LITERATURE CITED Allen, B. 1996. Moss diversity in Maine. Pp. 65-70 in Gawler, S. G., J. J. Albright, P. D. Vickery, & F. C. Smith, Biological Diversity in Maine: An assessment of status and trends in the terrestrial and freshwater landscape. Maine Natural Areas Program. Maine Department of Conservation. Augusta. Dibble, A. C., J. W. Hinds, A. M. Fryday, & N. G. Miller. 2007. Bryophytes and lichens of the alpine and subalpine zones of Mt. Katahdin, Maine, I. Overview

and Conservation Aspects. Manuscript in preparation. Fryday, A. M. 2006. New and interesting North American lichen records from the alpine and subalpine zones of Mt. Katahdin, Maine. Bryologist, in press. Hinds, J. W., A. M. Fryday, & A. C. Dibble. 2007. Bryophytes and lichens of the alpine and subalpine zones on Mt. Katahdin, Maine, III. Lichens. Manuscript in preparation. Kimball, K. D. & D. Weihrauch. 2000. Alpine vegetation communities and the alpine-treeline ecotone

boundary in New England as biomonitors for climate change. Pp. 93-100, in McCool, S. F., D. N. Cole, W. T. Borrie, & J. O'Loughlin (Comps.), Wilderness science in a time of change conference,Vol. 3: Wilderness as a place for scientific inquiry. U.S. Dep. Ag. For. Ser. Proc. Rocky Mountain Research Station-P-15-VOL-3. Miller, N. G. 1996. Liverwort diversity in Maine. Pp. 71-79 in Gawler, S. G., J. J. Albright, P. D. Vickery, & F. C. Smith, Biological Diversity in Maine: An assessment of status and trends in the terrestrial and freshwater

landscape. Maine Natural Areas Program. Maine Department of Conservation. Augusta. Miller, N. G. 2007. Bryophytes and lichens of the alpine and subalpine zones of Mt. Katahdin, Maine, II. Mosses and Liverworts. Manuscript in preparation. Miller, N. G., A. M. Fryday, & J. W. Hinds. 2005. Bryophytes and lichens of a calcium-rich spring seep isolated on the granitic terrain of Mt. Katahdin, Maine, U.S.A. Rhodora 107: 339-358.

SUMMARY

Katahdin (upper zones)

126 (58*)

78 (17*)

290 (0*)

MOSSES

HEPATICS

LICHENS

Maine

376 (Allen, 1996)

149 (Miller, 1996)

868 (Hinds, unpubl.)

39% of the Maine bryoflora

33% of the Maine lichen flora

* no. species reported by Kennedy Expedition

–

BRYOPHYTES FOUND

SUBALPINE- ALPINE ZONES OF MT. KATAHDIN2001 2004

0

50

100

150

200

250

2001 2002 2003 2004 other

hepatics

mosses

total

no

. s

pe

cie

s

SUMMARY

Two principal types of granite have been

recognized on Mt. Katahdin: a more resistant

type (the Summit facies) found at the higher

altitudes (above ca. 1200 m), and beneath it the

less resistant Doubletop facies, which erodes

rapidly into gravel and sand. No mafic dikes or

primary calcium carbonate rock have been found

in or associated with the Katahdin granite.

However, there are a few localized areas of base-

rich seepage. These presumably result from the

dissolution and transport in ground water of

secondary calcium carbonate precipitated in

bedrock cavities, joints, and fractures by

chemical reactions between calcium ions from

primary minerals in the granite (e.g., plagioclase

feldspar, epidote) and bicarbonate ions in water

(Miller et al. 2005). Places of base enrichment

are unusual in areas of granite bedrock. When

they occur, they are very localized.

Hypnum

5.991122Upper Saddle

Brook

5.461256Taber Gully

Stream

5.741411Caribou Spring

5.301356Saddle Spring

5.42930Cleftrock Pool

5.51883Chimney Pond

6.92, 6.621201SPRING SEEP

pHAltitude (m)Location

5.991122Upper Saddle

Brook

5.461256Taber Gully

Stream

5.741411Caribou Spring

5.301356Saddle Spring

5.42930Cleftrock Pool

5.51883Chimney Pond

6.92, 6.621201SPRING SEEP

pHAltitude (m)Location

SURFACE WATER pH: EAST SLOPE AND

TABLELAND, MT. KATAHDIN, FIELD

DETERMINATIONS

NORTH BASIN BASE-RICH SEEP

CHEMISTRY OF WATER SAMPLES FROM

THE NORTH BASIN SPRING SEEP,

CARIBOU SPRING, SADDLE SPRING, AND

TABER GULLY, MT. KATAHDIN, MAINE

LICHENS OF THE NORTH BASIN MINERAL-RICH

SEEP, MT. KATAHDIN. *Species always or often calcicoles

MOSSES AND LIVERWORTS FOUND NEAR THE

NORTH BASIN BASE-RICH SEEP, MT. KATAHDIN, ON

DRY ROCK, HUMUS BENEATH SHRUBS AND

HERBS,OR OTHER NON-SEEP HABITATS.

MOSSES GROWING AT BASE-RICH SEEP

HEADWALL OF NORTH BASIN, MT. KATAHDIN*Species characteristics of rich fens

pH alkalinity

(mg/L CaCO3)

conductivity (us/cm)

Cl¯ (ueq/L)

N032¯

(ueq/L) S04

2¯

(ueq/L) Al3+

( g/l) Ca2+

(mg/L) Mg2+

(mg/L) Na+

(mg/L) K+

(mg/L)

North Basin seep:

ceiling drip

(2003)

7.45

12.1

35.6

7.8

24.5

43.8

17.7

5.76

<0.05

0.85

0.20

same:

ceiling/ Philon-

otis drip

(2003)

7.45

12.1

35.8

7.5

31

43.8

17.1

5.86

<0.05

0.85

0.19

same:

ceiling

drip

(2004)

7.37

10.3

30.5

5.5

25

40.0

45.1

5.20 <0.05

0.16

0.81

Caribou

Spring

5.74 1.1 12.2 6.5 21 46.0 42.3 1.15 0.06 0.17 0.71

Saddle

Spring

5.30 0.04 9.55 7.1 21 43.0 105 0.50 0.05 0.17 0.62

stream: Taber

Gully

6.02

1

13.5

11.4

0

38.3

48

1.32

<0.05

0.50

0.50

TWO PATCHES OF WARNSTORFIA SARMENTOSA

GROWING DOWN OVER A GRANITE SLAB

HEADWALL OF NORTH BASIN, AREA OF SEEP

AND ASSOCIATED VEGETATION

NORTH BASIN, AREA OF BASE-RICH SEEP INDICATED

BY S

NORTH BASIN SEEP AND BEDROCK SLABS BELOW

SEEP

S

Warnstorfia sarmentosaPohlia wahlenbergii

Trichostomum tenuirostre*Plagiomnium ellipticum

Tortella tortuosa

var. fragilifolia

Loeskypnum wickesiae

*Philonotis fontana

*S. warnstorfii*Limprichtia revolvens

*S. subsecundum var.

subsecundum

*Hypnum lindbergii

S. russowii*Fissidens osmundioides

S. quinquefarium*Campylium stellatum

S. papillosumCalliergon stramineum

S. magellanicum*Bryum pseudotriquetrum

Sphagnum fuscumAulacomnium palustre

Sanionia uncinataAmphidium mougeotii

Warnstorfia sarmentosaPohlia wahlenbergii

Trichostomum tenuirostre*Plagiomnium ellipticum

Tortella tortuosa

var. fragilifolia

Loeskypnum wickesiae

*Philonotis fontana

*S. warnstorfii*Limprichtia revolvens

*S. subsecundum var.

subsecundum

*Hypnum lindbergii

S. russowii*Fissidens osmundioides

S. quinquefarium*Campylium stellatum

S. papillosumCalliergon stramineum

S. magellanicum*Bryum pseudotriquetrum

Sphagnum fuscumAulacomnium palustre

Sanionia uncinataAmphidium mougeotii

NORTH BASIN BASE-RICH SEEP

LECANORA CAESIOSORA

HYMENELIA cf. CARNOSULA

STEREOCAULON PLICATILEPROTOPARMELIA CUPREOBADIA

CATILLARIA CONTRISTANS

RHIZOCARPON ANAPERUM

FUSCIDEA GOTHOBURGENSIS

SIGNIFICANT LICHENSSIGNIFICANT LICHENS

LOESKYPNUM WICKESIAE

Scale bar in mm

Distribution ofLoeskypnumwickesiae (Grout) Tuom.

FOSSIL

(from L. Hedenaes, Lindbergia 18:131(1993),

with additions)AULACOMNIUM TURGIDUM

HERBARIUM SPECIMENS

PLEISTOCENE FOSSILS

5 mm5 mm

WARNSTORFIA SARMENTOSAPlants dioicous, with sporophyte (only

population of spore-producing plants

known in northeastern United States)

SIGNIFICANT MOSSESSIGNIFICANT MOSSES

CHIMNEY GULLY

KNIFE EDGE FROM BAXTER PEAK

DIGITAL ELEVATION

MODEL OF KATAHDIN AND

VICINITY. White area is

alpine tundra (treeless area

plus krummholz); dotted line

delimits area above 884 m

(the subalpine and alpine

zones). Key to landmarks:

CP (Chimney Pond), SB

(South Basin), BP (Baxter

Peak, alt. 1606 m [5267 ft]),

SP (South Peak), P (Pamola),

S (Saddle Slide), HP (Hamlin

Peak), NB (North Basin), HR

(Hamlin Ridge), HoP (Howe

Peaks), K (Klondike), NW

(Northwest Basin).

ALPINE PLANT

COMMUNITIES OF

KATAHDIN. AREA

ABOVE THE

ALPINE-TREELINE

ECOTONE(from Kimball and

Weihrauch, 2000).

BACKGROUND OBLIQUE AERIAL

PHOTOGRAPH OF MT.

KATAHDIN FROM THE

SOUTH (Holmberg Aerial

Survey, 1937; image

courtesy of Baxter State

Park Archives).

Key: BP-Baxter Peak, CP-

Chimney Pond, GB-Great

Basin, HR-Hamlin Ridge,

KE-Knife Edge, NB-North

Basin, P-Pamola, SB-

South Basin.

Mount Katahdin (45º54'N, 68º55'W) at 1606 m is the highest mountain in Maine and one of the highest in

northeastern North America. It is exceeded only by peaks of the Presidential Range in the White Mountains

of New Hampshire (e.g., Mt. Washington, 1917 m) and by Mt. Marcy in New York (1630 m). Katahdin has

about 13 km2 of alpine tundra and tundra-krummholz ecotone, as well as steep-sided rocky cirques, which

together otherwise occupy very small areas in the mountains of the northeastern United States.

Katahdin presents an island of alpine habitat about halfway between the equator and the North Pole. It is

ca. 190 km northeast of the Presidential Range in New Hampshire and farther still from the Green

Mountains of Vermont and the Adirondacks in northeastern New York. It is 360 km southwest of mountains

with alpine tundra on the Gaspé Peninsula, Québec. Katahdin has the greatest topographic relief of any

mountain east of the Rocky Mountain Front.

The first lichen and moss specimens from Katahdin were collected by Rev. Joseph Blake in 1856, and the

first liverworts by the Kennedy Expedition in 1900. However, much of high altitude Katahdin has been

unexplored or poorly explored for bryophytes and lichens. We spent 64 person-days between 2001 and 2004

investigating the bryophyte and lichen floras of the subalpine and alpine zones of Mt. Katahdin,

concentrating our efforts primarily on the Tableland and the eastern cirques. Katahdin and its forested

surroundings are parts of Baxter State Park (BSP), and we were required to obtain authorization for this

study from BSP through a special application process.

Our research goals were to: (1) determine what bryophytes and lichens are present in the subalpine and

alpine zones, (2) relate these data to the distribution of the species elsewhere in North America and

worldwide, (3) determine habitats and substrata occupied by Katahdin bryophytes and lichens, and (4)

provide this baseline information to BSP managers to help them develop conservation programs and

coordinate them with recreational use.

Miller focused on bryophytes, Hinds and Fryday on lichens, and Dibble assisted in collecting both groups

and characterized habitat types based in part on vascular plants. Bryophytes were determined by Miller,

lichens by Hinds and Fryday. We relocated most species from historic lists and added many others to the

documented flora. Resulting collections are deposited in the University of Maine Herbarium, the Herbarium

of Michigan State University, the Bryophyte Herbarium of the New York State Museum, and the herbarium of

the University of Maine at Fort Kent. We also examined Katahdin specimes of lichens and bryophytes

housed other herbaria. Further details are presented in Dibble et al. (2007), Fryday (2006), Hinds et al.

(2007), and Miller (2007).

TABLELAND, SADDLE, AND HAMLIN PEAK,

NORTHWARD FROM BAXTER PEAK

[From Rhodora

COLLINSKENNEDY

FERNALD

CHURCHILLWILLIAMS

, vol. 3, pl. 26 (1901)]

KENNEDY EXPEDITION BOTANISTS

AT CAIRN ON BAXTER PEAK

BACKGROUND

We logged 64 person-days during the summers of 2001-04 conducting an authorized bryological and lichenological survey of areas above 884 m on Katahdin (max. alt. 1607 m), a steep-sided,

somewhat isolated mountain in northern New England, with ca. 13 km2 of alpine tundra and well-developed cirques. Prior floristic research, mainly from expeditions in 1900 (Kennedy and

Collins), 1939 (Degelius), and 1949 and 1954 (Schuster), yielded records for 64 mosses, 53 liverworts, and 61 lichens. We confirmed most of these and added many more from field and

herbarium studies. Totals now stand at 126 mosses, 78 liverworts, and 290 lichens. Our collections contained 15 lichens previously unknown in North America (Buellia miriquidica, Catillaria

contristans, Fuscidea gothoburgensis, Hymenelia cyanocarpa, H. cf. carnosula, Lecanora caesiosora, 'Lecidea' commaculans, Metamelanea umbonata, Micarea coppinsii, M. marginata,

Porpidia superba f. sorediata, Rhizocarpon amphibium, Ropalospora lugubris f. sorediata ad int., Scoliciosporum intrusum, Stereocaulon plicatile), 39 others (including three macrolichens)

new to northeastern United States, and at least two undescribed species. Richest habitats for rare lichens were southeast-facing bedrock slopes and rocks near areas of late snow-lie.

Mosses rare in, or unrecorded for, the Northeast included Hygrohypnum smithii, Loeskypnum wickesiae, Neckera oligocarpa, Pohlia drummondii, P. tundrae, Pseudoleskea radicosa, Tortella

tortuosa var. fragilifolia, and Warnstorfia sarmentosa. Acidic fens associated with snowbeds on irrigated east-facing slopes just below alpine tableland contained many mosses also present

in lowland fens. Several interesting discoveries were at or near an alpine calcareous spring-seep on the North Basin headwall, a habitat not noted in earlier botanical work and an anomaly on

a mountain consisting of granite, which lacks carbonate minerals. Calcite produced by the interaction of calcium ions from primary granite minerals and bicarbonate ions in groundwater is

the favored explanation. Our findings on Katahdin suggest that inventories of other mountains in the northeastern United States will reveal much undiscovered cryptogam diversity.

ABSTRACTABSTRACT

Norton G. Miller, New York State Museum, Biological Survey, Albany, New York, 12230-0001, USA

Alan M. Fryday, Michigan State University, Herbarium, Department of Plant Biology, East Lansing, Michigan, 48824-1312, USA

James W. Hinds and Alison C. Dibble, University of Maine, Department of Biological Sciences, Orono, Maine, 04473-5751, USA

Norton G. Miller

Alan M. Fryday

James W. Hinds Alison C. Dibble

Unexpected Bryophyte and Lichen Biodiversity

Documented for the Alpine and Subalpine Zones

of Mt. Katahdin, Maine, U.S.A.

Unexpected Bryophyte and Lichen Biodiversity

Documented for the Alpine and Subalpine Zones

of Mt. Katahdin, Maine, U.S.A.