Fire Effects Information System (FEIS)

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• Introductory
• Distribution and occurrence
• Botanical and ecological characteristics
• Fire ecology
• Fire effects
• Management considerations
• References

INTRODUCTORY

	AUTHORSHIP AND CITATION FEIS ABBREVIATION NRCS PLANT CODE COMMON NAMES TAXONOMY SYNONYMS LIFE FORM FEDERAL LEGAL STATUS OTHER STATUS
	Cladonia stygia	C. stellaris
	© Sylvia and Stephen Sharnoff

AUTHORSHIP AND CITATION:
Munger, Gregory T. 2008. Cladonia spp. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov /database/feis//lichens/claspp/all.html [].

FEIS ABBREVIATION:
CLASPP
CLAARB
CLAMIT
CLARAN
CLASTE
CLASYG

NRCS PLANT CODE [135]:
CLST60
CLADO3
CLAR60
CLMI60
CLRA60
CLST5

COMMON NAMES:
Common names are rarely used for reindeer lichens, so this report uses scientific names when referring to individual species. Nevertheless, the following common names were occasionally encountered in the published literature:

for Cladonia arbuscula:
tree reindeer lichen
littletree reindeer lichen
shrubby reindeer lichen
woodland reindeer lichen

for Cladonia mitis:
green reindeer lichen
spineless reindeer lichen

for Cladonia rangiferina:
gray reindeer lichen
graygreen reindeer lichen

for Cladonia stellaris:
star-tipped reindeer lichen
alpine reindeer lichen
star reindeer lichen

for Cladonia stygia:
black-footed reindeer lichen
Styx reindeer lichen

TAXONOMY:
Many systematists place reindeer lichens in the genus is Cladonia P. Browne [3,4,56,64,124], although some systematists place reindeer lichens in the genus Cladina [24,135]. Reindeer lichens are in the family Cladoniaceae [3,4,24,56,64,124,135].

The taxonomic status of Cladonia spp. is in question [124]. A study of phylogenetic relationships within Cladonia and Cladina species based on DNA sequences, morphology, and chemical analyses led some researchers to conclude that Cladina is best treated as a subgenus of Cladonia [4,124].

The reindeer lichens discussed in this review are listed below.

Cladonia arbuscula (Wallr.) Flotow [4,64,124], tree reindeer lichen
    Cladonia arbuscula subsp. arbuscula (Wallr.) Flot. [3,64]
    Cladonia arbuscula subsp. beringiana (Ahti) N.S. Golubk. [3,23,64]
    Cladonia arbuscula subsp. boliviana (Ahti) Ahti & DePri
    Cladonia arbuscula subsp. imshaugii (Ahti) Ahti & DePriest
    Cladonia arbuscula subsp. pachyderma (Ahti) Ahti & DePriest
    Cladonia arbuscula subsp. squarrosa (Wallr.) Ruoss
    Cladonia arbuscula subsp. stictica Ruoss [64]

Cladonia mitis Sandst. [4,64,124], green reindeer lichen

Cladonia rangiferina (L.) Weber ex F.H. Wigg. [4,64,124], gray reindeer lichen
    Cladonia rangiferina subsp. abbayesii (Ahti) Ahti & DePriest
    Cladonia rangiferina subsp. grisea Ahti
    Cladonia rangiferina subsp. rangiferina (L.) Weber ex F.H. Wigg.

Cladonia stellaris (Opiz) Pouzar & Vezda [4,24,124,135], star-tipped reindeer lichen

Cladonia stygia (Fr.) Ruoss [4,64,124], black-footed reindeer lichen

SYNONYMS:
for Cladonia arbuscula:
Cladina arbuscula (Wallr.) Hale & Culb. [24,85,135]
Cladonia arbuscula var. arbuscula (Wallr.) Flot. (cited in [64])
Cladonia sylvatica (L.) Hoffm. [56]

for Cladonia mitis:
Cladina mitis (Sandst.) Hustich [24,85,135]
Cladonia arbuscula subsp. mitis (Sandst.) Ruoss
Cladonia arbuscula var. mitis (Sandst.) Sipman (cited in [64])

for Cladonia rangiferina:
Cladina rangiferina (L.) Nyl. [24,85,135]
Cladonia rangiferina var. abbayesii Ahti
Cladonia rangiferina var. rangiferina (L.) Weber ex F.H. Wigg. (cited in [64])

for Cladonia stellaris:
Cenomyce stellaris Opiz (cited in [64])
Cladina stellaris (Opiz) Brodo [22]

for Cladonia stygia:
Cladina stygia (Fr.) Ahti [24,135]

DISTRIBUTION AND OCCURRENCE

• GENERAL DISTRIBUTION
• HABITAT TYPES AND PLANT COMMUNITIES

Cladonia arbuscula is widely distributed in Canada and the northern half of the United States. In western North America, C. arbuscula occurs in Alaska [11,12] and western Canada [54] south to Oregon and west to Idaho, western Montana, Wyoming [84,85], and Colorado [23]. Cladonia arbuscula is uncommon in the Pacific Northwest and rare in the Southwest [85]. In eastern North America, C. arbuscula occurs as far south as the northern Great Lakes states [133,143] and New England [37,47]. Cladonia arbuscula occurs as far east as Labrador [49] and Newfoundland [1]. Although C. arbuscula was reported from North Carolina's coastal plain in 1931 [142], more recent sightings were lacking as of 2008.

Cladonia mitis occurs in Alaska, Canada, and the northern fringe of the United States [18,84,117,127]. In western North America, C. mitis occurs as far south as the southern Willamette Valley [61,85], the northern Cascade Range [38], Idaho, western Montana, and Wyoming [84,85]. Cladonia mitis is uncommon in the Pacific Northwest and rare in the Southwest [85]. In eastern North America, C. mitis occurs in the northern Great Lakes region [65,133,143], New Jersey [117], southeastern New York [37], Labrador [49], Newfoundland [1], Quebec [48,90], and the boreal and arctic vegetation zones in Ontario [3].

Cladonia rangiferina is relatively widespread in the arctic and temperate zones of the United States and Canada [3,23]. There are reports of C. rangiferina near its apparent southern distributional limits of Oregon, Montana [85], Minnesota [65], Wisconsin [133], Michigan [62,143], West Virginia [35], and New Jersey [93].

Cladonia stellaris and C. stygia are widespread in the arctic and boreal regions of North America [3,23]. Cladonia stellaris is rare along maritime coasts [3] but is reported in New Jersey [117], Michigan's Upper Peninsula [143], and Wisconsin [133]. Cladonia stygia is most common at the northern boreal timberline zone. It occupies habitats from Kodiak Island, Alaska, east to Newfoundland and south to Queen Charlotte Islands, British Columbia, Idaho, northern Minnesota, the Appalachian Mountains, and western North Carolina [5,143].

HABITAT TYPES AND PLANT COMMUNITIES: Perhaps the most commonly mentioned reindeer lichen habitat in North America is the lichen woodland or forest, usually dominated by white spruce (Picea glauca) or black spruce (P. mariana) (e.g., [2,3,6]). However, reindeer lichens occupy a wide variety of habitats. On the Queen Charlotte Islands, C. arbuscula subsp. beringiana occurs in or near bogs or fens and less frequently over rocks in full sun [23]. In northern Ontario, C. arbuscula habitat includes rock outcrops, rocky lakeshores, jack pine (Pinus banksiana) forests, muskegs, and exposed peatlands [2,3]. In southeastern New York, C. arbuscula is found in pitch pine (P. rigida) barrens and plains [37]. In the Pacific Northwest, C. mitis occurs on rock outcrops and talus slopes [85], and is noted in abandoned fields in eastern Minnesota [65]. Cladonia rangiferina occupies tundra habitats in Alaska [11,12,18], but on the Queen Charlotte Islands it occurs in open, dry habitats, usually with patches of moss [23]. In the Pacific Northwest, C. rangiferina occurs in xeric lodgepole pine (P. contorta) forests [53]. In northwestern New Jersey, C. rangiferina was described in rock crevices with litter accumulation [93]. Cladonia stellaris is reported from dry, open habitats in British Columbia [23] and Newfoundland [1]. Cladonia stygia is most frequent in peatlands, particularly treeless bogs [5].		C. rangiferina in oak-pine litter © Charles S. Lewallen
Reindeer lichens are also noted in oak (Quercus spp.) woodlands in southwestern Illinois, western Kentucky, and western Tennessee [52] and in grassland-heathland communities on Nantucket Island, Massachusetts [42].

The vegetation classifications listed below describe plant communities in which reindeer lichens are dominant.

• black spruce/lichen forest subtype with a "well-developed lichen carpet" of primarily Cladonia stellaris, C. mitis, and C. rangiferina, recognized by the Society of American Foresters [34]
• jack pine/lichen forest subtype, characterized by a dense reindeer lichen ground layer, also recognized by the Society of American Foresters [82]

• white spruce-black spruce/willow/mountain cranberry (Salix spp./Vaccinium vitis-idaea)/C. rangiferina forest type [144]
• white spruce/dwarf birch (Betula nana)/reindeer lichen vegetation unit [138]
• white spruce/thinleaf alder/kinnikinnick/snow lichen (Alnus incana subsp. tenuifolia/Arctostaphylos uva-ursi/Stereocaulon spp.)-reindeer lichen woodland [43]
• white spruce/reindeer lichen woodland vegetation type on gently rolling, stabilized sand dunes in Kobuk Valley National Park [102]
• mature black spruce-white spruce/bog birch (B. glandulosa)/lichen forest [46]
• black spruce/willow/splendid feather moss (Hylocomium splendens)/C. rangiferina forest type
• black spruce/prickly rose/horsetail (Rosa acicularis/Equisetum spp.)/C. rangiferina forest community in the Porcupine River and upper Yukon River drainages [144]
• black spruce/sphagnum (Sphagnum spp.)-reindeer lichen type [46,138]
• black spruce/moss-lichen mature forest community [46]
• black spruce/splendid feather moss-reindeer lichen vegetation type [138]
• black spruce/reindeer lichen vegetation type in Kobuk Valley National Park [102]
• black spruce/reindeer lichen vegetation [138]
• black spruce/lichen rangeland cover type, recognized by the Society for Range Management [129]
• bog blueberry-black crowberry (V. uliginosum-Empetrum nigrum)-dwarf birch-Labrador tea (Ledum spp.)-mountain cranberry/cetraria lichen (Cetraria spp.)-reindeer lichen-snow lichen (Stereocaulon paschale) tundra vegetation type in the Bering Land Bridge National Preserve [103]
• lichen tundra rangeland cover type, recognized by the Society for Range Management [130]

• polytrichum moss (Polytrichum spp.)-reindeer lichen association [35]

• Rocky Mountain lodgepole pine/bog Labrador tea (Pinus contorta var. latifolia/L. groenlandicum)/Cladonia mitis community type [127]

• Rocky Mountain lodgepole pine/velvetleaf blueberry (V. myrtilloides)/reindeer lichen forest community type in west-central Alberta [30]

• jack pine-black spruce/bog Labrador tea/reindeer lichen community
• jack pine/velvetleaf blueberry/C. mitis community
• jack pine/C. mitis community in northeastern Alberta and northern Saskatchewan [28]

• black spruce/C. stellaris midseral, upland forest association [73]

• sheep-laurel (Kalmia angustifolia)/C. stellaris-C. rangiferina raised bog community in southeastern Labrador [49]

• black spruce/reindeer lichen vegetation type in northern Manitoba [146]

• black spruce/sheep-laurel/reindeer lichen forest type [87]
• sheep-laurel-blueberry (Vaccinium spp.)/reindeer lichen maritime lichen-heath association
• sheep-laurel/rhacomitrium moss (Rhacomitrium lanuginosum)-reindeer lichen maritime lichen-heath association [1]

• black spruce/mountain cranberry/Schreber's moss (Pleurozium schreberi)-C. mitis community in central Saskatchewan
• jack pine/mountain cranberry/Schreber's moss-C. mitis community in central Saskatchewan [105]

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

• GENERAL BOTANICAL CHARACTERISTICS
• RAUNKIAER LIFE FORM
• REGENERATION PROCESSES
• SITE CHARACTERISTICS
• SUCCESSIONAL STATUS
• SEASONAL DEVELOPMENT

Vegetative regeneration is considered the predominant method of reindeer lichen reproduction. Thallus fragments or microscopic particles with both algal and fungal components can potentially produce a new lichen body. Given the complex reproductive biology and very small size of lichen reproductive structures or fragments, natural regeneration studies and observations are difficult. Webb [141] indicated that reindeer lichen regeneration in burned or logged boreal forests occurred through propagules or surviving fragments that were usually too small to see. Johnson [67] reported that C. mitis and C. rangiferina regeneration in the first years after fire was likely from thallus fragments [67].

Dispersal: Lichen fragments and microscopic fungal and algal particles can be dispersed by wind, water, or animals [55]. In grasslands and forests of northeastern Germany, most dried lichen thallus fragments dispersed less than 8 inches (20 cm) from the parent source. The maximum wind-dispersal distance of C. arbuscula fragments was 14 inches (35 cm) in a dry sandy grassland and 27 inches (68 cm) in an open, early-seral Scots pine (Pinus sylvestris) forest. Lichen cushions disturbed and/or moved by animals dispersed a maximum distance of 32 feet (9.7 m) [60].

Growth: Reindeer lichens are slow growing and long lived. Average growth rates of 4.8 to 11.1 mm/year [41] and average ages of over 100 years [1] have been reported. Growth rates do, however, vary with lichen age, lichen species, site conditions, and herbivory.

Three growth stages exist throughout the reindeer lichen lifespan. The 1st stage, the growth-accumulation period, lasts an average of 10 years but can vary from 6 to 25 years. During this stage, size increases every year, and no part of the podetium dies. In this 1st period of growth, internodes grow 10 to 15 times the height attained in the 1st year of life. During the 2nd stage, the renovation period, podetium height still increases, but internode death occurs at the base. Since decay of the internode is not immediate, height still increases some. This stage may continue for several decades and may exceed 100 years. During the 3rd stage, the withering period, the podetium decays at the base faster than internodes lengthen. This stage of growth lasts approximately 10 to 20 years [1].

Slow reindeer lichen growth has been documented in several locations in the northern hemisphere. On the Seward Peninsula of northwestern Alaska, the average growth rates for C. stellaris, C. rangiferina, and C. arbuscula were 5.0 mm/year, 5.3 mm/year, and 5.4 mm/year, respectively. The growth-accumulation period averaged 11.1 years for C. stellaris, 5.9 years for C. rangiferina, and 10.7 years for C. arbuscula [98]. In forested peatlands of northern Alberta, C. mitis grew an average of 4.8 mm/year [41]. In diverse areas of northern Russia, the average annual height growth was 3.9 mm/year for C. stellaris, 5.9 mm/year for C. rangiferina, and 4.5 mm/year for C. arbuscula. Growth-accumulation periods averaged 12 to 14 years for C. stellaris, 8 to 9 years for C. rangiferina, and 9 to 11 years for C. arbuscula (des Andreev 1939, cited in [1]).

Reindeer lichen growth is greatest on ungrazed, humid, somewhat sheltered sites with relatively long growing seasons. In Newfoundland, the average height of reindeer lichens was greatest in heath stands, followed by shrub-dominated subalpine barrens, and then black spruce forests. Reindeer lichens were shortest in bog stands. Within similar habitat types, ungrazed stands were typically taller than grazed stands (review by Ahti [1]). Nevertheless, reindeer lichen growth following sublethal disturbances may be relatively rapid compared with the growth of newly established individuals. In Sweden, Skunke [123] simulated the effect of reindeer grazing on C. stellaris by clipping to varying heights. Six to 8 years after cutting mats to 0.8 to 1 inch (2-3 cm) heights, C. stellaris recovered to a "completely grazable" condition. When clipped down to the "gelatinous material", C. stellaris was not "grazable" 14 years later [123].

Climate: In general, reindeer lichens occur in cool to cold climates. McCune and Geiser [85] indicated that reindeer lichens occurred on cool, moist sites in the Pacific Northwest. In Canada, reindeer lichens occur in tundra, boreal, cool-temperate, and cool meso-thermal climates [106]. In Quebec, reindeer lichen are subject to short, cool summers and long, cold winters [10]. Cladonia stellaris is an indicator of alpine tundra and boreal climates in coastal British Columbia [72]. Cladonia mitis occurs in arctic to temperate climates with continental tendencies [3].

Species-specific differences exist among reindeer lichens in their relative affinities for maritime and continental climates. In British Columbia, Goward and Ahti [54] found C. arbuscula subsp. beringiana and C. rangiferina "to be indifferent to continentality" and "present in coastal and highly continental inland regions alike". However, Ahti [1] indicated that C. mitis is much less common in maritime-influenced than in continental climates of Newfoundland.

Most literature suggests reindeer lichen abundance is greatest on dry sites, although a review by Ahti and Hepburn [2] indicated that high relative humidity is important to reindeer lichen growth, and McCune and Geiser [85] indicated that reindeer lichens occupied cool, moist sites in the Pacific Northwest. As of this writing (2008), research relating reindeer lichen abundance to precipitation data was lacking.

Moisture: Although reindeer lichens require a relatively humid climate, their ability to absorb water directly from the atmosphere allows them to colonize and even dominate habitats with soils that are too droughty, and often too cold, to support vascular plants [2]. Much of the available literature indicates that reindeer lichen abundance is greatest on dry sites. However, reindeer lichens in Newfoundland can be found on sites ranging from dry to wet and from very poor to moderately rich in fertility [87].

Many studies suggest that reindeer lichens are most common on dry or well-drained sites [2,30,32,82]. In coastal British Columbia [72] and in open-canopy forests in northern Canada [106], reindeer lichens are common on well-drained, water-shedding sites. In central British Columbia's Rocky Mountain lodgepole pine (Pinus contorta var. latifolia) forests, reindeer lichens occurred on mesic and xeric 50- to 100-year-old burned sites. Cover on mesic sites did not change appreciably after 100 years; however, on xeric sites, reindeer lichen cover increased with stand age, and researchers doubted "an equilibrium state had been attained" [26]. The Rocky Mountain lodgepole pine/velvetleaf blueberry/reindeer lichen forest type in west-central Alberta was restricted to sandy, well-drained, coarse-textured soils [30]. In the boreal zone of central and western Ontario, reindeer lichens were more common on shallow, dry soils or rocky outcrops than on poorly drained soils. On the Slate Islands of Lake Superior, reindeer lichens were more than twice as frequent and more than 10 times as abundant in forests with "dry" moisture regimes than those with "medium" moisture regimes [32]. In the Great Lake states, reindeer lichens are considered characteristic groundlayer components only in the driest, least productive jack pine forests [82]. Similarly, in Newfoundland, the black spruce/sheep-laurel/reindeer lichen forest type is characteristic of the driest, least fertile sites [87].

Rarely do reindeer lichens occur in standing water or on saturated substrates. In relatively wet habitats, reindeer lichens typically occupy only the driest microsites, often exploiting subtle topographic variations [2]. Within black spruce muskegs in western Alaska's upper Kuskokwim River region, reindeer lichens occur on hummocks and other dry sites [39]. In British Columbia, reindeer lichens occur occasionally in nutrient-poor wetlands on "topographic prominences subjected to regular desiccation" [72]. A review of indicator plants in Canada states that "in wet areas and peatlands, reindeer lichens occupy micro-topographic prominences subject to regular desiccation, such as the tops of sphagnum hummocks" [106]. In West Virginia heath bogs, C. rangiferina occurs on slightly raised areas of peat subject to periodic drought [35].

Several published accounts suggest there are interspecific differences in reindeer lichen abundance relative to site moisture. Cladonia arbuscula, although frequently found mixed with C. mitis in northern Ontario, is more common in moist habitats [2]. Rowe [109] characterized C. rangiferina as a xerophytic species in the southern boreal forests of Saskatchewan and Manitoba. According to Ahti [1], C. rangiferina may be the most moisture-tolerant of the reindeer lichens in Newfoundland but also occupies the driest sites. In southeastern Labrador, C. rangiferina is more abundant than C. mitis on moist sites [51]. A review suggests that C. stellaris prefers the driest sites, while moisture tolerances of other reindeer lichens are broader [106]. Cladonia stygia has been confused with C. rangiferina and the two are often found together; however, based on observations made in Sweden, C. stygia may be more tolerant of moisture, even enduring seasonal inundation [5].

Although evidence clearly indicates that reindeer lichen colonize and persist on dry sites, presence of dense lichen mats may alter soil surface moisture conditions. Rouse and Kershaw [107] found that soil moisture under lichen mats in mature northern Ontario black spruce forests was at least 40% greater than soil moisture in adjacent 3- and 16-year-old burns with minimal lichen cover, suggesting that lichen ground cover reduced soil moisture evaporation.

Substrate: Since lichens absorb water and mineral nutrients from the air through the outer surface of the thallus, they are not as dependent on soil as vascular plants. Published accounts of substrates bearing reindeer lichens in North America include mineral rock [3,84,85], soil, humus [85,106], duff [57], raised peat [35], organic debris [142], and coarse woody material [3,50,106]. According to Ahti and Hepburn [2], reindeer lichens require at least a thin soil layer for attachment, so they cannot colonize bare rock. Cooper [29] indicated that establishment of reindeer lichen mats on rock surfaces on Isle Royale, northern Michigan, may be facilitated by prior establishment of rhacomitrium moss (Rhacomitrium canescens). Apparently, the moss provides a means of attachment for the lichen mat on rock surfaces with few crevices [29].

The literature reports several instances of possible reindeer lichen affinities for specific substrates. C. stellaris has an affinity for infertile sand or bedrock substrates, while other reindeer lichens have somewhat broader substrate tolerances [106]. A study in east-central Alberta examined the diversity and abundance of lichens on downed woody debris in boreal quaking aspen (Populus tremuloides)-mixed-conifer forests. The frequency of C. mitis was as high as 20% on downed woody debris in the most advanced stage of decay, with nearly 100% humification. Frequency of C. mitis was lower, usually less than 5%, on downed woody debris in less advanced stages of decomposition [33]. In boreal forests of northwestern Ontario, reindeer lichen (C. rangiferina, C. mitis, and C. stellaris) establishment after fire and logging occurred almost exclusively on organic materials such as conifer needles, cone scales, woody debris, dead moss, crustose lichens, overturned ground cover, accumulations of fine organic matter, or shallow organic soils. Cladonia rangiferina occasionally colonized textured rock faces. Reindeer lichens were not observed on mineral soil on sites disturbed 2 to 16 years earlier [141]. On an 8-year-old burned site in northern Minnesota, C. mitis and C. rangiferina "preferred" dead wood [66].

Reindeer lichens often grow in soils and substrates ill-suited for other plants and bryophytes. Reindeer lichens may be found in habitats underlain by permafrost [13,25,46,138,139]. In recently abandoned agricultural fields in eastern Minnesota, C. mitis and C. rangiferina occurred on sandy, low-nutrient soils with delayed successional advancement [65]. Throughout Canadian forests, reindeer lichens are good indicators of acidic, nutrient-poor, coarse-textured, shallow, extremely dry to very dry soils [72,106]. Nevertheless, reindeer lichens in Newfoundland can be found, albeit at reduced abundance, on sites moderately rich in fertility [87].

Elevation: Few elevational ranges for reindeer lichens have been reported. McCune and Geiser [85] indicated that C. rangiferina is found mostly at "low to middle" elevations in the Pacific Northwest, while C. mitis and C. arbuscula are typically limited to "low" elevations. Cladonia stygia occurs between 33 and 490 feet (10-150 m) on British Columbia's Queen Charlotte Islands [23].

Shade tolerance: Reindeer lichens are generally considered shade intolerant, and their importance typically decreases with increasing canopy cover [19,72]. In the Slate Islands of Lake Superior, Ontario, reindeer lichen frequency was more than 10% in "dry regime" forests with less than 45% crown cover, while frequency was 2% or less in forests with greater than 65% crown cover [32]. On black spruce-dominated sites in northern Manitoba, reindeer lichens formed a nearly continuous carpet in openings, in sparsely wooded areas, and/or where trees were small. In denser areas, other lichens and mosses dominated [146]. In late-seral lodgepole pine/reindeer lichen forests of north-central British Columbia, splendid feather moss dominated microsites with a canopy leaf area indices that were significantly greater (P<0.05) than those of reindeer lichen-dominated microsites [128].

Cladonia rangiferina may be the most shade tolerant of the reindeer lichens [19,51]. However, in Rocky Mountain lodgepole pine forests in west-central Alberta, cover of C. rangiferina and C. mitis were not significantly different (P<0.05) between sites with 45.3% to 52.3% canopy cover and those with 72.8 to 87.0% canopy cover [100]. Tegler and Kershaw [131] suggest that heat tolerance and not shade tolerance dictates differences in C. stellaris and C. rangiferina distributions. Cladonia stellaris is more likely to grow in hot, open habitats, while C. rangiferina is more likely to grow in cool, shady habitats. In the pine barrens of south-central Wisconsin, C. mitis occurred in moderately open, warm microsites somewhat protected from xeric conditions by surrounding vegetation, while C. rangiferina occurred in relatively shady, cool microsites with mesic conditions [77].

General: Available literature suggests that reindeer lichens tolerate the later stages of primary succession in some locations and nearly all stages of secondary succession, although some species-specific preferences exist. On Isle Royale in northern Michigan, reindeer lichens, including C. arbuscula, C. rangiferina, and C. stellaris, were dominant in the late stages of rock-surface succession. These species remained, more or less, through subsequent stages of succession on rockshore habitats that were developing from heath communities to jack pine-black spruce forests. In more mesophytic forest sites, reindeer lichens were likely to be replaced by mosses. In xerophytic areas with sparse jack pine, reindeer lichens were likely to remain abundant on rocky surfaces in tree interspaces [29].

Several studies report reindeer lichens in early-seral forests. In quaking aspen woodlands of east-central Alberta, C. mitis is characterized as an early-seral species [33]. In jack pine-black spruce forests in northern Ontario, C. stellaris is typically replaced by splendid feather moss beginning around 40 to 50 years after fire [145]. About 40 to 60 years after stand initiation in Alaskan upland black spruce forests, reindeer lichens invade and may cover up to 20% of the ground layer. In this successional stage, trees are replacing tall shrubs, and there may be as many as 6,000 saplings and mature trees per ha and 12,000 seedlings per ha [137].

Many studies report reindeer lichens in mid- and late-seral communities. A review indicates that in central and southern North American black spruce/lichen forests, a "well-developed lichen carpet" of primarily C. mitis, C. rangiferina, and C. stellaris characterizes the intermediate successional stage [34]. In upland white spruce forests of Alaska's interior, reindeer lichens are generally common in stands over 170 years old [78]. In lowlands of east-central Ontario, Cladonia rangiferina was associated with unlogged and old logged stands [27]. Where site conditions restrict tree establishment and growth, reindeer lichens may persist for extremely long periods without fire. In west-central Alberta, the lodgepole pine/velvetleaf blueberry/reindeer lichen forest type was considered an "edaphic climax" on droughty soils where spruce and fir regeneration were lacking [30]. Payette [94] found dense black spruce/reindeer lichen krummholz sites in northern Quebec that had not burned for 1,500 years or more. Laboratory experiments indicated that reindeer lichen ash may promote nutrient leaching in subarctic spruce-lichen woodlands and thus slow postfire regeneration [40]. For more on reindeer lichen postfire succession, see Fire as a regeneration process and Discussion and Qualification of Lichen Response.

Generally, C. mitis occupies earlier seral habitats than C. rangiferina and C. stellaris. A review reports that mature northern boreal lichen woodlands in Canada "have a ground cover of almost pure C. stellaris". Isolated C. mitis and C. rangiferina podetia occur in the mats, but C. mitis and C. rangiferina are more important in earlier succession, possibly because of the relatively slow growth rate of C. stellaris [69]. In eastern Canada, C. rangiferina is the last species of three (C. stellaris, C. mitis, and C. rangiferina) to drop out as light levels diminish under increasing canopy cover [19]. During succession from bare sand to white spruce/lichen woodlands along the east coast of Hudson Bay, C. mitis established earlier than C. stellaris, which eventually succeeded it in groundlayer dominance [44]. In northern Ontario, C. stellaris is more abundant in later than in earlier stages of lichen woodland succession. Outside of maritime coastal habitats and sites disturbed by fire and caribou grazing, C. stellaris generally succeeds other reindeer lichens and is dominant in "climax" stands [3]. Miller [88] indicated that C. stellaris and C. rangiferina were climax species that appeared 40 or more years after fire in the taiga, but that C. mitis may establish earlier.

Many suggest that reindeer lichens may help maintain forest openings. In different-aged burned sites within northern Quebec's black spruce/lichen woodlands, Morneau and Payette [90] suggested that continuous lichen mats preserved canopy openings by restricting black spruce recruitment. Sedia and Ehrenfeld [117] also found that reindeer lichen mats within severely burned portions of New Jersey's pitch pine forests restricted vascular plant colonization and development, maintaining canopy openings for many decades.

SEASONAL DEVELOPMENT:
As of 2008, published information on the seasonal development of reindeer lichens was apparently lacking.

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

FIRE ECOLOGY OR ADAPTATIONS:
Fire adaptations: Reindeer lichens typically colonize burned sites from thallus fragments dispersed from unburned areas. While colonization by microscopic particles with both algal and fungal components may be possible, it was not reported in the literature and likely is not easily detected. According to Yarranton [145], initial postfire colonization occurs by "long range" dispersal of thallus fragments from populations outside the burned area. Once established, reindeer lichens expand vegetatively. After prescribed fires in Sweden, there was very little reindeer lichen colonization of burned sites by the 5th postfire year, but researchers noted "a few small fragments that dispersed onto some of the burned plots from unburned surroundings" [112].

Many studies have reported reindeer lichen survival in protected sites that were either missed by fire or burned with extremely low severity [28,108,116]. For more on reindeer lichen regeneration and/or survival on burned plots, see Fire Effects.

Fire regimes: Reindeer lichens are an easily ignitable surface fuel that aids in fire spread; however, these surface fuels alone rarely support extreme fire behavior or high-severity fires. Periodic fire maintains canopy openings and the early- to midseral habitats that reindeer lichens prefer. In many habitats, reindeer lichens are eventually replaced, often within 100 years, by mosses, other lichens, and/or vascular plants. However, in some habitats, tree regeneration is restricted by site conditions, so reindeer lichens may persist indefinitely in the absence of fire. Differences in site conditions may partially explain why mean fire-return intervals in reindeer lichen habitats range from 40 to over 500 years.

Fuel characteristics/fire behavior: Reindeer lichens dry readily, becoming a highly flammable fuel during warm, dry conditions [78]. In park-like forests, sun exposure to the "well-aerated (reindeer lichen) tubular body with a high surface-to-volume ratio" makes reindeer lichen potentially the most rapidly desiccated surface fuel in Canadian forests. When submerged in water, reindeer lichen moisture content can increase by 400%, and when drying in shaded conditions, as much as 63% of its moisture can be lost in less than 2 hours (Van Wagner 1969, cited in [96]). In bogs of southeastern Labrador, researchers report that "fire tends to burn preferentially along the lichen-covered ridges and hummocks" [49]. Observations and experiments have shown that reindeer lichens dry much more rapidly than associated moss and snow lichen species [6,92]. In the laboratory, C. rangiferina dried "rapidly" after soaking when compared to associated moss-layer bryophytes from the New Jersey pine barrens [91], and the drying rate of reindeer lichen samples was 1.7 times that of the wetting rate [92]. Based on estimations from measurements made during diurnal wetting and drying, Pech [97] determined that in the absence of cloud cover, C. rangiferina moisture content was reduced to the previous day's minimum by 10:00 am, when overnight dew was light, or by noon, when dew was heavy [97].

Although reindeer lichens burn readily when dry and form a substantial component of the surface fuel bed, fire severity and behavior are strongly dependent on the species and density of the overstory.

Experimental fires in primarily black spruce/lichen forests in the Northwest Territories indicated that potential fire spread rates and severity depend on overstory characteristics. Continuous lichen cover in these forests is flammable but does not, by itself, produce high fireline intensity even during favorable burning conditions. In canopy openings, the spread rate through the lichen mat is generally slow. Lichen bulk density is generally too low to support downward heat sufficient to ignite the entire surface layer before the flaming front passes and too high to support rapid fire spread. Even with wind, "a high-intensity flame-radiation surface fire does not develop". Typical residence times are 30 seconds or less. However, high-intensity fires can be produced when black spruce are ignited. In open black spruce stands, trees typically produce branches that extend to ground level. When black spruce moisture content declines to its typical summer moisture content of 80%, flames from surface fires burn quickly into the crowns under dry, windy conditions. Winds help to spread surface fires and ignite spot fires. Crown fires preheat lichen mats beyond the fire front, increasing the fire spread rate [6].

Jack pine/lichen forests rarely burn in the high-intensity fires described for black spruce stands. Self-pruning in aging jack pines produces crowns well above the surface. Fires in the lichen layer rarely produce surface fire intensities capable of igniting crowns or causing tree mortality [6].

Findings were similar during experimental fires in lodgepole pine stands in British Columbia, some of which contained a substantial reindeer lichen surface fuel component. While reindeer lichens were easily ignited and able to carry fire on the surface, the fire had low intensity and severity. When compared to mosses and lodgepole pine needles, reindeer lichens had the lowest bulk density, lowest surface-to-volume ratio, and lowest heat content. Surface fuels containing a substantial reindeer lichen component carried fire better than those without reindeer lichens, but fires in forests with reindeer lichens were of low severity and exposed very little mineral soil. The researcher concluded that lodgepole pine stands with primarily moss and lichen surface fuels were at low risk of crown fire in the absence of strong winds and/or ladder fuels [76].

Fire as a regeneration process: In the majority of reindeer lichen habitats, periodic fire is necessary to maintain groundlayer dominance in canopy openings. Reindeer lichen persistence in the absence of fire depends on forest and site conditions. Where site conditions restrict tree establishment and growth, reindeer lichens may persist for extremely long periods without fire. Payette [94] found dense black spruce/reindeer lichen krummholz sites in northern Quebec that had not burned for 1,500 years or more.

Eventual replacement of reindeer lichens by mosses and other associated lichens is described in black spruce habitats from the Northwest Territories to Newfoundland and Labrador. On sites sampled in a postfire chronosequence in western Labrador, reindeer lichen cover was greatest, 54%, on 40-year-old sites. Cover was much lower, 11%, in 20-year-old stands. Reindeer lichen cover was 1.3% in 140-year-old stands; reindeer lichen had been replaced successionally by Schreber's moss and knight's plume moss (Ptilium crista-castrensis) [120]. On rich, moist, black spruce forest sites in Newfoundland, reindeer lichens are generally succeeded by mosses and may even be excluded from postfire succession once vascular plants and mosses establish. However, in dry forests near the alpine or maritime treeline, reindeer lichens may be long-lasting after fire [1]. Replacement by mosses is described in 2 reviews: Damman and Johnston [34] suggested that without recurring fire, black spruce/lichen forests eventually become closed-canopy black spruce/feather moss (Hylocomiaceae) forests; Kershaw [69] similarly reported that in the absence of fire, tree density increases and eventually eliminates lichen ground cover. However, both reviews indicated that black spruce/lichen forests burn readily, and rarely are fires so infrequent that closed canopies develop [34,69].

Reindeer lichens dominated the ground flora in black spruce/lichen forests burned between 20 and 60 years ago but declined and were replaced by snow lichen when stands reached about 130 years old in the Abitau-Dunvegan Lakes region of Northwest Territories [81]. In taiga habitats of north-central Canada, fire is considered important to reindeer lichen regeneration. In late stages of succession, reindeer lichens are replaced by vascular plants and feather mosses due, at least in part, to increased amounts of surface litter. Fire removes litter as well as much of the late-successional vegetation; after this clearing process lichens can reestablish and eventually flourish [89]. While the time required for lichen establishment and dominance were not given in these papers, this topic is addressed in other reports (see Discussion and Qualification of Lichen Response).

Insect outbreaks and fire may work in concert to maintain canopy openings and reindeer lichen dominance. In southeastern Quebec, black spruce/reindeer lichen forest patches within a closed-canopy black spruce/feather moss forest were created by the combined effects of fire together with defoliating spruce budworm and European sawfly outbreaks [95]. The insects severely restrict black spruce reproduction, since they preferentially feed on reproductive buds [119]. Fires in infested stands thus cause tree mortality during a time when regeneration potential is low, producing open stands. Recurrent fire in these stands would likely maintain the black spruce/reindeer lichen type, whereas fire exclusion would likely advance development of the black spruce/feather moss forest [95]. Temporal scales were not described in the paper.

In some cases, canopy closure is restricted by factors other than fire; in these cases, reindeer lichens may not be replaced by mosses or vascular plants even after extremely long fire-free intervals. On several sites in northern Canada, reindeer lichen abundance appeared to be increasing after a century without fire. In black spruce/bog blueberry forests near Inuvik in the Northwest Territories, reindeer lichens were only abundant on sites with fire-return intervals substantially longer than 100 years. Cladonia mitis became dominant in stands burned 120 to 200 years ago, and C. stellaris and C. rangiferina were dominant in stands over 200 years old [17]. On burned black spruce/C. stellaris forests in northern Quebec, C. stellaris biomass was about 2,500 kg/ha on 47-year-old burned sites, nearly 6,000 kg/ha on 110-year-old burned sites, and 6,677 kg/ha on 140-year-old burned sites [10]. In burned areas of northern Saskatchewan, northern Manitoba, and the Northwest Territories, the average biomass of "high-value" lichens, including C. stellaris, C. mitis, and C. rangiferina, was nearly twice as great in stands over 120 years old as in stands 76 to 120 years old [114]. In northern Quebec, a dendro-ecological investigation indicated that a black spruce/lichen krummholz type was converted to a tundra/lichen community due to a lack of black spruce regeneration. Before a fire in 1750, the site was occupied by krummholz black spruce that regenerated primarily through layering. Postfire seedling establishment was severely limited. The subsequent development of a thick lichen mat may have further inhibited black spruce establishment [8].

Fire frequency/ecology: The few studies reporting average fire-return intervals in reindeer lichen habitats indicate a large range, about 40 to over 500 years. In jack pine/reindeer lichen forests in the Athabasca Plains region in northern Saskatchewan/northeastern Alberta, the mean fire-return interval averaged 40.4 years for 6 study sites. Fire scars spanned dates from 1783 to 1972. Average fire-return intervals calculated for individual sites ranged from 28 to 54 years, suggesting somewhat patchy burn patterns [28]. Mean fire-return intervals were similar in jack pine and black spruce forests near the northern boreal forest-tundra boundary in northern Quebec. Ground layers were dominated by either C. mitis or C. stellaris at the 18 study sites. Eight fires occurred between 1773 and 1988, including both crown fires and surface fires, although crown fires were considered more common. Mean fire-free intervals ranged from 41 to 85 years for individual sites; the overall mean fire-free interval was 54.8 years. On average, 1.4% of the study area burned each year, indicating a fire-rotation period of 70 years. After 1851, three fires accounted for more than 60% of the burned area, while the other fires were much smaller and burned less than 10% of the area. At the time of the study (1988), stand ages ranged from 35 to 150 years, and 67-year-old stands were most common [36]. Fire-return intervals may exceed 500 years in reindeer lichen habitats in coastal Labrador ([51] and sources therein).

The following table provides fire regime information that may be relevant to reindeer lichens. Included are only those vegetation communities for which published information confirms, or strongly implies, reindeer lichen occurrence. Because site tolerances and distributions for reindeer lichens are incomplete and many reindeer lichen habitats do not burn, such as rocky outcrops and talus slopes, other reindeer lichen habitats may also be missing from the fire regime table. Find further fire regime information for the plant communities in which these species may occur by entering the species' names in the FEIS home page under "Find Fire Regimes".

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON LICHEN
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• LICHEN RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF LICHEN RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

Studies suggest reindeer lichen survival on burned sites is most likely in moist, protected refugia and/or during a quickly burning fire. Rouse [108] reported that "the lichen mat is always consumed when fire reaches it, but if the fire is fast-moving, patches are often missed." In burned black spruce stands in northern Saskatchewan, some C. stellaris and C. rangiferina survival occurred in "small relic areas that had escaped fire" [116]. In jack pine/reindeer lichen woodlands of northeastern Alberta and northern Saskatchewan, surface and surface/crown fires generally resulted in lichen mortality but some survival occurred near the fire margin and in moist depressions [28]. The moisture-retaining properties of reindeer lichen mats may increase reindeer lichen survival potential. In northwestern Alaska, reindeer lichens commonly grow in or on a moss layer, which retains some moisture even under dry conditions. Reindeer lichen pieces dislodged by caribou trampling are more susceptible to drying and may be more susceptible to fire-caused mortality than undisturbed mats, which retain some moisture at the base [99].

Very low-severity fire may kill only the top portion of the lichen mat, allowing for some reindeer lichen survival. In a Scots pine forest in Sweden, burning treatments were evaluated on a continuous reindeer lichen layer. In the lowest-severity treatment, fire spread over the plot, but smoldering combustion was immediately extinguished with a thick blanket. Although appearing dead, by the end of the 5-year observation period new basal growth was observed in C. rangiferina in the lowest-severity treatment. Additional treatments that allowed smoldering for 5 minutes, 15 minutes, and 1 hour completely consumed the reindeer lichen mat, and no subsequent reindeer lichen growth was observed [112]. While experimental treatments may or may not represent realistic conditions, it is possible some reindeer lichens survive after extremely low-severity fires. However, 1 to 11 years after "low intensity" surface fires in Scots pine stands in Lithuania, C. rangiferina and C. arbuscula were present only in adjacent unburned plots [83].

It is often difficult to ascertain from fire studies whether or not postfire reindeer lichen recovery is the result of off-site dispersal or expansion of small surviving pieces in the burned area. Given the potential small size of regenerating or surviving pieces and slow increase in size and abundance of reindeer lichens, they may be overlooked in early postfire communities [67]. Miller [88] indicated that during initial regeneration after removal of all vegetation, including reindeer lichens, primary thalli were inconspicuous and scattered. Vascular plant sampling techniques may not be sufficient to detect early postfire reindeer lichen regeneration.

Many other fire studies indicate that reindeer lichens may not appear on burned sites for a decade or more. A European review reported that while small reindeer lichen thalli may occur 10 years after a fire, between 25 and 40 years were required before reindeer lichens were fully developed [78]. In burned black spruce forests of northern Saskatchewan, C. stellaris and C. rangiferina were rare in areas burned less than 30 years earlier [116]. When a chronosequence of 10 burned black spruce/C.stellaris stands was visited in northern Quebec, a 47-year-old stand was the youngest on which C. stellaris was recorded [10]. In forested peatlands of northern Alberta, where C. mitis was the dominant lichen, sites burned within 20 years of sampling had 0% to 20% lichen cover. Average lichen cover in 25- to 65-year-old burned sites was not significantly different (P>0.05) from that on mature sites (over 70 years old). Despite substantial variation in lichen cover among sites of all ages, postfire recolonization on peatlands was considered rapid compared to that on other sites [41].

Studies have documented substantial variation, ranging from decades to centuries, in the time required to reach prefire abundance, peak levels of abundance, or surface layer dominance following fire. In a postfire black spruce/lichen chronosequence in the Abitau-Dunvegan Lakes region of the Northwest Territories, reindeer lichens dominated ground flora between roughly 20 and 60 years after fire, while the ground layer was dominated by snow lichen in 61- to 130-year-old stands and by mosses in stands older than 130 years [81]. In another black spruce/lichen chronosequence that spanned 3 to 70 postfire years in the southeastern Northwest Territories, Cladonia stellaris was present (0.1% cover) in 12-year-old stands but had greatest cover (46-47%) in 49- and 70-year-old stands [70]. Maximum lichen cover where C. mitis dominated the ground layer occurred about 45 years after fire in jack pine/reindeer lichen woodlands of the Athabasca Plains region in northern Saskatchewan/northeastern Alberta [28]. In eastern Newfoundland, reindeer lichen cover was much greater on 38-year-old than 23-year-old burned black spruce-sheep-laurel stands. Reindeer lichen cover was low in early postfire succession, averaging 5% in 23-year-old stands, but increased to 79% in 38-year-old stands [20]. In subarctic black spruce forests in western Labrador, reindeer lichen cover was greatest, 54%, in 40-year-old burned stands when compared to 2-year-old, 18-year-old, 80-year-old, and 140-year old burned stands [120].

Studies indicate that fire severity, moisture conditions, grazing intensity, and vegetation type can affect postfire recovery of reindeer lichens. Investigations after a prescribed fire in young jack pine stands in northern Ontario indicated that C. rangiferina recovery may be affected by fire severity. Although initially "eliminated" by the fire, plots with less than 1.2 inches (3 cm) of duff removal had some C. rangiferina regeneration 6 to 10 years after fire. Plots with more than 1.2 inches (3 cm) of duff removed had no C. rangiferina in the same time frame [79]. In Rocky Mountain lodgepole pine forests in central British Columbia, reindeer lichens occurred on mesic and xeric 50- to 100-year-old burned sites. Cover on mesic sites did not change much after 100 years; however, on xeric sites, reindeer lichen cover progressively increased with stand age, and researchers doubted that "an equilibrium state had been attained" [26]. After years of studying caribou in the Kaminuriak Lake region of Nunavut, Miller [88] suggested that fire severity and intensity of caribou grazing may affect reindeer lichen recovery on burned sites more than time since fire.

Only 1 study to date (2008) compares reindeer lichen responses to fire from different vegetation types: This research, conducted in New Brunswick, suggests vegetation type may affect reindeer lichen recovery patterns and timing. Cladonia rangiferina was more abundant and occupied more ground cover earlier in burned jack pine than in mixed-hardwood stands. Average cover was 18% in 37-year-old jack pine stands and less than 0.5% in 37-year-old mixed-hardwood stands [80].

Species-specific responses: Within a vegetation type, species-specific differences in recovery patterns after fire are evident. In studies that compared reindeer lichen species recovery or establishment on burned sites, C. mitis was typically found earliest on burned sites. Generally, C. rangiferina occupied sites soon after C. mitis colonization. The pattern and timing of the appearance of C. stellaris and C. arbuscula on burned sites were less clear. Uggla [136] considered postfire recovery faster by C. rangiferina and C. arbuscula than by C. stellaris.

Treeless bogs: In a study of 4 bog hummocks in southeastern Labrador burned 6 to 83 years earlier, C. mitis and C. rangiferina were first recorded in a 22-year-old burn, and C. stellaris was first recorded in a 50-year-old burn [49].

Lodgepole pine: In burned lodgepole pine stands in north-central British Columbia, C. mitis and C. rangiferina occurred in 0- to-50-year-old stands, becoming groundlayer dominants in 50- to-150-year-old stands. Cladonia stellaris was present, but less abundant, in young and medium aged stands. In the 100- to 150-year-old and over 150-year old stands, C. arbuscula occurred with low abundance. The importance of C. mitis and C. rangiferina decreased as stand age exceeded 150 years [31].

Spruce: Cladonia mitis was often the first reindeer lichen to colonize and eventually dominate burned spruce forests. In black spruce/bog blueberry forests near Inuvik, Northwest Territories, C. mitis became dominant around 120 to 200 years after fire. C. stellaris and C. rangiferina did not dominate until 200 or more years after fire [17]. In northern Saskatchewan black spruce forests, C. stellaris and C. rangiferina may reach prefire abundance within 90 to 120 years of fire [116].

In burned spruce/lichen stands in northern Quebec, C. mitis occurred on 4-year-old burned sites. Cladonia rangiferina and C. stellaris were not detected until 14 years after fire. A continuous C. mitis lichen carpet occurred on a 38-year-old burned site, but cover was greatest on the 65-year-old burned site and declined in older stands. Cladonia stellaris cover was relatively low in early postfire stands but reached 70% to 80% on 130- and 250-year-old burned sites [90].

A postfire chronosequence in tundra, boreal forest, and ecotone habitats in northern Quebec showed that Cladonia mitis dominated in the first 50 years after fire but was replaced by C. stellaris on sites unburned for 90 years or more [9].

Cladonia mitis, C. rangiferina, and C. arbuscula appeared earlier in postfire succession than did C. stellaris in subalpine black spruce and open black spruce-tamarack (Larix laricina) sites in interior Newfoundland. Cladonia arbuscula and C. rangiferina occurred with high frequency but low cover by the 10th postfire year. Cover was still less than 20% on sites burned 35 to 36 years earlier. Cladonia stellaris occurred on 14-year-old burned sites, but increases in cover and frequency were negligible as time since fire increased to 36 years. On another site, cover of C. mitis, C. rangiferina, and C. stellaris in a 35-year-old burned stand was about half that in an adjacent unburned site. Cladonia mitis and C. rangiferina cover increased rapidly between 30 and 60 years after fire. At about 80 years after fire, C. stellaris began to dominate. Caribou grazing may delay or inhibit the development of this later successional stage of C. stellaris dominance [16].

In late-seral black spruce forests in southeastern Labrador, low site productivity may allow C. stellaris to persist indefinitely. On 10- to 12-year-old burned sites, C. mitis and C. rangiferina may occur. Of these early postfire species, C. mitis expands more rapidly, reaching maximum cover 40 to 60 years after fire, and C. rangiferina reaches a lower peak cover 75 to 85 years after fire. Cladonia stellaris may establish on 35-year-old stands, but maximum cover is not reached until 80 to 90 years after stand establishment. On productive sites where closed canopies develop, C. stellaris abundance declines rapidly in shady, mesic conditions that provide prime Schreber's moss habitat. On drier, less productive sites, C. stellaris may persist as the dominant ground cover until the next fire [51].

Jack pine-spruce: Cladonia mitis and C. stellaris were found on burned jack pine-spruce before C. rangiferina. In a chronosequence study in the southeastern Northwest Territories and northern Saskatchewan, C. mitis and C. stellaris biomass and cover peaked in 80 to 100-year-old stands. Cladonia rangiferina cover never equaled that of C. mitis and C. stellaris [132].

In central Quebec's black spruce and jack pine boreal forests, C. mitis and C. rangiferina were present with greater abundance and sooner after fire than C. stellaris. Ground layers in stands about 50 years old were dominated by Cladonia mitis and C. rangiferina, while C. stellaris dominated the ground layer in 71- and 110-year-old stands [48].

In jack pine-black spruce forests of northern Quebec, C. mitis was the ground layer dominant in postfire stands less than 67 years old, while in stands over 132 years old, C. stellaris dominated [36]. Severity of a crown fire affected reindeer lichen abundance on 57-year-old jack pine-black spruce forests in northwestern Quebec through its effect on seed sources and tree regeneration. Sites were sampled within areas of closed-canopy regeneration and in more open forests. Surface fire severity was considered comparatively uniform, as humus thickness was similar in open- and closed-canopy areas. Differences in reindeer lichen cover were likely most affected by canopy regeneration. Cladonia mitis was significantly more abundant in open- than closed-canopy plots, while C. rangiferina and C. stellaris were significantly more abundant in closed-canopy than open plots (P<0.001) [7]. Citing work by Lechowicz and Adams [77], Arseneault [7] suggested that differences in abundance may have been due to Cladonia mitis's greater tolerance of high evaporation rates in full sunlight.

• The effects of experimental fires in an Alaskan black spruce/feather moss community
• Forest floor and plant responses to experimental fires in an Alaskan black spruce/feather moss community

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

Large numbers of caribou, causing heavy grazing and trampling, can impact reindeer lichen habitats by exposing mineral soil, altering successional trajectories, and potentially setting succession back many decades [21]. Cladonia stellaris does not withstand heavy grazing and trampling [1]. See Successional Status for further information.

Small mammals: In a review, Sharnoff reported that reindeer lichens are eaten by southern red-backed voles [118]. It is possible that other small mammals feed on reindeer lichens.

Palatability/nutritional value: While reports suggest that reindeer lichens are nutritionally inferior to arboreal lichens and vascular plants, they are still preferred by caribou over other forage, at least in winter. A review by Ahti and Hepburn [2] suggests that reindeer lichens alone probably cannot sustain caribou, although there was no time frame provided for this assertion. A study of the chemical composition of caribou forage plants near Inuvik, Northwest Territories, showed that the protein content of reindeer lichens was significantly lower than that of vascular plants [115]. It is possible that energy conservation during winter foraging may compensate for the lower nutritional content.

OTHER MANAGEMENT CONSIDERATIONS:
Pollution: Reindeer lichens are sensitive to pollution and have been used as pollution load indicators [134]. Cladonia mitis, C. rangiferina, and C. arbuscula are rated sensitive to heavy metals and acid rain in the USDA Forest Service's Pacific Northwest region [134]. Reindeer lichen tissues contained heavy metals and sulfur oxides as far as 15 miles (24 km) downwind from pulp and paper mills in International Falls, Minnesota. The concentration of pollutants, however, was lower in terrestrial reindeer lichens than in arboreal lichen species [15].

Mountain pine beetle: Preliminary results from a study in west-central British Columbia suggest that lodgepole pine death due to a mountain pine beetle epidemic may negatively impact terrestrial reindeer lichens. Decreased abundance of reindeer lichens corresponded to increases in kinnikinnick and, to a lesser extent, Schreber's moss. Researchers speculated that increased abundance of these species in areas of reduced lodgepole pine canopy cover may be due to increased light and nutrient availability and/or increased forest floor and soil moisture. Although supporting evidence is needed, there is some concern that more mountain pine beetle outbreaks, possibly exacerbated by climate change, may have deleterious impacts on forage lichens utilized by caribou populations in the region [110].

Tree regeneration: Some evidence suggests that reindeer lichens may inhibit tree seedling establishment but facilitate growth of those seedlings that do establish. Houle and Filion [63] found that lichen removal in white spruce/C. stellaris forests in northwestern Quebec resulted in greater white spruce seedling establishment but reduced seedling growth, particularly in juveniles less than 8 inches (20 cm) tall. They also observed that nearly all seedlings emerged from the edges of the lichen-removal quadrats or close to the lichen mat. Based on these results and a review of the literature, researchers speculated that reindeer lichens inhibited white spruce seedling establishment through allelopathy, physical impediment, altered soil temperature, and/or light capture. They suggested that small openings in the lichen mat may provide favorable recruitment microsites through increased seed retention, protection from predators, and/or increased soil moisture. Growth of juveniles may be increased by the water-holding capacity or nutrient leaching of the lichen mat [63]. In contrast to these results, jack pine and white spruce seedlings watered for 17 weeks through mulches of Cladonia rangiferina or C. stellaris had significantly reduced growth compared with controls mulched with peat moss (P<0.05). The researcher suggested that reindeer lichens may interfere with jack pine and white spruce seedling development, although the mechanism was not determined [45].

Vascular plant development: In a long-abandoned (≥50 years) agricultural field in west-central Wisconsin, brittle prickly-pear (Opuntia fragilis) flowered more when cooccurring with C. mitis and C. rangiferina. The reindeer lichen presence lowered summer surface soil temperatures by up to 7 °F (4 ºC), which may have improved soil moisture conditions [14].

REFERENCES:

1. Ahti, T. 1959. Studies on the caribou lichen stands of Newfoundland. Annals of the Botanical Society. Vanamo. 30(4): 1-44. [18901]

2. Ahti, T.; Hepburn, T. L. 1967. Preliminary studies on woodland caribou range, especially on lichen stands, in Ontario. Res. Rep. (Wildlife) No. 74. Toronto, ON: Ontario Department of Lands and Forests, Research Branch. 134 p. [13294]

3. Ahti, Teuvo. 1964. Macrolichens and their zonal distribution in boreal and arctic Ontario, Canada. Annales Botanici Fennici. 1: 1-35. [66197]

4. Ahti, Teuvo; DePriest, Paula T. 2001. New combinations of Cladina epithets in Cladonia (Ascomycotina: Cladoniaceae). Mycotaxon. 78: 499-502. [66201]

5. Ahti, Teuvo; Hyvonen, Soili. 1985. Cladina stygia, a common, overlooked species of reindeer lichen. Annales Botanici Fennici. 22: 223-229. [66202]

6. Alexander, M. E.; Stocks, B. J.; Lawson, B. D. 1991. Fire behavior in black spruce-lichen woodland: the Porter Lake project. NOR-X-310. Edmonton, AB: Forestry Canada, Northwest Region, Northern Forestry Centre. 44 p. [18823]

7. Arseneault, Dominique. 2001. Impact of fire behavior on postfire forest development in a homogeneous boreal landscape. Canadian Journal of Forest Research. 31(8): 1367-1374. [66204]

8. Arseneault, Dominique; Payette, Serge. 1992. A postfire shift from lichen-spruce to lichen-tundra vegetation at tree line. Ecology. 73(3): 1067-1081. [18741]

9. Arseneault, Dominique; Villeneuve, Normand; Boismenu, Claire; LeBlanc, Yves; Deshaye, Jean. 1997. Estimating lichen biomass and caribou grazing on the wintering grounds of northern Quebec: an application of fire history and Landsat data. Journal of Applied Ecology. 34: 65-78. [28449]

10. Auclair, Allan N. D. 1985. Postfire regeneration of plant and soil organic pools in a Picea mariana-Cladonia stellaris ecosystem. Canadian Journal of Forest Research. 15(1): 279-291. [66004]

11. Barker, Marilyn H. 1994. SRM 916: Sedge-shrub tundra. In: Shiflet, Thomas N., ed. Rangeland cover types of the United States. Denver, CO: Society for Range Management: 136-137. [67506]

12. Barker, Marilyn H. 1994. SRM 918: Tussock tundra. In: Shiflet, Thomas N., ed. Rangeland cover types of the United States. Denver, CO: Society for Range Management: 138. [67508]

13. Beilman, David W. 2001. Plant community and diversity change due to localized permafrost dynamics in bogs of western Canada. Canadian Journal of Botany. 79: 983-993. [40118]

14. Bennett, James P.; Bomar, Charles R.; Harrington, Cynthia A. 2003. Lichens promote flowering of Opuntia fragilis in west-central Wisconsin. The American Midland Naturalist. 150: 221-230. [66205]

15. Bennett, James P.; Wetmore, Clifford M. 1997. Chemical element concentrations in four lichens on a transect entering Voyageurs National Park. Environmental and Experimental Botany. 37: 173-185. [66207]

16. Bergerud, Arthur T. 1971. Abundance of forage on the winter range of Newfoundland caribou. The Canadian Field-Naturalist. 85: 39-52. [14759]

17. Black, R. A.; Bliss, L. C. 1978. Recovery sequence of Picea mariana - Vaccinium uliginosum forests after burning near Inuvik, Northwest Territories, Canada. Canadian Journal of Botany. 56(17): 2020-2030. [7448]

18. Bliss, L. C. 1988. Arctic tundra and polar desert biome. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. New York: Cambridge University Press: 1-32. [13877]

19. Bloom, Robin G.; Mallik, Azim U. 2004. Indirect effects of black spruce (Picea mariana) cover on community structure and function in sheep laurel (Kalmia angustifolia) dominated heath of eastern Canada. Plant and Soil. 265(1-2): 279-293. [52344]

20. Bloom, Robin G.; Mallik, Azim U. 2006. Relationships between ericaceous vegetation and soil nutrient status in a post-fire Kalmia angustifolia-black spruce chronosequence. Plant and Soil. 289(1/2): 211-226. [66677]

21. Boudreau, Stephane; Payette, Serge. 2004. Caribou-induced changes in species dominance of lichen woodlands: an analysis of plant remains. American Journal of Botany. 91(3): 422-429. [56150]

22. Brodo, Irwin M. 1976. A new combination for Cladonia stellaris. The Bryologist. 79(3): 363-364. [66210]

23. Brodo, Irwin M.; Ahti, Teuvo. 1996. Lichens and lichenicolous fungi of the Queen Charlotte Islands, British Columbia, Canada. 2. The Cladoniaceae. Canadian Journal of Botany. 74(7): 1147-1180. [66211]

24. Brodo, Irwin M.; Sharnoff, Sylvia Duran; Sharnoff, Stephen. 2001. Lichens of North America. New Haven, CT: Yale University Press. 795 p. [90750]

25. Brown, R. J. E. 1983. Effects of fire on the permafrost ground thermal regime. In: Wein, Ross W.; MacLean, David A., eds. The role of fire in northern circumpolar ecosystems. Scope 18. New York: John Wiley & Sons: 97-110. [18506]

26. Brulisauer, Alfred R.; Bradfield, Gary E.; Maze, Jack. 1996. Quantifying organizational change after fire in lodgepole pine forest understorey. Canadian Journal of Botany. 74(11): 1773-1782. [27523]

27. Brumelis, G.; Carleton, T. J. 1989. The vegetation of post-logged black spruce lowlands in central Canada. II. Understory vegetation. Journal of Applied Ecology. 26(1): 321-339. [7864]

28. Carroll, S. B.; Bliss, L. C. 1982. Jack pine - lichen woodland on sandy soils in northern Saskatchewan and northeastern Alberta. Canadian Journal of Botany. 60(11): 2270-2282. [7283]

29. Cooper, William S. 1913. The climax forest of Isle Royale, Lake Superior, and its development. II. Botanical Gazette. 55(2): 115-140. [11538]

30. Corns, I. G. W. 1983. Forest community types of west-central Alberta in relation to selected environmental factors. Canadian Journal of Forest Research. 13(5): 995-1010. [691]

31. Coxson, Darwyn S.; Marsh, Janet. 2001. Lichen chronosequences (postfire and postharvest) in lodgepole pine (Pinus contorta) forests of northern interior British Columbia. Canadian Journal of Botany. 79: 1449-1464. [40594]

32. Cringan, Alexander Thom. 1957. History, food habits and range requirements of the woodland caribou of continental North America. In: Transactions, 22nd North American wildlife conference; 1957 March 4-6; Washington, DC. Washington, DC: Wildlife Management Institute: 485-501. [15651]

33. Crites, Susan; Dale, Mark R. T. 1998. Diversity and abundance of bryophytes, lichens, and fungi in relation to woody substrate and successional stage in aspen mixedwood boreal forests. Canadian Journal of Botany. 76: 641-651. [29073]

34. Damman, A. W. H.; Johnston, William F. 1980. Black spruce. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 11-14. [49853]

35. Darlington, H. Clayton. 1943. Vegetation and substrate of Cranberry Glades, West Virginia. Botanical Gazette. 104(3): 371-393. [64491]

36. Desponts, Mireille; Payette, Serge. 1992. Recent dynamics of jack pine at its northern distribution limit in northern Quebec. Canadian Journal of Botany. 70(6): 1157-1167. [66219]

37. Dirig, Robert. 1994. Lichens of pine barrens, dwarf pine plains, and "ice cave" habitats in the Shawangunk Mountains, New York. Mycotaxon. 52(2): 523-558. [66220]

38. Douglas, George W.; Bliss, L. C. 1977. Alpine and high subalpine plant communities of the North Cascades Range, Washington and British Columbia. Ecological Monographs. 47: 113-150. [9487]

39. Drury, William H., Jr. 1956. Bog flats and physiographic processes in the upper Kuskokwim River region, Alaska. Contributions from the Gray Herbarium No. 178. Cambridge, MA: Harvard University, The Gray Herbarium. 127 p. [12996]

40. Dubreuil, M. A.; Moore, T. R. 1982. A laboratory study of postfire nutrient redistribution in subarctic spruce-lichen woodlands. Canadian Journal of Botany. 60: 2511-2517. [7887]

41. Dunford, Jesse S.; McLoughlin, Philip D.; Dalerum, Fredrik; Boutin, Stan. 2006. Lichen abundance in the peatlands of northern Alberta: implications for boreal caribou. Ecoscience. 13(4): 469-474. [67385]

42. Dunwiddie, Peter W. 1997. Long-term effects of sheep grazing on coastal sandplain vegetation. Natural Areas Journal. 17(3): 261-264. [27443]

43. Dyrness, C. T.; Viereck, L. A.; Foote, M. J.; Zasada, J. C. 1988. The effect on vegetation and soil temperature of logging flood-plain white spruce. Res. Pap. PNW-RP-392. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 45 p. [7471]

44. Filion, Louise; Payette, Serge. 1989. Subarctic lichen polygons and soil development along a colonization gradient on eolian sands. Arctic and Alpine Research. 21(2): 175-184. [66221]

45. Fisher, R. F. 1979. Possible allelopathic effects of reindeer-moss (Cladonia) on Jack pine and white spruce. Forest Science. 25: 256-260. [66222]

46. Foote, M. Joan. 1983. Classification, description, and dynamics of plant communities after fire in the taiga of interior Alaska. Res. Pap. PNW-307. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 108 p. [7080]

47. Forman, Richard T. T. 1998. Common bryophytes and lichens of the New Jersey Pine Barrens. In: Forman, Richard T. T., ed. Pine Barrens: ecosystem and landscape. New Brunswick, NJ: Rutgers University Press: 407-424. [50794]

48. Fortin, Marie-Josee; Payette, Serge; Marineau, Kim. 1999. Spatial vegetation diversity index along a postfire successional gradient in the northern boreal forest. Ecoscience. 6(2): 204-213. [36002]

49. Foster, D. R.; Glaser, P. H. 1986. The raised bogs of south-eastern Labrador, Canada: classification, distribution, vegetation and recent dynamics. Journal of Ecology. 74(1): 47-71. [70176]

50. Foster, David R. 1984. Phytosociological description of the forest vegetation of southeastern Labrador. Canadian Journal of Botany. 62(5): 899-906. [15356]

51. Foster, David R. 1985. Vegetation development following fire in Picea mariana (black spruce)-Pleurozium forests of south-eastern Labrador, Canada. Journal of Ecology. 73(2): 517-534. [7222]

52. Fralish, James S.; Franklin, Scott B.; Close, David D. 1999. Open woodland communities of southern Illinois, western Kentucky, and middle Tennessee. In: Anderson, Roger; Fralish, James S.; Baskin, Jerry M., eds. Savannas, barrens, and rock outcrop plant communities of North America. Boston, MA: Cambridge University Press: 171-189. [51448]

53. Franklin, Jerry F.; Dyrness, C. T. 1973. Natural vegetation of Oregon and Washington. Gen. Tech. Rep. PNW-8. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 417 p. [961]

54. Goward, Trevor; Ahti, Teuvo. 1997. Notes on the distributional ecology of the Cladoniaceae (lichenized Ascomycetes) in temperate and boreal western North America. Journal of the Hattori Botanical Laboratory. 82: 143-155. [66225]

55. Hale, Mason E., Jr. 1961. Lichen handbook: A guide to the lichens of eastern North America. Washington, DC: Smithsonian Institution Press. 178 p. [9926]

56. Hale, Mason E., Jr.; Culberson, William Louis. 1970. A fourth checklist of the lichens of the continental United States and Canada. Bryologist. 73(3): 499-543. [19940]

57. Hammer, Samuel. 1994. Cladoniaceae americanae exsiccatae. Mycotaxon. 52(2): 475-493. [68879]

58. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2005. Interagency fire regime condition class guidebook. Version 1.2, [Online]. In: Interagency fire regime condition class website. U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy; Systems for Environmental Management (Producer). Variously paginated [+ appendices]. Available: http://www.frcc.gov/docs/1.2.2.2/Complete_Guidebook_V1.2.pdf [2007, May 23]. [66734]

59. Heatwole, Harold. 1966. Moisture exchange between the atmosphere and some lichens of the genus Cladonia. Mycologia. 58: 148-156. [19524]

60. Heinken, Thilo. 1999. Dispersal patterns of terricolous lichens by thallus fragments. Lichenologist. 31(6): 603-612. [66226]

61. Holt, Emily A.; Severns, Paul M. 2005. The effects of prescribed burning on wet prairie lichen communities. Natural Areas Journal. 25(2): 130-136. [54650]

62. Host, George E.; Pregitzer, Kurt S. 1992. Geomorphic influences on ground-flora and overstory composition in upland forests of northwestern lower Michigan. Canadian Journal of Forest Research. 22: 1547-1555. [19671]

63. Houle, Gillies, Houle; Filion, Louise. 2003. The effects of lichens on white spruce seedling establishment and juvenile growth in a spruce-lichen woodland of subarctic Quebec. Ecoscience. 10(1): 80-84. [66229]

64. Index Fungorum. 2013. ISF-Index Fungorum [Online]. Surrey, UK: Royal Botanic Gardens Kew, Mycology Section; Aukland, New Zealand: Landcare Research-NZ, Mycology Group; Beijing, China: Institute of Microbiology, Chinese Academy of Science (Custodians). Available: http://www.indexfungorum.org [2016, March 14]. [29113]

65. Johansson, P.; Reich, Peter B. 2005. Population size and fire intensity determine post-fire abundance in grassland lichens. Applied Vegetation Science. 8(2): 193-198. [61616]

66. Johansson, Per; Wetmore, Clifford M.; Carlson, Daren J.; Reich, Peter B.; Thor, Goran. 2006. Habitat preference, growth form, vegetative dispersal and population size of lichens along a wildfire severity gradient. The Bryologist. 109(4): 527-541. [66443]

67. Johnson, E. A. 1981. Vegetation organization and dynamics of lichen woodland communities in the Northwest Territories, Canada. Ecology. 62(1): 200-215. [19244]

68. Kari, Priscilla Russell. 1987. Tanaina plantlore. Dena'ina K'et'una: An ethnobotany of the Dena'ina Indians of southcentral Alaska. 2nd ed. [Revised]. Anchorage, AK: U.S. Department of the Interior, National Park Service, Alaska Region. 205 p. [67343]

69. Kershaw, K. A. 1977. Studies on lichen-dominated systems. XX. An examination of the northern boreal lichen woodlands in Canada. Canadian Journal of Botany. 55(4): 393-410. [69098]

70. Kershaw, K. A.; Rouse, W. R.; Bunting, B. T. 1975. The impact of fire on forest and tundra ecosystems. ALUR Report 74-75-63. Ottawa, ON: Department of Indian Affairs and Northern Affairs, Arctic Land Use Research Program. 81 p. [69581]

71. Klein, David R. 1982. Fire, lichens, and caribou. Journal of Range Management. 35(3): 390-395. [10898]

72. Klinka, K.; Krajina, V. J.; Ceska, A.; Scagel, A. M. 1989. Indicator plants of coastal British Columbia. Vancouver, BC: University of British Columbia Press. 288 p. [10703]

73. Klinka, K.; Krestov, P. V.; Chourmouzis, C. 2002. Classification and ecology of the mid-seral Picea mariana forests of British Columbia. Applied Vegetation Science. 5(2): 227-235. [47096]

74. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1). Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior. 72 p. On file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. [66741]

75. LANDFIRE Rapid Assessment. 2007. Rapid assessment potential natural vegetation groups (PNVGs): associated vegetation descriptions and geographic distributions. Washington, DC: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab; U.S. Geological Survey; Arlington, VA: The Nature Conservancy. 84 p. [66533]

76. Lawson, Bruce D. 1972. Fire spread in lodgepole pine stands. Missoula, MT: University of Montana. 119 p. Thesis. [6920]

77. Lechowicz, Martin J.; Adams, Michael S. 1974. Ecology of Cladonia lichens. II. Comparative physiological ecology of C. mitis, C. rangiferina, and C. uncialis. Canadian Journal of Botany. 52: 411-22. [66243]

78. Lutz, H. J. 1956. Ecological effects of forest fires in the interior of Alaska. Tech. Bull. No. 1133. Washington, DC: U.S. Department of Agriculture, Forest Service. 121 p. [7653]

79. Lynham, T. J.; Wickware, G. M.; Mason, J. A. 1998. Soil chemical changes and plant succession following experimental burning in immature jack pine. Canadian Journal of Soil Science. 78(1): 93-104. [29822]

80. MacLean, David A.; Wein, Ross W. 1977. Changes in understory vegetation with increasing stand age in New Brunswick forests: species composition, cover, biomass, and nutrients. Canadian Journal of Botany. 55: 2818-2831. [10106]

81. Maikawa, E.; Kershaw, K. A. 1976. Studies on lichen-dominated systems. XIX. The postfire recovery sequence of black spruce-lichen woodland in the Abitau Lake region, N.W.T. Canadian Journal of Botany. 54(23): 2679-2687. [7225]

82. Majcen, Zoran; Gagnon, Gilles; Benzie, John. 1980. Jack pine. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 8-9. [49850]

83. Marozas, Vitas; Racinskas, Jonas; Bartkevicius, Edmundas. 2007. Dynamics of ground vegetation after surface fires in hemiboreal Pinus sylvestris forests. Forest Ecology and Management. 250(1-2): 47-55. [68190]

84. McCune, Bruce. 1982. Lichens of the Swan Valley, Montana. Bryologist. 85(1): 13-21. [21743]

85. McCune, Bruce; Geiser, Linda. 1997. Macrolichens of the Pacific Northwest. Corvallis, OR: Oregon State University Press. 386 p. [28654]

86. McRae, D. J.; Lynham, T. J. 2000. Fire management impacts on boreal forest processes and health. In: Conard, Susan G., ed. Disturbance in boreal forest ecosystems: human impacts and natural processes: International Boreal Forest Research Association--1997 annual meeting proceedings; 1997 August 4-7; Duluth, MN. Gen. Tech. Rep. NC-209. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Research Station: 365-372. [68044]

87. Meades, W. J.; Moores, L. 1989. Forest site classification manual: A field guide to the Damman forest types of Newfoundland. Forest Resources Development Agreement FRDA Report 003. St. Johns, NF: Environment Canada. 295 p. [49220]

88. Miller, Donald R. 1976. Biology of the Kaminuriak population of barren-ground caribou. Part 3. Taiga winter range relationships and diet. Canadian Wildlife Service Rep. Series No. 36. Ottawa, ON: Environment Canada, Wildlife Service. 42 p. [13007]

89. Miller, Donald Ray. 1976. Wildfire and caribou on the taiga ecosystem of northcentral Canada. Moscow, ID: University of Idaho. 129 p. Dissertation. [40270]

90. Morneau, Claude; Payette, Serge. 1989. Postfire lichen-spruce woodland recovery at the limit of the boreal forest in northern Quebec. Canadian Journal of Botany. 67(9): 2770-2782. [9270]

91. Moul, Edwin T.; Buell, Murray F. 1955. Moss cover and rainfall interception in frequently burned sites in the New Jersey pine barrens. Bulletin of the Torrey Botanical Club. 82(3): 155-162. [14550]

92. Mutch, R. W.; Gastineau, O. W. 1970. Timelag and equilibrium moisture content of reindeer lichen. Res. Pap. INT-76. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 8 p. [14623]

93. Niering, William A. 1953. The past and present vegetation of High Point State Park, New Jersey. Ecological Monographs. 23(2): 127-148. [64426]

94. Payette, Serge. 1980. Fire history at the treeline in northern Quebec: a paleoclimatic tool. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 126-131. [16053]

95. Payette, Serge; Bhiry, Najat; Delwaide, Ann; Simard, Martin. 2000. Origin of the lichen woodland at its southern range limit in eastern Canada: the catastrophic impact of insect defoliators and fire on the spruce-moss forest. Canadian Journal of Forest Research. 30(2): 288-305. [36509]

96. Pech, Gy. 1989. A model to predict the moisture content of reindeer lichen. Forest Science. 35(4): 1014-1028. [66261]

97. Pech, Gyula. 1991. Dew on reindeer lichen. Canadian Journal of Forest Research. 21: 1415-1418. [16540]

98. Pegau, Robert E. 1968. Growth rates of important reindeer forage lichens on the Seward Peninsula, Alaska. Arctic. 21: 255-259. [66262]

99. Pegau, Robert E. 1969. Effect of reindeer trampling and grazing on lichens. Journal of Range Management. 23: 95-97. [66264]

100. Pharo, Emma J.; Vitt, Dale H. 2000. Local variation in bryophyte and macro-lichen cover and diversity in montane forests of western Canada. The Bryologist. 103(3): 455-466. [38066]

101. Pouzar, Zdenek; Vezda, Antonin. 1971. Cladonia stellaris (Opiz) Pouz. et Vezda, the correct name for Cladonia alpestris (L.) Rabenh. Preslia. 43(2): 193-197. [66267]

102. Racine, Charles H. 1976. Flora and vegetation. In: Melchior, Herbert R., ed. Biological survey of the proposed Kobuk Valley National Monument. Final Rep. CX-900-3-0136. Change Order No. 3. Fairbanks, AK: U.S. Department of the Interior, National Park Service; University of Alaska, Alaska Cooperative Park Studies Unit, Biological and Resource Management Program: 39-139. [69499]

103. Racine, Charles H.; Anderson, J. H. 1979. Flora and vegetation of the Chukchi-Imuruk area. In: Melchior, Herbert R., ed. Biological survey of the Bering Land Bridge National Monument. Revised final report. National Park Service contract no. CX-900-3-0316. Fairbanks, AK: University of Alaska, Alaska Cooperative Park Studies Unit, Biological and Resource Management Program: 38-113. [86975]

104. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford, England: Clarendon Press. 632 p. [2843]

105. Rettie, W. James; Sheard, John W.; Messier, Francois. 1997. Identification and description of forested vegetation communities available to woodland caribou: relating wildlife habitat to forest cover data. Forest Ecology and Management. 93: 245-260. [27753]

106. Ringius, Gordon S.; Sims, Richard A. 1997. Indicator plant species in Canadian forests. Ottawa, ON: Natural Resources Canada, Canadian Forest Service. 218 p. [35563]

107. Rouse, W. R.; Kershaw, K. A. 1971. The effects of burning on the heat and water regimes of lichen-dominated subarctic surfaces. Arctic and Alpine Research. 3(4): 291-304. [18737]

108. Rouse, Wayne R. 1976. Microclimatic changes accompanying burning in subarctic lichen woodland. Arctic and Alpine Research. 8(4): 357-376. [18738]

109. Rowe, J. S. 1956. Uses of undergrowth plant species in forestry. Ecology. 37(3): 461-473. [8862]

110. Saperstein, Lisa. 1996. Winter forage selection by barren-ground caribou: effects of fire and snow. In: Brown, Kent; Cichowski, Debbie; Edmonds, Janet; Seip, Dale; Stevenson, Susan; Thomas, Don; Wood, Mari, eds. Proceedings of 6th North American caribou workshop; 1994 March 1-4; Prince George, BC. In: Rangifer. Tromso, Norway: Nordic Council for Reindeer Research; 9(Special Issue): 237-238. [62094]

111. Schaefer, James A.; Pruitt, William O., Jr. 1991. Fire and woodland caribou in southeastern Manitoba. Wildlife Monograph No. 116. Washington, DC: The Wildlife Society, Inc. 39 p. [15247]

112. Schimmel, Johnny; Granstrom, Anders. 1996. Fire severity and vegetation response in the boreal Swedish forest. Ecology. 77(5): 1436-1450. [27371]

113. Scotter, George W. 1967. Effects of fire on barren-ground caribou and their forest habitat in northern Canada. Proceedings, 32nd North American Wildlife Conference. 32: 246-259. [16851]

114. Scotter, George W. 1971. Fire, vegetation, soil, and barren-ground caribou relations in northern Canada. In: Slaughter, C. W.; Barney, Richard J.; Hansen, G. M., eds. Fire in the northern environment--a symposium: Proceedings; 1971 April 13-14; Fairbanks, AK. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Range and Experiment Station: 209-230. [15730]

115. Scotter, George W. 1972. Chemical composition of forage plants from the Reindeer Preserve, Northwest Territories. Arctic. 25(1): 21-27. [16563]

116. Scotter, George Wilby. 1964. Effects of forest fires on the winter range of barren-ground caribou in northern Saskatchewan. Wildlife Management Bulletin. Series 1. No. 18. Ottawa, ON: Canadian Wildlife Service, National Parks Branch, Department of Northern Affairs and National Resources. 111 p. [28989]

117. Sedia, Ekaterina G.; Ehrenfeld, Joan G. 2003. Lichens and mosses promote alternate stable plant communities in the New Jersey pinelands. Oikos. 100(3): 447-458. [47208]

118. Sharnoff, Stephen. 1993. Use of lichens by wildlife in North America: A preliminary compilation. [Publisher unknown]. 20 p. On file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. [21464]

119. Simard, Martin; Payette, Serge. 2005. Reduction of black spruce seed bank by spruce budworm infestation compromises postfire stand regeneration. Canadian Journal of Forest Research. 35(7): 1686-1696. [61214]

120. Simon, Neal P. P.; Schwab, Francis E. 2005. Plant community structure after wildfire in the subarctic forests of western Labrador. Northern Journal of Applied Forestry. 22(4): 229-235. [61221]

121. Sims, H. P.; Bruce, N. G. 1966. The ecological effects of prescribed burning on jack pine sites southeastern Manitoba. Internal Report MS-27. Winnipeg, MB: Canada Department of Forestry, Forest Research Laboratory. 24 p. [69688]

122. Skoog, Ronald Oliver. 1968. Ecology of the caribou (Rangifer tarandus granti) in Alaska. Berkeley, CA: University of California, Berkeley. 699 p. Dissertation. [37914]

123. Skuncke, Folke. 1969. Reindeer ecology and management in Sweden. Biological Papers of the University of Alaska. No. 8. Fairbanks, AK: University of Alaska, Institute of Arctic Biology. 82 p. [69580]

124. Stenroos, Soili; Hyvonen, Jaakko; Myllys, Leena; Thell, Arne; Ahti, Teuvo. 2002. Phylogeny of the genus Cladonia s.lat. (Cladoniaceae, Ascomycetes) inferred from molecular, morphological, and chemical data. Cladistics. 18: 237-278. [68276]

125. Stergas, R. L.; Adams, K. B. 1989. Jack pine barrens in northeastern New York: postfire macronutrient concentrations, heat content, and understory biomass. Canadian Journal of Forest Research. 19: 904-910. [8629]

126. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]

127. Strong, W. L. 2002. Lodgepole pine/Labrador tea type communities of western Canada. Canadian Journal of Botany. 80(2): 151-165. [51690]

128. Sulyma,Randy; Coxson, Darwyn S. 2001. Microsite displacement of terrestrial lichens by feather moss mats in late seral pine-lichen woodlands of north-central British Columbia. The Bryologist. 104(4): 505-516. [66283]

129. Swanson, J. David. 1994. SRM 904: Black spruce-lichen. In: Shiflet, Thomas N., ed. Rangeland cover types of the United States. Denver, CO: Society for Range Management: 128. [67494]

130. Swanson, J. David. 1994. SRM 911: Lichen tundra. In: Shiflet, Thomas N., ed. Rangeland cover types of the United States. Denver, CO: Society for Range Management: 133-134. [67501]

131. Tegler, B.; Kershaw, K. A. 1980. Studies on lichen-dominated ecosystems. XXIII. The control of seasonal rates of net photosynthesis by moisture, light, and temperature in Cladonia rangiferina. Canadian Journal of Botany. 58: 1851-1858. [66286]

132. Thomas, Don C.; Kiliaan, H. P. L. 1998. Fire–caribou relationships: (IV) recovery of habitat after fire on winter range of the Beverly herd. Tech. Rep. Series No. 312. Edmonton, AB: Canadian Wildlife Service, Prairie and Northern Region. 115 p. [69053]

133. Thomson, John W., Jr. 1942. The lichen genus Cladonia in Wisconsin. The American Midland Naturalist. 27: 696-709. [19941]

134. U.S. Department of Agriculture, Forest Service. [2008]. National lichens and air quality database and clearinghouse: Pacific Northwest lichen sensitivity ratings by species, [Online]. U.S. Department of Agriculture, Forest Service, Northwest Alliance for Computational Science and Engineering (Producer). Available: http://gis.nacse.org/lichenair/index.php?page=sensitivity [2008, August 14]. [70878]

135. U.S. Department of Agriculture, Natural Resources Conservation Service. 2016. PLANTS Database, [Online]. Available: https://plants.usda.gov /. [34262]

136. Uggla, Evald. 1974. Fire ecology in Swedish forests. In: Proceedings, annual Tall Timbers fire ecology conference; 1973 March 22-23; Tallahassee, FL. No. 13. Tallahassee, FL: 171-190. [18974]

137. Van Cleve, Keith; Viereck, Leslie A. 1981. Forest succession in relation to nutrient cycling in the boreal forest of Alaska. In: Fire and succession in conifer forests of North America. New York: Springer-Verlag: 185-211. [50633]

138. Viereck, Leslie A. 1975. Forest ecology of the Alaska taiga. In: Proceedings of the circumpolar conference on northern ecology; 1975 September 15-18; Ottawa, ON. Fairbanks, AK: U.S. Forest Service, Department of Agriculture, Pacific Northwest Forest and Range Experiment Station. Supplement: 22 p. [7315]

139. Viereck, Leslie A. 1980. Black spruce-white spruce. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 84-85. [50019]

140. Viereck, Leslie A. 1982. Effects of fire and firelines on active layer thickness and soil temperatures in interior Alaska. In: Proceedings, 4th Canadian permafrost conference; 1981 March 2-6; Calgary, AB. The Roger J.E. Brown Memorial Volume. Ottawa, ON: National Research Council of Canada: 123-135. [7303]

141. Webb, Elizabeth T. 1998. Survival, persistence, and regeneration of the reindeer lichens, Cladina stellaris, C. rangiferina, and C. mitis following clearcut logging and forest fire in northwestern Ontario. In: Lankester, Murray; Racey, Gerald; Timmermann, Tim, eds. Proceedings of the 7th North American caribou conference; 1996 August 19-21; Thunder Bay, ON. In: Rangifer. Special Issue. No. 10: 41-47. [65843]

142. Wells, B. W.; Shunk, I. V. 1931. The vegetation and habitat factors of the coarser sands of the North Carolina coastal plain: an ecological study. Ecological Monographs. 1(4): 465-520. [19926]

143. Wetmore, Clifford M. 1990. Lichens of Pictured Rocks National Lakeshore, Michigan. Michigan Botanist. 29(1): 19-26. [17926]

144. Yarie, John. 1983. Forest community classification of the Porcupine River drainage, interior Alaska, and its application to forest management. Gen. Tech. Rep. PNW-154. Portland, OR: US Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 68 p. [69497]

145. Yarranton, G. A. 1975. Population growth in Cladonia stellaris (Opiz.) Pouz. and Vezda. New Phytologist. 75(1): 99-110. [69097]

146. Zoltai, S. C.; Tarnocai, C. 1971. Properties of a wooded palsa in northern Manitoba. Arctic and Alpine Research. 3(2): 115-129. [9778]