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Wildlife, Animals, and Plants
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Introductory
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
ABBREVIATION :
CLASPP
SYNONYMS :
Cladonia alpestris = C. stellaris
Cladonia arbuscula = C. sylvatica
SCS PLANT CODE :
NO-ENTRY
COMMON NAMES :
reindeer lichen
lichen
reindeer moss lichen
caribou lichen
TAXONOMY :
The most widely accepted genus name for reindeer lichens is Cladonia
Wigg. (subgenus Cladina (Nyl.) Vain) [13,35,40]. The characters derived
from the mycobiont (the fungal member of a lichen) have been given most
weight in the recognition of orders, families, and genera of lichen.
The characters used to separate these categories are now almost
exclusively mycological, but genera largely based on such features have
not found general acceptance [13]. Some authorities separate the
reindeer lichens into the genus Cladina, while others refer to it as
subgenus Cladina of the genus Cladonia [40]. In this report taxonomy
will follow the latter, with emphasis on the four species of reindeer
lichens listed below:
Cladonia alpestris (L.) Rabenh.
Cladonia arbuscula (Wallr.) Rabenh.
Cladonia rangiferina (L.) Nyl
Cladonia mitis Sandst.
LIFE FORM :
Lichen
FEDERAL LEGAL STATUS :
See OTHER STATUS
OTHER STATUS :
Cladonia perforata is federally listed as endangered [45].
COMPILED BY AND DATE :
Julie L. Tesky, August 1992.
LAST REVISED BY AND DATE :
NO-ENTRY
AUTHORSHIP AND CITATION :
Tesky, Julie L. 1992. Cladonia spp. In: Remainder of Citation
DISTRIBUTION AND OCCURRENCE
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
GENERAL DISTRIBUTION :
Reindeer lichens have a circumpolar distribution. In North America,
they occur throughout Alaska, Canada, and the northern United States
southward to Florida and Alabama [11,19].
National Park occurrences are listed below by species.
Cladonia alpestris
ACAD APOS BELA CAKR DENA DETO DEWA GAAR GATE
GLAC GRSO HOSP INDU ISRO KATM KOVA OLYM PIRO
SACR SHEN SLBE THRO VOYA YELL
Clandonia arbuscula
ACAD APIS BELA CAKR DENA DEWA GAAR GLAC GRSM
ILMI ISRO OLYM PIRO VOYA YELL YUCH
Cladonia rangiferina
ACAD APIS BELA CAKR CUVA DENA DEWA GAAR GRSM
INDU ISRO KOVA MORA NOAT NOCA OLMY PIRO SACR
SARA SHEN SLBE THRO VOYA YELL YUCH
Cladonia mitis
ACAD APIS CAKR GAAR GLAC GRSM INDU ISRO KATM
OLYM PIRO SACR SLBE THRO VOYA YELL
ECOSYSTEMS :
FRES10 White - red - jack pine
FRES11 Spruce - fir
FRES18 Maple - beech - birch
FRES19 Aspen - birch
FRES20 Douglas-fir
FRES22 Western white pine
FRES23 Fir - spruce
FRES24 Hemlock - Sitka spruce
FRES25 Larch
FRES26 Lodgepole pine
FRES28 Western hardwoods
FRES44 Alpine
STATES :
AL AK AR CA CO CT DE FL GA ID
IL IA MD ME MI MN MT NH NJ NY
NC ND OR PA RI SC SD UT VT VA
WA WI WY AB BC MB NB NF NT NS
ON PE PQ SK YT
ADMINISTRATIVE UNITS :
SEE GENERAL DISTRIBUTION
BLM PHYSIOGRAPHIC REGIONS :
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
13 Rocky Mountain Piedmont
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands
KUCHLER PLANT ASSOCIATIONS :
K001 Spruce - cedar - hemlock forest
K002 Cedar - hemlock - Douglas-fir forest
K003 Silver fir - Douglas-fir forest
K004 Fir - hemlock forest
K005 Mixed conifer forest
K008 Lodgepole pine - subalpine forest
K013 Cedar - hemlock - pine forest
K014 Grand fir - Douglas-fir forest
K015 Western spruce - fir forest
K020 Spruce - fir - Douglas-fir forest
K021 Southwestern spruce - fir forest
K093 Great Lakes spruce - fir forest
K094 Conifer bog
K095 Great Lakes pine forest
K096 Northeastern spruce - fir forest
K097 Southeastern spruce - fir forest
K106 Northern hardwoods
K017 Black Hills pine forest
K108 Northern hardwoods - spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak - pine forest
K111 Oak - hickory - pine forest
SAF COVER TYPES :
1 Jack pine
5 Balsam fir
12 Black spruce
13 Black spruce - tamarack
18 Paper birch
30 Red spruce - yellow birch
31 Red spruce - sugar maple - beech
32 Red spruce
33 Red spruce - balsam fir
34 Red spruce - Fraser fir
35 Paper birch - red spruce - balsam fir
37 Northern white-cedar
38 Tamarack
107 White spruce
201 White spruce
202 White spruce - paper birch
203 Balsam poplar
204 Black spruce
205 Mountain hemlock
206 Engelmann spruce - subalpine fir
251 White spruce - aspen
252 Paper birch
253 Black spruce - white spruce
254 Black spruce - paper birch
SRM (RANGELAND) COVER TYPES :
NO-ENTRY
HABITAT TYPES AND PLANT COMMUNITIES :
Reindeer lichens commonly occur as the dominant or codominant ground
cover on open sites dominanted by white spruce (Picea glauca), black
spruce (P. mariana), paper birch (Betula papyrifera), or jack pine
(Pinus banksiana). The white spruce/Sphagnum spp.-Cladonia spp.
community type described by Foote [12] is characterized by woodland or
open forests often occurring on valley bottoms or on north-facing slopes
where ice-rich permafrost is present. In this community type, Cladonia
rangiferina and C. arbuscula grow intermixed with Sphagnum spp.,
Schreber's moss (Pleurozium schreberi), and Polytrichum spp. [12]. The
black spruce/bog birch (Betula glandulosa)/Cladonia community type
decribed by Viereck [39] is fairly common on well-drained sites and near
treeline in interior, southwest, and northwest Alaska. Published
classification schemes identifying Cladonia spp. as ground cover
dominants or codominants are as follows:
Classification, description and dynamics of plant communities following
fire in the taiga of interior Alaska [12].
A preliminary classification system for vegetation of Alaska [39].
VALUE AND USE
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
WOOD PRODUCTS VALUE :
NO-ENTRY
IMPORTANCE TO LIVESTOCK AND WILDLIFE :
Terrestrial lichens are important in the winter diet of caribou. In the
forest zones of Ontario, the winter diet of caribou is made up almost
exclusively of reindeer lichens [2]. In some places these lichens
constitute over 50 percent of the caribou diet [2,21]. The winter diet
of Newfoundland caribou is made up mostly of Cladonia mitis and C.
rangiferina and to a lesser extent C. alpestris. Within the range of
the Newfoundland caribou C. mitis seems to be the most important
food-lichen. In Norway reindeer seem to prefer pastures with plenty of
C. rangiferina to those with plenty of C. alpestris [41]. Reindeer
lichens are also eaten by caribou throughout the summmer, although to a
lesser extent [2].
PALATABILITY :
Reindeer lichens are highly palatable to both barren ground and woodland
caribou [2]. Cladonia rangiferina contains bitter fumarprotocetraric
acid while C. alpestris and C. mitis do not contain this acid. It is
questionable whether this difference in reindeer lichens affects their
palatability to reindeer [41].
NUTRITIONAL VALUE :
The nutritive value of reindeer lichens is rather poor. The protein
content, calcium and phosphorous levels, and vitamin content are very
low. The calcium/phosphorous ratio is rated as poor [2,34]. The crude
fat in reindeer lichens is mainly lichen acids, which are largely
indigestible. However, the nitrogen-free extract of these lichens is
thought to be highly digestible. Crude fiber content is high. Reindeer
lichens are high in digestible carbohydrates, mostly in the form of
complex starches, and therefore are a good source of energy [17,34].
Ahti [2] listed the percent chemical composition of Cladonia species as
follows:
Carbohyrates= 93.4 to 94.4 percent
minerals= 1.1 to 1.5 percent
protein= 2.6 to 2.9 percent
fat= 1.8 to 2.2 percent
Although caribou can survive the winter on an exclusive reindeer lichen
diet, the deficiency of protein usually results in a sharp decrease in
body weight as the animal breaks down its own muscular tissue to
compensate for the protein deficiency [30,34]. Caribou appear to be
able to balance the low protein content of the reindeer lichens by
including in their diet a portion of the nitrogen-fixing lichens of the
genera Stereocaulon and Peltigera which have relatively high protein
contents [18]. Additionally, caribou can digest reindeer lichens more
efficently than other ruminants can [17]. The crude fiber is broken
down by rumen bacteria and protozoa which liberate large amounts of
energy [34].
COVER VALUE :
NO-ENTRY
VALUE FOR REHABILITATION OF DISTURBED SITES :
NO-ENTRY
OTHER USES AND VALUES :
NO-ENTRY
MANAGEMENT CONSIDERATIONS :
Reindeer lichen growth is so slow that it should not be regrazed by
caribou for 2 to 5 years if the grazing is moderate, or 10 to 15 years
if the grazing is intense. Under intense grazing the ground may be
almost totally depleted of reindeer lichens, with only the basal part of
the podetia (upright growing branches) remaining [30]. At the Taltson
River Region in the Northwest Territories, Cladonia alpestris had an
average annual growth rate of 0.11 inch (3.4 mm) per year while C.
rangiferina had an average annual growth rate of 0.14 inch (4.1 mm) per
year [31]. C. alpestris has many lateral branches compared with C.
rangiferina or C. mitis, and could produce more forage annually even
though the annual linear growth rate may be less [25]. C. alpestris,
however, can not withstand heavy grazing and trampling. Therefore, it
can be the main reindeer lichen food only in areas with a comparatively
small caribou population [41].
High relative humidity is especially important for reindeer lichens.
Studies have shown that a 9-month fully dry period is lethal to Cladonia
rangiferina, but under natural conditions much shorter periods are
critical [2].
BOTANICAL AND ECOLOGICAL CHARACTERISTICS
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
GENERAL BOTANICAL CHARACTERISTICS :
Reindeer lichens are slow-growing, long-lived, densely branched ground
lichens with short numerous outer branchelets [2]. These lichens often
form clumps or mats composed of a large number of podetia [30]. The
podetia are slender, elongated, and branched in whorls. They are often
densely intertangled in large colonies [30]. The slender thalli of
Cladonia rangiferina are hollow stems of very low density which are
finely branched and not only have a high surface to volume ratio, but
also have branches advantageously distributed to carry fire [28].
RAUNKIAER LIFE FORM :
NO-ENTRY
REGENERATION PROCESSES :
The dispersal of reindeer lichens mainly occurs by means of thallus
fragments and to a much smaller extent by ascopores. Wind is the most
important dispersal agent. Reindeer lichens grow vegetatively by
annually producing new growth at the top of the podetium, which
lengthens the internode formed in previous years [2].
The podetium of Cladonia spp. passes through three growth stages. The
first stage, called the growth-accumulation period, lasts an average of
10 years but can vary from 5 to 25 years. During this stage no part of
the podetium dies off. During the second stage, called the
growth-renewal period, the podetium grows at its highest rate but dies
off at the base at a rate equal to the growth. This stage often exceeds
100 years. During the third stage, the podetium degeneration period,
the podetium dies off at a greater rate than it grows. This stage may
also exceed 100 years. Factors that probably contribute to variation in
lichen growth include: the age of the podetium; prior disturbance by
animals; and site conditions such as substrate, drainage, and exposure
[25].
SITE CHARACTERISTICS :
Reindeer lichens typically occur in submontane to alpine environments.
They are scattered to plentiful in the open or in open-canopy forest and
tundra on well-drained, water-shedding sites with shallow and/or
coarse-skelatal soils. In these environments they often occur on soil
and sometimes on rocks, stumps, and logs [19]. Cladonia arbuscula and
C. mitis are the most competitive reindeer lichens on rock [2].
Northern boreal forests offer climatically optimal conditions for
reindeer lichen growth largely because of slow plant succession and
little competition from other plant forms. Dry reindeer lichen
woodlands and lichen bogs are characteristic throughout the northern
boreal forest belt [2]. On the Slate Islands of Lake Superior, Ontario,
reindeer mosses grow best on dry open sites. Here, they occur more than
twice as frequently in dry regime forest types than in moist regime
forest types of the same composition and are more than ten times as
abundant in the dry types [8].
Cladonia rangiferina has a wider ecological amplitude than other
reindeer lichens and is thus more common than the others in less
favorable habitats such as wet bogs and shaded woods. It is very common
all over northern Ontario, being the most widely distributed ground
lichen in the area. Even in the Cladonia alpestris stands of the
northern boreal forest it is considerably more plentiful than C. mitis
or C. arbuscula [2].
Cladonia arbuscula prefers a moister and more shaded habitat than most
reindeer lichens but is frequently found mixed with C. mitis. C. mitis
is commonly one of the dominants on rocks, in lichen woodlands, and on
dry bog hummocks [2].
Soils and climate: Reindeer lichens commonly occur on moist to very
dry, sandy, nitrogen-poor soils [19] on shallow humus layers or dry peat
[2]. They are adapted to a cool, moist climate. Most reindeer lichens
avoid calcareous soils and prefer the acid humus of podzolic soils
during germination [32]. Reindeer lichens have been found on sites with
pH values ranging between 4.5 and 5.5 [32]. Since reindeer lichens are
able to take up moisture from the air, the underlying soil is not as
important a source of moisture as it is to vascular plants. Reindeer
lichens, therefore, can colonize and become a dominant floral element on
soils too shallow or sterile to support higher plants, provided that
humidity is sufficently high for lichen growth and temperature is
sufficently low to inhibit competitors [2].
Plant associates: Reindeer lichens are commonly found associated with
the following species: whortleberry (Vaccinium myrtillus), rock
cranberry (V. vitris-idaea), bog blueberry (V. uliginosum), lowbush
blueberry (V. angustifolium), bog birch, sheep laurel (Kalmia
angustifolia), common bearberry (Arctostaphylos uva-ursi), black
crowberry (Empetrum nigrum), Stereocaulon paschale, and Schreber's moss
(Pleurozium schreberi) [6,7,29].
SUCCESSIONAL STATUS :
Facultative Seral Species (C. mitis)
Obligate Climax Species (C. alpestris, C. arbuscula, C. rangiferina)
Reindeer lichens are shade intolerant [19]. Cladonia mitis is an early
to mid-seral species, while C. alpestris, C. arbuscula, and C.
rangiferina are late-seral to climax species [25]. C. mitis is generally
the first reindeer lichen to become established in postfire succession
of white spruce and black spruce stands. This lichen dominates for 30
to 40 years and is then replaced by other reindeer lichens such as C.
alpestris and C. rangiferia. If the canopy becomes closed, reindeer
lichens are generally replaced by the shade-tolerant mosses such as
mountain fern moss (Hylocomium splendens) and Schreber's moss [2].
SEASONAL DEVELOPMENT :
NO-ENTRY
FIRE ECOLOGY
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
FIRE ECOLOGY OR ADAPTATIONS :
Reindeer lichens are not well adapted to fire. They are highly
flammable and may take 30 to 100 years or more to recover to prefire
densities [32].
POSTFIRE REGENERATION STRATEGY :
NO-ENTRY
FIRE EFFECTS
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
IMMEDIATE FIRE EFFECT ON PLANT :
Reindeer lichens can survive cool fires but are almost always killed by
severe fire [16,26,40]. In the black spruce zone, lichens generally
burn poorly in early morning or near sunset, even on hot days, whereas
at mid-day they flare up with almost incredible heat and flame.
Humidity changes in the microclimate at ground level and dehydration of
the lichens appear to be the most likely factors involved. Fifty
percent or more of the lichens may survive if there is not much organic
litter to retain the fire [16]. Following fire in a black spruce
community, all the Cladonia spp. survived a light burn, whereas none
survived a heavy burn [38].
DISCUSSION AND QUALIFICATION OF FIRE EFFECT :
PLANT RESPONSE TO FIRE :
Reindeer lichens recover very slowly after fire [22]. The length of
time required for full recovery varies with species, the extent and
intensity of the fires, and site and microclimatic condition, but an
average of 40 to 50 years appears to be a conservative estimate [22,40].
Based on annual growth rates of 0.14 and 0.16 inch (4.1 and 4.9 mm) for
C. alpestris and C. rangiferina, it has been estimated that these
species would require nearly a century to reach prefire abundance [21].
After fire the first reindeer lichen to become established is Cladonia
mitis. The second reindeer lichen phase is generally dominated by C.
alpestris, C. rangiferina, or C. arbuscula [22,40].
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE :
One hundred forty years after a severe wildfire in Sweden, Cladonia
alpestris still showed no recovery, but did show good recovery only 20
years after a light, controlled burn. Slow recovery rates were reported
from the Seward Peninsula of Alaska, while rapid recovery rates have
been reported from the open lichen woodlands of Newfoundland, where the
climate is warmer than in Alaska [40].
In a black spruce and jack pine woodland in northwestern Manitoba and
northeastern Saskatchewan, Cladonia mitis became established in less
than 40 years after fire, while C. alpestris and C. rangiferina first
appeared in stands greater than 40 years of age. A dense growth of
reindeer lichens was found in stands that had not burned for at least
150 years. In south-central Alaska it took 30 to 40 years for C.
rangiferina and C. arbuscula to recover after fire [25]. In a postfire
black spruce-lichen vegetation type of interior Alaska, cover values for
Cladonia species were 32 percent in 26- 50-year-old stands, and 41
percent in stands greater than 100 years [25]. In a lightly burned
75-year-old stand with a open canopy of spruce and occasional jack pine,
C. alpestris made up 32.8 percent of the total ground cover and C.
rangiferina made up 1.7 percent [16].
FIRE MANAGEMENT CONSIDERATIONS :
Reindeer lichens are highly flammable. Cladonia rangiferina collected
from Ely, Minnesota, and ovendried had a heat value of 4,360 cal/g [15].
They dry rapidly during periods of low atmospheric humidity because of
the absence of roots, water storage tissues, and low resistance to water
loss. Reindeer lichens resemble dead litter more than live tissue in
their susceptibility to fire. Continuous mats of reindeer lichen
present an uninterrupted surface along which a fire spreads. Lichen
mats also typically accumulate tree and shrub litter which adds to the
flammability [3]. In black spruce-Cladonia alpestris woodland, litter
suspended in the lichen mat added 20.5 percent dry weight to the total
combustible material present above the soil [3].
While the destruction of reindeer lichens may have an immediate effect
on the winter range of caribou, some studies indicate that at least
infrequent fire is necessary to maintain optimum lichen cover [40]. In
the northern boreal lichen belt, lichen supplies could be increased by
burning Sphagnum fuscum peatlands, treeless bogs, or wooded muskegs.
The result of severe fire is an almost solid reindeer lichen stand in
some 40 to 50 years. Because black spruce and mosses regenerate more
slowly than lichen on these sites, good lichen growth persists for at
least 100 years [2]. Light burning has been suggested as a method to
improve reindeer range in Scandinavia [40].
REFERENCES
SPECIES: Cladonia (Cladina) spp. | Reindeer Lichen
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Index
Related categories for Species: Cladonia (Cladina) spp.
| Reindeer Lichen
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