage and Mertens, 1968; Mitchell and Dean, 1978;
Scoggan, 1978). We recognize four achene surface types in Poly-
gonum section Polygonum: smooth, roughened, papillose and
striate-papillose. Smooth surfaces, which have no ornamentation,
are characteristic of P. raii and P. ramosissimum (Figures 1A & 1B).
In such species as P. fowleri and P. caurianum the achene surfaces
have no ornamentation, but appear roughened (Figures IC & ID).
The achene surface of P. achoreum has previously been described as
granular; however, the surface is actually densely papillose (Figures
2A & 2B). The most common achene surface type in the section,
striate-papillose, which is characteristic of all species previously
included in the P. aviculare complex, has never been adequately
described. Previously, this surface type has been referred to as dull,
striate or punctate. However, it is evident even under relatively low
magnification that the surface consists of longitudinal rows of papil-
lae (Figures 2C & 2D).

Many species of Polygonum section Polygonum produce elon-
gated, exserted fruits in late season in addition to normal fruits,
produced throughout the growing season. The late-season fruits,
which are 2-3 times as long as normal fruits, are usually olivaceous,
lanceolate, smooth surfaced and have a very thin seed coat. In most
species fruits occur only late in the growing season, i.e., after late
September; however, P. caurianum ssp. hudsonianum S. J. Wolf &
McNeill has a preponderance of them as early as August |. Failure
to distinguish late-season fruits has led to a proliferation of names in
the section. For example, P. exsertum Small is actually P. ramosis-
simum with late season fruits (Gleason, 1952; Mertens and Raven,
1965). Although late-season fruits cause some difficulty in species
identification, normal fruits are also usually present and diagnostic
perianth and vegetative features are still also evident.

462 Rhodora [Vol. 88

Figure |. Scanning electron micrographs of Polygonum achenes. A & B: smooth
type. C & D: roughened type.

KEY TO THE SPECIES OF POLYGONUM SECT. POLYGONUM IN CANADA

|. Achene smooth or somewhat roughened, never papillose.
2. Achene smooth and shiny.

3. Fruiting perianth white, at least at the margins, appearing +
petaloid, spreading at maturity; achenes 3.5-5 mm long,
exserted; plants homophyllous, usually glaucous; mari-
RING SURCIES 6528p ydhwuowunn pax soe cles ees 13. P. raii.

3. Fruiting perianth yellow-green throughout, appressed to
achene at maturity; achenes 2-3.5 mm long, included;
plants heterophyllous, glabrous; inland species, often
WECUY 5 4.00040 00% Leer eer Tee 14. P. ramosissimum.

2. Achenes irregularly roughened and dull.
4. Leaves lanceolate, oblanceolate or linear, 5—12 times as long

as broad.
5. Plants heterophyllous; leaves acute; pedicels 2.5-3.5 mm
long, exserted from the ochreae................00 i

1986] Wolf and McNeill — Polygonum 463

Figure 2. Scanning electron micrographs of Polygonum achenes. A & B: papillose
type. C & D: striate-papillose type.

5. Plants homophyllous; leaves obtuse; pedicels 2 mm long,
included in the OCHICIS seu eine ees 12. P. prolificum.

4. Leaves oblong, ovate or obovate, 2-4 times as long as broad.
6. Achenes 3-4 mm long, beaked, with 3 sub-equal concave
sides; fruiting perianth with cucullate tips enclosing
achene at maturity; leaves 1-5 cm long, elliptic to
elliptic-obovate, acute to obtuse; plants much branched
tHTOURNOUL +6440 vace eee Loy wpe baw 9. P. fowleri.

6. Achenes 1.5-2.5 mm long, beakless, with | narrow concave
and 2 broad + convex sides; fruiting perianth flat or
flared at tip, rarely cucullate, the achene generally
exserted at maturity; leaves less than 1.5 cm long, obo-
vate, rounded to obtuse; plants branching mainly at the

a ee eae neeee ee ees P. caurianum.

464 Rhodora [Vol. 88

7. Achenes 1.5-2.2 mm long; perianth lobes flared at
maturity; plants of Mackenzie District, N. W. T.,
westward to Alaska............ 6. ssp. caurianum.

7. Achenes 2-2.5 mm long; perianth lobes overlapping at
maturity; plants of Hudson’s Bay eastward to the St.
Lawrence estuary ........... 7. ssp. hudsonianum.

1. Achenes papillose, the papillae visible at 30 magnification.
8. Achene uniformly papillose, yellow-green to tan; fruiting peri-
anth divided 1/3 or less to the base ...... 1. P. achoreum.
8. Achene striate-papillose, chestnut to dark brown; fruiting
perianth divided 1/2 or more to the base.

9. Plants heterophyllous.
10. Leaves ovate-lanceolate to narrowly elliptic, acute;
plants prostrate; ochreae brown, lacerate...........
iat Renesas Mee enka eso ena 3. P. aviculare.
10. Leaves elliptic to ovate, obtuse; plants erect; ochreae
oles GA! C21 | - ac cr no ae a 8. P. erectum

9. Plants homophyllous.
11. Leaves linear to lanceolate, 5-9 times as long as

OHNO: 20a sceeoaeneeeoeieeesaws 11. P. neglectum.
11. Leaves oblong, ovate or obtuse, 2-4 times as long as
broad.

12. Achenes with 3 convex sides; perianth tips cucullate;
ochreae silvery, largely entire; lower stems some-
WHALWOOEY ssaxceauvsn eee eens 5. P. buxiforme.

12. Achenes with | narrow concave and 2 convex sides;
perianth tips flat or flared, not cucullate; ochreae
brown, lacerate; lower stems often wiry, but not
woody.

13. Achenes 2.4-2.8 mm long.......... 4. P. boreale.
13. Achenes 1.5-2.2 mm long.

14. Fruiting perianth divided to below the middle, the
segments usually overlapping; leaves narrowly
elliptic, 10-25 mm long; native species of shores
and beaches of eastern Canada..............

ee er ee ee eer ene er 10. P. franktonii.

14. Fruiting perianth divided to the middle or less,
the segments rarely overlapping; leaves elliptic-
oblanceolate, 5-20 mm long, widespread weedy
species of dry disturbed areas ...............

pes kGeS SESS eters wees es 2. P. arenastrum.

1986] Wolf and McNeill — Polygonum 465

1. Polygonum achoreum Blake

Plants erect, homophyllous, glabrous, often whitish from powdery
mildew infection; stems moderately branched, mostly from near the
base, the branches mostly ascending, somewhat woody near the
base, up to 50 cm long. Leaves short petiolate, oval to elliptic, apex
rounded, 8-35 mm long. Ochreae 4-10 mm long becoming brown
and lacerate. Inflorescence 1-3 flowered. Fruiting perianth 2.6-3.5
mm long, divided less than 1/3 to the base, the segments overlap-
ping, the margins yellow to green, the apices cucullate. Achene
2.4-3.3 mm long, yellow-green to tan, uniformly papillose, with |
narrow concave and 2 broad + convex sides, included in the persist-
ent perianth. Late-season fruits common, exserted, up to 5 mm
long, olivaceous, smooth.

DIsTRIBUTION: Very common in disturbed areas such as roadsides
and sidewalks, throughout temperate Canada.

CHROMOSOME NUMBER: 2n = 40, 60.

ACHENE TYPE: Papillose.

Polygonum achoreum is one of the most common and distinctive
members of Polygonum section Polygonum and is almost certainly
native as suggested by Fernald (1950). It is characterized by its
elliptic, rounded leaves; distinctive yellow-green, bottle-shaped
fruiting perianth which is divided 1/3 to the base or less; and its
yellow-green to tan, uniformly papillose achenes which have one
narrow and two broad convex sides. The only species with which it
has been confused is P. erectum. Léve and Léve (1982) considered
P. achoreum a subspecies of P. erectum and Mertens and Raven
(1965) considered it synonymous with the latter. However, except
for the erect habit and broad leaves of these two species, they bear
little resemblance to each other. They differ with respect to achene
size, shape, color and surface texture as well several features of the
fruiting perianth and ochreae. In particular, P. erectum has con-
spicuous silver ochreae, striate-papillose achenes with three concave
sides, and a very deeply divided fruiting perianth. In addition, P.
achoreum is homophyllous while P. erectum is markedly hetero-
phyllous.

2. Polygonum arenastrum Boreau

Plants prostrate, homophyllous, forming dense mats, glabrous;
stems densely branched, 5-50 cm long. Leaves subsessile, elliptic to
oblanceolate, acute to obtuse, 5-20 mm long, all approximately

466 Rhodora [Vol. 88

equal in size. Ochreae to 4 mm long, becoming brown and lacerate.
Inflorescence 2-3 flowered. Fruiting perianth 1.8-2.6 mm long, the
segments divided 1/2 way to the base, rarely overlapping, the mar-
gins white to pink, the apices flat. Achene 1.6-2.2 mm long, dark
brown, striate-papillose, with one narrow concave and two broad
convex sides, the tips slightly exserted from persistent perianth. Late
season fruits common, prominently exserted from the perianth, up
to 4 mm long, olivaceous, smooth.

DISTRIBUTION: Common in such disturbed sites as roadsides,
vacant lots and sidewalks throughout most of temperate Canada.

CHROMOSOME NUMBER: 2n = 40.

ACHENE TYPE: Striate-papillose.

Polygonum arenastrum is the most common member of Polygo-
num section Polygonum and was probably introduced from Europe
(McNeill, 1981). It is characterized by its prostrate, matted habit,
homophyllous leaves, striate-papillose achenes with one narrow
concave and two broad convex sides, and fruiting perianth in which
the segments are divided 1/2 way to the base and do not overlap. It
is readily distinguished from P. aviculare and P. buxiforme by
achene size and shape as well as several features of the fruiting
perianth. Additionally, P. aviculare is heterophyllous and P. buxi-
forme has conspicuous, silver ochreae and a relatively thick and
woody stem.

3. Polygonum aviculare L.

Plants prostrate to sub-erect or loosely spreading, heterophyllous,
glabrous; stems freely branched, up to 2 m long. Leaves short-
petiolate to subsessile, ovate-lanceolate to narrowly elliptic, acute,
the largest on the main stem 25-60 mm long, progressively reduced
on the branches. Ochreae 4-8 mm long, becoming brown and lacer-
ate. Inflorescence 3-6 flowered, mostly restricted to the axils of
upper reduced leaves. Fruiting perianth 2.9-4.0 mm long, the seg-
ments divided 3/4 to the base, overlapping, the margins white to
pink, the apices flat. Achene 2.2-3.0 mm long, dark brown, striate-
papillose, with three + subequal concave sides, the tip + slightly
exserted from the persistent perianth. Late-season exserted fruits
very rare.

DISTRIBUTION: Common on such disturbed sites as roadsides,
vacant lots and sidewalks throughout most of temperate Canada.

CHROMOSOME NUMBER: 2n = 40, 60.

ACHENE TYPE: Striate-papillose.

1986] Wolf and McNeill — Polygonum 467

Polygonum aviculare is a common, introduced weed throughout
most of temperate Canada. Its distinctive features have been enu-
merated in a number of studies (Styles, 1962; Mertens and Raven,
1965; McNeill, 1981). It is a heterophyllous species with striate-
papillose achenes that have three sub-equal sides, and a deeply
divided perianth in which the segments overlap in fruit. Addition-
ally, the flowers are generally restricted to the upper portions of the
branches, in the axils of reduced leaves. The heterophyllous habit,
distinctively shaped achenes and perianth, and distinctive inflores-
cence of P. aviculare readily distinguish it from the other species of
the section, particularly P. arenastrum, with which it is most often
confused.

4. Polygonum boreale (Lange) Small

Plants prostrate, homophyllous, glabrous; stems sparsely branched,
3-25 cm long. Petioles less than 3 mm long, leaves obovate, acute,
5-20 mm long. Ochreae 1-3 mm long, silver-brown, slightly lacer-
ate. Inflorescence 2-5 flowered. Fruiting perianth 2.8-3.5 mm long,
the segments divided 3/4 to the base, overlapping, the margins white
to pink, the apices flat. Achenes 2.4-2.7 mm long, dark brown,
striate-papillose, with one narrow concave and two convex sides,
the tips slightly exserted. Late-season fruits unknown.

DISTRIBUTION: Known from Greenland, but probably also pres-
ent in boreal eastern Canada.

CHROMOSOME NUMBER: 2n = 40.

ACHENE TYPE: Striate-papillose.

We, like Mertens and Raven (1965), are unable to account for this
taxon in North America; however, since it occurs in Greenland, it
can probably be expected to occur in boreal Canada. Fernald (1950)
and Léve and Léve (1956) recognized it in North America; Glea-
son (1952) however, did not. Styles (1962) gave its distribution as
the British Isles, Greenland, Iceland and Northern Scandinavia, and
suggested that North American plants also fit its description. Plants
of this species are relatively diminutive, and some resemble Polygo-
num arenastrum; however, P. boreale differs from P. arenastrum in
its sparsely branched habit, larger fruiting perianth in which the
segments are deeply divided and overlap, and larger fruits. The
name P. boreale has been erroneously applied to a much larger,
heterophyllous taxon that has spathulate, long petiolate leaves, a
much larger fruiting perianth, and much larger achenes which have
three concave sides (Fernald, 1959; Live and Love, 1956; Styles,

468 Rhodora [Vol. 88

1962). However, we have examined Lange’s type of P. boreale and
are unable to account for Styles’ (1962) description of this taxon.
Polygonum boreale sensu Lange is homophyllous with obovate
short petiolate leaves, a smaller fruiting perianth, and smaller
striate-papillose achenes which have one narrow concave and two
convex sides. It is evident that these two taxa are considerably
different and Styles’ (1962) concept of P. boreale is not congruent
with Lange’s original concept of P. boreale. However, since Lange’s
name and type have priority and this species can probably be
expected in Canada, we have included it in this treatment.

5. Polygonum buxiforme Small

Plants prostrate, homophyllous, glabrous to slightly glaucescent;
stems much branched, prominently ribbed, somewhat woody at
lower nodes, up to 2 m long. Leaves distinctly short-petiolate, gray-
glaucescent, subcoriaceous, obovate to oblanceolate, obtuse, 5-25
mm long. Ochreae to 5 mm long, hyaline-silvery, entire to only
slighlty lacerate. Inflorescence 2-6 flowered. Fruiting perianth 2-3
mm long, divided 2/3 to 3/4 to the base, the segments overlapping,
the margins white to pink, the apices cucullate, lower perianth
asymmetric with a pouch-like protrusion. Achene 2.0-2.8 mm long,
dark brown, striate-papillose, with three concave sides, one side
slightly broader and the achene thus appearing heart-shaped, wholly
included in the persistent perianth. Late-season fruits common,
prominently exserted, up to 4.5 mm long, olivaceous, smooth.

DISTRIBUTION: Relatively common in disturbed sites such as
roadsides, vacant lots, and sidewalks throughout most of temperate
Canada, particularly the Prairie Provinces.

CHROMOSOME NUMBER: 2n = 60.

ACHENE TYPE: Striate-papillose.

Polygonum buxiforme is acommon and distinctive native, weedy
species which heretofore had only been recognized in local floras
(Savage and Mertens, 1968; Mitchell and Dean, 1978). It is charac-
terized by its rather stout, somewhat woody stem, gray-glaucescent,
sub-coriaceous, obovate, distinctly petiolate leaves, very deeply
divided perianth which has cucullate apices and a pouch-like pro-
trusion; and striate-papillose, heart-shaped achenes. It is readily
distinguished from P. aviculare by its homophyllous habit and from
P. arenastrum by its distinctively shaped achenes and fruiting peri-
anth. In the Atlantic Provinces, P. buxiforme is sometimes confused
with P. fowleri; however, the latter has roughened rather than

1986] Wolf and McNeill — Polygonum 469

striate-papillose achenes, and occurs on maritime beaches rather
than on dry, disturbed sites.

6. Polygonum caurianum Robins. ssp. caurianum

Plants prostrate, homophyllous, glabrous, frequently purple;
stems filiform, moderately branched mostly from the base, 2-15 (20)
cm long. Leaves short petiolate, greenish-purple, elliptic, rounded to
obtuse, 3-12 mm long. Ochreae less than 3 mm long, brown, lacer-
ate. Inflorescence 2-6 flowered. Fruiting perianth 1.6-2.3 mm long,
the longest segments 1.0-1.6 mm long, divided at least 2/3 to the
base, rarely overlapping, the margins purple, the apices flat and
flared. Achene |.5-2.2 mm long, dark brown to purple, roughened,
with one narrow concave and two broad, + convex sides, exserted.
Late season fruits common, exserted, up to 3.5 mm long, elongated,
olivaceous to purple, smooth to roughened.

DISTRIBUTION: Relatively common on gravel bars and waste
places, Mackenzie District, Northwest Territory westward through
Alaska.

CHROMOSOME NUMBER: Unknown.

ACHENE TYPE: Roughened.

This distinctive taxon is relatively common from the Mackenzie
District, N. W. T. westward through Alaska. It is distinguished by
its diminutive habit, filiform stems, purple color, branching from
the base, relatively small fruiting perianth and small, roughened
achenes. Although it may appear similar to Polygonum arenastrum,
P. caurianum ssp. caurianum is distinguished by its deeply divided
perianth, roughened achenes, purple color and filiform habit. In
addition to geographical distribution, ssp. caurianum is distingh-
ished from ssp. hudsonianum by its smaller stature, achenes, fruit-
ing perianth, leaves and petioles, its filiform stems and _ its
non-overlapping, purple perianth segments with their flat and flared
tips. The only other species in the region that ssp. caurianum could
be confused with is P. fowleri; however, this species has much
larger, beaked achenes with three concave sides and a much larger
fruiting perianth with cucullate tips.

7. Polygonum caurianum Robins. ssp. hudsonianum S. J. Wolf &
McNeill, ssp. nov. TYPE: QUEBEC, CANADA, Fort George, Baie
James, Rampant, terrasse de sable, sec, 30 Aout 1950, E. Lep-
age 12863, DAO (Figure 3).

470 Rhodora [Vol. 88

HELE Ha

*

THRELLUUGOLGM EGE VQGREU AGL OES SE GH

Polygonum caurianum Robinson
subsp. hudsonianum §.J. Wolf &
McNeill

S.J. Wolf 6 J, McNeill 1983
FLORE DU QUEBEC, Canada.
1 ay BE At Tie
: Nem joiygonw: buxiforme Smasd
P A Loco Wort George, Baie Janes, que

Coll, par Vabbé Ernest Lepoge

ident

Figure 3. Holotype of Polygonum caurianum ssp. hudsonianum S. J. Wolf &
McNeill.

Plantae prostratae, isophyllae, glabrae virides usque vix rubentes;
caules aliquantum crassi, modice ramosi pro parte maxima e basi,

1986] Wolf and McNeill — Polygonum 471

6-30 cm longi, internodiis comparate longis. Folia viridia usque
rubentia, elliptica usque angusto-obovata, rotundata usque obtusa,
6-20 mm longa, basi saepe in petiolum distinctum attenuata. Och-
reae minus quam 3 mm longae, hyalinae usque brunneae, vix lacera-
tae. Inflorescentia 2-6 floribus praedita. Perianthium in fructu
2.6-3.3 mm longum, segmentis longis 1.3-2.1 mm_ longissimis,
2/3-3/4 ad basim fissis, imbricatis, marginibus albis usque roseis,
apicibus planis usque vix cucullatis. Achenium 2.0-2.5 mm longum,
fuscum, exasperatum, lateribus duobus latis + convexis et uno
latere angusto concavo, pro parte maxima exsertum; fructus tempo-
ris serotini abundantes, ad 4.5 mm longi, elongati, olivacei usque
brunnei laeves usque exasperati. Chromosomatum numerus ignotus.

Plants prostrate, homophyllous, glabrous, green to slighlty red-
dish; stems relatively stout, moderately branched, mostly from the
base, 6-30 cm long, internodes relatively long. Leaves green to red-
dish, the base frequently tapering to a distinct petiole up to 4 mm
long, elliptic to narrowly obovate, rounded to obtuse, 6-20 mm
long. Ochreae less than 3 mm long, hyaline to brown, slightly lacer-
ate. Inflorescence 2-6 flowered. Fruiting perianth 2.6-3.3 mm long,
the longest segments 1.3-2.1 mm long, divided at least two thirds to
the base, overlapping, the margins white to pink, the apices flat to
somewhat cucullate. Achene 2.0-2.5 mm long, dark brown, rough-
ened, with one narrow concave and two broad + convex sides,
mostly exserted. Late-season fruits very common, exserted, up to
4.5 mm long, elongated, olivaceous to brown, smooth to roughened.

DISTRIBUTION: Relatively uncommon on gravelly shores, Hudson
and James Bays, eastward to Labrador and the St. Lawrence
estuary.

CHROMOSOME NUMBER: Unknown.

ACHENE TYPE: Roughened.

This new subspecies of Polygonum caurianum is relatively
uncommon and occurs from Hudson and James Bays eastward to
Labrador and the St. Lawrence estuary. Although Hultén (1968)
did not recognize this taxon; his distribution map of P. caurianum
from Hudson Bay eastward reflects most of its known distribution.
Polygonum caurianum ssp. hudsonianum also occurs in northern
Quebec and Labrador (see distribution map of P. caurianum, Por-
sild and Cody, 1980). Based on Fernald’s (1950) description and
distribution of P. boreale, particularly in Cote Nord, Quebec and
the shores of Hudson Bay, we suspect that P. caurianum ssp. hud-
sonianum is at least partly included in his concept of P. boreale. In

472 Rhodora [Vol. 88

addition to its geographical distribution, ssp. hudsonianum is dis-
tinguished from ssp. caurianum by a number of chiefly quantitative
features including its larger stature, achenes, fruiting perianth and
leaves, longer petioles, stouter stems and overlapping, white to pink
perianth segments which have slightly cucullate tips. Additionally,
ssp. hudsonianum has a large number of elongated late-season
fruits, even in early August. As Davis and Heywood (1963) note,
subspecies are chiefly differentiated by a number of largely quantita-
tive characters, usually in conjunction with geographical disconti-
nuities. We therefore feel it appropriate to recognize the eastern race
of P. caurianum at the subspecific level.

8. Polygonum erectum L.

Plants erect, heterophyllous, glabrous; stems stout, up to 75 cm
long, sparingly branched, with relatively long internodes. Petioles
distinct, less than 4 mm long; leaves elliptic to obovate, obtuse, the
largest on the main stems 25-60 mm long, shorter on the branches.
Ochreae 5-10 mm long, hyaline-silvery, entire to slightly lacerate.
Inflorescence 2-3 flowered. Fruiting perianth 2.8-3.2 mm long, the
segments divided 3/4 to the base, overlapping, yellow-green, the tips
cucullate. Achene 2.4-3.0 mm long, light to dark brown, striate-
papillose, with three + equal concave sides, wholly enclosed by the
persistent perianth. Late-season fruits uncommon, exserted, up to 5
mm long, olivaceous, smooth.

DISTRIBUTION: A very rare species of waste places across Canada,
but particularly localized in southern Ontario and central Alberta.

CHROMOSOME NUMBER: Unknown.

ACHENE TYPE: Striate-papillose.

Polygonum erectum is a rare but distinctive native species fre-
quently confused with P. achoreum. Most collections of P. erectum
in Canada and the eastern United States are pre-twentieth century,
with a few from the 1940’s and 1950’s. It is distinguished by its erect,
heterophyllous habit; broad leaves; conspicuous, silver ochreae;
deeply divided perianth and striate-papillose achenes. Most herba-
rium specimens labeled P. erectum are actually P. achoreum; for
example, the vouchers for published chromosome counts of P. erec-
tum at BSUH by Moore et al. (1970) are in fact P. achoreum. Love
and Léve (1982) considered these taxa to be related and treated
them as subspecies of P. erectum. Mertens and Raven (1965) could

1986] Wolf and McNeill — Polygonum 473

make no distinction between the two and therefore considered P.
achoreum synonymous with the earlier P. erectum. However,
although superficially similar in habit, the two species differ with
respect to achene shape, size, color and surface texture, as well as in
their ochreae and fruiting perianth. The heterophyllous habit,
striate-papillose achenes with three concave sides, and deeply
divided fruiting perianth of P. erectum contrast with the homophyl-
lous habit, uniformly papillose achenes with one concave and two
convex sides, and slightly divided fruiting perianth of P. achoreum.

9. Polygonum fowleri Robins.

Plants prostrate to ascending, homophyllous, sub-succulent, gla-
brous; stems moderately branched, up to 50 cm long. Leaves short
petiolate, reddish to green, often sub-succulent, elliptic to elliptic-
obovate, acute to obtuse, 10-45 mm long. Ochreae less than 4 mm
long, lacerate, brown. Inflorescence 2-5 flowered, the pedicels
included in the ochreae. Fruiting perianth 3.5-4.5 mm long, the
segments divided 3/4 to the base and overlapping, the margins white
to pink, the apices cucullate. Achene 3.0-4.0 mm long, beaked,
chestnut to dark brown, roughened, with three sub-equal concave
sides, wholly included in the persistent perianth. Late-season fruits
common, much like normal fruits but with only the tip elongated,
exserted, olivaceous to light brown, smooth to roughened.

DISTRIBUTION: Common on sandy or gravelly seashores on both
the west and east coasts of Canda and the St. Lawrence estuary, but
relatively rare in the southern Hudson and James Bay regions.

CHROMOSOME NUMBER: 2n = 40, 60.

ACHENE TYPE: Roughened.

Polygonum fowleri is a common native species of both the west
and east coasts of Canada. It is distinguished by its frequently sub-
succulent habit, elliptic to elliptic-obovate leaves, deeply divided
fruiting perianth with overlapping, cucullate segments, and rela-
tively large, beaked, roughened achenes with three concave sides.
Polygonum fowleriis relatively uncommon in the Hudson and James
Bay regions, where it has been confused with P. caurianum ssp.
hudsonianum. However, the latter is distinguished by its considera-
bly smaller beakless achenes which have one narrow concave and
two convex sides. Polygonum buxiforme is also sometimes confused
with P. fowleri; however, the former has much smaller, beakless,
striate-papillose achenes and conspicuous silver ochreae.

474 Rhodora [Vol. 88

10. Polygonum franktonii S. J. Wolf & McNeill. sp. nov. TYPE:
NOVA SCOTIA, CANADA, Digby County, coast road, St. Ber-
nard to New Edinburg, J. McNeill 5172, 2 Sept. 1976, DAO
(Figure 4).

Plantae erectae, isophyllae, glabrae; caules internodiis comparate
longis, vix ramosi ramis plerumque a basi orientibus, 10-15 cm
longi. Folia angusto-elliptica, acuminata, 10-25 mm longa; petioli
ad 3 mm longi. Ochreae ad 6 mm longae, argenteae usque brunneae
solum vix laceratae. Inflorescentia 2-3 floribus praedita. Perian-
thium in fructu 2.1-2.8 mm longum, segmentis 3/4 ad basin fissis,
imbricatis, marginibus albis usque roseis, apicibus planis. Achenia
2.0-2.4 mm longa, fusca, striato-papillosa, lateribus duobus latis
convexis et uno latere concavo praedita, apicibus vix exsertis. Fruc-
tus temporis serotini ignoti. Chromosomatum numerus 2n = 60.

Plants erect, homophyllous, glabrous; stems with relatively long
internodes, sparingly branched, mostly from near the base, 10-50 cm
long. Petioles up to 3 mm long; leaves narrowly elliptic, acuminate,
10-25 mm long. Ochreae to 6 mm long, silver to brown, sub-
lacerate. Inflorescence 1-3 flowered. Fruiting perianth 2.1-2.8 mm
long; the segments divided 3/4 to the base, overlapping, the margins
white to pink, the apices flat. Achenes 2.0-2.4 mm long, dark
brown, striate-papillose, with one narrow concave and two broad
convex sides, the tips slightly exserted. Late season fruits unknown.

DISTRIBUTION: Maritime and freshwater beaches, dunes and
shores in eastern Canada.

CHROMOSOME NUMBER: 2n = 60.

ACHENE TYPE: Striate-papillose.

This distinctive new species, named for Clarence Frankton, is
restricted to native habitats on beaches and shores in eastern Can-
ada. It is an erect, homophyllous hexaploid with narrowly elliptic
leaves and relatively long internodes. Although quite distinct,
Polygonum franktonii shares some features with both P. aviculare
and P. arenastrum. Its fruiting perianth, although shorter than that
of P. aviculare, is deeply divided and the segments overlap. How-
ever, its achenes, which have one narrow concave and two broad
convex sides, are similar to those of P. arenastrum.

11. Polygonum neglectum Besser

Plants prostrate to ascending, homophyllous, glabrous; stems
slender, flexuous, sparingly branched, up to 50 cm long, internodes

1986] Wolf and McNeill — Polygonum 475

gee eee REET UH a US
Se ee aie ea
t

+

; %
nT TY MT

NedL THAIN aR SS Me

: an nn oe aa a
~—¥-

‘AULA Ug uA gt ne HE
PITTI

—_

NOVA SCOTIA, CANADA

Polygonum franktonii S. J. Wolf and McNeill

Digby County: Coast Rd., St. Bernard to New
Edinburgh. Gravel and rocks just above high
tide level.

J. McNeill 5172 2 September 1976

Figure 4. Holotype of Polygonum franktonii S. J. Wolf & McNeill.

relatively long. Petioles less than 2 mm long; leaves linear-
lanceolate, acute, 8-30 mm long. Ochreae to 4 mm long, silver to
brown, becoming lacerate. Inflorescence 2-3 flowered. Fruiting
perianth 1.9-2.8 mm long, the segments divided 1/2 to 2/3 to the
base, free, the margins white to pink, the apices flat. Achene 2.0-2.3
mm long, dark brown, striate-papillose, with one narrow concave

476 Rhodora [Vol. 88

and two broad convex sides, exserted from the persistent perianth.
Late season fruits common, prominently exserted, up to 4.5 mm
long, olivaceous, smooth.

DISTRIBUTION: Common in such disturbed sites as roadsides,
vacant lots and sidewalks throughout most of temperate Canada.

(CHROMOSOME NUMBER: 2n = 40, 60.

ACHENE TYPE: Striate-papillose.

Polygonum neglectum is a relatively common and apparently
native species that has usually been included in P. arenastrum. Fer-
nald (1950), who recognized three varieties of P. arenastrum, consi-
dered P. neglectum the typical form while Mitchell and Dean (1978)
recognized it at the specific level. Although P. arenastrum and P.
neglectum are somewhat similar, P. neglectum differs by its more
ascending, slender, relatively unbranched stems which have longer
internodes, linear leaves, a slightly longer and more divided fruiting
perianth, and larger achenes. Additionally, since P. arenastrum is
uniformly tetraploid and introduced while P. neglectum is native
and has both tetraploid and hexaploid chromosome races (Wolf and
McNeill, 1986) we recognize it at the specific level.

12. Polygonum prolificum (Small) Robins.

Plants erect to ascending, homophyllous, glabrous; stems with
numerous upcurving branches, 15-80 cm long. Leaves short petio-
late, oblanceolate, rounded to obtuse, 8-30 mm long, green to
bluish-green, rugose-veiny. Ochreae to 3 mm long, silver to brown,
lacerate. Inflorescence 2-4 flowered, the pedicels less than 2 mm
long, included within the ochreae. Fruiting perianth 2.2-2.8 mm
long, the segments divided 3/4 to the base, overlapping, the margins
white to pink, the apices cucullate. Achene 2.5 mm long, included in
the persistent perianth, brown, roughened (occasionally smooth),
with three subequal concave sides. Late season fruits common,
exserted, up to 4 mm long, lanceolate, olivaceous, smooth.

DISTRIBUTION: Uncommon in waste places and saline shores
throughout temperate Canada.

CHROMOSOME NUMBER: 2n = 60.

ACHENE TYPE: Roughened.

Polygonum prolificum is a relatively uncommon native species
which has often been confused with P. ramosissimum. Mitchell and
Dean (1978) considered it a variety of the latter. However, P. ramo-
sissimum is heterophyllous, considerably larger, yellow, has larger

1986] Wolf and McNeill — Polygonum 477

achenes and fruiting perianths, and very long exserted pedicels. In
contrast, P. prolificum is homophyllous, smaller, and has rounded,
veiny leaves, upcurved branches, roughened achenes and the flowers
are not clustered in the upper branch tips.

13. Polygonum raii Bab.

Plants prostrate to erect, homophyllous, glaucescent to glacous
and slightly fleshy; stems sparingly branched, mostly from the base,
up to 75 cm long. Leaves short petiolate, lanceolate to narrowly
elliptic, acute, 10-40 mm long. Ochreae to 5 mm long, brown, 3-5
nerved, lacerate. Inflorescence 2-6 flowered. Fruiting perianth
3.5-5.5 mm long, divided 3/4 to the base, the segments broadly
petaloid and + overlapping, the margins white to pink, the apices
flat and flared. Achene 3.5-5 mm long, chestnut to brown, smooth,
with three subequal concave sides, exserted. Late season fruits
unknown.

DISTRIBUTION: Relatively common on maritime beaches, dunes
and shores throughout Atlantic Canada.

CHROMOSOME NUMBER: 2n = 40.

ACHENE TYPE: Smooth.

Polygonum raii is a distinctive maritime species of beaches and
shores throughout Atlantic Canada. It is distinguished by it glau-
cous habit, very large petaloid perianth segments, and large smooth
achenes. The only species it is likely to be confused with is P. glau-
cum which occurs farther south. However, the latter has narrow
leaves, much smaller achenes and perianth, and much longer (up to
1 cm) 8-10 nerved ochreae.

14. Polygonum ramosissimum Michx.

Plants erect, heterophyllous, glabrous to sub-glaucescent, yellow-
ish; stems moderately branched, prominently ridged, stiff, to 1.5 cm
diameter at the base, to 2 m long. Leaves short petiolate, yellow-
green, lanceolate, acute to acuminate, the largest on the main stem
to 60 mm long, progressively reduced upward. Ochreae to 5 mm
long, silver to brown, lacerate. Inflorescence 2-4 flowered, largely
restricted to upper branches among reduced leaves; pedicels 2.5—3.5
mm long, exserted from the ochreae. Fruiting perianth 2.8-3.7 mm
long, the segments divided 3/4 to the base and overlapping, the
margins yellow-green, the apices cucullate. Achene 2.5-3.5 mm

478 Rhodora [Vol. 88

long, brown, smooth (rarely roughened), with three subequal con-
cave sides, included in the persistent perianth. Late-season fruits
very common, prominently exserted, up to 15 mm long, lanceolate,
olivaceous, smooth.

DISTRIBUTION: Common in waste places, and particularly in saline
soils and shores throughout temperate Canada, especially the
Prairie Provinces.

CHROMOSOME NUMBER: 2n = 60.

ACHENE TYPE: Smooth.

This very distinctive species is common throughout most of Can-
ada, particularly the Prairie Provinces. Its erect, yellow, thick stems
up to 2 meters tall; heterophylly; long, lanceolate leaves; long pedi-
celed flowers restricted to the upper branches; and smooth achenes
distinguish it from the other members of Polygonum section Polyg-
onum. The only species it has been confused with is P. prolificum,
which has been considered a variety of P. ramosissimum (Mitchell
and Dean, 1978). However, P. prolificum is homophyllous, consid-
erably smaller in stature, green, has smaller achenes and fruiting
perianths, and very short pedicels included within the ochreae. The
late season fruits of P. ramosissimum, up to 15 mm long, are the
longest of any species in the section. We concur with Gleason (1952)
and Mertens and Raven (1965) that P. exsertum is a late-season
form of P. ramosissimum.

ACKNOWLEDGMENTS

Financial support from NSERC Canada is gratefully acknowl-
edged. We also thank the curators and staff of the herbaria con-
sulted for supplying loans and/or accommodating visits.

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GLEASON, H. A. 1952. The New Britton and Brown Illustrated Flora of the North-
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versity Press. Stanford, CA.
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1986] Wolf and McNeill — Polygonum 479

Love, A. AND D. Love. 1956. Chromosomes and taxonomy of eastern North
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, AND . 1982. IOPB chromosome number reports. LXXIV. Taxon
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MCNEILL, J. 1981. The taxonomy and distribution in eastern Canada of Polygo-
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STYLES, B. T. 1962. The taxonomy of Polygonum aviculare and its allies in Bri-
tain. Watsonia 5: 177-214.

Wo Lr, S. J. AND J. MCNEILL. 1986. Cytotaxonomic studies on Polygonum sec-
tion Polygonum in eastern Canada and the adjacent United States. Canad. J.
Bot. (in press).

DEPARTMENT OF BIOLOGY
UNIVERSITY OF OTTAWA
OTTAWA, ONTARIO KIN 6N5

RARE VASCULAR PLANTS OF
GROS MORNE NATIONAL PARK,
NEWFOUNDLAND, CANADA

ANDRE BOUCHARD, STUART HAY, CLAIRE GAUVIN
AND Y VES BERGERON

ABSTRACT

In Gros Morne National Park, Newfoundland, 43 vascular plant taxa have been
identified as significantly rare; they are examined in subgroups of species sharing
similar special habitats. These divisions correspond to a general classification of land
regions and biophysiographic areas characterizing the Park. Aspects of range pat-
tern, floristic affinity, ecology and community composition of the populations are
discussed within each subgroup. Information from sample populations is treated by
principal coordinate ordination to illustrate relationships between rare plant com-
munities and abiotic variables.

Key Words: Rare plants, biophysiographic areas, ordination, phytogeography,
Gros Morne National Park, Newfoundland

INTRODUCTION

The west coast of Newfoundland has a well established reputation
for its interesting vascular flora. The importance of this area was
first recognized by Fernald in the early part of this century. His
pioneering explorations led to many important discoveries related
in the pages of Rhodora (Fernald, 1911, 1926-27, 1933) and resulted
in two landmark papers concerning the phytogeography of the
region (Fernald, 1924, 1925). The presence of so many special dis-
junct and endemic elements in the flora of this coast has raised
exciting problems in plant geography which continue to stimulate
discussion and provoke interpretation. Furthermore, when exam-
ined in the greater context of the Gulf of St. Lawrence region, the
west coast of Newfoundland (Figure 1) harbours a high proportion
of these critical taxa. Drury (1969), Morisset (1971) and Rousseau
(1974) have surveyed the problems and hypotheses which have made
this area the focus of attention of botanists and plant geographers
for more than half a century.

Gros Morne National Park was created in 1973 to preserve the
most significant and representative sector of the west coast of New-
foundland (Figure 2). The outstanding landscape features and
important natural resources of the Park (Airphoto Analysis Assoc.,

481

482 Rhodora [Vol. 88

GROS MORNE

Haves so
eo NATIONAL PARK
——_ Comme Brook

Tay
ro a
a 4 ———— =
Figure |. Location of Gros Morne National Park, Newfoundland, in the Gulf of

St. Lawrence

1975; Bouchard, 1975) make this area one of the most distinctive
parts of the Natural Land Region of Canada—Highlands of West-
ern Newfoundland (Falkner & Carruthers, 1974).

Study and protection of rare plants have attracted considerable
attention over the last decade. In North America, both Canada and
the United States have embarked on programs of evaluation and
preservation of their rare native flora. Since 1975, the Botany Div-
ision of the Museum of Natural Sciences of Canada has directed
and published reports for eight provinces. Similarly, in eastern Can-
ada, Forillon National Park, Québec (Morisset, P. 1974, 1979.
Parks Canada report, Ottawa) and Cape Breton Highlands National
Park (Hinds, H. 1982. Parks Canada report, Ottawa) have been the
subject of unpublished reports on their rare vascular flora.

Identification and study of the rare plants of Gros Morne
National Park were undertaken to underline their importance in the
Newfoundland flora and to provide the basis for necessary protec-
tion, future park planning and nature interpretation.

1986] Bouchard et al. — Gros Morne Nat’l Park 483

PREVIOUS FLORISTIC STUDIES

The western coast of Newfoundland has attracted many notable
botanists. Bachelot de la Pylaie, a French naturalist, made two trips
to Newfoundland (Leroy, 1957), the first in 1816 and the second in
1819-20. He spent fifteen days at St. Georges’s Bay and eight days at
Ingornachoix. This first locality is south and the latter is north of
Gros Morne National Park. Unfortunately the vascular plant sec-
tion of his flora was not published, but his important herbarium
collections are kept at the National History Museum of Paris. At
the end of the last century, A. C. Waghorne did some collecting in
regions southeast of the park area. His findings and studies of other
botanists were published in the form of a flora (Waghorne, 1893,
1895, 1898).

In the first part of this century M. L. Fernald directed a series of
botanical trips in western Newfoundland. Within Gros Morne
National Park, his explorations were limited mainly around Bonne
Bay where he made a remarkable contribution to our knowledge of
the flora and was the first to find 20 of the Park’s 43 rare vascular
plants. Many of these rare and localized plants of western New-
foundland and of other areas in the Gulf of St. Lawrence, especially
endemic and disjunct elements, were the basis for his famous
nunatak theory, discussed in “Persistence of plants in unglaciated
areas of Boreal America” (Fernald, 1925).

Another important contribution to the knowledge of the flora of
Newfoundland was made by E. Rouleau of the Université de
Montréal. In addition to collecting herbarium specimens from all
over the province, he compiled previous collection records and pre-
pared distribution maps. He published three checklists of the vascu-
lar plants of Newfoundland (Rouleau, 1949, 1956, 1978); he
discovered two of the Park’s rare vascular plants.

During this same period, other botanists such as Elkington, Jan-
son, Kimball and Penson discovered seven more of the rare vascular
plants in the Park. Creation of Gros Morne National Park favoured
further botanical studies carried on by our research group (Bou-
chard et al., 1977, 1978; Bouchard & Hay, 1974, 1976a, b); discovery
of fifteen of its rare plants resulted from these studies.

METHODOLOGY AND CRITERIA

An approach similar to that developed for the “Rare Vascular
Plants of Québec” (Bouchard et al., 1983) was employed. An

484 Rhodora [Vol. 88

exhaustive listing of all the vascular plants known from Gros Morne
National Park was extracted from the “Flora of St. Barbe South
District” (Bouchard et al., 1978). This information was amended
with more recent revisions and additions. Working from this cata-
logue, each taxon was examined for potential rarity. The great
majority was quickly excluded from further consideration for the
following reasons: common or evidently widespread taxa, intro-
duced alien species or varieties, hybrid taxa, misidentified speci-
mens, and names which have been placed in synonymy with more
widespread or common species. Following this initial screening, all
taxa on the residual list of candidates were evaluated in order to
select the most significant or exceptional rare elements in the flora
of Atlantic Canada. Two general categories exist: (a) those species
of very restricted distribution such as endemic and disjunct ele-
ments, and (b) species at the limit of their native range or species
showing a more widespread distribution, but which occur only in
disparate, localized colonies in Newfoundland.

We consider all of the 43 species treated as rare in Gros Morne to
be rare for Newfoundland. It is noteworthy that 39 of these plants
have also been designated as rare in other provinces, states or Gulf
of St. Lawrence National Parks.

All published information concerning the flora of Newfoundland
was considered in evaluating the rare status of the selected plants.
The distribution maps of the island’s flora, compiled by E. Rouleau
(unpublished files at the Marie-Victorin Herbarium, Université de
Montréal), were especially useful at this stage. Particular attention
was paid to the west coast because of the unique flora associated
with the limestone or serpentine bedrock and the arctic-alpine ele-
ment found on the Long Range Plateau of the Northern Peninsula.

Fieldwork in late July and early August of 1984 permitted us to
locate and sample populations of 39 of these rare plants, with the
objective of describing the community and the habitat for each
population. Some additional sampling information concerning
numbers of colonies, surface area occupied, slope, aspect, and alti-
tude has been compiled (Bouchard et al., 1985a, and CISTI).

Ateach site, soils were sampled for pH, cation exchange capacity,
base concentration, organic carbon, total nitrogen, conductivity,
and exchangeable H and AI following standard procedures
(McKeague, 1978; Thomas, 1982).

‘Depository of Unpublished Data, CISTI, National Research Council of Canada,
Ottawa, Ont., Canada KIA 0S2

1986] Bouchard et al. — Gros Morne Nat’l Park 485

Species are here grouped to correspond to the classification of
general biophysiographic areas of the Park which have been earlier
defined for analysis of the entire vascular flora (Bouchard et al.,
1978, 1985b). Descriptive profiles resuming total range, Newfound-
land distribution, pertinent published maps, general habitat and
rare status in North America for the 43 rare taxa are contained in an
Appendix which has been deposited at CISTI. This Appendix also
includes a complete bibliography and the list of voucher specimens
(Bouchard et al., 1985a), compiled from the following herbaria,
BM, CAN, DAO, GH, MT and NFLD. Nomenclature followed is
taken from the “Vascular Flora of Gros Morne National Park”
(Bouchard et al., 1985b), based largely on Scoggan (1978-79), with
updated synonymy following Kartesz and Kartesz (1980).

To study the relationships between rare plant communities, an
ordination of rare plant stands, based on presence and absence of
associated species, was performed using principal coordinate analy-
sis (Gower, 1966) on a similarity matrix computed with Jaccard’s
coefficient (Legendre & Legendre, 1983). The ordination techniques
were selected because of their performance at high B diversity, as
was the case with our data. The two first axes, representing respec-
tively 10% and 8% of the total variablility, were used for interpreta-
tion. Spearman’s rank correlations (Nie et al., 1975) between abiotic
variables and rare plant stand positions were computed for this
purpose.

RESULTS AND DISCUSSION

The 43 rare vascular plants of Gros Morne National Park are
distributed in the four land regions comprising different biophysio-
graphic units (Bouchard et al., 1978, 1985b) of the Park territory as
follows: Coastal Plain (11), Limestone Escarpments (12), Alpine
Plateau (15), and Serpentine Tableland (5).

Coastal Plain

On the coastal plain, 11 rare vascular plants are found within the
following biophysiographic units: tidal flats and brackish shores,
peatlands, and freshwater areas.

Tidal Flats and Brackish Shores

In this biophysiographic unit, the rare species Solidago semper-
virens L. and Zannichellia palustris L. (halophytic plants with
widespread distributions, particularly southward along the Atlantic

Rhodora [Vol. 88

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1986] Bouchard et al. — Gros Morne Nat’l Park 487

seaboard) reach the northern limit of their distribution and are
known from only a few widely scattered localities. A third rare
species, Elymus virginicus L., is also included in this category
because the Bonne Bay plants are referable to FE. virginicus var.
halophilus (Bickn.) Wieg., a halophytic variety found along the
Atlantic seacoast.

As described by Bouchard et al. (1978), the only saltmarsh, tidal-
flat or well-developed brackish shoreline communities in the Park
occur in the eastuaries around the arms of Bonne Bay and in part of
St. Pauls Inlet. No populations of these plants have been located
north of Bonne Bay.

Of the three rare species, only Zannichellia palustris is a true
aquatic, occurring in tidal pools in the species-poor tidal mud-flats.
Elymus virginicus and Solidago sempervirens occur in sporadic
populations on brackish sand or gravel strands. Despite the fact that
these sites are influenced by sea water, salinity as measured by con-
ductivity (Jackson, 1958) is relatively low. This finding can be
explained by fresh water seepage in the wet shoreline habitats and
by leaching in the dry habitats such as gravel strands. Some stands
seem to be more stabilized with presence of humus accumulation
while others are unstable habitats with low organic accumulation
(Bouchard, et al., 1985a). Colonies are found at very few sites and
the plants occur in low numbers, along with the common character-
istic maritime species Glaux maritima L., Ligusticum scoticum L.,
Plantago maritima L. and Triglochin maritima L. Admixed are the
aggressive widespread adventives Agropyron repens (L.) Beauv.,
Agrostis stolonifera L. and Festuca rubra L.

Peatlands

The six rare vascular plants of the peatlands, Cypripedium regi-
nae Walt., Drosera linearis Goldie, Eriophorum gracile W. D. J.
Koch, Parnassia glauca Raf., Rhynchospora capillacea Torr., and
Salix serissima (Bailey) Fern., are widespread mostly boreal species
that are either transcontinental or have important western ranges
from Alberta or Saskatchewan to Newfoundland. F. gracile is also
found in Eurasia.

The six rare vascular plants were found mainly in eutrophic and
very poorly drained sites such as American larch scrub and sedge
fens (Bouchard et al., 1985a). Although oligotrophic bogs are com-
mon in Gros Morne National Park, eutrophic peatlands are rather

488 Rhodora [Vol. 88

scarce. These plant communities are relatively rich, containing
many species such as Habenaria dilatata (Pursh) Hook. [ Platan-
thera dilatata (Pursh) Lindl. ex Beck var. dilatata], Potentilla fruti-
cosa L., Primula mistassinica Michx., Rhamnus alnifolia L’Heér.,
and Thalictrum pubescens Pursh. Several other eutrophic sites were
sampled which lacked these rare plants. Eriophorum gracile seems
to occupy a more acidic (pH 4.8) microhabitat with associated seep-
age from serpentinic soils (Ca: Mg ratio 0.51; Bouchard et al.,
198Sa).

These six plants are all found within slowly evolving plant com-
munities that should maintain their present composition and
structure.

Freshwater Areas

The two rare vascular plants of the aquatic communities, Pota-
mogeton amplifolius Tuckerm. and Ranunculus hyperboreus Rottb.,
are widespread transcontinental species. R. hyperboreus, which also
occurs in Greenland and Eurasia, has not been observed in Gros
Morne since its discovery in 1940 by Penson. P. amplifolius is a
subdominant of an aquatic plant community dominated by P. epi-
hydrus Raf. and Sparganium angustifolium Michx. [S. emersum
Rehmann] (Bouchard et al., 1985a). This well-established commun-
ity colonizes one shallow acidic pond with a muddy or rocky bottom
located on the coastal plain.

Limestone Escarpments

Gros Morne National Park is especially rich in rare calcicolous
vascular plants. In the limestone escarpment land region are found
Androsace septentrionalis L., Antennaria cana (Fern. & Wieg.)
Fern. [A. alpina (L.) Gaertn. var. media (Greene) Jepson], A. gas-
pensis (Fern.) Fern. [A. neglecta Greene var. gaspensis (Fern.)
Cronq.], Arabis drummondii A. Gray, Arnica tomentosa Macoun
[A. alpina (L.) Olin & Ladau var. tomentosa (Macoun) Cronq.],
Carex petricosa Dewey var. misandroides (Fern.) Boivin, Festuca
rubra L. var. prolifera (Piper) Piper ex Robinson, F. saximontana
Rydb., Gymnocarpium robertianum (Hoffm.) Newm., Sphenopho-
lis intermedia (Rydb.) Rydb. [S. obtusata (Michx.) Scribn. var.
major (Torr.) K. S. Erdman], Woodsia alpina (Bolton) S. F. Gray
and W. glabella R. Br.

1986] Bouchard et al. — Gros Morne Nat’l Park 489

Three species, Androsace septentrionalis, Woodsia alpina, and
W. glabella are circumpolar with southern disjunct populations; the
latter two occur as far south as New York, Vermont and Maine.
Four other species, Arabis drummondii, Festuca rubra var. prolif-
era, F. saximontana, and Sphenopholis intermedia are essentially
transcontinental, with southern stations in United States. S. inter-
media, for instance, occurs as far south as Arizona, Texas and
Florida. Arnica tomentosa is a western alpine species represented by
disjunct populations in Newfoundland. Several of these rare vascu-
lar plants with arctic or cordilleran affinities, such as Androsace
septentrionalis, Arnica tomentosa, Woodsia alpina, and W. gla-
bella, seem to have become isolated from their main distribution
areas, according to the geohistorical scenario discussed by authors
such as Drury (1969), Morisset (1971), Rousseau (1974), Miller and
Thompson (1979) and Marquis (1981). The arctic-alpine species
may have preceded glacial movement southward and survived in
coastal refugia or periglacial tundra zones. The cordilleran species
could have migrated to the eastern part of the continent by a discon-
tinuous tundra corridor at the southern margin of the glacier, as
outlined by Rousseau (1953). In the late-glacial period, climatic and
edaphic conditions may have been suitable for rapid plant migration
(Miller & Thompson, 1979) and widespread colonization by these
arctic-alpine elements on such favourable periglacial habitats as raw
morainic deposits (Rousseau, 1974). With subsequent climatic
change and reinvasion by more competitive boreal flora, disjunct
elements may have survived in relict habitats such as alpine sum-
mits, cold sea cliffs, and unstable serpentine or limestone rock. On
the other hand, Belland (Belland, R. 1981. M. Sc. Thesis, Memorial
University, Newfoundland) has evoked the original nunatak theory
(Fernald, 1925) of survival in ice-free, alpine refugia as a possible
explanation for the presence in Bonne Bay of widely-disjunct,
arctic-alpine and cordilleran bryophytes. He cites recent glacial geo-
logical evidence by Grant (1977a, b) in support of Wisconsin-age
nunataks in the study area.

Four other calcicolous species of these limestone outcrops which
occur in eastern North America are Antennaria cana, A. gaspensis,
Carex petricosa var. misandroides, and Gymnocarpium robertia-
num (s. str.). The latter is an amphi-Atlantic species; the three
former ones are endemic to eastern Canada. Carex petricosa var.

490 Rhodora [Vol. 88

misandroides is restricted to Québec (Gaspé, Mistassini and Arc-
tic) and to Newfoundland (west coast). Antennaria cana and A.
gaspensis, endemics to the Gulf of St. Lawrence area, are found only
in southeastern Québec (Gaspé and Anticosti) or in western
Newfoundland.

Endemism around the Gulf of St. Lawrence has attracted the
attention of botanists since the explorations of Fernald and Marie-
Victorin. Although many of the endemic taxa proposed by these
authors have been discredited by subsequent systematic work, oth-
ers have been maintained as credible species or varieties, isolated by
intense local selection. These Antennaria species illustrate develop-
ment of distinctive endemic characters, even though their taxo-
nomic status may remain questionable. Authors such as Scoggan
(1978-79) have treated them as hardly separable variants, referable
to widespread, polymorphic northern species.

Except for Antennaria cana, a calcicolous species found on a rich
serpentinic site (Ca:Mg ratio 1.1, Bouchard et al., 1985a), the 11
other species are found on limestone cliffs and talus, characterized
by rapidly-drained to well-drained soils. The richness of these rare
vascular plant communities varies generally according to their
stability. For instance, Festuca rubra var. prolifera was found in a
well-stabilized talus community of 38 species dominated by Junipe-
rus communis L., whereas Sphenopholis intermedia was found on
an eroding scree slope with 18 accompanying species. This stability
or lack thereof is reflected in the carbon content of the soils; well-
stabilized sites have a thick humus developed on the underlying
limestone in contrast to less stabilized sites (Bouchard et al., 1985a).
On the other hand, a few well-protected and rather stable microhab-
itats found in the crevices of limestone cliffs are very poor in vascu-
lar plants. For example, Woodsia alpina and W. glabella are found
in stands characterized by only three species each. All these com-
munities include numerous other calcicolous species such as Ane-
mone multifida Poiret, A. parviflora Michx., Arnica lonchophylla
Greene ssp. chionopappa (Fern.) Maguire, Cypripedium calceolus
L., Dryas integrifolia Vahl, Oxytropis johannensis (Fern.) Fern. [O.
campestris (L.) DC. var. johannensis Fern.], Potentilla nivea L., and
Saxifraga aizoon Jacq. [S. paniculata P. Miller].

Most of the rare plants restricted to limestone colonize constantly
disturbed habitats for which they are well adapted. Periodically, a
population will be destroyed by a talus landslip while another will
establish a foothold as new habitats are created. These natural

1986] Bouchard et al. — Gros Morne Nat’l Park 49]

cycles should maintain the presence of these rare vascular plants in
the Park. Future field search may eventually re-discover popula-
tions of Androsace septentrionalis, not observed since its discovery
in 1929 by Fernald and populations of Carex petricosa var. misan-
droides, not observed since its discovery in 1958 by Elkington.

Alpine Plateau

In the alpine plateau land region of Gros Morne National Park,
there are [5 rare vascular plants. All are found in the tundra barrens
biophysiographic unit, which is taken to include special micro-
habitats such as zones of snow accumulation and sheltered alpine
ravines and slopes occurring adjacent to the more extensive,
exposed plateau barrens (Bouchard et al., 1978).

Rare species in the alpine flora include Athyrium distentifolium
Tausch ex Opiz var. americanum (Butters) Boivin, Cassiope hyp-
noides (L.) D. Don, Epilobium anagallidifolium Lam., Gnaphalium
norvegicum Gunn., G. supinum L., Lycopodium alpinum L., Oxy-
ria digyna (L.) Hill, Poa fernaldiana Nannf., Salix herbacea L.,
Saxifraga rivularis L., S. stellaris L. var. comosa Poiret [S. foliolosa
R. Br. var. foliolosa], Sibbaldia procumbens L., Thelypteris lim-
bosperma (All.) H. P. Fuchs, Vahlodea atropurpurea (Wahlenb.)
Fries [Deschampsia atropurpurea (Wahlenb.) Scheele] and Viola
palustris L. Most of these species belong to the arctic-alpine floristic
element, characterized by widespread, arctic-circumpolar distribu-
tions whose ranges extend southward in the eastern part of the
continent to isolated or disjunct alpine habitats such as occur in the
Long Range Mountains of western Newfoundland, on the alpine
summits of the Shickshocks on the Gaspé Peninsula and on the
alpine peaks of New England. Ten of these species, including Cassi-
ope hypnoides, Epilobium anagallidifolium, Lycopodium alpinum,
Oxyria digyna, Salix herbacea, Saxifraga rivularis, S. stellaris var.
comosa, Sibbaldia procumbens, Thelypteris limbosperma and Viola
palustris typically share this general distribution pattern. The/ypte-
ris limbosperma is an exceptional case, in that it shows a striking
east/west disjunction for North America, in an otherwise undis-
rupted circumpolar distribution. The only known colony of this
species, which is isolated in the large gap between western North
America and Europe, was discovered in Gros Morne National Park
and has been the subject of previous papers by Bouchard and Hay
(1976a) and Bouchard et al. (1977).

492 Rhodora [Vol. 88

Three of the rare plants of this biophysiographic unit, Gnapha-
lium norvegicum, G. supinum and Vahlodea atropurpurea, have
boreal to arctic amphi-Atlantic ranges bridging northeastern North
America and northwestern Europe.

Athyrium distentifolium var. americanum is an essentially west-
ern alpine fern with a few, highly disjunct, rare populations in the
Gulf of St. Lawrence region.

Only one rare endemic, Poa fernaldiana, is confined to the tundra
barrens unit of the upland plateau region. This distinctive grass is
rare throughout its restricted range on alpine summits in the Gulf of
St. Lawrence region and New England.

The isolated occurrence of this arctic or cordilleran flora in dis-
junct, alpine outposts of eastern North America has been discussed
in connection with similar elements which occur as disjunct popula-
tions on the limestone escarpments in the park. An alternative
hypothesis, that of long-distance dispersal to disjunct eastern sta-
tions, cannot be disregarded for the mainly cordilleran ferns Athy-
rium distentifolium var. americanum and Thelypteris limbosperma
(Bouchard et al., 1977). Wagner and Rouleau (1984) suggested this
same possibility for another rare cordilleran fern in Newfoundland,
Polystichum X scopulinum D. C. Eaton.

All of the 15 rare species studied in this biophysiographic unit
have been recorded from snowbed zones (zabois) or more sheltered
alpine ravines and slopes, which contrast strikingly with the exposed
tundra vegetation on the adjoining plateau barrens. Damman (1983)
emphasized that species such as Cassiope hypnoides, Oxyria digyna
and Salix herbacea are restricted to the snowbeds of the Northern
Long Range and similar sites in the Serpentine Range of the West-
ern Newfoundland Ecoregion. The cool, seepy conditions of these
habitats favour richer herbmat communities often dominated by
many sub-alpine species such as Clintonia borealis (Ait.) Raf., Cop-
tis groenlandica (Oeder) Fern. [C. trifolia (L.) Salisb. ssp. groenlan-
dica (Oeder) Hultén], Cornus canadensis L., Deschampsia flexuosa
(L.), Trin., Dryopteris spinulosa (O. F. Muell.) Watt and Solidago
macrophylla Pursh. More characteristic of these sites is the
arctic-alpine flora composed of relatively abundant species such as
Carex bigelowii Torr., C. stylosa C. A. Meyer, Diapensia lapponica
L., Juncus trifidus L., Loiseleuria procumbens (L.) Desv. and
Phyllodoce caerulea (L.) Bab., which often grade into these sites
from the exposed tundra on the adjoining plateau barrens. Except

1986] Bouchard et al. — Gros Morne Nat’ Park 493

for the Cassiope hypnoides colony found on the gabbro talus, the
soils sampled are relatively acidic with moderately decomposed
organic accumulation (Bouchard et al., 1985a).

Belland (1983) has reported similar findings for the widely-
disjunct bryophyte flora which is also restricted to the alpine
snowbed communities in Gros Morne. The special ecological condi-
tions created in these micro-habitats due to accumulation of late-
lying snow and a significantly shorter growing season have been
studied in comparable situations in northern Québec (Filion &
Payette, 1982) and on the Gaspé Peninsula (Boudreau, F. 1981.
M. Sc. Thesis, Université Laval, Québec). The rare arctic-alpine
plants are well adapted to this micro-environment and they may
illustrate a relict flora which has persisted since the late-glacial
period.

Unquestionably, the highland plateau region of the Park remains
relatively unexplored botanically, a fact which may explain the con-
centration of apparently rare species based on what we presently
known of the flora. Two rare species included here, Lycopodium
alpinum and Thelypteris limbosperma, were new additions to the
flora of Newfoundland (Bouchard & Hay, 1974, 1976a; Bouchard et
al., 1977). Two other remarkable arctic-alpine plants, Saxifraga stel-
laris var. comosa and S. rivularis, are newly reported for St. Barbe
South District, previous records being known only from the tip of
the Northern Peninsula. Finally, another three species, Athyrium
distentifolium var. americanum, Epilobium anagallidifolium and
Poa fernaldiana constitute new records for the Gros Morne
National Park area of St. Barbe South District. With further explo-
ration, perhaps other arctic-alpine species will be added to the flora
of the Park. [René Belland found an additional rare plant, Ranun-
culus pedatifidus Sm., in an alpine ravine on Gros Morne Mountain,
27 August 1984 (substantiating specimen deposited at NFLD, pers.
comm., 24 January 1985).] By the same token, some of the species
presently considered to be rare may prove to be more frequent
throughout the alpine plateau region.

Serpentine Tableland

The Tableland is treated as a distinct land region comprising one
biophysiographic unit (Bouchard et al., 1978, 1985b), including the
altered gabbro outcrops and talus of the mafic highland which
intergrade with the serpentine massif.

494 Rhodora [Vol. 88

The five rare vascular plants of the Tableland are Arenaria mar-
cescens Fern. [Minuartia marcescens (Fern.) House], Danthonia
intermedia Vasey, Eleocharis nitida Fern., Festuca altaica Trin. in
Ledeb. and Salix arctica Pallas. Three of these, Danthonia interme-
dia, Eleocharis nitida, and Festuca altaica, are mostly boreal species
occurring transcontinentally in widely-scattered, isolated localities.
Populations of the two grasses become more frequent towards the
western part of their range, whereas Eleocharis nitida is more widely
represented in the east. Salix arctica has a continuous circumpolar
distribution with isolated populations occurring southward, in the
eastern part of the continent, on the serpentine massifs of western
Newfoundland and on the Gaspé Peninsula. Arenaria marcescens
is an endemic taxon of northeastern North America. It is rare
throughout its restricted range, being confined to serpentine expo-
sures of western Newfoundland, the Gaspé Peninsula and Ver-
mont. It is perhaps better interpreted as a disjunct variety of
cordilleran affinity to the western species A. obtusiloba (Rydb.)
Fern. [ Minuartia obtusiloba (Rydb.) House].

Of these rare species, only Arenaria marcescens is found exclu-
sively on serpentine substrate. Salix arctica, although restricted to
this habitat in the Gulf of St. Lawrence, is not strictly serpentinico-
lous elsewhere in its range. Similarly, although they are known
chiefly from this area in the Park, the other three species may colon-
ize different types of rocky or peaty barren.

Arenaria marcescens is a frequent component of the sparse vege-
tation colonizing open rock and gravel polygon barrens. Character-
istic species recorded from the sample stand include Armeria
maritima (P. Miller) Willd. ssp. /abradorica (Wallr.) Hultén,
Carex scirpoidea Michx., Rhododendron lapponicum (L.) Wahlenb.,
Salix glauca L., Senecio pauperculus Michx. and Silene acaulis (L.)
Jacq. (Bouchard et al., 1985a). The rare Danthonia intermedia and
Festuca altaica are found on screes and talus in the protective cover
of dense, stunted scrub communities dominated by tolerant woody
species such as Juniperus communis, J. horizontalis Moench, Larix
laricina (Du Roi) K. Koch and Potentilla fruticosa. The sample
population of Eleocharis nitida was found in an open, gravelly seep
on a serpentine moraine at the base of the Tableland. The Mg
concentration and the Ca:Mg ratio (Bouchard et al., 1985a) of
sampled sites for four of the rare species is consistent with those
reported for serpentine soil throughout the world (Proctor &
Woodwell, 1975). Although Salix arctica was collected in 1973 and

1986] Bouchard et al. — Gros Morne Nat’l Park 495

1978, we were unable to re-locate stands of this rare willow in 1984.

The five rare species are well established in a diversity of habitats
unique to the Tableland. The sparse vegetation of these barren was-
telands and their unique ecology have been discussed for Gros
Morne National Park by Bouchard et al. (1978) and Dearden
(1979). Because of the restrictive ecological factors, the serpentine
flora, including the rare species, is highly adapted to this particular
environment. Arenaria marcescens is quite common on the Table-
land, but populations of the four other species are few in number.

The cordilleran fern. Adiantum pedatum L. ssp. calderi Cody [A.
pedatum var. aleuticum Rupr.], although restricted to serpentine in
Newfoundland, is fairly widespread and frequent in its habitat and
was not treated as rare in the Park. It may still warrant considera-
tion in a forthcoming study of Newfoundland’s rare plants.

ORDINATION OF RARE PLANT STANDS

To present an overview of the rare plant communities and to
clarify their relationships to abiotic variables, a principal coordinate
ordination of the data (Bouchard et al., 1985a, and CISTI) was
performed (Figure 3). Five groups are easily identified: the coastal
plain land region comprising the tidal flats and brackish shores, the
coastal plain peatlands; the limestone escarpments land region; the
alpine plateau land region; and the serpentine tableland land region.
Three abiotic variables are correlated with the first axis: pH —.5483
(p < .001); Mg —.4144 (p < .01); altitude .6185 (p < .001) and four
with the second axis: cation exchange capacity .4767 (p < .05);
Mg .5043 (p < .05); Na .4698 (p < .05); drainage .6350 (p < .001).

On the first axis, soil pH and altitude are important abiotic varia-
bles. The oligotrophic sites are on the right, especially associated
with the alpine plateau land region, while the more eutrophic sites
are on the left with the limestone escarpment land region and the
serpentine tableland land region. The rare plant stands on peatlands
occupy an intermediate position because they are not found on
oligotrophic peats, which are abundant in the Park, but are con-
fined to restricted eutrophic sites (fens). The tidal flat stands fall in
the center of the ordination because they share few species with the
other plant communities. On the right are found the high altitude
arctic-alpine plant communities. Altitude is also an important factor
contributing to the presence of rare plants in the limestone escarp-
ments land region. Within this land region, the four stands on the

496 Rhodora [Vol. 88

296 axis

SERPENTINE @ #33
TABLELAND

\\ COASTAL *2
PLAIN

peatiands |
weed
NED ALPINE PLATEAU

tidal flats
and
brackish shores

Ae

COASTAL .27
PLAIN

fo

LIMESTONE ESCARPMENTS

1S axis

Figure 3. Principal coordinate ordination and biophysiographic affinities for the
rare plant communities. Stand numbers refer to sample populations based on data in
Bouchard et al. (1985a, and CISTI).

right (21, 23, 24 and 25) are all situated at an elevation higher than
600 m while the other stands are usually at an elevation of less than
100 m (Bouchard et al., 198Sa).

On the second axis, drainage is an important abiotic variable. The
rare plant stands on limestone, characterized by rapidly-drained to
well-drained soils, fall in the lower part of the ordination, while the
stands on very poorly-drained, organic soils are in the middle and
upper part. These organic soils are also characterized by a higher
cation exchange capacity. The magnesium content of the serpentine
soils accounts for the segregation of the Tableland stands from
those of the limestone escarpments land region. Some integration

1986] Bouchard et al. — Gros Morne Nat’l Park 497

exists between stands on these latter basic substrates because these
open unstable habitats, characterized by their low plant cover, are
colonized by many of the same pioneer species.

The ordination graphically illustrates a clear segregation of rare
plant colonies of Gros Morne National Park according to commun-
ity composition and important abiotic variables. Occurrence of
many of the exceptional species in the Park can be directly attrib-
uted to the presence of special restricted habitats which are reflected
in the ordination.

CONCLUSION

Gros Morne National Park has an important heritage of rare
vascular plants. We have treated as rare 43 taxa among a total
vascular flora of about 700 species. Despite the somewhat subjective
nature of what constitutes rarity, these taxa are considered to be the
most significant examples because they are species which are rare
not only within the Park territory, but they have very few known
populations in Atlantic Canada.

Rare plants owe their status to a complex variety of interrelated
geohistorical, genetic and ecological factors (Brouillet, 1985; Drury,
1980; Stebbins, 1980). In Gros Morne National Park, they are for
the most part restricted to special, localized habitats such as tidal
flats, limestone escarpments or talus, serpentine barrens, eutrophic
peaty areas and alpine snowbeds. Stebbins (1980) described this
concept as “ecological islands” of isolated or marginal populations
of rare species established and persisting on special, restricted
habitats.

The primary importance of these rare plants lies in the complex
biogeographic problems and interpretations which they pose. For
example, Antennaria gaspensis, Arenaria marcescens and Poa fer-
naldiana are rare endemics of northeastern North America. The
isolated populations of Arnica tomentosa, Athyrium distentifolium
var. americanum and Thelypteris limbosperma show remarkable
eastern disjunctions from their main range of distribution in the
western part of the continent. The arctic-alpine species Salix herba-
cea, Saxifraga rivularis and Sibbaldia procumbens occur isolated
far to the south of their arctic-circumpolar ranges on a few alpine
summits such as the Long Range Mountains. The peripheral popula-
tions of Elymus virginicus and Solidago sempervirens, occurring at

498 Rhodora [Vol. 88

the northern limit of their native range, appear to be rare in New-
foundland. Finally, a few species such as Danthonia intermedia,
Drosera linearis and Eriophorum gracile, though widely dispersed
continentally, are rare in Newfoundland and several other provinces
or states.

The only practical means of protecting these rare plant popula-
tions is to preserve the special habitats where they occur. Except for
Arenaria marcescens, which is relatively common on the serpentine
barrens, few populations of these rare species are known in the
Park. Some extreme cases such as Androsace septentrionalis, Carex
misandroides var. petricosa, Eriophorum gracile and Thelypteris
limbosperma are known from only one population, and some have
not been reobserved for many years. Fragile habitats such as the
limestone cliffs or talus and the alpine snowbeds must receive par-
ticular attention if they are to be preserved.

Bratton and White (1981) discussed the problems involved in
management of rare or endangered plant populations after the
initial steps have been taken to establish a preserve. For Gros
Morne National Park, a management or monitoring program of the
populations seems neither necessary nor advisable because of the
remoteness of the rare plant colonies. Attempts at management
could in fact be more detrimental than beneficial, especially where
the habitats are particularly fragile. The authorities should, how-
ever, protect the sites and show prudence about revealing their exact
locations. Nevertheless, they should facilitate population studies
such as that done on Cypridepium passerinum Rich., in Pukaskwa
National Park (Keddy et al., 1983).

The intrinsic appeal of rare plants should not be overplayed by
the nature interpretation program of the Park. With the exception
of the Showy Lady’s-slipper, the rare plants of Gros Morne
offer little visual appeal to all but the most ardent botanist. Rather,
the accent should be placed on the fascinating geohistorical expla-
nations for the establishment and persistence of these species, and
the biogeographical problems raised by the persence of such a diver-
sity of endemic, peripheral and disjunct elements in the Park’s vas-
cular flora.

ACKNOWLEDGMENTS

We wish to thank particularly E. Rouleau (MT) for making avail-
able his collection records and distribution maps of the Newfound-
land flora. Specimen verification and taxonomic assistance came

1986] Bouchard et al. — Gros Morne Nat’! Park 499

from S. Aiken (CAN), D. Britton (OAC), and P. Hoch (MO). Cor-
respondance with G. Argus (CAN), M. Chater (BM), W. Cody
(DAO), S. Downie (ALTA), J. McNeill (OTT), J. and A. Reddoch
(CAN) and S. Wolf (MO) helped in the evaluation of various taxa.
The suggestions by G. Argus, K. Pryer and two anonymous review-
ers were helpful in improving the manuscript. Soil samples were
analyzed by C. Camiré, Université Laval. Computer processing
work was done by A. Leduc, Université de Montréal. Typing of
the manuscript was done by C. Blanchard, Y. Bourget, E. Lemaire
and H. Levert. We are also grateful for field assistance from R.
Belland, J.-L. Bourdages and F. Tremblay. Financial assistance
came from Parks Canada (research contract GM-83-20) and D.
LeSauteur facilitated access to the Park.

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502 Rhodora

A. B.

JARDIN BOTANIQUE DE LA VILLE DE MONTREAL
4101 EST, RUE SHERBROOKE

MONTREAL (QUEBEC) CANADA HIX 2B2

A. B., S. H.

INSTITUT BOTANIQUE DE L'UNIVERSITE DE MONTREAL
4101 EST, RUE SHERBROOKE

MONTREAL (QUEBEC) CANADA HIX 2B2

c.G,

CENTRE DE RECHERCHES ECOLOGIQUES DE MONTREAL
UNIVERSITE DE MONTREAL

5858 CHEMIN COTE-DES-NEIGES, BUREAU 400

CASE POSTALE 6128, SUCCURSALE “A”

MONTREAL (QUEBEC) CANADA H3C 3J7

Y. B.
DEPARTEMENT DE SCIENCES BIOLOGIQUES

UNIVERSITE DU QUEBEC A MONTREAL, C. P. 8888, SUCC. A,

MONTREAL (QUEBEC) CANADA H3C 3P8

[Vol. 88

NEW MESOAMERICAN SPECIES OF
CYPERUS (CYPERACEAE)!

GORDON C. TUCKER

ABSTRACT

Four species of the genus Cyperus are described, illustrated, and compared to their
nearest relatives. These are: C. matudae and C. breedlovei (Subgenus Protocyperus)
from the Pacific slope of Chiapas, Mexico; and C. wilburii and C. svensonii (Subge-
nus Cyperus) from the lowlands of the Isthmus of Tehuantepec, and from the
uplands of Chiapas, Guatemala, Honduras, and Nicaragua, respectively.

Key Words: Cyperaceae, Cyperus, new species, Mexico, Central America

Four new species of Cyperus have been detected in a study of the
genus in Mexico (Tucker, G. C. 1985. Unpubl. Ph. D. thesis, Duke
University, Durham, N. C.). These new species are members of the
two largest subgenera of Cyperus: subgenus Protocyperus K. A.
Lye, in which the plants have trigonous achenes, digitate clusters of
spikelets, and eucyperoid (non-Kranz) anatomy (Lye, 1981); and
subgenus Cyperus, in which the plants have trigonous achenes, spi-
cate inflorescences, and chlorocyperoid (Kranz) anatomy (Raynal,
1973). A detailed account of the characteristics of the subgenera
appeared in Kiikenthal (1935-36) and Tucker (1983).

Cyperus matudae G. C. Tucker sp. nov. TYPE: MEXICO. Chiapas, Mt.
Ovando, pine land, 1250-2370 m, July 1938, Matuda 2553
(HOLOTYPE: MICH; ISOTYPES: F, GH, LL, NY). (Figure 1).

Plantae perennes, 50-75 cm alta. Culmi triquetri, scabri in angu-
lis, 1.3-2.5 mm crassi. Bracteae 6-8, 3-30 cm longae, 2-10 mm latae.
Radii 10 vel 11, ad 15 cm longi; radii secundarii ad 4 cm longi; radii
tertiarii ad 15 mm longi. Spiculae digitatae, lineari-lanceolatae,
compressae, 7-15 mm longae, 2.3-3.5 mm latae. Squamae deciduae,
oblongo-ellipticae, 2.2-2.4 mm longae, cuspidae apicali 0.4-0.8 mm
longo. Stamina tria. Stigmata tria. Achenia trigona, ellipsoidea,
1.8-2.0 mm longa, circa 1.2 mm lata. Cypero chorisantho C. B.
Clarke affinis.

Perennials, 50-75 cm tall. Rhizomes horizontal, indurate, clothed

‘Contribution number 495 from the New York State Science Service.

503

504 Rhodora [Vol. 88

Figure |. Cyperus matudae. a. plant; b. spikelet; ¢. scale (abaxial view); d.
achene.

1986] Tucker — Cyperus 505

with fibrous leaf bases. Culms triquetrous, scabrous on the angles
only, especially above, 1.3-2.5 mm thick. Leaves flat to subtly pli-
cate, about as long as the culm, 5-15 mm wide, the margins and
keels scabrellate. Inflorescence bracts 6-8, 3-30 cm long, 2-10 mm
wide, vertical to ascendent at (60) 75°. Rays 10-11, up to 15 cm
long; secondary rays 3-8 per primary ray, 10-40 mm long; tertiary
rays 3-15 mm long. Spikelets digitate in groups of 1-3, linear-
lanceolate, 7-15 mm long, 2.3-3.5 mm wide (including the apices of
the scales), greenish brown, deciduous at maturity; rachilla slightly
flexuous, wingless, persistent. Scales 8-16, oblong-elliptic, 2.2-2.4
mm long (the excurved cusp 0.4-0.8 mm long), 1.4-1.6 mm wide,
laterally 3-4 nerved, reddish to brownish green, medially 3-7
nerved, the midvein green, smooth or scabrellate distally. Stamens
3; anthers 0.9-1.2 mm long, the setose apices of the connectives an
additional 0.1-0.2 mm long. Styles 0.6-0.8 mm long; stigmas 3, (1.2)
1.6-2.6 mm long. Achenes trigonous, ellipsoid, 1.8-2.0 mm long,
about 1.2 mm wide, sessile, apiculate, papillose, brown.

Endemic to southern Chiapas, Mexico, Cyperus matudae is
known from only two collections made near the Guatemalan
border. Plants with mature achenes have been collected in July and
August. Cyperus matudae is placed in subgenus Protocyperus K. A.
Lye, because of its trigonous achenes and digitately arranged spike-
lets. Both known collections of this new species had been identified
previously as “C. diffusus Vahl” [= C. laxus Lam.], a widespread
neotropical species (Tucker, 1983). Cyperus matudae differs strik-
ingly from C. /axus in features of the achenes and scales, and in its
more highly branched inflorescences. Cyperus laxus has only pri-
mary and secondary rays, while all available specimens of C. matudae
have tertiary rays as well. This species is named for Professor Eizi
Matuda (1894-1978), specialist on the flora of Mexico, who made
both known collections of this species.

ADDITIONAL SPECIMEN: Mexico. Chiapas, Escuintla, Finca Fuarez, 12 Aug. 1937,
Matuda 1937 (GH, MICH).

Cyperus breedlovei G. C. Tucker sp. nov. TYPE: MEXICO. Chiapas,
on Hwy. 211 about 25 km N of Huixtla, in crevices on rocky
roadcut in forest with Bursera, 700 m, 28 July 1984, Tucker &
Schwartz 2686 (HOLOTYPE: DUKE; ISOTYPES: BH, CAS, ENCB, F,
GH, K, MEXU, MICH, MO, NY, US, WIS). (Figure 2).

506 Rhodora [Vol. 88

Figure 2. Cyperus breedlovei. a. plant; b. spikelet; c. scale (abaxial view); d
achene.

Plantae perennes, 25-50 cm alta. Bracteae 3-5, ad 12 cm longae;
bractea longissima erecta vel valde adscendens. Spiculae digitatae,
1-5 (7), lineari-oblongae, 4-15 mm longae, 1.8-2.4 mm latae.

1986] Tucker — Cyperus 507

Squamae oblongo-lanceolatae, mucrone apicali 0.2-0.4 mm longo.
Stamina tria. Stigmata tria. Achenia trigona, ellipsoidea, 1.2-1.4
mm longa, usualiter 0.6 mm lata. Cypero chalarantho Pres! affinis.

Perennials, 25-50 cm tall. Rhizomes indurate, about 2 mm thick.
Culms solitary or 2-3 together, trigonous and smooth below, trique-
trous and scabrellate (or smooth) above, 0.8-1.2 mm thick. Leaves
1-5, conduplicate, 10-40 cm long, 2-3.5 mm wide, the margins and
keels scabrellate. Inflorescence bracts 3-5, up to 12 cm long, 1-2.5
mm wide, the margins and keels scabrellate, the longest erect or
strongly ascendent (appearing as a continuation of the culm). Rays
2-8, up to 4 cm long, scabrellate; secondary rays absent or 1-4, up
to | cm long. Spikelets digitate, 1-5 (7), linear-oblong, compressed,
reddish brown, 4-15 mm long, |.8-2.4 mm wide; rachilla straight,
wingless, persistent. Scales 7-16, oblong-lanceolate, 1.8-2.3 mm
long (including the mucronate apex of 0.2-0.4 mm), I1-1.2 mm wide,
laterally reddish brown, 2-3 nerved, the margins sometimes sparse-
ly ciliate-scabrellate, medially green, 3-5 nerved, the midvein sca-
brellate, deciduous. Stamens 3; anthers about | mm long, the setose
connective apices about 0.2 mm long. Styles 0.2-0.4 (0.6) mm long;
stigmas 3, 2.3-2.5 mm long. Achenes trigonous, ellipsoid, 1.2-1.4
mm long, about 0.6 mm wide, apiculate, cuneate to substipitate,
puncticulate, brown.

Endemic to southern Chiapas, Cyperus breedlovei is known from
only three collections from the type locality: crevices in steep moist
ledges in tropical forest at about 700 m elevation. Flowering and
fruiting collections have been made in July and October. Cyperus
breedlovei is distinguished from all other Mexican and Central
American species of subgenus Protocyperus by the combination of
linear-oblong spikelets, reddish brown mucronate scales, and ellip-
soid apiculate achenes. The oblong-lanceolate scales nerved nearly
to the margins, the straight mucronate apices of the scales, and the
ellipsoid achenes indicate that the new species is most closely related
to C. chalaranthus Presl of northwestern South America (Tucker,
1983). Cyperus breedlovei differs from C. chalaranthus in having
the longest inflorescence bract erect to ascendent (at 60°), and the
scales reddish brown and scabrellate along the midvein; in C. chala-
ranthus the bracts are horizontal to ascendent (at 30°) and the scales
light green and smooth. This species is named for Dennis E. Breed-
love, diligent investigator and collector of the flora of Chiapas, who
gathered the first collection.

508 Rhodora [Vol. 88

ADDITIONAL SPECIMENS: Mexico. CHIAPAS: Mpio. Motozintla, 25-27 km NE of
Huixtla on the road to Motozintla [Hwy. 211], 7 Oct. 1972, Breedlove 28603 (DS); 15
miles N of Huixtla on Rte. 211, 28 July 1984, Wilbur 35843 (DUKE).

Cyperus wilburii G. C. Tucker sp. nov. TyPE: MEXICO. Oaxaca, 8.5
km W of Niltepec on Hwy. 200 at the turnoff to Cerro Iguana,
31 July 1984, Tucker 2756 (HOLOTYPE: DUKE; ISOTYPES: BH, C,
CAS, ENCB, F, GH, K, MICH, MO, NY, US). (Figure 3).

Plantae annuae caespitosae, (5) 10-25 (30) cm alta. Radii simpli-
ces; spicae subdigitatae. Spiculae compressae, (3.5) 5-6.5 (7) mm
latae, stramineae vel fulvae. Squamae (3.8) 4-4.6 mm longae, (2.4)
2.6-3.2 (3.6) mm latae, cuspide apicali (1.1) 1.3-1.5 (1.6) mm longa.
Achenia obovoidea, stipitata, trigona, apiculata, 1.7—2.2 mm longa,
1-1.4 mm lata. Cypero compresso L. affinis.

Caespitose annuals, (5) 10-25 (30) cm tall. Roots fibrous, rhizome
none. Culms trigonous, smooth, 0.5-1.5 mm thick. Leaves (1) 3-4,
(1) 12-20 (30) cm long, 0.5-3 mm_wide, conduplicate, the margins
and keels scabrellate distally. Inflorescence bracts (2) 3-5 (6), (2)
8-15 (25) cm long, 0.5-4 mm wide, the margins and keels scabrel-
late, ascendent at 45-60° (the longest bract in some plants erect,
looking like a continuation of the culm). Rays 1-3 (5), unbranched,
smooth, 1.5-5 cm long, occasionally lacking, the inflorescence in
such plants a loose cluster of 3-6 spikelets; rachis smooth, 1.5-3.5
(6) mm long. Spikelets (1) 3-7 (10), oblong to oblong-lanceolate,
strongly compressed, (10) 14-20 (32) mm long, (3.5) 5-6.5 (7) mm
wide, stramineous to tawny brown; rachilla slightly geniculate,
brown to reddish brown, persistent, 0.5-0.8 mm wide, 0.2-0.3 mm
thick, the hyaline wings 0.1 (-0.2) mm wide, successive scale scars
2-2.4 (2.6) mm apart on the same side. Scales 10-16 (30), ovate-
oblong, (3.8) 4-4.6 mm long, (2.4) 2.6-3.2 (3.6) mm wide, the cusp
straight to slightly excurved (1.1) 1.3-1.5 (1.6) mm long, the medial
part of the scale greenish, smooth, weakly 3-5 nerved, laterally
stramineous to reddish brown, conspicuously 2-nerved midway
between the center and margins, deciduous. Stamens 3; filaments
4-5 mm long; anthers oblong, 0.8-0.9 mm long, the connective
apices minute. Styles 2-2.5 mm long; stigmas 3, about 0.6 mm long.
Achenes trigonous, obovoid, stipitate, 1.7-2.2 mm long, 1.0-1.4 mm
wide, apiculate, the base 0.3-0.4 mm long and about as wide, the
angles acute, the faces concave, puncticulate, brownish.

Endemic to Mexico, Cyperus wilburii occurs in the Pacific Coast-
al lowlands of the Isthmus of Tehuantepec in the State of Oaxaca.

1986] Tucker — Cyperus 509

Figure 3. Cyperus wilburii. a. plant; b. spikelet; c. scale (abaxial view); d. achene.

Apparently uncommon and known from only five collections, it
grows in disturbed open soil among thorny shrubs and cacti and in
pastured thorn savannas from sea level to about 100 m. A single

510 Rhodora [Vol. 88

flowering collection has been made in June; collections made from
July through October have mature achenes.

Cyperus wilburii is classified in subgenus Cyperus, as indicated by
its trigonous achenes and spicate inflorescences. It is closely related
to the pantropical and warm temperate C. compressus L., as clearly
indicated by its annual habit, loosely clustered spikelets, cuspidate
scales, and trigonous obovoid achenes. To the unaided eye, C. wil-
burii is easily distinguished from C. compressus by its wide (5.0-6.5
mm), stramineous to tawny spikelets; in C. compressus the spikelets
are only 2.0-3.0 (3.6) mm wide and greenish white. In addition, the
achenes of C. wilburii are stipitate and larger (1.7-2.2 mm long,
1—1.4 mm wide) than the sessile achenes of C. compressus (1-1.5 mm
long, 0.9-1.1 mm wide). This new species is named for Robert L.
Wilbur, Professor of Botany at Duke University.

ADDITIONAL SPECIMENS: Mexico. OAXACA: Ixtepec, 200 ft., 17 Aug. 1935, Fisher
35258 (ARIZ, F, MO, NY, SMU, TEX, US); along the hwy. to Tehuantepec, 6 km N of
Salina Cruz, 30 June 1958, King 9/7 (MICH, TEX, US); San Geronimo, 5 Oct. 1933,
Mell 2115 (Ny, US); Route 200, about 5.3 miles W of Niltepec near turnoff to Cerro
Iguana, Wilbur 35922 (DUKE, ENCB).

Cyperus svensonii G. C. Tucker sp. nov. TyPE: MEXICO. Chiapas,
just E of Colonia Aztlan on Hwy. 195 about 1.7 km N of the
Pan American Hwy., along gully in cornfield at about 1000 m,
24 July 1984, Tucker 266] (HOLOTYPE: DUKE; ISOTYPES: CAS,
ENCB, F, GH, K, MEXU, MICH, MO, NY, US, WIS.) (Figure 4).

Plantae perennes, caespitosae. Culmi trigoni, laevi. Bracteae
horizontales vel reflexae. Spicae sessiles, dense cylindricae, albidae.
Spiculae ellipsoideae vel oblongo-ellipsoideae, deciduae; rachillae
alatae. Squamae obovatae. Achenia trigona, oblongo-ellipsoidea
usque ellipsoidea, brunnea usque rubro-brunnea. Cypero regiomon-
tano Britt. affinis.

Perennials, 15-55 cm tall. Rhizomes indurate. Culms loosely ces-
pitose, trigonous, smooth, 1-2 mm thick. Leaves 3-9, plicate, 10-35
cm long, 2.5-4 mm wide, the margins and veins scabrous. Inflores-
cence bracts (3) 4-6, horizontal to reflexed downward parallel to the
culm, 3-15 (25) cm long, 1-5 mm wide, the margins and veins sca-
brous. Inflorescence composed of (2) 4-5 (8) oblong-ellipsoid to
oblong-ovoid, densely cylindrical spikes (3) 10-15 (25) mm long,
5-12 mm wide. Spikelets (30) 60-120, ellipsoid to oblong-ellipsoid,
subterete, 2-4.5 mm long, (0.7) 0.8-1.2 mm wide; rachilla decidu-
ous, with hyaline wings 0.3-0.4 mm wide. Scales 2 (—4), obovate,

1986] Tucker — Cyperus 511

Figure 4. Cyperus svensonii. a. plant; b. spikelet; c. achene.

512 Rhodora [Vol. 88

mucronulate, 2.2-3.2 mm long, |.2-2 mm wide, 3-nerved and green-
ish (rarely off-white) medially, 3-4 nerved and whitish (greenish-
white when immature) laterally, persistent. Stamens 3; anthers
oblong to oblong-ellipsoid, 0.4-0.5 (0.7) mm long, the connective
apices at most 0.1 mm long. Styles (0.4) 0.6-0.8 (1.2) mm long;
stigmas 3, 0.8-1.4 mm long. Achenes trigonous, oblong-ellipsoid to
ellipsoid, 1.5-1.8 mm long, 0.6-0.7 mm wide, stipitate to substipi-
tate, acuminate, apiculate, puncticulate, browi: to reddish brown.

The known stations of Cyperus svensonii include the uplands of
Chiapas, Guatemala, Honduras, and Nicaragua, in moist open
situations in pine or oak forests, from (800) 1200-2300 m. The
species is frequent throughout its range, especially in the montane
valleys of Chiapas. Fruiting specimens have been collected from late
June until November.

Cyperus svensonii is the only species of the genus in the New
World that has densely cylindric whitish spikes. The densely cylin-
drical sessile spikes indicate a close relationship with C. regiomon-
tanus Britt. In fact, several collections of this species were previously
identified as “C. regiomontanus.” However, C. svensonii has ellip-
soid to oblong-ellipsoid spikelets with obovate scales that are later-
ally whitish; C. regiomontanus has subulate-lanceolate spikelets and
ovate scales that are laterally greenish to brownish. The two species
are Clearly allopatric: C. svensonii occurs in moist upland sites from
(800) 1200-2300 m from Chiapas to Nicaragua; C. regiomontanus
occurs in beach thickets and brushy pastures at or near sea level
along the Pacific from Sinaloa, Mexico to Ecuador. The new species
is named for Henry K. Svenson (1902-1986), lifelong student of the
Cyperaceae and author of several works on the North American and
neotropical genera of the family.

ADDITIONAL SPECIMENS: Mexico. CHIAPAS: Mpio. Tenejapa, paraje of Sibanil
Ha’, Breedlove 6073 D (Ds, NY); just W of San Cristobal, King 300] (MICH, TEX, UC);
20 km S of Tapilula, Tucker 2630 (DUKE); near Huixtan, Tucker 2209 (DUKE, ENCB);
20 km W of Comitan, Tucker 2679 (DUKE, ENCB); San Benito, Miranda 1889 (MEXU);
5 km N of Bochil, Tucker 2635A (DUKE). Guatemala. Depto. Huehuetenango, “Los
Pinitos,” Steyermark 48192 (F, LL); Depto. Santa Rosa, Santa Rosa, Heyde & Lux
3542 (M, MICH, US); Depto. Chimaltenango, near Chimaltenango, Standley 59055 (F).
Honduras. Depto. Morazan, Rio Yeguare, Glassman 1817 (F); Depto. Comayagua,
vicinity of Siguatepeque, Yuncker 563] (F, GH, MICH); Depto. Morazan, Montafia
Uyuca, Molina R. 12789 (F). Nicaragua. Depto. Esteli, along road from Hwy. | to
San Nicolas, Kral 69478 (MO) and Stevens 17860 (Mo); Depto. Jinotega, Mesas
Moropotente, Henrich & Stevens 429 (DUKE, MO).

1986] Tucker — Cyperus 513

ACKNOWLEDGMENTS

I am grateful for National Science Foundation Grant no. NSF-
BSR-83-03070 which provided support for field work in Mexico. I
thank Prof. Robert L. Wilbur for his advice and support during my
graduate studies, Owen M. Schwartz for assistance and pleasant
company in the field, Charlotte M. Taylor for preparing the illustra-
tions in this paper, Norton G. Miller, Norton H. Nickerson, Joan Y.
Nickerson, and two anonymous reviewers for helpful editorial
comments; and the curators of the following herbaria for access to
specimens: A, ARIZ, ASU, B, BM, BR, C, CAS, CCNL, CR, CU,
DAV,..DS, .DUKE, E, ECON, ENCB, F,. FLAS, G, GH, HAL,
IBUG, K, LCU, LE, LL, M, MEXU, MICH, MO, MSC, NASC,
NY, NYS, PENN, PH, PMA, POM, PR, PRC, RSA, S, SD, SMU,
TENN, TEX, UC, US, UTEP, WIS, WVA, YU, and Z.

LITERATURE CITED

KUKENTHAL, G. 1935-36. Cyperus. In: L. Diels, Ed. Das Pflanzenreich 4 (20,
Heft 101): 1-622.

Lye, K. A. 1981. Two new subgenera of Cyperus. Nord. J. Bot. 1: 57-61.

RAYNAL, J. 1973. Contribution a la classification de la sous-famille des Cyperoi-
deae. Adansonia, sér. 2, 13: 145-171.

Tucker, G.C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and
Panama. Syst. Bot. Monogr. 2: 1-82.

BIOLOGICAL SURVEY

NEW YORK STATE MUSEUM

THE STATE EDUCATION DEPARTMENT
ALBANY, NEW YORK 12230

THE TAXONOMIC POSITION OF
VACCINIUM SEM PERVIRENS (ERICACEAE)

LEONARD J. UTTAL

ABSTRACT

Vaccinium sempervirens Rayner and Henderson is assigned to
Vaccinium sect. Herpothamnus.

Key Words: Vaccinium sempervirens, sect. Herpothamnus, South Carolina

In the original description of Vaccinium sempervirens Rayner
and Henderson (1980), a rare evergreen shrub of central South
Carolina, the authors did not discuss the affinity of the species. They
remarked on its resemblance to Gaylussacia brachycera (Michaux)
Gray and Vaccinium crassifolium Andrews and that it had been
misidentified as one or the other by different botanists. By assigning
this species to section Herpothamnus (Small) Sleumer (Sleumer,
1941), confusion is eliminated.

Sect. Herpothamnus, as here circumscribed, consists of two low
repent to repent-ascending shrubs (Vaccinium crassifolium and V.
sempervirens) endemic to the coastal plain of the southeastern Uni-
ted States. The berry is shiny black to black-purple, round, sweet,
5-locular and 5-seeded, the seeds nutlike; the 5-lobed calyx is articu-
late with a very short pedicel, the corolla pink to white, urceolate
with 5 short lobes, the flowers few in compact |-4 flowered racemes
usually borne in lower leaf axils, or sometimes terminal; leaf innova-
tions are mostly from upper leaf axils. Both species have nodular
lignous roots. Vaccinium crassifolium has strictly repent stems while
in V. sempervirens the stems vary from repent to repent-ascending
to strictly ascending. The two species may be separated by the
following key:

1. Leaves 7-12 (20) mm long, 3-10 mm wide, about 20 per dm of
branchlet; stem always repent, rooting at nodes...........
Ee ee ee er re rr er ere er ree V. crassifolium

2. Leaves 2.2-5.5 cm long, |.3-3.0 cm wide, about 12 per dm of
branchlet; stems repent and node-rooting to ascending, often

CIP TOOUNE <xind Kevws eevee eenva Uae RENE V. sempervirens

Vaccinium crassifolium: sandy pinelands, barrens, or peaty
pocosins, outer coastal plain from southeast Virginia to Georgia
(the type species).

ag Be

516 Rhodora [Vol. 88

Vaccinium sempervirens: A proposed endangered species pres-
ently known only from Atlantic white-cedar communities on seep-
age slopes of Shealy’s Pond, an impoundment in the sand hills of
Lexington County, South Carolina, 4.5 miles southwest of Edmund,
Congaree Girl Scout Camp and at Peach Tree Rock Preserve, a few
miles to the east. The areas are under protective supervision of the
Nature Conservancy. This site is isolated to the west of the range of
V. crassifolium.

Conclusions are based upon (1) telephone consultation with Mr.
D. A. Rayner, who knows the species well, (2) a tototype (VPI), (3)
the original description which includes excellent illustrations of the
plant, and (4) personal observation in the field.

Vaccinium sempervirens is misidentified in Radford, Ahles, and
Bell (1968) as Gaylussacia brachycera (Michaux) Gray, a species
which does not naturally occur south of Virginia.

ACKNOWLEDGMENT

I thank Mr. D. A. Rayner for freely sharing his store of informa-
tion concerning this handsome new evergreen creeper of horti-
cultural potential.

LITERATURE CITED

Raprorp, A. E., H. E. AHLES AND C. R. BELL. 1968. Manual of the Vascular
Flora of the Carolinas. University of North Carolina Press, Chapel Hill.

Raynor, D. A. AND J. HENDERSON. 1980. Vaccinium sempervirens (Ericaceae), a
new species from Atlantic white-cedar bogs in the sandhills of South Carolina.
Rhodora 82: 503-507.

SLEUMER, H. 1941. Vaccinoideen Studien. Bot. Jahrb. Sonder/-Abdr. 71: 375-510.

DEPARTMENT OF BIOLOGY
VIRGINIA POLYTECHNIC INSTITUTE AND STATE UNIVERSITY
BLACKSBURG, VA 24061-0794

NEW ENGLAND NOTE

NOTES ON THE FLORA OF
WINDHAM COUNTY, VERMONT

PETER F. ZIKA AND ELIZABETH H. THOMPSON

ABSTRACT

New stations or re-locations of historical stations for 15 species of rare plants have
been found along the Connecticut and West Rivers in southeastern Vermont. Platan-
thera flava and Rhexia virginica are proposed as state threatened species.

Key Words: rare or threatened plants, Vermont flora

INTRODUCTION

“I have long felt sure that the extreme southeastern corner of
Vermont, embracing the southern half of Windham County, would
prove arich field for botanical exploration. As it is not only next to
Massachusetts but has a warm sandy soil, like that of eastern Mas-
sachusetts and southeastern New Hampshire, I expected to find a
similar flora. Accordingly during my vacation this last summer I
was located in Vernon, on the banks of the Connecticut [River], and
was there about two months exploring the country around.” Thus
wrote A. J. Grout in an unpublished manuscript about his first
extensive collecting trip to Windham Co. in 1895. Although John
Carey and a few other botanists had collected in southeastern Ver-
mont in the early 1800s (Oakes, 1842), Grout was the first to provide
good specimens of the rarities of the area. (Of course, many of these
are rare only in Vermont, and are abundant in southern and coastal
New England.)

Grout discovered several important botanical areas in the 1890s,
including South Pond and Sunset Lake in Marlboro, Kenny Pond
in Newfane (now far less interesting due to adam on the outlet), and
Lily Pond in Vernon. The south-flowing Connecticut River divides
Vermont from New Hampshire, on the eastern edge of Windham
County, and in the broad valley of the Connecticut in Brattleboro
and Vernon, Grout made many of his most interesting finds.

Around the turn of the century W. H. Blanchard and F. Blan-
chard made extensive collections in Windham Co., primarily in the
floodplain of the Connecticut River. Their work and Grout’s was

517

518 Rhodora [Vol. 88

incorporated in the Vermont floras of Brainerd et al. (1900) and
Eggleston et al. (1915). Little has been published about the central
Connecticut River flora since then, except in New Hampshire.

Neighboring Cheshire Co. in New Hampshire is floristically sim-
ilar to Windham Co. (D. Boufford, unpublished manuscript). Crow
and Storks (1980) pointed out that nearly 60 of New Hampshire’s
rare, threatened or endangered species are confined to the Connecti-
cut River and its valley. Unfortunately for Vermont botanists, New
Hampshire territory includes the width of the Connecticut River
and its diverse aquatic flora.

The West River, a tributary of the Connecticut, drains most of the
Green Mountain plateau and foothills in northern Windham Co.
Below a massive flood control dam in Jamaica the West River is ina
narrow valley with cobble- and boulder-lined grassy shores. The
riverbanks have scattered thickets of Alnus crispa (Ait.) Pursh and
Salix sericea Marsh., and an extensive population of Sanguisorba
canadensis L. The last is wholly restricted to the shores of the West
River in Vermont. Most of the rare plants in the West River drain-
age were first collected by L. A. Wheeler between 1911 and 1917
(e.g., Wheeler, 1912, 1913, 1914, 1919). The West is quite different
from the Connecticut, and still provides excellent botanizing.

RECENT WORK

Contemporary collectors in Vermont have attempted to docu-
ment extant populations for all the native plants reported in our
flora. Using the Vermont checklist (Atwood et al., 1973) and label
data on specimens at GH, HNH, MASS, MO, NEBC, SJFM, and
VT, we attempted to re-locate all the rare native species recorded in
Windham Co. (A rare plant is one known from ten or fewer con-
temporary sites statewide.) Rediscoveries and new stations through
1984 are reported below, as is the first Vermont record for the
European adventive Verbascum lychnitis L.

Table | is a list of species known to occur in Vermont only in
Windham Co. at the time of this writing. The majority of these are
in the town of Vernon or along the banks of the West River.

Taxa we searched for unsuccessfully in Windham Co. (and else-
where in Vermont) are given in Table 2. However, more field work
is likely to show that many of these palnts are still extant in the
county.

1986]

Table 1. Native Vermont species currently known to occur only in Windham Co.

New England Note

Aster vimineus

Cassia nictitans
Crotalaria sagittalis
Hypericum gentianoides
Isoetes engelmannii
Juncus acuminatus
Juncus greenei
Lespedeza hirta
Panicum oligosanthes
Panicum rigidulum
Paspalum ciliatifolium
Platanthera flava

Polygala polygama
Polygala sanguinea
Quercus coccinea
Quercus ilicifolia
Rhexia virginica
Sanguisorba canadensis
Scirpus ancistrochaetus
Solidago odora

Tillaea aquatica
Triphora trianthophora
Viola lanceolata

Table 2. Native species documented at least once from Windham Co., with no
known current stations in Vermont.

Arabis drummondii

Aristida longispica var. geniculata

Asclepias tuberosa
Astragalus alpinus
Baptisia tinctoria
Carex davisii
Cynoglossum boreale
Cyperus diandrus
Cyperus engelmannii
Cyperus ferruginescens
Desmodium rotundifolium
Eleocharis olivacea
Epilobium palustre
Eupatorium fistulosum
Fimbristylis autumnalis
Helianthus strumosus
Juncus militaris

Juncus secundus

Lechea minor

Lechea villosa
Lespedeza X nuttallii
Lespedeza steuvii
Myosotis laxa
Myriophyllum humile
Najas gracillima
Oenothera cruciata
Polygonum careyi
Pterospora andromedea
Rosa nitida

Rumex altissimus
Sagina decumbens
Sericocarpus asteroides
Sisyrinchium atlanticum
Subularia aquatica
Utricularia radiata
Xyris difformis

NOTEWORTHY FINDS

Chimaphila maculata (L.) Pursh In Vermont this species of dry
woods has only been found near the Connecticut River. It is much
more common to the south and west. The last Vermont collection
was in 1903. In 1984 three stations were located: a small population

520 Rhodora [Vol. 88

at the base of Black Mountain, Dummerston (Thompson &
Rawinski 760 VT); about 50 plants in dry oak woods in South
Vernon (Zika 8406 VT); and a third site in Springfield, Windsor Co.

Cornus florida L. This attractive plant is at the northern limit of
its range in the Connecticut River valley in southern Vermont and
New Hampshire. A new station for the species was located ona rich
wooded slope in Rockingham (Thompson & Zika 8352 NEBC, VT).
An estimated 20 mature fruiting trees were seen. About 200 young
sprouts formed dense patches of ground cover in the area.

Corylus americana Walt. About 15 shrubs are in dry oak woods
near the railroad right-of-way NE of Fox Hill, Vernon, Windham
Co., Vt. (Thompson & Zika 8383 VT). This is a new county record,
and the only extant site east of the Green Mountains in Vermont.

Elymus villosus Muhl. ex Willd. A rare species in Vermont, last
collected in 1922. An estimated 250 stems were found in a floodplain
forest on the West River in Brattleboro (Zika 7533 VT), growing
with E. wiegandii Fern. and Matteuccia struthiopteris (L.) Todaro.

Elymus wiegandii Fern. Also a rare species in Vermont. About
50 stems were observed in a floodplain forest along the West River
in Brattleboro (Zika 7532 NEBC, VT), growing with the E£. villosus
and Matteuccia struthiopteris.

Juncus marginatus Rostk. A small population is on the sandy
shore of a small unnamed pond in Rockingham, Windham Co., Vt
(Zika 7797 VT). A larger colony is present on the banks of the West
River, on ledges and in peaty wet soil (Thompson & Zika 8360 VT).
Only one other station in Vermont is known to be extant, a small
colony on the banks of the Huntington River in the Champlain
Valley.

Muhlenbergia sobolifera (Muhl.) Trin. About 100 stems are on
a dry rocky forested slope in Rockingham (Thompson & Zika 8353
VT). This is the first record east of the Green Mountains in
Vermont.

Muhlenbergia tenuiflora (Willd.) BSP. About 500 plants are on
a dry rocky wooded slope in Westminster (Thompson & Zika 8348
VT). This is the only extant population in eastern Vermont.

1986] New England Note 521

Platanthera flava (L.) Lindl. A total of 100 plants was found
in the towns of Newfane and Brookline on a short stretch of the
West River (Zika 7553, 7554 VT). Sterile as well as flowering indi-
viduals were inconspicuous, mixed in with the dense low herbs on
the cobble shore.

Podostemum ceratophyllum Michx. The status of this sub-
merged aquatic in Windham Co. has long been unclear. It was first
reported by C. C. Frost from Brattleboro, presumably in the West
River (Eggleston et al., 1915), but no vouchers were located and
recent searches failed to find the species in the city limits. Podoste-
mum was collected in the West River in 1907 in the town of Jamaica
by F. Dobbin (Dobbin, 1907), and was visited there by botanists
through the 1920s. In recent decades Dobbin’s station was not re-
located. The extreme flow fluctuations caused by the installation of
two flood-control dams upriver of the Jamaica site were believed to
have destroyed the population (Countryman, 1978; Philbrick &
Crow, 1983). A new station was discovered on one of the numerous
short rapids on the West River, downriver of Jamaica in Newfane.
Between 30 and 60 cobblestones in the deepest part of the riverbed
were covered with Podostemum (Zika 7562 VT).

Quercus ilicifolia Wang. Scrub oak is restricted to Windham
Co. in Vermont. A colony of about 50 vigorous shrubs was located
under a powerline clearing on the west side of Black Mountain, near
the West River (Zika 7544 VT). Subsequent field work showed the
species was scattered on the southern and western slopes of Black
Mountain, above the powerline, to an elevation of 1100 feet
(Thompson & Rawinski 758 NEBC). This is the only known extant
population in Vermont.

Rhexia virginica L. Vermont’s only extant station is at Lily
Pond in Vernon, where about 200 plants are concentrated on a
short, grazed stretch of the northern shoreline (Zika 6534 VT).

Stellaria alsine Grimm Three colonies of 100 to 300 stems were
found in springs along a small unnamed creek between Vt. Route
142 and the Connecticut River in Vernon (Thompson & Zika 8373
VT). The only other extant colony in Vermont is in Ludlow, Wind-
sor Co.

522 Rhodora [Vol. 88

Verbascum lychnitis L. A weed introduced from Europe and
known mostly to the south and west of Vermont. A small colony
was found in 1982 on the west side of U.S. Route 5 in Dummerston,
just south of the Putney town line (Zika 6266 VT). This is the first
Vermont record of this species.

Viola lanceolata L. Although a common and somewhat weedy
species in southern New England, lance-leaved violet is currently
quite rare in Vermont. The only extant population is about 20
plants on barren ground at the rim of a borrow pit on Lily Pond
Road in Vernon (Zika 7080 VT). A few more clumps can be found
nearby on the shore of Lily Pond. The entire population may be
threatened by grazing cows.

THREATENED STATUS OF PLATANTHERA FLAVA AND RHEXIA VIRGINICA

Platanthera flava was formerly a widespread species in Vermont,
with 25 historical stations represented in herbaria. Eggleston et al.
(1915) considered P. flava locally common in Vermont; Dole (1937)
listed it as occasional. Only two historical stations along the West
River in Windham Co. have been seen in recent years. Many histor-
ical stations in the Lake Champlain valley have disappeared. The
cause of the apparently drastic decline of the species in Vermont is
unknown.

One of the two historical stations for Rhexia was discovered by L.
Henry Potter in Wallingford, where a housing development has
since destroyed the colony (Potter, pers. comm.). The second sta-
tion, in Vernon, is extant but is threatened by grazing cows.

Due to declining populations, Platanthera flava and Rhexia vir-
ginica are recommended for threatened species status in Vermont
(10 V.S.A., Chapter 123).

ACKNOWLEDGMENTS

We are grateful to the Vermont Chapter of The Nature Con-
servancy and the Pringle Herbarium for funding which supported
this field work.

1986] New England Note $23

LITERATURE CITED

Atwoop, J. T., W. D. CoUNTRYMAN, R. A. Jervis, D. H. MILLER, F. C. SEYMOUR,
AND M.L. SMITH. 1973. Check List of Vermont Plants. Vermont Botanical
and Bird Clubs.

BRAINERD, E., L. R. JONES, AND W. W. EGGLESTON. 1900. Flora of Vermont. Free
Press Association, Burlington, Vt.

COUNTRYMAN, W.D. 1978. Rare and Endangered Plant Species in Vermont. The
New England Botanical Club in cooperation with the U.S. Fish & Wildlife
Service, Newton Corner, Massachusetts.

Crow, G. E. AnD I. M. Srorks. 1980. Rare and endangered plants of New
Hampshire: a phytogeographic viewpoint. Rhodora 82: 173-189.

Dossin, F. 1907. Rediscovery of Podostemum ceratophyllum in Vermont. Rho-
dora 9: 220.

Dote, E. J., Ep. 1937. The Flora of Vermont. Third ed. Free Press Printing Co.,
Burlington, Vt.

EGGLESTON, W. W.,G. L. KirK, AND J.G. UNDERWOOD. 1915. Flora of Vermont.
Vermont Agricultural Experiment Station Bulletin 187: 139-258.

Oakes, W. 1842. Catalogue of Vermont plants. Jn: Thompson, Z. History of
Vermont, Natural, Civil, and Statistical. Published by the author, Burlington,
Vt., pp. 173-208.

PHILBRICK, C. T. AND G. E. Crow. 1983. Distribution of Podostemum cerato-
phyllum Michx. (Podostemaceae). Rhodora 85: 325-341.

WHEELER, L. A. 1912. Rare plants of the West River Valley. (Abstract). Bull. Vt.
Bot. Club 7: 13.

1913. Additions to the flora of West River: Valley in 1912. (Abstract).

Bull. Vt. Bot. Club 8: 14-15.

1914. New West River plants. Bull. Vt. Bot. Club 9: 27.
1919. Additions to West River Valley flora. Bull. Vt. Bot. and Bird Clubs

4-5: 8.

P.-E, Z:

PRINGLE HERBARIUM
BOTANY DEPARTMENT
UNIVERSITY OF VERMONT
BURLINGTON, VT 05405

E.H. T.

VERMONT NATURAL HERITAGE PROGRAM
THE NATURE CONSERVANCY

138 MAIN ST.

MONTPELIER, VT 05602

BOOK REVIEW

Proctor, G. R. 1985. Ferns of Jamaica. vi + 631 pp. British
Museum (Natural History), London (£50.00).

The Ferns of Jamaica is the culmination of over thirty years of
study by the author of the native and naturalized pteridophytes of
the island. The concise descriptions of 609 species, varieties and
hybrids along with notes on their distributions and habitats will
serve as an invaluable reference for those wishing to identify Jamai-
can pteridophytes for a long time to come.

Seventy-five of the eighty-three genera recognized are illustrated.
Especially noteworthy are the fine line drawings of twenty of the
twenty-eight species of Elaphoglossum by Charles D. Clare. Other
large and difficult genera such as Adiantum, Thelypteris, Asple-
nium, Diplazium, Polypodium and Grammitis received considera-
bly fewer illustrations by comparison.

Twenty maps plotting the distributions of thirty-five species are
interspersed throughout the text. The author does not indicate why
these species are mapped whereas the other 544 are not.

A conspectus of the major taxa is included rather than a key to
families. Once a plant has been identified to family, there are keys to
subfamily, genus and species. The key to the subfamilies of Poly-
podiaceae sens. lat. appears to be workable for the most part; how-
ever, it is not possible to arrive at the Athyrioideae when keying out
Diplazium plantaginifolium.

The systematic treatment is an interesting blend of a rather liberal
philosophy in recognizing pteridophyte entities with a conservative
approach to recognizing genera and families. All ferns with slender-
pedicellate sporangia which have a vertical annulus are placed in the
Polypodiaceae sens. lat. While some might argue that the subfami-
lies should be raised to families, the question of rank seems less
important than the composition of the groups. In most cases Proc-
tor’s placement of genera into higher taxa agrees with that of other
authors (e.g. Christensen, 1938; Holttum, 1949; Crabbe et al., 1975;
and Tryon and Tryon, 1982), however, a few assignments are
worthy of mention. All of the above authors place Saccoloma in the
dennstaedtioid ferns. Proctor places it in the Lindsayoideae. Sim-
ilarly, all align the genus Hemidictyum with the athyrioid ferns.

525

526 Rhodora [Vol. 88

Proctor places it with the asplenioid ferns. Since descriptions of the
subfamilies are omitted and since it is difficult to determine the
characters of the subfamilies from the key (one subfamily is keyed
out in nine different places), one cannot determine what evidence
the author considered in reaching these systematic conclusions.

Despite these criticisms, the book stands as a major contribution
to tropical American Pteridology. Anyone interested in New World
ferns or in the flora or phytogeography of the Caribbean would
find this book worth the investment.

LITERATURE CITED

CHRISTENSEN, C. 1938. Filicinae. In: Manual of Pteridology, F. Verdoorn, Ed.
522-550. Nijhoff, The Hague.

CRABBE, J. A., A. C. JERMY, AND J. M. MICKEL. 1975. A new arrangement for the
pteridophyte herbarium. Fern. Gaz. 11: 141-162.

Ho.ttum, R. E. 1949. The classification of ferns. Biol. Beviews 24: 267-296.

Tryon, R. M. AND A. F. TRYON. 1982. Ferns and Allied Plants. Springer-Verlag,
Heidelberg.

DAVID S. CONANT

DEPARTMENT OF NATURAL SCIENCE
LYNDON STATE COLLEGE
LYNDONVILLE, VERMONT 05851

NEBC AWARD
FOR THE SUPPORT OF BOTANICAL RESEARCH
IN NEW ENGLAND, U.S.A.

Debra A. Dunlop was selected to receive the 1986 New England
Botanical Club award in support of botanical research in New Eng-
land for her proposal “Biosystematics of Carex section Scirpinae
(Cyperaceae).” Ms. Dunlop, a Ph.D. candidate working with Dr.
Garrett E. Crow at the University of New Hampshire, will use the
award to travel to alpine areas of New York, Vermont, New Hamp-
shire and Maine and to limestone and serpentine sites in south-
eastern Canada to collect material for anatomical, cytological and
micromorphological studies and for crossing experiments.

The New England Botanical Club will again offer an award of
$1000 in support of botanical research to be conducted in the New
England region during 1987. This award is made to stimulate and
encourage botanical research on the New England flora and to
make possible visits to the New England region by those who would
not otherwise be able to do so. The award will be given to the
graduate student submitting the best research proposal dealing with
field studies in systematic botany and plant ecology, but proposals
for research in other areas of botany will also be considered. This
award is not limited to graduate students at New England institu-
tions, nor to members of the New England Botanical Club. Papers
based on this research must acknowledge the NEBC’s support, and
it is encouraged that they be submitted to Rhodora, the Club’s
journal, for possible publication—subject to standard review
processes.

Applicants should submit a proposal of no more than three dou-
ble spaced pages, a budget (the budget will not affect the amount of
the award), and a curriculum vitae. Two letters, one from the stu-
dent’s major professor, in support of the proposed research are also
required. Proposals and supporting letters should be sent before 28
February 1987 to:

Awards Committee
The New England Botanical Club
22 Divinity Avenue
Cambridge, MA 02138
The recipient of the award will be notified by 30 April 1987.

527

THE 1986 JESSE M. GREENMAN AWARD

The 1986 Jesse M. Greenman Award has been won by David H.
Lorence for his publication “A Monograph of the Monimiaceae
(Laurales) in the Malagasy Region (Southwest Indian Ocean),”
which appeared in the Annals of the Missouri Botanical Garden,
volume 72, number |. The publication is derived from a Ph.D.
dissertation from Washington University under the direction of
Alwyn H. Gentry. This comprehensive account of fifty-five species,
all endemic to the region, is based on herbarium records and exten-
sive fieldwork. It includes information on pollination syndromes in
relation to floral morphology, cytological investigations, population
biology, vernacular names, and economic uses.

The Award is named for Jesse More Greenman (1867-1951), who
was Curator of the Missouri Botanical Garden Herbarium from
1919 until 1943. A cash prize of $250 is presented each year by the
Garden, recognizing the paper judged best in vascular plant or
bryophyte systematics based on a doctoral dissertation that was
published during the previous year. Papers published during /986
are now being considered for the 19th annual award, which will be
presented in the summer of 1987. Reprints of such papers should be
sent to:

Greenman Award Committee
Department of Botany

Missouri Botanical Garden

P.O. Box 299

St. Louis, MO 63166-0299, U.S.A.

In order to be considered for the 1987 award, reprints must be
received by | July, 1987.

528

MEETING ANNOUNCEMENT
NEW ENGLAND BOTANY GRADUATE STUDENTS

The second New England Botany Graduate Student Meeting will
be hosted by the Department of Botany and Plant Pathology at the
University of New Hampshire, Durham, NH on Saturday 14 March
1987. Attendance is open to all. Paper presentations will be largely
restricted to graduate student research (completed or in progress)
representing all areas of botany (systematics, ecology, reproductive
biology, anatomy, physiology, etc.). Time slots for paper presenta-
tions are limited and prior registration is required. Abstracts are due
by 15 February 1987. For additional information and abstract
forms, contact:

Debra Ann Dunlop

Department of Botany and Plant Pathology
Nesmith Hall

University of New Hampshire

Durham, NH 03824

(603) 862-3531

WILLIAM WIRT CALKINS
MOSS AND LICHEN COLLECTIONS

The Chicago Academy of Sciences houses the moss and lichen
collections of William Wirt Calkins. There are several thousand
specimens, including 12 exsicatti volumes. The locales represented
include Florida, Illinois, Virginia, Ohio and others. The collections
were made between 1870 and 1910. This collection has not appeared
in scientific literature and is available for study. Inquiries should be
sent to:

Ron Vasile

c/o Chicago Academy of Sciences
2001 North Clark Street

Chicago, IL 60614

529

REVIEWERS OF MANUSCRIPTS
JULY 31, 1985-JULY 31, 1986

The editors of RHODORA are grateful to each of the following
specialists for their participation in the review process.

Ralph M. Adams
Ihsan Al-Shehbaz
Burton R. Anderson
John H. Beaman

C. John Burk
Christopher S. Campbell
Paul M. Catling
Arthur Cronquist
Antoni W. Damman
Thomas C. Dent
Lesley M. Eastman
Leslie A. Garay

C. Barre Hellquist
Walter H. Hodge
Richard Howard
Edwin H. Ketchledge
Robert Kral

Walter H. Lewis
Leslie J. Mehrhoff
Norton G. Miller

Douglass H. Morse
William Niering
Gerald B. Ownbey
Thomas S. Patrick
Anna M. (Nancy) Reid
Anton A. Reznicek
Ruben P. Sauleda
John C. Semple
Charles J. Sheviak
Bruce A. Sorrie
Peter J. Stevens
John L. Strother
Fred Taylor
Francis Trainor

B. L. Turner
Charles H. Uhl
Mary M. Walker
Daniel B. Ward
Gerald A. Wheeler

Hovdora

JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB

NORTON H. NICKERSON, Editor-in-Chief
JOAN Y. NICKERSON, Managing Editor

Associate Editors

DAVID S. BARRINGTON RICHARD A. FRALICK
A. LINN BOGLE GERALD J. GASTONY
DAVID E. BOUFFORD C. BARRE HELLQUIST
WILLIAM D. COUNTRYMAN MICHAEL W. LEFOR
GARRETT E. CROW ROBERT T. WILCE
VOLUME 88
1986

Che Netu England Botanical Club, Inc.

Harvard University Herbaria, 22 Divinity Ave., Cambridge, Mass. 02138

INDEX TO VOLUME 88

New scientific names are in bold face

Achene morphology of Polygonum
section Polygonum (Polygonaceae) in
Canada. Synopsis and, 457-479

Achene morphology. Taxonomy of
Carex section Folliculatae using,
399-403

Additions to the vascular flora of Kent
Island, New Brunswick. 441-443

Aerial shoots of the common marsh reed
Phragmites australis (Poaceae). Dis-
persion pattern of, 325-330

Allred, Kelly W. Studies in the Aristida
(Gramineae) of the southeastern Uni-
ted States. IV. Key and conspectus.
367-387

Al-Shehbaz, Ihsan A. New wool-alien
Cruciferae (Brassicaceae) in eastern
North America: Lepidium and Si-
symbrium. 347-355

Announcements:

Calkins, William Wirt: Collection of
mosses and lichens 529
Greenman, Jesse M.:
1985 award and 1986 notice 154
1986 award and 1987 notice 528
Hesler Visiting Professorship 154
Joint Field Meeting, 1986 295
NEBC:
1985 award and 1986 notice 155
1986 award and 1987 notice 527

New England Botany Graduate
Student Meeting: 1986, 156; 1987,
529

Aquifoliaceae from Haiti. A new spe-
cies of Ilex, 435-439

Aristida (Gramineae) of the southeast-
ern United States. IV. Key and con-
spectus. Studies inthe, 367-387

Aristida purpurescens var. tenuispica
(Hitchc.) Allred, comb. nov. 383

mf Pe

Aristida purpurescens var. virgata (Trin.)
Allred, comb. nov. 383
Astereae. The cytogeography of Chry-
sopsis mariana (Compositae:): Survey
over the range of the species. 261-266
Baker, Les see Harms, Vernon L.
Barrington, David S. The morphology
and cytology of Polystichum X potteri
hybr. nov. (= P. acrostichoides X P.
braunii). 297-313
Bergeron, Yves see Bouchard, Andre
Book Reviews:
Bogs of the Northeast 149
Botany and Natural History of Pana-
ma: La Botanica e Historia Natu-
ral de Panama. The, 150-151
Cytotaxonomic Studies of the Ferns
of Trinidad 421-422
Ferns of Jamaica 525-526
Field Manual of the Ferns and Fern-
Allies of the United States and
Canada. A, 293-294
Michigan Flora, Part II Dicots
424
Bouchard, Andre, Stuart Hay, Claire
Gauvin and Yves Bergeron. Rare
vascular plants of Gros Morne Na-
tional Park, Newfoundland, Canada.
481-502
Bowles, Marlin L. see Sheviak, Charles
J
Brassicaceae in eastern North America:
Lepidium and Sisymbrium. New wool-
alien Cruciferae, 347-355
Briggs, Errol C. see Effron, Michael
Briggs, Timothy L. see Davis, Andrew N.
Campbell, Christopher S., Norman C.
Famous and Michael G. Zuck. Pol-
lination biology of Primula laurenti-
ana(Primulaceae) in Maine. 253-260

423-

534

Canada. Carex oligocarpa (Cypera-
ceae), a rare sedge in, newly discov-
ered in Quebec. 427-433

Canada. Rare vascular plants of Gros
Morne National Park, Newfoundland,
481-502

Canada. Synopsis and achene mor-
phology of Polygonum section Poly-
gonum (Polygonaceae) in, 457-479

Carex mackenziei new for Saskatche-
wan: Additional rare inland stations
for two seacoast salt marsh species.
Plantago maritima and, 315-323

Carex oligocarpa (Cyperaceae), a rare
sedge in Canada newly discovered in
Quebec. 427-433

Carex purpurifera Mack. (Cyperaceae).
Cytology of, 141-147

Carex section Folliculatae using achene
morphology. Taxonomy of, 399-403

Carex striata, the correct name for C.
walteriana (Cyperaceae). 405-406

C. (arex) walteriana (Cyperaceae). Carex
striata, the correct namefor, 405-406

Carter, Richard and Charles E. Jarvis.
Re-evaluation and lectotypification of
Scirpus retrofractus L. 451-456

Castillo, Rafael and Richard Evans
Schultes. New combinations in the
Solanaceae. 291-292

Catling, P. M. see Reznicek, A. A.

Chinnappa, C. C. see Semple, John C.

Chrysopsis mariana (Compositae: Aste-
reae): Survey over the range of the
species. The cytogeography of, 261-
266

Connecticut. Holocene fruit, seed and
leaf flora from riverine sediments near
New Haven, 229-252

Connecticut. Seasonal succession and
vertical distribution of phytoplankton
in Candlewood Lake, 331-346

Cruciferae (Brassicaceae) in eastern
North America: Lepidium and Sisym-
brium. New wool-alien, 347-355

Current status of Magnolia virginiana in
Massachusetts. 357-365

Cyperaceae, a rare sedge in Canada

Rhodora

[Vol. 88

newly discovered in Quebec. Carex
oligocarpa, 427-433

Cyperaceae. Carex striata, the correct
name for C. walteriana, 405-406

Cyperaceae. Cytology of Carex purpu-
rifera Mack. 141-147
Cyperaceae. New Mesoamerican spe-

cies of Cyperus, 503-513

Cyperus breedlovei G. C. Tucker sp.
nov. 505

Cyperus (Cyperaceae). New Mesoa-
merican species of, 503-513

Cyperus matudae G. C. Tucker sp.
nov. 503

Cyperus svensonii G. C. Tucker sp.
nov. 510

Cyperus wilburii G. C. Tucker sp. nov.
508

Cypripedium acaule (Orchidaceae). The
pollination biology of, 445-450

Cytogeography of Chrysopsis mariana
(Compositae: Astereae): Survey over
the range of the species. The, 261-266

Cytology of Carex purpurifera Mack.
(Cyperaceae). 141-147

Cytology of Polystichum X_ potteri
hybr. nov. (= P. acrostichoides X P.
braunii). The morphology and, 297-
313

Davis, Andrew N. and Timothy L. Briggs.
Dispersion pattern of aerial shoots of
common marsh reed Phragmites aus-
tralis (Poaceae). 325-330

Davis, Richard W. The pollination
biology of Cypripedium acaule (Or-
chidaceae). 445-450

Del Tredici, Peter see Primack, Richard
B.

Dispersion pattern of aerial shoots of
the common marsh reed Phragmites
australis (Poaceae). 325-330

Effron, Michael and Errol C. Briggs.
Rediscovery of Isotria verticillata
(Willd.) Raf. (Orchidaceae) in Ver-
mont. 407-408

Ericaceae. The taxonomic position of
Vaccinium sempervirens, 515

Eurasian adventives in the flora of west-

1986]

ern Massachusetts. Four rare, 413-
417

Famous, Norman C. see Campbell,
Christopher S.

Fernald, Merritt Lyndon. A Tribute
to, New England Botanical Club 800th
meeting. 157-228

First record of Poa bulbosa L. (Poaceae)
for Vermont. The, 419-420

Flora of Windham County, Vermont.

Notes onthe, 517-523

Four rare Eurasian adventives in the
flora of western Massachusetts. 413-
417

Freeda, Stanley J. and Peter A. Siver.
Seasonal succession and vertical dis-
tribution of phytoplankton in Can-
dlewood Lake, CT. 331-346

Gagnon, Daniel see Hay, Stuart G.

Gauvin, Claire see Bouchard, Andre

Goldenaster. The cytogeography of
Chrysopsis mariana (Compositae:
Astereae): Survey over the range of
the species. 261-266

Gramineae of the southeastern United
States. Studies in the Aristida, IV.
Key and conspectus. 367-387

Grayum, Michael H. and Marian F.
Rohman. Four rare Eurasian adven-
tives in the flora of western Massa-
chusetts. 413-417

Gros Morne National Park, Newfound-
land, Canda. Rare vascular plants
of, 481-502

Haiti. A new species of Ilex (Aquifoli-
aceae) from, 435-439

Harms, Vernon L., Donald F. Hooper
and Les Baker. Plantago maritima
and Carex mackenziei new for Sas-
katchewan: Additional rare inland
stations for two seacoast salt marsh
species. 315-323

Hay, Stuart see Bouchard, Andre

Hay, Stuart G. and Daniel Gagnon.
Carex oligocarpa (Cyperaceae), a rare
sedge in Canada newly discovered in
Quebec. 427-433

Hedyotis rosea Rafinesque and a new

Index to Volume 88 535

combination in Houstonia. Nomen-
clatural notes on, 389-397

Hehre, Edward J. see Mathieson, Arthur
Cc.

Hendry, Edward see Primack, Richard
B.

Holocene fruit, seed and leaf flora from
riverine sediments near New Haven,
Connecticut. 229-252

Hooper, Donald F. see Harms, Vernon
3

Houstonia. Nomanclatural notes on
Hedyotis rosea Rafinesque and a new
combination in, 389-397

Houstonia rosea (Rafinesque) Terrell,
comb. nov. 395, fig. 394

Ilex (Aquifoliaceae) from Haiti. A new
species of, 435-439

Ilex blancheana W. Judd, sp. nov. 436,
fig. 437

Inland stations for two seacoast salt
marsh species. Plantago maritima and
Carex mackenziei new for Saskatche-
wan: Additional rare, 315-323

Isotria verticillata (Willd.) Raf. (Orchi-
daceae) in Vermont. Rediscovery of,
407-408

Jaltomata glandulosa Castillo and
Schultes comb. nov. 292.

Jaltomata vestita Castillo and Schultes
comb, nov. 292

Jarvis, Charles E. see Carter, Richard

Judd, Walter S.A new species of Ilex
(Aquifoliaceae) from Haiti. 435-439

Keenan, Philip E. New stations for
Platanthera flava and Triphora trian-
thophora and other observations.
409-412

Kent Island, New Brunswick. Additions
to the vascular flora of, 441-443

Lectotypification of Scirpus retrofractus
L. Reevaluation and, 451-456

Lepidium and Sisymbrium. New wool-
alien Cruciferae (Brassicaceae) in
eastern North America: 347-355

Magnolia virginiana in Massachusetts.
Current status of, 357-365

Maine. Pollination biology of Primula

536 Rhodora

laurentiana (Primulaceae) in, 253-260
Manhart, James R. Cytology of Carex
purpurifera Mack. (Cyperaceae).
141-147
Marsh reed Phragmites australis (Poa-
ceae). Dispersion pattern of aerial
shoots of the common, 325-330
Massachusetts. Current status of Mag-
nolia virginiana in, 357-365
Massachusetts. Four rare Eurasian ad-
ventives in the flora of western, 413
417
Mathieson, Arthur C. and Edward J.
Hehre. A synopsis of New Hamp-
shire seaweeds, 1-139
Mcllraith, Alexander L. Additions to
the vascular flora of Kent Island, New
Brunswick. 441-443
McNeill, John see Wolf, Steven J.
Menapace, Francis J. see Wujek, Daniel
E.
Merinthopodium vogelii Castillo and
Schultes comb. nov. 292
Mesoamerican species of Cyperus (Cy-
peraceae). New, 503-513
Morphology and cytology of Polysti-
chum X potteri Aybr. nov. (= P.
acrostichoides X P. braunii). The,
297-313
New Brunswick. Additions to the vas-
cular flora of Kent Island, 441-443
New combinations in the Solanaceae.
291-292
New England Botanical Club 800th meet-
ing. A Tribute to Merritt Lyndon
Fernald. 157-228. Remarks: 157-198;
letters: 198-228
New England Notes:
First record of Poa bulbosa L. (Poa-
ceae) for Vermont. The, 419-420
Four rare Eurasian adventives in the
flora of western Massachusetts.
413-417
New stations for Platanthera flava
and Triphora trianthophora and
other observations. 409-412
Notes on the flora of Windham County,
Vermont. 517-523
Rediscovery of Isotria verticillata
(Willd.) Raf. (Orchidaceae) in Ver-
mont. 407-408

[Vol. 88

New Hampshire seaweeds.
of, 1-139

New Mesoamerican species of Cyperus
(Cyperaceae). 503-513

New species of Ilex (Aquifoliaceae) from
Haiti. A, 435-439

New stations for Platanthera flava and
Triphora trianthophora and other ob-
servations. 409-412

New wool-alien Cruciferae (Brassicaceae)
in eastern North America: Lepidium
and Sisymbrium. 347-355. Key to
species of Lepidium 349

Newfoundland, Canada. Rare vascular
plants of Gros Morne National Park,
481-502

Nomenclatural notes on Hedyotis rosea
Rafinesque and a new combination in
Houstonia. 389-397

North America: Lepidium and Sisym-
brium. New wool-alien Cruciferae
(Brassicaceae) in eastern, 347-355

Notes on the flora of Windham County,
Vermont. 517-523

Orchidaceae. The pollination biology
of Cypripedium acaule, 445-450

Orchidaceae in Vermont. Rediscovery
of Isotria verticillata (Willd.) Raf.,
407-408

Orchids, the prairie fringed: A pollinator-
isolated species pair, 267-290

Phragmites australis (Poaceae). Disper-
sion pattern of aerial shoots of the
common marsh reed, 325-330

Phytoplankton in Candlewood Lake, CT.
Seasonal succession and vertical dis-
tribution of, 331-346

Pierce, Lee S. and Bruce H. Tiffney.
Holocene fruit, seed and leaf flora
from riverine sediments near New
Haven, Connecticut 229-252

Plantago maritima and Carex macken-
ziei new for Saskatchewan: Addition-
al rare inland stations for two seacoast
salt marsh species. 315-323

Platanthera flava and Triphora trian-
thophora and other observations. New
stations for, 409-412

Plantanthera praeclara Sheviak and
Bowles sp. nov. 278, fig. 269

Poa bulbosa L., (Poaceae) for Vermont.

A synopsis

1986]

The first record of, 419-420
Poaceae. Dispersion pattern of aerial
shoots of the common marsh reed

Phragmites australis, 325-330
Poaceae for Vermont. The first record
of Poa bulbosa L., 419-420

Pollination biology of Cypripedium a-
caule (Orchidaceae). The, 445-450
Pollination biology of Primula laurenti-
ana (Primulaceae) in Maine. 253

260

Pollinator-isolated species pair, A. The
prairie fringed orchids: 267-290

Polygonaceae in Canada. Synopsis and
achene morphology of Polygonum
section Polygonum, 457-479

Polygonum caurianum Robins. ssp. hud-
sonianum S. J. Wolf & McNeill ssp.
nov. 469, fig. 470

Polygonum franktonii S. J. Wolf &
McNeill sp. nov. 474, fig. 475

Polygonum section Polygonum (Poly-
gonaceae) in Canada. Synopsis and
achene morphology of, 457-479

Polystichum X potteri Barrington Aybr.
nov. 298, fig. 300

Polystichum X potteri Ayhr. nov. (= P.
acrostichoides * P. braunii). The
morphology and cytology of, 297-313

Prairie fringed orchids: A_ pollinator-
isolated species pair. The, 267-290

Primack, Richard B., Edward Hendry
and Peter Del Tredici. Current status
of Magnolia virginiana in Massachu-
setts. 357-365

Primula laurentiana (Primulaceae) in
Maine. Pollination biology of,
253-260

Primulaceae in Maine.
ology of Primula laurentiana,
260

Quebec. Carex oligocarpa (Cyperaceae),
a rare sedge in Canada newly dis-
covered in, 427-433

Rare sedge in Canada newly discovered
in Quebec. Carex oligocarpa (Cype-
raceae), a, 427-433

Rare vascular plants of Gros Morne
National Park, Newfoundland, Canada.
481-502. Alpine plateau 491; coastal
plain 485; limestone escarpments 488;

Pollination bi-
253=

Index to Volume 88

537

serpentine tableland 493

Rediscovery of Isotria verticillata ( Willd.)
Raf. (Orchidaceae) in Vermont. 407-
408

Re-evaluation and lectotypification of
Scirpus retrofractus L. 451-456

Reviewers, List of 530

Reznicek, A. A. and P. M. Catling.
Carex striata, the correct name for
C. walteriana (Cyperaceae). 405-406

Riverine sediments near New Haven,
Connecticut. Holocene fruit, seed and
leaf flora from, 229-252

Rohman, Marian F. see Grayum, Michael
H.

Salt marsh species. Plantago maritima
and Carex mackenziei new for Sas-
katchewan: Additional rare inland sta-
tions fortwo, 315-323

Saskatchewan: Additional rare inland
stations for two seacoast salt marsh
species. Plantago maritima and Carex
mackenziei new for, 315-323

Schultes, Richard Evans see Castillo,
Rafael

Scirpus retrofractus L. Re-evaluation
and lectotypification of, 451-456

Seasonal succession and vertical distri-
bution of phytoplankton in Candle-

wood Lake, CT. 331-346
Seaweeds. A synopsis of New Hamp-
shire, 1-139

Sedge in Canada newly discovered in
Quebec. Carex oligocarpa (Cypera-
ceae), arare, 427-433

Semple, John C. and C. C. Chinnappa.
The cytogeography of Chrysopsis mari-
ana (Compositae: Astereae): Survey
over the range of the species. 261-266

Sheviak, Charles J. and Marlin L. Bowles.
The prairie fringed orchids: A polli-
nator-isolated species pair. 267-290

Sisymbrium. Lepidium and, New wool-
alien Cruciferae (Brassicaceae) in
eastern North America: 347-355

Siver, Peter A. see Freeda, Stanley J.

Solanaceae. New combinations in the,

291-292

Southeastern United States. Studies in
the Aristida (Gramineae) of the, IV.
Key and conspectus. 367-387

538

Studies in the Aristida (Gramineae) of
the southeastern United States. IV.
Key and conspectus. 367-387

Synopsis and achene morphology of
Polygonum section Polygonum (Poly-
gonaceae) in Canada. 457-479.
Key to the species 462

Synopsis of New Hampshire seaweeds.
A, 1-139. N. H. coastal zone 9; phe-
nology and longevity 21; physiological
ecology and distributional patterns of
select seaweeds 34; species composi-
tion 20.

Appendix: (Station numbers, coordi-
nates and descriptions): Bellamy
River 128; Blackwater River and
adjacent tributaries 137-138; Brown
River and adjacent tributaries 136-
137; Cocheco River 128-129: Great
Bay 127; Great Bay estuary system
Piscataqua River 121-125; Hamp-
ton Harbor 139; Hampton-Seabrook
estuary, Hampton River and adja-
cent tributaries 134-136; Knowles
Island and Mill Creek areas 138;
Lamprey River 130; Little Bay 125-
127; nearshore open coast between
southern Maine and New Hamp-
shire 120; Oyster River 130-131;
Salmon Falls River 131-132;
Squamscott River 133-134; Winni-
cut River 134.

Distribution maps: 66-119

Tables: Chlorophyceae 39; Phaeophy-
ceae 42; Rhodophyceae 48; sum-
mary of collection sites 54.

Taxonomic position of Vaccinium sem-
pervirens (Ericaceae). The, 515

Taxonomy of Carex section Folliculatae
using achene morphology. 399-403

Terrell, Edward E. Nomenclatural notes
on Hedyotis rosea Rafinesque and a
new combination in Houstonia.
389-397

Thompson, Elizabeth H. see Zika, Peter
F,

Rhodora

[Vol. 88

Tiffney, Bruce H. see Pierce, Lee S.

Triphora trianthophora and other ob-
servations. New stations for Platan-
thera flava and, 409-412

Tucker, Gordon C. New Mesoameri-
can species of Cyperus (Cyperaceae).
503-513

United States, southeastern. Studies in
the Aristida (Gramineae) of the, IV.
Key and conspectus. 367-387

Uttal, Leonard J. The taxonomic posi-
tion of Vaccinium sempervirens (Eri-
caceae). 515

Vaccinium sempervirens (Ericaceae).
The taxonomic position of, 515

Vascular flora of Kent Island, New
Brunswick. Additions to the, 441-443

Vermont. Notes on the flora of Wind-
ham County, 517-523
Vermont. Rediscovery of Isotria verti-

cillata (Willd.) Raf. (Orchidaceae)
in, 407-408

Vermont. The first record of Poa bul-
bosa L. (Poaceae) for, 419-421

Windham County, Vermont. Notes on
the flora of, 517-523

Wolf, Steven J. and John McNeill. Syn-
opsis and morphology of Polygonum
section Polygonum (Polygonaceae) in
Canada. 457-479

Wool-alien Cruciferae (Brassicaceae) in
eastern North America: Lepidium and
Sisymbrium. New, 347-355

Wujek, Daniel E. and Francis J. Mena-
pace. Taxonomy of Carex section
Folliculatae using achene morphology.
399-403

Zika, Peter F. and Elizabeth H. Thomp-
son. Notes on the flora of Windham
County, Vermont. 517-523

Zika, Peter F. First record of Poa bul-
bosa L. (Poaceae) for Vermont. The,
419-420

Zuck, Michael G. see Campbell, Chris-
topher S.

Vol. 88, No. 855, including pages 297-426, was issued July 30, 1986

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RHODORA October 1986 Vol. 88, No. 856

CONTENTS

Carex oligocarpa (Cyperaceae), a rare sedge in Canada newly discovered in
Quebec

Stuart G. Hay and Daniel Gagnon ; ecm ee k tume ga Nf
A new species of Ilex (Aquifoliaceae) from Haiti

Walter S. Judd fom "NF Sachi Sa!
Additions to the vascular flora of Kent lana: New Beunewisl

Alexander L. Mcllraith ; Pet is Sa
The pollination biology of Choviesdtun sande (Ovehidecenal

Richard W. Davis : : : ; «2 eS
Re-evaluation and vetotypilication of Seite nelcalbatiut L;

Richard Carter and Charles E. Jarvis ; 451

Synopsis and achene morphology of Suasien seco piiieeee
(Polygonaceae) in Canada

Steven J. Wolf and John McNeill wig fakes 457
Rare vascular plants of Gros Morne National Park, Newfoutilliead; Canada

André Bouchard, Stuart Hay, Claire Gauvin and Yves Bergeron. § 2 4Od
New Mesoamerican species of Cyperus (Cyperaceae)

Gordon C. Tucker , i eek, «eRe
The taxonomic position of Vestiahen sempervirens (Ritekeenss

PO OROI MOE mig gt ey gaa

NEW ENGLAND NOTE
Notes on the flora of Windham County, Vermont
Peter F. Zika and Elizabeth H. Thompson PDair em ea grein Waele Pg
BOOK REVIEW
Ferns of Jamaica

David S. Conant i : 4 ‘ ; : e : : : : §25
NEBC 1987 Award Notice : ; ; 3 ‘ : ; ‘ : Keke 4)
Jesse M. Greenman Award ‘ : : : ; : ; : ; 2 ee
Botany Graduate Student Meeting Oe Cs EEN OM 50 Vo) geet oT
Calkins Moss and Lichen Collections : : ; > ; ‘ : ; $29
List of Reviewers ; é : ; ; . : ; ; : . ; 530
Index to Volume 88 , : : 3 : ; ‘ : ; : . , 533
Instructions to Contributors : : : ; . Y : % é : 539

Statement of Ownership : SR Ty Sgr Wig Pee E grt se alianrn te RTS eR