(DRAFT) - Taxonomy
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
TAXONOMY
NAME - LOUSEWORT, FURBISH
OTHER COMMON NAMES - LOUSEWORT, FURBISH; LOUSEWORT, FURBISH'S; WOOD BETONY, MISS FURBISH'S;WOOD BETONY and ST. JOHN RIVER
ELEMENT CODE -
CATEGORY - Angiosperm
PHYLUM AND SUBPHYLUM - MAGNOLIOPHYTA,
CLASS AND SUBCLASS - MAGNOLIOPSIDA,
ORDER AND SUBORDER - SCROPHULARIALES,
FAMILY AND SUBFAMILY - SCROPHULARIACEAE,
GENUS AND SUBGENUS - PEDICULARIS,
SPECIES AND SSP - FURBISHIAE,
SCIENTIFIC NAME - PEDICULARIS FURBISHIAE
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Furbish Lousewort
Pedicularis furbishiae S. Wats.
KINGDOM: Plant GROUP: Angiosperm
DIVISION: Magnoliophyta CLASS: Magnoliopsida
ORDER: Scrophulariales FAMILY: Scrophulariaceae
Pedicularis furbishiae S.Wats. (34) is an herbaceous perennial in
the Figwort Family (Scrophulariaceae). The leaves, which grow in a
rosette on the ground and (in reproducing individuals) alternately
along the stem, are rather fern-shaped, i.e., they are long, narrow,
and deeply cut (pinnatifid to pinnately-compound). Most leaves are
between 4 and 10 cm in length, although they may range up to 20 cm.
Soft, pale hairs cover most leaves; some leaves are edged with a
silvery line. Leaves are usually bright green, but those exposed to
high light levels assume a reddish cast due to the accumulation of
anthocyanins. Reproducing plants send up flowering stalks (scapes)
0.1 - 1 m tall. The flowers are in a dense cylindrical cluster (a
raceme) at the tip of the stem some stems send off lateral branches
which bear racemes as well. Toothed, green bracts subtending each
flower dominate the inflorescence, with the individual flowers
projecting horizontally. The flowers are tubular, with a five-toothed
Taxonomy - 1 (DRAFT) - Taxonomy
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
calyx and yellow petals forming the corolla. The corolla is
two-lipped at its apex, with an opening for pollinator access. The
flowers are approximately 2 cm long. They may be found from mid-July
to late August: the flowers at the base of the inflorescence ripen
first, followed by those towards the apex. After pollination, they
develop into ovoid capsules which protrude from the calyx and open in
September. The oblong, flattened seeds have a greyish, loose, and
pitted seed coat, and are approximately 2 mm in length. [Paragraph
references: 01,12,16]
Kate Furbish, botanist and wildflower artist, discovered this
strange Pedicularis on the shores of the St. John River at Van Buren,
Aroostook Co., ME in 1880 (02). From her specimens, Sereno Watson of
Harvard University described it as a new species and named it in her
honor (34). The validity of the taxon has never been questioned (06),
and there are thus no synonyms (04). Indeed, L.W. Macior, a
specialist on the genus Pedicularis, considered P. furbishiae to be
the most endemic of the more than 500 known members of the genus (03).
Common names for Pedicularis furbishiae include Furbish lousewort (05
and many others), Furbish's lousewort (01), Miss Furbish's Wood Betony
(01), and the St. John River Wood Betony (05,06).
The type specimen, and others, are deposited at the Gray
Herbarium of Harvard University (07). Other specimens are at the New
England Botanical Club Herbarium (Cambridge, MA); the University of
Maine at Orono Herbarium; the University of Wisconsin--Madison
Herbarium; the National Herbarium of Canada, Ottawa; the New Brunswick
Museum in St. John; and the Fowler Herbarium of Queen's Univ.,
Kingston, N.B.; as well as several other herbaria in the U.S. and
foreign countries (07). Illustrations of the plant may be found in
the recovery plan (05), Maine's Planning Report (01), and New
England's Rare, Threatened, and Endangered Plants (06). The latter
also has a color photo of the plant. Miss Furbish's watercolor of her
namesake is part of a collection donated to Bowdoin College Library,
Brunswick, Maine.
Pedicularis furbishiae's relationship to its congeners is
unclear. Just what its nearest relative is, and how it arrived in
the St. John Valley (which has only been deglaciated for about 10,000
years) are mysteries (08). Morphology, floral structure, and
phenology discount any close relationship with the two other
Pedicularis species of eastern North America (09). It may instead be
related to a western North American or a central-east Asian species
(08).
Taxonomy - 2 (DRAFT) - Status
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
STATUS
Coded Status
E: Federal Endangered
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The Furbish lousewort (Pedicularis furbishiae) has been
designated an Endangered species pursuant to the Endangered Species
Act of 1973 (50 CFR 17.12; P.L. 93-205, 87 Stat. 884; 16 U.S.C.
1531-1540), as amended. The species has this status wherever found
including the State of Maine and the Province of New Brunswick.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Maine
UNOFFICIAL LIST: Designation of 13 critical areas for the
Furbish lousewort.
STATE STATUTES: Critical Areas Act, Title 5, Chapter 312.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
The lousewort is listed as Endangered pursuant to the Endangered
Species Act of the Province of New Brunswick. In addition, the
Canadian Committee on the Status of Endangered Wildlife in Canada
(COSEWIC) has declared the species endangered. This species is also
found listed in the IUCN Red Book (1978).
ECONOMIC STATUSES:
No known commercial values exist, however, the lousewort has
educational, scientific, and aesthetic value to man.
76/06/16:41 FR 24524/24572 - Proposed rule
Status - 1 (DRAFT) - Status
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
78/04/26:43 FR 17910/17916 - Listing as Endangered
83/12/08:48 FR 55100/55102 - Five year review
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
INLAND AQUATIC
SOCIETY OF AMERICAN FORESTRY TYPES
SAF TYPE STAGE CLOSURE
White spruce balsam fir young tree
White spruce balsam fir mature tree
Aspen-paper birch young tree
LAND USE -
Residential
Cropland and Pasture
Evergreen Forest Land
Mixed Forest Land
Nonforested Wetland
NATIONAL WETLAND INVENTORY CODES
NWI NWICLS NWIMOD NWISPEC
Riverine, upper perennial RS1
Palustrine SS1
COMMENTS ON HABITAT ASSOCIATIONS -
The St. John River is characterized by high banks of gravel and
alluvium, large seasonal flow variations, and ice jams due to its
northward flow (01). These conditions create a shifting, unstable
bank where entire stretches of vegetation are occasionally removed by
high water and ice. Denudation and revegetation appear to be
characteristic of the habitat.
Pedicularis furbishiae grows in the geolittoral zone of these
shores. This shrub- and herb-dominated zone is below the forest edge
and above the river bed, and is maintained in a semi-open state by
occasional ice scour and annual high water (01,05,09,11). Most
populations are on north-facing banks, overshadowed by a tree canopy,
so that they receive little direct sun. Furbish's lousewort is most
commonly found in rivershore "seeps", where groundwater trickles out
of the gravelly soil, but is also found in slightly drier areas (01,
05,11,12).
Because of the annual inundation, this rivershore community can
be classified as a wetland (19). Using the USFWS Classification
system (19), most sites for Furbish's lousewort would be classified as
follows: SYSTEM = Palustrine, CLASS = Shrub-Scrub Wetland, SUBCLASS =
Broadleaf Deciduous, WATER REGIME = Temporarily flooded, WATER
CHEMISTRY = Fresh-circumneutral, SOIL = Mineral (20,21). Rarely, a
small population occurs on ledge outcrops (01,12), in which case it
would be classified in the "bedrock" subclass of the "Rocky Shore"
class.
With the exception of one small population in New Brunswick,
P. furbishiae is restricted to the immediate rivershore. Adjacent
habitats which may affect lousewort habitats are in most cases forests
of the Eastern Spruce-Fir Association (01,05,09,21,28). Conifers such
as Picea rubens, Picea glauca, Abies balsamea, and Thuja occidentalis
predominate; the hardwoods Acer spicatum, Betula lutea, and Populus
balsamifera are occasional (21). On the upper reaches of the St. John
River, shoreland zoning restricts cutting of these forests immediately
adjacent to the open shore (exact width of buffer varies with
Habitat Associations - 1 location), so while these are not virgin forests, they are mostly
mature stands. At a few populations, thin forests of the Aspen-Birch
Association (28), or a mixed forest of Aspen-Birch and Spruce-Fir,
border the rivershore. These are generally narrow strips of young
trees between the rivershore and adjacent agricultural fields or
between the rivershore and the adjacent road. As discussed under the
"Reasons for Status" narrative, ideal lousewort habitat is rivershore
that is bordered by the native spruce-fir forest. Removal of these
forests is detrimental to the populations (05,13). Populations which
are adjacent to agricultural fields are threatened only if the buffer
strip of trees between the field and river is removed or severely
thinned.
Known Environmental Parameters:
Temperature: Temperature in Allagash, ME ranges from 37-85
degrees F in June, 45-89 degrees F in July, and 41-86 degrees F in
August 1984 (12). (The warm spell in June was unusual.) However, the
north- or northwest-facing banks on which Furbish's lousewort usually
grows are noticeably colder (up to 10 degrees F) than the surrounding
uplands. Air temperature at a lousewort population from mid-July to
late-August 1977 ranged from 8 to 32 degrees C (46-89 degrees F), with
a daily average of 13 to 23 degrees C (55-73 degrees F) (09).
Elevation. P. furbishiae populations grow at approximately 500
ft. to 800 ft. above mean sea level (29). At each site, however, the
plants are restricted in elevational distribution to a swath 1-9 m
wide (11). This band varies with respect to distance below forest
edge and distance above river bed, and may represent former hydrologic
events (11).
Slope: Furbish's lousewort typically grows on steep slopes. At
six such populations studied in 1983, the average slope ranged from
64% to 107% with slopes up to 127% locally. Populations may, however,
be found on sandy or ledgy flats of less than 15% slope (11).
Aspect: Almost all populations are found on the east or south
bank, and thus face NW, N, or NE. The three or four populations on
the other bank face E to SE (01,12). Almost all known stations are
shaded for most of the day either by aspect or due to the overhanging
trees (01).
Tree Canopy Cover: Six sites studied in 1983 had 23 to 59% of
the overhead sky obscured by tree canopy; one exceptional site had no
adjacent canopy (11).
Soils: Riverbank soils are derived from glacio-fluvial and
glacio-lacustrine deposits (15) and are mostly gravel and sand.
Samples from seven sites were sandy loams or loamy sands in texture,
with 2 - 29% gravel (11). Soils at three of the seven sites were
continually saturated by groundwater flow. While other sites were
drier, the maximum soil water tension recorded was only -72 centibars.
Thus, while the soils are very well drained, groundwater keeps the
soils saturated to fairly moist. Soil pH ranges from neutral to
slightly acid (7.0 - 5.2); but 50% of these samples were between 6.6
and 7.0, closer to neutral (09). Soils at the three Canadian stations
had pH values of 7.2, 7.4, and 7.8 (22). The soils are high in
calcium but low in nitrates and ammonia as well as organic matter
(09).
Cover: P. furbishiae occurs in vegetated areas of at least 50
pct. cover (11). It is generally found in areas which have been
undisturbed long enough to develop a ground cover of moss or of
graminoid litter over moss. It is, however, absent from the most
densely vegetated areas. At most sites the dominant shrubs are under
2 m high and do not form a continuous cover; Furbish's lousewort
tends to occupy the gaps (11,12).
Conclusion: Furbish's lousewort occupies a narrow "window" not
Habitat Associations - 2 only in space but also in time. Studies to date have shown that while
the species does not colonize areas newly scoured by ice, neither does
it succeed where lack of disturbance allows dense shrub cover to
develop (11).
Habitat Associations - 3 (DRAFT) - Food Habits
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
FOOD HABITS
TROPHIC LEVEL -
AUTOTROPH
Food Habits - 1 (DRAFT) - Environment Associations
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G
G Aquatic Features: Springs [flowing]
G Terrestrial Features: Rock outcrops
G
G
G
G
G
G
G
Environment Associations - 1 (DRAFT) - Life History
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
LIFE HISTORY
HABIT:
Herb (01,05,23).
LIFE CYCLE:
Perennial. How long individuals live is unknown, since the
plants are impossible to age and demographic studies have followed
individuals for only two years to date. The studies to date have,
however, confirmed that the plant is an iteroparous perennial (11,16),
and indicate that individuals take at least three years to reach
flowering size. A field population from seed sown in the fall of 1981
contained several individuals which bloomed and set seed in 1984 (12).
TYPE OF REPRODUCTION:
Sexual. P. furbishiae is an obligate outcrosser (09) and
reproduces only by sexual means (09,01).
REPRODUCTIVE PHENOLOGY:
Germination Dates: Mid-late June through August (11,12).
Leafing Dates: Late May, when the water recedes (01,11).
Budding Dates: Late June (11,12).
Anthesis Dates: Mid-July to late August (09,11,12,16).
Fruiting dates: Late August to September (01,12,16).
Seed/Fruit Dispersal Dates: September (perhaps through winter)
(16).
Furbish's lousewort has a rosette growth form in the vegetative
state, with reproductive plants producing scapes up to 1 m tall. The
plants leaf out very early, before the overstory shrubs (01); maximum
basal leaf area is generally attained in late June (12). Developing
scapes are first visible in mid to late June. First flowering date
varies with season; in the warm, dry summer of 1983, the first flowers
opened on 12 July, whereas in cool wet 1984 anthesis did not occur
until approximately 10 days later (12). Capsules open in September,
but when the majority of seeds are actually dispersed is unclear (16).
SEX OR SPORE STATUS:
Monoclinous (09,23).
POLLINATION, SPORE AND SEED DISSEMINATION:
Pollen is disseminated solely by the bumblebee, Bombus vagans
(09). This species is a common bumblebee of eastern North America
which forages on many other species in this area. However, individual
bumblebees display high constancy to Pedicularis furbishiae, and the
plant species appears not to be pollinator limited (09,16).
Presumably seed is disseminated by wind and water (07,16).
Dispersal patterns and distances are unknown. Numerous seedlings are
commonly found beneath adult plants (07,12); like most plants, it
appears that a majority of P. furbishiae seeds fall close to the
parent. The light sculptured seed coat suggests wind or water
dispersal, and seeds placed in a jar of riverwater float for 24 hours
before sinking (12,16). Since the plant's habitat is frequently
inundated by spring floods, water may be an important dispersal agent.
Life History - 1 (DRAFT) - Life History
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
SEED BIOLOGY:
Only one year's seed production has been analyzed.
Inflorescences from five populations were sampled in 1983 to assess
seed production. Sites vary in the number of flowers initiated, the
level of pre-dispersal predation, and the number of seeds matured
(16). Averaging over all sites, however, the average plant matured
2 inflorescences (out of 5 initiated), initiated 25 flowers per
inflorescence, and matured 6.7 capsules from those 25 flowers. The
average healthy capsule contains 25.4 seeds. Thus, each reproducing
plant produced, on the average 340 seeds. Note, however, that there
is great variation within and between sites. At one site, for
example, 98% of the scapes were preyed upon by rabbits, and virtually
the whole seed crop was lost. Other populations lose inflorescences
to deer predation. A significant source of predation is the larva of
the lepidopteran Amblyptilia picta Wals. This predator, which devours
the seeds inside the developing capsule, destroyed approx. 40 pct.
of the developing capsules in 1983 (16). Spittlebugs reportedly
infested up to 91% of all scapes in 1977 (09), but only 30% of the
scapes in 1983 were infested; furthermore, while the spittlebug's
presence was obvious, it did not significantly reduce seed set
compared to scapes that escaped infestation (16). Predispersal seed
predation may be an important factor in maintenance of populations
(16).
Seeds required cold stratification before germination (10,12).
Seeds stratified on moist filter paper, in a refrigerator, for at
least 6 weeks, germinate readily (12). Field germination rates are
also high; over 100 seedlings were counted in an area 6 square dm
(12).
POPULATION BIOLOGY:
With only two years' data on a multi-year-lived perennial, our
knowledge of the population biology of P. furbishiae is still
incomplete. Results to date indicate the following:
Factors limiting the species have not been clearly identified.
The plants are not commonly found among dense shrub cover, and this
may be one limiting factor; the species appears to do best where the
bank is maintained in a semi-open state (11). Pre-dispersal seed
predation may be limiting replacement of individuals, growth of
existing populations, and/or establishment of new populations (16).
In 1983, within-season mortality both of seedlings and of established
plants averaged approximately 25% among sites (11). A more important
source of mortality appears to be destruction of colonies during
ice-out. In the one year for which we have data (1984), within-site
mortality ranged from 19% to 100%, and averaged 75% over all sites.
Three sites were removed completely (12). Year to year variation
undoubtedly exists; and original sampling may have been
inadvertently biased towards precarious sites.
Population density varies from site to site. At the densest
population, densities were 3.5 - 8.5 individuals per square meter,
except for one very dense site with 17.2 plants per square meter:
these figures include seedlings, and figures for established plants
would be lower (11).
Life History - 2 (DRAFT) - Life History
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
Population trends are extremely difficult to assess in this
species. A 1980 survey estimated the population at 5000 individuals,
including both U.S. and Canadian stations (13). Since 1980, severe
ice jams and floods have decimated several large colonies (12); on the
other hand, additional colonies have been discovered upriver (25,26).
In 1984, 4287 flowering stems were counted in the same area in which
4968 flowering stems had been counted in 1980, representing a decrease
of 14% (26). (The 1984 survey did not attempt a complete population
estimate.) Populations fluctuated wildly from 1980 to 1984 within the
area counted, however. In the downriver two-thirds of the range,
populations declined, some precipitously; but in the upriver third
(above Dickey), the discovery of new populations and the increased
counts at most known stations yielded an overall upriver increase
which almost offset the losses downriver. Note, however, that whether
the population upriver is really increasing is unclear, since the area
is relatively inaccessible, and previous population counts left many
stretches of riverbank uncovered.
Causes of mortality in P. furbishiae are not yet well understood.
Local ice scour and flooding remove some individuals every spring; but
these floods appear to be, on the whole, beneficial in maintaining the
habitat (11).
P. furbishiae appears to have good recovery potential if the
integrity of the St. John's watershed remains intact (05,11).
Recovery would be more difficult (and impossible in the true sense of
restoring natural processes) if the hydrologic regime of the river
were altered or if significant stretches of the immediate overstory
were removed. Zoning, if the current laws are enforced, should
prevent the latter. At least one source suggests that Furbish's
lousewort cannot be considered "recovered" until the natural
hydrologic regime of the upper St. John River is permanently protected
(33).
ECOLOGICAL/EDAPHIC FACTORS:
P. furbishiae grows on well-drained sandy loam banks with
moderate to abundant groundwater seepage (01,05,09). The upper St.
John River is underlain by Seboomook slate and graywacke, with some
Cary's Mill's carbonates in the downstream portion (31), but glacial
deposits form the substrate at almost all lousewort stations (01,09).
A very few stations occur on or near vertically fissured slate
outcrops (01,12). The complex postglacial stratigraphy of the upper
St. John River is dominated by lacustrine and alluvial sediments (31).
Parent material differs among sites, and varies in texture from gravel
outwash to para-glacial alluvium. Lacustrine sediments, including
some extensive clay deposits, occur at sites of former ice-jammed
lakes (31).
The classification of soils at particular lousewort sites is
problematic since soil maps for northwestern Aroostook County, where
most of the populations occur, have not been published. In general
terms, however, the soils along the St. John River fall into three
orders (32): Spodosols, the most well-developed soils, on older,
well-drained surfaces; Inceptisols, younger alluvial deposits, showing
weak horizonation; and Entisols, very young deposits with little soil
development. Soil series at the downriver sites (where soil maps are
Life History - 3 (DRAFT) - Life History
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
available) include the Van Buren and Caribou series (32). Other
series are likely involved upriver.
Soils at Furbish's lousewort sites are typically wet to very wet,
but also well-drained. The species is not found in areas of standing
water, although at several sites the soils are constantly saturated
with groundwater which seeps out of the banks and drains to the river
(11). Even at the drier sites, the maximum soil water tension
recorded was -72 centibars (11).
Soil texture at lousewort sites is fairly constant, falling
within the sandy loam to loamy sand range (11). The percentage of
gravel is the main factor distinguishing sites: It ranged from 2% to
29% (11). Soil pH is neutral to slightly acid. Fifty percent of the
samples from Maine stations were between 6.6 and 7.0 pH, although the
range extended down to pH 5.2 (09); pH at Canadian stations is higher,
7.2 to 7.8 (22). The soils are high in calcium but low in nitrogen
and organic matter (09).
In elevation, P. furbishiae is generally restricted to the steep
bank below the forest edge and above the river beach. Plants are
occasionally found one to four meters inside the forest edge (12) as
well as on sandy flats (11,12). Slopes at 6 sites studied in 1983
ranged up to 127% locally; the average slopes in the lousewort zone at
these sites was 64% to 107% (11). Almost all populations face NW, N,
or NE, since they grow on the south or east bank, and thus receive
direct sunlight for only a small part of the day. Most stations are
shaded further by trees overhanging the top of the bank. Canopy cover
from these trees obscures 23% to 59% of the overhead sky at the
elevations where the louseworts grow (11).
Disturbance appears to be important in regenerating habitat for
P. furbishiae. Ice scour and spring floods create open areas which
may then gradually revegetate, and they maintain other stretches of
bank in a semi-open state. While Furbish's lousewort is typically
found in areas which have developed a ground cover of moss or
graminoid litter (i.e., not the most highly disturbed sites), it is
absent from the most densely vegetated areas. At most sites, the
dominant shrubs are under 2 m high and do not form a continuous cover;
the lousewort tends to occupy the gaps (11,12).
TROPHIC STATUS:
P. furbishiae is phototrophic, but like most members of the
genus, and closely related genera, it is hemiparasitic at least in
seedling stages. Seedlings germinate and produce green leaves without
host attachment, but remain stunted and eventually die (10,24). The
plant is not host-specific, and has been grown in the greenhouse on
wheat, clover, and other plants (10,24,12). Root connections between
P. furbishiae and other plants have not been found in the field:
whether the root connections disappear after seedling stages or are
present, but difficult to excavate, is unknown. Longer greenhouse
studies may resolve this question. In the field, preliminary evidence
suggests that the hemiparasitism is not a strongly limiting factor:
seedling plots established in 1981 and 1983 shows moderate germination
and survivorship rates. In the 1981 plot, for example, while first
year growth was not followed, by the second year over 140 plants were
present on a 2 square meter plot, out of 500 seeds sown (12). The
Life History - 4 (DRAFT) - Life History
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
major implication for growth in the field is that Furbish's lousewort
cannot colonize newly scoured banks that are bare of vegetation (11).
CHARACTERISTIC DOMINANCE:
Component (21).
COMMUNITY ECOLOGY:
P. furbishiae grows in a riparian shrub/herb community. This
association, located below the forest edge and above the river bed,
is characterized by inundation of all or most of its vertical extent
at ice-out, with localized ice-scour; steep, unstable sand and gravel
banks; and wet but well-drained soils low in nitrogen and organic
matter but high in calcium (21).
A diversity of shrubs and herbs is commonly found with Furbish's
lousewort. [Nomenclature follows Gray's Manual of Botany (23).]
Dominant shrubs include Alnus spp., Salix spp., Cornus stolonifera,
Diervilla lonicera (at drier sites), and Myrica gale (at wetter
sites). The most common graminoids are Calamagrostis canadensis and
various sedges (Carex spp.). Equisetum arvense is abundant at most
sites. Common native forbs include Solidago spp., Aster spp., Rubus
pubescens, Viola spp., Fragaria virginiana, Thalictrum polygamum,
Conioselinum chinense, and Anemone canadensis. Several introduced
"weeds" are common, and indicate the disturbed nature of the
community: Vicia cracca is common everywhere, as are Trifolium
repens, T. pratense, Chrysanthemum leucanthemum and Taraxacum
officinale. Most remarkable about the community, however, are the
numerous rare or unusual "fugitive" calciphiles such as Castilleja
septentrionalis, Hedysarum alpinum var. americanum, Astragalus alpinus
var. brunetianus, Tanacetum huronense var. johannense, Listera
auriculata, Arnica mollis, Gentiana amarella, etc. These plants and
others (34 species in all) found here are considered rare in Maine,
New England, or the United States. (Listera auriculata, for example,
is under review for possible federal listing) (01). Most of these
species are commoner on rivers further north or in western North
America (28). Furbish's lousewort is not just one rare species in an
otherwise typical habitat, but is part of a highly unusual rivershore
community.
Particular community associations vary among and within sites.
Soil moisture is a major factor in the differences among sites, while
within a site the community is structured mostly by an elevational
gradient from the forest edge to the rivershore (21). Six
intergrading classes may be described along these major axes of
variation: boreal forest, wet steep banks, dryish upper banks, dryish
lower banks, semi-open wet flats, and flat sandy thickets (21). These
classes are not, however, discrete communities: this riparian shrub
community is phytosociologically cohesive (21).
Succesional trends in the community have not been documented.
The periodic hydrologic disturbance of the riverbank is presumably
important in successional patterns. If the successional trends are
from sparser to denser vegetation, as is true for certain disturbance
communities (30), the hydrologic action may be important for
maintaining the semi-open conditions that the species seems to require
(11).
Life History - 5 (DRAFT) - Life History
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
SPECIES INTERRELATIONSHIPS:
None specific. While the species is hemiparasitic, at least in
seedling stages, it is not host specific and appears to attach readily
to whatever hosts are present (10,24).
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 6 (DRAFT) - Management Practices
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Maintaining/Controlling Water Flow
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Restricting/regulating human use of habitats
Beneficial Land Acquisition
Beneficial Restricting Timber Harvest
Beneficial Transplanting Wild Eggs/Wild Seeds
Adverse Rural Residential/Industrial Areas
Existing Rural Residential/Industrial Areas
Adverse Strip mining
Existing Strip mining
Adverse Developing/maintaining stream bank vegetation
Existing Developing/maintaining stream bank vegetation
Adverse
Existing
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
A major reason for the rarity of Pedicularis furbishiae is its
natural endemism, i.e., its complete restriction to the shores of the
St. John River. Even within this small geographical range
(approximately 130 linear miles), only certain areas are suitable (see
Habitat narrative): habitat availability may be a limiting factor
(10) (note, however, that this is still unclear). Furthermore, the
natural hydrologic cycle of the river both destroys extant populations
(whole or in part) and creates habitat for new populations (01,05,11).
Thus, overall populations must be large enough to withstand years of
high mortality such as in 1984 when mortality averaged 75% over all
sites and 3 colonies were completely removed due to ice-out (12) (See
Biology Narrative, Population Biology).
Several human factors aggravate this natural rarity. In the
downriver two-thirds of its range, agricultural and residential
development of the river terraces has destroyed the tree canopy which
would normally be present above the rivershore shrub zone. This
canopy is an integral part of P. furbishiae's habitat, and this
clearing of land for fields and homes is thought to be one reason for
its scarcity in the lower part of its range (05,13). Agricultural
crops on the river terraces include potatoes and oats. Only when the
terraces are cleared too close to the river is this a problem: at the
Fort Kent site, for example, the high, steep wooded bank between
backyards and the river provides adequate protection for a large
population (01).
A similar problem is the clearing of riverbank trees for a view
of the river. This seems to be increasing, as more people are
building homes and camps close to the rivershore (06). At least three
populations have been significantly damaged by clearing for scenic
views in the last four years (14).
Forest clearing in the St. John River basin, and its effects on
Management Practices - 1 (DRAFT) - Management Practices
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
surface drainage, may indirectly affect lousewort populations. The
mean daily flow and size of the annual flood have both increased over
the last several decades (15). Valley residents, including local
loggers, attribute these changes to increased timber harvest and
clearcuts (14); whether these increases are actually correlated with
cutting rates and/or weather patterns is unknown. At any rate, the
increase in the size of the annual spring flood means an increased
rate of disturbance to the riverbanks. Pedicularis furbishiae appears
to need a delicate balance between disturbance and revegetation (11),
and whether the hydrologic changes will seriously upset that balance
remains to be seen.
The effects of water quality on the populations are unknown. The
highest quality waters are upriver of Fort Kent, where the bulk of the
population occurs; lower quality water in the downstream portion of
the range corresponds to the widely spaced, smaller populations (17).
Many factors other than water quality could be responsible for this
pattern.
Inherent characteristics which may contribute to the species'
rarity are also unknown. A small gene pool (founder effects) is
possible if colonization of the St. John River was a chance and
one-time event; but no studies of genetic variation within the species
have been done. Life-cycle characteristics ("bottlenecks")
contributing to the species' rarity are being studied (11,12,16) but
several years' data will be necessary before we can draw conclusions.
In summary: Pedicularis furbishiae is naturally rare because of
its endemism. Human impacts on the river known to increase the rarity
are 1) agricultural clearing and residential development in the lower
two-thirds of its range, and 2) bank clearing for "scenic views",
throughout the range but especially around Allagash. The latter
appears to be increasing while the former appears stable.
The major future threat to the species is alteration of the
hydrologic regime (05,06). A dam (such as the formerly proposed
Dickey-Lincoln School Lakes Project) would have two major effects:
1) inundation of some populations and 2) cessation of the hydrologic
disturbance regime which appears necessary for habitat regeneration
(05). Comparable "disturbance" communities on Swedish rivers have
been gradually eliminated after flow stabilization (18).
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1983. Recovery Plan for the Furbish
Lousewort. U.S. Fish and Wildlife Service, Newton Corner, MA. 31 pp.
Actions necessary for recovery of the Furbish lousewort include:
(1) Protection of the St. John River Riparian Ecosystem, which
involves, among other things, utilizing existing state legislation to
protect habitat. For example, implement the 1982 St. John River
Resource Protection Plan, for unorganized portions of the river, which
prohibits commercial and residential development, water impoundments,
utility facilities, regulates timber harvesting, and restricts
construction of roads, bridges, and gravel pits.
Management Practices - 2 (DRAFT) - Management Practices
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
(2) Maintain the 28 known colonies and establishing ten new
reproducing colonies on the upper St. John River by means of
transplanting wild seed.
(3) Utilize land agreements and purchases to secure habitat. Ensure
that community zoning standards provide adequate protection for the
species.
(4) Establish and maintain international cooperation for protection
efforts.
(5) Monitor the population and conduct species biology studies and
habitat surveys.
(6) Develop management recommendations for individual colonies.
All of the actions above are currently underway or completed.
Management Practices - 3 (DRAFT) - References
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Gawler, S.C. 1983. Furbish's Lousewort (Pedicularis furbishiae
S. Wats.) in Maine and its relevance to the Critical Areas Program.
Planning Rep. No.13, State Planning Office, Augusta, ME. 69 pp.
02 Furbish, K. 1881. A botanist's trip to "The Aroostook". Am.
Nat. 15:469-470.
03 Macior, L.W. 1977. Physiological studies on Pedicularis
furbishiae, Allagash, Maine. U.S. Army Corps of Engineers, Waltham
MA. 24 pp.
04 Kartesz, J.T. and R. Kartesz. 1980. A Synonymized Checklist of
the Vascular Flora of the United States, Canada, and Greenland.
Vol. II., The Biota of North America. Univ. of North Carolina
Press, Chapel Hill. 498 pp.
05 U.S. Fish and Wildlife Service. 1983. Furbish Lousewort Recovery
Plan. U.S. Fish and Wildlife Service, Newton Corner, MA. 31 pp.
06 Crow, G.E. 1982. New England's Rare, Threatened, and Endangered
Plants. U.S. Gov. Print. Off., Washington, D.C. 129 pp.
07 Stirrett, G.M. 1980. The status of Furbish's Lousewort,
Pedicularis furbishiae S. Wats., in Canada and the United States.
Canadian Wildlife Service, Ottawa, Ont. 78 pp.
08 Macior, L.W. 1981. The Furbish Lousewort--weed, weapon, or
wonder? The Amer. Biol. Teacher 43:323-326.
09 Macior, L.W. 1978. The pollination ecology and endemic adaptation
of Pedicularis furbishiae S. Wats. Bull. Torrey Bot. Club
105:268-277.
10 Macior, L.W. 1980. The population ecology of Furbish's Lousewort
(Pedicularis furbishiae S. Wats.). Rhodora 82:105-111.
11 Gawler, S.C., D.M. Waller, and E.S. Menges. In prep.
Environmental factors affecting establishment and growth of
Pedicularis furbishiae S. Wats., a rare endemic of the St. John
River Valley, Maine. Botany Dept., Birge Hall, Univ. Wis.,
Madison, WI 53706.
12 Gawler, S.C., E.S. Menges, and D.M. Waller. Unpublished data.
Field season notes of 1983-1984. Botany Dept., Birge Hall, Univ.
Wis., Madison, WI 53706.
13 Richards, C.D. 1980. Report on monitoring populations of
Furbish's Lousewort, Pedicularis furbishiae, along the St. John
River in northern Maine and New Brunswick during the summer of
1980. Off. Endang. Spec., U.S. Fish and Wildlife Service, Newton
Corner, MA. 14 pp.
14 Gawler, S.C. 1983. Unpublished data. Field notebook. Botany
Dept., Birge Hall, Univ. Wis., Madison, WI 53706.
15 Gawler, S.C. 1984. Unpublished manuscript. Hydrologic and
vegetational interactions along the upper St. John River, Maine.
32 pp.
16 Menges, E.S., D.M. Waller, and S.C. Gawler. 1984. Seed set and
seed predation in Pedicularis furbishiae, a rare endemic of the
St. John River. Submitted to Am. J. Bot.
17 New England River Basins Commission. 1981. St. John River Basin
Overview. Public Review Draft. 141 pp.
References - 1 (DRAFT) - References
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
18 Grelsson, G. and C. Nilsson. 1980. Colonization by Pinus
sylvestris of a former middle-geolittoral habitat on the Umealven
River in northern Sweden, following river regulation for
hydro-electric power. Holarctic Ecol. 3:124-128.
19 Cowardin, L.M., V. Carter, F.C. Golet, and E.T. LaRoe. 1979.
Classification of wetland and deepwater habitats of the United
States. U.S. Fish and Wildlife Service. FWS/OBS-79/31.
Washington, D.C. 103 pp.
20 U.S. Geological Survey. Hydrologic records for the St. John River
at Ninemile, Dickey, and Fort Kent, from begining to record to
present. U.S. Geological Survey, Augusta, ME.
21 Gawler, S.C. 1983. Unpublished manuscript. Community relations
of Pedicularis furbishiae S. Wats. Botany Dept., Birge Hall, Univ.
Wis., Madison, WI 53706.
22 Brown, D.C. 1982. Summer study of potential ecological reserves.
A report to the Environmental Council of New Brunswick. Dept. Nat.
Resour., Fredericton. 24 pp.
23 Fernald, M.L. 1950. Gray's Manual of Botany. Eighth ed.
American Book Co., New York. 1632 pp.
24 Wheeler, C.E. 1980. Seedling development and root parasitism in
Pedicularis furbishiae S. Wats. M.S. Thesis, Univ. Akron. 61 pp.
25 Gawler, S.C. 1982. Report on landowner contact program for
Furbish's Lousewort (Pedicularis furbishiae) in the St. John River
Valley, Maine. Off. Endang. Spec., U.S. Fish and Wildlife Service,
Newton Corner, ME. 23 pp.
26 Menges, E.S., S.C. Gawler, and D. Lawler. In prep. Population
fluctuations in Pedicularis furbishiae, 1980-1984.
27 Hurst, L.A., E.W. Knobel, and B.H. Hendrickson. 1917. Pages 7-46
+ map. 19th Report, Field Operations of the Bureau of Soils. U.S.
Department of Agriculture.
28 Garrison, G.A., A.J. Bjugstad, D.A. Duncan, M.E. Lewis, and D.R.
Smith. 1977. Vegetation and Environmental Features of Forest and
Range Ecosystems. U.S. Department of Agriculture, Agric. Handbk.
475. 68 pp.
29 U.S. Geological Survey. Topographic Maps, 15' series. Maine:
Eagle Lake, St. Francis, Allagash, Round Pond, Rocky Mountain, and
Van Buren quadrangles. U.S. Geological Survey, Washington, D.C.
30 White, P.S. 1979. Pattern, process and natural disturbance in
vegetation. Bot. Rev. 45:229-299.
31 Kite, J.S. 1983. Late quaternary glacial, lacustrine, and
alluvial geology of the upper St. John River basin, northern Maine
and adjacent Canada. PhD. Dissertation, Univ. Wis. 339 pp.
32 Arno, J.R. 1964. Soil survey of Aroostook County, Maine:
northeastern part. Soil Conservation Service, U.S. Department of
Agriculture, Series 1958 no. 27. 80 pp., 176 sheets.
33 Gawler, S.C. 1984. Unpublished data. Botany Dept., Birge Hall,
Univ. Wis., Madison, WI 53706.
34 Watson, S. 1882. Proc. Am. Acad. 17:375.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 Gawler, S.C. 1983. Furbish's Lousewort (Pedicularis furbishiae)
References - 2 (DRAFT) - References
Species LOUSEWORT, FURBISH
Species Id ESIS705003
Date 13 MAR 96
in Maine and its relevance to the Critical Areas Program. Planning
Rep. No.13, State Planning Office, Augusta, ME. 69 pp.
02 Menges, E.S., S.C. Gawler, and D. Lawler. In prep. Population
fluctuations in Pedicularis furbishiae, 1980 - 1984.
03 Richards, C.D. 1980. Report on monitoring populations of
Furbish's Lousewort, Pedicularis furbishiae, along the St. John
River in northern Maine and New Brunswick during the summer of
1980. Off. Endang. Species, U.S. Fish and Wildlife Service,
Newton Corner, MA. 14 pp.
04 Gawler, S.C. 1985. Personal communication. Botany Dep., Birge
Hall, Univ. Wisconsin, Madison, WI 53706.
References - 3