Vegetation of Amchitka 

Island, Aleutian Islands, 

Alaska

GEOLOGICAL SURVEY PROFESSIONAL PAPER 648
Vegetation of Amchitka 

Island, Aleutian Islands, 

Alaska

By HANSFORD T. SHACKLETTE and others

L. W. DURRELL 

JAMES A. ERDMAN 

JOHN R. KEITH 

WILLIAM M. KLEIN

HILDUR KROG 

HERMAN PERSSON 

H. SKUJA 

WILLIAM A. WEBER

GEOLOGICAL SURVEY PROFESSIONAL PAPER 648

An account of the plant species and plant 

communities that occur in an oceanic 

island environment

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1969
UNITED STATES DEPARTMENT OF THE INTERIOR 

WALTER J. HICKEL, Secretary

GEOLOGICAL SURVEY 

William T. Pecora, Director

Library of Congress catalog-card No. 76-602502

For sale by the Superintendent of Documents, U.S. Government Printing Office 

Washington, D.C. 20402 - Price $1 (paper cover)
CONTENTS

Abstract.__________________________________________

Introduction, by Hansford T. Shacklette_______________

Objectives and limitations of these studies.________

Botanical studies that have been conducted. _______

Nomenclature, and notes on collections____________

Acknowledgments. _ _ ____________________________

Description of the island and its vegetation, by Hansford 

T. Shacklette-__--__-_-__-______-_____-______

Geography.____________________________________

Climate.______________________________________

Physiography and geology.______________________

Soils.__-__-___-__-_-___--________-_______--__-

Plant habitats and plant communities-____________

Page 

1 

1 

1 

3 

3

3

4

4

4

5

5

9

Plant species of Amchitka Island.____________________ 34

Annotated list of vascular plants, by Hansford T.

Shacklette, James A. Erdman, and John R. Keith. 34 

Annotated list of bryophytes, by Herman Persson and

Hansford T. Shacklette-_--_--_--_-----_---__- 40

Annotated list of lichens, by William A. Weber, 

James A. Erdman, and Hildur Krog.___________ 47

Some algae and other protists, by H. Skuja___-___- 50 

Selected references-_________________________ 55

Soil fungi and algae, by L. W. DurrelL.___________ 55

Observations on the flora, by William M. Klein____ 57

References cited-____-_____________-_---_-____--__-- 58

Index._________-___-___-_____--_---__-_-_________- 61

ILLUSTRATIONS

[Plates follow index] 

PLATE 1. Drawings of alga species.

2. Drawings of algae and other protists. Page 

FIGURE 1. Map of Amchitka Island, Alaska, showing locations of vegetation study sites.-______._-__.__._..___.-- ._-- 2

2-35. Photographs:

2. Section of a peat deposit with an upper and two lower ash layers that are marked by a thick growth

of the moss Pogonatum urnigerum,- _______________-___--_--_-_---___---_---_____-_------ 8

3. Effects of soil fertility on the growth of the grass Calamagrostis nutkaensis. _____________________ 8

4. Zonation of Honckenya and Senecio at Makarius Bay._________________________________________ 12

5. Sand dune with cover of Elymus community at the seashore, Rifle Range Point-_________________ 13

6. Sea-cliff habitat on the Pacific Ocean coast at Makarius Bay.__-__________-____-----__--_--___ 14

7. Rock pinnacles ("sea stacks"), Makarius Bay __________________________---_---__-___----___ 14

8. Aerial photograph of the Empetrum heath in an area of many lakes.______-__--_---__-____-_-___ 15

9. Hummocky Empetnim-Carex-lichen community in the vicinity of Cyril Cove____________________ 16

10. A well-developed moss mound in a Cladonia-Carex meadow near Cyril Cove_____________________ 17

11. Ecotone of an Empelrum-Carex-lichQH community and a Cladonia-Carex meadow, near Cyril Cove__ 17

12. Organic bird-perch mound showing characteristic columnar summit composed of Poa arctica subsp.

williamsii, near White House Cove.______________-______--_--______---_---_--____----_- 18

13. Streamside community near Cyril Cove____________________________-__-___--____--__-----__- 19

14. Peat-bank community in the Empetrum heath near Constantine Harbor.________________________ 20

15. Frost scar in the Empetrum heath near Cyril Cove.______-___-_-___-_____-_-__--__-___-_--_-_ 20

16. Spring issuing from the margin of a large soligenous bog in a broad valley near Cyril Cove________ 21

17. Wet sedge-meadow community at the margin of a large lake near Cyril Cove____________________ 22

18. Wet sedge meadow near Cyril Cove.______________________----____-___--_-_---_--___------- 22

19. Ice ridge on the leeward bank of a large lake near Cyril Cove._________________________________ 23

20. Small pool supporting a Juncus-Eriophorum community, surrounded by a Cladonia-Carex meadow,

near Cyril Cove._____________________________________________________________________ 24

in
IV CONTENTS

FIGURES 2-35. Photographs Continued Page

21. Bottom of a dry pool near Constantine Harbor bearing a dense growth of Subularia aquatica. ______ 24

22. Vertical view of a Siphula-Scapania community in a shallow rock-bottomed pool near Crown Re?fer

Point__________________________________________-.___-__.__________ 25

23. Deeply entrenched stream in the Empetrum heath near Chitka Point.__________________________ 25

24. Fragments of a roadside bank that has been severely eroded by wind___________________________ 26

25. Lupinus nootkatensis and Deschampsia caespitosa subsp. orientalis growing on the graveled shoulder

of an unused road near Constantine Harbor.____________________________________________ 26

26. The discontinuous heath habitat, with scattered patches of the Salix-Empetrum community________ 27

27. High-altitude land surfaces that are strongly influenced by solifluction processes, between Chitka

Cove and Andesite Point._____________________________________________________________ 29

28. Rock stripes that have developed on a moderate slope between Chitka Cove and Andesite Point. __ 30

29. Fell-field at the summit of a mountain near Chitka Cove______________________________________ 30

30. Solifluction terraces on the high plateau near Rim Point,_____________________________________ 30

31. Abrupt transition from fell-field to discontinouus heath at Chitka Cove_________________________ 31

32. Low-altitude rock outcrop that is used as a bird perch, near Banjo Point._______________________ 32

33. High-altitude bedrock outcrop near Chitka Cove____________________-___-___-_-_____________- 32

34. Rock outcrops on a mountain summit about 1 mile west of Chitka Cove________________________ 32

35. Sitka spruce (Picea sitchensis) that was planted in front of a club building during World War II___. 33

TABLE

Page 

TABLE 1. Fungi and algae cultured from soil samples taken on Amchitka Island, Alaska_______.__-__-_______-_-_ 56
VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

By HANSFORD T. SHACKLETTE and others

ABSTRACT

An Empetrum heath blankets the low plateaus of Amehitka 

Island, except where interrupted by streams, lakes, sand 

dunes, and man-made disturbances. This heath is composed 

of low shrubs of Empetrum, Vaccinium, Loiseleuria, and varying amounts of many species of sedges, grasses, forbs, lichens, 

and bryophytes. Lakes and pools contain the aquatics Hippuris, 

fsoetes, Ranunculus, Limosella, and Subularia, and poorly 

drained areas around them support communities composed 

largely of Carex or Sphagnum. Moss mounds are abundant 

throughout, and bird-perch mounds occur near the coasts. The 

steep cliffs along the shores support a distinctive flora in which 

Saxifraga, Potentilla, Drafter, Eitrhynchium, and RamaUna are 

predominant. Dense stands of Elymus grow intermixed, at 

places, with Anemone and Ligusticum on the margins of the 

plateaus near the sea coasts. Beaches of sand and cobbles 

commonly have a band of Honckcnya near the water and populations of a tall Senecio among the driftwood. Sand dunes are 

vegetated with dense stands of Elymus and other grasses or, if 

pebbly, with colonies of Rhacomitrium, Schistidium, Ulota, and 

Andreaea.

At altitudes between 400 and 600 feet the Empetrum mats 

lie in strips on unstable pebbly or rocky mineral soil; Salix, 

Loiseleuria, Lusula, Cladonia, and Thatnnolia may also be common components of the mats. On the unstable soil between the 

strips, a few mosses (Andreaea, Rhacomitrium, and Schistidium), Koenigia, Juncus, and Saaeifraga grow as scattered 

colonies or individual plants.

Solifluction ridges and frost-moved cobble deposits at altitudes between 600 and 900 feet are sparsely colonized with intermixed Draba, Sibbaldia, Diapensia, Lupinns, Potentate, Conostomum, Luzula, Veronica, and Rhacomitrium. Mountain 

streamsides and late-melting snow beds have extensive cover 

of Ant7ielia, Nardia, and Harsupella, with Saxifraga, Geum, 

Pyrola, Leptarrhcna, Hippuris, Vahlodea, and other vascular 

plants.

At altitudes between 900 and 1,100 feet most areas are covered 

with boulder fields in which only a few heath mats of Empetrum, 

Salijc, Rhododendron, Vaccinium, and Loiseleuria occur. On the 

highest peaks, where winds commonly are very strong, only 

Carex circinnata and Agrostis horealis grow among the holders 

and outcrops that support Umbilicaria, Ocrolechia, Parmelia, 

and other saxieolous lichens.

The following numbers of plant taxa are reported to occur 

on the island: Vascular plants, 198; bryophytes, 153, lichens, 

84; fungi, 25; and algae, 173; total taxa reported, 633. These 

taxa are distributed among the 15 habitats and 41 plant communities that are described in this report. Numbers of taxa that 

are reported for the first time to occur on Amchitka Island 

are: Vascular plants, 47; bryophytes, 145; lichens, 75; fungi,

25 (no earlier reports known) ; and algae, 173 (no earlier reports known).

The following taxa were found for the first time in the Aleutian Islands: Vascular plants Diapensia lapponica subm>. 

obova-ta and Poa alpina; bryophytes Aongstroemia. longipes, 

Anastrophyllum sphenoloboides, Anomobryum conoinnatum, 

Brachytliecium acutum, B. salebrosum, Bryum arctic-urn, B. pollens, B. salinum, Dicranella subulata, Gymnocolea inftata, 

Hypnum imponen-s, Lophozia opacifolia, Lophozia incisa, Mielichhofcria macrocarpa, Polytrichum sphaerothecium, Spheno7-obus sacoatulus, and Tortula mucronifolia ; fungus Nidi-la 

Candida.

Of the taxa listed above, the following were also found for 

the first time in Alaska: Anastrophyllum sphenoloboides, Brachythecium acutum, Hypnum imponcns, Lophosia opacifolia, 

Nidula Candida, Polytrichum sphaerothecium; and Sphenololus 

saccatulus (first report of its occurrence in North America).

Five new species and a new variety of protists are described 

in this report Chla-mydomonas alasJcensis, C. subangulosa, Oscillatoria facttis, Pseudanabaena- granulifera, Raphidonemagracile, and Synechococcus cedrorum var. pallidus.

INTRODUCTION

By HANSFORD T. SHACKLETTE 

OBJECTIVES AND LIMITATIONS OF THESE STUDIPS

The studies reported in this paper were designed to 

inventory land and aquatic plants that grow on Amchitka island and to describe the relationships of plant 

species and plant communities to climatic and edapHc 

factors. The fresh-water and soil algae and the fungi 

listed in this report were collected incidental to other 

studies; therefore, the lists do not represent the total 

microflora, The fungus Nidula Candida (Peck) White, 

collected during these studies, was reported by Keeves 

(1968) ; other collections of fleshy fungi have not be-m 

identified. Marine algae were not included in this study.

At the time of the investigations, access to the mountainous parts of the island was difficult because of poor 

roads and generally inclement weather; according^, 

only a short time was spent at some mountain sites. 

Many sites on other parts of the island were studied in 

detail in both summer and winter. Plant communities 

in all major kinds of environments were examined after 

traveling on foot or by truck or helicopter to 40 study 

sites (fig. 1). Dense fog in summer and very strong ccM
VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA


 OJ OJ OS "-1

II33 a 

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<D w

§?o

> ^:'
INTRODUCTION

winds in winter at times prevented effective field studies, and good-quality photographs were difficult to obtain under the generally overcast sky. Quantitative ecological studies that were begun are incomplete and are 

not discussed in this report.

BOTANICAL STUDIES THAT HAVE BEEN CONDUCTED

The first known botanical investigation of Amchitka 

Island was made by W. H. Ball of the U.S. Coast Survey on July 30, 1873. His collection of plants is in the 

U.S. National Herbarium and the Gray Herbarium of 

Harvard University (Hulten, 1960, p. 12). No other 

plant collections were made until June 11-12,1929, when 

Misao Tatewaki and K. Takahashi visited Constantine 

Harbor. This area was visited by Yoshio Kobayashi in 

the summer of 1931. The reports of these Japanese botanists were given by Akiyama (1933), Tatewaki and 

Kobayashi (1934), and Satake (1932). Eric Hulten 

collected plants on this island July 9,1932, near "Square 

Bluff" (Bird Cape) and in the middle of the north 

shore. Reports of these collections, as well as sight records, are given in his "Flora of the Aleutian Islands" 

(Hulten, 1960). In 1936 O. J. Murie collected plants 

on Amchitka Island while studying the avifauna. 

Hutchison (1942, p. 783) spent 2 clays at Constantine 

Harbor in the late summer of 1936 and collected about 

50 species of flowering plants. These specimens are in 

the British Museum (Hutchison, 1937, p. 144-145). On 

August 22, 1949, George A. Llano collected lichens 

and bryophytes on this island.

The first systematic collecting of bryophytes, lichens, 

and vascular plants on Amchitka Island was done by 

H. T. Shacklette, in connection with U.S. Geological 

Survey investigations, in November and December 1965, 

August 1966, and June and July 1967. E. E. C. Clebsch 

collected vascular plants and made ecological observations on the island in late June 1967, and A. J. Sharp 

collected bryophytes in early July of the same year. J. 

A. Erdman, M. A. Heifner, J. R. Keith, W. M. Klein, 

and H. T. Shacklette collected plants and conducted 

ecological studies in July 1967; these studies were continued in August 1967 by Klein and Heifner, assisted by 

R. T. Ward and W. L. Slauson. In September and October 1967 and in January and early February 1968, C. 

A. Amundsen and Clebsch continued their ecological 

studies on the island. Shacklette and Erdman collected 

plants and studied the phytoecology of the island February 5-20, 1968. These studies were continued by 

Keith, Heifner, and Slauson during March 19-26, 1968, 

and by R. L. Dix, Kenneth Bierly, Richard Cooley, 

Heifner, David Putman, Skuson, and Walter Younkin 

for various periods during the summer of 1968.

NOMENCLATURE, AND NOTES ON COLLECTIONS

The nonmenclature of the plant species in this report 

generally adheres to that of the following sourcer: 

Vascular plants, Hulten (1968); mosses, Crum, Steers, 

and Anderson (1965); lichens, Hale and Culberson, 

(1966); and liverworts, algae, and fungi, from various 

sources as judged authentic by Herman Persson, H. Skuja, and L. W. Durrell. Some species found on Amchitka 

Island are not listed in the generally accepted references 

given above. For these species the nomenclature follows the usage of specialists in the particular taxonomic group.

Most plants listed in this report were collected by 

Shacklette, and they bear his collection numbers. He 

made approximately 790 numbered collections on Amchitka Island, each at least in triplicate, and one specimen of each is held in the U.S. Geological Survey herbarium in Denver, Colo., for eventual deposit in the U.S. 

National Herbarium. The other two specimens of each 

collection were distributed as follows: Vascular plants 

and bryophytes to the Herbarium of the Swedish Natural History Museum, Stockholm, and to the Herbarium 

of the University of Michigan Museums, Ann Arbor; 

and lichens to the Herbarium of the Botanical Museum, 

Oslo, and to the Herbarium of the University of Colorado Museum, Boulder. The algae collections were segregated from the bryophyte and Isoetes collections, ard 

the segregates are either being held at the University of 

Uppsala or being cultured from soil samples at Colorado 

State University.

ACKNOWLEDGMENTS

This research was supported by the Advanced Besearch Projects Agency (ARPA) of the Department 

of Defense and was monitored by Hansford T. Shacklette under ARPA Order No. 938. Dr. Stanley Ruby 

and Lt. Col. Charles Akard, both with ARPA, were 

most helpful in expediting this work, and Lieutenant 

Colonel Akard also assisted in the first field study. 

Messrs. W. S. Twenhofel, R. E. Davis, R. H. Morris, 

and L. M. Gard, all of the U.S. Geological Survey, assisted with administrative matters and field studies, and 

their cooperation is greatly appreciated. The author 

thanks Messrs. Ted Toren and Joe Brown of the U.S. 

Atomic Energy Commission for logistical support on 

Amchitka Island in June, July, and August 1967 and 

in February and March 1968. Mr. Odd Clemmetson 

helped in many ways with housing and transportation 

on the island during all periods of field study. The author also thanks Mrs. Jessie Bowles for her assistarce 

in the preparation of specimens and manuscript.

350-274 O - 69 - 2
VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

The critical studies of Amchitka Island specimens 

made by Drs. L. W. Durrell, Eric Hulten, Hildur Krog, 

Herman Persson, H. Skuja, and W. A. Weber included 

in this report were supported entirely by their own institutions. The author gratefully acknowledges this 

support and the invaluable assistance of these 

specialists.

DESCRIPTION OF THE ISLAND AND ITS VEGETATION

By HANSFORD T. SHACKXETTE 

GEOGRAPHY

Amchitka Island, the southernmost of the Kat Islands, 

has an area of 114.1 square miles and is the fifth largest 

Aleutian Island (Coats, 1956, p. 86). The island is 

about 35 miles long, in a northwesterly direction, and 

is 3 to 5 miles wide (Powers and others, 1960, p. 522). 

The Bering Sea is to the north, and the Pacific Ocean, 

to the south; therefore, the island is in a region of turbulent winds and waters.

The settlement of the island by the Aleuts occurred 

more than 4,000 years ago, and until Russian occupation 

the island apparently supported a relatively large native 

population (Powers and others, 1960, p. 552). The locations of 40 villages are indicated on a map by Guggenheim (1945, p. 22-23), who reported that many sites 

had been plundered during the military occupation of 

World War II. All known sites of Aleut huts (barabaras) are along the coasts, commonly on sandy deposits. At least one barabara was still habitable in February 1968, but all that remains to indicate the location 

of most huts is a rectangular pattern in the vegetation.

The Russian expeditions to the Aleutian Islands in 

the 18th century were for the purpose of obtaining furs 

of sea otter (Barbeau, 1958, p. 28-32). A Russian village, inhabited by Aleuts and people of Russian descent, was established at Constantine Harbor and apparently flourished for many years; however, when 

Hutchison (1937, p. 145) visited the island in 1936, it 

was entirely uninhabited, although the Russian church 

was in a good state of preservation. At present, the only 

remaining evidence of this early habitation is a small 

cemetery at the head of Constantine Harbor in which 

the most recent tombstone bears a death date of 1930.

In 1913 the entire Aleutian Island chain was set aside 

as a National Wildlife Refuge under the supervision 

of the Fish and Wildlife Service, U.S. Department of 

the Interior. From January 12,1943, until early November 1950, Amchitka Island was used as a military base 

and, at times, was occupied by thousands of troops. 

Many square miles of the eastern part of the island are 

today littered with the ruins of military buildings, most 

of which are uninhabitable. During this occupation, an

extensive network of roads and three airplane runways 

were built in the general area of Constantine Harbor, 

and one road was constructed from this ar°»a to the 

northwest point of the island a distance of about 30 

miles. Apparently, there have been no permanent residents or commercial establishments on the island since 

the military occupation of World War II.

From 1946 to 1963 the island was visited from time 

to time by personnel of the Fish and Wildlife Service, 

the U.S. Weather Bureau, and the U.S. Geological 

Survey (Coats, 1956, p. 84; Powers and others, 1960, 

p. 552). In 1964 work was begun to prepr.re a site 

for nuclear testing, and in October 1965 an 80-kiloton 

nuclear device, buried at a depth of 2,300 feet, was detonated (Army Research and Development Fewsmagazine, 1966).

No native land mammals are on the island. Norway 

rats were introduced probably during World War II 

and are now abundant on the eastern half of the island, 

occurring most commonly along the coasts. Foxes, as 

well as feral dogs remaining from the military occupation, were exterminated after 1945 by the Fish and 

Wildlife Service as a measure to encourage birds to nest 

on the island. Sea otters, Stellar's sealions, and harbor 

seals are common in some coves and on the beaches. 

There are no ants on the island (Hulten, 1960, p. 29), 

mosquitoes (if present) are rare, and earthworms are 

not common, but certain species of Diptera and Hymenoptera are numerous. The avifauna was reported by 

Murie (1959), Kenyon (1961), and others; waterfowl, 

ocean birds, and eagles are abundant, and ptarmigans 

are frequently seen on the heath. The only trees on the 

island are several Sitka spruce (Picea sitchensis} , now 

about 3 feet high, that were planted in front cf military 

buildings during the war. Several species of weedy 

herbs were probably brought to the island by man, 

but they constitute an insignificant part of the present 

flora.

CLIMATE

The Amchitka Island climate is maritime, being cool, 

stormy, and cloudy throughout the year (Arctic 

Weather Central, 1950). Average temperatures range 

from 0°C in January and February to 9°C in August 

(extreme low, 9.5°C in January; extreme high 18.5°C 

in July). Partial to complete cloud cover is rearly constant; it ranges from 99.7 percent of the tine in July 

to 96.8 percent of the time in October. During the summer, fog occurs more than 50 percent of the time, and 

complete overcast occurs more than 65 percent of the 

time. In the winter, there is less fog and overcast. Average wind speeds range from 22 knots in January to 14 

knots in June and July (monthly maximum^ of 52-100
DESCRIPTION OF THE ISLAND AND ITS VEGETATION

knots). Precipitation averages 35 inches annually, including snowfall of 70 inches. The above summary of 

the climate is based on records made from 1943 to 1948 

at the airbase (alt 225 ft); doubtless, the mountainous 

part of the island is colder and windier.

In this study, soil temperatures were measured with a 

recording thermograph for 9-day periods in each of 4 

months. For these periods, averages of hourly temperatures, measured at a depth of 20 cm (centimeters) 

below the surface of the organic mat (a depth considered to be within the zone of greatest root activity), 

follow: June, 5.0°C; July 7.7C C; August, 7.2°C; and 

February, 0°C.

Barometric pressures fluctuate frequently and often 

greatly. During the 2-week period February 9-19, 1968, 

high pressures of 29.48, 28.70, and 29.82 inches of mercury alternated with low pressures of 28.23 and 28.15. 

Abrupt changes in weather generally accompanied 

these barometric fluctuations.

PHYSIOGRAPHY AND GEOLOGY

The physiography and geology of Amchitka Island 

were described by Coats (1956) and by Powers, Coats, 

land Nelson (1960). The discussion that follows is based 

on these reports.

The three landforms on the island are, from east to 

west, low plateaus, rugged mountains, and high 

plateaus (fig. 1). This varied topography is thought to 

have been developed from plateaus of low relief that 

were somewhat disturbed by block faulting and were 

modified by dissection to greatly different degrees by 

marine, stream, and glacial erosion.

Elevated wave-cut platforms, at altitudes ranging 

from 135 to about 500 feet, constitute the lower plateaus 

of the island. Several successive marine benches commonly are present; many are poorly drained and have 

lakes of different sizes and shapes. On eastern Amchitka Island, glacial boulders occur beneath marine 

gravels at altitudes of more than 100 feet.

The mountainous part of the island is between Chitka 

Point and Windy Island and is a sinuous ridge that 

divides the drainage to the Pacific Ocean and to the Bering Sea. This ridge has several summits exceeding 1,100 

feet in altitude, and the summits iare separated by four 

passes at altitudes of about 750 feet. Five prominent 

spurs extend from the main ridge and form divides between major compound amphitheater valleys that head 

in the main ridge. Parts of the ridge and spur summits 

are .gentle slopes eroded across the local rock structure 

and are remnants of an erosional surface of very low 

relief, but it is not known whether the cut surface has 

been glaciated or even whether it is subaeriial or submarine in origin. At the base of the steep slopes leading

from the headwalls of the amphitheater valleys g,re 

benches or remnants of valley floors at altitudes of 700 

to 800 feet in most places. Most of the bedrock surface 

of the benches is grooved and striated, and tarns s.re 

present on some benches and valley floors.

High plateaus occur in the segment of the island 

northwest of Windy Island, at altitudes between 700 

and 800 feet. Two separate areas of undissected surface 

form the two mesas at the west end of the island. The 

high-plateau surface is underlain by colluvium of 

angular rock fragments.

The oldest rocks, probably early Tertiary in age, r.re 

volcanic ash, tuff, breccia, and lava flows generally of 

andesitic composition that apparently were largely 

erupted and emplaced under the ocean. These rocks 

were deformed, uplifted, and eroded before and during 

deposition of the interbedded conglomerate, basaltic 

breccia, and tuff of a later sequence of Oligocene or possibly early Miocene age. Erosion reduced the mass to a 

surface of low relief, now at an altitude of about 1,100 

feet, that was possibly a submarine shoal. Marine sards 

and cobble conglomerate are now exposed at altitudes 

of up to 600 feet.

During late Tertiary and Quaternary times, uplift of 

at least 500 feet, and possibly more than 1,100 feet, took 

place differentially and spasmodically. Abandoned pm 

cliffs and beach deposits are inferred to be structurally 

dislocated remnants of shoreline features of a late Pleistocene interglaciial sea level, judged from the fact tl at 

the fauna of the beach deposit is characteristic of water 

warmer than that which now surrounds the island.

SOILS

The soils of Amchitka Island have been insufficiently 

studied to be classified by soil series; hence, at present 

they can be placed only in the great soil groups to which 

they appear to belong. The following outline of these 

soil groups that are represented by the soils on this 

island is based on the classification of Alaskan soils by 

Kellogg and Nygard (1951, p. 33) and on certain concepts of tundna soils on Kodiak Mand that were proposed by Kieger and Wunderlich (1960, p. 27~32).

A. Zonal soils

1. Tundra

a. Without permafrost (Ando soils of Rieger rnd

Wunderlich) 

B. Intrazonal soils

1. Bog

2. Half Bog 

C. Azonal soils

1. Alluvial

a. From local alluvium

2. Lithosol

3. Regosol
6 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

Eieger and Wunderlich (1960, p. 27) classified certain 

soils as Ando soils; these same soils were classified 

earlier by Kellogg and Nygard (1951) as "Tundra 

without permafrost." Inasmuch as microclimates and 

soils on Kodiak Island are similar in many respects to 

those on Amchitka Island, it is possible that some of the 

Tundra without permafrost soils of the latter island 

should be named Ando soils. They are, however, not so 

strongly influenced by volcanic ash as are those of 

Kodiak Island.

In 1960 the Soil Survey Staff, U.S. Department of 

Agriculture, published a new system of soil classification and in 1967 they expanded and refined the system 

in a supplement. Their definition of soil requires that 

plant life be supported; however, an area covered with 

water that supports only floating plants or with rocks 

that support only lichens is not considered to have soil. 

The lower boundary of soil is considered to be (Soil 

Survey Staff, 1960, p. 1) "the vague lower limit of 

common rooting of the dominant native perennial 

plants, or the vague lower limit of the genetic horizons, 

whichever is the deeper." Within the limits of this 

definition, an extensive system of classification and 

nomenclature was developed on the premise that soils 

should be classified by their properties rather than by 

their genesis. (For a review, see Shacklette, 1961.) If 

the soils of Amchitka Island are to be fully classified 

by the system of the Soil Survey Staff, a much more 

intensive study of their properties than has yet been 

made will be necessary. This new system classifies soils 

into so many more groups than is possible under the 

old system that only examples considered to be typical of 

a Great Soil Group are classified below that is, there 

probably is more than one subgroup in an order on 

Amchitka Island. The correlation of the old and the 

new systems of classification, insofar as it can be done 

at present, is given below.

Tundra without permafrost. If these soils on 

Amchitka Island are to be related to Ando soils, the 

following classification is suggested (Soil Survey Staff, 

1960, p. 136-140; 1967, p. 89-90) : Order, Inceptisols; 

suborder, Andepts; great group, Cryandepts; and subgroup, Lithic Cryandepts. If, however, the small 

amount of ash in their horizons as a whole is considered 

to be most characteristic, they are classified as follows 

(Soil Survey Staff, 1960, p. 136-143; 1967, p. 107-108) : 

Order, Inseptisols; suborder, Umbrepts; great group, 

Cryumbrepts; and subgroup, Lithic Cryumbrepts.

Bog and Half Bog. Because the new system is not 

yet complete, these soils can be classified only as being

in the order Histosols (Soil Survey Staff, 196C, p. 247- 

248). However, some of the Half Bog soils may belong 

to a different order, and they could be classified further 

if sufficient data were at hand.

Alluvial. These soils cannot be correlated with the 

new system on the basis of available data.

Lithosols. The Amchitka Island lithosols are classified as follows (Soil Survey Staff, 1960, p. 136-139; 

1967, p. 93): Order, Inseptisol; suborder, Aquepts; 

great group, Cryaquepts; and subgroup, Histic Cryaquepts.

Regosols. These soils, at both high and low altitudes 

on Amchitka Island, are classified as follows (Soil Survey Staff, 1960, p. 136-143; 1967, p. 98): O"der, Inseptisol ; suborder, Ochrepts; great group, Cryochrepts; 

and subgroup, Lithic Cryochrepts.

The most recent worldwide classification cf soils is 

that of Ball (1967), who devised a system for coding 

soils found in terrestrial biological communities that 

may be proposed for conservation in the International 

Biological Program. This classification recognizes 13 

categories of soil that are designated by letter symbols 

with number subscripts. By eliminating the categories 

of soils that do not occur on Amchitka Island, the author has prepared a key to the soils of Amchitha Island, 

using the system of Ball (1967, p. 123-124), as follows:

1. Soil with dominantly organic surface horizon at least 

50 cm deep. If total soil depth less than 50 cm, then 

surface organic horizon directly succeeded by unaltered rock soil type O.

Soil without dominantly organic surface horizon or 

with organic surface horizon succeeded by mineral 

soil at less than 50 cm depth 2.

2. Well drained (that is, no evidence of strong impedance or waterlogging above 40 cm depth) 3. 

Poorly drained (that is, evidence in mottled colors 

of strong impedance or waterlogging nearer surface than 40 cm) Soil type P2.

3. Immature profile (that is, with weakly developed 

shallow soil formation, possibly with little biological activity) 4.

Well-developed horizon sequence with moderate to 

strong biological activity 5.

4. Immaturity resulting from climatic factors; for example, very low rainfall or temperature, or both 

soil type Ii

Immaturity resulting from lack of time for soil formation to proceed; for example, on recent alluvium, 

dune sands, and eroded surfaces soil type I2
DESCRIPTION OF THE ISLAND AND ITS VEGETATION

5. Shallow or simple profiles of A horizons overlying

parent material soil type F3 

A(B) C profiles a soil type F4 

ABC profiles; that is, with a horizon accumulation of

clay or iron oxides, or both soil type F5

Because the classification of the Soil Survey Staff 

(1960, 1967) cannot at present be fitted to all soils on 

the island, subsequent parts of this report will use the 

classification based primarily on Kellogg and Nygard's 

(1951) that was presented earlier, and the system of 

Ball (1967).

In 1946 Kellogg and Nygard (1951, p. 41, 43), of the 

U.S. Department of Agriculture, obtained soil samples 

and data for a profile description of a smooth, fairly 

well drained slope on Amchitka Island. They classified 

the soil as Tundra without permafrost and gave the 

following description of the soil profile:

Aoo and Ao. 5 to 0 inches, dark-reddish-brown peaty mat. 

Ai. 0 to 5 inches, nearly black humus-rich silty very fine 

sandy loam; the slick moist soil is held together by 

many roots. 

B2. 5 to 10 inches, nearly black granular silty very fine

sandy loam. 

B3. 10 to 15 inches +, dark-grayish-brown silty very

fine sandy loam. In the lower part, horizontal streaks

suggest ash layers. 

C. At 30 inches, yellowish-brown silty very fine sandy loam.

The low plateaus of Amchitka Island, because of 

their varied relief, bear soils that include all the soil 

groups on the island. The soil profile given above is 

probably typical of soils of the Empetrwm, heath on the 

undulating plateau surfaces. Land surfaces of steeper 

slope may have Half Bog soils and soil intergradations 

between Half Bog and Bog soils at the base of the slope. 

Small areas of local Alluvial soil are common near 

streams, and Bog soils occur at lake margins and on 

nearly flat areas of the plateaus. Sand dunes near the 

coasts and beach sands are considered to be Kegosols. 

Shallow Lithosols develop on rock outcrops, both along 

the coasts and inland.

The mountain section of Amchitka Island, because of 

the predominance of rock outcrops and colluvium, contains most Lithosols. Bog and Half Bog soils, however, 

have developed along some of the high mountain streams 

and around the tarns of the large amphitheaters. Much 

of the total area of unconsolidated surficial deposits at 

high altitudes supports litle or no vegetation because of 

instability produced from steepness of slope and frost 

action; these deposits cannot be considered to be soil.

1 (B) and B horizons are defined (Ball, 1967, p. 121) as, "Horizons 

lying between A horizons and the parent material C horizons, which showweathering and release of iron oxides without their transportation, (B) ; 

or which are horizons of deposition of transported sesquioxides and/or 

clay, B."

The surficial material on the high plateaus, owing to 

less relief, has greater stability than that on the mountain section, and large areas of sparse vegetation occur 

on the Lithosol of the rock fields. Interrupted vegetation mats, in which Empetrum, Vaccinium^ and Saliw 

predominate, develop a thin organic soil in and under 

them, but the profile of this soil has little in common 

with that of the Tundra soils at lower altitudes.

The author did not observe permafrost in any section 

of Amchitka Island. Kellogg and Nygard (1951, p. 11, 

135) stated that there is none in the Aleutian Islands. 

However, local areas of perennially frozen ground possibly occur under the late-melting snow beds in the 

mountains of Amchitka Island, and Lithosols at the 

margins of permanent snowfields on some of the higher 

Aleutian Islands are probably underlain by frozen 

ground.

All soil profiles of the low plateaus of Amchitka Island examined in the present study contained an ash 

layer 1-1.5 cm thick in the upper part of the B horizon 

and two ash layers of about the same thickness in the 

lower part of the B horizon. A soil profile on Tanaga 

Island, about 115 miles east of Amchitka Island, was 

reported by Anderson and Bank (1952, p. 84) to have 

somewhat similar, but more numerous, ash layers. There 

have been no volcanoes on Amchitka Island since the 

present soil was formed, but recently active volcanoes 

on other islands in this region may have produced this 

ash. Powers (1958, p. 67) reported the activity of Cerberus Volcano on Semisopochnoi Island, which is about 

40 miles north of Amchitka Island, as follows: 1772, 

smoke; 1790-92, smoke; 1830, smoke; 1873, active. He 

referred to smoke as "a word used in records; it refers 

to steam or steam-and-ash clouds." Therefore, it is 

possible that the ash deposits on Amchitka Island came 

from the Semisopochnoi Volcano or from volcanoes on 

other islands nearby. Powers (1958, p. 67) reported for 

Little Sitkin Island, "1776, active; 1828, smoke;" and 

for Segula and Kiska Islands, "steaming intermittent."

A trench that was dug through a peaty soil deposit of 

the Empetrum heath on Amchitka Island in 1965 exposed three ash layers in the profile. This trench was 

examined again in July 1967; a moss, Pogonatum umigerum, had colonized only the ash layers. The peat part 

of the profile bore no mosses or other vegetation; therefore, the growth of the moss on the ash made the ash 

layers very conspicuous (fig. 2). The "preference" of 

this moss for a mineral soil was known from other observations, but only rarely is such a specific requirement 

for a particular substrate demonstrated so clearly.

Ash layers in the profile described above contained 

preserved stems of sedge and Empetrum. These stems 

were removed from the peat and ash matrix; radiocar-
8 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

FIGURE 2. Section of a peat deposit with an upper (a) and two 

lower (;&) ash layers that are marked by a thick growth of the 

moss Poffonatum urnigerum. Timber (2 in. X 4 in.) in upper 

center provides scale. No other plants grew on the vertical 

bank, which was formed by excavation approximately 2 years 

before this photograph was made on July 28,1967.

bon dates of samples from the top, middle, and lower 

ash layers were 750 ±250 years, 1,950±250 years, and 

I,740=fc250 years B.P. (before present), respectively. 

The middle and lower ash layers were so close together, 

and the stems in them were so intermixed, that satisfactory separation of the stems was not possible. The dates 

of these stem samples, therefore, were averaged as 1,845 

years B.P. (Shacklette and Rubin, 1969).

Although mechanical properties of soils related to engineering operations have 'been studied on Amchitba 

Island, results have not been published. Such studies, 

because of the broad definition of soil that is used, have 

but little relevance to ecological investigations.

An evaluation of the soils of this island as media for 

the support of native plants is best made by observing 

the vegetation that grows on the soils. From this standpoint, the soils appear to be fertile, because most land 

surfaces especially those of the low plateaus are covered with dense mats of vegetation. This dense plant 

growth, however, does not indicate that the soils meet 

the requirements for optimum growth of most plant 

species that are present, nor does it indicate that the soils 

are fertile, as judged by agricultural standards. The 

effects of a soil that is more fertile than the average soil

on the island can be seen by the vigorous growth of many 

plant species on bird-perch mounds, calcareous dunes, 

kitchen middens, barabara sites, and recently formed 

soil banks (fig. 3). The Amchitka Island soils, if compared with agricultural soils or with the soils of most 

temperate-zone plant communities, are low in nutrient 

elements, especially calcium, phosphorus, and nitrogen. 

Kellogg and Nygard (1951, p. 41) gave a chemical 

analysis of a soil sample from Amchitka Island and 

wrote, "The data * * * show a high exchange capacity 

but a very low base saturation. The magnesium is low, 

but very high in relation to calcium. Available phosphorus is exceedingly low." They (1951, p. 43,116) reported 

the following phosphorus content, in parts per million, 

of samples from this island:

Ash of a sedge-grass sample __ ____ 24 

Soil samples [presumably not ashed] :

Aw horizon______ 12 

An horizon_________ 2 

B to Chorizons [4 samples] 0

A similar trend in calcium and potassium content of 

soils and plants from Amchitka Island was reported by 

Kellogg and Nygard. They also reported a carbon-nitrogen ratio (1951, p. 14) of 15 to 1 in the AI soil horizon, 

as compared with that of 26 to 1 in the B2 horizon, which 

suggests the formation of peat in place accompanied by 

a loss of nitrogen by leaching.

FIGURE 3. Effects of soil fertility on the growth of the grass 

Calamagrostis nutkaensis. Soil banks had been formed in digging-in a pyramidal tent; later a latrine was located on this 

tent site and was closed February 4, 1944. The grass is least 

vigorous on the thin rocky soil in the foreground, more vigorous on the soil banks, and most vigorous on the latrine site, 

where the grass tussocks are about 1 meter high. Photographed February 16, 1968
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 9

Soil profile samples were collected from an Empetrum heath on Amchitka Island by Shacklette on August 13,1965. Descriptions of these samples, which were 

taken at 4.5-cm intervals of a soil profile 0.5 m (meter) 

thick; their percentages of ash; and their chemical contents, expressed as percent of ash, follow:

Sample description Ash CalciumPotas- Phos- SO4 

sitim phorus radical

Living heath plants, mixed __ . _ _ .

Dead heath plants (Aoo horizon). ....

Peat--. 

Mucky peat.-.. -- _ ... _ --- __ --..

Peaty muck.. ___ - ________ - Muck (including ash layer) ..- ____ .

Muck ---.--_--.-.-- __ -.-- __ _ ...

Muck (including ash layer) -------- _ . Muck (including ash layer) - __ ..--..

Muck..-. _ -. __ _ _ ---- _ .-----.

Organic and mineral soil

 --- 5.7

.--- 6.2

.... - 11.0

...... 13.7

...... 26.7

...... 57.2

...... 21.5

44.2 

..... . 45.0

...... 31.1

46.5

7.5

8.2

3.8

3.5

3.4

3.7

2.2

3.3 

2.3

1.7

2.1

7.2

2.2

1.7

1.7

1.4

1.3

1.1

.9 

1.1

1.0

.6

2.0

1.6

.8

.9

.4

.16

.6

.3 

.2

.4

.3

3.6£ 4.1*

3.24

3.93

2.14

.93

2.05

1.33 

1.17

1.17

.97

These samples were analyzed by T. F. Harms and 

C. S. Papp, U.S. Geological Survey, Denver, Colo.

The concentration of these elements in plants is 

clearly shown by these data, and a greatly abbreviated 

biogeochemical cycling of these elements is indicated. 

That is to say, although the supply of these essential 

nutrients is deficient in the pedosphere as a whole, the 

amounts that are present are held where they have the 

greatest beneficial effects on plants in the upper root 

zones and in the plants themselves. The trends in nutrient-element concentration in plants and soil horizons 

shown in the samples just described were supported by 

those found by analyses of other similar samples.

PLANT HABITATS AND PLANT COMMUNITIES

Plant communities of the Aleutian Islands have been 

described by Hulten (1933, 1960), Tatewaki and Kobayashi (1934), Walker (1945), and Bank (1951,1956). 

These authors, however, almost entirely neglected the 

bryophytes and lichens in the communities or associations that they established; yet these two groups of 

cryptogams are very important and, at places, dominant 

elements of plant communities in the Aleutian Islands. 

One objective of the present study was to give full consideration to all these groups of plants in describing the 

vegetation of Amchitka Island; therefore, the ecological units of the former investigators were not adopted 

in full, although there is a general correspondence of 

many plant communities described in this paper to those 

of the earlier writers.

The organization of vegetation that is given in this 

paper is intended neither to form a hierarchical system 

nor to establish plant communities as formal taxonomic 

entities in a phytosociological system. It is, on the other 

hand, a method of naming groups that are easily recognized in the field to be composed of species that have

nonrandom association, although many species, if considered individually, appear to be somewhat randomly 

distributed throughout one or more types of habitat. 

Groups of plants that are composed largely of vascular 

plants, of bryophytes, or of lichens were given equal 

consideration in establishing the communities. Some 

communities that are described are much larger than 

others in a real extent and number of component species. The size of a community is determined largely by 

the areal extent of habitat factors in particular combinations. Quantitative studies of the compositional elements of a community have not been completed; the 

present establishment of the communities is subjective 

to a high degree and is likely to be modified when quantitative data are available.

The vegetation of Amchitka Island can be described 

conveniently by presenting the species according to 

their natural occurence in plant communities, then relating these communities to distinct habitats in which 

they occur. Causal factors in the development of plant 

communities may be implied in this method of describing the organization of the vegetation, but they are 

not stated more positively than this investigation warrants. Detailed measurements of physical factors of the 

habitats are at present almost entirely lacking; likewise, the responses of many plant species to these factors are largely unknown. Within these limitations 

imposed on ecological interpretations, however, the 

communities and habitats may be described in such a 

manner that the kinds of substrates and microclimates 

essential for the formation of the communities are suggested. At the same time, the occurrence of individual 

species in only one or in several communities is an indication of their ecological amplitude, whatever their 

specific requirements and tolerances may be.

Some plant communities are separated by ecotones 

that are as great in extent as the adjacent communities 

themselves; hence, the exact delimitations of each community can be made only arbitrarily. An example of 

communities that have this relationship are the Empetrum-Carex-lichen and the Careoe-lichen meadow communities. Similarly, the latter community may intergrade, through a broad ecotone, with a wet-sedge 

meadow community.

The vegetation on this island is mainly one layered; 

there are but few examples of one plant species regularly overtopping another, if a few tall herbs and some 

of the minute liverworts and mosses are excluded. There 

are no native trees, and the low shrubs are intertwined 

with herbs, mosses, and lichens in a manner that allows 

one, then another, to lie on top of the intricate mat of 

vegetation. For example, Empetrum may overgrow, or 

be overgrown by, certain mosses and lichens. The pros-
10 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

trate willows have leafy stems that may temporarily 

suppress mosses and lichens, but eventually these 

stems are completely buried by the moss and lichen 

mat. In some places, saxicolous mosses and lichens grow 

over each other.

An adequate description of community dynamics on 

Amchitka Island must await further study. If phases 

of succession are believed to be evident, however, they 

are so stated. The roles of some biotic, edaphic, and 

microclimatic agents in the establishment and perpetuation of certain communities are strongly indicated by 

characteristics that can be observed in the field. Likewise, the interaction of species in a community can. at 

places, be appraised. The interpretations of community 

dynamics presented in this paper are, however, only 

provisional.

The names of plant communities have been derived 

from either their characteristic species or their distinctive site features, according to which terminology is 

more useful for recognizing the communities in the 

field. In this paper, usage of the term "characteristic 

species" agrees in general with that of Costing (1956, p. 

74-75) and of other writers who include in this term the 

species with a high degree of fidelity (exclusiveness) 

to a particular community. An estimate of the quantitative characters abundance and cover, however, was also 

used in some of the more complex communities to select 

species that are considered to be characteristic of a community, even though their fidelity to the community is 

low. For example, Elymus armarius subsp. irwllis var. 

mollis is considered to be a characteristic species of two 

communities, being dominant in one and codominant 

with Ligusticum and Anemone in the other; moreover, 

it is listed as occurring in several other communities. 

Similarly, Empetrum nigrum subsp. nigrum is a characteristic species in two communities and occurs in nine 

others. To ignore these species as characteristic of some 

communities because their degree of fidelity is low 

would not be a realistic use of the word "characteristic" 

in its less restrictive meaning and would not convey the 

importance of the species in the community. If quantitative data were available, the importance of species in 

a community could be more accurately stated than by 

using the term "characteristic species" in these ecological discussions.

In contrast to the examples given above, some of the 

species that grow on sea cliffs and in snow beds occur 

only in a single type of community. They have, therefore, the highest degree of fidelity and, also, are considered to be characteristic species.

Other species that occur in ia community also are 

listed. The species list given for each community is complete only insofar as collected specimens and field notes 

indicate the presence of the species. An enumeration of 

all species that occur in some communities would add 

new names to the list but perhaps would not contribute 

significantly to a better characterization of the 

community.

The land habitats of Amchitka Island and the communities of plants that occur in these habitats are outlined as follows:

1. iStrand

a. Honckenya-Senecio community

2. Dune

ia. Poa-Deschampsia-Festuca community 

 h. Elymus community 

c. Rhacomitrium-Schistidium-Ulota community

3. Sea cliff

a. Eurhynchium-Pucainellia-Caloplaca community

b. Potentillea-Draba-Sawifraga community

c. Xanthoria-Ramalina, community

d. Elymus-Ligusticum-^nemone community

4. Empetrum heath

a. Empetrum-Ccwese-Iichen community

b. Cladonia-Carew meadow community

c. Moss mound community

d. Organic bird-perch mound community

e. iStreamside community

f. 'Peat-bank community

g. Frost-scar community 

6. Bog

a. Sphagnum bog community

b. Philonotis-Pamassia community

c. Scapania-Nardia^Marsupella community

6. Marsh

a. Wet sedge-meadow community 

b. Caltha-Claytonia community

7. Lake

a. Hippuris-Ranunculus community

b. Isoetes-Ranunculus-Limosella community

8. .Pool

a. Juncus-Eriophorum community 

b. Subularia-CaMitriche community 

c. SiphulOrScapama community

9. Stream

a. Fontinalis-Ranunculus community

10. Buderal

a. Calamaffrostis community

b. Lupinus-Amica community

c. Equisetum-Sagina-Ceratodon community

11. Discontinuous heath

a. Salix-Empetrum community

b. Andreaea-Schistidium-Rhaoomitrium community

12. iSolifluction

a. Fell-field community

b. Solifluction terrace community

13. Alpine meadow

a. <Snow-bed community

b. Alpine meadow community

14. Inland bedrock

a. Low-altitude bedrock community 

b. High-altitude bedrock community
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 11

15. Miscellaneous habitats, substrates, and communities 

a. Carcasses 

b. Imported wood 

c. Plantings 

d. Burned areas

The most recent classification of vegetation on a 

worldwide basis is that of Fosberg (1967), who devised 

a system for use in coding vegetation of terrestrial biological communities that may be proposed for conservation in the International Biological Program. This system uses a combination of vegetational features that 

may be grouped under "structure" (the arrangement in 

space of the components of vegetation), and "function" 

(features that suggest special adaptation to past or 

present environmental situations), and relies on "composition" (the species comprising the vegetation) only 

for the lowest categories. In this system, vegetation is 

first classified in one of three "Primary Structural 

Groups" closed, open, and sparse or desert.

I have constructed an outline of classification, based 

on Fosberg's system, for the plant communities of 

Amchitka Island that eliminates the categories of vegetation that do not occur on this island. Fosberg's system, as presently organized, does not permit all plant 

communities of this island to be fully classified to 

Formation or Subformation. The outline that follows 

classifies these communities only insofar as is possible 

without devising new categories.

PRIMARY STRUCTURAL GROUP 1 

[Closed vegetation (crowns or peripheries of plants touching or overlapping)]

CLASS 1C. Dwarf scrub (closed predominantly woody vegetation

less than 0.5 m high). 

GROUP 1C1. Evergreen dwarf scrub. 

FORMATION 1C12. Evergreen broad sclerophyll dwarf

scrub. 

SUBFORMATION 1C12C. Microphyllous evergreen

dwarf scrub. 

Community: EmpetrumCarex-lichen.

GROUP 1C2. Deciduous dwarf scrub. 

FORMATION 1C21. Deciduous orthophyll dwarf scrub. 

SUBFORMATION !C21a. Deciduous orthophyll dwarf

scrub. 

Community: Salix-Empetrum. 

CLASS 1L. Tall grass (closed herbaceous vegetation more than

1 m high, predominantly graminoid).

GROUP 1L2. Seasonal tall grass (turning brown in winter). 

FORMATION 1L21. Seasonal orthophyll tall grass.

Communities: Elymus-LigusticumAnemone; Elymus.

CLASS 1M. Short grass (closed herbaceous vegetation, less than 

1 m high, predominantly graminoid).

GROUP 1M2. Seasonal short grass.

Community: Organic bird-perch mound. 

FORMATION 1M21. Seasonal orthophyll short-grass

meadows.

Communities: Poa-Deschampsia-Festuca; wet sedge meadow; Calamagrostis. 

FORMATION 1M22. Seasonal orthophyll marsh.

Community: Juncus-Eriophorum.

CLASS IN. Broad-leafed herbaceous vegetation (closed vegetation predominantly of broad-leafed herbaceous 

plants). 

GROUP 1N2. Seasonal broad-leafed herbaceous vegetation.

Community: Honckenya-Senecio. 

FORMATION 1N21. Seasonal broad-leafed herbaceous

vegetation.

Communities: Streamside; CalthaClaytonia; alpine meadow; lowaltitude bedrock. 

CLASS 1O. Closed bryoid vegetation.

GROUP 1O1. Closed bryophyte vegetation.

Communities: Moss mound; peat bank; Scapania-Nardia-Marsupella. 

FORMATION 1O11. Sphagnum bog.

Community: Sphagnum bog. 

FORMATION 1O12. Moss meadow.

Communities: PhUonotis-Parnassia;

snow bed.

GROUP 1O2. Closed lichen vegetation. 

FORMATION 1O21. Lichen bog.

Community: Siphula-Scapania. 

FORMATION 1O22. Lichen meadow.

Community: Cladonia-Carex meadow. 

CLASS IP. Submerged meadows (vegetation of rooted aquatic

herbs). 

GROUP 1P1. Evergreen submerged meadows.

Community: Subularia-Callitriche. 

FORMATION 1P11. Evergreen watergrass.

Communities: Isoetes-RanunculusLimosella; Fontinalis-Ranunculus.

GROUP 1P2. Seasonal submerged meadows (plants, at least

their shoots, disappearing in winter). 

FORMATION 1P21. Seasonal watergrass.

Community: Hippuris-Ranunculus.

PRIMARY STRUCTURAL GROUP 2

[Open vegetation (plants or tufts of plants not touching, but crowns not separated by 

more than their diameters; plants, not substratum, dominating the landscape)]

CLASS 2G. Steppe (open herbaceous vegetation, tufts or plants 

discrete, yet close enough to dominate the 

landscape). 

GROUP 2G2. Seasonal steppe.

Communities: Potentilla-Draba-Saxifraga; 

Equisetum-Sagina-Ceratodon.

CLASS 2H. Bryoid steppe.

GROUP 2H2. Open lichen vegetation.

Community: Xanthorea-Ramalina.

350-274 O - 69 - 3
12 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

PRIMARY STRUCTURAL GROUP 3

[Sparse vegetation or desert (plants so scattered that substratum dominates 

landscape)]

CLASS 3B. Desert scrub (scattered shrubs in an otherwise bare

or only ephemerally vegetated landscape). 

GROUP 3B1. Evergreen desert scrub. 

Community: Fell-field. 

CLASS 3C. Desert herbaceous vegetation (scattered herbaceous

plants only). 

GROUP 3Cl. Evergreen desert herbaceous vegetation.

Communities: Eurhync hium-Puccinellia-Caloplaca; Andreaea-Schistidium-Rhacomitrium; 

high-altitude bedrock. 

FORMATION 3C12. Evergreen psammophyte desert.

Community: Rhacomitrium-Schistidium-Ulota. 

FORMATION 3C13. Lichen tundra sparse phases.

Community: Solifluction terrace.

The occurrence of most species in the communities 

was determined by collecting specimens in the field. 

Some of the easily recognized species were, however, included in the community descriptions on the basis of 

sight records. In both the community descriptions that 

follow and the annotated check lists of species, the communities are designated by letters and the habitats by 

numbers; therefore, the communities and habitats in 

which each species was found can easily be determined. 

The community name is followed by the community 

classification according to the system of Fosberg (1967).

The great soil group, or groups, and the soil classification, according to the system of Ball (1967) of the 

substrate on which the community occurred, follow the 

community classification. The approximate pH value, 

or range in values, is given for the substrates of most 

communities. The pH of the water in which aquatic or 

semiaquatic communities occur also is provided. These 

values were determined for soils by making a slurry, 

or extract, consisting of two parts, by volume, neutral 

water and one part soil or other substrate and by 

measuring pH with a potentiometer. The pH of water 

was measured directly with the same instrument. These 

tests were performed at base camp soon after the 

samples were collected.

The photographs of habitats and communities were 

made by Shacklette, except as noted.

1. STRAND HABITAT

Occurs around the island wherever a shoreline bench 

bears a deposit of sand, pebbles, or cobbles. The presence of abundant driftwood in this habitat indicates 

that this habitat is inundated by sea water from time 

to time.

a. Honckenya-Senecio community (1N2).

Soil: Kegosol; type ?2; pH, 6.8-7.0. This community forms a distinct zone of dense vegetation 

near sea level (fig. 4). The Senecio may be as much 

as 15 dm (decimeters) high at places and so closely 

spaced that passage through the community is 

difficult. Stands of Lathyrus occur on sand banks 

of the beach, and scattered plants of Galium commonly grow in the piles of driftwood.

Characteristic species: Honckenya peploides 

subsp. major and Senecio pseudo-arnica.

Other species: Cerastium fischerianum, Ceratodon purpureus, Claytonia sibirica, Cratoneuron 

filicinum, Eurhynchiwn praelongum, Galvwm 

aparine, G. trifidvm subsp. cohimbianwn, Heracl&u/m lanatu/m, Juncus a/rcticus subsp. sitchensis, 

Lathyrus maritimus subsp. pubescens, Mertensia 

maritima,) Peltigera spuria, Rumex obtusifolius, 

Sagina occidentalis, and Stellaria calycantha 

subsp. calycantha.

2. DUNE HABITAT

This habitat type designates sandy and pebbly deposits that occur most commonly on top of seacoast 

bluffs that are from 25 to 100 feet above sea level, where 

the sand is moved by wind.

a. Poa-Deschampsia-Festuca community (1M21).

Soil: Kegosol; type P2 ; pH, 6.7 6.8. On elevated 

beach deposits of sand and shells that are 100 to

FIGURE 4. Zonation of Honckenya (nearest the bare beach) and 

Senecio at Makarius Bay. Photographed June 17,1967.
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 13

200 feet above the present sea level. Calcium derived from the shells probably favors the development of this community that is composed almost 

entirely of grasses and Equisetum.

Characteristic species: Poa eminens, Deschampsia beringensis, and Festuca rubra subsp. aucta.

Other species: Agrostis exarata, Bromus sitchensis var. aleutensis, Elymus arena/ring subsp. 

mollis, Eguisetum arvense, and Poa lanata. 

b. Elymus community (1L21).

Soil: Kegosol; type Lz pH, 6.6-7.0. On dunes 

composed of dark-colored sand derived from volcanic rocks (fig. 5).

Characteristic species: Elymus arenaris subsp. 

mollis var. mollis. This community is dominated by 

a dense growth of this grass which at places forms 

tussocks that are as much as 12 dm high and are 

separated from each other a distance of 14-30 cm. 

Inasmuch as the distinct tussocks occur on steep 

slopes, soil slippage probably causes their separation. This community may extend to the sandy 

beaches and dominate the old beach terraces, as is 

shown in figure 5; more commonly, however, it occupies dunes at slightly higher altitudes.

Other species: Bryophytes that are able to grow 

on the almost bare soil between the grass tussocks 

include Antitrichia curtipendula, Bryum pseudotriquetrum, Cephaloziella arctica, Ceratodon purpweus, Eurhynchium praelongum, Lophocolea 

cuspidata, Nardia scalaris, Plagiothecium roeseanum, PogonaPum umigerum, PoJila cruda, Rhytidiadelphus squarrosus, and Tor-tula mucronifolia. 

The flowering plants Claytonia sibirica, Geum macrophyllum subsp. macrophyllum, and Heracleum

FIGURE 5. Sand dune with cover of Elymus community at the 

seashore, Rifle Range Point. Grass clumps (at top) are about 

3 feet high. Photographed February 17,1968.

lanatum are sparsely scattered among the tussocks. 

The lichens Caloplaca sp. and Peltigera comma, were 

found on moss-covered driftwood in this community.

c. Rha£omitrium-)Schistidium-Ulota community 

(3C12).

Soil: Regosol; type I2 ; pH, 6.8-6.9. On dunes 

composed of dark-colored sand, pebbles, and cobble. 

Rhacomitrium and Ulota grow on the cobbles and 

pebbles, and Schistidium grows on the sand. This 

community and its unattached moss polsters of 

Schistidium, were described and illustrated by 

Shacklette (1966).

Characteristic species: Rhacomitrium lanuginosum, Schistidium apocarpum, and Ulota 

phyllantha.

Other species: Andreaea rupestris, Brachythecium albicans , Ceratodon purpureus, Drepanocladus uncinatus, Lecanora atra, L. aleutica, 

Lecidea macrocarpa, Ligusticum scoticum subsp. 

hultenii, Pogonatum urnigerum, Rhacomitrium, fascicidare, R. ericoides, Stereocaulon alpinum, Tortula mucronifolia and Umbilicaria proboscidea.

3. SEA-CLIFF HABITAT

The island is almost completely bordered with precipitous sea cliffs of andesite, breccia, and other igneous 

rocks. At a few locations on the southeast end of the 

island, the cliffs are composed of sedimentary deposits 

that give but little support for plants. The cliffs range 

in height from only a few feet to at least 100 feet above 

sea level on the central and southeastern parts of the 

island; commonly, they are higher on the northwestern 

shores. Plant communities on these cliffs are subjected 

to waves and spray of sea water and have but little protection from the strong winds (fig. 6). The cliffs are 

partly covered with ice from time to time during the 

winter months.

a. Eurhynchium-Puccilennia-Caloplaca community 

(3C1).

Very little, or no, soil present. Occurs on beach 

boulders, rock pinnacles, and on cliff faces to the 

cliff summits (fig. 7). The characteristic species of 

this community have great tolerance to salt water, 

and the moss and the grass are nitrophilous in that 

they grow very luxuriant where manured by birds.

Characteristic species: Eurhynchium praelongum, PuGcinellia langeana subsp. alaskana, and 

Caloplaca granulosa.

Other species: Blastenia sp., Bryoerythrophyllum recurvirostrum, BueUia punctata, Draba hy-
14 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

-. vS«***»je " ' i>«*«a.-ii-j»«fcJC££S

FIGTJBE 6. Sea-cliff habitat on the Pacific Ocean coast at Makarius Bay. Dense growths of marine algae cover the rocks of 

the intertidal zone. The Elymus-Ligusticum-Anemone community covers the upper slopes and summits of the cliffs. Photographed June 17,1967.

perborea. Lecanora aleutica, L. straminea, Physcia 

caesia, PotentiUa villosa, Schistidium maritimum, 

Ulota phyllantha, Verrucaria maura, Xanthoria 

candelaria, and X. elegans.

b. PotentUla-Draba-/Saxifraga community (2G2).

Soil: Lithosol, in rock crevices; type I2 ; pH, « 

7.0. Occurs on the vertical cliff faces and rock pinnacles from about 10 feet above the beach, or sea 

level, to the tops of the cliffs. The vascular plants 

are rooted in rock crevices, and the bryophytes 

most commonly occur under shallow shelves of the 

cliff faces (figs. 6,7).

Characteristic species: PotentiUa villosa, Drdba 

hyperborea, and Saxifraga bracteata.

Other species: Amphidium lappomcum, Barfiida cylindrica, Bartramia ithyphylla, Bryum lapponicum, B. stentotrichum, Cardamine umbellata, 

Cephaloziella arctica, Claytonia sibirica. Cochlea/rid 

officmalis subsp. oblongifolw, Gratoneuron filicinum, Cystopteris fragUis subsp. fragilis, Eurhynchium praelongum, Lophocolea cuspidata, Luzvla 

tundricola, Marchantia polymorpha^ Mielichhoferia maorocarpa, Montia fontana subsp. fontana, 

Platydictya jtmgermannioides, Plantago maritima 

subsp. juncoides, Pohlia cruda^ Pottia heimii, 

Rhacomitrivm fasciculare, Sagina crassicaulis, 

Schistidium maritimum, Taraxacum trigonolofium, TortuZa nmcrontfolw, and Ulota phyllaritha.

c. XanthoricL-Ramolina community (2H2).

Soil: Lithosol, or rock surfaces; type I2. Occurs 

principally near the summits of high cliffs and on 

"sea stacks" offshore. Xanthoria appears to require 

fertilization by birds, and Ramalina grows only 

where there is free air movement; both requirements commonly are met on cliff summits (fig. 7).

Characteristic species: Xanthorea candelarw, 

Ramalina scoparia, and R. almquistii.

Other species: Amblystegiwn serpens, Cardamine umbellata, Caloplaca granulosa, Cystopteris 

fragilis subsp. fragttis, Eurhynchium praelongvm, 

Ochrolechia frigida, Parmelia saxatilis, P. sulFIGUBE 7. Rock pinnacles ("sea stacks"), Makarius Bay. 

Xanthorea-Ramalina community is on and near the summit, 

the Potentilla-Dralta-Samfraga community is near the middle, 

and the Eiirhynchium-Puccinellia-Caloplaca community is 

near the base of the higher pinnacle. Grass clumps (foreground) are about 6 inches high. Photographed June 17, 1967.
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 15

cata, Potentilla villosa, Physcia caesia, P. dubia, 

JSchistidiwn maritimum, and Ulota phyllanttia.

d. Elymus-Ligusticum-Anemone community (1L21).

Soil: Lithosol, and tundra without permafrost; 

type F4 ; pH, 6.4-6.6. On sloping faces of the less 

steep cliffs, and at the brink of vertical cliffs (fig. 

6). The soil generally is highly organic but at 

places may be sandy or rocky. This community is 

dominant also on large flat-topped sea stacks 

around the island, where Elymus commonly is 

more abundant than either Anemone or Ligusticwm.

Chacteristic species: Elymus arenariiis subsp. 

mollis var. mollis, Ligusticum scoticum subsp. Jvultenii, and Anemone narcissiflora subsp. villosissima.

Other species: Agrostis alaskana, Amblystegiwm 

serpens, AtKyrium filiw-femina subsp. cyclosorwm, 

Brywn stenotrichum, Calamagrostis nutkaensis, 

Conioselinum chinense, Dactylorhiza aristata,

EurhyncMwn praelongwn, Festuca rubra, Hierochloe odorata, Leptoforyum pyriforme, Ochrolechia 

frigida, Plantago maritima subsp. juncoides, Ranimculus occidentalis subsp. in&idaris, Salix arctica 

subsp. crassijulis, and Viola langsdorffi.

4. EMPETRUM HEATH HABITAT

The undulated surfaces of the old elevated marine 

platforms that occur on the southeastern third of the 

island are covered with a heath composed largely of 

Empetrwm, with sedges, grasses, lichens, and mosses occurring in various proportions. This habitat occurs also 

in low-lying valleys in the mountainous part of the 

island, and near Aleut Point, at altitudes ranging from 

about 100 to 600 feet above sea level. The plant communities at places are very complex in composition and 

structure, commonly having broad ecotones and many 

interruptions (fig. 8).

FIGURE 8. Aerial photograph of the Empetrum heath in an area of many lakes, with the Empetr urn-Car ex-lichen community predominating. A wet sedge-meadow community surrounds the two small lakes in left center. At least 14 well-developed moss 

mounds are shown. The trail of a tracked military vehicle is clearly visible in the upper left, although it was made at least 

20 years ago. The area shown here isi on the low plateau about midway between Rifle Range Point and Crown Reefer Point. 

Large moss mounds are about 25 feet in diameter. Photographed October 20, 1965, by the U.S. Air Force.
16 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

a. Empetrum-Carex-\\ohsx\. community (lC12c).

Soil: Tundra without permafrost; type O; pH, 

5.9-6.0. This community is characteristic of the 

summits and slopes of the low ridges and commonly 

is hummocky and interdigitated with ecotonal 

variants and interrupted by pools and lakes. At the 

upper altitudinal limit, the continuity of the community is broken by areas of virtually bare pebbly 

ground. In the more hummocky areas, dense stands 

of Cladoma pacifica occupy the depressions that 

surround the hummocks composed of Empetrum, 

Carex, and Sphaerophorus (fig. 9). At other places, 

Empetrum, several species of pleurocarpous 

mosses, and lichens form a dense mat of vegetation 

that is fairly smooth and has few surface depressions. Herbaceous flowering plants and Lycopodium are scattered throughout this community, 

but they rarely form dense stands. Commonly, the 

substrate is a fibrous peat, ranging in thickness 

from a few centimeters to at least 6 dm, which 

overlies fragmented andesite and breccia bedrock, 

or, at places, deposits that resemble till, colluvium, 

or old beach gravel. A thin (2-6 cm) layer of 

bluish-gray to grayish-brown mineral soil generally occurs on these rock substrates. The organic 

mat and the underlying peat and muck are nearly 

saturated with water throughout the year, even 

during the infrequent days in summer when the 

surface vegetation is dry and crisp.

FIGURE 9. Hummocky Empetrum-Carex-lichen community in 

the vicinity of Cyril Cove. The light snow cover emphasizes 

the irregularity of the surface. Photographed December 2 

1965.

Characteristic species: Empetrum nigrum subsp. 

nigrum, Carex pluriflora, C. macrochaeta, Cladonia 

pacifica, and Sphaerophorus globosus.

Other species: Alectoria ochroleuca, Anemone 

narcissiflora subsp. villosissima, Antitrichia curtipendula, Arnica unalaschcensis, Aulacomnium 

palustre, Brachythecium acutum, B. asperrimum, 

Calamagrostis nutkaensis, Calypogeia sphagnicola, 

Campanula lasiocarpa subsp. lasiocarpa, Cardamine umbellata, 'Carex anthoxanthea, C. dioica 

subsp. gynocrates, C. lyngbyaei, C. stylosa, Cassiope lycopodioides, Chrysanthemum arcticum 

subsp. arcticum, Cladonia l>ellidiflora, C. coccifera, 

C. gracilis, Coptis trifolia, Comicularia divergens, 

Cornus suecica, Dicranum elongatum, D. groenlandicum, D. howellii, Elymus arenarius subsp. 

mollis var. mollis, Equisetum arvense, Erigeron 

peregrinus subsp. peregrinus, Eurhynchium pnccelongum, Festuca l>rachyphylla, F. rubra, F. rubra 

subsp. aucta, Galium trifidum subsp. columbianum, 

Geum calthifolium, Hylocomium splendens, 

Hypnum dieckii, Linnaea borealis subsp. borealis, 

Listera cordata var. nephrophylla, Lobaria linita, 

Loiseleuria procumbens, Lophozia cf. groenlandica, Luzula multiflora subsp. multiflora var. 

frigida, Lycopodium annotinum subsp. annotinum, 

L. clavatum subsp. clavatum* L. selago subsp. 

selago, Microlepidozia mdkinoana, Mnium glabrescens, Mycoltlastus alpinus, Mylia taylori, Ochrolechia frigida, Orthocaulis binsteadii, Pedicularis 

chamissonis, Peltigera canina, P. malacea, Phleum 

commutatum var. americanum* Plagwthecium undulatum, Platanthera chorisiana* P. convallariaefolia, P. dilatata, Pleurozium schreberi, Pogonatum alpinum, Polygonum viviparum, Primula 

cuneifolia subsp. saxifragifolia, Ptilium cristacastrensis, Pyrola asarifolia var. purpurea, 

Rhacomitrium lanuginosum, Rhytidiadelphus 

loreus, R. triquetrus, Rubus arcticus subsp. stellatus, R. chamaemorus, Salix arctica subsp. crassijulis, S. cyclophylla, Saxifraga punctata subsp. insularis, Sphagnum capillaceum, S. subsecundum, 

Sphenolobus saccatulus, Thamnolia vermicularis, 

Tofieldia coccinea, Trientalis europaea subsp. arctica, Vaccinium vitis-idaea, Vahlodea atropurpurea subsp. latifolia, Veronica stelleri, and Viola 

langsdorffii.

b. Cladonia-Carese meadow community (1O22).

Soil: Tundra without permafrost; type O; pH, 

5.1-6.5. This community generally occurs adjacent 

to the Empetrum-Caresc-lichen. community, and the 

two communities merge in a narrow to wide eco-
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 17

tone. The chief distinctions of the Cladonia-Careso 

meadow community from the adjacent community 

are its location on the more level areas or lower 

parts of gentle slopes, the less prominent role of 

Empetrum, and greater abundance of Carese lyngbyaei. In general, forbs are more plentiful, and 

the vegetation mat is wetter in this community than 

in the Empetrum-Carese-lichen community (fig. 10,

11).

Characteristic species: Cladonia pacifica, Careso 

lyngbyaei, and C. macrochaeta.

FIGURE 10. Well-developed moss mound in a Cladonia-Carese 

meadow near Cyril Cove. Mound is about 5 feet high. Photographed June 10,1967.

FIGURE 11. Ecotone of an Empetrum-Carex-llchen community 

(irregular patches of snow-covered vegetation) and a CladoniarCarex meadow, near Cyril Cove. Contrast between the 

two communities was enhanced by a light snowfall, the 

snow having melted on the wetter, Cladonia-Carex meadow. 

Strip of snow-covered vegetation (center foreground) is about 

18 inches wide. Photographed December 7,1965.

Other species: Brachythecium cisperrimum, Calamagrostis nutkaensis, Calypogeia trichomanis, 

Cardamine wribellata, Carew antTuxwmthea, Ceratodon purpweus, Claytonia sibirica, Deschampsia 

beringensis, Diplophyllum albicans, Drepanocladus revolvens, Empetrum nigrum subsp. nigrwm, 

Eriophorwn russeolum subsp. rufescens, Festuca 

brachyphylla, Gymffiocolea, inflata, Jimcus trigtumis subsp. albescens, Lophozia cf. groenlandica, 

Lycopodium ann^tinum subsp. annatinum, Mylia 

taylori, Oncophorus wahlenbergii, Orthocaulis 

binsteadii, Petasites frigidus, Pinguicida vulgaris 

subsp. macroceras, Platanthera convallariaefolia, 

P. dilatata, Pogonatwn alpinum, Scapania irrigua, 

S. pcdudosa,) Trichophorum caespitosum, and Viola 

langsdorffii.

c. Moss mound community (1O1).

Conical mounds covered with vegetation are 

characteristic of many of the central and western 

Aleutian Islands. These mounds were first reported 

by Hulten (1933, p. 17-18) ; he considered them to 

be bird-perch mounds caused by the deposition of 

bird manure. This explanation of their origin was 

accepted by the following subsequent investigators: 

Hutchison (1937); Walker (1945); Bank (1951), 

1956); and Fraser and Barnett (1959). These 

mounds were described in more detail in a later 

publication by Hulten (1960, p. 29-30).

There are two basic types of conical mounds on 

Amchitka Island moss mounds (fig. 10) and rock 

mounds (fig. 33). If either of these mounds is used 

extensively by birds for perches, the heavy manuring by the birds causes a change in the vegetation 

of the mound. Many of the original plant species are 

killed by the manure, and coprophilous species 

that can thrive in the enriched substrate occupy the 

mound. Bird-perch mounds are characterized by a 

dense turf of grass at the summit (figs. 12, 32).

On the low plateaus the thousands of moss 

mounds are the most conspicuous features of the 

landscape (figs. 8, 10). The mounds range in size 

from single polsters of moss 15 to 20 cm high to conical plant communities as much as 2 m high and 10 

m in diameter. They are organic throughout and 

originate from single polsters of vertical-growing 

mosses. The manuring by birds plays no part in 

their early development. The plant community on 

the mounds is largely composed of the same species 

that grow on the adjacent heath. The substrate, 15 

cm below the mound apex, ranges in pH from 5.1 

to 6.5.
18 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

A well-developed moss mound has a zonal arrangement of plant species that surrounds the apex 

because certain combinations of species follow in 

succession up the sides of the mound. The invading 

plants eventually suppress the vertical-growing 

mosses at the apex, thus destroying the impetus toward development of the mound, and the mound 

finally begins to break apart. The zonal arrangement of species in a well-developed moss mound is 

given below.

Apex of mound: One of the following species 

ordinarily forms the dominant vegetation Rhacomitrium lanuginosum, Dicrcmum elongatum, D. 

groenlandicum, D. howellii, D. fuscescens, and Oncophorus wcMenbergii. Intermixed with these 

mosses are a few strands of leafy liverworts, including Anastrophyllum sphenoloboides, Bazzania tricrenata, Calypogeia sphagnicola, Cephalozia bicuspidata, C. leucantha, Microlepidozia maJcwuxma, 

Od&ntoschisma elongatum, Orthocaulis binsteadii, 

Riccardia latifrons, and Sphenolobus minutus. Usually, a spreading mat of Antitrichia curtipendula 

eventually grows over the vertical-growing moss of 

the apex. This moss gives the summit of the mound 

a conspicuous yellow color and tends to restrict 

further vertical development of the mound.

Sides of mound: Calamagrostis nutkaensis, Carex 

macrochaeta, C. pluriflora, Cladonia amaurocraea, 

C. arbuscula subsp. beringiana, C. bellidiflora, C. 

pacifica, C. scabrmscula, Coptis trifolia, Comicularia divergens, Cornus suecica, Drepanocladus uncinatus, Eurhynchium praelongum, Gymnocolea 

inflata, Hylocomium splendens, Loiseleuria procumbens, Mycoblastus alpinus, Mylia taylori, Parmelia omphalodes, Peltigera aphthosa, Pleurozium 

schreberi, Pogonatum alpinum, Polytrichum commune, P. strictum, Ptilidium ciliare, Rhytidiadelphus loreus, R. triquetrus, Rubus arcticus subsp. 

stellatus, R. chamaemorus, Scdix arctica subsp. 

crassijulis, Sphaerophorus globosus, Tofieldia coccinea, Vaccinium vitis-ida&a subsp. minus, and 

Vahlodea atropurpurea subsp. latifolia.

Base of mound: Empetrum nigrum subsp. 

nigrum, Calamagrostis nutkaensis, and Carex 

lyngbyaei.

d. Organic bird-perch mound community (1M2).

Substrate pH, 6.4-6.7. The mounds that support 

this community (fig. 12) are in ihv Empetrum heath 

habitat and are organic throughout, in contrast to 

the bird-perch mounds formed over boulders and 

the conical projections of bedrock that occur at 

higher altitudes on Amchitka Island. The vegetation of the two types of mounds is composed of virFIGURE 12. Organic bird-perch mound showing characteristic 

columnar summit composed of Poo, arctica subsp. williamsii, 

near White House Cove. The grass on the sides of the mound 

is mostly Calcrniagrostis nutkaensis, but a few plants of Elymus arenarius subsp. mollis var. mollis are present. Columnar 

summit is about 8 inches in diameter. Photographed June 11, 

1967.

tually the same species, if the mounds are equally 

manured. Organic bird-perch mounds are located 

mostly along the coasts, but because the island is 

narrow, sea birds at some inland locations have 

caused these mounds to develop. These mounds 

originate as moss mounds that are described in 4c; 

when the mounds reach a certain height (probably 

about 6 dm) birds may begin to use them as lookout perches. Bird manure favors the growth of 

coprophilous plants and, at the same time, kills the 

noncoprophilous species that formerly covered the 

apex and sides of the moss mound. The verticalgrowing mosses and the characteristic Antitrichia 

at the summit are replaced by 'a dense cylindrical 

turf composed of a grass species. On mounds where 

Dicranum elongatum forms the apex, however, the 

moss summit may persist for an unknown length of 

time, but it becomes abnormally green.

In time, the influence of the manure extends 

down the aides of the mound, killing most species 

of mosses and lichens; these plants are replaced 

by Calamagrostis and several species of forbs. 

Eventually, the sides of the mound collapse, leaving a cylinder of grass turf 6 to 9 dm high protruding from an irregular, hummocky base (fig. 12). 

On neighboring Rat Island, foxes were observed to 

have caused virtually the same transformation of 

moss mounds, but the fox-mound lookouts do not 

have cylindrical summits.

Listed below are the species of plants found on 

these mounds.

Apex of mounds: Festuca brachyphylla, Poaarctica subs. wiUiamsiip Puccinellia langeana
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 19

subsp. alasTcana, Rinodina turfacea, Cladonia 

alpestris, Eurhynchium praelongum, and Dicranum elongatum.

Sides of mound: AntitricTiia curtipendula, Brachythecium alhicans, C alamagrostis nutTcaensis, 

Cladonia scabriuscula, Diplophyllum albicans, 

Drepanocladus uncinatus, Elymus arenarius subsp. 

mollis var. mollis, Eurhynchium praelongum, 

Heracleum lanatum, Macrodiplophyllum plicatum, 

Peltigera canina, Ranunculus occidentalis subsp. 

insularis, Rhacomitrium lanuginosum, Rinodina 

turfacea, Sagina crassicaulis, Sphaerophorus globosus, Ulota phyllantha, Xanthoria candelaria, and 

some species listed in 4c that have not yet succumbed to the effects of the manure.

Base of mound: C alamagrostis nutTcaensis, 

Caresc lyngbyaei, Empetrum nigrum subsp. nigrum, IcmadophUa ericetorum, Poly podium vulgare subsp. occidentals, and Psoroma hypnorum.

e. Streamside oonimunity (1N21).

Soil: Tundra without permafrost and alluvial; 

type O; pH, 5.0-6.5. The narrow, deeply entrenched 

streams of gentle slopes and valleys support a distinctive plant community that is especially conspicuous in June, when the bright green of the 

streamside plants contrasts strongly with the brown 

tones of the adjacent Cladonia-Carese meadows. 

The vegetation is composed principally of tall forbs 

and sedges. An understory of lesser forbs, bryophytes, and lichens that are adapted to reduced 

light grows on the steep stream banks (fig. 13).

Characteristic species: Athyrium filix-femina 

var. cyclosorum, Carex lyngbyaei, Heracleum lanatum, Geum macrophyttum subsp. macrophyllum, 

and Dryopteris dttatata subsp. americana.

Other species: Bracliythecium asperrimum, 

Bryum pollens, B. pseudotriquetrum, Calliergon 

sarmentosum, Calypogeia trichomanis, Campylium 

stellatum, Cardamine umbellata, Cephalozia fiicuspidata, C. media, CTiiloscyplius pallescens, Claopodium crispifolium, Climacium dendroides, Conioselium chinense, ConocepJialum conicum, Cystopteris fragilis subsp. fragilis, Dicranum howellii, 

Diplophyllum albicans, Eurhynchium praelongum, 

Fritillaria camschatcensis, Lecanora castanea, Luzula parviflora subsp. parviflora, Marsupella emarginata, Mnium glabrescens, M. pseudopunctatum, 

Nardia scalaris, Peltigera aphthosa, P. canina, 

Petasites frigidus, Platydictya jungermannioides, 

Ranunculus occidental-is subsp. insularis, Riccardia 

pinguis, Saxifraga punctata subsp. insularis, Scapania paktdosa, 8. undulata, Sphagnum girgensohnii, S. teres, and Streptopus amplexifolius.

FIGUBE 13. iStreamside community near Cyril Cove. The stream 

has been dammed by the aquatic moss FontinaMs; the small 

pool above the dam is about 45 cm deep. Streptopus amplexifolius is just beginning to grow on the top of the dam. The 

vegetation on the streambanks includes Carex lyngbyaci, Heracleum lanatum, Ranuculus occidentalis subsp. insularis, and 

Athyrium filix-femina subsp. cyclosorum. Photographed 

June 14, 1967.

f. Peat-bank community (lOl).

Soil: Tundra without permafrost; type O; pH, 

5.4-6.1. Numerous peat banks were formed on this 

island by excavations related to military operations 

during World War II. The bare banks, commonly 

1 to 2 m high, were opened to primary invasion by 

plants that were adapated to a saturated organic 

substrate, strong winds, and full exposure to light. 

Apparently, only bryophytes and a few lichens are 

early invaders, for after 20 years vascular plants 

have become established in only a few locations. 

Areas of seepage on the vertical or overhanging 

surfaces may support conical or curtainlike growths 

of bryophytes (principally Cephalozia bicuspidata) at the drip points (fig. 14). All the naturally 

formed peat banks observed had passed the primary stages of colonization, and their plant communities resembled those of the Empetrum heath. 

The flowering plant Majanthemum dilatatum, 

however, appears to grow only on old well-vegetated peat banks, never on the more nearly level 

areas of the heath.

Characteristic species: Pogonatum alpinum, 

Polytrichum commune, Cephalozia bicuspidata, 

and Dicranella heteromalla.

Other species: Agrostis alaskana, Anomobryum 

concinnatum, Brachythecium albicans, Bryum arc350-274 O -
20 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

FIGURE 14. Peat-bank community in the Empetrum heath near 

Constantine Harbor. The elongated bryophyte colonies, composed principally of Ccphalozia Mcmpidata, form at drip 

points on the saturated bank that is about 2.5 m high. The 

colony of large mosses, shown near the upper left, is composed of Polytrichum sphaerothecium. Photographed June 5, 

1967.

ticum, Blasia pusilla, Bryoerythrophyllum recurvirostrum, Calypogeia mriilleriana, Cephalozia ambigua, G. pleniceps, Cephaloziella arctica, Cladonia 

degenerans, G. gracilis, Dichodontium pellueidum, 

Dicranetta heteromalla, D. subulata, Diplophyllum 

albicans, Lophozia alpestris, L. cf. groenlandica, 

L. indsa, Majanthemum dilatatum, Mnium glabrescens, M. insigne, Nardia scalaris, Orthocaulis 

floerkei, Pannaria pezizoides, Plagiothecium cf. 

undulatum, Plectocolea cf. obovata^ Pogonatum 

contortum, P. umigerum, Pohlia annotina, P. cf. 

cruda, P. nutans, Polytrichum formosum, P. 

sphaerothecium, Rhacomitrium fasciculare, Scapania scandica, Sphagnum, compactum, S. papillosum, and Toninia Idbulata.

g. Frost-scar community (3C2).

Soil: Tundra without permafrost, or lithosol; 

type ?2; pH 6.0-6.3 Frost scars as much as 1 m or 

more in diameter occur throughout the Empetrum 

heath from which the organic mat has been removed, and mineral soil is exposed. Frost heaving 

is active during the winter and is intensified by 

frequent cycles of freezing and thawing of the soil 

(fig. 15). This unstable substrate of soil, gravel, 

and cobbles, or at places angular bedrock fragments, provides a suitable niche for species unable 

to compete with the vegetation of the unbroken 

heath.

Characteristic species: Antennaria dioica, Cardam-ine umbellata, Deschampsia beringensis, and Oncophorus woMenbergii.

Other species: Cephaloziella arctica, Chiloscyphus pallescens, Dicranella palustris, Dicranum 

fuscescens, Diplophyllum albicans, Epilobium 

glandulosum, Geum calthifolium, lonaspis epulotica, Lecidea cf. vernalis, Leptobryum cf. pyriforme, 

Luzula multiflora subsp. multiflora var. frigida, 

Microlepidozia mdkinoana, Mnium insigne, M. 

pseudopunctatum, Nardia scalaris, Philonotis 

fontana^ Rhytidiadelphus loreus, Riccardia sinuata, 

Rubus arcticus subsp. stellatus, Scapania paludosa, 

/Sibbaldia procumbens, Stellaria calycantha subsp. 

calycantha, 8. sitchana var. bongardiana, and Veronica serpyllifolia subsp. humifusa.

FIGURE 15. Frost scar in the Empetrum heath near Cyril Cove. 

The light snow cover had fallen before the soil had frozen; 

subsequent frost heaving is indicated by the cracks (black 

lines) in the snow. Field notebook gives indication of scale. 

Photographed November 29,1965.

5. BOG HABITAT

Most bogs on this island are soligenous and occur on 

gentle to moderately steep slopes. The borders of lakes 

and pools do not support bogs, probably because the 

strong winds agitate the water too violently for Sphagnum, and other bog plants to grow; therefore, there are 

no floating bogs on the island. At a few locations the 

bogs have an unstable surface and may be termed "quaking bogs." Bogs are more common and of larger size on 

the low plateaus of the eastern part of the island than 

in the mountains, where they are ordinarily restricted 

to small areas in drainage ways.

a. /Sphagnum, bog community (1O11).

Soil: Bog; type O; pH, 6.1-6.2. Water pH, 

« 5.9. These bogs are neither abundant nor large, 

and at places they intergrade with wet sedge
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 21

meadows. Isolated colonies of Sphagnum, commonly S. capillaceum and /S. subsecundum, occur in 

small water-filled depressions in the Empetrum 

heath. Typically, this community is best developed 

in seepage channels or broader areas of gentle slope 

(fig. 16), where stands composed largely of several 

species of Sphagnum limit the invasion of other 

plants.

FIGURE 16. Spring issuing from the margin of a large soligenous 

bog in a broad valley near Cyril Cove. A wet sedge-meadow 

community is in the foreground, and a Sphagnum bog community is in the center and background of the picture. At 

the time this photograph was made, water temperature in the 

spring was 3.3°C5 and under the ice of nearby pools, 0.7°C; 

air temperature was -2.3 °C. Photographed February 10, 1968.

Characteristic species: Sphagnum teres, IS. squarros-um, 8. magellanicum, IS. compactum, S. papillosum, and /S. girgensohnii.

Other species: Aulacomnium palustre, Brachythecium asperrimum, Calypogeia sphagnicola, 

Careso anthoxanthea, C. lyngbyaei, C. pluriflora, 

Dicranum angustum, Drepanocladus revolvens, 

Eriophorum russeolum subsp. rufescens, Mnium 

pseudopunctatum, Philonotis americana, Pinguicula vulgaris subsp. macroceras, Plantago maritime subsp. juncoides, Riccardia latifrons, 

Selaginella selaginoides, and Siphula ceratites.

b. Philonotis-Parnassia community (1O12).

Soil: Bog; type P2 ; pH, 6.5-6.8. Water pH, 6.3- 

7.0. Requirement for the development of this community appears to be a saturated inorganic substrate of very gentle slope through which water 

moves. The water is less acid than that of 

Sphagnum bogs. The largest observed community 

of this type is on sandy soil below the large spring

near Makarius Bay5 where an almost unbroken mat 

of Philonotis covers an area of a hectare or more.

Characteristic species: Philonotis americana and 

Parnassia kotzebuei.

Other species: Amblystegium serpens, Carex kelloggii, C. lyngbyaei, Cerastium fontanum subsp. 

triviale, Dicranella heteromalla, D. subulata, Epilobium sertulatum, Juncus triglumis subsp. albescens, 

Jungermannia sphaerocarpa, Limosella aquatica, 

Lophozia cf. groenlandica, Lupinus nootkatensis, 

Platanthera convallariaefolia, P. dttatata, Pogonatum umigerum, Pottia heimii, Saxifraga punctata subsp. insularis, and Veronica serpyllifolia 

subsp. humifusa. 

c. Scapania-Nardia-Marsupella community (1O1).

Soil: Bog; type P,; pH, 6.1-6.8. Although small 

colonies of each of the characteristic species occur 

in wet areas of the Empetrum heath, these three liverworts commonly form a distinctive community 

that occupies drainage channels and wet areas only 

at altitudes above 400 feet. These species also grow 

in the snow-bed community, but there they have 

different vascular-plant associates. The brownish 

red to bright red of the Scapania and Marsupella 

and the bright green of the Nardia make the community conspicuous, even at a distance. The saturated organic mat formed by this community is as 

much as 20 cm thick.

Characteristic species: Scapania paludosa, Nardia scalaris, and Marsupella emarginata.

Other species: Anthelia julacea, Aulacomnium 

palustre, Brachythecium asperrimum, Dicranella 

palustris, Drepanocladus revolvens, Lycopodium 

annotinum subsp. annotinum, L. sabinaefolium 

var. sitchense, Oncophorus ^vahlenberg^^ Philonotis fontana, and Saxifraga punctata subsp. 

insularis.

6. MARSH HABITAT

A marsh on this island can be distinguished from a 

bog during any season of the year by its abundance of 

sedges and f orbs, whereas a bog has a preponderance 

of bryophytes. If the substrate of a marsh is stirred 

up, even in winter, it gives off a strong odor of hydrogen 

sulfide. The substrate of a bog however, is virtually 

odorless. Probably chemical compounds in the bryophytes of bogs inhibit the production of hydrogen 

sulfide by bacteria.

a. Wet sedge-meadow community (1M21).

Soil: Bog and Half Bog; type O; pH, 5.9-6.2. 

Water pH, «=* 6.3. This is the most abundant community in the marsh habitat and occurs typically 

at the borders of lakes (figs. 8, 17), but it also
22 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

FIGURE 17. Wet sedge-meadow community at the margin of a 

large lake near Cyril Cove. The projecting mats of Carex 

are as much as 2 m from base to tip and 30-60 cm wide 

but are being undercut and are thus destroyed by wave action. 

A line of new sedge growth is in the center of the picture. 

Numerous projections of sedge mats encircle some large lakes 

and give a fimbriate appearance to the shoreline, as seen in 

aerial photographs. Photographed June 13, 1967.

develops on very wet slopes and beside streams in 

th&Empetru/mhesith (fig. 18).

Characteristic species: Carex lyng~byaei, O. pluriflora, C. anthoxanthea, C. nwcrochaeta, Eriophorum russeolum subsp. rufescens, and Juncus 

triglumis subsp. albescens.

Other species: Aongstroemia longipes, Aulacomniwn palustre, Claytonia sibirica, Drepanocladus fluitans, D. revolvens, Epilobium glandulosmn, Festuca rubra, Galium trifidvm, subsp. 

columbianum, Geum macrophyllum subsp. macrophyllwn, Lophozia alpestris, Mnium glabrescens, 

Nardia scalaris, Pedicularis chamissonis, Pellia 

neesiana, Philonotis americana, Plantago mcucrocarpa, Pleurozium schreberi, Polytrichum longiFIGURE 18. Wet sedge meadow near Cyril Cove. This area has 

a dense stand of Car ex lyng~byaei forming a wide border by 

the stream. A Fontinalis-Ranunculus community is in the 

stream. Photographed November 23,1965.

setwn, Rcmunculus occidentalis subsp. nelsoni, 

Rhinanthus minor subsp. borealis, Rhytidiadelphus loreus. Sphagnum girgensohnii, S. papillosum, /S. squarrosum, S. teres, and Viola langsdorffii.

b. Caltha-Claytonia community (1N21).

Soil: Half Bog; type P2 ; pH, 6.1-6.9. Water pH, 

6.2-6.9. This community occurs most commonly on 

a wet inorganic substrate that has sufficient movement of water through it to prevent freezing in 

winter. An area about 2 hectares in extent is 

occupied by this community near Constantine 

Harbor, and many smaller areas support this community where water upwells through the Empetrwm 

heath mat.

Characteristic species: CcHiha, palustris subsp. 

asarifolia and Claytonia sibirica.

Other species: Careoc lyngbyaei, Epilofiium 

glandulosvm, E. sertulatum, Petasites frigidus, 

RammculMs occidentalis subsp. insularis, and 

Rumex obtusifolius.

7. LAKE HABITAT

In this report, lakes are distinguished from pools 

by their being sufficiently large for wind to strongly 

agitate the surface and waves to pound the shore, thus 

influencing the kind of community that can develop 

in and around them (fig. 17). The ice on some large 

wind-swept lakes affects the formation of the lake banks.
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 23

If a sudden lowering of the air temperature to a point 

below freezing occurs when gale-velocity winds are 

blowing, water is lifted from the lake surface, frozen in 

midair, and dropped on the leeward side of the lake. 

A covering of rough ice then develops from the leeward 

to the windward side of the lake until the entire lake 

surface finally is frozen, with thicker ice on the leeward 

side.The thick ice appears to exert pressure on the 

vegetation mat of the bank such that this mat (largely 

unfrozen) is pushed out and up to form the elevated 

rims characteristic of many lakes (fig. 19). Moreover, 

the suspended inorganic materials in the windblown 

water are deposited on the banks. This process, especially active in the more shallow lakes, tends to deepen 

the lakes and to elevate the lake banks.

Hamelin and Cook (1967, p. 99) evoked a different 

process in explaining the formation of lake ramparts: 

"A rise in temperature causes the ice to expand, induces 

considerable stresses and crowds the edge of the shore, 

the material [gravel and cobbles, in their illustration] 

is pushed, overriding the low shore, for many feet 

where it is deposited upon contraction or melting of the 

ice." This process may also be active on Amchitka 

Island.

Two kinds of lakes are on the low plateaus of the 

island those that have a geologic origin, are relatively 

deep, and have a bedrock or cobble bottom, and those 

that develop on the peneplaned area of the heath 

habitat, are shallow, and have a muck or soil bottom 

(Powers and others, 1960, p. 526). With but few 

exceptions, only the latter type of lake contains vascular

.*-4£f.f- 

> 

^

" jfe *. '^. j .««: 

;f *V "V

%;#) t^, i

*.

FIGURE 19. Ice ridge on the leeward bank of a large lake near 

Cyril Cove. The water near the distant shore is not frozen. 

Strong winds lift this water, which is frozen in midair and 

then dropped on the leeward bank and the lake surface. 

Top of the tundra bank is about 6 feet above water level. 

Photographed December 3,1965.

plants, which are emergent or immersed bottom-rooted 

aquatics.

a. Hippuris-Ranunculus community (1P21).

Soil: Type P2 ; pH, 5.8-6.3. Water pH, 5.1-6.7. The 

Hippuris grows in water as much as 1 m deep; in 

summer the upper part of the plant emerges from 

the water. Dense stands of this plant occur in some 

lakes. Commonly, the Ranunculus is intertwined 

with the Hippuris.

Characteristic species: Hippuris vulgaris and Rwnunculus trichophyUus var. trichophyllus.

Other species: None, although a band of sedges 

(principally Carex lyngbyaei} may surround the 

lake and extend into the water.

b. Isoetes-Ranunculus-LimoseUa community (1P11).

Soil: Type P2 ; pH, 6.4-6.6. Water pH,« 6.6. The 

plants of this community commonly grow in lake 

water 45 to 60 cm deep. Isoetes grows completely 

submerged in the deeper water, but the other characteristic species grow in shallow water near the 

shore and may be emergent. All are tenaciously 

rooted in mud of the lake bottom and grow throughout the year.

Characteristic species: Isoetes muricata subsp. maritima, Ranunculus reptans, and Limosella aquatica.

Other species: Commonly none; however, at one lake 

Scapania subalpina and Sphagnum squarrosum 

grew as submerged aquatics. At some lakes Ranunculus hyperboreus subsp. hyperboreus grows on 

mud along the shore.

8. FOOL HABITAT

Pools usually are formed in depressions of the vegetation and peat mantle on the Empetrum heath, although some are in depressions of bedrock or colluvium, 

and some are formed by bryophyte dams (fig. 13). Because the pools have a periodic fluctuation of water 

level, emergent aquatic plants most commonly grow in 

them and in the mud along the pool banks.

Small pools appear to become enlarged by the 

process of "swamping," and when they reach a diameter 

of about 3 m (if in an exposed location) wind becomes 

active in "digging" them deeper. The agitated water 

holds much silt or humus from the pool bottom in 

suspension, and the wind blows this water from the pool 

to the adjacent heath. Deposits of silt on the leeward 

side of pools was observed to be sufficiently thick to 

obscure the green color of the Empetrum mat at a distance of 3 m from the pool.
24 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

a. Juncus-Eriophorum community (1M22).

Soil: Tundra without permafrost and Alluvial; 

type P2 ; pH, 5.0-6.4. Water pH, 4.8-6.3. This 

community commonly occurs in shallow pools on 

the Empetrum heath (fig. 20) and less commonly 

in roadside pools and very wet areas of stream 

alluvium. The vegetation along the pool shores is 

included here because it is generally inundated 

after heavy rains or snow melt.

Characteristic species: Juncus triglumis subsp. 

aTbescens and Eriophorum rus s eolum subsp. 

rufescens.

Other species: Alopecurus aequalis, Aulacomnium palustre, Campylium stellatum, Oarex kelloggii, Chiloscyphus pdttescens, Deschampsia 

caespitosa subsp. orientalis, Diplophyllum albicans, Drepanocladus revolvens, Juncus ensifolius, 

Marchantia polymorpha, MarsupeUa emarginata, 

Mnium gldbrescens, Nardia scalaris, Parnassia 

kotzebuei, Pinguicula vulgaris subsp. macroceras, 

Pogonatum alpinum, Ranunculus tricTiophyllus 

var. trichophyllus, Rhacomitrium aciculare, Riccardia pinguis, /Scapania paludosa, &. undulata, 

Selaginella selaginoides, /Sparyanium hyperfioreum. Veronica americana, and V. serpyllifolia 

subsp. humifusa.

b. /Subularia-Gallitriche community (1P1).

Soil: Lithosol; type P2 ; pH, ^ 6.6. Water pH, 

6.9-7.2. This community occurs in shallow seasonal 

pools that have bedrock bottoms. The two characteristic plants can grow as submersed or emergent aquatics or can persist on damp soil if the 

pools dry up (fig. 21).

FIGURE 20. Small pool supporting a Junctts-Eriophorum community, surrounded by a Cladonia-Carex meadow, near Cyril 

Cove. Sedges in the pool extend about 1 foot above water surface. Photographed June 17, 1967.

FIGURE 21. Bottom of a dry pool near Constantine Harbor 

bearing a dense growth of Subularia aquatica that was 

flowering abundantly. Callitriche anceps grows in similar 

shallow rock-bottomed pools. Utility pole (background) provides scale. Photographed June 17, 1967.

Characteristic species: /Subularia aquatica and 

Callitriche anoeps.

Other species: Alopecurus aequalis, Bryum sp., 

Carex kelloggii, Cladonia bettidiflora, and Ranunculus reptans.

c. Siphula-Scapania community (1O21).

Soil: Tundra without permafrost and Lithosol; 

type P2 ; pH 6.0-6.4. Water pH, 6.3-6.6. Common 

in small (60-120 cm in diameter) pools on the Em>- 

petrum heath, but most extensive in shallow bedrock 

pools of larger size. /Scapania forms a, dense dark 

mat on the pool bottom, generally in 5 to 15 cm of 

water, and Siphula grows as an emergent aquatic 

for much of the year (fig. 22). Both plants can 

thrive, however, if the pool dries up. Siphula is the 

most hydric lichen on the island and is the only one 

observed to grow completely submersed.

Characteristics species: Siphula ceratites and 

Scapania paludosa.

Other species: Anthelia julacea, Cladonia 

amaurocraea, Isoetes sp., Lophozia incisa, and 

Sphagnum compactum.

9. STREAM HABITAT

Most streams are narrow, swift, and short; and the 

water, generally clear and colorless, has percolated 

through the vegetation mat. The streams flow throughout the year at a, fairly steady rate, even under an ice 

covering, except for periods of greatly increased flow
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 25

FIGURE 22. Vertical view of a Siphula-Scapania community in 

a shallow rock-bottomed pool near Crown Reefer Point. Dark 

colonies of Scapania can be seen on pebbles in the water and 

at the bases of the Siphula thalli. White thalli are about 2 

inches long. Photographed February 18, 1968.

following heavy rainfall and snow melt. They commonly are deeply entrenched in the vegetation and peat 

mantle (fig. 23) and have a bedrock, gravel, or colluvium 

bottom. At places the vegetation mat has closed over the 

stream trenches and f ormed "bridges" that may not support the weight of a man. The trenches are periodically

FIGURE 23. Deeply entrenched stream in the Empetrum heath 

near Chitka Point. The narrow stream flows on a bedrock bottom between nearly vertical banks, 1 to 2 m high, of peat and 

vegetation. Photographed November 22, 1965.

filled with snowdrifts in winter and are a hazard to foot 

travelers. The streams appear to have been entrenched 

by the growth and deposition of vegetation at their margins ; thus they differ from "haggs" of the British moors, 

defined by Pearsall (1950, p. 249) as erosion channels in 

peat. Streams draining broad valleys generally have alluvial fans at their mouths that are bordered by sedge 

marshes (fig. 18), whereas those draining the more 

nearly level plateaus descend over the sea cliffs in a 

series of rapids.

a. Fontinalis-Ranunculus community (1P11).

Soil: Type I2 ; pH, 6.5-6.9. Water pH, 6.8-7.2. 

Occurs commonly in rapid, entrenched streams 

where the light intensity is low. Fontinalis grows 

attached to bedrock or stones in water as much as 

1 m deep and develops fronds as much as 60 cm 

long. Brachythecium and Hygrohypnum also grow 

attached to rocks, but they form dense mats in shallow water, especially in cascades. These mosses are 

shown forming a dam in figure 13. The Ranunculus 

is rooted in soil and produces underwater flowers 

and fruits throughout the winter.

Characteristic species: Fontinalis neomexicana 

and Ranunculus trichophyllus var. trichophyllus.

Other species: Brachythecium asperrmwm, Bryoerythrophyllum recurvirostnmi^ Chiloscyphus 

pallescens, Hygrohypnum bestii, H. ochraceum, 

Scapania irrigua, and S. undulata. Montia fontana 

subsp. fontana grows in the shallow water of cascades but not in deeply entrenched streams. Shallow 

parts of the stream margins, if not shaded by high 

banks, support the emergent aquatics Mimulus guttatus, Veronica americana, and Carex lyngbyaei.

10. RUDEEAL HABITAT

Soils of this habitat are classified only by type. Many 

substrates have the surface qualities of a lithosol or 

regosol, and some substrates are not soil. The extensive 

disturbance of the ground surfaces by military operations and by recent construction on the island has opened 

many new areas for colonization by plants that are 

adapted to a mineral soil and that can withstand the 

effects of microclimatic extremes. The species of the 

ruderal communities are, with very few exceptions, native plants that also grow in other habitats on the island 

but that commonly grow more vigorously in the areas 

which have been disturbed by man. For example, the 

largest plants of Lupinus nootkatensis on the island 

grow along roadsides, although smaller plants occur in 

many habitats. Loiseleuria procumbens commonly is a 

depauperate plant in the Empetrum-Carex-lichen community ; yet a plant that grew in the center of an unused
26 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

gravel road had, in about 20 years, formed a dense prostrate shrub 30 cm in diameter.

The communities of this habitat described below are 

not so clearly .defined and discrete at all places as are 

some of the more "natural" communities on the island 

probably because many of the component species of the 

ruderal communities have a wide ecological amplitude. 

There is, nevertheless, a tendency toward grouping of 

certain species in niches that may be only slightly separated spatially.

a. Calamagrostis community (1M21).

Soil: Type I2 ; pH, «5.9. In the construction of 

roads during World War II, the organic mantle and 

some of the mineral soil below it were pushed aside 

to form banks 1/2-'L m high that paralleled the roads. 

These banks are covered with an almost unbroken 

turf of tall grass intermixed with very few forbs. 

After about 20 years of dominance by the grass, 

there is now some invasion on the banks of the low 

plateaus by Empetrum, and at a few places the 

banks are somewhat similar in appearance to the 

Empetrum-Caresc-lichen community. At altitudes 

of about 400 to 600 feet, wind has severely eroded 

the soil banks so that only fragments of the banks 

remain. The fragments are held in place only so 

long 'as the turf is sufficiently strong to resist wind 

erosion (fig. 24) .

Characteristic species: Calamagrostis nutkaensis.

Other species: Antennaria dioica, Bryum 

stenotrichum, Caresc macrochaeta, Cephalozia bicuspidata, Ceratodon purpureus, Dicranella subulata, Ditrichum heteromallum, Gentiana amarella 

subsp. acuta var. plebeja, Heracleum lanatum.

FIGURE 24 Fragments of a roadside bank severely eroded by 

wind. This remnant of a bank, held in place by a dense turf 

of Calamagrostis nutkaensis, is near Sea Otter Point. 

Bank, including turf, is about 4 feet high. Photographed 

November 22, 1965.

Hylocomium splendens, Lusula multiflora subsp. 

multi-flora var. frigida, L. parmflora subsp. parviflora, Nardia scalaris, Pogonatum umigerum, 

PoMia nutans, Primula cuneifolia subsp. saxifragiand Rhacomitrium ericoides.

b. Lupinus- Arnica community (3C2).

Soil: Type I2 ; pH, 6.7-7.0. The community is 

common along many roadsides and on disturbed 

areas; it is very showy in midsummer because of 

the large size of the plants and the profusion of 

flowers on the two characteristic species (fig. 25).

FIGTJEE 25. Lupinus nootJcatensis and Deschampsia caespitosa 

subsp. orientalis growing on the graveled shoulder of an unused road near Constantine Harbor. Lupine flower spikes are 

about 8 inches high. Photographed June 15, 1967.

These plants grow in the loose gravel and mineral 

soil of road shoulders, relatively dry ditches, and 

similar sites. Other species occupy the wetter ditches 

and lower parts of roadside banks. The substrate 

is not greatly compacted and is subjected to strong 

frost heaving in winter. Heavy rains severely erode 

the surfaces; therefore, bryophytes and lichens are 

rare, but large colonies of the moss Philonotis occur 

at some places in the wet roadside ditches.

Characteristic species : Lupinus nootkatensis and 

Arnica unalaschcensis.

Other species : Achillea borealis, Anaphalis margaritacea, Angelica lucida, Antennaria dioica^ 

Cardamine umbellata, Cerastium fontanum subsp. 

triviale, Deschampsia caespitosa subsp. orientalis^ 

Epilobium sertulatum, Gentiana aleutica^ Marchantia polymorpha, Mnium glabrescens, Dactylorhiza aristata, Philonotis americana, Polygonum 

viviparum, and Potentilla egedii subsp. grandis.
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 27

c. Equisetum-Sagina-Ceratodon community (2G2).

Soil: Type I2 ; pH, 5.8-6.5. The characteristic 

substrate of this community has a hard, very compact, often stony surface that resists frost heaving. 

This community occurs on unused well-compacted 

roads, graveled parking lots and service areas, fractured concrete slabs, and cracked macadam runways. Some species are rooted in fine-gravel accumulations 2 cm or less thick or unbroken concrete 

or macadam surfaces. Many old gravel-road surfaces have an almost unbroken growth of Equisetum. Sagina occidentalis forms dense rosettes 

rooted in pavement cracks or very compact road 

centers. Rhinanthus, Luzula, Hieracium, and Lycopodium occur most commonly on well-packed 

stony animal trails.

Characteristic species: Equisetum arvense, 

Sagina occidentalis, and Ceratodon purpureus.

Other species: Antennaria dioica, Braehythecium acutum, B. albicans, B. asperrimum, Bryum 

stenotrichum, Cladonia pyseidata, Drepanocladus 

uncinatus, Empetrum nigrum subsp. nigrum, Eurhynchium praelongum, Hieracium gratiile var. 

alaskanum, Lecidea macrocarpa, Loiseleuria procumbens, Luzula multi-flora subsp. kobayasii, L. 

parviflora subsp. parviflora, Lycopodium selago 

subsp. selago, Nardia scalaris, Ochrolechia frigida, 

Placopsis gelida, Platanthera chorisiana, Poa alpina, P. turneri, Polytrichum formosum, Rhacomitrium ericoides, Rhinanthus minor subsp. borealis, 

Sagina intermedia, Saxifraga punctata subsp. 

insularis, Sibbaldia procumbens, Stellaria calycantha subsp. calycantha, Stereocaulon intermedium, S. tomentosum var. alpestre, and Veronica 

serpyllifolia subsp. humifusa.

11. DISCONTINUOUS HEATH HABITAT

A distinct change in habitats occurs at an altitude of 

about 400 feet, where the extensive Empetrum heath 

of the low plateaus becomes fragmented into small 

strips and patches that alternate with almost bare 

ground (fig. 26). This disruption of the heath mat 

probably is caused mostly by wind erosion, but frost 

heaving doubtless accelerates the process, once the vegetation cover is broken. Hulten (1960, p. 40-41) wrote, 

"It is quite clear that the mosaic of plant communities 

in the Aleutians is to a large extent regulated by the 

wind." He noted a very sharp change in vegetation at 

levels where the full force of the wind was felt, which 

he reported to be the boundary that separates the 

meadows from the heath. He referred to this boundary 

as the "wind plane." On the low plateaus of Amchitka

FIGURE 26. Discontinuous heath habitat, with scattered 

patches of the Saliso-Empetrum community. Large mound 

(center) is a densely vegetated bedrock outcrop. The area is 

between Sea Otter Point and Chitka Point at an altitude of 

about 400 feet. Bedrock outcrop is about 15 feet wide. Photographed November 22,1965.

Island, the boundary between these two communities 

is determined more by the degree of drainage than by 

wind action. The heath at many places extends almost 

to the lowest points of sheltered valleys, if drainage is 

adequate. Conversely, throughout the highest levels of 

the low plateaus, sedge meadows occur everywhere that 

drainage is poor. The streamside community and the 

alpine meadow community are the ones that most 

clearly require shelter from wind.

The striking boundary on Amchitka Island that is related to wind is the ecotone between the Empetrum 

heath and the alpine region; it is about 400 to 600 feet 

above sea level. At this altitude, wind erosion probably 

is more active than frost action in determining which 

plant communities survive (fig. 24).

The strips of vegetation ordinarily are eroded on the 

windward side until the roots and subterranean stems 

of the dwarf shrubs are exposed and finally die, but on 

the leeward side, new plant growth slowly extends the 

mat outward. Two lichen species, Ochrolechia frigida 

and Mycoblastus alpinus, are associated with the dead 

margins of the strips; however, it was not determined 

whether the lichen growth causes, or is the result of, 

death of the vegetation at the edge of the strips. The 

vegetation mats, therefore, appear to "move," in time, 

with the wind, but this supposition has not been proved 

by actual measurements. The concept of patches of vegetation "moving" in this manner was described by Watt 

(1947) and by Pearsall (1950, p. 83) as observed in 

British moorlands, and the floristic composition and 

successional changes in these mats were given by Metcalfe(1950,p.53).

350-274 O - 69 - 5
28 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

There is no one direction of prevailing winds on 

Amchitka Island as a whole. In the mountains and high 

plateaus, topography exerts a controlling effect on prevailing wind directions at specific locations. On slopes, 

the strongest winds commonly are upslope, regardless of 

compass direction, and the vegetation mats tend to 

erode on the lower side and advance on the upper side. 

On more nearly level areas the local topographic features appear to determine the direction of this "movement." More careful observations than have yet been 

made and measurements throughout the year are necessary if these processes are to be defined more accurately.

The substrate between and under the vegetation 

patches is composed of volcanic rock fragments in a 

matrix of fine mineral "soil" and is subjected to moderate frost heaving, but patterned ground generally is not 

formed. This substrate between vegetation patches is 

largely devoid of vascular plants, although some scattered bryophyte colonies occur. Bedrock outcrops in this 

area are discussed separately in this report.

a. Salisc-Empetrum community (lC21a).

Soil: Tundra without permafrost and Lithosol; 

type F3 ; pH, 6.0-6.1. The fabric of the vegetation 

patches is composed largely of interwoven stems 

and branches of Salim and Empetrum, which hold 

in place a thin organic soil that supports a few 

herbs, lichens, and mosses. Some of the woody 

stems of Salix arctica subsp. crassijulis are almost 

1 cm in diameter and appear to be many years old. 

The shrubs form roots throughout the length of 

their buried stems; therefore, they may continue to 

grow at the stem tips while dying at the basal end 

of the stems.

In the more (sheltered locations of the discontinous heath, areas between the heath patches develop 

a conspicuous growth of lichens. Species of Cladonia grow on the summits and sides of the small 

hummocks, and dense stands of Thamnolia are 

present in the depressed areas between the 

hummocks.

. Characteristic species: Salix arctica subsp. crassijulis, 8. rotundifolia, and Empetrum nigrum subsp. 

nigrum.

Other species: Alectoria cf. irvingii, A. ochroleuca, Campanula lasiocarpa subsp. lasiocarpa, Cassiope lycopodioides, Cerastium fischerianum, Cladonia alpestris, C. pacifica, Dicranum howellii, 

Epilobium behringianum, Gymnamitrion corallioides, Hieracium triste, Loiseleuria procumbens, 

Luzula arcuata subsp. unalascheensis, L. tundricola, Lycopodium alpinum, L. selago subsp. selago, 

Mycoblastus alpinus, Ochrolechm frigida, Peltigera aphthosa, Polygonum viviparum, Rhacomitrium fasciculare, R. lanuginosum, Sphaerophorus 

globosus, and Thamnolia vermicularis.

b. Andreaea-Schistidium-Rhacomitrium community 

(3C1).

Soil: Lithosol; type I; pH, 5.8-6.0. Although 

the area between the heath mats that is covered 

with rock fragments and soil may appear to be 

barren, small moss polsters grow on many rocks 

and in the mineral soil (fig. 26). Andreaea generally is attached to small rocks, but at places it 

grows on the soil, and the polsters are undercut 

by wind erosion (Shacklette, 1966, p. 349). 

Schistidium and Rhacomitrium ordinarily grow on 

rock fragments, and frost heaving and thawing 

only naise and lower the rocks without disrupting 

the moss polsters.

Characteristic species: Andreaea rupestris, Schistidium apocarpum, and Rhacomitrium, lanuginosum.

Other species: A few colonies of Brachythecium 

albicans, Conostomum tetragonum, and Drepanocladus uncinatus and rosettes of Draba aleutica and 

Cerastium aleuticum, which characteristically occur on solifluction terraces, are widely scattered 

through this community.

12. SOLIFLUCTION HABITAT

At altitudes between 600 and 1,100 feet, the effects of 

strong frost action are conspicuous in all but the most 

sheltered locations (figs. 27, 28). The land surface is 

composed largely of scree slopes and colluvial boulder 

fields in which the surface rocks are arranged in terraces, rock stripes, <and weakly developed boulder polygons. Solifluction terraces and rock stripes occur in 

pebbly ground and are presently active in reshaping the 

ground surface, but observations of the weak boulder 

polygons were inconclusive in determining whether the 

process of rearrangement was currently active, or 

whether the polygons are relics of frost iaction of former 

times. High-altitude rock outcrops and sheltered valleys 

are discussed separately in this report.

a. Fell-field community (3B1).

Soil: Lithosol; type I; pH, 6.0-7.0. This community is most extensive on the high plateaus of 

the western part of Amchitka Island (fig. 1), but 

smaller areas occur throughout the mountains. 

Lithosol development is very slight, and many rock 

surfaces have only a partial covering of lichens 

(fig. 29).
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 29

* **.

v

-*: .* 

Ste

FIGURE 27. High-altitude land surfaces that are strongly influenced by solifluction processes, between Chitka Cove and Andesite 

Point. A light snow cover emphasizes the surface features. Slopes are covered with an almost continuous vegetation mat; 

they develop a microrelief in which the ridges and furrows parallel the direction of slope. (Compare with fig. 28.) Photographed February 7,1968.

Small patches and fragments of a heath community 

are scattered through some boulder fields; these 

are more common on the steep slopes, where the 

rare Rhododendron camtsckaticwin subsp. camtschaticwn is a component of the vegetation patches. 

On the nearly level high plateaus, Lupinus is the 

principal initiator of vegetation mats. Single plants 

become established between the boulders on the 

almost barren plateaus, and mosses and lichens grow 

in the loose debris and soil that accumulates in the 

shelter of these plants. Vegetation mats, as much 

as 10 dm in diameter, that contain a significant 

number of heath plants were observed to have developed around large and obviously very old lupine 

plants.

Characteristic species: Salioo rotundifolia, S. cyclophylla, Cassiope lycopodioides, Empetrum nigrum subsp. nigrum, and Lupinus nootkatensis.

Other species: Alectoria ochroleuca, A. pubescens, 

Andreaea rupestris, AntitricMa curtipendula, Arctoa fulvella, Bryum stenotrichum, Calypogeia

sphagnicola, Campanula lasiocarpa subsp. lasiocarpa, Carese circinnata, Cephalozia cf. bicuspidata, 

C. pleniceps, Cetraria nigricans, Conostomum tetragonwn, Diapensia lapponica subsp. obovata, Dicranum JioweUii) Diplophyllum albicans, D. taxifolium, Drepanocladus revolvens, D. uncinatus, Hypogymnia subobscura, Luzula tundricola, Ochrolechia 

frigida, Parmelia omphalodes, Pohlia nutans, Rhacowdtrium brevipes, R. ericoides, R. lamiginosum, 

Rhododendron camtschaticum subsp. camtschaticum, Salix arctica subsp. crassijuZis, Scapania 

scandica, Sibbaldia procumbens, /Sphaerophorus 

globosus, Vmbilicaria proboscidea, Vaccinium 

uliginosum subsp. mwropTiyllum, and Veronica 

grandiflora.

b. Solifluction terrace community (3C13).

Soil: Lithosol; type Ii; pH, 6.6-6.7. The'ground 

surface occupied by this community is composed of 

pebbles, generally not more than 3 cm in diameter, 

in a matrix of very loose wet mineral soil. Frost
VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

FIGURE 28. Rock stripes that developed on a moderate slope 

between Ohitka Cove and Andesite Point. A light drifted snow 

cover emphasizes the surface features. Distance between the 

summits of adjacent ridges is about 30 cm. Ridges are perpendicular to the direction of slope. (Compare with fig. 27.) 

Photographed February 7,1968.

FIGURE 29. Fell-field at the summit of a mountain near Ohitka 

Cove. Very little soil forms at this site; vegetation is sparse, 

and strong winds remove most of the fine rock particles. 

Photographed February 7, 1968.

action has arranged these materials into rock stripes 

(fig. 28) and terraces (fig. 30) that are perpendicular to the direction of slope. The brinks of the low 

terraces may have a weakly developed strip of heath 

plants, but the pebbly ground of rock stripes supports only single plants or polsters spaced widely 

apart. Lichens can scarcely become established on 

the most unstable surfaces, and only one moss, 

Conostomum tetragonwn, is well adapted to this 

substrate.

FIGURE 30. Solifluction terraces on the high plateau near Rim 

Point. The terraces support depauperate heath mats, and the 

stony ground between the terraces bears a scattered cover of 

Lusula tundricola and a few other plants. An animal trail 

leads from the foreground to the helicopter. Photographed 

June 11, 1967.

Characteristic species: Potentilla hyparctica, 

Koenigia islandica, Cerastium aleuticum, Draba 

aleutica, Juncus biglumis, and Conostomum 

tetragonum.

Other species: Andreaea rupestris, Antitrichia 

curtipendula, Brachythecium aTbicans, Ca/rdamine 

bellidifolia, Cassiope lycopodioides, Cladonia alpestris, Claytonia arctica, Dicranum howellii, 

Diapensia lapponica subsp. obovata, Diplophylhtm albicans, Drepanocladus uncinatus, Herberta 

adunca, Hypnum plicatulm, Jungermannia atromrenSj Lu&ula tundricola, Marsupella emarginata, 

Poa lanata var. vivipara, Ranunculus occidentalis 

subsp. insularis, Rhacomitrium fasciculare, R. lanuginosum, Sagina intermedia, Salix cyclophylla, 

Saxifraga punctata subsp. insularis, /S. foliolosa 

var. foliolosa^ Sphagnum compactum, Stellaria 

ruscifolia subsp. aleutica, Stereocaulon vesuvianum, and Tritomaria quinquedentata.

13. ALPINE MEADOW HABITAT

In the narrow valleys in the mountains, wind velocity 

is greatly reduced, and in some places snow accumulations may persist until mid-June. The valleys afford 

protection from strong frost action, desiccating winds, 

and severely cold weather and thereby permit growth 

of species that are not found elsewhere on the island. 

The wettest parts, of the valleys, where snow melt water 

saturates the ground, and the margins of small streams 

that drain the slopes and valleys (fig. 31) support snowbed communities. The more mesic valley walls, slopes,
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 31

FIGURE 31. Abrupt transition from fell-field (lower right) to 

discontinuous heath at Chitka Cove. The streams (foreground) are bordered with snow-bed communities; a large 

wet sedge meadow surrounds the lake; and a moss mound 

marks the narrow Empetrum heath above the sea cliffs. 

Photographed August 16,1966.

and streamside terraces bear a profusion of showy 

flowering plants, ferns, and mosses that characterize 

the alpine meadows. Lichens are not abundant in this 

habitat. The ecotone between the alpine meadow and 

the solifluction terraces, or boulder fields, is at places 

very narrow; for example, the distance (measured up 

the steep valley sides) from the stream bed, through 

the alpine meadow, to the almost bare terraces and 

rock fields may be as little as 10 m.

a. Snow-bed community (1O12).

Soil: Type P2 ; pH, 5.8-6.2. Water pH, 5.4-6.6. 

The substrate of this community is saturated with 

water, and large patches of the gray liverwort 

Anthelia make the community conspicuous. This 

liverwort is characteristic of snow-bed communities in many parts of the northern hemisphere. The 

yellow-flowered Sataifraga hirculus grows in the 

liverwort mats with the diminutive Hippuris montana and the rare shrub Geum pentapetalum. Colonies of Sphagnum and other hydric bryophytes 

complete the mosaic of the snow-bed community, 

which extends along some of the small streams that 

flow through the discontinuous heath habitat (fig. 

31). Many species listed below were found only in 

this community.

Characteristic species: Anthelia julacea, Scapania paludosa, Saxifraga hirculus, and Leptarrhena pyrolifolia.

Other species: Andreaea rupestris, Carex bigelowii, C. physocarpa, Cephalozia ambigua, C. bicuspidata, Dicranella palustris, Diplophyllum 

albicans, Drepanocladus revolvens, Geum pentapetalum, Gymnocolea inflata, Hippuris montana, 

Lophocolea cuspidata, Lophozia alpestris, Marsupella emarginata, Mylia anomala, Nardia compressa, N. scalaris, Oligotrichum hercynicum, O. 

parallelum, Osoyria digyna, Pellia neesiana, Primula cuneifolia subsp. saxifragifolia, Radula polyclada, Rhytidiadelphus sguarrosus, Riccardia pinguis, Sparganium hyperboreum, Sphagnum compactum, S. papillosum, S. squarrosum, Takakia 

ceratophylla, and Veronica serpyllifolia subsp. 

humifusa.

b. Alpine meadow community (1JST21).

Soil: Tundra without permafrost or Half Bog; 

type F5 ; pH, 6.2-6.4. Although all vascular plants 

of this community (except Pyrola minor} were also 

found in the Empetrum heath, the distinctive features of this community are in the predominance of 

large-flowered forbs, the sparsity of Empetrum 

and other shrubs, and the lesser importance of 

lichens, sedges, grasses, and bryophytes in the 

structure of the community. This community is 

limited to the higher altitudes, generally above 600 

feet, in well-sheltered valleys; at lower altitudes 

the Empetrum, heath predominates in the valleys.

Characteristic species: Achillea borealis, Arnica 

unalaschcensis, Chrysanthemum arcticum subsp. 

arcticum, Claytonia sibirica, Geum calthifolium, 

and Ranunculus occidentalis.

Other species: Carex pluriflora, Cephalozia ambigua, C. bicuspidata, Cetraria ericetorum, Cladonia rangiferina, Cornus suecica, Drepanocladus 

revolvens, D. uncinatus, Empetrum nigrum subsp. 

nigrum, Erigeron peregrinus, Listera cordata var. 

nephrophylla, Lophozia alpestris, L. opacifolia, 

Lupinus nootkatensis, Nardia geoscyphus, N. scalaris, Plagiothecium, undulatum, Plantago maritima 

subsp. juncoides, Platanthera convallariaefolia, P. 

dilatata, Pleurozium schreberi, Polygonum vimparum, Pyrola minor, Rhytidiadelphus loreus, 

Sphaerophorus globosus, Sphagnum lindbergii, 

S. magellanicum, Thelypteris limbosperma, T. 

phegopteris, and Vahlodea atropurpurea subsp. 

latifolia.

14. INLAND BEDROCK HABITAT

Bedrock outcrops at inland locations throughout 

the island support plant communities different from 

those of the sea cliffs. The inland outcrops are remnants 

of an earlier landform and are resistant to erosion. At 

places they project only slightly above the general 

ground level; elsewhere they form irregular masses or 

symmetrical cones that are conspicuous landmarks 

(figs. 32, 33). The plant communities of the outcrops 

are low growing and do not completely cover the rock
32 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

FIGURE 32. Low-altitude rock outcrop that is used as a bird 

perch, near Banjo Point. The thick turf at the summit is 

composed of Festuca firachyphylla. Photographed August 17, 

1966.

FIGURE 33. High-altitude bedrock outcrop near Chitka Cove. 

The fell-field (foreground) has patches of heath plants on 

the unstable rocky substrate. Although much of this conical 

outcrop is covered with talus, the loose rocks are not subjected to strong frost action because of the underlying bedrock. Outcrop is about 40 feet wide at the base. Photographed 

August 16,1966.

surfaces, unless fertilized by birds. The species composition of the communities appears to be controlled 

largely by factors associated with altitude; accordingly, two communities are recognized.

a. Low-altitude bedrock communities (1N21).

Soil: Weakly developed Lithosol; type I2 ; pH, 

6.0-6.6. Sites of these communities range in altitude from about 150 to 600 feet above sea level; outcrops at lower altitudes are on the seacoasts. Some 

conical outcrops are used by birds as perches and 

are recognizable by the thick turf of grass at the 

summits and sides of the mounds (fig. 32).

Characteristic species: Veronica stelleri, Cassiope lycopodioides, Tofieldia coccmea, and Salix 

rotv/ndifolia.

Other species: Agrostis borealis, Alectoria ochroleuca, A. pubescens, Andreaea rupestris, Angelica 

lucida, Antennaria dioica, Antitrichia curtipendula, Bacidea sp., Campamda chamissonis, C. 

lasiocarpa subsp. lasiocarpa, Oaresa macrochaeta, 

Oeratodon purpureug, Cetraria cucullata, Chrtfsanthemu/m arcticum subsp. arcticum, Oornicularia 

divergens, Dicranum kowellii, Drepanocladus uncinatus, Empetrum nigrum subsp. nigrwn, Festuca 

bracJiypJiylla^ Gentiana aleutica, Heracleum lanatum, lonaspis epulotica var. arctica, Letidea 

aleutica, L. macrocarpa, Loiseleuria procumbens, 

Lophozia cf. wenzelii, Luzula arcuata subsp. unalaschcensis, Lycopodiwn sabinaefolium var. 

sitchense, Lycopodium selago subsp. selago, Mnium 

glabrescens, Nardia scalaris, Ochrolechia frigida, 

Pamnelia saxatilis, P. sulcata, Peltigera aphthosa, 

Pertusaria coriacea, Placopsis gelida, Platanthera 

chorisiana, Poa arctica subsp. williamsii, Pogonatum urnigerwn, Polytrichum junipervrwm, 

Primula cu/neifolia subsp. saxifragifolia, Ramalina 

almguistii, Rhacomitrium fasciculare^ R. lanuginosum, Rhizocarpon Tiochstetteri, JSalios arctica subsp. 

crassijulis, S. cyclopJiylla, Thamnolia vermicularis, 

Umbilicaria cylindrica, U. proboscidea. Viola 

langsdorffii, and XantJioria candelaria.

b. High-altitude bedrock community (301).

Soil: Weakly developed Lithosol; type I; pH, 

6.4^6.6. This community contains few vascular 

plants but many cryptogams; a large part of the 

rock surfaces, however, is not vegetated (fig. 34). 

Mosses and lichens can colonize the relatively stable 

rocks of outcrops, whereas the rocks of adjacent

FIGURE 34. Rock outcrops on a mountain summit about 1 mile 

west of Chitka Cove. Strong winds and almost constant fog 

characterize the summits. Vegetation consists of one species 

of Carex, one of Affrostis, and several species of lichens and 

mosses. Photographed July 29,1967, by J. A. Erdman.
DESCRIPTION OF THE ISLAND AND ITS VEGETATION 33

solifluction terraces and fell-fields are so unstable, 

due to frost action, that fewer of these plants can 

grow on them. The conical outcrops at high altitudes (fig. 33) are used very little as perches by 

birds. At high altitudes, sea birds and ptarmigans 

do not congregate, and raptorial and passerine birds 

are not abundant; therefore, coprophilous species 

of plants are not common on high-altitude outcrops.

Characteristic species: Carex drcinnata, Umbilicaria proboscidea, and Agrostis borealis.

Other species: Alectoria cf. irvingii, A. nigricans, 

Andreaea rupestris, Anthelia julacea, Cephdlo&ia 

leuoantha^ Cerastium aleuticum, Cetraria ericetorum, Cladonia bettidiflora, Conostomum tetragonum, Cornicularia divergens, Diplophyllum 

albicans, Empetrum nigrum subsp. nigrwn, Festuca brachyphylla, Gymnomitrion corallioides, 

Haematomma ventosum subsp. lapponicum, Hypnum cupressiforme, Lecidea armeniaca,, L. elegantwr, L. flavocaerulescens, L. macrocarpa, Lophozia 

cf. groenlandica, L. incisa, Luzula tundricola, Mycoblastus alpinus, Ochrolechia frigida, Orthocaulis 

binsteadii, Parmelia alpicola, P. omphalodes, Pertusaria hultenii, Platysmatia lacwwsa, Pogonatuan 

alpinum, Pohlia nutans, Rhacomitrium fasciculare, 

R. lanuginosum, Rhizocarpon atroalbescens, R. 

geographicwn, Sphaerophorus globosus, SphenoloIrus minutus, Thelypteris limbosperma, and 

Umbilicaria proboscidea.

15. MISCELLANEOUS HABITATS, SUBSTRATES, AND 

COMMUNITIES

a. Carcasses.

Nitrophilous mosses of the family Splachnaceae 

most commonly grow on dung of herbivorous mammals. These animals do not occur on Amchitka 

Island, however, and the only splachnaceous moss 

that is present, Tetraplodon mnioides, grows on decomposed carcasses of birds and rats. This moss can 

be found on any part of the island (except possibly 

at high altitudes) where carcasses happen to lie. 

Tetraplodon colonies are conspicuous because of the 

density and reddish-brown color of the sporophytes 

and the abundance of greenish-yellow spores that 

are produced. Byrum stenotrichum was found 

growing on a rat carcass, and Pleuro&nt/m schreberi, 

on a bird carcass, but these mosses do not commonly 

grow on this type of substrate. No other plants were 

found on carcasses.

b. Imported wood.

No native trees grow in the Aleutian Islands; 

therefore, all wood on the islands was brought in by

man or by ocean currents. Military and other operations on Amchitka Island have resulted in the importation of much wood for buildings, utility poles, 

and other uses. The plants which grow on this 

wood are as follows: Baeomyces placophyllus, 

Brachyihecium acutmn,, Ceratodon pwpureus, 

Cetraria ciliaris, Drepanocladus wtcinatus, 

Eurhynchium praelongum, Hypogynvrda enteromorpha, H. pkysodes, Lecanora attophana, Lecidea 

cf. symmicta, L. vernaLis (on tar paper over a 

wooden roof), Mnium glabrescens, M. pseudopunctatum, Parmelia saxatilis, P. sidcata, Parmeliopsis ambigiM, and Pertusaria sp.

c. Plantings.

Several Picea sitchensis trees planted in front of 

buildings during World War II are still living, are 

about 1 m high, and had about 3 cm of new growth 

in 1967 (fig. 35). Similar plantings of this species 

were observed to have been made on Shemya Island, and four of the trees were still alive in 1967. 

On Amchitka Island the window boxes of a military building were planted with Anemone narcissiflora subsp. villosissima, and these plants are still 

living. No other evidence of plant cultivation was 

seen on this island.

d. Burned areas.

An area of Empetrum heath approximately 1 

square kilometer in extent was burned during 

World War II. Apparently, only the aerial parts

FIGURE 35. Sitka spruce (Picea sitchensis) planted in front of 

a club building during World War II. Tree is about 3 feet 

high. Photographed February 19,1968.
34 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

of plants were burned, whereas the saturated substrate protected the crowns and roots. After 20 years 

the burned area still contrasts strongly to adjacent 

unburned areas because of the abundance of Calamagrostis nutkaensis and other grasses that revegetated the area after the burn. The grass disclimax 

that develops after a heath fire was described by 

Pearsall (1950, p. 145, 155). Lichens, including 

Cladonia cocci/era, C. degenerans, C. gracilis, C. 

pacifica, C, pyxidata, and Empetrum nigrum subsp. 

nigrum are beginning to revegetate the burned area. 

Probably many more years will be required for this 

area to recover completely from the fire damage.

PLANT SPECIES OF AMCHITKA ISLAND 

ANNOTATED LIST OF VASCULAR PLANTS

By HANSFOKD T. SHACKLETTE, JAMES A. EKDMAN, and JOHN R.

KEITH

Duplicates of specimens cited in this list have been 

identified or their names verified, with but few exceptions, by Dr. Eric Hulten (written commun., 1967), and 

the nomenclature follows that of his new flora (1968). 

Species from Amchitka Island that were reported by 

Hulten (1960) but were not found during this study 

are cited, but without habitat notes. Collection numbers, 

in italics, are those of Shacklette unless otherwise specified; they follow the species name. The habitat and 

plant community from which a collection was taken, 

designated by numbers and letters as set forth in the 

section of this report on the description of the island 

and its vegetation, follow the collection number. One 

asterisk before the species name indicates that this is a 

report of the first collection of this species on Amchitka 

Island; two asterisks indicate a first collection of the 

species in the Aleutian Islands.

BORAGINACEAE

Mertensia maritima subsp. maritime Hult. 8196, la. This 

plant occurs only on sandy beaches and was found at 

only a few locations on the island.

CALLITRICHACEAE

*Callitriche anceps Fern. 7631, 8b. Grows in shallow roadside pools; rare. 

Callitriche hermaphroditica L. (Hulten, 1960, p. 257).

CAMPANULACEAE

Campanula chanvissonis Federov. 75^9, 14a; 7550, 14a. 

Found only at altitudes above 300 feet. The common form 

has violet corollas, but at places colonies of many plants 

with lavender to pink corollas occur. Flowers in August.

Campanula, lasiocarpa subsp. lasiocarpa Hult. 7397, 4a; 

7653, 14a; Erdman 580, 12a. Most abundant in the Empetrum heath, but it occurs also throughout the mountains 

except on the highest peaks. Flowers in August.

CAPRIFOLIACEAE

Linnaea borealis subsp. toorealis Hult. 7298, 4a; 7500, 4a; 

Erdman 564, 4a. Occurs commonly on banks in a mat of 

Empetrum and mosses; flowers in mid-August.

CARYOPHYLLACEAE

Cerastium aleuticum Hult. 7573, 12b; 8226, 14b; Erdman

563, lib. Found only on unstable soil at high altitudes:

flowers in mid-August. 

Cerastium ^eeringianum var. teeringianum Hult. (Hulten.

1960, p. 178). 

Cerastium fischerianum Ser. 7496, la; 7995, lla. This plant

has slender light-red stems that are very brittle.

* Cerastium f&ntanum subsp. triviale (Link) Jalas. 7470,

lOb; 8040, 5b. Common on roadsides and sand dunes. 

SoncJcenya peploides subsp. major (Hook.) Hult. 7490, la.

Found only on sandy or rocky beaches; forms a zone of

dense vegetation near the high-tide level. 

Saffina crassicaulis S. Wats. 76.18, 4d; 7619, 4d. Found only

in wet crevices of sea cliffs. 

Sagina intermedia Fenzl. 7575, 12b; 8037, lOc; Erdman 578,

12b. Occurs on salifluction slopes and rocky roadsides:

forms very dense rosettes on hard soil or in pavement

cracks.

*Sagina occidentalis S. Wats. 7471, lOc; 7654, la. Found on

upper beach terraces and on old gravel roads. 

Stellaria calycantha subsp. calycantha Hult. 7443, 4g; 7633,

la ; 8034, lOc. On disturbed soil at low altitudes. 

Stellaria media (L.) Vill. (See Hulten, 1960, p. 176.)

*Stellaria ruscifolia subsp. aleutica Hult. 7580, 12b; 8223, 

12b. A rare plant of solifluction slopes. Leaves rigid, 

glaucous, and sharp pointed. Flowers very sparingly, in 

mid-August.

Stellaria sitchana var. tongarMana (Fern.) Hult. 75^6, 4g. 

On frost scars and other sites of disturbed soil in the 

Empetrum heath.

COMPOSITAE

Achillea torealis Bong. 7^39, lOb. Common on roadsides and 

on disturbed soil in the Empetrum heath.

Anaphalis margaritacea (L.) Benth. & Hook. f. 7423, lOb. 

Grows in dense colonies on compacted road shoulders. 

Probably an adventive species.

Antennaria dioica (L.) Gaertn. 7255, 14a; 7395, lOc; 7440, 

lOb; 7613, 4g; 7871, lOa. Characteristic of frost scars and 

other areas of disturbed soil, but was not found above an 

altitude of 800 feet

Antennaria pallida E. Nels. Cited by Tatewaki and Kobayashi (1934, p. 84) as A. alpina Gaertn.; Hulten (1960, 

p. 346) considered this citation to refer to A. pallida but 

did not cite a collection from Amchitka Island.

Arnica unalaschcensis Less. 7^79, lOb. Abundant on roadsides and disturbed soil of the Empetrum heath. Begins 

to flower in early August.

Chrysanthemum arcticum subsp. arcticum Hult. 7625, 4a. 

Common throughout the island except at high altitudes. 

Begins to flower in late July.

Erigeron peregrinus subsp. perearinus Hult. 7469, 4a. The 

large pink flowers are showy and resemble some cultivated 

asters. Abundant and widespread; flowering from late 

July to mid-August.

*Hieracium gracile var. alasJcanum Zahn. 7433, lOc. Not common ; found only at altitudes of no more than 200 feet.
PLANT SPECIES OF AMCHITKA ISLAND 35

Hieracium triste Willd. 7561, lla. Not common; found only 

at altitudes above 300 feet.

*Petasites frigidus (L.) Franch. 7624, 4e; 7845, 4b. Not common. Flowers the first of June, before the leaves are well 

developed.

Scnecio pseudo-arnica Less. 7503, la. Abundant on all sandy 

or pebbly beaches; at places forms dense stands of tall 

(as much as 1.3 m) succulent plants; never found inland. 

Flowers in mid-August.

Taraxacum trigonolotitm Dahlstedt. 7491, 3b. Usually grows 

in crevices of sea cliffs, but occasionally occurs on roadsides. Tatewaki and Kobayashi (1934, p. 89-90) described 

a new species, T. aleiiticum Tatewaki & Kitamura, from 

Amchitka Island, but Hulten (1960, p. 366) considered it 

to be T. trigonoloftum.

CORNACEAE

Cornus eatwdcnsis L. x succica L. (See Hulten, 1960, p.

274-275.) No specimens of Cornus that matched Hulten's

description of this hybrid were found. 

Cornus suecica L. 7404, 4a. Abundant in the moss and lichen

mats at medium to low altitudes. Flowers in mid-June to

early July and produces bright-red fruits abundantly.

CRUCIFERAE

Aphragnnts esclisclwltzianus Andrz. (See Hulten, I960, p. 

206.)

*Cardamine bellidifolia L. Erdman 567, 12b. Rare; apparently restricted to unstable, nearly bare, rocky soil at high 

altitudes.

Cardamine umbellata Greene. 7474, lOb; 7846, 4b; 7903, 3c; 

7997, 4e. Abundant in a wide variety of habitats at altitudes below 400 feet.

Cochlearia offlcinalis subsp. oftlongifolia (DC.) Hult. 7614- 

3b. Occurs typically on the thin wet soil cover of coastal 

cliffs.

*Draba aleutica Ekman. 7571, 12b. Rare; apparently restricted to solifluction slopes, where it forms dense 

rosettes.

Drafter hyperborea (L.) Desv. 7615, 3b; 8022, 3b. A characteristic species of crevices in sea cliffs. Flowers in June 

and July; the large petals are cream colored.

*Siil>ularia aquatica L. 8028, 8b. Found flowering the middle 

of June in a shallow rock-bottomed pool. Petals of the 

minute stellate corolla are white on the upper side and 

light blue on the lower side.

CYPERACEAE

Carex antlioxantliea Presl. 7465, 4b; 7657, 6a. Grows on 

ridges in the Empctrum heath and in wet sedge meadows; 

nonstooling; never forms large colonies.

*Carcx bigeloicii Torr. 7587, 13a. Apparently very rare, occurring only in snow-bed communities. Tatewaki and 

Kobayashi (1934, p. 108) reported C. rigida Good, from 

Amchitka Island; this report probably should be referred 

to C. bigelowii.

Carex circinnata C. A. Mey. 7606, 14b; 7891, 12a ;8228, 14b. 

This is one of two vascular plant species that can endure 

the most severe weather of mountain summits, and although it fruits very sparingly, at places it forms large 

colonies.

*Carex dioica subsp. gynocratcs (Wormsk.) Hult. Klein 2710, 

4a. Found on a hummock in the center of a pond.

Carex kclloggii W. Boott. 7485, 8a; 8031, 8b. Commonly occurs in wet sandy soil at low altitudes, where it forms 

dense stools.

Carex lyngbyaei Hornem. 7451, 6a; 7^67, 4a; 7473, 4b; 

7537, 4c; 7658, 6a; 8199, 5b. Occurs in many different 

habitats at low altitudes and ranges in height from about 

10 cm at unfavorable locations to nearly 2 m where g~owing in streamside alluvium.

Carex macrochaeta C. A. Mey. 7478, lOa; 7610, 14a. Widespread and abundant on the low plateaus.

Carex nesophila Holm. Questioned as occurring on Amchitka Island by Hulten (1960, p. 122-123) on the 1 <?sis 

of the report of C. macrochaeta var. subrigida Kuek. 

apiid Macoun by Tatewaki and Kobayashi (1934, p. 307).

Carex pluri flora Hult. 7536, 4c; 7656, 6a; 8221, 13b. Common in the Empetrum heath.

Carex saxatilis subsp. laxa (Trautv.) Kalela. 7586, 13a. Apparently rare; found only in a snow-bed community.

Carex stylosa C. A. Mey. 8185, 4a. Found only in the Empetrum heath on the low plateaus.

Erioplwrum russeolum subsp. rufescens (Anders.) Hyl. 

7428, 6a; 7534, 8a; 7655, 6a; 8026, 6a. The heads range 

in color from white to slightly tawny. The rhizomer are 

deeply buried in a tough layer of fibrous peat.

Eriophorum russeolum var. ma jus Sommier. Collected on 

Amchitka Island by Hutchison (1937, p. 229) who wrote, 

"A very handsome species of bog-cotton with large pale 

golden tufts." This collection was identified by Hulten 

but was not recorded from Amchitka Island in his fora. 

The species was reported from Amchitka Island by Tatewaki and Kobayashi (1934, p. 110). It was not found 

during the present study, although especially sought. 

*Triclwphorum caespitosum (L.) Hartm. 8283, 4b. Found 

only in moss and lichen mats on the lower terrace at 

Chitka Cove.

DIAPENSIACEAE

**Diapcnsia lapponica subsp. obovata (F. Schm.) Hult. 7570, 

12b; Klein 2829, 12a. This collection provides an iirwrtant range extension for the species. Found growing 

among andesite pebbles of a wet solifluction slope ard in 

a fell-field. The flowering season was past in mid-August.

EMPETRACEAE

Empetrum nigrum subsp. nigrum Hult. 7295, 4a; 7535, 4c; 

8054. 14a, Erdman 547, 4a. The most abundant and characteristic plant on the island. It flowers in early May, 

and on June 5 the green fruits were almost full size.

EQUISETACEAE

Equisetum arrense L. 7^34, lOc; 7^95, 2a; 7626, 4a; 7914, 

lOc. Abundant and widespread in many habitats, but best 

developed as colonies on sand dunes and disturbed mir^ral 

soil. The sporangia mature in early June and are produced in abundance.

ERICACEAE

Cassiopc lycopodioides (Pall.) D. Don. 7254, 14a; 7567,12a; 

7852, 12b; 8053, 14a. Common in the Empctrum heath, but 

usually depauperate. At high altitudes it is a major component of the discontinuous heath patches. Flowei*s in 

June and early July.
36 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

Loiseleuria procumbens (L.) Desv. 7264, 14a; 7867, lOc. 

Common and well developed in the Empetrum heath; 

flowers profusely in mid-June.

*R1iododendron camtschaticum subsp. camtschaticum Hult. 

8216, 12a. Apparently occurs only in small patches of 

heath near the mountain tops. Flowers in late July.

Vaccinium uliginosum subsp. micropliyllum Lange. 8217, 

I2a. Found only with Rhododendron; neither flowering 

nor fruiting.

Vaccinium vitis-idaea subsp. minus (Lodd.) Hult. 7296, 4c. 

Most commonly occurs in moss mats of the Empetrum 

heath and on moss mounds. Flowers very sparingly and 

none was observed to bear fruits. Some leaves turn brilliant red and persist through the winter.

GENTIANACEAE

*Cfentia-na amarella subsp. acuta var. plebeja ( Cham. & 

Schleoht.) Hult. 7400, lOa. Found only on a disturbed 

soil bank. The corolla is dull blue to lavender and is produced in early August.

*Gentiana aleutica Cham. & Schlecht. 7862, lOb; 7683, lOb: 

8191, 14a. Two color forms occur together one with 

white, the other with dull purple, corollas. Flowers in 

late July and early August.

GERANIACEAE

Geranium erianthum DC. (Hulten, I960, p. 256, listed as a 

sight record only.) This very conspicuous plant was 

sought during the present study without success. If it 

occurs on this island now, it is rare. It was not found on 

Rat Island, but was common on Shemya Island.

GRAMINEAE

Agrostis alaskana Hult 7515, 4f. Found only on a peat bank 

near Constantine Harbor. Considered by Klein, in his 

discussion of plant species in this report, to be A. borealis 

Hartm.

*Agrostis borealis Hartm. 8230, 14b. Found only at altitudes 

above 900 feet.

* Agrostis eorarata Trin. 7438, 2a; 7794, 2a. The culms of this 

plant spread horizontally and are closely appressed to the 

sandy deposits on which they grow.

Alopecurus aequalis Sobol. 7460, 8b; Erdnmn 555, 8a. Found 

only in shallow pools and other very wet locations.

Bromus sitehensis var. aleutensis (Trin.) Hult. 7492, 2a. 

On sand and shell deposits at South Bight.

Galamagrostis nutkaensis (Presl) Steud. 7213, 4c; 7291, 4c: 

7294, 4c; 7383, lOa; 7519, 3d; 75,38, 4c. The most common 

and widespread grass on the island. It forms almost pure 

stands on roadside banks, but is not abundant at high 

altitudes.

Descharnp&ia alpina (L.) Roeni. & Schult. Cited from Amchitka Island by Tatewaki and Kobayashi (1934, p. 97). 

Hulten (1960, p. 85), however, doubted that this species 

occurs in Alaska. i

Deschampsia beringensis Hult. 7475, 4b; 8188, 2a. Common 

in open areas of the Empetrum heath. The spikelets are 

purplish red. Considered by Klein, in the section on plant 

species in this report, to be D. caespitosa (L.) Beauv.

Deschampsia caespitosa subsp. orientalis Hult. 7412, 8a 

Occurs at the margins of shallow pools, as well as on compacted roads.

Elymus arenarius subsp. mollis var. mollis Hult. 7289, 2b; 

7520, 3d. Occurs in large stands only near the coasts.

although single plants are widespread at low altitudes. 

At some locations it forms many dense cylindrical stools, 

as much as 0.5 m high, that are separated by narrow 

passages between the stools. The development of this 

unusual type of growth pattern was observed only on 

slopes and probably is caused by soil creep. 

Festuca brachyphylla Schult. 7612, 14a; 7999, 14b. A coprophilous species on this island, found only on the summits 

of bird-perch mounds, where it forms a very dense turf. 

Considered by Klein, in his section on plant species in this 

report, to be F. ovina var. braohyphylla (Schult.) Piper.

*Festuea nibra L. 7297, 6a; 7408, 4a; 7518, 3d. Found in 

several habitats on the low plateaus.

Festuca rubra subsp. aucta (Krecz. & Bobr.) Hult. 7408, 4a ; 

7463, 4a; 8189, 2a. Vegetative plants are abundart in the 

Empetrum heath; fruiting plants are not common in the 

heath, but are abundant at some sites of distur*]^! soil. 

Klein, in his section on plant species in this re'wrt, includes this form in the species listed above.

Hierochloe odorata (L.) Wahlenb. 7849, 3d. Occurs as individual plants or small colonies in the Empetrum heath.

Phleum oommutatum var. americanum (Fourn.) Hult. 7466, 

4a. Common 'in the Empetrum heath and on disturbed soil 

at low altitudes.

**Poa alpina L. 8032, lOc. This collection represents an important range extension for this species. The specimens grew 

in the rocky soil of an old trail, and Hulten (written commun., 1968) remarked that they were "The smallest specimens I have seen." Klein, in his section on plant species 

in this report, placed Poa hispidula, P. komarovii, P. 

lanata, and P. turneri and their varieties in the P. alpina 

species complex.

*Poa aretica subsp. idlliamsii (Nash) Hult. 7609, 14a. Apparently a coprophilous plant. Found only on the summit 

of a bird-perch mound.

Poa eminens Presl. 7342, 2a; 7493, 2a; 8190, 2a. The short 

blades are thick and rigid, and the entire plant light purple. Found only on the deposit of sand and shells at South 

Bight and on sand dunes near Crown Reefer Point.

Poa lifepidula var. aleuttoa, Hult. (See Hulten, 1960, p. 

94-95.)

Poa Jiispidula var. wmpara, Hult. (See Hulten, 1960, p. 95.)

Poa, malacantha Kom. (P. komarovii Roshev., Hulten, 1960, 

p. 96).

*Poa lanata Scribn. & Merr. 8187, 2a. Found only on the deposit of sand and shells at South Bight.

*Poa lanata var. viin-para Hult. 7566, 12b. Found only in 

gravelly soil of a roadside ditch at an altitude of about 

650 feet.

Poa turneri Scribn. 8186, lOc; Erdman 548, lOc. Fonnd only 

on rock rubble on an old road near Constantine Harbor.

Puccinellia langeana subsp. alaskana (Scribn. £ Merr.) 

S0rens, 7504. 3a; 7617, 4d. Apparently a coprophilous species. Found only in drain channels of bird perches on the 

coastal cliffs and on the summits of bird-perch mounds.

Trisetum spicatum subsp. alaskanum (Nash) Hult. (See 

Hulten, 1960, p. 90.) Tatewaki and Kobayashi (1934) recognized both this taxon and the variety molle Scribn. & 

Merr. in the Amchitka Island flora.

Vahlodea atropurpurea subsp. latifolia (Hook.) Pors. 7464, 

4a; 8220, 13b. Occurs in the Empetrum heath and in alpine meadows. Considered by Klein, in his section on 

plant species in this report, to be Deschampsia atropurpurea (Wahl.) Sheele.
PLANT SPECIES OF AMCHITKA ISLAND

Vahlodea atropurpurea subsp. paramushirensis (Kudo) 

Hult. (See Hulten, 1968, p. 115; Amchitka Island occurrence indicated by a dot on the distribution map.)

HALORAGACEAE

*Hippuris montana Ledeb. 8227, 13a; 8261, 13a. This plant 

has the appearance of a Oalium or a Poffonatum; the 

stems are about 10 to 15 cm long, but only the leafy part, 

about 1 to 4 cm long, protrudes from the mud. This plant 

was also found in a Sphagnum mat of a small mountain 

stream that issued from a spring.

Hippuris vulgaris L. 7627, 7a. Common in lakes and pools 

at low altitudes; the leafy stems commonly emerge from 

the water, which can be as much as 1 m deep. 

Hyriophyllum spicatum L. (See Hulten, 1960, p. 268.) Not 

found during the present study, although extensively 

sought. Its superficial resemblance to the common Ranunculus trichophyllus may have caused it to be 

overlooked.

IRIDACEAE

Iris setosa subsp. setosa Hult. Reported to occur at Chitka 

'Cove (sight record, by W. M. Klein).

ISOETACEAE

Isoetes muricata subsp. maritima (Underw.) Hult. 7373, 7b; 

7396, 7b; Erdman 575, 7b. Found only in lakes, growing 

in water about 0.5 m deep. The plants are robust, with 

conns as much as 3 cm in diameter; many sporelings also 

were found.

Isoetes sp. 8192, 8c. A slender plant with only slight corm development; occurs in shallow rock-bottomed pools. Its 

aspect and habitat suggest that it may be a different taxon 

from that listed above.

JUNCACEAE

Juncus arcticus subsp. sitctiensis Engelm. 7634, la - A common species at wet sites; occurs at low to medium-high 

altitudes.

* Juncus biglumis L. 7576, 12b; Erdman 579, 12b. These collections are from unstable soil at high altitudes. The 

plants are from 1 to 8 cm high.

*Juncus ensifolius Wikstr. 7685, 8a; 7882, 8a. Found only at 

pool margins at low altitudes. Even when sterile, plant 

can be recognized by its ensiform leaves.

*Juncus trialiimis subsp. albescens (Lange) Hult. 7484, 8a; 

8210, 8a. A densely caespitose species that grows in shallow pools at low altitudes. The culms are about 15 cm 

high.

Luzula arcuata subsp. unalaschcensis (Buchenau) Hult. 

7563, lla; 7608, 14a; 8194, 14a. Found only at altitudes 

above 600 feet.

Luzula multi flora subsp. kobayasii (Satake) Hult. 7461, lOc. 

Found on disturbed mineral soil near Constantine Harbor.

Luzula multiflora subsp. multi flora var. frigida (Buchenau) 

'Sam. 7419, lOa; 7659, 4a. Specimens of the multifiora 

complex are abundant on disturbed sites on the low 

plateaus.

Lusula parviflora subsp. parviflora Hult. 7472, lOa; 7686, 

4e; 8030, lOc. Found only at low altitudes on banks and 

roadsides; not common.

Luzula tundricola Gorodk. 7883, 12a; 7907, 12b; 7994, Ha. 

Found only at altitudes above 1,100 feet in exposed fellfields and on solifluction terraces.

Luzula wahleribergii subsp. piperi (Oov.) Hult. Amchit>a 

Island occurrence shown on the distribution map by Hulten (1968, p. 297). Listed from this island as L. divaricata 

Watson by Tatewaki and and Kobayashi (1934, p. 112).

LEGUMINOSAE

Lathyrus maritimus subsp. pubescens (Hartm.) C. Reg^l. 

7489, la. Common on beaches and sand banks near the 

coasts; flowers and fruits are produced abundantly.

Lathyrus palustris subsp. pilosus (Cham.) Hult. Hulten 

(1960, p. 255) wrote, "Amchitka, according to Tatew. & 

Kobay." This report was verified by Hulten (1968, p. 67f). 

Not found during the present study, although often 

sought.

Lttpinus nootkatensis Donn. 7^86, lOb; 8044, 5b; 8228, 13b. 

An abundant and very showy plant, especially common 

along roadsides; occurs at all altitudes. Two colonies 

of plants found had pale-pink to nearly white flowers; 

these may be referred to f. leucanthus Lepage.

*Trifolium repens L. One clone of this adventive plant was 

found on a disturbed site by W. L. Slauson and M. A. 

Heifner.

LENTIBULARIACEAE

Pinffuicula vulffaris subsp. macroceras (Link) Calder & 

Taylor. 7414, 5a ; Erdman 753, 4b. Occurs only at low altitudes at very wet sites in the Empetrum heath.

LILIACEAE

Fritillaria camschatcensis (L.) Ker-Gawl. Klein 2718, 4e. 

Abundant in streamside communities throughout the low 

plateaus.

*Majanthemum dilatatum (How.) Nels. & Macbr. 7866, 4f.

Found only on peat banks and is only locally abundant.

Streptopus amplexifolius (L.) DC. 7543, 4e. Found only in

streamside communities, but was not abundant at any

location.

Tofleldia coccinea Richards. 7262, 14a; 7410, 4a. Common 

in a wide variety of habitats throughout the island.

LYCOPODIACEAE

Lycopodium alpinum L. Keith 108, lla. Found only in moss

and liverwort mats in a small stream. 

Lycopodium annotinum subsp. annotinum Hult. 7236, 5c;

7284, 4a; 7680, 4a ; Keith 107, 4a. Locally abundant in the

heath of the low plateaus. 

Lycopodium olavatum subsp. clavatum Hult. 7283, 4a;

7681, 4a; Keith 109, 4a. Grows with the species listed

above. 

Lycopodium sabinaefolium var. sitchense (Rupr.) Fe~n.

7237, 5c; 7263, 14a. Found commonly on low-a'ltitrde

bedrock outcrops. 

Lycopodium selago subsp. selago Hult. 7253, 14a; 7432, lOc;

7870, 4a. Widely distributed, although not abundant,

throughout the island.

ONAGRAGEAE

Epilobium behringianum Haussk. 7560, lla. Apparently common on the island; however, because of confusion in nomenclature with the following species, its abundance is 

not known with certainty.

Epilobium glandulosum Lehm. 7448, 6a; 7547, 4g. A robust 

plant as much as 40 cm high if growing in wet sedge meadows ; much smaller if growing in the Empetrum heath.
38 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

Klein, in his section on plant species in this report, refers 

this species to E. boreale Haussk.

Epilobiwm sertulatum Haussk. 7436, lOb; 7437, lOb; S039, 

5b. A low-growing plant occurring along roadsides and on 

disturbed soil. Plants at a site may either have green 

leaves and pale-pink flowers or be dark red throughout.

ORCHIDACEAE

Dactylorliiza aristata (Fisch.) Soo. 7417, 3d; 8027, lOb. 

A showy species that is locally common at low altitudes, 

especially on compacted mineral soil. The lip is pale 

magenta spotted with purple; other petals are deep lavender. One white-flowered specimen was found (Klein 1744)-

Listera cordata var. nephrophylla (Rydb.) Hult. 7416, 4a; 

7679, 4a. This very small orchid occurs as scattered individuals in the moss mat of banks in the Empetrum heath 

and in alpine meadows. Plants with either green or purplish-brown flowers grow intermixed.

* 'Platanthera chorisiana (Oham.) Rchb. 741S, 4a; 7497, 

14a ; 8210, lOc. Found only on lithosol or on sparsely vegetated mineral soil. The petals and sepals at anthesis are 

greenish yellow with narrow whitish margins; they soon 

'become brown tipped or brown dotted and streaked.

Platanthera convallariaefolia (Fisch.) Lindl. 7426, 4b. This 

and P. dilatata are common in wet places at low altitudes; 

usually both species occur intermixed. The flowers are 

green.

Platanthera, dilatata (Pursh) Lindl. 7468, 4a. Very similar 

to P. convallariaefolia, but has pure-white flowers. The 

flowers of Amchitka Island specimens are not fragrant, 

contrary to the report of Hutchison (1937, p. 231).

Platanthera tipuloides var. behringiana (Rydb.) Hult. (See 

Hulten, 1968, p. 323.)

PAPAVERACEAJE

Papaver alaskanum Hult. Collected in the mountains of 

Amchitka Island; the specimen is preserved in the 

Colorado State University herbarium.

PLANTAGINACEAE

*Plantago macrocarpa Chain. & Schlecht. 7427, 6a. Occurs

in wet sedge meadows on the low plateaus. 

Plantaao maritima subsp. juncoides (Lam.) Hult. 7660, 5a; 

797^, 3b. Occurs in bogs and on soil of coastal cliffs.

POLYGONACEAE

*Koenigia islandica L. 7574, 12b; Erdman 562, 12b; Erdman 

577, 12b. Found only on wet mineral soil of solifluction 

slopes at high altitudes.

*Oxyria digyna (L.) Hill. 7582, 13a. Rare; found only at

high altitudes. 

Polygonum viviparum L. 7435, lOb; 7562, lla; 8218, 13b.

Widely distributed but nowhere abundant on the island. 

Rumex fenestratus Greene. (See Hulte'n, 1960, p. 167-170.)

Hutchison (1937, p. 232) reported this species to be

"very luxuriant and fairly common on Amchitka."

* Rumex obtusifolius L. 7544. la. This European and Asiatic 

species is adventive in the Aleutian Islands (Hulten, 1960, 

p. 168). It occurs as isolated individuals on wet disturbed 

soil, usually in areas of human habitation.

POLYPODIACEAE

Athyrium filix-femina subsp. cyclosorum (Rupr.) Christens.

7517, 3d; 76S7, 4e. The common tall fern of s c.reamside

communities; it also occurs widely scattered through the

Empetrum heath. 

Cystopteris fragilis subsp. fragilis Hult. 7690, 4e; 7904, 3c.

Most common in crevices of the sea cliffs, but also occurs

on shaded peat banks. 

Dryopteris dilatata subsp. americana (Fisch.) Fult 7688,

4e. A tall fern that occurs sparingly in streamside

communities. 

Gymnocarpium dryopteris (L.) Newm. (See Hul^n, 1960,

p. 50, as Dryopteris linneana C. Chr.) 

^Polypodiutn vulgare subsp. occidentale (Hook.) Hult. 8268,

4d. Found only at the base, of a bird-perch mound above

the Pacific Ocean cliffs near the entrance to South Bight.

On February 8 the leaves were green and b^re many

sori, although the sporangia were empty. 

Thelypteris limbosperma (All.) Fuchs. 8244, 14b; 8245, 13b.

Found only at high altitudes; in alpine meadows or rock

slides. 

Thelypteris phegopteris (L.) Slosson. 8243, 13b. Found only

in alpine meadows.

PORTULACACEAE

*Claytonia arctica Adams. Klein 2820, 12b. Found only in 

a fell-field community; occurs as scattered individuals. 

The corolla is white.

Claytonia sibirica L. 7406, 4b. Very abundant, usually in 

wet locations; on low plateaus and in alpine meadows. 

Plants with white or pink petals occur together. The 

white-flowering form was named var. albiflorc, by Tatewaki and Kobayashi (1934, p. 34) and was reported from 

Amchitka Island.

*Montia fontana subsp. fontana Hult. 7632, 3b; 7684, 9a; 

8198, 3b. Found only in cascades and on dripping sea cliffs.

PRIMULACEAE

Primula cuneifolia subsp. saxifragifolia (Lehm.) Sm. & Forrest. 7421, lOa ; 7430, 4a ; 7583, 13a ; 7848, lOa ; 7869, 4a ; 

Erdman 551, 14a. Occurs at all altitudes on the island and 

is most abundant on disturbed soil. Flowers in late May 

and early June on the low plateaus and in late July and 

early August at high altitudes. The variety albiflora 

Koidzumi was reported for Amchitka Island by Tatewaki 

and Kobayashi (1934, p. 72).

Trientalis europaea subsp. arctica (Fisch.) Hult. 7501, 4a. 

Common in the Empetrum heath at low altitudes; flowers 

in mid-August. Tatewaki and Kobayashi (19^?4, p. 73- 

74) described the variety alentica and designated the Amchitka Island specimen as the type. Hulten (1960, p. 

298) considered this variety to be "a commo^ form of 

the plant."

PYROLACEAE

*Pyrola asarifolia var. purpurea (Bunge) Fern. 7611, 4a. 

Only one plant was found, although the species vas sought 

extensively. Flowers in mid-August.

*Pyrola minor L. 8219, 13b. Found only along streams in the 

mountains; locally abundant. Flowers the latter part 

of July.
PLANT SPECIES OF AMCHTTKA ISLAND 39

BANUNCULACEAE

Anemone narcissi flora subsp. villosissima (DC.) Hult. 7413, 

3d; 7844, 4a. One of the earliest flowering herbs, and 

very abundant and conspicuous throughout the low 

plateaus.

CaltJia palustris subsp. asarifolia (DC.) Hult. 7200, 6b; 

1629, 6b. Produces new leaves under water in November 

and December that emerge in early spring; fully developed 

leaves and flowers are present in early June.

Copt is trifolia (L.) Salisb. 7222, 4a; 7628, 4c; 8052, 4a. The 

firm shiny leaves persist through the winter, some becoming red, and are replaced by new leaves at the time of 

flowering in late June.

*R<munculus Jiyperboretis subsp. hyperboreiis Hult. 7429, 

7b. Found only by a small pool, growing on mud; flowers 

the first half of August.

Ranunculus occidentalis subsp. insiilaris Hult. 7909, 12b. 

Common throughout the island, except at high altitudes.

Ranunculus occidentalis subsp. nelsoni (DC.) Hult. 7450, 

6a. Found only in wet sedge meadows.

Ranunculus reptans L. 7598, 7b; 7399, 7b; Erdman 553, 

8b. Grows as a submersed or emergent aquatic, or on 

muddy lake shores. The leaf shape ranges from linear 

to spatulate, and is very similar to that of Limosella 

aquatica.

Ranunculus trichophyllus var. trichophyllus Hult. 7206, 8a; 

7444, 9a; 7674, 9a; 8266, 9a. Grows submersed in rapidly 

flowing streams, in pools of wet sedge meadows, and in 

lakes.

ROSAOEAE

Geum calthifolium Menzies. 7462, 4a. Abundant on the Empetrutn heath and in alpine meadows; flowers from late 

July to mid-August.

Geutn macrophyllum subsp. macrophyllum Hult. 7449, 6a. 

One of the few tall forbs on the island, reaching a height 

of 1 m at places. It may be an adventive species (Hulten, 

1960, p. 247).

Geum pentapetaltim (L.) Makino. 8215, 13a. This low plant 

occurs only at high altitudes in wet snow-bed communities. The white flowers appear in mid-July.

Potentilla hyparctica Malte. 7572, 12b; 7605, 12b; 7993, 

12b. This species is restricted to unstable soil at high 

altitudes.

Potentilla egedii subsp. grandis (Torr. & Gray) Hult. 7630, 

lOb. Found only on gravelly soil of a lake shore. Flowers 

in mid-August.

Potentilla villosa Pall. 7487, 3c; 7913, 3c; 8021, Be. A very 

conspicuous plant with silvery pubescent leaves and stems; 

the yellow flowers are produced from early June to midAugust. Found only on sea cliffs, where it is abundant.

Rul)us chamaemorus L. 7405, 4a. Abundant in the Empetrum mat; fruits freely, the fruits not yet ripe at the end 

of August.

Ruftnx arctlcus subsp. stellatus (Sm.) Boiv. emend. Hult. 

7405, 4a; 7476, 4g. The red flowers are produced the first 

week of August, but none that produced fruit was 

observed.

Sibbatdia procumbens L. 7411, 4g; 7987, 12a; 8029, lOc. Occurs throughout the island; usually grows on disturbed 

mineral soil.

BUBIACEAE

*Galium aparinc L. 8195, la. Found only in ruderal habitats,

usually in driftwood deposits.

Galium trifldum subsp. columbianum (Rydb.) Hult. 7652, 

la; 7678, 4a; 8045, 6a. Found only at low altitudes; most 

common in deposits of driftwood or in wet sedge meadows 

near the ocean.

SALICACEAE

Salix arctica subsp. crassijulis (Trautv.) Skvortz. 7420, 3d; 

7551, 14a; 79S8, 12a; 7996, lla; Erdman 549, 4a. The 

largest native woody plant on the island. The prostrate 

stems are as much as 0.5 m long and 1 cm in diameter. 

Widely distributed, but most abundant at sites near the 

ocean.

Salix cyclophylla Bydb. 7548, 14a; 1851, 12b; 7884, ITa; 

8209, 4a. Widely distributed, but most abundant in 

patches of discontinuous heath at high 'altitudes.

Salix polaris subsp. pseudopolaris (Flod.) Hult. Collected 

on this island by Tatewaki and Takahashi, according to 

Hultfin (I960, p. 161).

Salix rotundifolia Trautv. 7558, 14a; 7S85, 12a; 7586, l£a; 

7998, lla; Erdman 560, lla. Found only at altitudes of 

800 feet or more; commonly grows with S. cyclophylla.

SAXIFBAGACEAE

*Leptarrhena pyrolifolia (D. Don) Ser. 7584, 13a; 8222, 13a.

Bestricted to high-altitude snow beds; locally common.

Flowers in mid-July. 

Parnassia kotzebuei Cham. & Schlecht. 7477, 8a;8042, 5b.

Found only on wet mineral soil at low altitudes. Flowers

from mid-July to mid-August. 

Saxifraga bracteata D. Don. 74S8, 3b; 7616, 3b; 7881, f b;

7912, 3b. A common and characteristic plant of coastal

cliffs.

*Saxifraga foliolosa var. foliolosa Hult. 7579, 12b. Rare; 

found only on wet solifluction slopes. The plant bears 

many bulbils in the branches of the inflorescence.

*Saxifraga liirculus L. 8225, 13a. Found only in moss and 

liverwort mats of snow beds. This is the only report of 

this species from the central and western Aleutian 

Islands.

Saxifraga pimctata subsp. insularis Hult. 7445, 4e; 7577, 

12b; 7578, 12b;7868, lOc; 7985, 4a; 8043, 5b; 8824, 12b; 

Erdman 554, 5b. Common and widespread, usually occurs in sheltered locations.

SCBOPHULARIACEAE

*Castitteja unalaschcensis (Cham. & Schlecht.) Malte. 8lc,uson, 15c. Found growing with the introduced Pieea 

sitcliensis that is shown in figure 35.

*Limosella aquatica L. 7221, 7b; 7394, 7b; 8200, 5b. Was not 

found flowering. The vegetative organs are similar to 

those of Subularia aquatica and to submersed forms of 

Ranunculus reptans.

Mimulus guttatus DC. 7622, 9a. Found flowering in late 

July to mid-August in shallow streams of the Empetr'wi 

heath and in cascades at the coastal cliffs.

Pedicularis chamissonls Stev. 7422, 4a; Erdman 552, 6a. 

The large spikes of reddish-purple flowers, produced in
40 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

late July to early August, are conspicuous in the 

Empetrum heath.

*Rliinanthus minor subsp. ~borealis (Sterneck) Love. 7431, 

lOc; Erdman 574, 6a. Grows in disturbed mineral soil at 

low altitudes; flowers in late July and early August.

Veronica americana Schwein. 1239, 9a; 7446, 9a; 1516, 8a. 

Common along margins of swift streams. Flowers from 

late July until December, the latest flowers being produced entirely under water.

Veronica grandiflora Gaertn. 79861), 12a. A rare plant of 

high-altitude unstable lithosol. The dark-violet flowers 

are borne on very small plants in mid-June.

Veronica serpyllifolia subsp. humifusa (Dickson) Syme. 

7403, 8a; 75^5, 4g; 7581, 13a; 8033, lOe; 8041, 5b. Abundant on mineral soil at low altitudes; flowers all summer.

Veronica stelleri Pall. 7.^99, 14a; Erdman 550, 14a; Erdman 

561, 12b. A small pubescent plant that grows on disturbed soil and lithosol. Bears lavender campanulate 

flowers from late July to mid-August.

SELAGINELLACEAE

*Selaginella selaginoides (L.) Link. 7415, 5a; 8191, 8a. 

Found only as scattered plants on the low plateaus.

SPABGANIACEAE

Sparganium Jiyperboreum Laest. 7585, 13a; 8193, 8a. The 

plants are very slender and depauperate and grow on the 

mud at snow-bed sites and in shallow pools. Flowers 

sparingly in late July.

UMBELLIFEBAE

Angelica lucida L. 7.J80, lOb. A common plant of the Empetrum heath; flowers in early to mid-August. 

Conioselinum chinense (L.) BSP. 7623, 4e; 8197, 3d. Occurs

as scattered plants on banks and streamsides; flowers

sparingly in late July and early August. 

Heracleum lanatum Michx. 7635, la. An abundant coarse

plant of many habitats on the low plateaus. 

Ligusticum scoticum subsp. hultetiii (Fern.) Calder &

Taylor. 7502, 2c. Found only on soil banks at the top of

sea cliffs; locally abundant.

VIOLACEAE

Viola langsdorffli Fisch. 7442, 6a; 7^98, 14a; 7677, 4a; 78^7, 

4b. Common to locally abundant in the Empetrum heath 

and alpine meadows; flowers from early June to midAugust.

ANNOTATED LIST OF BRYOPHYTES

By HERMAN PEBSsoN 2 and HANSFOBD T. SHACKLETTE

Duplicate packets of specimens cited in this list have 

been examined, and the species have been identified by 

me (Persson), except as noted. I (Persson, 1968) have 

also examined the entire collection of many specimens, 

if critical specimens were found in the duplicated 

packets, and have published an account of the new or 

otherwise noteworthy bryophytes from Amchitka 

Island. Many collections, especially of liverworts, are

2 Paleobotaniska avdelningen, Naturhistoriska Riksmuseum, Stockholm, Sweden.

mixtures of several to many species; the same collection 

number is used for all members of each species that 

were segregated from the mixture.

In the list that follows, the collection numbers, in 

italics, following the species name are those of Shacklette, except as noted. The habitat and plant community 

in which a species was found, designated by numbers 

and letters as set forth in the section on the island and 

its vegetation, follow the collection numbers. The habitat notes and community assignments were provided by 

Shacklette. Species listed as newly reported for the 

Aleutian Islands (Persson, 1968) are preceded by one 

asterisk; those newly reported for Alaska are indicated 

by two asterisks.

I (Persson) am indebted to Dr. T. Amakawa, Shyukan High School, Fukuoka, Japan, for the determination of a Plectocolea that was in poor condition for 

study; to Dr. Eustace W. Jones, Oxford University, 

Oxford, England, for confirmation of my determinations of two Anthelia specimens; to Dr. N. Kitagawa, 

Nara University of Education, Nara, Japan, for confirmation of my determination of a Cephaloziella specimen and for determination or confirmation of some 

critical and poorly represented Lophozia specimens of 

the section Ventricosae; to Dr. Elsa Nyholm, Naturhistoriska Riksmuseum, Stockholm, Sweden, for determination of a Sphagnum specimen; to Dr. E. R. B. 

Little, Kingston Upon Hull, England, for naming two 

Riccardia latifrons specimens and confirming the determination of another; and to Dr. M. Mizutani, Hattori 

Botanical Laboratory, Obi-Nichinan-shi, Japan, for his 

critical study of a Riccardia specimen.

Identification of the Amchitkan hepatic specimens 

has been rather difficult because these plants cften are 

represented only by single stems that creep through 

tufts of other bryophytes; this condition of growth is 

common in alpine and similar regions. In addition, the 

sterility of the bryophytes, particularly the hepatics, 

is especially pronounced in these specimens and doubtless is caused by the severe climate that is charrcteristic 

of the Aleutian Islands.

HEPATICAE 

Order TAKAKIALES 

TAKAKIACEAE

Takakia ccratophylla (Mitt.) Gro. [Lepidosia ce~atophylla 

Mitt.]. 7591, 13a. Collected August 16, 1966, and first 

segregated from a packet of Andreaea rupestris by Dr. 

Howard Crum. This plant grew in a drain of a snow-bed 

community at an altitude of about 650 feet and was 

associated with large colonies of Anthelia julacea, 8capania paltidosa, Sphagnum compaction, and other snowbed bryophytes. Reported for Amchitka Island by Sharp 

and Hattori (1967), Hattori, Sharp, Mizutani, and Iwatsuki (1968), and Persson (1968).
PLANT SPECIES OF AMCHITKA ISLAND 41

Order JUNX3KERMANNTALES 

ADELANTHAOBAE

Odontoschisma elongatum (Lindto.) Evans. 7338, 4c. Found 

only on the summit of a small moss mound that developed on wet frost-heaved soil.

ANTHELIACEAE

Anthelia julacea (L.) Dum. 7235, 5c; 7564, 13a; 7592, 13a; 

7593, 13a; 8011, 14b; 8019, 14b. A characteristic species of 

drains from snow-beds at high altitudes and very conspicuous because of the large gray colonies that it forms. 

Dr. E. W. Jones has kindly confirmed the determinations 

of Nos. 7592 and 7593.

CALYPOGEIAGEAE

Calypogeia miilleriana (Schiffn.) K. Mull. 7261«, 4f. On a 

shaded peat bank above a small stream in the Empetrum 

heath, with Cephalozia media, Mniutn glabrescens, and 

Peltigera canina.

Calypogeia sphagnioola (Arn. & Perss.) Warnst. & Loeske. 

7209, 5a; 7214, 4a; 7258, 4c; 7341, 4c; 7455, 5a; 8281, 12a. 

In bogs at low altitudes, intermixed with Sphagnum 

capillaceitm, S. magellanicum, and Riceardia latifrons 

and on small moss mounds with Dicranum elongatum.

Calypogeia triohomanis (L). Corda. 7540, 4b; 7862, 4e. In 

the Empetrum heath, intermixed with Oncophorux wahlenbergii, Diplophyllum albicam, and Oymnocolea 

inflata.

CEPHALOZIACEAE

Cephalozia ambigua Massal, 7524, 4f; 7564, 13a; 8232, 13b. 

On a peat bank in the Empetrum heath, intermixed with 

Mnium glabrescens, Lophozia cf. groenlandica, Dicranella 

Jieteromalla, Nardia scalaris, and Scapania sca-ndica, and 

at altitudes of about 500 feet with Lophozia alpestris and 

Nardia geoscyphus.

Cephalosia biscuspidata (L.) Dum. 7258, 4c; 7265, 4f; 7300, 

lOa; 7340, 4e; 7386, lOa; 7522, 4f; 7526, 4f; 7528, 4f; 

7564, 13a; 7688, 4f; 7670, 4f; 7981 (cf.), 12a; 7.978, 4e; 

8233, 4e; 8235, 13b; 8262, 4f. One of the most common 

leafy liverworts on the island; grows in a wide variety 

of habitats. It is the principal component of the pendant 

bryophyte colonies of a peat bank shown in figure 14.

Cephalozia teucantha Spruce. 7242, 4c; 7258, 4c; 8281, 14b. 

Found on the summits of moss mounds, intermixed with 

Dicranum elongatum, D. groenlandic-um, Microlepidozia 

makinoana, Cephalozia bicuspidata, Anastrophyllum 

sphenoloboides, and Sphenolobus minutus; also found on 

bedrock outcrops.

Cephalosia media Lindb. 7261a, 4c; 8087, 4e; 8091, 4e; 8275, 

4e. Found on shaded peat banks, intermixed with Mnium 

glabrescens, Calypogeia miilJeriana, and Peltigera canina.

CepJialozia pleniceps (Aust.) Lindb. 7874, 4f; 7899, 12a; 

8264, 4f. Found on peat banks and in a mixture of leafy 

liverworts from a fell-field community.

CEPHALOZIELLIACEAE

Cephalosiella arctica Bryhn & Douin. 7300, 4g; 7345, 2b; 

7640, 3b ; 7641, 3b ; 7642, 3b ; 7643, 3b ; 7644, 3b ; 7673, 4f. 

Grows as dense poMers on sandy banks above the seashore and in wet crevices of breccia sea cliffs, intermixed 

with Bryum lapponicum, Schtetidium maritimum, Barbulacylindrica, and Bartramia itJiyphylla.

HEBBERTACEAB

Herberta adunca (Dicks.) Gray [H. hutchinsiae (Gottsche 

& Rabenh.) Evans]. 7567'a, 12b. Found only at the base 

of a boulder on a solifluction terrace, with Tritomaria 

quinquedentata, Marsupella ema-rginata, Diplophyllum 

albicans, and Jungermannia atrovirens.

JUNGERMANNIACEAE

Jungermannia atrovirens Dum. 7567a, 12b. Found only on a 

wet solifluction terrace, growing with Herberta adunca 

and its associated species.

Jungermannia sphaerocarpa Hook. 8208, 5b. Found only on 

ridges in wet sandy soil, mixed with Pogonatum urnigerum, LopJiozia cf. groenlandica, and Dicranella subvlata.

Mylia anotnala (Hook.) Gray. 7564- 13a. Found only in a 

drain channel at an altitude of 650 feet near Chitka Foint, 

intermixed with SpJMgnum papillosum, 8. squarroswn, 

Anthelia julacea, CepJialozia ambiffita, Pellia neesiana, 

Oymnocolea inflata, Lophozia alpestris, and Cephalozia 

bicuspidata.

Mylia taylori (Hook.) Gray. 7335, 4c; 7338, 4c; 7424, 4b. 

Found only on moss mounds in the Empetrum heatt and 

in wet areas of the heath, where it formed tufts with 

Lophozia cf. groenlandica and Orthocaulis binsteadii.

Nardia compressa (Hook.) Gray. 8282, 13a. Found only on 

the bottom of a snow-bed pool, where it formed a dense 

polster.

Nardia geoscyphus (DeNot.) Lindb. 8232, 13b; 8235, 13b. 

Found inly in alpine meadows; grows with Lophozia 

opacifolia.

Nardia scalaris (Schrad.) Gray. 7265, 4f; 7300, 4g; 7359, 

2b; 7375, 14a; 7387, lOa ; 7389, 8a; 7523% 4f; 7524, 4f; 

7526, 4f; 7593, 13a ; 7667 4f; 7668, 4f; 7841, 4f; 7842, 4f; 

7877, 4f; 7879a, 4f; 8036, lOc; 8050, 6a; 8232, 4e; 8233, 

4e; 8358, 4f. Abundant in many habitats.

Plectocolea cf. obovata (Nees) Mitt. 7525, 4f. Found only 

in a peat bank community. I (Persson) had, with 

hesitation, identified this specimen as P. obovata. This 

specimen was examined by Doctor Amakawa (written 

commun., 1968), who expressed his opinion that is must 

 belong to this species.

LEPIDOZIACEAE

Bazzania tricrenata (Wahlenb.) Trevis. 7227, 4c. 7341, 4c.

Found only on moss mounds in the Empetrum heath. 

Microlepidozia maJcinoana (St.) Hatt. [Lepidozia t~ichooladus K. Mull. Mierolepidozia silvatica (Evans) Jnrg.l.

7242, 4c; 7261a, 4g; 734^, 4c; 7343, 4a. Found only in the

Empetrum heath.

LOPHOCOLEACEAE

Chiloscyphus pallescens (Ehrh.) Dum. 7177, 9a; 717?, 9a; 

7202, 4g; 728S, 9a; 7391, 8a; 7857, 4e. Forms mats in 

shallow rapid streams with Brachythecium asperriwum; 

also grows on frost scars with Scapania paludosa, and 

along pool banks in the Empetrum heath.

Lophocolea cuspidata (Nees) Limpr. 7352, 2b; 7353, 2b; 

7639, 3b; 7644, 3b; 7863, 13a. Grows as pure colonies or 

intermixed with Plagiotheeium roeseanum on sandy 

banks above the seashore; also in crevices of wet breccia 

sea cliffs, and with Rhytidiadelphus squarrosus and 

Radula polyclada in a stream margin at Bird
42 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

LOPHOZIACEAE

**Anastrophyllum sphenoloboides Sehust. (See Sdiuster, 1969. 

p. 741.) 7242, 4c; 7243, 4c; 7258, c. per., 4c. Found only 

at the summits of small moss mounds, intermixed with 

Dicranum groenlandicum, Cephalozia leucantha, and 

Microlepidozia makinoana. This species was represented 

in the entire material by only a few stems scattered 

through tufts of other bryophytes. I (Persson), with 

hesitation, identified it as Lophozia porphyroleuca 

(Nees) Schiffn.; the leaves were similar to those of the 

latter species, and the trigons of the leaf cells were very 

prominent. Later, I found a fertile stem with a perianth 

in one of the specimens (No. 7258). Dr. N. Kitagawa 

(written commun., 1968) examined this specimen and 

wrote, "Shacklette No. 7258, which was identified by you 

as Lophozia porphyroleuca, seems to be very different 

from the latter species in the brownish coloration of the 

plants, the rounded leaves (not longer than wide), and 

especially in the paroicous inflorescence. I have never before seen such a species. I think that it is a new species 

related to Isopaches bicrenatus, from which it differs 

chiefly in the female bract and the perianth mouth. However, the material is too scanty to describe a new 

species."

Several weeks later, Dr. Kitagawa, while examining 

specimens in Dr. Sinske Hattori's herbarium, discovered 

that the Amchitkan hepatic was a species of Anastrophyllum which Dr. Rudolph M. Schuster had discovered 

earlier in northern Greenland. Dr. Schuster (written 

commun., 1968) examined the fertile specimen (Shacklette No. 7258) and confirmed Dr. Kitagawa's identification of it.

*Gymnocolea inflata (Huds.) Dum. forma. 7888, 4c; 75^0, 

4b ; 7564, 13a. On wet areas in the Empetrum heath, intermixed with Oncophorus wahlenbergii, Calypogeia trichomanes, and Diplophyllum albican-s, and in drain channels at an altitude of 650 feet.

In my first report of these collections from Amchitka 

Island (Persson, 1968), I referred them to Gymnocolea 

acutiloba (Kaal.) K.M. After a renewed study of the extremely polymorphous Gymnocolea inflata complex, I have 

formed the opinion that G. acutiloba represents only a 

modification of G. inflata; this opinion is shared with 

several other taxonomists.

The Amchitkan form seems to resemble even more 

closely the form that Kitagawa (1966), in his monograph 

of Japanese Lophoziaceae, called G. marginata (St.) Hatt, 

under which he placed the entire material of Japanese 

Gymnocolea and stated that this taxon is endemic to Japan 

(Hokkaido, Honshu, and Kyushu). The synonyms that he 

gives are of interest: Sphenolobus marginatus St., 1917; 

Cephalozia montana Horik., 1932; Gymnocolea, montana 

(Horik.) Hatt, 1948; Gymnocolea inflata (Huds.) Dum. 

subsp. montana (Horik.) Hatt., 1958; and Gymnocolea 

montana (Horik.) Hatt. var. acuta Hatt. According to 

Kitakawa (1966, p. 113), the species grows "on muddy 

soil and mires at high altitudes."

I have studied good material of the Japanese Gymnocolea 

and, with Kitagawa, have observed that it is possible to 

find forms in some specimens which, according to Kitagawa 

(1966, p. 113). ". . . are quite similar to G. inflata, another 

polymorphic species." Kitagawa (1966, p. 111-112) also 

stated, "Gymnocolea contains only three species, G. inflata,

G. acutiloba, and G. marginata, all of which range from the 

arctic to temperate regions in the Northern Hemisphere. 

They are all polymorphic and related so closely to each 

other that G. acutiloba and G. marginata could be regarded 

as varieties or subspecies of G. inflata." He added (1966, 

p. 113), "However, most forms including the type specimens 

of G. marginata and G. montana are rather definitely different from G. inflata in the following features: T/te plants 

are less flaccid, the leaves are broader and more or less 

concave (sometimes even canaliculate), and the leaves are 

rather regularly arranged (leaves on a shoot spread at 

similar angles) and dorsally secund. These differences seem 

to be sufficient to make G. marginata specifically distinct 

from G. inflata."

The Amchitkan Gymnocolea (the first report of tMs genus 

in the Aleutian Islands) has the broad, more or less concave 

leaves of the Japanese form. With the form that has been 

called G. acutiloba, it shares the only important character 

of this taxon the fairly acute leaf lobes. It is noteworthy 

that Hattori (1958) described a variety acuta of G. montana 

which Kitagawa (1966) placed as a synonym of G. marginata but otherwise did not describe. The Aleutian form 

differs from both G. acutiloba, and G. marginata in the distinctly stronger walls, especially at the corners, of its leaf 

cells. The broad leaves, as found in the Amchitkan material, 

are occasionally found also in European specurens; the 

only leaf of Gymnocolea inflata which Macvicar (1960) 

illustrated in his well-known hepatic flora of Great Britain 

could as well have been taken from a flgxire of the Japanese 

Gymnocolea. The floras also commonly note that the leaves 

can be concave and even canaliculate. Because th^ habitat 

of G. inflata ranges from xerophytic to very hyt^ophytic, 

the species is extremely variable. Apart from the more 

commonly used handbooks and floras, the work of Jb'rgensen 

(1934) on the hepatics of Norway is recommended, for Ms 

treatment of the ecology and taxonomy of G. inflata. Unfortunately, this outstanding study of northern hepatics is 

written in Norwegian and, therefore, is too littT e known 

outside Scandinavia.

Gymnocolea inflata is widely distributed in the Northern 

Hemisphere; it is found, among other places, in Siberia, 

Greenland, and Iceland. In North America it occurs throughout the continent; its northernmost known legality in 

Alaska is Mount McKinley National Park, and it is also 

reported from Yukon Territory.

In summarizing, I consider the Gymnocolea inflata complex to be extremely plastic, and I believe that o^her taxa 

in the genus probably will prove to be environmental modifications. Experiments in the growth of these taxa on substrates of different pH, possibly in water culture, would be 

of great interest, especially because G. acutilola is generally regarded as one of the so-called copper mouses. 

Lophozia alpestris ( Schleich.) Evans, 7524, 4f; 7564, 13a ; 

7593, 13a; 7665, 6a; 8282, 13b; 8288, 13b; 8*34, 13b; 

8235, 13b. Commonly occurs in alpine meadows or snowbed communities, but it was also found in a wet sedge 

meadow with Pellia neesian-a and on a peat bank. 

Lophozia cf. groenlandica (Nees) Bryhn. 7424, 4b; 7425, 

4b; 752^, 4f; 7525, 4f; 7528, 4f; 8018, 14b ; 8202, 5b. Most 

common on the low plateaus but also found en a highaltitude bedrock outcrop. No. 7525 was named by Dr. 

Kitagawa, who also studied No. 7528 and agreed with 

me (Persson) on its identification. Kitagawa (written
PLANT SPECIES OF AMCHITKA ISLAND 43

commun., 1968) wrote that the specimens "agree well 

with [R.M.] Schuster's description of the species, except 

for the somewhat larger leaf cells."

*Lophosia incisa (Schrad.) Dum. (See Persson, 1968.) 7879, 

4f; 8018, 14b; 8019, 14b; 8285, 8c. Found on peat banks 

and andesite rock outcrops at both high and low altitudes.

**Lophozia opacifolia Culmann. (See Persson, 1968.) 8233, 

13b; 8234, 13b; 8235, 13b. Found only on wet soil by a 

stream in an alpine meadow, intermixed with Lophozia, 

alpestris, Nardia geoscyphus, and Cephalozia T)icuspidata.

Lophozia cf. icenzelii (Nees) Steph. 7556, 14a; 7568, 12a. 

(Specimen 7568 was identified by Dr. Kitigawa.) Found 

in patches of turf on a solifiuction terrace and on a highaltitude bedrock outcrop.

Orthooaulis Mnsteadii (Kaal.) Buch. 7338, 4c; 7340, 4c; 

7341, 4c; 7424, 4b; 7425, 4b; 8281, 14b. Found in the 

Empetrum heath, usually intermixed with many other 

bryophytes on moss mounds, and on a high-altitude bedrock outcrop.

OrthocauUs floerkei (Web. & Mohr) Buch. 8262, 4f. Found 

only on a peat bank above a small spring at Chitka Cove.

SphenoloMs mimttus (Crantz) Steph. 1241, 4c; 7242, 4c; 

7244, 4c; 1341, 4c; 8281, I4b. Found on moss* mounds in 

the Empetrum heath and on high-altitude bedrock outcrops.

**SpJieno]o1)us saccatulus (Lindb.) K.M. 1906-14 [Junoermannia minuta var. grandis Gottsche Mss.; J. riaida Lindb. 

j8 grandis Lindb. 1879; J. saccatula Lindb. 1883; Cephaloziopsis saccatula Schffn. 1893; Sphenolobus minutus var. 

grandis S. Arn. & O. Martensson 1959]. 7424, 4a. Found 

in the Empetrum heath in tufts' of Pogonatum alpinum, 

intermixed with Lophosia cf. oroenlandica, Mylia taylori, 

and Orthocaulis olnsteadii. This is the first report of the 

occurrence of this species in North America. 

Tritomaria quinquedentata (Huds.) Buch. 7569, 12b; 7567a, 

12b. Occurs as small round polsters on loose unstable 

soil among the pebbles of solifiuction slopes.

MARSUPELLACEAE

Gymnomitrion corallioides Nees. 7554, lla; 7594, 14b; Erdman 612, lla. Found as single polsters on almost bare, 

rocky soil of the discontinuous heath habitat and on highaltitude rock mounds.

Marsupella emarginata (Ehrh.) Dumort. 7220, 8a; 7234, 

5c; 7567a, 12b; 7569, 12b; 7854, 4e. Found as extensive 

colonies in a Philonotis-Parnassia bog, and common in 

smaller quantities at many other sites.

PTILIDIACEAE

PtiUdi-um ciliare (L.) Nees. 7338, 4c. Found at only one location on a moss mound. Surprisingly, this conspicuous 

liverwort was not found at other locations; apparently, 

it is actually rare on the island.

RADULACEAE

Radula polyclada Evans. 7863, 13a; 7864, 13a. Found only 

at the margin of a small stream at Bird Cape, with 

Rhytidiadelphus squarrosus and Lophocolea cuspidata.

SCAPANIACEAE

Diplophyllum albicans (L.) Dum. 7241, 4f; 7334, 4g; 7392, 

8a ; 7540, 4b ; 7567, 12a ; 7569, 12b; 7593, 13a; 7900, 12a ;

7978, 4e; 8016, 14b; 8019, 4d; 8280, 14b. Grows in a wide 

variety of habitats, usually as single stems internixed 

with other bryophytes.

Diplophyllum taxi folium (Wahlenb.) Dum, 7853,12a. Found 

only in a volcanic rubble fell-field.

Macrodiplophyllum plicatum (Lindb.) H. Perss. (See Persson, 1968.) 8010, 4d; 8016, 4d; 8017, 4d; 8020, 4d. Found 

on an andesite bird-perch mound at an altitude o* 950 

feet.

Scapania irrigua (Nees) Dum. 7482, 4b; 8264, 9a. Grows on 

wet peat banks and in small drain channels in peat.

Scapania paludosa K. Mull. 7202, 4g; 7217, 13a; 721?, 5c; 

7225, 4b; 7226, 4b ; 7459, 8c; 7589, 13a; 7864, 4e; 8212, 8a. 

Occurs at wet locations on the Empetrum heatl ; at 

places forms mats in small pools that support Siphula 

ceratites; also in snow-bed communities.

Scapania scandica (Arn. & Buch) Macvic. 7265, 4f; 7524, 

4f; 7525, c. per., 4f; 7879, 4f; 7999, 12a. Found in a peatbank community, associated with Mnium glabrescens, 

Cephalozia atnbigua, and Lophozia cf. groenlandica-; on 

an almost bare peat and soil bank with Pogonatum 

urnigerum, Cephalozia bicuspidata, and Nardia scalaris; 

and in a fell-field community.

Scapania suoalpina (Nees) Dum. 8293, 7b. Found orly in 

a rock-bottomed pond; grows submersed with Limosella 

aguatica.

Scapania undulata (L.) Dum. 7287, 9a; 7288, 9a; 7390. 8a; 

7856, 4e. Found in water of small pools with Nardia 

scalaris and Chiloscyplius pallescens, and in streams with 

Brachj/thecium asperrimum and Chiloscyphus pallescens.

Order METZGERIALES

ANEURAOEAE

Riccardia latifrons Lindb. 7338, 4c; 7455, 5a. On a small 

moss mound, with Ptilidlum ciliare, Mylia taylori, 

Odontoschisma elongatum, and Orthocaiilis binsteadii; 

also found in a bog with Sphagnum magellanicum, 8. 

squarrosum, and S. teres. Dr. E. R. B. Little (written 

commun., 1968) wrote, "No. 7338 is good Riccwdia 

latifrons, and I am almost sure that No. 7455 is the same 

species."

Riccardia pinguis (L.) Gray. 7185, 8a; 7199, 4e; 7593 13a. 

Occurs in very wet locations at pool and stream margins 

and in drains of snow beds.

Riccardia simiata (Dum.) Trevis. 7201, 4g. Found on wet 

mineral soil of a frost scar in the Empetrum heath. The 

determination of this specimen was confirmed by 

Dr. E. R. B. Little and also, with some hesitation by 

Dr. M. Mizutani.

BLASIACEAE

Blasia pusilla L. 7667, 4f. Found only in a peat-bank community with Nardia scalaris, Dicranella suoulata, and 

Cephalozia, Mcuspidata.

DILAENAOEAE

Pellia neesiana (Gottsche) Limpr. 7564, 13a; 7665, 6a. 

Found in wet sedge meadows with Lophozia alpestris and 

in snow-bed communities with Sphagnum papillosum, S. 

squarrosum, Anthelia julacea, Mylia anomala, and other 

bryophytes.
44 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

Order MARCHANTIALES 

CONOGEPHALACEAE

ConocepJwilum conicum (L.) Dum. 7198, 4e. Found only in 

streamside communities on sheltered banks in the Empetriim heath; easily identified in the field by its spicy odor 

after it has been crushed.

MAROHANTIACEAE

Marchantia polymorpha L. 7285, 3b; 7402, 8a; 7604, lOb. 

Not common; found only on the mud bottom of a drained 

pool, in an open steel barrel, and on thin soil of coastal 

cliffs.

SPHAGNOBRYA

SPHAGNACEAE

Sphagnum capillaceum (Weiss) Schrank. 7214. 4a. Found 

only near the base of a moss mound in the Empetrtim 

heath.

Sphagnum compaotum Lam. & DC. 7590,13a ; 7676, 8c; 7872, 

4f; 7970, 5a ; 7992, 12b; 8236, 13a. Found in the drain of a 

snow bed, and on wet soil in the Empetrum heath.

Sphagnum girgensohnii Buss. 7662, 6a; 7858, 4e. A principal 

component of bogs on the low plateaus.

Sphagnum lindoergii Schp. 8237, 13b; 8238, 13b. Found only 

in small streams of the alpine meadows.

Sphagnum magellanicum Brid. 7453, 5a; 7455, 5a; 8236,13b. 

A component of bogs on the low plateaus; also found in 

small streams of alpine meadows.

Sphagnum papillosum Lindb. 7564, 13a; 7661, 6a; 7663, 5a; 

7672, 4f; 7969, 5a. Most common at high altitudes in 

drains from snow beds, but also found on wet peat banks 

and wet sedge meadows of the low plateaus. Dr. Elsa 

Nyholm kindly determined specimen No. 7969.

Sphagnum squarrosum Sw. ex Crome. 7452, 5a; 7564, 13a; 

7661, 6a; 8293, 7b. A principal component of bogs on the 

low plateaus; also found in wet sedge meadows and in 

snow-bed communities.

Sphagnum subsecundum Nees ex Sturm. 7215, 4a. Found 

only near the base of a moss mound in the Empetrum 

heath.

Sphagnum teres (Schimp.) Angstr. ex C. Hartm. 7454, 5a; 

7663, 6a; 8260, 4e. Abundant in bogs and wet sedge meadows of the low plateaus; usually bright green in color.

ANDREAEOBRYA

ANDREACEAE

Andreaea rupestris Hedw. 7192, lib; 7256, 14a; 7363, 2c; 

7591, 13a; 7893, 12b; 8280, 14b. Found at all altitudes; 

usually grows on rocks, but at places in the mountains it 

forms mats in streams of alpine meadows and in snow-bed 

drains (Shacklette, 1966).

EITBRYA

AMBLYSTEGIACEAE

AmUystegium serpens (Hedw.) B.S.G. 8203, 5b; 8204, 3d. 

Found on a turf bank above the beach near Makarius Bay, 

growing with Leptobryum pyrifonne and Bryum stenotrichum; also found in a wet sandy 'borrow pit with Philonotis americana.

Oalliergon sarmentosum (Wahlenb.) Kindb. 8047, 4e. Found 

on a concrete spillway at a large spring near Makarius 

Bay.

Campylium stellatum (Hedw.) C. Jens. 7975, 4e; 8212, 8a. 

Submerged in a small pool in the Empetrum heath, growing with Drepanocladus revolvens and Scapania pnludosa; 

also found at the margin of a small stream in tl > heath.

Cratoneuron fllicinum (Hedw.) Spruce. 7636, la; ??44, 3b; 

7645, 3b; 7647, 3b; 7983, 3b. Found on dripping s«a cliffs, 

commonly with Vlota phyllantha, Bryum lapponteum, 

Schistidium maritimum, and Platydictya jungermannioides; also found at other sites on the seashore.

Drepanocladus fiuitans (Hedw.) Warnst. 8284, 6?. Found 

only in wet moss-lichen mats with Polytrichum tongisetum, on the lower terrace at Cyril Oove.

Drepanocladus revolvens (Sw.) Warnst. 7219, IFa; 7226, 

5c; 7567a, 12a; 7975, 13b; 8212, 8a. Occurs at rool margins, in streams from snow beds and alpine rreadows, 

and in fell-fields on the mountain summits.

Drepanocladus uncinatus (Hedw.) Warnst. 7188, lib; 7189, 

lib ; 7249, 14a ; 7327, 4c ; 7358, 2c; 7380, lOc; 75<>7a, 12b; 

7887, 12a; 7973, 15b; 8006, 4d; 8007, 4d. Abundant in 

many different habitats and distributed throughout the 

island.

Hygrohypnum oestii (Ren. & Bryhn ex Ren.) Holz. ex 

Broth. 7178, 9a; 7621, 9a; 8048, 9a. Found only at the 

margins of, or submersed in, swift streams on the low 

plateaus, commonly growing with Hygrohypnum ochraceum, Brachytheciwn asperrimum, and Chiloscyphus 

pallescens.

Hygrohypnum ochraceum (Turn, ex Wils.) Loes-ke. 7178, 

9a; 8269, 9a. Found associated with the spec'es listed 

above; also found with Bryoerythrophyllum recurvirostrum on a limonite deposit in a small stream.

Platydictya jungermannioides (Brid.) Crum. [Amolystegietta sprucei (Bruch) Loeske]. 7180, 3b; 7182, 3b; 7646, 

3b; 8275, 4e. Occurs in crevices and under small rock 

shelves of breccia sea cliffs, at two locations growing with 

Mielichhoferia macrocarpa, (Shacklette, 1967) ; also found 

on soil at the margin of a small stream in the heath.

AULACOMNIACEAE

Aulacomnium palustre (Hedw.) Schwaegr. 7231, 5c; 7233, 

5c; 7402, 8a; 7482, 6a ; 7663, 6a; 7850, 6a. A common moss 

in Sphagnum bogs, wet sedge meadows, and liverwort 

mats in streams.

BARTRAMIAOEAE

Bartramia ithyphylla Brid. 7641, 31). Found only on thin soil

over breccia sea cliffs. 

Conostomum tetragonum (Hedw.) Lindb. 7594, 14b; 8231,

12a. Found as small polsters on unstable rocky soil and

bedrock mounds at high altitudes. 

Philonotis americana Dism. 7441, lOb, 7457, 5a; 7482, 6a;

8201, 5b. Found in a wet sandy borrow pit, near Makarius

Bay, where it formed a dense mat over a large area; also

found on wet soil of a roadside ditch and in a wet sedge

meadow. 

Philonotis fontana (Hedw.) Brid. 7202, 4g; 7203, 4g; 7217,

5c. On wet disturbed mineral soil, commonly forming

large mats.
PLANT SPECIES OF AMCHITKA ISLAND 4-5

BRACHYTHECIACEAE

**Brachythecium acutum (Mitt.) Sull. (See Persson, 1968.) 

7380, lOc; 7850, 4a; 7373, 15b. Found in Empetrum mats 

near Constantine Harbor, on broken macadam runways, 

and on wet boards in a World War II building.

BrackytJiecium albicans (Hedw.) B.S.G. 7-Z89, lib; 7358, 2c; 

1381, lOc; 1981, 4d; 8002, 12b; 8291, 4f. Commonly found 

on sparsely vegetated rocky or sandy soil, with Rhacomitrlum lanuginosum, R. ericoides, Drepanocladus uncinatus, 

and Ceratodon purpureus, but may also be found in other 

habitats.

Brachythecium asperrimutn (Mitt.) Sull. 7-Z77, 9a ; 1224, 5c; 

7287, 9a ; 7343, 4a ; 7380, lOc; 7456, 5a ; 7691, 9a ; 7860, 9a ; 

7983, 9a. Occurs in a wide variety of habitats on the low 

plateaus.

Eurhynchium praelongum (Hedw.) B.S.G. 1191, 4e; 1216, 

3a ; 7258, 4c; 7269, 3a ; 7270, 3a; 1211, 3a; 7343, 4a ; 7350, 

2b ; 1351, 2b ; 7355, 2b ; 7382, lOc ; 7649, 3b ; 7902, 4e ; 7973, 

15b; 7977, 4e. This species is widespread at low altitudes 

and is usually abundant on sea cliffs. At places on the 

Pacific Ocean coast cliffs, it is the only bryophyte found.

BRYACEAE

*Anomo1)ryum concinnatum (Spruce) Lindb. [A. ftliformis 

(Dicks.) Solms var. concinnatum (Spruce) Loeske]. 

(See Persson, 1968.) 7670, 4f. Found only on a wet peat 

bank near Constantine Harbor, with CepJialozia bicuspidata, Bryoerytliropliyllum recurvirostrum, Polilia nutans, 

Pogonatum urnigerum, and Bryum arcticum.

*Bryum arcticum (B. Br.) B.S.G. (See Persson, 1968.) 

7673, 4f. Found only in the community listed immediately 

above.

*Bryiim pattens Sw. sens. lat. 7855, c. fr., 4e. Found only 

in a streamside community about 100 feet above sea level 

at Bird Cape, growing with Mnium glabrescens, Plectocolea cf. obovata, Polilia nutans, and Scapania undulata. 

Bryum pseiidotriqtietrum (Hedw.) Gaertn., Meyers & 

Scherb. 1199, 4e; 73^8, 2b. Found at the margin of a small 

stream in the heath, growing with Climaciiim dendroides 

and Riccardia pinguis; also found on a sandy bank at the 

seacoast.

*Bryum lapponicum Hag Kaur. [B. salinum Hag.] (See Persson, 1968.) 7542, 3b; 7640, 3b; 7650, 3b. Apparently restricted to sea cliffs that are subjected to salt spray or 

waves.

Bryum, stenotrich urn C. Mull. [B. inclinatum (Brid.) 

Bland.]. 7281, 12a ; 7483, lOa ; 7565, 15a; 7648, 3b; 8035, 

lOc; 8206, 3d. This moss, with Tetraplodon mnioides and 

Pleurosium schreberi, grew on decomposed animal carcasses ; it grows also on mineral and organic soil on 

Amchitka Island.

Leptobryum pyriforme (Hedw.) Wils. 8205, 3d; 8286, cf., 

4g. Found on a soil-covered sea cliff at Makarius Bay, 

with Amblystegiuni serpens and Bryum stenotrichum; 

also found on mineral soil of a frost scar.

*Mielichlioferia macrocarpa (Hook, ex Drumni.) Bruch & 

Schinip, ex Jaeg. & Sauerb. (See Persson, 1968.) 1181, 

3b; 1182, 3b; 1541, 3b; 8208, 3b. Shacklette (1967) discussed this "copper moss" as it occurs on Amchitka 

Island, where it is found on sea cliffs, growing only on 

narrow rock strata that have a greater concentration of 

metals than that found in adjacent strata. This moss was

first collected in Alaska on Sitkalidak Island by Eyerdam

(Persson, 1968) not on Amchitka Island, as erroneously

stated by Shacklette (1967). 

Polilia annotina (Hedw.) Loeske. 8258, 4f. Found only near

Constantine Harbor, on a peat and soil bank. 

PoMia cruda (Hedw.) Lindb. 7354, 2b; 1651, 3b; 8264, cf.,

4f. Found near the seacoast on sandy soil, breccia clifs,

and peat banks. 

Polilia nutans (Hedw.) Lindb. 7300, lOa; 7672, 4f; 7899,

12a; 7901, 12a; 8018, 14b. Found on soil and peat banks at

low altitudes, commonly growing with Pogonatum

urnigerum and Ditrichum heteromallum.

OLIMACIACEAE

Climaciiim dendroides (Hedw.) Web. & Mohr. 7194, 4^; 

7195, 4e. Found at only one location on wet soil by a 

small stream in the Empetrum heath. Apparently scare"; 

this distinctive moss is very easy to recognize in the fieM.

DICRANACEAE

AmpMdium lapponicum (Hedw.) Schimp. 1183, 3b. Fou^d 

only at Cyril Cove, growing below a breccia shelf with 

Pottia lieimii, a few feet above sea level.

*Aongstroemia longipes (Sommerf.) B.S.G. 8051, 6a. Fourd 

only in a wet sedge meadow near Makarius Bay.

Arctoa fulvella (Dicks.) B.S.G. 7853, c. fr., 12a. In a highaltitude fell-field, growing with Diplophyllum taxifoliv.m 

and Rhacomitriutn brevipes.

Diohodontium pellucidum (Hedw.) Schimp. 1521, 4f. On a 

wet peat bank at Constantine Harbor, growing with 

Cephalozia bicuspidata, Dicranella heteromttlla, and 

other bryophytes.

Dicranella heteromalla (Hedw.) Schimp. 7524, 4f; 7526, 4f; 

7528, 4f; 7877, 4f; 8050, c. fr., 5b. Commonly found on wet 

peat banks; also found on wet sandy soil near Cleveng-er 

Lake.

Dicranella palustris (Dicks.) Crundw. ex E. F. Warb. 

[Anisothecium squarrosum (Stark.) Lindb.]. 1201, 4<?r; 

7205, 4g; 1218, 5c ; 7588, 13a ; 7859, 5c; 7984, 5c. Found in 

a wide variety of habitats that range in altitude from the 

low plateaus to snow beds.

*Dicran-ella subulata (Hedw.) Schimp. (See Persson, 196P ) 

7385, lOa; 7386, lOa; 7668, 4f; 8201, 5b; 8203, 5b. Tl is 

moss is fairly common on the island, and it is surprising 

that it has not been found earlier in the Aleutian Islands.

Dicranum angustum Lindb. 7453, 5a. Found only in a lo^valtitude bog, with Sphagnum magellanicum, S. squarrosum, and 8. teres.

Dicranum etongatum Schleich. ex Schwaegr. 7258, 4c; 

7339, 4c; 7340, 4c; 8246, 4d. A common species at the 

summits of moss mounds. It was found on one orgaric 

bird-perch mound, where it was abnormally dark gre-^n 

and very robust.

Dicranum fuscescens Turn. 1316, 4c; 7333, 4ff- Found on 

moss mounds, although it is not a common component of 

the mound flora, and on frost scars.

Dicranum groenlandicurn Brid. 7242, 4c. The dominant moss 

on the summits of some small moss mounds.

Dicranum hoicellii Ren. & Lesq. 7233, 4a; 7252, 14a; 7325, 

4c; 7326, 4c; 7977, 4e; 8003, 12b. Very common in a wide 

variety of habitats. According to Persson (1968), probably all Aleutian collections named D. scoparium belong 

to this species.
46 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

Onoophorus wahleribergii Brid. 7223, 5c; 1/333, 4g; 75^0, 4b. 

This moss forms hard, black conical polsters in wet 

heath and in sedge meadows that can develop into small 

moss mounds 0.5 m in diameter and as much as 40 cm 

high. These polsters are saturated with water and freeze 

into hard icy lumps that one often stumbles over while 

walking on the otherwise resilient vegetation mat.

DITBICHACBAB

Ceratodon purpureus (Hedw.) Brid. 7280, lOc; 7348, 2b; 

7359, 2c ; 7376, 14a; 7378, lOc ; 7402, 4b ; 7483, lOa ; 8289, 

la. Common on compacted soil and on concrete and macadam walks and roads ; also found on peat deposits and 

other organic substrates.

Ditrichum heteromallum (Hedw.) Brid. 7300, lOa; 8050, 

c. fr., 5b. Found in a dense growth of Calamagrostis 

nutkaensis on a roadside bank of mineral soil, and on 

wet sandy soil in a PMlonotte-Parnassia community.

ENTODONTACEAE

Pleurozium schreberi (Brid.) Mitt. 7309, 4a ; 73-77, 4c ; 7328, 

4c; 7514, 15a; 7663, 6a; 7664, 6a. Very abundant as a 

component of the Empetrum heath mat.

FONTINALACEAE

Fontinalis neomexicana Sull. & Lesq. 7176, 9a; 7240, 9a; 

7447, 9a; 8207, 9a. Grows submersed in swift clear 

streams of the Empetrum heath. The "fronds" are as 

much as 60 cm long, and some of them fruit abundantly.

GBIMMIACBAB

Rhacomitrium aciculare (Hedw.) Brid. 7186, 8a. Only one 

polster was found; it was growing with Riccardia pinguis 

on the bottom of a small pool that had been drained.

Rhacomitrium brevipes Kindb. ex Mac. 7853, 12a; 8267, 12a. 

One of the two polsters found had formed a dense cylindrical cushion about 8 cm high in a fell-field community, 

where it had entrapped much windblown silt.

Rhacomitrium ericoides Brid. [R. canescen-s (Hedw.) Brid. 

var. ericoides (Brid.) B.S.G.] 7357, 2c; 736-7, 2c; 7279, 

lOc; 7666, lOa; 7890, 12a. Not common; found on sand 

dunes near the coast (Shacklette, 1966), on hard soil 

of a gravel road, and in a fell-field community.

Rhacomitrium fasciculare (Hedw.) Brid. 7373, 14a; 7508, 

3b; 7556, lla; 7873, 4f; 800.7, 12b; 8880, 14b. Found on 

sea cliffs, low- and high-altitude rock mounds, and solifluction terraces.

Rhacomitrium lanuginosum (Hedw.) Brid. 7.790, lla; 7.79-7, 

lib; 7207, 4c; 7241, 14a; 725.7, 14a; 73.74, 4c; 73.75, 4c; 

7362, 2c; 7372, 14a ; 7568, 12b; 7889, 12a; 7892, 12a ; 7901, 

12a; 7990, 12a; 8020, 4d. Very abundant throughout the 

island. Probably initiates mound formation more commonly than any other moss on the island.

Schistidium apooarpum (Hedw.) B.S.G. [Grimmia apocarpa 

Hedw.]. 7366, 2c. Unattached polsters of a spheroidal 

shape that were composed of this species were described 

from Amchitka Island (Shacklette, 1966), growing on 

a pebbly sand dune near Cyril Cove.

Sohistidium maritimiim B.S.G. [Grimmia maritima Turn.]. 

7267, 3a ; 7272, 3a ; 7274, 3c; 7507, 3c; 7649, 3b; 7650, 3b; 

8025, 3a. Apparently a strict halophyte; grows only on 

rocks that receive sea spray or waves.

HYLOCOMIACBAB

Hylocomium splendens (Hedw.) B.S.G. 7208, 4c; 7228, 4a; 

7307, lOa; 73.72, lOa; 7329, 4c; 7331, 4c; 7649, 4c. Found 

only on the low plateaus, usually mixed with ti e common 

mosses of the heath, or on moss mounds; at places it is the 

dominant moss on the collapsed summits of old mounds.

HYPNACBAB

Hypnum cupressiforme Hedw. (See Persson, 1P38.) 8007, 

14b. Found only on an andesite bird-perch mound near 

Buoy Point, growing with Drepanoctodus uno'natus and 

Pogonatum alpinum.

**Hypnum dieckii Ren. & Card. (See Persson, 1968.) 7850, 4a. 

Found only in the Empetrum heath near Consta ntine Harbor, growing with Aitlacomnium palustre, Brachythecium 

acutum, and Pogonatum alpinum.

Hypnum plicatulum (Lindb.) Jaeg. 8002, 12b. Found only 

at the margin of a vegetation mat on a solifluction terrace.

Ptilium crista-castrensis (Hedw.) DeNot. 7230, 4a; 7306, 

4a. Usually sparse, but at several locations on the Empetrum heath it was observed to form almost pure colonies 

5 sq m or more in size.

LBUCODONTACEAB

Antitrichia curtipendula (Hedw.) Brid. 7209, 4c; 7210, 4d; 

7248, 14a ; 7250, 14a; 7313, 4c ; 7318, 4c; 732?, 4c; 7330, 

4c ; 7331, 4c; 73/A 2b ; 7890, 12b ; 8002,12b; 8008, 4d. Common, and at places very abundant, on the Empetrum 

heath ; also found on bird-perch mounds and on solifluction 

terraces. Its yellow color makes it conspicuous as the 

summit moss on many "mature" moss mourns, visible 

even on aerial photographs (fig. 8). W. A. Weber (oral 

commun., 1967) suggested that in the treeless Aleutian 

Islands moss mounds substitute for trees in providing the 

elevated habitat characteristic of this moss throughout 

its range.

MNIACBAE

Mnium glabrescens Kindb. (See Persson, 1968.) 72616, 4e; 

7388, lOb; 7402, 8a; 7483, 4a; 7524, 4f; 758(7, 4f; 7528 

4f; 7862, 4e; 8264, 4e; 8284, 6a. A common species on many 

different substrates throughout the island.

Mnium insigne Mitt. 7261o, 4g; 8288, 4f. Foun-l on stony 

soil of a frost scar hi the Empetrum heath and on a shaded 

peat bank.

Mnium pseudopunwtatum Bruch & Schimp. (See Persson, 

1968.) 7196, 4e; 7203, 4g; 7260, 4e; 7458, 5a. F<nnd only at 

low altitudes, usually on very wet soil and in bogs.

ORTHOTBICHACEAB

Ulota phyttantha Brid. 7271, 3a; 7273, 3c; 7364, 2c; 7507, 

3h; 7638, 3b; 7911, 3b; 8024, 4d. Found on rocks near 

the sea, and, at one location, on an organic bird-perch 

mound. A facultative halophyte.

PLAGIOTHBCIACEAE

Plagiothecium roeseanum B.S.G. 7353, 2b. Found only on 

sandy banks above the beach at Cyril Cove, growing with 

Eurhynchium praelongum, Lophocolea cusfMata, and 

Pohlia cruda.

Plagiothecium undulatnm (Hedw.) B.S.G. 7227, 4a; 8239, 

13b; 8273, cf., 4f. Occurs sparingly as single strands in
PLANT SPECIES OF AMCHITKA ISLAND 47

the moss mat of the Empetrum heath and in alpine 

meadows.

POLYTRICHACEAE

Oligotrichum Jiercynicum (Hedw.) Lam. & DC. 8240, 13a. 

Found on mineral soil in the bed of a stream that flowed 

from a snow field.

Oligotrichum parattelum (Mitt.) Kindb. 823%, 13a. Found 

only on soil by a stream that flowed from a snow bed at 

Chitka Cove.

Poffonatum alpinum (Hedw.) Rohl. 1229, 4a: 7252, 14b; 

1311, 4a; 7384, cf., 4f; 7402, 8a ; 7425, 4b ; 7523, 4f; 7841, 

4f; 7850, cf., 4a; 8006, cf., 14b; 8257, 4f. Found on low 

terraces of the Empetrum heath, usually growing in the 

thick moss mat, and on high-altitude bedrock outcrops. 

At one location it grew with Marchantia polymorpha on 

the peat bottom of a drained pond.

Pogonatum contortum (Brid.) Loeske. (See Persson, 1968.) 

7206, 4f. Found only on the vertical walls of a narrow 

deep trench that had been dug in peat during World II.

Poffonatum urnigerum (Hedw.) P. Beauv. 7261, 14a; 7265, 

4f; 7300, lOa; 7355, 2b; 7356, 2c; 7385, lOa; 7481, lOa; 

7672, 4f; 8201, 5b; 8202, 5b; 8213, lOa; 8214, lOa. This 

moss commonly grows on disturbed soil and peat banks 

(flg.2).

Polytrichum commune Hedw. 7332, 4c; 7377, 4f. On moss 

mounds and peat banks of the low plateaus.

Polytrichum formosum Hedw. 7241, 4f; 7299, 4f; 7381, cf., 

lOc. On peat banks and broken macadam pavement.

Polytrichum juniperinum Hedw. 7374,14a. Apparently rare; 

found only on an andesite outcrop at an altitude of about 

400 feet.

Polytrichum longisetum Brid. [P. yracile Sm.]. 8284, 6a. 

Found only in wet moss-lichen mats on the lower terrace 

at Cyril Cove, with Drepanocladus flultans. 

**Polytrichum sphaerothecium (Besch.) Broth. [Pogonatum 

sphaerothecium Besch.]. (See Persson, 1968.) 7529, 4f. 

This is the first collection of this moss in Alaska, and 

apparently the second collection in North America. Persson 

(1968) made the observation that this moss, in its worldwide distribution, is often associated with Bryoxiphium 

norvegicum and that these mosses are found only in nonglaciated regions of the world.

Polytrichum strictum Menz. ex Brid. 7335, 4c. Found only 

on a small moss mound.

POTTIACEAE

Barbula cylindrica (Tayl. ex Mack.) Schimp. ex Boul. 764^, 

3b;-7642, 3b; 7644, 3b. Found only on breccia cliffs of the 

seacoast.

Bryoerythrophyllum recurvirostrum (Hedw.) Chen. 7671, 

4f; 8270, 9a; 8276, 3a. Found growing with Anomobryum 

concinnatum, Pohlia nutans, Pogonatum urnigerum, and 

Dicranella subulata on a peat bank at Constantine Harbor 

and in a spring on a deposit of limonite, where it was 

heavily incrusted with iron compounds.

Pottict heimii (Hedw.) Fiirur. ex Hampe sens, lat, 7184, 

3b; 8203a., 5b; 8204, 3b. Found growing with Amphidium 

lapponicum on breccia sea cliffs at Cyril Cove and on 

sandy soil of a borrow pit near Makarius Bay, associated 

with Philonotis americana. It also grew on turf banks at 

Makarius Bay, associated with Aniblystegium serpens, 

Leptobryum pyriforme, and Brytim stenotrichum.

*Tortula mucronifolia Schwaegr. (See Persson, 1968.) 75SV, 

2b; 7531, 2b; 7532, 2b; 8247, 3b. In loose sand of dun?s 

near the coast and on breccia sea cliffs.

RHYTIDIACEAE

Rhytidiadelphus loreus (Hedw.) Warnst. 7204, 4g; 7228, 4a;

7232, 4a; 7308, 4a; 7327, 4c; 7664, 6a. Abundant, especially

in wet locations in the Empetrum heath. 

Rhytidiadelphus squarrosus (Hedw.) Warnst. 7344, 2Ii;

7863, 13a. On sandy soil near the seacoast, growing among

dense stools of Elymus, and in a snow-bed community. 

Rhytidiadelphus triquetrus (Hedw.) Warnst. 7309, 4a. A

common species in the moss mats on the low plateaus.

SPLAOHNACEAE

Tetraplodon mnioides (Hedw.) B.S.G. 75^4, 15a; 7880, 15a; 

7968, 15a; 8242, 15a. Entirely restricted to decomposed 

carcasses of birds and rats; it forms dense polsters tint 

fruit profusely.

THUIDIAOEAE

Claopodium crispifolium (Hook.) Ren. & Card. 8274, ^eFound only on soil at the margin of a stream in the heath 

near Cyril Cove.

ANNOTATED LIST OF LICHENS

By WILLIAM A. WEBER,' JAMES A. EBDMAN, and HILDUR KEO?*

Shown below is the only published list of lichens that grow on 

Amchitka Island, and all species on the list are new records for 

this island, unless otherwise noted. To our knowledge, the only 

former Amchitkan collections of significance are those made 

by Eric Hulten on July 9, 1932, and those of George A. Llano 

made on August 22, 1947. Inventories of lichens collected frcm 

the Aleutian Island chain were published by Hedrick (1936) and 

Degelius (1937).

Duplicates of specimens cited in this list that were collect?d 

by Hansford T. Shacklette in November and December 1965 

were identified by Krog. All subsequent collections were identified by Weber, except those of Stereocaulon determined by I. 

Mackenzie Lamb, Harvard University, Cambridge, Mass.; several packets of Umbilicaria sent to George A. Llano, National 

Science Foundation, Washington, D.C.; and Parmeliopsis amMgua identified by Mason E. Hale, Jr., Smithsonian Institution, 

Washington, D.O. Identification of the rocks on which the 

saxicolous lichens grew was made by W. D. Quinlivan, U.S. 

Geological Survey.

Nomenclature follows that of Hale and Culberson (1966) insofar as possible. Collection numbers, in italics, are those of 

Shacklette, unless otherwise specified; they follow the species 

name. The habitat and plant community from which a species 

was taken, designated by numbers and letters, as set forth 

in the section "Description of the Island and Its Vegetation" of 

this report, follow the collection number.

CLADONIACEAE

Baeomyces placophyllus Ach. Brdma/n 646, 15b. On a wood°n 

sill of a World War II hut above Constantine Harbor.

Cladonia alpestris (L.) Rabenh. Hutchison s.n.; 8023, 4d; 

Erdman 613, 12b. Hutchison (1937, p. 144-145) reported:

3 University of Colorado Museum, Boulder, Colo. 

 * Botanical Museum, Oslo, Norway.
48 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

''The yellow-grey Cladonia alpestris, one of the reindeer 

mosses, formed the groundwork of the moors [of Amchitka], into which the foot crunched as if walking on 

snow. It was mixed with empetrum * * *." She undoubtedly was referring to the dominant Cladonia pacifica, recently described by Ahti (1961) in his treatment 

of the reindeer-lichen complex. Cladonia alpestris occurs 

widely scattered over the heath mat, often in patches, but 

it is less abundant than C. pacifica.

Cladonia amaurocraea (F16rke) Schaer. Erdman 653, 4c; 

Erdman 666, Sc. Uncommon ; found on a few moss mounds 

and at the edge of a tundra pool.

Cladonia arbiiscula subsp. Tjeringiana Ahti [C. sylvatica 

(L.) Hoffm.]. St evens s.n.; 7539, 4c. On the summit of a 

tundra mound above Cyril Cove. Merrill (1929, p. 42) 

stated that "D. H. Stevens found the species [C. sylvatica 

subsp. sylvestris Oed.] at Amehitka and Ilac Islands, 

and it has come to hand from Tanana crossing." No habitat was given.

Cladonia bellidiflora (Aeh.) Schaer. 7245, 4c ; 7332, 4c; 1513, 

4a; 7596, cf., 14b; 7602, 14b; 8012, 4a; Erdman 595, 8b. 

Widespread but not abundant; found in a variety of 

habitats.

Cladonia coccifera (L.) Willd. 7301, 15d; 7510, 4a; 7511, 

4a; 7512, 4a; Erdman 620, 15d. Fairly uncommon; however, it is more abundant and conspicuous in burned 

areas than elsewhere.

Cladonia degenerans (Fl6rke) Spreng. [C. cerasphora 

Vain.]. 7303, 15d; Erdman 608, in part, 4f. Uncommon 

throughout the Empetrum- heath.

Cladonia gracilis (L.) Willd. 7302, 15d; 7505, 4a; 7669, 4f; 

Erdman 608, in part, 4f; Erdman 609, in part, 4f; Erdman 

617, 15d ; Erdnian 618, 15d. Occurs in a variety of habitats 

on the low plateaus, but is common only in burned areas.

Cladonia pacifica Ahti [C, impexa Harm.]. 7210, 4c ; 7323, 

4c ; Erdman 568, 4a. The dominant lichen in the Empetrum 

heath; forms pure clumps in surface depressions, thus 

giving a dotted appearance to the ground surface as seen 

from the air (fig. 8).

Cladonia puxidata (L.) Hoffm. 7290, lOc; Erdman 619-A, 

15d. Sparse, except in burned areas, where it is abundant.

Cladonia rangiferina (L.) G. H. Web. ex Wigg. Erdman 

582, 13b. Observed only on a thick Empetrum mat in a 

small valley above Chitka Cove; apparently uncommon.

Cladonia scabriuscula (Del. ex Duby) Leight. 7211, 4c; 7324, 

4c; Erdman 624, 4d. A component species of moss mounds.

LECANORACEAE

HaemaJomma ventosum subsp. lapponicum (Has.) Laund. 

[H. lapponicum Has.]. Erdman 586, 14b. On a high-altitude rock outcrop.

Icmadopliila, ericetorum (L.) Zahlbr. Erdman 633, in part, 

4d. At the base of an organic bird-perch mound.

lonaspis epulotioa var. arctica (Lynge) H. Magn. Erdman 

639, in part, 14a; Erdman 645, 4g. On a latite boulder 

above Constantine Harbor and on basalt andesite scoria 

rocks in a frost scar.

Lecanora aleutica. H. Magn. ex Hedrick. 7365, 2c; Erdman 

630-D, 3a. Common on boulders, primarily on those along 

the beaches. I. Mackenzie Lamb examined specimen 7365. 

which grew on a sand dune, and wrote (written commun., 

1968), "Apparently a Lecanora species that has gone

from a crustose to semi-fruticose growth due to sand

abrasion and deposition." 

Lecanora allophana (Ach.) Rohl. [L. subfusca (L.) Ach.,

in part]. Erdman 632, 15b. On a small plank in the margin

of a wet sedge meadow. 

Lecanora atra (Hnds.) Ach. Erdman 641, 2c. C ~\ a small

'boulder in a sand-dune blowout. 

Lecanora oasta<nea (Hepp) Th. Fr. Erdman 67f, 4e. With

bryophytes in a streamside community. 

Lecanora straminea (Wahlenb.) Ach. Erdman 630-B, 3g.

On andesite beach rocks. 

OchrolecMa friffida (Sw.) Lynge. Hulten s.n. (Degelius,

1937, p 126) ; 7277, 3d; 7555, 14a; 7597, 14b; 7895, 12a;

7898, 12a; Erdman 565, 4a; Erdman 566, 4a; Erdman

610, lla; Erdman 663, 14b; Erdman 673, lOc. Occurs in

a wide range of habitats, at places overgrowing the heath

mat. Degelius (1937, p. 126) stated that it "seems to be a

very common species on these [Aleutian] islands." This

taxon is highly polymorphic. 

Placopsis gelida (L.) Linds. [Lecanora gelida (L.) Ach.].

Hulten s.n. (Degelius, 1937, p. 125) ; 7£97, 14a; Erdman

596, lOc. Common, especially on loose cobb^s of old

roadbeds.

LECIDEACEAE

Bacidea sp. Erdman 643, 14a. On andesitic tuff breccia rocks.

Lecidea cf. aleutica Degel. Erdman 629, in part, 14a. On 

andesite sill or flow boulders.

Lecidea armeniaca (DC.) Fr. Erdman 592, 14b. Common 

on high-altitude basaltic rocks above Chitka Cove.

Lecidea elegantior H. Magn. Erdman 589, 14b. On highaltitude basaltic rocks above Chitka Cove. Not listed by 

Hale and Culberson (1966) ; first reported for North 

America by Weber and Viereck (1967).

Lecidea flavocaerulescens Hornein. Erdman 587, 14b. Common on high-altitude basaltic rocks above Chitka Cove.

Lecidea macrocarpa (DC.) 'Steud. [L. platycarpa Ach.; 

L. steriza (Ach.) Vain.]. Erdman 590, 14b; Erdman 597, 

lOc ; Erdman 639, in part, 14a; Erdman 642, 2c. On breccia 

rocks, extending from sea level to high-altitud? fell-fields.

Lecidea cf. symmicta (Ach.) Ach. Erd-man 604, 15b. Found 

on the wind-eroded wooden frame of a World War II 

hut.

Lecidea vernalis (L.) Ach. Erdman 603, 15b ; cf. Erdman 667, 

4g. On tar-paper roofing of a hut at high altitude and in 

a frost scar of the Empetrum heath.

Mycoblastus alpinus (Fr.) Kernst. [M. sanguinarius (L.) 

Norm., in part]. 7597, 15b; Erdman 566, 4a; Erdman 611, 

lla; Erdman 652, 4c. Occasionally found intermixed with 

OchrolecMa frigida.

Rhisocarpon atroalbescens (Nyl.) Zahlbr. Erdman 591, 14b. 

Common on high-altitude basaltic rocks ab^ve Chitka 

Cove.

RMzocarpon geographicum (L.) DC. Hulten «.«. (Degelius, 

1937, p. 117) ; Erdman 661, 14b. On an andesite outcrop of 

a scree ridge above Chitka Cove.

RMzocarpon hochstetteri (Korb.) Vain. Hulten s.n. (Degelius, 1937, p. 117) ; Erdman 629, in part, 14a; Erdman 639, 

14a. Apparently common on rock outcrops throughout the 

island.

Toninia lobulata (Somin.) Lynge. Erdman 640, 41 This 

lichen was found covering the peat banks of an old gun 

emplacement.
PLANT SPECIES OF AMCHITKA ISLAND 49

PANNARIACEAB

Pannaria pezizoides (G. Web.) Trev. 7843, 4f; 7865, 4f;

Erdman 601, 4f. Collected from soil or peat banks at both

ends of the island. 

Psoroma hypnorum (Vahl) S. Gray. Erdman 633, in part,

4d. At the base of an organic bird-perch mound.

PARMELIACEAE

Cetraria ciliaris Ach. [C. orbata (Nyl.) Fink]. Erdman 572, 

15b. On a felled utility pole; possibly an adventive species 

introduced during World War II occupation.

Cetraria cucullata (Bell.) Ach. Erdman 656, 14a. Found 

near the summit of a large rock mound.

Cetraria ericetorum Opiz [C. crispa (Ach.) Nyl.; C. islandica var. crispa (Ach.) Nyl.]. Erdman 581, 13b; Erdman 

615, 14b. Observed and collected only above Chitka Cove; 

apparently uncommon.

Cetraria niaricans (Retz.) Nyl. 7897, 12a. In the rubble of a 

high-altitude rock field.

Hypogymnia enter omorplia (Ach.) Nyl. 7282, 15b. On a 

bridge timber near Cyril Cove; possibly introduced with 

construction materials.

Hypogymnia pJiysodes (L.) Nyl. [Parmelia physodes (L.) 

Ach.; P. duplicata var. doughlasiocla Gyeln.]. Erdman 

570, 15b. On a felled utility pole.

Hypogymnia subobscura (Vain.) Poelt [Parmelia subobscura Vain.]. 7897, 12a. In the rubble of a high-altitude 

rock field, mixed with several fruticose lichens.

Parmelia alpicola Th. Fr. Erdman. 601, 14b. On boulders 

at high altitudes.

Parmelia omphalodes (L.) Ach. 7603, 14b; 7888, 12a; Erdman 655, 4c. On rocks at high altitudes.

Parmelia saxatilis (L.) Ach. 7277«, 3c; Erdman 606, 15b; 

Erdman 621, 14a. On rock outcrops above or on the beach, 

and on wooden supports of a World War II shed.

Parmelia sulcata Tayl, 7275, 3c; Erdman 605, 15b; Erdman 

625, 14a. On breccia outcrops above or on the beach, 

and on wooden supports of a World War II shed.

Parmeliopsis ambigua (Wulf.) Nyl. Erdman 631, 15b. On 

a small plank at the edge of a wet sedge meadow.

Platysmatia lacunosa (Ach.) Culb. & Culb. [Cetraria lacunosa, Ach.]. 7601, I4b. On a rock mound above Chitka 

Oove.

PELTIGERACEAE

Peltigera apJitJwsa (L.) Willd. 7259, 4e; 7319, 4c; 7321, 4c; 

Erdman 638, 14a ; Erdman 651, lla. A fairly common terricolous lichen on shaded, often vertical, substrate surfaces.

Peltiaera canina (L.) Willd. [P. oanina var. membranacea 

Ach.; P. membranacea (Ach.) Nyl.]. Hulten s.n. (Degelius 

1937, p. 110) ; 7261c, 4e; Erdman 623, 4d; Erdman 634, 

4a; Erdman 669, 2b. A common terricolous lichen on 

shaded, often vertical, substrate surfaces.

Peltiaera malacea (Ach.) Funck. Erdman 635, 4a. At the 

base of a heath bank above Cyril Cove.

Peltiaera, spuria (Ach.) DC. [P. canina var. spuria (Ach.) 

Schaer.]. 7637, la. Found on the soil of a second beach 

terrace at Constantine Harbor.

PERTUSARIAOEAE

Pertusaria coriacea (Th. Fr.) Th. Fr. Erdman 

a flow breccia outcrop.

2, 14a. On

Pertusaria hultenii Erichs. Erdman 588, 14b. Saxicolous;

on basaltic outcrop above Chitka Cove. 

Pertusaria sp. Erdman 571, 15b. Corticolous; on a utility

pole.

PHYSCIACEAE

Buellia punotata (Hoff.) Mass. [B. pullata Tuck.], Erdman 

630-C, in part, 3a. On andesite beach boulders.

Physcia caesia (Hoffm.) Hampe. 7276, 3c; Erdman 630-C, 

in part, 3a. On sea cliffs and beach boulders.

Physcia dubia (Hoffm.) Lett. 7276, 3c. Found growing with 

P. caesia.

Rinodina turfacea (Wahlenb.) Korb. 7982, 4d; Erdman 671, 

4d. A nitrophilous lichen found on a few organic birdperch mounds.

SPHAEROPHORAGEAE

SphaeropJiorus globosus (Huds.) Vain. Hulten s.n. (legelius, 1937, p. 107) ; 7246, 4c ; 7320, 4c; 7595,14b ; 7599, 14b ; 

7897, 12a ; 7991, 14b; 8014, 4d ; 8015, 4d; Erdman 569, 4a; 

Erdman 583, 13b. Degelius (1937, p. 107) described this 

species as very common throughout the Aleutian Islands. 

It is the dominant lichen on the summits and ridge^ of 

the hummocks in the Empetrum heath.

S TEREOCAULAOEAE

Stereocaulon alpimim Laur. 7360, 2c. On a sand dune above 

Cyril Cove.

Stereocaulon intermedium (Sav.) H. Magn. Erdman 599, 

lOc. Common on a gravel roadbed south of Gonstartine 

Harbor. I. Mackenzie Lamb (written conimun., 1968) 

considered it to be "a stunted, subcrustose state of 8. 

intermedium."

Stereocaulon tomentosum var. alpestre Flot. [8. tom-entosum 

Fr.]. Erdman 600, lOc. Associated with 8. intermedium 

on a gravel roadbed south of Constantine Harbor.

Stereocaulon vcsuvianum Pers. [8. denudatum Plk.-; 8. 

vesuvianum var. denudatum (Florke) Lamb]. Erdman 

612, 12b. On upland slopes above the Pacific Ocean, opposite Chitka Cove.

STICTACEAE

Lobaria, linita (Ach.) Rabenh. [Sticta pulmonaria var. 

linita, (Ach.) Tuck.]. Hulten s.n. (Degelius, 1937, p. 108), 

Erdman 636, 4a. At the base of a heath bank above Cyril 

Cove.

TELOSCHI STAGE AE

Blastenia sp. Erdman 630-C, in part, 3a. On andesite beach 

boulders.

Caloplaca aranulosa (Mull. Arg.) Jatta. Erdman 594: 3a. 

A conspicuous saxicolous crustose lichen; common on 

supralittoral beach boulders and sea stacks.

Caloplaca sp. (Ferruginea group). Erdman 668, 2b. Or the 

end of a large driftwood log.

XantJwria, candelaria (L.) Th. Fr. [TeloscMstes ca*tdelariits (L.) Fink]. 7369, 4d; Erdman 627, 14a; Erf man 

630-C, in part, 3a; Erdman 649, 14a. On rock outcrops 

fertilized by birds.

Xanthoria elegans (Link) Th. Fr. [Caloplaca. elegans 

(Link) Th. Fr.]. Erdman 637, 3a. Occurs on beach boulders and sea cliffs. Tatewaki and Kobayashi (1934, p. 6) 

stated that "This saxicolous association is developed on 

such precipitous cliffs that the vascular plants cannot
50 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

grow. The beautiful red colour of Caloplaca elegans 

adorns the monotonous gloomy rock surface."

UMBILICABIACBAE

Umbilicaria cylindrioa (L.) Del. [Gyrophora cylindrica 

(L.) Ach.]. 7367, 14a; Erdman 628, 14a. On an andesite 

boulder pile near Cyril Cove. Although Llano (1950) cited 

no collections from Amchitka Island, he did state (p. 

11) that, in the Aleutian Islands, U. cylindrico, is "the 

dominating Umbilicaria on all exposed lava rocks, in 

association with U. proboscidea, U. hyperborea, and U. 

torrefacta * * *."

Umbilicaria proboscidea (L.) Schrad. 73670, 14a; 7506, 2c; 

7607, 14b; 7894, 12a; Erdman 585, 12a; Erdman 602, 

14b; Erdman 650, 14b; Erdman 660, 14a; Erdman 664, 

14a. Common on rocks in a wide variety of habitats.

USNEACBAE

Alectoria cf. irvingii Llano. 8013, 14b- Found on an andesite bird perch on a high peak north of Buoy Point. We 

have not compared this collection with the type from 

Anaktuvuk Pass, but it appears to agree well with the 

type description that is, it resembles Comicularia divergens, medulla Pd-, with long fissural pseudocyphellae.

Alectoria nigricans (Ach.) Nyl. 7596, 14b. Mixed with Cladonia bellidiflora at the base of an andesite rock mound 

south of Chitka Cove.

Alectoria oehroleuca (Hoffm.) Mass. 7193, 14a; 7368, 14a; 

7553, 14a; 7553, 14a; 7886, 12a; Erdman 567, 4a. A conspicuous fruticose lichen that apparently is of wide ecological amplitude, for it occurs as scattered clumps on 

breaks in the Empetrum heath, as well as on rocks ait the 

highest altitudes on the island.

Alectoria pubescens (L.) R. H. Howe [Parmelia lanata 

(L.) Wallr.]. 7371, 14a; 7857, 12a; Erdman 657, 14a. On 

rocks in outcrops or rubble fields, in many places mixed 

with other fruticose lichens.

Comicularia divergens Ach. [Alcctoria diver gens (Ach.) 

Nyl.]. 7247a 14a; 7598, 14b; Erdman 614, 14b; Erdman 

647, 4a; Erdman 654, 4c; Erdman 659, 14a. Widespread, 

but most common on rock mounds and outcrops.

Ramalina almquistii Vain. 7370, 14a; 7906, 3c; Erdman 626, 

14a. On breccia cliffs and inland outcrops.

Ramalina scoparia Vain. 7078, 3c; 7505, 3c. Commonly 

occurs with R. almquistii.

Siphula ceratites (Wahlenb.) FT. 7266, 8c; 7675, 8c.; Erdman 665, 8c. Forms dense colonies along the edges of 

small pools.

Thamnolia vermicularis (Sw.) Ach. ex Schaer. IT. subuliformis (Bhrh.) W. Culb.]. 7257, 14a; 7559, 14a; Erdman 672, 4a. Common throughout the island in a wide 

variety of habitats but is best developed in the uplands, 

where it forms pure stands in the eroded Empetrum 

heath. All material fluoresces under ultraviolet light.

VERRUCARIACEAE

Verrucaria maura Wahlenb. ex. Ach. Erdman 630-A, 3a. 

A dominant but inconspicuous lichen on the rocks of the 

beaches. In describing the plant communities of the 

Aleutian Islands, Tatewaki and Kobayashi (1934) placed 

Verrucaria in the supralittoral belt, which is just above 

the mean high-water level, where the plant is washed by 

occasional high tides. They stated (p. 6), "Verrucaria

maura, the calcivorous [sic] lichen, is distribrted over 

the supralittoral rocks, giving a dark aspect to this belt."

SOME ALGAE AND OTHER PROTISTS 

By H. SKUJA S

Collections of algae from three types of habitat on 

Amchitka Island were sent to me for identificat: on. The 

habitat descriptions and the annotated list, including 

descriptions of new taxa, follow.

The following taxa, found in a liverwort polster 

(mostly Marsupella emarginata) , were collected (No. 

7234} by H. T. Shacklette, November 22, 1965, from a 

small stream near Chitka Point:

CYANOPHYCEAE

Chroococcus varius A. Br. 

Gloeocapsa magma ( Breb.) Kiitz. 

ScMzothria caloicola (Ag.) Gom. 

ScMzothriae lardacea (Ces.) Gom. 

Scytonema mirabile (Dillw.) Born. 

Stigonema minutum (Ag.) Hass. 

Stigonema ocellatum Lyngb.

DESMIDIACEAE

Actinotaenium cucurbita (Breb.) Telling. Cells28/u-30/t long, 

13p-15p broad. (PI. 1, fig. 1.)

Cosmarium cyclicum Lund. var. arcticum Nordst.

Cosmarium decedens (Reinsch) Racib. Cells 45^-48/t long, 

24/4-26/t broad, and 17/X-19/H thick; breadth of isthmus 

18/X-20/*. (PI. 1, fig. 7.)

Cosmarium obliquum Nordst. Cells 20^-23/11 long, 17/i-18/t 

broad, and 13^-14/t thick; breadth of isthmur* 14/t-16jtt. 

(PI. 1, figs. 8-9.)

Cosmarium tatricum Racib. Cells 4lM-52p lonf, 23/i-27j«. 

broad, and 17/*-20/i thick; breadth of isthmus 18/t-21/t. 

(PI. 1, figs. 4-6.)

Cosmarium variolatum Lund. Cells 45^-52/1 long, 25/i-27/u 

broad, and 17/*-20/i thick; breadth of isthmus 12/t-15/i. 

(PI. 1, figs. 2, 3.)

Cylindrocystis brebissonii Menegh.

Cylindrocystis brebissonii var. minor W. et G. S West.

Cylindrocystis crassa De Bary.

Mesotaenium micrococcum Kiitz.

Staurastrum punctulatum Breb. var. pygmaeum (Breb.) 

W. et G. S. West. A quadrangular form. Length 36/t-40M, 

breadth 34/t-40/t; isthmus 18/i-20ju broad; cell wall uniformly granulate, granules minute and acute.

DIATOMACEAE

Achnanthes spp.

Caloneis silicula (Ehrnb.) Cl.

Epithemia turgida (Ehrnb.) Ktitz.

Eunotia arcus Ehrnb.

Eunotia gracilis (Ehrnb.) Rbh.

Eunotia lunaris (Ehrnb.) Grun. var. capitata Gr^n. Length

50/i-86/*, breadth 4/t. 

Eunotia praerupta Ehrnb. var. muscicola Boye-Pe4:.

5 Uppsala Universitets Institution fOr Systematisk Botanik, Uppsala, 

Sweden.
PLANT SPECIES OF AMCHITKA ISLAND 51

Eunotia praerupta var. bidens (W. Sm.) Grun. Cells 15/4-

75/4 long, 5/4-10/4 wide; striae 10-12 in 10/4. 

Eunotia robusta Ralfs var. diadema (Bhrnb.) Ralfs. 

Frustulia rhomboides (Bhrnb.) D.T. var. saxonica (Rbh.)

D.T. 

Gomphonema angustatum (Kiitz.) Rbh. var. prod<uctum

Grun. Length 38/4-45/4, breadth lO/i-12/4. 

Melosira distans (Bhrnb.) Kiitz. var. alpigena Grun.

Breadth 7/4-9/4, height of the whole cell 10/4-12/4; striae

obsolete, valvae conspicuously dotted. 

Pinnularia gibba Bhrnb. 

Pinnularia lata (Br6b.) W. Sm. Length 85/4-103/4, breadth

28/4-30/4; striae 4 in 10/t. 

Pinnularia microstauron (Bhrnb.) Cl. 

Pinnularia viridis (Nitzsch) Bhrnb. var. rupestris

(Hantzsch) Cl. 

Tabellaria flocculosa (Roth.) Kiitz.

DINOPHYCBAB

Gloeodinium montanum Klebs. Diameter of cells with integument 41/4-58/4, without integument 25/4-38/4; integument or cell wall thick and lamellose, colorless or yellowish or brownish; cell content brown; 2-celled aggregates 

with integument 57/4-60/4X60/4-85/4. (PI. 1, figs. 20-23.)

Peridinium cinctum (O.F.M.) Ehrnb.

The following taxa were found in a shallow pool on 

andesite bedrock at a quarry near Banjo Point, with 

Isoetes sp. (44 chromosomes). Collected by H. T. Shacklette, July 28,1967.

OYANOPHYCBAE

Aphanothece castagnei (Br6b.) Rbh.

Oscillatoria amoena (Kiitz.) Gom. Trichomes straight, with 

more or less attenuated and frequently curved ends, 4/4-6/4 

broad, not constricted or only slightly constricted at the 

crosswalls, and grayish blue green or olivaceous and 

pale violet; cells nearly quadrate or slightly shorter, or 

longer than broad, granulated at dissepiments; end cells 

capitate, with broadly conical apex. (PI. 2, fig. 25a-d.)

Oscillatoria- amphibia Ag. Trichomes nearly straight or 

coiled, 2/1-2.5/4 broad, ends nonattenuated and not capitate, 

not constricted at the joints, and pale blue green; cells 

4/4-7.5/4 long, with two granules at the septa; end cells 

rounded.

Synechococcus maior Schrot. Cells oblong or ellipsoidal with 

rounded apices, 19/4-21/t broad, and 24/4-34/4 long; contents 

homogeneous and blue green.

CHLOROPHYCBAB s.l.

Chlorella elUpsoidea Gerneck.

Chlorella vulgaris Beyerinck.

Cosmarium humile (Gay) Nordst.

Cosmarium laeve Rbh. forma. Cells 24/t-26/4 long, 17/4-18/4 

broad, and 1V-12/4 thick, with 4/4-5/t broad isthmus; certain specimens have one semicell of the type, the other 

one of var. septentrionale Wille. (PI. 1, fig. 13.)

Cosmarium margaritiferum Menegh.

Cosmarium meneghini Br6b. var. concinnum Rbh. Cells. 

25/4-27/t long, 19/4-20/t broad, and 13/4-14/4 thick; apex 

13/4-14/4, isthmus 4/4-5/4 broad; cell ends truncate or often

slightly convex, rarely conspicuously depressed. (PI. 1,

fig. 12.)

Cosmarium nitidulum De Not.

Cosmarium subcrenatum Hantzsch forma. Cells 30/4-33/4 

long, 25/4-26/4 broad, and 17.5/4-18.5/4 thick; isthmus 11/4- 

12/i broad; semicells subtrapezo-rectangular, with flattened 4-crenate ends and 4-5 crenae at each side; the low 

frontal swelling above the isthmus with generally 5, 

rarely 4 or 6, strong rows of 3-5 angular granules. 

(PI. 1, fig. 11.)

Cosmarium subpachydermum Schmidle forma. Cells 38/«^-41/4 

long, 28/4-32/4 broad, and 20/4-22/4 thick; isthmus 13/4-14/4 

broad; cell wall scrobiculate, but without a refractive 

thickening in the center of each semicelL (PL 1, fig. 14.) 

Compare also C. pseudonitidulum Nordst.

Cosmarium subundulatum Wille. Cells 60/4-65/4 long, 42ju--45/i 

broad, and 28/i thick; isthmus 20ju-22/i broad; semicells 

high semicircular with a 10-undulate margin; membrane 

distinctly punctato-scrobiculate with a large but low 

lenticular inflation in the center of each semicell. (PI. 1, 

fig. 10.)

Cylindrocystis brebissonii Menegh.

Cylindrocystis crassa De Bary.

Euastrum ansatum Ehrnb. Cells 78ju-87ju long, 39/4-44/4 wide, 

and 27/t-29ju thick; apex 20ju-22/4, isthmus 13/4-15ju broad.

Euastrum Udentatum Nag. Cells 53ju-61/t long, 24/4-26/t 

broad, and about 20ju thick; apex 22ft, isthmus 10/4-12ju 

broad.

Euastrum denticulatum (Kirchn.) Gay. Cells 33/4-35/t long, 

22ju-26/i broad, and 15ju-17ju thick; apex 16ju-20/4, isthmus 

5/4-6/4 broad.

Euastrum denticulatum var. quadrifarium Krieger. C^lls 

about 20/t long, 160 broad, and 10/t thick; apex 13/4, isthmus 

4/t broad.

Euastrum didelta Ralfs. Cells 125/i-135/t long, 64/t-66/t bread, 

and 37ju-39ju thick; apex 30ju-31ju, isthmus 20/t-21/4 broad.

Euastrum elegans (Br6b.) Kiitz. A typical form; c^lls 

30ju-32ju long, 20/1-21/4 broad, and 12/t thick; isthmus 5/t-6/t 

broad.

Euastrum oblongum (Grev.) Ralfs.

Euastrum pectinatum Br4b. Cells 74/t-76/t long, 46/t-48/t 

broad, and 34/4-35/t thick; width of the polar lobe 27/4-?9ji4, 

width of the isthmus 19/i-21/t.

Hormidium flaccidum A. Br. Filaments «=> 4/4-5/t broad, not 

constricted at the crosswalls; cells 2-3% times longer 

than wide; chloroplast a unilateral plate with a pyrenoid.

Mougeotia sp. ster.

Oocystis parva W. et G. S. West. Solitary or in families of 

2-4 cells, inclosed by the enlarged mother cell wall; c^lls 

ellipsoidal or fusiform with pointed poles, 3/i-7/t bread, 

7/4-10/t long; chloroplasts 1-2 parietal discs, usually without pyrenoids. (PI. 1, fig. 15.)

Pediastrum braunii Wartm. Coenobia 7- to 10-celled, cells 

10/4-16/t in diameter.

Scenedesmus acutus (Meyen) Chod.

Scenedesmus ecornis (Ralfs) Chod.

Scenedesmus spinosus Chod.

Scotiella nivalis (Shuttlew.) Fritsch. Cells solitary, 

ellipsoidal, 12/4-14/4 broad and 20/4-24/4 long; cell vail 

colorless, with 6-8 longitudinal ribbons or low wirgs; 

single parietal chloroplast with one pyrenoid.

Zygogonium ericetorum Kiitz.
52 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

DIATOMACEAE

Caloneis silicula var. longissima Schirschow. Cells linear,

with an evident widening in the middle, 80/i-85/j long and

lO/i-14/i broad; striae about 10 in 10/i.

Caloneis silicula var. alpina Cl. Cells oblong, with conspicuously triundulate sides, 32^-37/j long and 12/j-15,u

broad. 

Cymbella aequalis W. Sm. Cells 17^-25^ long and

broad; striae 13-15 in lO/i. 

Cymbella, angustata (W. Sm.) 01. 

Cymbella angustata var. hybrida (Grun.) R. Ross. 

Cymbella cuspidata Kiitz. Cells 50^-62^ long and 15

23/j broad; striae 10-13 in 10/j. 

Cymbella delicatula Kiitz. Cells 25/^-36^ long and

broad. 

Cymbella gracilis 01. Cells 56/i-60/i long and 8ju-10,u broad;

striae 12-13 in 10,u. 

Cymbella hebridica (Greg.) Grun. Cells 29/t-37/i long and

7{j.-8/ji broad; striae 10-12 in 10/i. 

Cymbella naviculiformis Anersw. Cells 38^-50^ long and

12ju-16,u, broad; striae 14-18 in 10/i. 

Cymbella perpusilla A. 01. Cells 16^-20^ long and

broad.

Cymbella pusilla Grun. 

Cymbella tumidula Grun. Cells 28/J-32/J long and

broad.

Cymbella ventricosa Kiitz. 

Eunotia diodon Ehrnb. Cells 52,u-70/j long and 10^-15^ wide ;

striae 12-14 in 10p. 

Eunotia pectinalis (Kiitz.) Rbh. var. minor (Kiitz.) Rbh.

Cells 10|U-17/j long and about 4/t wide ; striae 15-18 in 10/i. 

Eunotia praerupta var. bidens (W. Sm., Grun. Cells 15/J-75/J

long and S/i-lO/i wide; striae 10-12 in 10/t. 

Eunotia robusta Ralfs var. diadema (Ehrnb.) Ralfs. Cells

47,u-50ju long and 20^-21/1 broad, with 6-undulate dorsal

margin; striae about 10 in 10/t. 

Eunotia septentrionalis Ostr. Cells 200-23,11 long, 4^-5/1

broad; striae 17-18 in 10/i. 

Fragilaria construens (Ehrnb.) Grun. var. binodis (Ehrnb.)

Grun.

Fragilaria construens var. subsalina Hust. 

Fragilaria nitssohioides Grun. 

Frustulia rJiomboides var. saxonica (Rbh.) D.T. Cells 50^-

105/t long and 13,u-200 broad. 

Gomphonema angmtatum var. productum Grun. 

Hantzschia amp7iioxys (Ehrnb.) Grun. 

Melosira distans var. alpigena Grun. Breadth 6^i-10(H, height

of the whole cell 50-17,u; striae 16-18 in 100; valvae

conspicuously dotted.

Navicula contenta Grun. Cells 80-140 long and 20-30 broad. 

Navicula cryptocephala Kiitz. Cells 250-400 long and 50-80

broad; striae 16-18 in 10/i.

Navicula minima Grun. Cells 10^-16^ long and 3/i-4^ broad. 

Navicula minima var. atomoides (Grun.) 01. Cells lO^i-16/i

long and 3/t 4/t broad. 

Neidium afflne (Ehrnb.) 01. var. minus Cl. 38^-45^ long and

8^-10^1 broad; striae 20-23 in 10,u. 

Neidium iridis (Ehrnb.) 01. var. amphigomphus (Ehrnb.)

V.H. Cells 7(V-130/* long and 28^-33^ broad; striae 16-18

in WfjL.

Nitsscliiapalea (Kiitz) W. Sm. 

Nitsschia stagnorum Rbh.

Pinnularia appendiculata (Ag.) Cl. Cells 28/4-33^ long and

4/1-6/1 broad; striae about 16 in 10/i. 

Pinnularia brevicostata Cl. Cells 83/i-92/i long and1 13^-15^

broad; striae 8-9 in 10/j. 

Pinnularia divergens W. Sm. Cells 58/i-66(U long anc1 15/i-17/i

broad; striae 10-12 in 10/*. 

Pinnularia gentilis (Donk.) Cl. Cells 200ju-205/i long, 34/i36/i broad; striae 6-7 in 10/i. 

Pinnularia gibba Ehrnb. Cells 65/I-90/I long, lO/i-lSa broad;

striae about 10 in 10/i. 

Pinnularia interrupta W. Sm. 

Pinnularia lata (Breb.) W. Sm. Cells 80fj.-85fi long and 30/i33/* broad; striae 3 4 in lO^u. 

Pinnulariamaior (Kiitz.) Cl. 

Pinnularia microstauron (Ehrnb.) Cl. Cells 34^-72/i long

and Sju-ISM broad; striae 10-12 in 10/i. 

Pinnularia microstauron forma biundulata O. MiHl. Cells

S-ifi-72/j. long and 8/*-13/i broad; striae 10-12 in IQu. 

Pinnularia viridis (Nitzsch) Ehrnb. 

Pinnularia viridis var. sudetica (Hilse) Hust. 

Stauroneis anceps Ehrnb. Cells 48/*-57/* long and 10/*-13)U

broad; striae about 20-23 in 10/i. 

Stauroneis anceps forma linearis (Ehrnb.) Cl.

CHRYSOPHYCEAE

(Only cysts were found.)

The following taxa were found in a tundra lake about 

50 cm deep near Cyril Bay, with Isoetes sp. (66 chromosomes) . Collected by H. T. Shacklette, July 28,1967.

CYANOPHYCEAE

Lyngbya rivulariarum Gom. 

Merismopedia elegans A. Br. 

Merismopedia glauca (Ehrb.) Nag. 

Nostoc microscopicum Carm. 

Oscillatoria amoena (Kiitz.) Gom. 

Oscillatoria amphibia Ag. 

Oscillatoria facilis Skuja, sp. nov.

Trichomata solitaria vel in strata expansa, tenuia, 

arachnoidea, olivacea vel sordide fusco-viridia agglomerata, 

aut plusminusve recta aut subflexuosa et elongata, ad genicula leniter constricta, apice breve attenuata e4; paullo 

curvata, l/u-1.3/u crassa; cellulis diametro trichomatis 2-4 

plo longioribus, protoplasto homogeneo sed cum cl romatoplasmate a centroplasmate sat limitato, pallide griseoaerugineo vel olivaceo et subfusco; cellula apicali elongate 

conoidea, superne parum obtusata. (PL 2, fig. 26a-e.)

Trichomes single or forming a flat arachnoid, ol'vaceous 

or brownish-green thallus, more or less straight or slightly 

flexuous, slightly constricted at the cross-walls, l/j-1.3/* 

broad, and short at the ends but clearly attenuated and 

curved; cells 2-4 times as long as wide, with a homogeneous, 

pale-grayish-blue-green or olivaceous and brownish protoplast ; end cell elongate, conoid with slightly obtusa te apex, 

not capitate and without a calyptra. (PI. 2, flg. 26«-e.)

This species resembles O. neglecta Lemm. but 1 as both 

curved and attenuated apices, longer cells, and a different 

color of thallus and trlchomes; compare also O. exit's Skuja 

(Skuja, 1964, p. 51, table 7, figs. 1-3), which is considerably 

thicker, more constricted at the crosswalls, and of a different 

color. Also different are O. deflesea W. et G. S. West, O. subtilissima Kiitz., O. limnetica Lemm. with var. acicularis 

Nygaard, and others.
PLANT SPECIES OF AMCHITKA ISLAND

Oscillatoria grossegranulata Skuja.

Trichomes solitary among other filamentous algae, elongate, nearly straight or slightly flexuous, not attenuated or 

curved at the ends, and distinctly constricted at the crosswalls, thus torulose, 5.5/J-6/* broad; cells cylindrical, 1-1% 

times longer than wide, 6/^-12^, rarely 16/*, long with clearand pale-olivaceous or light-grayish-blue-green coarsely 

granulated protoplast, and at times with granulated septa. 

(PI. 2, fig. 28.) 

Oscillatoria splendida Grev.

A very characteristic species, with straight or flexuous 

trichomes 1.7/*-2/j broad which usually are not constricted at 

the crosswalls but are rather short and abruptly attenuated 

at the more or less curved ends; cells 2^1 times longer than 

broad, with often granulated septa and a clear blue-green 

homogeneous protoplast; end cell very elongated and 

pointed, more or less bent, capitate at the apex, without 

calyptra. (PI. 2, fig. 27.)

The thallus of this alga also has a characteristic intense 

odor which resembles that of fertile soil. 

Oscillatoria, sp.

Trichomes nearly straight or slightly bent, unconstricted 

at the joints and not attenuated or capitate at the ends, 

0.5/4-0.7/* broad; cells mostly 2-4 times, rarely 5 times, as 

long as wide, with a homogeneous pale-blue-green protoplast 

but with 2 granules at the dissepiments; end cell simply 

rounded.

Only a few trichomes seen; probably a small form of 

0. amphibia. (PL 2, fig. 29.)

Phormidium frigidiim F. E. Fritsch. 

Pseudanabaena catenata Lauterb. 

Pseudanabaena granulifera Skuja, sp. nov.

Trichomata solitaria, inter algas benthonicas sparsa, sat 

brevia, plerumque 6-20-cellularia, 15/^-50^ longa, fere recta 

vel leviter arcuata, 1.5ju-1.8/i crassa, apices versus non 

attenuata, ad dissepimenta hyalina sat crassa evidenter 

constricta, motu proprio praedita; cellulis 1-1% plo raro 

2% plo longioribus quam latis, breve cylindricis, lateribus 

baud concavis; protoplasto pallide olivaceo vel dilute 

viridi-aerugineo, granulato, granulis sat grossis praecipue 

peripheriter locatis; cellula apicali superne late conoidea. 

(PL 2, figs. 30-32.)

Trichomes solitary among other benthonic algae, short, 

usually 6- to 20-celled and 15/j-SO/j long, mobile, nearly 

straight or slightly curved, 1.5ju-1.8/i broad, not attenuated 

at the ends, distinctly constricted at the joints, dissepiments 

hyaline, rather thick ; cells short-cylindrical l-l%-2% times 

longer than wide, with pale olivaceous or light-greenish-blue 

coarsely granulated protoplast, granules mostly peripheral ; 

end cell at apex conoid. (PI. 2, figs. 30-32.)

This species perhaps resembles most nearly P. minuta 

Skuja forma (Skuja, 1956, p. 70, table 6, figs. 27-28), from 

which it is distinguished by its considerably shorter and 

smaller trichomes and differently colored, coarsely granulated protoplast. The typical P. minuta (Skuja, 1948, p. 57, 

table 5, figs. 6-7) is also closely allied but is broader and 

has more rounded cells usually with homogeneous or only 

finely granulated protoplast. Compare further P. galeata 

Bocher and P. bipes Bocher.

Synechococcus cedrontm Sauv. var. pallidus Skuja var. nov.

Cellulae solitariae, motu proprio praeditae, singulae vel

geminatae, breve cylindricae vel ellipsoideae, apicibus

rotundatis, 2.5^-3/1 raro 3.5/i latae et 3.5^-6^ longae; membrana tenuissima, perspicua; contentu cellularum pallidissime aerugineo, fere achroo, homogeneo vel granulis subtilis, 

sparsis ornato; multiplicatio fit divisione transversal!. 

(PL 2, fig. 24.)

A forma genuina cellulis paullo minoribus, colore prllidissimo primum differt.

Cells solitary, mobile, single or two together after division, 

short cylindrical or ellipsoidal with rounded apices, 2.5;*- 

3.5/i broad and 3.'5ju-6/i long; membrane delicate and hyaline; 

contents very pale blue green, almost colorless, homogeneous 

or with some minute granules; multiplication by transverse 

division. (PL 2, fig. 24.)

This variety is well characterized by its smaller size and 

the very pale protoplast.

Compare also S. notatus Skuja forma (Skuja, 1964, p. 24, 

table 1, figs. 39-40), which is considerably larger, entirely 

colorless, and with a differently constituted protoplast. 

Synechococcus maior Schrot.

CHLOROPHYCEAE s.l.

Ankistrod&smus braunii (Nag.) Collins var.

Cells may be fusiform but usually slightly curved or 

lunate with more or less acutely pointed ends, solitary or 

after division (forming of autospores) in pairs or in forrs 

contacting each other, 7/i-14/i long, 1/^-2^ broad; cell wall 

thin, smooth, and colorless; chloroplast single, parietal, 

usually unilateral and with a small rounded medial excavation, without a pyrenoid. (PL 2, fig. 49.)

Only a few cells were seen. Considerably smaller than the 

typical A. braunii, but otherwise similar; compare w'th 

Vischer's (1920) description. Perhaps also it is a relative of 

some Koliella species (Hindak, 1963). 

Ankistrodesnuts spiralis (Turner) Lemm. 

Chaetoplaca nom. nov. genericum. Syn. Chaetopedia Skuja

1948 non Pascher 1939. Fain. Chaetoplaeaceae nom. nov.

fam. 

Chaetoplaca crassiseta Skuja var. puclla Skuja (Skuja,

1964, p. 110, table 16, figs. 29-33). Has characteristic 4-

and 8-celled colonies. 

Chlamydomionas alaskensis Skuja, sp. nov.

Cellulae rotundato ellipsoidales et ovoideae vel interdum 

fere globosae, 12^-18^-21(1 longae, 10/j-15/*-l9/i latae, 

fiagellis binis aut cellula e aequilongis aut paullo brevioril 'is 

praed-tae; membrana modice crassa, plerumque distin"te 

porosa raro fere laevi, achroa, papilla antica parva n^nnunquam vix evoluta instructa; chromatophoro parietal! 

crasso, bursaeformi, multipartite sed cum partibus inter se 

dense contiguis, pyrenoide destitutis; stigmate fuscorutro, 

med-ocri, breve baculiformi in parte anteriore; vacuolis 

contractilibus binis, parvis in polo antico ad basin flagellorum, nucleo nucleolato media in cellula sito. Multiplies tio 

divisione protoplast! cellulae matricali in duas vel frequenter quattuor cellulas filiales. (PL 2, figs. 33^11.)

Cells rounded ellipsoidal or sometimes almost globose, 

12ju-18/i-2V long and 10^-15/^-19^ wide, with two fiagella 

which are as long as or slightly shorter than the cell; cell 

wall moderately thick, distinctly porous or almost smooth, 

colorless, with anterior semicircular small papilla which 

sometimes are entirely lacking; chloroplast parietal, thick 

but divided into a number of separate closely joined bod! <?s, 

without a pyrenoid ; eyespot brownish red, small, short sublinear, in the anterior half of the cell; contractile valuoles 

two, small, at the base of the flagella; nucleus more or less
VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

centrally placed. Asexual reproduction by the division of 

the cell protoplast usually into 4 or more, rarely 2, daughter 

cells or zoospores. (PI. 2, figs. 33-41.)

C. alasTcensis is in the subgenus Chloromonas. This species 

is similar to C. vulgaris Anachin but is considerably smaller 

and has a porous cell wall, a smaller papilla, and a higher 

placed, shorter eyespot. It is also somewhat like C. jemtlandica Skuja but differs in being almost twice as large 

and in having a porous cell wall, shorter flagella, and a 

higher placed sublinear eyespot. Should also be compared 

with some other chloromonads. 

Chlamydomonas subangulosa. Skuja sp. nov.

Cellulae subanguloso-ovales vel late-ellipsoideae, 13/i-16ju 

longae, 9/i-14/* latae, flagellis binis cellulae circiter aequilongis vel paullo brevioribus instructae ; membrana delicata, 

achroa levisve, papilla antica mediocri hemisphaerica vel 

truncato-rotundata praedita ; chromatophoro poculiformi in 

parte anteriore extus longitudinaliter striato-carinato, parte 

postriore valde incrassato et pyrenoide depresso-globoso sat 

rnagno donato; stigma te fuscorubro mediocri, rotundo vel 

elliptico in parte anteriore; vacuolis contractilibus binis in 

polo antico ad basin flagellorum locatis; nucleo nucleolato 

in excavatione chromatophori sito. Multiplicatio divisione 

protoplast! cellulae maternae plerumque in 4 cellulas flliales. 

(PI. 2, figs. 42-46.)

Cells slightly rectangular or broad-ellipsoidal, 13ju-16ju 

long and 9/t-14/* wide, with paired flagella which are as 

long as, or slightly shorter than, the cell; cell wall thin, 

smooth and colorless, with anterior semicircular mediumsized papilla; chloroplast cup shaped, striped on the outside 

of the anterior part, the posterior portion massive and with 

one rather large pyrenoid; eyespot brownish red, small, at 

the anterior end of the chloroplast. Asexual reproduction 

by division of the cell protoplast to form 4 zoospores or 

daughter cells. (PL 2, figs. 42-46.)

This species is in the subgenus Euchlamydomon-as, and 

is in some respects most similar to C. angulosa Dill but 

differs in its smaller size, its only medium-sized papilla, 

and the longitudinally-striped chloroplast on the outside, 

punctiforrn stigma, and the not angular but more or less 

rounded smaller pyrenoid. C. leptobasis Skuja is somewhat 

similar, but it is smaller, more rounded-ovoid, with longer 

flagella, and it has a thinner, simple cup-shaped chloroplast 

with a small globose pyrenoid. C. subangulosa should also 

be compared with C. saxonensis Skuja and with some other 

species of the subgenus Euchlamydomonas. 

Chlorococcum humicola (Nag.) Rbh.

Typical form. Cells globose, solitary or often gregarious 

in small amorphous clumps, variable in size within the same 

aggregation namely 7^-20^-25^ in diameter; cell wall thin 

or moderately thick, colorless and smooth but in an older 

state sometimes more or less evidently lamellose; chloroplast cup shaped, thin, covering the whole wall or nearly 

so, with one pyrenoid in a thickened part of the plastid. 

Reproduction by the forming of usually 8-16, rarely as many 

as 32, aplanospores in a mother cell. (PI. 1, figs. 16-19.) 

Closterium acutum Breb. Typical form; cells lOO/i-123/i

long and 4/*-4.5/t broad. 

Euastrum denticulatum (Kirchn.) Gay. 

Euastrum elegans (Br6b.) Kiitz. 

Gloeocystis rupestris (Lyngb.) Rbh. Compare also Skuja

(1964, p. 106, table 15, fig. 11.)

Hortnidium flaccidum A. Br.

Mougeotia sp. ster. .

Oedogonium sp. ster.

Pedict strum braunii Wartm.

Planctococcus sphaerocystiformis Korschik.

Free-floating spherical colonies, 25ju-30/i in dirmeter; the 

gelantinous nonlamellate and colorless envelope relatively 

wide, including a central group of 4 globose cells, 9/j-12ju in 

diameter. Cells have a thin smooth wall and a thin or only 

moderately thick parietal multipartite chloroplast covering 

most of the wall; and lack a pyrenoid. (PI. 2, fig. 47.)

Probably not a separate intraspeciflc taxon but only a 

4-celled form of P. sphaerocystiformis. 

Raphidonema gracile Skuja, sp. nov.

Cellulae solitariae vel post divisionem factern binatae, 

elongatae et cylindraceo-fusiformes, 1.7ju-2.3ju latae, 12^-25/u, 

longae, apicibus subito angustatis, polls subacuto rotundatis, 

plerumque plusminusve curvatis; membrana tenui, achroa 

laevisve; chromatophoro parietali, plusminusve unilateral!, 

sine pyrenoide sed cum excavatione ventral! med'a in cellula 

ubi nucleus locatus. Propagatio fit cellularum divisione in 

duas partes. (PI. 2, fig. 48o-f.)

Cells free floating, solitary, or in pairs after division, 

elongate fusiform to somewhat cylindrical, with slightly 

tapering and subacute rounded ends, usually more or less 

curved, 1.7ju-2.3ju broad and 12/A-25//. long; cell wall thin, 

smooth, and colorless ; chloroplast single, parietal, and more 

or less unilateral, without a pyrenoid but commonly with a 

rounded ventral sinus in the middle where the nucleus is 

located. Reproduction by a transverse division into two 

daughter cells that remain connected for some time. (PI. 2, 

flg.48«-/.)

This form perhaps most nearly resembles F.oliella corcontica Hindak (Hindak, 1963, p. 107, table 2 (20), fig. 1), 

although it differs considerably because of its smaller and 

less cylindrical cells. I believe now that the rnair differences 

between Koliella Hindak and Raphidonema Lagerheim are 

not of a general but of an entirely gradual character and, 

therefore, of only secondary value taxonomically. Compare 

in this respect also the key to both genera by Hindak (1963, 

p. 99) and the figures in his plates 1-6, figures 19-24). 

Scenedesmus aoutus (Meyen) Chod. 

Scenedesmus armatus (Chod.) G. M. Smith 

Scenedesmus brevispina (G. M. Smith) Chod. 

Scenedesmtis ecornis (Ralfs) Chod. 

Scenedesmus quadricauda (Turp.) Breb. 

Scenedesmus quadricauda var. maximus W. et G. S. West 

Scenedesmus spinosiis Chod. 

Sphaerozosma granulatum Roy et Biss. 

Staurastrum margaritacetim (Ehrnb.) Menegh. Granulation

of the cell wall more or less reduced.

EUGLENOPHYCEAE

Anisonema acinus Duj.

Euglena pisc-ifonnis Klebs.

Euglena viridis Ehrnb. var. olivacea Klebs.

Menoidium incur vum (Fres.) Klebs.

CHRYSOPHYCEAE

Bodo minimus Klebs. 

Honas uniguttata Skuja. 

Pleuromonas jaculans Perty.
PLANT SPECIES OF AMCHITKA ISLAND 55

DIATOMACEAE

Achnanthes minutissima Ktitz.

Cyml) lla austriaca Grun.

Cynibelld delicatula Kiitz.

Cymbellahebridica (Greg.) Grun.

Cymbella ventricosa Kiitz.

Fragilaria construens var. Mnodis (Ehrnb.) Grun.

Melosira distans var. pfaffiana (Beinsch) Grun.

Neidium duMum (Ehrnb.) Cl. Cells 40/i-48/i long, 10/i-12/t

broad. 

Neidium iridis var. ampMffomphus (Ehrnb.) V. H. Cells

80^-120/1 long, 27/i-30/i broad. 

Nitzschia gracilis Hantzsch. 

Pinnularia cardinaliculus Cl. A slightly larger form length

of the cells 100^-115/1*, breadth 21/t-22/i; striae 9-10 in 10/j. 

Pinnular'M dactylus Ehrnb. Length of the cells 230/i-250/tt,

breadth 46/1-48/u,; striae 5-6 in 10/t. 

Pinnularia gentilis (Donk.) Cl. 

Pinnularia gi~b~ba Ehrnb.

Pinnularia gibba forma sul)undulata A. Meyer. 

Pinnularia intermpta W. Sm. 

Pinnularia interrupta forma minutissima Hust. 

Pinnulariamaior (Kiitz.) Cl. 

Pinnularia microstauron (Ehrnb.) Cl. 

Pinnularia microstauron var. ambigua Meist. 

Pinnularia viridis (Nitzsch) Ehrnb. 

Stauroneis alabamae Heiden var. angulata Heid. Length

75/1-80^, breadth 17/j. According to Cleve-Euler (1953,

p. 211-212) Heiden's taxon is identical with 8. nol)iUs

Schumann, but see also Cleve (1894, p. 148). 

Stauroneis anceps Ehrnb. Length of the cells 63/i-71/i,

breadth 14^-15/i; striae about 25 in 10/t. 

Stenopterobia intermedia (Lewis) Fricke. Length of the

cells 200/1-215/tt, breadth ll/i-13/j; costae short, 4-5 in lO/i;

transverse striae very delicate, about 20 in 10/tt. 

Tal>ellariaflocculosa (Roth.) Kiitz.

CHLOROMONADOPHYCEAE

Vacularia virescens Cienk. Mostly encysted. 

CRYPYOPHYCEAE

Cryptaulax milgaris Skuja. 

Cyathomonas truncata (Fres.) Fisch.

DINOPHYCEAE

Peridinium cunningtonii Lemm. tab. contactum Lef. 

RHIZOPODA

Paulinella chromatopJiora Lauterb. The ovoid test with a 

low collar around the aperture, 29/t-31/u. long and 25/t*-26/j 

wide; cyanells 2, about 25/t long and 4.5/i-5/x broad.

HYPHOMYCETES

Tetracladium marchalianum De Wildem.

Mycelium of this swamp species was found on decaying 

leaves of Isoetes sp. The evidently septate hyphae are 

sparsely or moderately branched and irregularly undulate 

or are more or less nodulose (constricted always at the 

septa), colorless or pale brownish ; older hyphae as much as 

6/1 broad, the terminal hyphae about 1.5/tt broad, with cells 

that are 3-8 times as long as broad ; sporo- or conidiophores 

lateral, relatively short and usually unicellular thus without septa, the terminal parts slightly capitate with many

short, budlike protruding branchlets from which subsequently the terminal more or less tetraradiate hyaline spore 

or conidium arises. (PI. 2, figs. 50-53.)

SELECTED REFERENCES 

[For references cited, see general listing of references]

Cleve-Euler, Astrid, 1951-55, Die Diatomeen von Schweden uni 

Finnland: Kgl. Svenska Vetenskapsakad., Handl. ser. 4; v. 2, 

no. 1, 151 p., 1951; v. 3, no. 3, 143 p., 1952; pt. 2, v. 4, no. 1, 

149 p., 1953; pt. 3, v. 4, no. 5, 240 p., 1953; pt. 4, v. 5, no. 4, 

217 p., 1955.

Croasdale, H. T, 1955, Freshwater algae of Alaska; Pt. 1, Sonre 

Desmids from the interior: Farlowia, v. 4, no. 4, p. 513-563.

 1956, Freshwater algae of Alaska; Pt. 1, Some Desmic's 

from the interior; Pt. 2, Actinotaenium, Micrasterias, and 

Cosmarium: Am. Microscop. Soc. Trans., v. 75, no. 1, p. 1-77.

 1957, Freshwater algae of Alaska; Pt. 1, Some Desmids 

from the interior; Pt. 3, Cosmariae concluded: Am. Microscop. Soc. Trans., v. 76, no. 2, p. 116-158.

1958, Freshwater algae of Alaska; Pt. 2, Some new fonrs

from the plankton of Karluk Lake: Am. Microscop. So°.

Trans., v. 77, no. 1, p. 31-35. 

Hirano, M., 1968, Desmids of arctic Alaska: Kyoto Univ. Biol.

Lab. Contr. 21, p. 1-53. 

Krieger, Willi, and Gerloff, Johannes, 1962, Die Gattur*

Cosmarium, Pt. 1: Weinheim, Germany, J. Cramer, 112 p.

 1965, Die Gattung Cosmarium, Pt. 2: Weinheim, Germany, J. Cramer, p. 113-240.

Nilsson, Sv., 1964, Freshwater Hyphomycetes: Symbolae Bet. 

Upsaliensis, v. 18, no. 2,130 p.

Polunin, Nicholas, compiler and ed., 1947, Botany of the Canadian Eastern Arctic; Pt. 2, Thallophyta and Bryophyta: 

Canada Natl. Mus. Bull. 97, Biol. Ser. 26, 573 p.

Proschkina-Lawrenko, A. I., Zabelina, M. M., Kisselew, T. A., 

and Scheschukowa, V. S., 1951, Diatomovije vodorosli: Moscow, Opredelenie Presnovodnych Vodoroslei SSSR 4.

Ross, Robert, 1947, Freshwater Diatomeae (Bacillariophyta), p. 

178-233, in Polunin, Nicholas, compiler and ed., Botany of 

the Canadian Eastern Arctic, Pt. 2: Canada Natl. Mus. Bull. 

97, Biol. Ser. 26, 573 p.

Starmach, Karol, 1966, Cyanophyta-Sinice, Glaucophyta-Glauiofity, v. 2 of Flora Slodkowodna Polski: Warszaw, Panstwo^e 

Wydawnictwo Naukowe, 807 p.

Taylor, W. R., 1928, The alpine algal vegetation of the mountains of British Columbia: Acad. Nat. Sci. Philadelphia 

Proc., v. 80, p. 45-114.

 1934, The fresh-water algae of Newfoundland, Pt. 1: 

Michigan Acad. Sci., Arts, and Letters Papers, v. 19, p. 

217-280.

1935, The fresh-water algae of Newfoundland, Pt. 2:

Michigan Acad. Sci., Arts, and Letters Papers, v. 20, p.

185-230.

Vischer, W., 1920, Sur le polymorphisms de I'Ankistrodesmus 

Braunii (Naegeli) Collins: Schweizer. Zeitschr. Hydrologie, 

v. 1, p. 5-50.

SOIL FUNGI AND ALGAE

By L. W. DuEREix 6

The fungi and algae were cultured from soils that 

surrounded the roots of several grasses (Alopecurus

« Professor Emeritus of Botany and Plant Pathology, Colorado State 

University, Fort Collins, Colo.
56 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

aequalis, Deschampsia caespitosa, Phleum alpinum, 

and Trisetum spicatum) that had been collected for 

transplanting in a common garden in order to study 

their ecotypic variation. The grasses, together with the 

upper 3-4 inches of soil that adhered to their roots, were 

collected on August 7, 1967, from upland sites and were 

placed in plastic bags to keep them moist during shipment. While collecting the specimens, no special precautions were taken to prevent contamination of the soil 

by foreign microorganisms; the preliminary inventory 

of species that were found in the soil samples may, 

nevertheless, be indicative of the microflora at the sites. 

To isolate the fungi from the soil samples, small 

fragments of soil were planted on Rose-Bengal agar. 

This method of culture gives a more reliable index of 

the fungi that are present than does the old dilution 

technique. Table 1 gives a list of the fungi that were 

isolated from the 21 samples. Fungi were cultured from 

all but one of the samples, and two or more taxa were 

identified from more than half of the samples. Sample

7, in which six taxa representing as many genera were 

found, was the richest sample. In some of the genera, 

notably Phoma, Fvxarium, and Gephalosporium, it is 

difficult if not impossible to make determinations of 

species.

In my studies of Alaskaii soil samples from Peters 

Lake in the Brooks Range, Barter Island, the shores of 

the Arctic Ocean, and Matanuska, I have isolated 28 

species of fungi. Judged from those and hundreds of 

other samples from throughout the world, none of the 

taxa found on Amchitka Island can be considered to 

represent unique distributions all are widespread taxa. 

Much more intensive and careful sampling would be 

required to reveal unique elements of the microflora, if 

they are present.

Algae were cultured from 10 of the 21 soil samples by 

placing soil fragments on white quartz sane1 that had 

been autoclaved in small Erlenmeyer flasks. The sand 

was then wetted with modified Bristol-Roach solution 

(Bold, 1949) to add some nutrients and to stimulate

TABLE 1. Fungi and algae cultured from soil samples taken on Amchitka Island, Alaska 

[X indicates the presence, .... the absence, and ? the questionable presence of the organism in the soil sample)

Soil sample

Organism

Fungi 

Alternaria tennis auct. Wiltshire ......... ....

Aspergittus nidulans (Nidan) Winter..... ......................... . .....

terreus Thorn. ...-..-----......-...... --... ...

Cephalosporium sp... .--.------.....-.-.... --.-._._ . .

Curvularia sp_.... .................................... .

Fusarinm sp _ .......................... ....

Mucor corticolus Hagen. .......... .....

racemosus Fresenius... ........ .....

variant Povah.... __ ............... .. ....

sp......... ............................................................

Penicillium cambertii Thorn.. .................. _ ......

citrinum Thorn-----..-.--. . . .....

frequentans Westling _ ...... -----

oxaltimm Thorn. -...-..-.- . .....

Sp... ..-.--..__...._.______.__.._______. _.__.________._._._ . , ..

Phoma sp. ..............................

Pythium sp.---- -.-.

StachybotTysatTaCor&a...--^- ........... . .....

Stemphylium ilicis Tengwall. . . . . . .

Trichoderma viride Pers. ex Fries... --.-.

Verticillium sp. ...................... ...

Algae

Chlorella vulgaris Beyerinck.

Chlorococcum humicola (Naeg.) Rabenhorst..-.-

Palmella sp...-. ..

Phvrmidium antarcticum W. et G. S. West

Stichococcus subtilis (Kuetz.) Klercker..

Stigonema turfaceum (Berk.) Cooke __ .

1

X

y

y

2

y

y

3 4 5

?

y

y

6

y

7

y

y

y

y

8 9

y

y

10

y

y

y

11

X

12

y

y

13

y

14 15

y

X

16

X

X

?

X

17

X

X

X

X

X

18

X

X

X

X

19

X

X

20

X

21

X

1. Black sandy loam, pH 7.0. 

2. Black sandy loam, pH 6.0. 

3. Gray organic clay, pH 6.0. 

4. Gray organic clay, pH 5.9. 

5. Gray organic clay, pH 5.5. 

6. Black sandy loam, pH 6.4. 

7. Black sandy loam, pH 6.2.

SOIL SAMPLES

8. Black sandy loam, pH 5.9. 

9. Black sandy loam, pH 6.4. 

10. Black sandy loam, pH 6.5. 

11. Gray sandy loam, pH 6.5. 

12. Gray sandy loam, pH 6.6. 

13. Loam (muddy), pH 6.6. 

14. Dark sandy loam, pH 6.8.

15. Gray organic matter and clay, r*I 6.5. 

16. Gray sandy loam, pH 6.2. 

17. Gray organic matter and clay, pH 6.5. 

18. Black sandy loam, pH 6.2. 

19. Black sandy loam, pH 6.5. 

20. Gray organic matter and clay, pH 6.5. 

21. Gray organic matter and clay, pH 6.9.
PLANT SPECIES OF AMCHITKA ISLAND 57

growth. In about 60 days, algae, if present in the sample, 

will form a green growth on the sand. As in fungi, sample 7 was among the richest in algae of those examined, 

having four species that represent three genera. The 

absence of algae from 11 of the samples may have been 

because of limited soil sampling. Growth in a culture 

frequently originates from a single propagule; therefore, more extensive sampling at a site would have 

increased the likelihood of including these single reproductive bodies.

OBSERVATIONS ON THE FLORA »

By WILLIAM M. KLEIN 8

Since 1965 several Government agencies have supported studies of the vegetation and ecology of Amchitka Island. Botanists of the U.S. Geological Survey and 

others have collected extensively on the island, with 

vascular plants, bryophytes, and lichens having received the most attention. I collected vascular plants on 

this island during the summer of 1967 to provide taxonomic support for ecological investigations. These 

recent studies have resulted in the accumulation of a 

large number of plant specimens, and the Amchitka 

Island flora is now probably more thoroughly collected 

than that of any other island in the Near Island and 

Rat Island groups.

After I had critically studied some of these collections, it became apparent that the recent collecting efforts had brought the understanding of this flora to 

a new level and that certain binomials could no longer 

be applied. The observations in this report indicate the 

progress that has been made in these floristic studies 

and emphasize the need for a clarification of certain 

vascular plant names as applied to this flora.

The floristic studies of the Aleutian Islands have 

followed the usual course of events as described by 

Davis and Hey wood (1963). The first and so-called 

pioneering phase in the Aleutian Islands was led by 

Hulten (1960). In this stage taxonomic decisions must 

be based upon limited material, and judgments regarding the status of a particular taxon must at times be 

arbitrary. Through more comprehensive collections and 

herbarium studies knowledge of the flora enters the 

second or consolidation phase where many of the variants described as species are found to be variants of 

previously described taxa, Intensified exploration, also 

in this phase, may lead to the discovery of new species 

and may make possible conclusions regarding phytogeographic relationships. The floristic studies of Am7 This study was supported by U.S. Department of the Interior contract 

14-08-0001-10935.

8 Department of Botany and Plant Pathology, Colorado State University, Fort Collins, Colo.

chitka Island have now approached this second phase, 

as is indicated by the comments on the taxa that folk w. 

The third and fourth phases, biosystematic and encyclopedic, are mostly in the future, although some work 1 as 

now begun which might appropriately be called 

biosystematic.

The collections of the Gramiiieae were compared with 

those in the U.S. National Herbarium. The names tl at 

I suggest are based primarily upon these studies and reflect my own views regarding the application of specific 

and subspecific categories which may be summarized 

as follows:

1. When two taxa intergrade over a considerable pT.rt 

of their distribution range there can be, in my 

opinion, no real justification for recognizing thQ.m 

as distinct species. The occurrence of intermediate 

types commonly indicates genetic continuity between populations which are best treated as conspecific. A satisfactory treatment, however, vlll 

come only after the geography and pattern of 

variation are more thoroughly understood. Until 

such time, it seems best to follow a conservative 

course in the application of new names.

2. Intraspecific taxa should be recognized where this 

appears to be an appropriate way of dealing with 

the variation pattern. Recognition of such taxa 

usually requires intensive field and herbarium 

studies, and if possible these studies should be 

correlated with experimental findings. In irost 

instances where intraspecific taxa are distinguished, there is geographic and ecological dif ?-rentiation, and taxa can be delimited on the basis 

of morphology.

3. The occurrence of two or more distinct morphological forms in the same populations throughout the 

range of a taxon may indicate the polymorphic 

nature of the population rather than fundamental 

differences. The application of names to the virtually endless number of variants can only burien 

the taxonomic system and inevitably obscure the 

more important relationships.

Determinations of Epilobium were made by Dr. P. A. 

Munz of the Rancho Santa Ana Botanic Garden, Claremont, Calif. Collections of Salix were sent to Dr. G. W. 

Argus of the W. P. Fraser Herbarium, University of 

Saskatchewan, Saskatoon, Saskatchewan. Collections 

of Ca-rex were sent to Dr. F. J. Hermann of the U.S. 

Forest Service. The assistance of these specialists is 

gratefully acknowledged.

In the vegetation studies and the annotated lis^ of 

vascular plants of this report, Shacklette, Erdman, and 

Keith used the names that are given in Hulten's fora
58 VEGETATION OF AMCHITKA ISLAND, ALEUTIAN ISLANDS, ALASKA

(1968), except for some references to Amchitka Island 

taxa that were not collected in the present study. Hulten's 1968 nomenclature was adopted because of the advantages gained by following a comprehensive modern 

treatment of the Alaskan flora. Results of taxonomic 

studies that I report below are preliminary to more complete morphological and cytological investigations of 

certain critical groups of taxa from this island.

The following list includes only the families on which 

some critical work has been done and the taxa that have 

binomials whose proper application was in doubt. Suggested names for species of four families in the Amchitka Island flora follow.

CYPERACEAE

Hermann's determinations of species in this family agreed 

with those of Hultdn for all but one species, Carex kelloggii 

Boott. Two collections of this species, 7485 and 8031, were 

made by Shacklette. Collection 7485 was determined as C. 

Jiindsii C. P. Clark and collection 8031 as possibly C. 

aquatills Wahl., although it was noted that the material of 

the latter was too immature for a critical identification to be 

made.

GRAMINEAE

Agrostis borettlis Hartm. [A. ala-skana Hult. (Hult£n, 1960, 

p. 78; 1968, p. 99) ]. Agrostis alaskana is noted by Hultfin 

(1960, p. 77) to be the most common species of the genus 

in the Aleutian Islands. It intergrades with A. oorealis 

and should be placed in that complex.

Descliampsia atropurpurea (Wahl.) Sheele [Vahlodea 

atropurpurea (Wahl.) Fr. (Hulten, 1960, p. 88; 1968, 

p. 115)]. Vahlodea is treated as a segregate of 

Deschampsia in the U.S. National Herbarium and in most 

American works.

DescHiampsia caespitosa (L.) Beauv. [D. beringensix Hult. 

(Hulten, 1960, p. 85-87; 1968, p. 114)]. Deschampsia 

leringensls is cited by Hult6n (1960, p. 86) as "one of 

the most common grasses on the Aleutians." This species 

cannot be distinguished from D. caespitosa, and Kawano 

(1963) treats it in that complex. Hulten (1968) noted that 

many specimens show "hybrid influence from D. 

caespitosa."

Festuca ovina var. ~brachypliylla (Schult.) Piper [F. ~bracliyphylla Schult. (Hulten, 1960, p. 103-104; 1968, p. 168)]. 

Festuca frrachypliylla intergrades widely with F. ovina 

and affinities are best shown by maintaining it in this 

complex, which may also include F. ovina subsp. afosfcewsis Holmen (Hulten, 1968).

Festuca rulra L. [F. rubra subsp. acuta (Krecz & Bobr.) 

Hult. (Hulten, 1960, p. 104; 1968, p. 171)]. Festuca rulra 

is a highly variable species and some of the variants that 

have been described appear to represent expressions of 

polymorphic systems and should not be accorded formal 

taxonomic recognition.

Plileum alpinum L. [P. alpinum var. americanum Fourn. 

(Hulten, 1960, p. 74-75) ; P. commutatum Gandoger var. 

americanum (Fourn.) Hult. (Hult6n, 1968, p. 88)]. The 

recognition of infraspecific taxa and the application here 

of another specific epithet does not appear to be justified 

on the basis of existing evidence.

Poa alpina L. [P. Jiispidula Vasey; P. Jcomarovii Roshew; 

P. lanata (Scribn.) Merr.; P. turneri Scribn.]. An extremely complex group; the above-listed taxa cited by 

Hultgn (1969, 1968) for Amchitka Island probably can 

all be assigned to the P. alpina complex .All of these taxa 

produce viviparous forms that appear to be environmentally induced.

ONAGRACEAE

Epilotitim boreale Hausskn. [E. Iglandulosum Lehm. 

(Hultgn, 1960, p. 263-264; 1968, p. 690)]. Material determined by P. A. Munz to be E. boreale is very common on 

Amchitka Island, although this species is not renorted by 

Hultgn for any of the Aleutian Islands. It seems likely 

that this plant is being called E. glandulosum, which is 

reported to be common along the entire island chain.

SALICACEAE

Salix arctica Pall. [8. crassijulis Trautv. (Hult6n, 1960, 

p. 158-160) ; 8. arctica Pall, subsp. crassijulis (Trautv.) 

Skvortz. (Hulten, 1968, p. 340)]. Salix arctici, according to G. W. Argus (written commun., 1968), is a highly 

variable species and it is impossible at this point to 

distinguish intraspecific taxa.

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60 VEGETATION OF AMCHTTKA ISLAND, ALEUTIAN ISLANDS, ALASKA

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INDEX

A Page 

Achittea borealis............................ 26,31,34

Achnanthes minutissima...................... 55

spp -.._- __.-- __ __ 50 

Acknowledgments..........._.__--.... 3,40,57

Actinotaenium cucurbita.................... 50;pl.l

Adelanthaceae................................ 41

Age of ash layers, radiocarbon dating. -..._.__ 8 

Agrostis alaskana....................... 15,19,36,58

borealis............................. 32,33,36,58

exarata................................... 13,36

Alectoria divergent............................ 50

irvingii................................ 28,33,50

nigricans................................. 33,50

ochroleuca....................... 16,28,29,32,50

pubescent ............................ 29,32,50

Algae_.___.__._....__._-.___.. 60

cultured from, soil samples---.-..--------- 56

selected references........................ 55

soil.----...---------------.-------..-....- 66

Alluvial soil...._.__..................... 6

Alopecurusaequalis........................ 24,36,55

Alpine meadow community. ----.-------._.__ 10, 31

Alpine meadow habitat...................... 10,30

Alternaria tennis.............................. 56

Am blystegiaceae....---.-...-.....--.-----.-. 44

Amblystegiella sprucei......................... 44

Amblystegium serpens............. 14,15,21,44,45,47

Amphidium lappmicum................... 14,45,47

Anabaena variabilis........................... 56

Anaphalis margaritacea....................... 26,34

Anastrophyllum.............................. 42

sphendtoboides..--...-................. 18,41,42

Andreaceae................................... 44

Andreaea rupestris....... 13,28,29,30,31,32,33,40,44

Andreaea-Schistidium-Rhacomitrium community--.---.------.-------.... 10,25

Anemone..................................... 10

narcissiflora villosissima................ 15,16,39

Anemone-Elymus-Ligusticum community_.. 10, 16 

Aneuraceae .........................._... 43

Angelica lucida............................ 26,32,40

Anisonema acinus............................ 54

Anisothecium squarrosum..................... 45

Ankistrodesmusbraunii............. ....... 53; pi. 2

spir<dis................................... 53

Anomobryum condnnatum................. 19,45,47

filiformis.................................. 45

Antennaria dioica.................... 20,26,27,32,34

pallida .......___..-..___--...... 34

Antheba...................................... 40

julacea..................... 21,24,31,33,40,41,43

Antheliaceae....__................___. 41

Antitrichia.. ........._......___._..... 18

curtipendula............ 13,16,18,19,29,30,32,46

Aongstroemialongipes..-. _____......_... 22,45

Aphanothece castagnei......................... 51

Aphragmus eschscholtzianus................... 35

Arctoafulvetta.----........................... 29,45

Arnica unalaschcensis................... 16,26,31,34

Arnica-Lupinus community.................. 10,26

Ash layers-.................................. 7

Aspergillus nidulans.......................... 56

terreus.................................... 56

Athyrium felix-femina cydosorum........... 15,19,38

Aulacomniaceae.---.__..................... 44

Aulacomnmm palustre .......... 16,21,22,24,44,46

[Italic page numbers indicate major references]

B Page 

Bacidea sp._...._.___..______... 32,48

Baeomyces placophyllus...... .......-.---.. . 33,47

Barbula cylindrica......................... 14,41,47

Bartramiaithyphytta....................... 14,41,44

Bartramiaceae- ...____..._.... ..... 44

Bazzaniatncrenata.----...................... 18,41

Blasia pusitta .............................. 20,43

Blasiaceae.................................... 43

Blastenia sp.___....__...______.... 13,49

Bodo minimus................................ 54

Bog habitat-.....-.. -.---..--- ------ 10,20

Bog soil......_...........__-.____-..- 6

Boraginaceae...- -..-------..---..-----.. - 34

Brachytheciaceae_.-...-..---.-.-..----.-.-- 45

Brachythecium....... .........__........... 25

acutum... ....................... 16,27,33,45,46

albicans....................... 13,19,27,28,30,45

asperrimum...... 16,17,19,21,25,27,41,43,44,45

Bryaceae...-........ ............ - ... 45

Bryum sp.-.. .......-.. .. ... - 24

Bromus sitchensis aleutensis....-....----.....- 13,36

Bryoerythrophyllum recurvirostrum............ 13,

20,25,44,45,47 

Bryophytes, annotated list....... ...... 40

Bryoxiphium norvegicum...................... 47

Bryum arcticum.............................-19,45

inclinatum- .........___..... . ... 45

lapponicum... -...---....-..-..-..-- 14,41,44,45

pollens.................................... 19,45

pseudotriquetrum ......-.....-.-.---.. 13,19,45

*o/in«n»--.----- --..---.-- -------- 45

stenotrichum......... 14,15,26,27,29,33,44,45,47

Buellia pullata................................ 49

punctata......................-...---..... 13,49

Burned areas, plant communities...--------- 33

Odamagrottis................................. 18

community_ - - -- . - 10, SO

nutkaensis........... 15,16,17,18,19,26,34,36,46

CdlUergonsarmentosum......................- 19,44

Callitrichaceae..___.__...__........ 34

Callitricheanceps...---....................... 24,34

hermaphroditica........................... 34

Cattitriche-Subularia community. ___ - _ - - - 10,24 

Caloneis silicula. ............................ 50

silicula alpina .......................... 52

longissima........................---- 52

Caloplaca elegant..... ___.__....... - 49

gramdosa.............................. 13,14,49

sp........................................ 13,49

Caloplaca-Eurhynchium-Puccinetta community- 10, 13 

Caltha palustris asarifolia..................... 22,39

Caltha-Claytonia community._.._.... ... 10,22

Calypogeia mulleriana........................ 20,41

sphagmcola...................... 16,18,21,29,41

trictiomanis-........................ 17,19,41,42

Calypogeiaceae..-------.-......---.. .--.--- 41

Ca mpanula chamissonis... _ - 32,34

lasiocarpa lasiocarpa............. 16,28,29,32,34

Campanulaceae---.- _ _ . _. 34 

Campyliumstellatum---..._............... 19,24,44

Caprifoliaceae..-------..-. ----------- 34

Carcasses, plant communities ------------ S3

Cardaminebdlidifolia......................... 30,35

umbellata.................. 14,16,17,19,20,26,35

Pagi 

Corel____._._-_.____ ....... 57

anthoxanthea..................... 16,17,21,22,35

aquatilis........................... .. ... 58

bigelowii.................................. 31,35

circinnata-.-......................... 29,33,35

dioica gynocrates.......................... 16,35

hindsii.................................... 58

kdloggii............................. 21,24,35,58

lyngbyaei............ 16,17,18,19,21,22,23,25,35

macrochaeta................ 16,17,18,22,26,32,35

subrigida.............................. 35

nesoptiila........................ ... 35

physocarpa.... _ - - 31

pluriflora........................ 16,18,21,22,31

saiatUislaxa....................... ....... 35

ttylosa.................................... 16,35

Carex-Cladonia meadow community---------- 10, 16

Carer-lichen community, ecotones around.... 9

Carex-lichen-Empetrum community------- 10, 16, 17

Caryophyllaceae ......... 34

Cassiope lycopodioides............. 16,28,29,30,32,35

CastilUja unalaschcensis.............. . 39

Cephalosporium sp . .. .-.. ... 56 

Cephalozia ambigua..................... 20,31,41,43

bicuspidata ........... 18,19,26,29,31,41,43,45

Uucantha........................... 18,33,41,42

media..................................... 19,41

montana................. - - 42

pUniceps.............................. 20,29,41

Cephaloziaceae...----. ------- -- ------ 41

Cephalozietta............------------ ---- 40

arctica..............- ........... 13,14,20,41

Cephalozielliaceae.--------------------------- 41

Cephaloziopsis saccatula....................... 43

Cerastium aleuticum.................... 28,30,33,34

beeringianum beeringianum................ 34

fischerianum........................... 12,28,34

fontanumtwale.--.................... 21,26,34

Ceratodon purpureus..... 12,13,17,26,27,32,33,45,46

Ceratodon-Equisetum-Sagina community . - 10,27 

Cetraria ciliaris............................... 49

crispa.................................- 49

cacMflote - - - " 32,49

erfcetoram----------------------------- 31,33,49

islandica crispa....................-...... 49

lacunosa.....................------------- 49

nigricans..................... ----- - 29,49

orbata....................-...-............ 49

CAoeiopedJo.-.-------------------------------- 53

Chaetoplaca -- - 53

crassiseta puella..............-...-...... 53

Chttoscyphus pallescens........ 19,20,24,25,41,43,44

CMamydomonas alaskensis............... 53,54; pi. 2

angulosa..---------- ------------------- 54

jemtlandica.....................-------... 54

leptobasis.........-----.. --- ------- 54

saxonensis................... ... - --- 54

subangulosa. ..------------- - 54; pi. 2

vulgaris. ------------ --------- ------- 54

Cldorella ellipsoidea..-------.-.------- -- ------ 51

vulgaris...... ._ . . ---- --- 51,56

Chlorococcum tiumicola... -_ - 54,56; pi. 1 

Chloromonadophyceae.------- - ------ 55

Chloromonas.......----......----------------- 54

Chlorophyceae.---------------- -- ----- 51,53

Chroococcus varius............................ 50

61
62 INDEX

Page 

Chrysanthemum arcticum arcticum...... 16,31,32,34

Chrysophyceae.------------------------------ 52,54

Citraria ciharis.................................. 33

Cladoma alpestris.................... 19,28,30,47,48

amaurocraea........................... 18,24,48

arbuscula beringiana...................... 18,48

belhdiflora.................... 16,18,23,24,48,50

cerasphora................................... 48

coccifera............................... 16,34,48

degenerans............................. 20,34,48

gratilis............................. 16,20,34,48

impexa...................................... 48

paciflca....................... 16,17,18,28,34,48

pyxidata............................... 27,34,48

rangiferina................................ 31,48

Salix-Empetrum community---.---..-------- 28

scabriuscula .......................... 18,19,48

sylvatica..................................... 48

sylvestris................................. 48

Cladonia-Carex meadow community---------- 10, 16

Cladoniaceae..---------..--___...__._._--_ 47

Cladi>p\ora...................................... 56

Claopodium crisp'folium...................... 19,47

Claytonia arctica.............................. 30,38

sibinca.................... 12,13,14,17,22,31,38

albiflora................................. 38

Claytonia-Caltha community.-...-___.__ 10,22 

Climaciaceae..-... .....-.-..----.-..-.....--- 45

Climacium dendroides......................... 19,45

Climate of area..........-..-.--------.- ..-. 4

Closteriumacutum............................... 54

Cochlearia offlcinalis oblongifdia............... 14,35

Communities, alpine meadow.--..-_____-_-_- 10, 31

Andreaea-Schistidium-Rhacomitrium....... 10,28

Clamagrostis.............................. 10,26

Caltha-Claytonia.......................... 10,22

Cladonia-Carex meadow......._...._. 10,16

Elymus.................................. 10, IS

Ely'mus-Ligusticum-Anemone............. 10, IB

Empetrum-Carez-lich&i ................. lo,16

Equisetum-Sagina-Ceratodon.............. 10,27

Eurhynchium-Puccinellia-Caloplaca....... 10, IS

fell-field.. -....-- ...-.-..-.--....... 10,28

Fontinalis-Ranuncidus................... 10,25

frost-scar.-.--.---_,....--.....-..-._. 10,20

high-altitude bedrock................_, 10,32

Hippuris-Ranunculus.................... 10,2S

Honckenya-Senecio........................ 10, .72

Isoetes-Ranunculus-Limosella..... _..... 10,2S

Juncus-Eriophorum...................... 10,24

low-altitude bedrock.....__.__..___. 10,32

Lupinus-Arnica.......................... 10,26

miscellaneous plant. ........._........ S3

moss mound .._.-------..._.....-.- 10,77

organic bird-perch mound.. _____. __ 10, /8 

peat-bank..__._.______...._. 10, 19

Philonotis-Parnassia................... 10,2^,46

plant...____..______._.-._. 9 

burned areas...._-..---.._---...- S3

carcasses .._............._..... S3

imported wood-.---------..--.-....-- S3

plantings..-..__.-..- .-._...._- SS

Poa-Deschampsia-Festuca................. 10,12

PotentiUa-Draba-Sax:fraga................. 10, 14

Rhacomitrium-Schistidium- Ulota.......... 10,13

Salix-Empetrum.......................... 10,28

Scapania-Nardia-Marsupella ............ 10,21

Siphula-Scapania......................... 10,24

snow-bed...-.--___..._______... 10, SI 

solifluction terrace._________-...- 10,29 

Sphagnum bog.._.__._____._ 10,20 

streamside. --------_.-.______ 10,19

Subularia-Callitriche...................... 10,24

wet sedge-meadow...___...__.___ 10,£/ 

Xanthoria-Ramalina...................... 10,14

Compositae_..----..____.__...___ 34

Conical mounds, basic types._.............. 17

Conioselinum chinense..................... 15,19,40

Conocephalaceae...____.___...__ 44

Page 

Conocephalum conicum....................... 19,44

Conostomum tetragonum............. 28,29,30,33,44

Coptis trifolia.............................. 16,18,39

Cornaceae...............--...-...-.--.--....- 35

Cornicularia divergent................ 16,18,32,33,50

Cornus canadensis x suecica................... 35

suecica ........................... 16,18,31,35

Cosmarium cydicum arcticum................. 50

decedens............................... 50; pi. 1

humile.................................... 51

laeve................................... 51; pi. 1

septentrionale-...--................... 51

margaritiferum............................ 51

meneghini concinnum.................. 51; pi. 1

nitidulum................................. 51

obliquum.............................. 50; pi. 1

pseudonitidulum.......................... 51

subcrenatum........................... 51; pi. 1

subpachydermum....................... 51; pi. 1

subundulatum......................... 51; pi. 1

tatricum............................... 50; pi. 1

variolatum............................. 50; pi. 1

Cratoneuronfllicinum...................... 12,14,44

Cruciferae.---.--..---_.-.-__..--.-_...- 35 

Cryptaulazvulgaris----....................... 55

Crypyophyceae.-.-..-....-......-...--.__. 55

Curvularia sp..---------__.---__.. T -----_.- 56

Cyanophyceae__.................__. 50,51,52

Cyathomonas truncata......................... 55

Cylindrocystis brebissonii...................... 50,51

brebissonii minor.......................... 50

crassa..................... __..--.____ 50,51

Cymbetta aequalis............................. 52

angustata................................. 52

hybrida............................... 52

austriaca.................................. 55

cuspidala..............------- --------- 52

delicatula................................. 52,55

graalis.................................... 52

hebridica.................................. 52,55

naviculiformis............................. 52

perpusilla .............................. 52

pusitta.................................... 52

tumidula.................................. 52

eentricosa................................. 52,55

Cyperaceae..-_._____-----._-----------.-- 35,58

Cystopteris fragilis fragilis.................. 14,19,38

D 

Dactylmhiza aristata....................... 15,26,38

Deschamsia alpina............................ 36

atropurpurea. -------- . . __- 36,58

beringensU....................... 13,17,20,36,58

caespttosa.............................. 36,56,58

orientalis-......................... 24,26,36

Deschampsia-Festuca-Poa community -_.. 10, /2 

Description of island....--.-----.----- ... 4

Desmidiaceae.. -.----.--...-----------.. 50

Diapensia lapponica obovata................ 29,30,35

Diapensiaceae__-...---,.-_-... ..... 36

Diatomaceae....--_......_.. --..... 52,55

DichodonliumpeUucidum.............. ...--.. 20,45

Dicranaceae...-.__--------_----...-.---- 45

Dicranella heteromalla................ 19,20,21,41,45

palustris-.......................... 20,21,31,45

subulata................... 20,21,26,41,43,45,47

Dicranumangustum.......................... 21,45

elongatum....................... 16,18,19,41,45

fuscescens.............................. 18,20,45

groenlandicum.--------...... 16,18,41,42,45

howettii................. 16,18,19,28,29,30,32,45

scoparium................................ 45

Dilaenaceae....-...---------..---.----------- 43

Dinophyceae....--.---------------- ------- 51,55

Diplophyllum albicans........................ 17,

19,20,24,29,30,31,33,41,42,43 

taiifolium............................. 29,43,45

Discontinuous heath habitat-. . -._.- 10,27 

Ditrichaceae____________.___. 46

Page 

Ditrichumheteromallum ................. 26,45,46

Drabaaleutica............................. 28,30,35

hyperborea.---......................... 13,14,35

Draba-Saiifraga-Potentilla communr' "_ __... 10, 14

Drepanocladus fluitans_____________ 22,44 

reealvens................... 17,21,22,24,29,31,44

uncinatus................................. 13,

18,19,27,28,29,30,31,32,33,44,45,46

Dryopteris dilatata amencana.................. 19,38

linneana.................................. 38

Dune habitat __._____________ 10,/2

E 

Elymus...................................... 47

arenariusmollis........................... 13

mottis mollis.............. 1% 13,15,16,19,36

community------______..._____ 10, IS

sea stacks....------------_..-_.__- 15

Elymus-Ligusticum-Anemone comnrinity..--- 10, IB 

Empetraceae -----________.___..- 35

Ecotones, alpine region..--------------------- 27

between plant communities.----------_ 9

Empetrum nigrum nigrum.................... 10,

16,17,18,19,27,28,2^, 31,32,33,34,36 

Empetrum heath, alpine community..... _.... .31

discontinuous heath habitat- - __... __ 27 

frost-scar community.--------------------.-..20

pool habitat_._____.___._.-.-. 23 

Siphula-Scapania community....-----._- 24

soil profile_.-._...__..._____... 7

Sphagnum ........................... 21

vascular plants........... ..-.----.-..- S4

wet sedge-meadow community........... 22

Empetrum heath habitat... ----- ... 10,15

organic bird-perch mounds.... .... 18

Empetrum-Carex-lichen community .... 10, /6,17

ecotones-..--___.-...-..--....-.-....- 9

Loiseleuria procumbens................... 25

ruderal habitat-___.._.. ----- 25

Empetrum-Salix community..---...--......- 10,28

Entodontaceae ...__.__..-.. - - 46 

Epilobtum.................................... 57

behringianum............................. 28,37

boreale.................................... 38,58

glandulosum........................ 20,22,37,58

sertulatum ....................... 21,22,26,38

Epithemia turgida............................. 50

Equisetaceae.-..._....... --------------- 35

Eyuisetum.................................... 13

arvense............................. 13,16,27,35

Equisetum-Sagina- Ceratodon community.... - - 10, 87

Ericaceae -..-- ___-.--.---_-----._.--- 35

Erigeronperegrinus-.......................... 31

peregrinusperegrinus..... . ....... 16,34

Eriphorum russeolum majus.................. 35

russeolum rufescens.............. 17,21,22,24,35

Eriophorum-Juncus community-------------- 10,24

Euastrum ansatum........................... 51

bidentatum................................ 51

denticulatum.............................. 51,54

quadrifarium.......................... 51

didelta. ................................ 51

elegans.... .--.--.... .-...-...-.. ...- 51,54

oblongum. -------------. ----------- 51

pectinatum --.____-..------_----------- 51

Euchlamydomonas............................ 54

Euglenapisciformis........................... 54

viridis olieacea............................- 54

Euglenophyceae..--..-.-.- .-.-.- .- 54 

Eunotia arcus .............................. 50

diodon ............ - 52

gracilis.............................------- 50

lunaris capitata..........................- 50

pectinalis...................-...---------- 52

praeruptabidens.......................... 51,52

muscicola....................--------- 50

robusta diadema........................... 51,52

septentrionalU.... -------------------...-- 52
INDEX 63

Page 

Eurhynchium praelongum..................... 12,

13,14,15,16,19,27,33,45,46 

Eurhynchium-PuccineUia-Caloplaca community..---.----....--...-...-... 10, 13

Fell-field community.----.--.-.._-------- W,28

Festuca brachyphytta........... 16,17,18,32,33,36,58

ocina..................................... 58

alaskensis............................. 58

brachyphytta.......................... 36,58

rubra............................ 15,16,22,36,58

aucta........................... 13,16,36,58

Festuca-Poa-Deschampsia community. --_. 10, 1% 

Flora, observations.---.-------.----- .------ 57

Fontinalaceae...... _....................... 46

Fontinalis neomexicana........................ 25,46

Fontinalis-Ranunculus community. -. _...-- 10. S5 

Fragilaria construens binodis.................. 52.55

construens subsalina....................... 52

nitzschioides............................... 52

Fritillaria camschatcensis...................... 19,37

Frost action, solifiuction habitat.. ........ 28

Frost-scar community... ----.--..---------- 10,50

Frustulia rhomboides saxonica................. 51,52

Fungi, cultured from soil samples . _... 56 

Nidula Candida........................... 1

soil.._ - - - _..____ SB

Fusarium sp ._- - . ..... 56

O 

Galium....................................... 37

aparine................................... 12,39

trifidum columbianum............... 12,16,22,39

Gentiana aleutica........................... 26,32,36

amarella acuta plebeja..................... 26,36

Qentianaceae-... ..................._.-... 36

Geography of area.......-__...__.. .-.-. 4

Geology of area... - 5

Qeraniaceae...........-..----..........---... 36

Geranium erianthum.......................... 36

Qeum calthifolium....................... 16,20,31,39

macrophyllum macrophyllum........ 13,19,22,39

pentapetalum............................. 31,39

Oloeocapsa magma............................ 50

Oloeocystis ampla............................. 56

rupestris.................................. 54

Gloeodiniummontanum.-...-.-............ 51; pi. 1

Gomphonema angustatum productum.......... 51,52

Qramineae...__._..................... 36,58

collections..------- ----._.----.-..---. 57

Grimmia apocarpa.........----.-....---.--.-- 46

maritima................................. 46

Grimmiaceae.. ......_-----.--.----. 46

Gymnocarpium dryopteris..................... 38

Gymnocolea acutttoba.......................... 42

inflata........................... 17,18,31,41,42

montana ............................ 42

marginata................................. 42

montana......-.-.--.....-... ............. 42

acuta....... - _ . 42

Gymnomitrion corattioides...-------_..... 28,33,43

Gyrophora cylindrica...-.--.............--.... 50

H

Habitats, alpine meadow. 

bog-....--.-----.---

discontinuous health.

so

80

87

Empetrum heath .__......._....-.. IB

inland bedrock. _ - . . 37

lake...__....____.....____....... 88

land, associated plant communities, outlined-. ..-__. --------....... 10

miscellaneous .... .... _ -------- 11,33

marsh..-__--.--.- .--.-....._......... 81

plant..._. _ _ ___._..........--.-. 9

Page 

Habitats Continued

pool...__..........__--.....--..__... #3

ruderal-.............._.____...._... 85

sea-cliff--.-..---...-.--.- -----.-.. IS

solifluction...____________..... 88

strand.-.-....- --. . ... ...-- 1%

stream ---.------...--.-.-._.-...-..- 84

Haematomma lapponicum..................... 48

ventosumlapponicum..... ____....... 33,48

Half Bog soil...-.....-...-....-........- 6

Haloragaceae.------.-........._........... 37

Hantzschia amphioxys......................... 52

Heracleum lanatum.............. 12,13,19,26,32,40

Herberta adunca....................-.-------- 30,41

hutchinsiae.... ...._._.____....... 41

Herbertaceae--...... ....................... 41

Hieracium.................................... 27

gracilealaskanum................---...... 27,34

Mate...................................... 28,35

Hierochloe odorata............................. 15,36

High-altitude bedrock community --------- 10, 38

Hippuris montana............................ 31,37

vulgaris................................... 23,37

Hippuris-Ranunculus community. ----------- 10, S3

Honckenya peploides major.................... 12,34

Honckenya-Senecio community --------- 10, 1%

Hormidiumflaccidum......................... 51,54

Hygrohypnum................................ 25

buM...................................... 25,44

ochraceum................................ 25,44

Hylocomiaceae.--.---..._...._----------- 46

Hylocomiumsplendens.................. 16,18,26,46

Hyphomycetes. - . 55

Hypnaceae.---. ----._---.-...---.---------. 46

Hypnum cupressiforme........................ 33,46

dieckii.................................... 16,46

plicatulum................................ 30,46

Hypogymnia enteromorpha.................... 33,49

physodes.................................. 33,49

subobscura................-...----.-.--.-- 29,49

Icmadophilaericetorum................ ........ 19,48

Inland bedrock habitat............-.....--- W,S1

lonaspisepulotica.-..-------.................. 20

epuloticaarctica..-........................ 32,48

Iridaceae...--..--.---- -_..---..-----.---- 37

Iris setosa setosa............................... 37

Isoe'taceae..-------------------- ---------- 37

Isoetes muricata maritima..................... 23,37

sp... _ .... .-. -. 24,37,52 

Isoltes-Ranunculus-Limosella community.. -. - 10, S3 

Isopaches bicrenatus........................... 42

Juncaceae.-.---.- -..------..----- .. . 37

Juncus arcticus sitchensis...................... 12,37

biglumis.................................. 30,37

ensifolius................................. 24,37

triglumis albescens............... 17,21,22,24,37

Juncus-Eriophorum community-. . ... 10, 84 

Jungermanniaatrovirens...................... 30,41

minuta grandis...------------------------- 43

rigida /8 grandis.........-.......--.----..-. 43

saccatula.....................------------- 43

sphaerocarpa.............................. 21,41

Jungermanniaceae ._..... - 41

K 

Koenigia islandica............................. 30,38

Kolieaa....................................... 53

corcontica.......................---------- 54

Lake habitat- - -- - 10, *» 

Land habitats, associated plant communities,

outlined.-.. - ----- -------- 10

miscellaneous_ ._ - --- - - 11,33

Page 

Lathyrus maritimus pubescent................. 12,37

palustris pilosus.......................... 37

Lecanora aleutica...........______... 13,14,48

allophana................................. 33,48

atra........_____ ..______.__ 13,48

castanea.................................. 19,48

gelida..................................... 48

straminea................................. 14,48

subfusca.....-------------------------__ 48

Lecanoraceae.-------___- -____......... 48

Lecidea aleutica..........____ .. ...__ 32,48

armeniaca................................ 33,48

elegantior................................. 33,48

flavocaerulescens.......................... 33,48

macrocarpa...........-.......... 13,27,32,33,48

platycarpa................................ 48

symmicta..........................._... 33,48

vernaKt................................ 20,33,48

Lecideaceae..............___..____._. 48

Leguminosae.....-----.-------..-........-- 37

Lentibulariaceae. - 37

Lepidoiiaceratoptiylla .--.................... 40

trichodadus............................... 41

Lepidoziaceae.. __ __ __ __ _._.. .. ._._ 41

Leptarrhena pyrolifolia........................ 31,39

Leptobryum pyriforme................ 15,20,44,45,47

Leucodontaceae....----.._-.._- _ .____ 46

~Lichen-Empetrum-Carex community..... - 10, 16, 17

Lichens, annotated list. . __......... 1ft

Ligusticum................................... 10

scoticum hultenii....................... 13,15,40

Ligusticum-Anemone-Elymus community. 10, 15 

LDiaceae ......__...... ___--------- 37

Limosettaaquatica...................... 21,23,39,43

Limosella-Isoetes-Ranunculus community... -. 10, S3 

Linnaea borealis borealis....................... 16,34

Listera cordata nephrophylla. ....... _. __ 16,31,38

Lithosols, classification...-.--.-.-------- -.- 6

Lobarialinita................................. 16,49

Loiseleuria procumbens........ 16,18,25,27,28,32,36

Lophocolea cuspidata......... ._- 13,14,31,41,43,46

Lophocoleaceae ...._......______... 41

Lophozia..................................... 40

alpestris...................... 20,22,31,41,42,43

groenlandka............ 16,17,20,21,33,41,42,43

ineisa............................... 20,24,33,43

opacifolia.............................. 31,41,43

porphyroleuca........-__.__.___. 42

wenzelii.-................................ 32,43

Lophoziaceae...-....-_._................. 42

Low-altitude bedrock communities..... 10,3^

Lupinus...................................... 29

nootkatensis-..................... 21,25,26,29,37

Lupinus-Arnica community...__...__.... 10, 88

Luzula....................................... 27

arcuate unalaschcensis................. 28,32r 37

multiflora kobayasii....................... 27,37

multiflorafrigida................ 16,20,26, 37

parviflora parviflora................. 19,26,27,37

tundricola..................... 14,28,29,30,33, 37

wahlenbergiipiperi........................ 37

Lycopodiaceae -.. 37

Lycopodium.................................. 16 27

alpinum.................................. 28 37

annotinum annotinum.............. 16,17,21.37

clavatum clavalum........................ 16 37

sabinaefoliumsitchense................. 21,32,37

selagoselago.. ................. 16,27,28,32,37

Lyngbya rivulariarum......................... 52

M

Macrodiplophyllumplicatum.-.----------..... 19 43

Majanthemum dilatatum................... 19,20 37

Marchantiapolymorpha.............. 14,24,26,44 47

Marchantiaceae.-----.------ --------------- 44

Marsh habitat-.. 10 91

Marsupetta emarginata..... 19,21,24,30,31,41,43 50

Marsupetta-Scapania-Nardia community... - - 10 81
64 INDEX

Page

Marsupellaceae ..___...__..__.__- 43 

Melosira distans dlpigena.........--...-.-..... 51,52

distansp)'affiana................_. ... 55

Menoidium incunum......................... 54

Merismopedia elegans......................... 52

glauca.................................... 52

Mertemia mantima........................... 12

maritima maritima........................ 34

Mesotaenlum micrococcum.---------.......... 50

Microlepidozia makinoana............ 16,18,20,41,42

sllvatica................................... 41

Mielichhoferia macrocarpa ................ 14,44,45

Mimulusguttatus............................. 25,39

Mniaceae. .................................... 46

Mnium glabrescens........................... 16,

19,20,22,24,26,32,33, 41,43,45,46

insigne.................................... 20,46

 pseudopunctatum................ 19,20,21,33,46

Monas uniguttata............................. 54

Montia fontana fontana.................... 14,25,38

Montospora brevis............................. 56

Mortierella vinacea............................ 56

Moss mound community........_.___... 10, 17

Mougeotia.................................... 51,54

Mucor corticolus.............................. 56

racemosus................................. 56

varians................................... 56

sp...-._ - - __.........___. 56

Mycoblastus alpinus.............. 16,18,27,28,33,48

sanguinarius.............................. 48

Mylia anomala............................ 31,41,43

taylori........................... 16,17,18,41,43

Myriophyllum spicatum....................... 37

N

Names, flora, specific and subspecific categories............................... 57

Nardia co mp ressa............................ .31,41

geoscyphus............................. 31,41,43

scalaris-... 13,19,20,21, 22,24,26,27,31,32,41,43 

Nardia-Marsupella-Scapania community.-... 10, HI 

Navicula contenta................ ............. 52

cryptocephala....--....--................. 52

minima................................... 52

atomoides............................. 52

Neidium affine minus......................... 52

dubium................................... 55

iridis amphigomphus...................... 52,55

Nidula Candida.. ............................ 1

Nitzschia gracilis.............................. 55

palea..................................... 52

stagnorum................................ 52

Nonmenclature of plant species ............. 3

Nostoc microscopicum......................... 52

paludosum................................ 56

O

Ochrolechiafrigida........ 14,15,16,27,28,29,32,33,48

Odontoschisma elongatum................... 18,41,43

Oedogonium.................................. 54

Oligotrichum hercynicum.-_..-.-.............. 31,47

parallelum............................... 31,47

Onagraceae................................... 37,58

Oncophorus wahlenbergii-...... 17,18,20,21,41,42,46

Oocystispana.............................. 51; pi. 1

Orchidaceae..........._...................... 38

Organic bird-perch mound community.. - _. 10,18 

Orthocaulis binsteadii-............ 16,17,18,33,41,43

floerkei ................................. 20,43

Orthotrichaceae......................._.... 46

Oscillatoria amoena...................... 51,52; pi. 2

amphibia.... ...-.....-.. ..... 51,52,53; pi. 2

defleia.................................... 52

exilis ................................... 52

facilis.................................. 52; pi. 2

grossegranulata......................... 53; pi. 2

limnetica acicularis........................ 52

neglecta.............................. ..... 52

Page

Oscillatoria Continued

splendida............................... 53; pi. 2

subtilissima............................... 52

sp........................................ 53

Oxyria digyna................................. 31,38

P 

Palmetto, sp................................... 56

Pannaria pezizoides........................... 20,49

Pannariaceae..........._..........._.-... 49

Papaver alaskanum........................... 38

Papaveraceae.....-...-.---.__..-..--.-_ 38

Parmeliaalpicola............................. 33,49

duplicata douglasicola..................... 49

lanata.................................... 50

omphalodes. ........................ 18,29,33,49

physodes.................................. 49

saiatilis .......................... 14,32,33,49

subobscura................................ 49

sulcata. -------------------------- 14,32,33,49

Parmeliaceae.-.-------------.----.._...--.- 49

Parmeliopsis ambigua--.... ................... 33,49

Parnassia kctzebuei........................ 21,24,39

Parnassia-Philonotis community... ___ - 10, tl, 46 

Paroicous inflorescence ._............... 42

Paulinella chromatophora..................... 55

Peat-bank community. .---...--.---------. 10,19

Pediastrum braunii........................... 51,54

Pediculans chamissonis.................... 16,22,39

Pdlianeesiana ................. 22,31,41,42,43

Peltigera aphtltosa-................... 18,19,28,32,49

canina..--......... .......__... 13,16,19,41,49

canina membranacea............------..-- 49

spuria.-..--------.------.-.---------- 49

malacea... ......................._--..-. 16,49

spuria.... ...._. . .....__........ 12,49

Peltigeraceae................................. 49

Penicillium cambertii ........................ 56

citrinum.................................. 56

freguentans.....................-...--.--- 56

oxalticum.---------.-.-----------------.-. 56

sp................. _-- .-- -- .-- 56

Peridinium cinctum........................... 51

cunningtonii contactum....-..-..--.....-.- 55

Pertusaria coriacea..-.-......--.-----.-------- 32,49

hultenii................................... 33,49

sp---.... -. ...- 33,49

Pertusariaceae..-----.-...__-----.---.------ 49

Petasitesfrigidus -................... 17,19,22,35

Philonotis..................................... 26

americana....................... 21,22,26,44,47

fontana......-...--..--...-.-....---... 20,21,44

PMlonotis-Parnassia bog-..........--------... 43

community_.._._____.-.-.-.-. 10,27,46 

Phleumalpinum.............................. 56,58

alpinum americanum..................... 58

commutatum americanum.............. 16,36,58

Phomasp -----------------.-----..---...... 56

Phormidium antarcticum ...........-..-.-.-. 56

crouani...... .........__. . 56

frigidum.---.---.-----------------.....-.. 53

motte......--...----....--.......-----.... 56

subcapitatum..................... ----- 56

Phystia caesia.... ..-.--------..-----. ... 14,15,49

dubia..................................... 15,49

Physciaceae.----.----._.._. _......... 49

Physiography of area.--------.-.--..---..-.-- 5

Picea sitchensis............................. 4,33.39

Pinguicula vulgaris macroceras- ......... 17,21,24,37

Pinnularia appendiculata... .................. 52

brevicostata .............................. 52

cardinalicidus.......-.....--.-.-.-------.- 55

dactylus- ............ .......-...-.. .-. 55

divergens.................................. 52

gentilis ---------------------------------- 52,55

gibba- .............................. 51,52,55

subundulata.------..---......--..-... 55

interrupta.....--------------_.......---- 52,55

mtnufiwtma...................... - 55

Page 

Pinnularia Continued

lata---................................... 51,52

motor..................................... 52,55

microstauron................ .._.__ 51,52.55

ambigua......... ..................... 55

biundulata............................ 52

piridis ---------------------------- 52,55

rupestris........--....--.......-...-.. 51

sudetica.-...-.-..-.--................. 52

Placopsisgelida............................ 27,32,48

Plagiotheciaceae ..-.-.-------.--..-..-----._ 46

Plagiothecium roeseanum................... 13,41,46

undulatum ....................... 16,20,31,46

Planctococcus sphaerocystiformis............ 54; pi. 2

Plant communities.-.__.-.-...-.__....... 9

associated land habitats, outlined ...... 10

classifications...--....._..........-.--.- 11

miscellaneous ._..__.......__.... 11, SS

Plant habitats---..---------....------------- 9

Plant species-.....--. .... ........... 34

nomenclature-_........__ .__...... 3

Plantaginaceae.---...-_-.----.-----.--.---- 38

Plantago macrocarpa...... ... ... ----- - 22,38

maritima juncoides...... ......... 14,15,21,31,38

Platanthera chorisiana ................ 16,27,32,38

convallariaefolia.................. 16,17,21,31,38

dilatata.......................... 16,17,21,31,38

tipuloides behringiana..------------------- 38

Platydictya jungerman nioides ............ 14,19,44

Platysmatialacunosa ......-...-...--......- 33,49

Plectocolea .........__ .. - 40

obovata .............................. 20,41,45

PUuromonasjaculans. ...................... 54

Pleurozium schreberi. .......... 16,18,22,31,33,45,46

Poa alpina .............................. 27,36,58

arcticawilliamsii.-.................... 18,32,36

eminens. .......... ---- --.- 13,36

hispidula ............................... 36,58

aleutica............................... 36

vivipara ~..-.......--------..---.-..-. 36

komarovii. ---_.---. --------------------- 36,58

lanata.................................. 13,36,58

vivipara...-----.....--.-.-..-...-----. 30,36

malacantha................................ 36

turneri-. ........................... 27,36,58

Poa-Veschampsia-Festuca community 10, H 

Pogonatum...-........... .. .. 37

alpinum............. 16,17,18,19,24,33,43,46,47

contortum.......----.-.-......-...---..... 20,47

sphaerothetium...... ....----- - -- 47

urnigerum.--.-..- 7,13,20,21,2P, 32,41,43,45,47 

Pohlia annotina~~........................... 20,45

cruda .. . - - -- 13,14,20,45 

naten* - 20 26,29,33,45,47

Polygonaceae - ----------- ...-- ... 38

Palygonum viviparum................ 16,26,28,31,38

Polypodiaceae . ... ... - ------- 38

Polypodium vulgare octidentale................ 19,38

Polytrichaceae- -.. - ....... 47

Polytrichum commune..........-------. - 18,19,47

formosum.......-.----.-------......--- 20,27,47

gracile............................ ------ 47

juniperinum.....-----------.------------- 32,47

longisetum ........................... 22,44,47

sphaerothecium...------------------------ 20,47

strictum......--.------------------------- 18,47

Pool habitat------------------------- 10,23

Portulacaceae.---------------- ------ 38

Potentillaegedtigrandis...-------- --------- 26,39

hyparctica----- --------------- ---------- 30,39

viUosa.. . - .-- 14,15,39

Potentilla-Draba-Sa t ifraga community-------- 10, 14

Portia ftdww------------------------ 14,21,45,47

Pottiaceae ----------- -------------- 47

Primula cuneifolia saxifragifolia...... 16,26,31,32,38

cuneifolia saiifragifolia albiflora............ 38

Primulaceae - --.----- ------------ 38

Protists.... -..--__..----- ------------ B0

selected references.. - - --- BB
INDEX 65

Page 

Pseudanabaena bipes.......................... 53

catenata................................... 53

galeata.................................... 53

granulifera............................. 53; pi. 2

minuta................................... 53

Psoroma hypnorum........................... 19,49

Ptilidiaceae_......--...-.-.._........... 43

Ptilidium ciliare.............................. 18,43

Ptilium crista-castrensis....................... 16,46

Puccinellia langeana alaskana.............. 13,18,36

Puccinettia-Caloplaca-Eurhynchium community-.. ........................... 10,13

Pyrola asarifolia purpurea..................... 16,38

minor..................................... 31,38

Pyrolaceae................................... 38

Pythiumsp................................... 56

R

Radiocarbon dating, ash layers...------------ 7,8

RaduJapolyclada........................... 31,41,43

Radulaceae ..............._......._... 43

Ramalina almquistii....................... 14,32,50

on cliff summits-.-----..--------------.-- 14

scoparia.................................. 14,50

Ramalina-Xanthoria community ----------- 10,14

Ranunculaceae.-.-._-------.-.-_---------.-.. 39

Ranunculus hyperboreus hyperboreus.......... 23,39

occidentalis............................... 31

insularis..................... 15,19,22,30,39

nelsoni............................... 22,39

reptans................................ 23,24,39

trichophyllus.............................. 37

trichophyllus.................... 23,24,25,39

Ranunculus-Fontinalis community........... 10,25

Ranunculus-Hippuris community............ 10,23

Ranunculus-Limosella-Isoetes community.---- 10,23

Raphidonema................................. 54

gracile................................. 54; pi. 2

References, cited, general listing-.______ . 58

selected, algae and protists............... 55

Regosols ................................... 6

Rhacomitrium aaciulare....................... 24,46

breeipes................................ 29,45,46

canescens......................... ........ 46

ericoides...................... 13,26,27,29,45,46

fasciculare.............. 13,14,20,28,30,32,33,46

lanuginosum... 13,16,18,19,28,29,30,32,33,45,46 

Rhacomitrium-Andreaea-Schistidium community. ---.- ... - . 10,28 

Rhacomitrium-Schisiditum- Ulota community.. 10, / ? 

Rhinanthux.--................................ 27

minor borealis.......................... 22,27,40

Rhizocarpon atroalbescens..................... 33,48

geographicum............................. 33,48

hochstetteri................................ 32,48

Rhizopoda................................... 55

Rhododendron camtschaticum camtschaticum... 29,36 

Rhytidiaceae.-._....................__... 47

Rhytidiadelphus loreus............ 16,18,20,22,31,47

squarrosus....................... 13,31,41,43,47

triquetrus.............................. 16,18,47

Riccardia..................................... 40

latifrons. ........................ 18,21,40,41,43

pinguis....................... 19,24,31,43,45,46

sin uata................................... 20,43

Rinodina turfacea............................. 19,49

Rosaceae..-.. - -- __ _ ._ ___ 39 

Rubiaceae.................................... 39

Rubus arcticus stellatus................. 16,18,20,39

chamaemorus.......................... 16,18,39

Ruderal habitat.....-..-..----..-,.......... 10,25

Rumei fenestratus............................ 38

obtusifohux............................ 12, 22,38

Sagina crassicaulis......................... 14,19,34

intermedia........ ..................... 27,30,34

ocddentalis............................. 12,27,34

Page 

Sagina-Ceratodon-Equisetum community...... 10,27

Salicaceae..-.__.... .... ._.__..... 39,58

Salii......................................... 7,57

arctica.................................... 58

crassijulis... ........ 15,16,18,28,29,32,39,58

crassijulis .............................. 58

cyclophyUa....................... 16,29,30,32,39

polaris pseudopolaris...................... 39

rotundifolia......................... 28,29,32,39

Salix-Empetrum community.......... _.... - 10,28

Samples, soil-profile, chemical contents....... 9

Saiifraga bracteata............................ 14,39

foliolosafoliolosa.......................... 30,39

hirculus ................................ 31,39

punctata insularis .......... 16,19,21,27,30,39

Saxifraga-Potentilla-Draba community........ 10,.74

Saxifragaceae. ................................ 39

Scapania irrigua........................... 17,25,43

paludosa............. 17,19,20,21,24,31,40,41,43

scandica............................ 20,29,41,43

subalpina................................. 23,43

undulata ...................... 19,24,25,43,45

Scapania-Nardia-Marsupella community..... 10,2.7

Scapania-Siphula community...__...._.-._ 10,24

Scapaniaceae..-...----.--..----.-.-...-.-.-.- 43

Scenedesmusacutus........................... 51,54

armatus.................................. 54

brevispina................................. 54

ecornis.................................... 51,54

quadricauda............................... 54

maximus ........................... 54

spinosus.................................. 51,54

Schistidium apocarpum-------------------- 13,28,46

maritimum...................... 14,15,41,44,46

Schistidium-Rhacomitrium-Andreaea community.-.--.---------.-.--------.-. 10,28

Schistidium-Ulota-Rhacomitrium community.. 10,73 

Schizthrix calcicola........................... 50

lardacea...... ............_..-..--.--.... 50

Scotiella nitalis............................... 51

Scrophulariaceae........ _ _--.-.-------_---- 39

Scytonena mirabile.... - - - 50

Sea-cliff habitat.............................. 10,73

Selaginellaselaginoides.---..--....... -... 21,24,40

Selaginellaceae--.........._....... ----- 40

Selected references on algae and other protists-. 55 

Senecio pseudo-arnica.....-..----..... ...-.. 12,3.5

Senecio-Honckenya community - -..-..-..-- 10, 1% 

Sibbaldiaprocumbens--.--.............. 20,27,29,39

Siphulaceratites........................ 21,24,43,50

Siphula-Scapania community................. 10,24

Snow-bed community.-------.--------------- 10,37

Soils, algae...---.-,._--.-----.------------.-- 55

chemical composition _______.. --._ 8,9

classification .....___........ ..... 5

fungi.........-.._..----...--_------------ 55

Solifluction habitat...--.---...----------.- 10,28

Solifluction terrace community.....-...-....- 10,29

Sparganiaceae...--......--......--.---.-.-.-. 40

Sparganium hyperboreum...--.---.--.--... 24,31,40 

Sphaerophoraceae.......--..-....--....-..--. 49

Sphaerophorus globosus...... 16,18,19,28,29,31,33,49

Sphaerozosma granulatum..... ......--.-...--- 54

Sphagnaceae .............-....---.-...--.. 44

Sphagnum... ................................. 40

bog community.----.-.--------------- 10,20,44

capillaceum--....................... 16,21,41,44

compactum................ 20,21,24,30,31,40,44

Empetrum heath habitat...---.. ------ 21

girgensohnii- ... -_____--..___.._.- 19,21,22,44

lindbergii..- -.-.............--- .--.-.... 31,44

magellanicum................. 21,31,41,43,44,45

papillosum................... 20,21,22,31,41,43

squarrosum. ............ 21,22,23,31,41,43,44,45

subsecundum. -...--... .. ....-.-.-. 16,21,44

teres.......................... 19,21,22,43,44,45

Pige 

Sphenolobus marginalus....................... 42

minutus............................ 18,33,41,43

grandis............................... 43

saccatulus................................ I"),43

Splachnaceae.-. - - _ _ 47

Siachybotrys atra........_................... 56

Staurastrum margaritaceum................... 54

punctulatum pyjmaeum- ............... 50

Stauroneis alabamaeangulata...--------------- 55

anceps.................................... 5"?.55

linearis.................---........... 52

nobilis.---..-----.--..--.................. 55

Stellaria calycantha calycantha.... ....... 12,20,27,34

media..................................... 34

ruscifolia aleutica.......................... 30,34

sitchana bongardiana...................... 20,34

Stemphylium ilicis............................ 56

Stenopterobia intermedia....................... 55

Stereocaulaceae...----.._--- -_-- -- 49

Stereocaulon alpinum......................... 13,49

denudatu m................................ 49

inter medium..............................27,49

tomentosum............................... 49

alpestre............................... 27,49

vesuvianum..................--.--.--..... 30,49

denudatum............................ 49

Stichococcus subtilis..------------------------- 56

Stictapulmonarialinita-.._... ................ 49

Stictaceae.-....-----.----.----.------------- 49

Stigonema minutum.. ..-.--------.-........ 50

ocellatum.....-.-.--..-..---.--.........-- 50

turfaceum... ..-- .---. .-....-.--.. 56

Strand habitat ---------------------------- 10.72

Stream habitat. - - - -- -- 17,*4 

Streamside community. -- -- - 17, 19 

Streptopus amplexifolius...................... 19,37

Studies, floristic- --.--.--------------------- 57

Substrates, miscellaneous.~ --. . 33 

Subulariaaquatica......................... 24,?5,39

Subularia-Callitriche community-..-...-. 10,24 

Synechococcus cedrorum pallidus.....--..... 53; pi. 2

moi'or ----------------------------------- £1,53

notatus....-....---..---.-------.-------.- 53

Tabellaria flocculosa .....--...---.-...---.-. 51,55

Takakiaceratophylla.------................... F1,40

Takakiaceae-_.... . .--. .-... ... 40

Taraxacum aleuticum.---..-------..---.-...-- 35

trigonolobum.....-------........--.....--. 14,35

Teloschistaceae..-----.- _-.----...--.-.--.. 49

Teloschistes candelarius. .....---.---....----- 49

Tetracladium marchalianum................ 55; pi. 2

Tetraplodon mnioides...................... 33,45,47

Thamnolia ..-.--. .................. ----- 28

s ubidiformis ----------------------------- 50

vermicularis ...................... 16,28,32,50

Thelypterislimbosperma ................. 31,33,38

phegopteris . ................ . 31,38

Thuidiaceae....-----..-.--------------------- 47

Tofieldia coccinea..---------------- 16,18,32,37

Toninialobulata.............................. 20,48

Tortula mucroni folia....................... 13,14,47

Trichoderma viride ---------------------------- 56

Trichophorum caespitosum. ------------------- 17,35

Trientalis europaea arctica-............. ----- 16,38

europaea arctica aleutica-.. ... --_ - 38

Trifolium repens.............................. 37

Trisetum spicatum.....----------------------- 56

spicatum alaskanum...................... 36

alaskanum molle...................... 36

Tritomaria quinqwdentata------... ------- 30,41,43

Tundra without permafrost.---------------- 6

Tundra without permafrost, profile ___ 7

U 

Ulota phyllantha.................. 13,14,15,19,44,46

Ulota-Rhacomitrium-Schistidium community.- 10, IS 

Umbelliferae. __..-_ .......... - 40
66 INDEX

Page Page Page

Umbilicaria eylindnca,......................... 32,50 Vahlodea atropurpurea........................ 58 Viola langsdorffii.... .............. 15,1% 17,22,32,40

hyperborea................................ 50 atropurpurea latifolia ......_----- 16,18,31,36 Violaceae..._.. .--..-.....--. -..-..._. 40

proboscidea .................... 13,29,32,33,50 paramushirensis...................... 37

torr«/octo ----------------------- 50 Vascular plants, annotated list..... 34 W, X, Z

UmbUiranaoeae 50 Vegetation on island . ._. - 4 Wet sedge-meadow community............... 10,*/

sneaceae _-_. ........ ..-------.... Veronica, americana........................ 24,25,40 Wood, imported, plant communities ...-. 33

v grandiflora................................ 29,40 Xanthona, candelaria.................... 14,19,32,49

Vaccinium-..-............................ 7 serpylUfolia humifusa......... 20,21,24,27,31,40 elegans.................................... 14,49

uliginosummicrophyllum................. 29,36 stellen................................. ib,dA4U onseastacks 14

«>«« mtiS'idaea................................. iitnen ifi Tlerrwcorio maura............................. 14,50 " ----------------------- ib Xanthona-Ramalina community. .... 10, U

minus................................ 18,36 Verrucariaceae --.. -.. .. -. .- _ 50

Vacularia vireseens............... _ - _ 55 Verticillium sp .-.-.__ .. -- 56 Zygogonium ericetorum........................ 51
PLATES
PLATE 1

FIGURE 1. Actinotaenium cucurbita (Br6b.) Telling.

2, 3. Cosmarium variolatum Lund.

4-6. C. tatricum Racib.

7. C. decedens (Reinsch) Racib.

8, 9. C. obliquum Nordst.

10. C. subundulatum Willie.

11. C. subcrenatum Hantzsch forma.

12. C. meneghini Br6b. var concinnum Rbh.

13. C. laeve Rbh. forma.

14. C. subpachydermum Schmidle forma.

15. Oocystis parva W. et G. S. West.

16-19. Chlorococcum humicola (Nag.) Rbh.

20-23. Gloeodinium montanum Klebs.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 648 PLATE 1

DRAWINGS OF ALGA SPECIES
PLATE 2

FIGURE 24. Synechococcus cedrorum Sauv. var. pallidus, var. nov.

25. Oscillatoria amoena (Kutz.) Gom.

26. 0. facilis, sp. nov.

27. 0. splendida Grev.

28. 0. grossegranulata Skuja.

29. 0. sp. cf. amphibia Ag. forma. 

30-32. Pseudanabaena granulifera, sp. nov. 

33-41. Chlamydomonas alaskensis, sp. nov. 

42-46. C. subangulosa, sp. nov.

47. Planctococcus sphaerocystiformis Korschik.

48. Raphidonema gracile, sp. nov.

49. Ankislrodesmus braunii (Nag.) Collins var. 

50-53. Tetracladium marchalianum De Wildem.

U.S. GOVERNMENT PRINTING OFFICE: 1969 O 350-274
GEOLOGICAL SURVEY PROFESSIONAL PAPER 648 PLATE 2

DRAWINGS OF ALGAE AND OTHER PROTISTS