A PHYTOSOC!GLGSICAL $7qu or AN UPLIFTED MARINE BEACH Rance NEAR Pam? BARROW, AiASKA Thesis for the Degree of M. S. M!CHIGAN STATE COLLEGE John James Koranda 1954 '1 «3 3c; .33....» , This is to certifg that the thesis entitled A PHYTOSOCIOLOGICAL STUDY 01' AN [1?le MARIN! m RIDG Hm POINT WV, ALASKA presented by John James Koranda. has been accepted towards fulfillment of the requirements for M.— degree in m Datefl M //- / 0-169 A lHITOSOCIOLOGICAL oTUDY OF AN UPLIFTAU EARIHE BEACH RIDGE INS/tit POINT dAiuLON, ALASKA By John James fipranda AN ABDTRACT buhmitted to the School of Graduate Studies of hichigan state College of Agriculture and Applied Science in partial fulfillment of the requirements for the degree of hAbTbh OF SCIENCE Department of Botany and Plant Pathology Approved Triage This phytosociological study concerns the vegetation of an old uplifted marine beach ridge on the Arctic Coastal Plain, near Point Barrow, Alaska. lt was undertaken to obtain some information about the distributional patterns of Arctic plants, to study the particular conditions presented by an old uplifted marine beach ridge and, finally, as a study in phytosociological methods and techniques. The vegetational complex on the beach ridge was sampled quanti- tatively by list—quadrats placed randomly in'aibelt-transect. These quantitative data were evaluated by three statistical methods: (1) Comparisons of the observed frequency distribution with the Poisson series. Density histograms —- which diagrammed the actual number (2) of individuals occurring in each quadrat. Measures of aggregation —— statistical indices which evaluate (3) the amount of relative aggregation in the distribution of a species. Few species were found to have distributions that approximated the Poisson distribution. Frequencies occurred, in most cases, beyond the end of the theoretical distribution. The density histograms revealed relationships existing between the contour of the beach ridge, soil pH, and the distribution of many Species. The localized distributions of species, as shown on the histograms, were compared with the relative order indicated by the measures of aggregation. Close agreement between the histograms and 331293 and the measures of aggregation existed when an apprOpriate quadrat size and number were used. The effect of the number of quadrats and the size of the quadrat upon density, frequency, and the measures of aggregation were analyzed. Twenty-one quadrats were found to be as effective a sample as 100 if the object of the sampling was merely to determine the dominant Species. 1f phytosociological relationships were to be investigated, a larger sample was recommended. The relative order of the Species, in regard to the degree of aggregation, was affected by the number of quadrats used, eSpecially when micro-distributional factors were present. It appeared from this study of tundra vegetation, where habitat differences occur in relatively short distances, that a 1.25 percent sample of the total area was not detailed enough to describe accurately the distrib- utional aSpects of the vegetation. It was shown that density was prOportional to quadrat size, when density was defined as the number of individuals per quadrat. ’Frequency, however, did not react to quadrat size in the same manner. The degree of aggregation apparently influenced the change of frequency that occurred when the quadrat size was reduced. Such Species as Ericphorum scheuchzeri and §§xifraga cernua were cited as examples of this condition. The measures of aggregation were also affected by quadrat size and tended to misrepresent the degree of aggregation when smaller quadrats were used. This was especially evident when the species-were distributed very contagiously or locally. A comparison was made of two transects, which represented distinct areas on the beach ridge. On the basis of Species alone, the higher area could be defined as a more xeric area. The density differences of certain species occurring on these transects were also considered. A photographic method was used to measure the percentage cover for two species, Salix rotundifolia and Petasites frigidus. Salix rotundifolia exhibited low cover values on extremely'moist sites and The frequency of areas where frost action was relatively more severe. s. rotundifolia in the photographic sample was very close to the quadrat-frequency of this species. The relationship of percentage cover to density was determined for Petasites frigidus. This Species occurred on a hummock area where varying conditions of micro-relief were eXpressed in the size of the individual and, thus, the percentage of the quadrat covered. The varying conditions of micro-relief were effective in creating discrepancies in the density-percentage cover relationship. An analysis of the structure of the beach ridge and the component soils revealed several correlations with vegetative characteristics. Soil-frost phenomena were shown to create a habitat which at times excluded a Species but which, more often, merely caused a reduction in density. The particular conditions of active zone depth and snow cover were also discussed. The beach ridge was found to have a deeper active layer than either the marsh or polygonal habitats. The early withdrawal of snow cover was also indicated by the series of readings made in several sites and actual observation. LIDT UF thhhfihUbS Blackman, G. E. 19h2. statistical and ecological studies on the distri- bution of Species in plant communites. Ann. Bot. o: 351--30o. Uver-dispersion in grassland communities and the Jour. scol. 24: Clapham, A. Ho 1930. use of statistical methods in plant ecology. 232-’2510 Curtis, J. T. 195C. The interractions of certain analytic and synthetic phytosociological characters. .Ecology 31: h34~~455o Evans, F. C. 1952. The influence of size of quadrat on the distributional patterns of plant pOpulations. Contrib. Lab. Vert. Biol., Univ. of Rich. 54: l--lA. Fracker, S. B., and H. A. Brischle. 19AA. Measuring the local distri- bution of hibes. Ecology 25: 283--303. hopkins, D. M., and H. S. Sigafoos. 1950. -Frost action and vegetation patterns on Seward Peninsula, Alaska. Geol. burv. Bull. 971,-0. Flora of Alaska and Yukon. Parts I-IK. Lunds Hulten, Eric. 1941-1949- Universitats Arsskrift, h.F. Vols. 37-45. Johnson, D. w. 1919. Shore processes and shoreline deveIOpment. wiley and sons, New York. thinnies, N. G. 193h. The relation between frequency index and abundance as applied to plant pepulations in'a semi-arid region. Ecology 15: 263--282. holina, E. C. 1942. Poisson's Emponential Binomial Limit. D. Van hostrand Co., New York. ppsL l--L7. Uplifted beach ridges and first generation lakes in the hex. R. 1953. Unpublished data. Barrow, Alaska area. oigafoos, R. S. 1951. Soil instability in tundra vegetation. Jour. sci. 51: 281--298. Ohio whitford, P. B. l9b9. Distribution of woodland plants in relation to succession and clonal growth. Ecology 30: l99-208. ‘ A PHYTOSOCIOLOGICAL STUDY OF AN UPLIFTED MARINE BEACH hIDGE, NEAR POINT BARRON, ALASKA By John James Koranda A DISSERTATION Submitted to the School of Craduate Studies of Michigan State College of Agriculture and Applied Science in partial fulfillment of the requirements fer the degree of MASTER OF SCIENCE Department of Botany and Plant Pathology 1954 TASLE ACKNONLEDGhthS . . . LIST OF THAT FIGbAno . LIST or Tam TABLES . . -. . . LIST or Arm-mix Fist-has . . . LIST or APPENDIX TABLL-JS . . . INTRODUCTION . . . . Review of Literature . . . Description and history l‘lETHO mLOG Y o o o 0 Sampling of Vegetation . Photographic Sampling 3011 Sampling 0 o o o o TREATMENT OF DATA . . . Quadrat Data . . DISCUSSION AND ALALYSIS OF DATA . Comparison of Quadrat Numbers . Density and frequency . . heasures of aggregation . SURUhary o o 0 Comparison of Quadrat Size . Density and frequency . measures of aggregation . UF CUT. 1‘15th . . . . . . . . . . . . Area . . . . . . . . . . . . . . 0 O I. O . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ii ii iii vi g... 10 ll 13 13 l9 l9 19 2O 25 26 26 28 -r Summary Comparison of Transects Discussion of Photographic Sampling . Discussion of Soil-Vegetation Melationships SUMMARY LIST OF i‘LEFLliEIfiES . APPENDIX 30 31 33 37 53 57 61 AC KM) WLEDGNEN T5 The author wishes to express his sincere thanks to hr. Daniel Q. Thompson, who provided the opportunity for this study and constantly encouraged the writer during the progress of the field work. Grateful acknowledgment is also due to Dr. William B. Drew, major professor, for assistance and guidance in the preparation of manuscript and the innumerable other details that inevitably appear in the process- ing of scientific data. Dr. Ira L. Wiggins, Scientific Director of the Arctic Research Laboratory, graciously allowed the writer to use his herbarium and aided him is becoming familiar with the local flora. Drs. Charles L. Gilly and George W. Parmalee are to be sincerely thanked for their expert asSistance in the preparation of the manuscript. He is also extremely grateful to Dr. John W. Thomson, University of Wisconsin, for the lichen identifications which served as reference Specimens for this study. The moss specimens were graciously identified by Drs. William Steere and Howard Crum. The alga Specimen was checked by Dr. Gerald w. Prescott. TEXT'FIGURES I. II. III. IV. V. VI. VII. TEAT TABLnS l. 2. LIST OF TEAT FlGUhfiS Diagram of the Point Barrow Area adapted from an Aerial PhOtograph 0 o 0~ o o o o o 0 Percentage of cover as determined from h".x 5" photographs, Salix rotundifolia . . . . . Density—cover Relationship, Petasites frigidus Active Layer Depths During Summer, Transect 111 Active Layer Depths During Summer, Transect I HistOgram of Hummock and Frost Boil Occurrence pH Values - Transect I, Transect III . . . LIST OF TdAT TABLES Soil Data - mechanical Analysis - Chemical Analysis . . . . . . Snow Retreat in Various Habitats Point Barrow, Alaska Depth of Active Zone Transects Point Barrow, Alaska . . . . . . . . .. ii PAGE . 7 . 35 . 36 . 49 o 50 - 51 . 52 . A6 0 1+8 APPENDIA FIGURE I. II. III. IV. V. VI. VII. VIII. IX. XI. XII. XIII. XIV. XV. XV. XVI. AVII. XVIII. XIX. XX. LIST OF APPnthk FIGLRHS Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Density Histogram histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram Histogram of of of of of of of of of of of of of of of of of of of of Poa arctica . . . . Arctagrostis latifolia EriOphorum scheuchzeri Carex aquatilis . . . Luzula nivalis . . . Luzula confuse . . . HriOphorum angustifolium Saxifraga cernua . . Stellaria laeta . . . Petasites frigidus . . Senecio atrOpurpureus . Saxifraga punctata . . Potentilla emarginata . Saxifraga foliolosa . Vaccinium vitis-idaea . Saxifraga flagellaris . Thamnolia vermicularis Dactylina arctica . . Cetraria islandica . . SphaerOphorus globosus Cornicularia divergens PAGE 62 03 0A 05 66 67 68 69 7O 71 72 73 7h 75 76 76 77 78 79 80 .11}?qule FlGUnms‘ may; XXI. Density Histogram of Cetraria nivalis . . . 82 XXII. Density Histogram of Cladonia Sp. . . . . 83 XXIII. Density Histogram of Peltigera canina . . . 8A XXIV. Density Histogram of Lobaria linita . . . 85 XXV. Density Histogram of Parmelia omphalodes . . 8o XXVI. Density Histogram of Cladonia uncialis . . 87 XXVII. Density Histogram of Peltigera aphthosa . . 88 XXVIII. Density Histogram of Stereocaulon evolutoides' 89 XXIX. Density Histogram of Caloplaca subolivacea . 90 XXX. Density Histogram of Cetraria richardsonii . 91 XXXI. Frequency Distribution - Poa arctica . . . 92 XXXII. Frequency Distribution - Arctagrostis latifolia . . . . . . . . . . . 93 XXXIII. Frequency Distribution - Eriophorum scheuchzeri . . . . . . . . . . . 9A XXXIV. Frequency Distribution - Carex aquatilis . . 95 XXXV. Frequency Distribution - Luzula nivalis . . 96 XXXVI. Frequency Distribution - Luzula confusa . . 97 XXXVII. Frequency Distribution - EriOphorum angustifolium . . . . . . . . . . 98 XXXVIII. Frequency Distribution - Saxifraga cernua . 99 XXXIX. Frequency Distribution - Stellaria laeta . . 100 XL. Frequency Distribution - Petasites frigidus . . . . . . . . . . . . 101 XLI. Frequency Distribution - Senecio . . . . 102 atropurpureus . . APPENDIX FIGURE XLI]. XLIII. xuv. va. XLVl. XLVII. XLVIII. XLIX. L. LI. LII. LIII. LIV. LV. LVI. LVII. Frequency Distribution Frequency Distribution VitiS-idaea o o 0 Frequency Distribution emarginata . . . Frequency Distribution foliolosa . . . . Frequency Distribution vermicularis . . . Frequency Distribution Frequency Distribution Frequency Distribution uglobosus . . . . Frequency Distribution divergens . . . . Frequency Distribution Frequency Distribution Frequency Distribution Frequency Distribution canina . . . . . - Saxifraga punctata - Vaccinium - Potentilla - Saxifraga - Thamnolia - Dactylina arctica - Cetraria islandica - Sphaerophorus - Cornicularia Cetraria nivalis . Cladonia gracilis Cladonia sp. . . Peltigera Frequency Distribution - Lobaria linita . Frequency Distribution Frequency Distribution - Psoroma hypnorum . - Cetraria delisei . PAGE 103 10A 105 100 107 108 109 110 111 112 113 11h 115 116 117 118 APPEK DIX TABLcAS l. 2. 5. 6. 11. 12. 13. LloT 0F APPENDIX TABLnb Transect Ill - statistics . . . . . . Comparative Transect Illa . . . . . . Transect III and Transect IIIa Comparisons DenSity and Frequency 0 o o o o o o Transect III and Transect Illa Comparisons Leasures of Aggregation Grasses, Sedges, and Rushes. . . . . . Transect III and Transect llIa Comparisons Measures of Aggregation Miscellaneous Flowering Plants . . . . Transect III and Transect Illa Comparisons Measures of Aggregation LiChens o o o 0 o o o o o 0 V. o Transect III and Transect Illa Comparisons Measures of Aggregation LiChens o o o o o o o o o o o 0 Comparison of Transect I and Transect III . Transect IIIb . . . . . . . . . . Transect III and Transect IIIb . Density and Frequency . . . . . . . Transect III and Transect IIIb leasures of Aggregation Grasses, Sedges, and Hushes- . . . - . Transect III and Transect Illb Measures of Aggregation Miscellaneous Flowering Plants_ . . . . Transect III and Transect IIIb Measures of Aggregation- Lichens o o o o o o o o o o o . PAGmS 119 120 121 122 I23 12h 125 126 127 128 129 130 131 vi vii 1A. Transect III and Transect IlIb Comparisons Measures of Aggregation LiChenS o o o o o o o o o o o o 132 INTRODUCTION Ecological studies of Arctic plants have been mainly of a descrip- tive nature and few investigators have attempted to ascertain the phyto— sociological aspects of Arctic plant communities. The time required for a detailed phytosociological study often prohibits investigations of this type. The extremely short vegetative period and lack of facilities in the Arctic, in general, have deterred investigators from conducting pro- longed studies in these areas.‘ The Arctic Research Laboratory at Point Barrow, Alaska, under the Office of Naval Research, for several years has provided a base of Operations fer many informative and detailed studies which will add considerably to the knowledge of Arctic biOIOgy. This study, phytosociological in approach, was undertaken to obtain some information about the distributional patterns of Arctic Species, to study the particular conditions presented by an old uplifted marine beach ridge and, finally, as a study in phytosociological methods and techniques. Although only limited soil data were obtained, some of the relationships between plant species and the soil can be observed, where the soil is a relatively unstable medium because of soil-frost activity. Review of Literature Wiggins (1951) in his study of the distribution of vascular plants on polygonal ground near Point Barrow, Alaska, mentioned the interesting and unpredictable variations in the vegetational cover from site to site and the accompanying problems in the ecology of such micro-habitats. This study is concerned with the same beach ridge upon which one of Wiggins' sites was located. He feund that there was more variation in the habitats and, also, more vegetational variation on the beach ridge site than in any other of his study areas. The Sparser lichen cover on this ridge was also noted. Acock (19h0) conducted a study on raised shingle beaches and asso- ciated habitats in an inner fjord region in Spitzbergen. hany of the same species involved in his study were present in the vicinity of Point Barrow. He noted that soil-frost phenomena (fissure polygons), by causing soil heterogeneity, gave rise to a hyperdiSpersion of species that was similar to the banded communities occurring on the shingle beaches where differential snow cover seemed to be the determining ecological factor. Russell and Wellington (19AO) made detailed physiological and ecological studies on Arctic vegetation on Jan hayen Island, north of Iceland. They analyzed climatic and edaphic factors in view of varia- tions in the type of vegetation and found that shelter, time of snow retreat, water supply and available mineral nutrients were the most important factors. Nitrogen deficiency was found to be common in that area and in Arctic regions in general. An exhaustive study by Gelting (193A), on the vascular plants of East Greenland, described the differences in vegetation occurring between the outer coast and the inner fjord region. The phytogeography of num- erous Arctic species was discussed and ecological and distributional data were furnished. This paper affords an excellent comparison with North American Arctic studies in regard to circumpolar distributions of species. Bacher (1951) described the distributions of plants in the circum- polar area in relation to ecological and historical factors and discussed many of the Species that were mentioned in the aforementioned articles. Lindsey (1952) used photographic techniques in studying the ancient beaches above Great Bear and Great Slave Lakes, in the Canadian Arctic. Lichens and flowering plants were evaluated according to percentage cover obtained from photographic data. Several authors have studied soil—frost phenomena (cryopedologic features) in relation to vegetation in Alaska with very interesting results. Hepkins and Sigafoos (1950) and Sigafoos (1951, 1952) studied the relationship of soil-frost activities and vegetation occurring on the Seward Peninsula and defined the processes referred to by the general term frost action. Benninghoff (1952) discussed soil-frost and vegetation interactions in Alaska and outlined a vegetation-soil-frost cycle. hany other authors have conducted soilvfrost and vegetation studies in Alaska and some of these citations will be included in the list of references. In addition to those articles which were pertinent to this study, several recent publications were used as source material for phytosocio- logical formulae and computations. Curtis (1950) reviewed and clarified the usage of such terms as density, frequency, abundance, and many others. He obtained eXperimental data from artificial populations to illustrate the interrelationships of phytosociological characters. Evans (1952) analyzed the influence of the size of quadrat on the distributional patterns of plant pOpulations in his studies on an old- field community in southeastern Michigan. The size of quadrat was shown to have an effect upon frequency and abundance and to have a marked effect upon the measures of dispersion. One of the most extensive reviews of the quantitative aspects of plant distributions was made by Goodall (1952). A complete list of phytosociological references was included. Description and History of Study Area This study was conducted in the summer of 1953 at Point Barrow, Alaska, under contract with the Office of Naval Research and the Arctic Institute of North America. Reconnaissance, to attain familiarity with the local flora and to locate suitable study sites, was made during the previous summer, 1952. The study area was located in the vicinity of Point Barrow, Alaska, which lies at the northernmost extremity of the Arctic Coastal Plain (Latitude - N. 71° 18' 56", Longitude - w. 156° 36' 16"). The coast of the plain at this point is prolonged into a triangular area, bounded on the west by the Chukchi Sea and onthe east by the Beaufort Sea. The geomorphic history of the coastal plain was one of emergence and uplift, with progradation of the coastline occurring in the posteglacial period (Rex.1953). Intermittent uplift has produced a pattern of shorelines and barrier beaches. The beaches were, and still are,being formed by the combined activities of waves, currents, and ice action. As the coastal area was uplifted, the beaches were drained and raised, and various sediments such as loess were deposited upon the original sand and gravel. Permafrost (permanently frozen ground) is present through— out the coastal plain and is interestingly modified in these uplifted beach ridges. In the vicinity of the Naval Contractor's Base, near the Eskimo village of Barrow, several old beach ridges were quite evident. They occurred inland on the tundra and could be recognized by their slightly higher elevation and better drained surfaces. The northernmost of these ridges was 500 to 2,000 yards from the present Arctic Ocean beach. It was over two miles long, roughly "U" shaped, with the Open part of the "U" facing southpsoutheast (approx. 1520 E. of magnetic N.). This northernmost ridge was chosen as the site of the study area. Text Figure l is a diagram of the beach ridge and vicinity. The western portion of the ridge extended southwest, paralleling the coastline, and was transected by the drainage of a salt lagoon called Middle Salt Lagoon. 0n the eastern portion of the ridge, there was a high area that attracted attention because of its interesting and uncommon assemblage of Species in the immediate vicinity of the coast. This area was, very likely, the most xeric site on the eastern portion of the beach ridge. The ridge sleped gradually downward from the seaward side to the landward side which formed the inside of the "U" shaped structure of the entire ridge. The width of the ridge at the study area was slightly over one hundred meters. The inside perimeter of the ridge, within the basin enclosed by the ridge, had been modified by the action of waves as evidenced by several small terraces of gravel and a definite wave-cut margin, especially on the eastern portion of the ridge. There were also several persistent ponds, remnants of an old drained lake or lagoon, in the center of the basin enclosed by the beach ridge. " ampmz amps uochaom one ammo; ....'...‘. .‘300 I I I \ x . .l.-..l‘.. C 00!. :oowmq Conan gm \/ , Ix EOLM «030,??ch $3.3. 20;th page; I \l) ‘ ~I .ud V 4 H . ‘. .. c . .. .J no awn as: .H magmas same oxmq Loose gnonm oamom .Konmdd coowmq hood: sash mopmhnxxu o smmoo capon< METHODOLOGY dampling of Vegetation To sample the qualitative and quantitative vegetational changes that occurred on the beach ridge, a belt transect one meter wide and 100 meters in length was outlined with stakes and string line. 'The beach ridge, at the transect zone, was oriented LAO west of magnetic North. The transect heading was 55° East of magnetic North so that the transect was nearly at right angles to the long axis of the beach ridge. The transect began at the lower or inland side of the ridge and extended to the top or seaward side which was 2.7 meters higher than the lower end. The transect was surveyed by the plane table method so that the actual profile of the slepe was known. The area of the transect was sampled by the list-quadrat method. After preliminary trials, a quadrat size of 0.05 Squareemeter was decided upon since larger sizes gave frequencies that were too high. The quadrats were distributed through the one hundred square-meter area of the transect by a method designed to avoid bias and achieve a random placement of samples. Each square-meter of the transect was sampled with a 20 x 25 cm. quadrat. The placement of the quadrat in each square-meter was determined by drawing from ten lots, each of which represented a partic- ular position in the square-meter. The lots were rescrambled after each drawing. Each of these positions, actually 0.1 square-meter in area, was taken to have an inside and an outside position which was alternated with each quadrat. Thus, the sampling procedure was a combination of a random and a regular system of quadrat placement. The placement of the 100 quadrats was determined before the counting began. To facilitate the placing of the quadrat within the square-meter being sampled, a wooden frame with an inside area of one square-meter was constructed. The inside of the frame was divided into the ten pre- scribed positions by strings and the alternating portions were determined merely by placing the quadrat in the 0.1 squareemeter compartment so that it filled either the inside or the outside half. The quadrat frame, 20 x.25 cm. in size, was made of plexiglass strips 1/8 inch thick and one inch wide. On each 25 cm. side, three notches were filed, dividing the side into four quarters. Two sets of brass pins, 15 inches long, 3/32 inch in diameter, connected by 12 inches of fine, linen cord, were also used in the counting procedure. The counting procedure was as follows: (1) the quadrat was placed on the ground in the pre-determined position within the square-meter being sampled; (2) one set of brass pins was pushed into the ground next to the first set of notches and the string stretched taut across the quadrat; (3) the plants in the first quarter were then counted; (4) the second pair of pins was put into the ground next to the second set of grooves, and counts were made in the second quarter of the quadrat; (5) the first set of pins was then moved to the third set of grooves, and -the plants in the third and fourth quarters of the quadrat were counted. The string and pins also served to hold the quadrat frame tightly to the lO ground and to maintain a constant inside area during the counting. This method proved to be a very effective way of counting the stunted or diminutive vegetation characteristic of high Arctic regions. The counts for each quarterequadrat were kept separate on the data sheets for statistical reasons. Notes concerning the physical features of each quadrat were also recorded on the data sheets. One hundred quad- rats were counted on the 100 square—meter transect area; thus, the sanple was five percent of the total transect area. Nomenclature of the higher plants mentioned in this paper is in accord with Hultén (l9h1--l9h9). Representative lichen collections were identified by Dr. John Thomson, and these were used as reference speci- mens for quadrat determinations. Photographic Sampling Because certain Species presented growth forms that made it impossi- ble to obtain actual density counts, a photographic sampling technique was devised to evaluate these species. A A" xKS" Speed Graphic camera mounted on a sturdy tripod was used to photograph the 20 x.25 cm. quad- rat from a position directly overhead. The photographs were made on PluSeh film, through a green 11 filter, using a number 11 flashbulb at approximately two feet. The camera was focused so that the quadrat completely filled the ground glass. Twenty-five photographs were taken of quadrats located every fourth meter in the transect. The photographic sample was, therefore, a 1.25 percent sample of the total area of the ll transect. The prints of the quadrats were enlarged to sake the quadrat on the photographic paper the same size as the original quadrat. The quadrat size was scored on a plexiglass sheet, l/8 inch thick, and 12 inches square. This area was divided into 100 compartments. Per— centage cover was determined from the photographs for Salix rotundifolia and Petasites frigidus by the use of this plexiglass grid. In determining the percentage cover for these species, the grid was placed on the photograph and each Square containing a plant or a portion of a plant was counted as one unit. Salix rotundifolia cover values were determined because actual density could not be ascertained due to its prostrate, mat-like growth form. Cover values for Petasites frigidus were compared with actual density values for the quadrats that were photographed. The resulting data furnish a gauge of the effective- ness of cover values and their relationship to density. Soil Sampling Some of the differences in vegetational cover on natural areas are due to variations in soils and their physical and chemical makeup. There- fore, a series of trenches was dug at various points along the transect and soil samples were taken from the strata that were revealed. The soil' samples were analyzed both chemically and mechanically, whenever possible. The mechanical analysis of the soil samples was made using the hydrometer method according to the procedure outlined by Bouyoucos (1936). The Soils Testing Department, Michigan State College, performed the active, test for the elements in the chemical analysis. Three times during the progress of the study, soil samples were collected from a stratum just below the vegetative mat for immediate pH analysis. The samples were analyzed with a Beckman pH meter, Model G. These samples were taken at stations ten meters apart along the transect. Active layer depth measurements were also recorded several times during the summer at stations ten meters apart on the transect. A metal probe was pushed into the ground until the permafrost table was reached and then this depth was measured with a rule. Data for active layer transects in three different habitats are also considered in the discussion of soil and vegetation relationships. l3 TRLATMENT OF DATA Quadrat Data In analyzing the quadrat data, three methods were used to describe the distribution of each species of lichen and flowering plant on the transect.v The densities of the species occurring on this transect were determined by actual stem count. Density histograms were made for most of the species. Saxifraga caespitosa (six individuals in quadrat 45), Juncus biglumis and several lichen species were omitted. The histograms are included in Appendix Figures I to XXXI. On these histograms, the profile of the ridge was plotted and also the average pH value for the ten soil-sampling stations on the transect. The profile or elevation values are represented by the solid line while the pH values are symbol- ized by the dotted line. Each bar represents the number of individuals that occurred in the quadrat with the appropriate scale at the right of the histogram. The transect distance of one hundred meters is represented by the horizontal axis. Mean density of the species in the quadrats and the frequency of occurrence in the one hundred quadrats are also placed on the histogram. The distribution of a particular Species, whether it be aggregated or scattered uniformly, can be observed directly from these diagrams. The relationship of elevation and soil chemistry to species density is also evident. However, such factors as soil moisture, soil-frost pheno- nena (frost scars or "boils"), and hummocks which influence the lb micro-distribution of many of the Species, cannot be reasonably portrayed. The lichens presented an unusual growth form and densities do not have the same value when applied to them as when applied to the higher plants. However, for the purposes of denoting the quantitative changes that occurred on this beach ridge and transect, they can be used, although with certain reservations. Each distinct thallus, unconnected to another, was counted as one unit. The species present on the transect were not fragmented into minute pieces and, therefore, could easily be counted. This was especially true of Cetraria nivalis, Q. islandiga, Lobaria linita, Parmelia omphalodes and others. In the case of such fruticose species as Cornicularia divergens, §phaer0phorus globosus, and Dactylina arctica clumps were counted. These Species occurred in small clumps of thalli or lobes of a thallus and very likely these clumps represent individuals. The measurement of growth forms of this type is very difficult and any method used is subject to error. With the same observer doing all the counting, some of this error was minimized. Flowering species such as Luzula nivalis also presented a problem in arriving at a stem count. This Species grows in clumps of one to several rosettes of leaves. Each rosette was counted as an individual in this case, since it was theoretically capable of producing one or more flowering Spikes. The rest of the flowering plants counted were easily recognized as individuals. Qélig rotundifolia, as indicated before, was measured with the photographic method, in terms of percentage 15 of the quadrat covered. To study the distribution of the species on the transect and to evaluate this condition in terms of the relationship of the observed frequency distribution to the theoretical distribution eXpected for a random dispersion, the plates comparing these two distributions were prepared. Poisson's eXponential binomial limit tables by Molina (19A2) were used to plot the theoretical distribution line. Several of the lichen Species have been omitted in this series of plates. From these Appendix Figures (XXXII - LVIII), it can be seen that the distribution of few species approximates the random condition. The actual frequency class contributing the most to aggregation can be ob— served on the figures. I . In addition to the frequency distribution figures, four statistical indices or measures of aggregation were computed for all the Species. These four indices have been used by many investigators in phytosociology in an attempt to find a statistical measure that will effectively evalu- ate the aggregation occurring in plant distributions. These four measures are those prOposed by McGinnies (1934), Clapham (1936), Fracker and Brischle (19th). and Whitford (19A9). All of these measures use the basic frequency-density relation, P 3 100(l-e-d). Clapham (1936) used a characteristic of the Poisson distribution, the equivalence of the variance of the distribution to the mean, and called this measure the relative variance. Blackman (l9h2) called this measure the coefficient of dispersion. The relative variance 16 in the present paper is indicated by SZ/X; Departure from unity (l) is the measure of non-randomness, within the limits imposed by sample size. McGinnies (1934) compared the actual densities with theoretical densities at the observed frequency level. This measure is represented by D/d in the tables. Fracker and Brischle (IQAA) used the relationship Dad/d2 in describing the distributions of BEESE and included a very helpful table that can be used to obtain the expected density at a given frequency level according to the frequency-density equation above. In both of the measures described above, 2 represents the actual mean density of the Species and g_represents the theoretical mean density expected at the observed frequen- cy. Whitford's index employs the relationship between frequency and abundance (the ratio of the number of individuals found to the number of samples of occurrence). In the present paper, A represents this index, which is equal to lOO(D)/frequency2. The value 2, again, is equal to the observed mean density. These phytosociological indices or measures were very well outlined and explained by Curtis (1950). Tables listing these measures for individual Species are found in the Appendix. The Species are listed in the following categories: 1) grasses, sedges and rushes; 2) miscellaneous flowering plants; 3) and lichens. In order to ascertain the effect of quadrat size upon these measures of aggregation, the data for two quadrat sizes were compared. For convenience of discussion and comparison, the one-hundred 0.05 square— meter quadrats were designated as Transect Ill. The counts for each 17 0.05 square-meter quadrat were recorded by quarters on each data sheet and, by using the counts for the first two quarters of each quadrat, data for a 0.025 square-meter quadrat size were obtained. These smaller quadrats were designated as Transet IIIb. It was the intention at the beginning of the study to make a similar transect on an area adjacent to this high portion of the ridge so that the quantitative and the qualitative differences in vegetation might be observed. Because of a shortened field season, a transect as detailed as the original transect (III) could not be completed. Therefore, a transect (designated as I) was made approximately 1/8 mile north of Transect III on the same beach ridge. The number of samples in Transect I was reduced to 21 quadrats, 0.05 square-meter in size; these were Spaced in every fourth.meter-unit of the transect. The total area of Transect I was the same as Transect III, that is, 100 Square meters. To provide a basis for comparison of these two transects, a compa— rable series of quadrats was separated from the original transect (III) and designated as Transect Illa. The quadrats selected for this purpose were from every fourth meter on the transect area. The data concerned in these comparisons are listed in the Appendix Tables. Appendix Tables 1 and 2 contain the basic data for Transects III and IIIa. These data were used in making the computations for the succeeding tables. The comparisons of Transects III and Illa involving mean density, frequency, and the four measures of aggregation are contained in Appendix Tables 3 to 7. In Appendix.Table 8, Transects Illa l8 and I are compared. Comparisons between the 0.025 and the 0.05 square- meter quadrats (Transect Illb and III) are listed in Appendix Tables 9 to 1h. l9 DISCUboION AND ANALYSIS OF DATA Comparison of Quadrat Numbers Density and frequency. From the comparison of Transect III and Illa in Appendix Table 3, it can be seen that in the grasses, rushes, and sedges there is some shifting of the relative densities of certain Species, §.g., Ericphorum schuechzeri; but the two grasses showing the greatest relative density have retained their same position. Eriophorum scheuchzeri dr0ps from third place in Transect III to fifth place in Transect Illa, which consisted of only 21 quadrats. This can be explained by the restricted distribution of g, scheuchzeri which evidently was not sampled as effectively by the smaller number of quadrats and therefore, is under-represented. The tiny rush, Juncus biglumis, was not recorded in the 21 quadrat transect. The frequencies of several Species remain the same for both series of samples. In the case of the miscellaneous flowering plants in Appendix Table 3, the same five species are retained as the five densest Species with the usual shifting among the more Sparsely represented species. §§Xifraga cernua dropped from the most dense species on Transect III to the fourth most dense species on Transect Illa. An inSpection of the histogram for this species (Appendix Figure VIII) Shows that it occurred on the transect at three places which are correlated with relatively moister soil conditions. 0f the miscellaneous flowering plant species, §axifraga_cernua had the most discontinuous distribution. The distributions of Eriophorum scheuchzeri and Saxifraga cernua can be termed micro-distributions and it is evident that the reduced number of samples was not sensitive enough to record the actual density conditions for these Species. The lichens appear to be more effectively sampled by the reduced number of quadrats than the flowering plants since the comparison in Appendix Table 3 shows little disagreement, and then only in the minor or less conspicuous elements of the lichen flora. These comparisons indicate that the 2l-quadrat transect produced nearly the same results in regard to mean density and frequency as the larger sample. Poor representation is noted in the case of contagiously distributed species. Measures of aggregation. In analyzing the distributions of the Species on the transect according to the measures of aggregation, tables comparing the effect of both quadrat number and quadrat size Upon these measures have been prepared. Appendix Tables A to 7 are concerned with the comparison of the effect of the number of quadrats upon these measures. In Appendix Table A, the Species of grasses, rushes, and sedges are arranged in decreasing order of non—randomness, according to the four measures of aggregation. All of the measures, except Nhitford's index (A), give values that indicate a definite non-random condition. Fracker and Brischle's index, D—d/dz, showed the least difference in the arrange- ment of the species between the two transects. This index placed 222. arctica as the second most random species while according to the other 21 three measures, this Species is indicated to be either the second or the third most aggregated. A consideration of the histograms (Appendix Figures I - VII) for the grass, rush, and sedge species indicates that the relative order of non-randomness according to D—d/d2 is the most sensitive and best de- scribes the distributions of these Species. Luzula confusa, indicated the third most aggregated by D-d/dz, exhibited a sporadic distribution of densities which are correlated with its tussock type of growth form. Eriophorum angustifolium and g. scheuchzeri clearly Show the most aggre- gation on the histograms. All of the measures and both of the transects reveal the Same general order of species, except for the instances mentioned, and indi— cate, as did the mean density and frequency comparisons, that the 21- quadrat sample was nearly as efficient as the loo-quadrat sample. Appendix Table 5 lists the computations of the same four measures of aggregation for the miscellaneous flowering plants. There is not the agreement in this comparison between the measures and quadrat number, as there was in the grasses, rushes, and sedges. It is possible that the 0.05 square-meter quadrat was too small to evaluate these Species with the measures of aggregation. However, when the species indicated as the most aggregated or contagious by the measures of aggregation are examined on the histograms, the order of species in Appendix Table 5 appears to agree with the actual conditions. Again D-d/d2 agrees very closely with the relationships indicated by the histograms. Whitford's measure (A) 22 also approximates quite closely these relationships. Vaccinium vitis-idaea is indicated to be the most non-random species by the indices D—d/d2 and A. Its distribution, confined to the hummock tops in the upper portion of the transect, would seem to warrant this position. Potentilla emagginata, the most random species according to D—d/d2, has mean density almost the same as that of Vaccinium vitis- igggg, but its frequency is twice as large. One of the characteristics -of aggregated species is that for any given density, the frequency value is lower than would be expected. Petasites frigidus is another Species with a high mean density and a low frequency and is indicated to be very contagiously distributed by all of the measures. As with the grasses, rushes, and sedges, the more sensitive of the four measures are apparently Dw-d/d2 and A. The smaller number of quadrats, however, did not maintain the same relationships of the measures that were indicated in the larger sample. Appendix Tables 6 and.7 compare the effect of quadrat number upon the measures of aggregation for the lichen Species. All of the measures, except the index D/d, indicate that Stereocaulon evolutoides is the most non-random Species. The histogram (Appendix Figure XXIX) for this species shows that it occurred on only two portions of the transect. At both locations there was gravel at the surface of the ground. The high density in the first quadrat was associated with the gravel edge of the lower end of the transect. The relatively high density present on these micro-habitats was indicated by the measures in placing Stereocaulon 23 evolutoides in the most aggregated position. No eXplanation can be given for the disagreement of the index Q/d in these comparisons. It is noticeable that for the smaller sample of 21 quadrats, D/d gives values which indicate Stereocaulon evolutoides the most non-random, which is in agreement with the values indicated by the other indices and the lOO—quadrat transect. Fracker and Brischle's index, D—d/dz, does not ShOW'mUCh agreement between transects, except that Stereocaulon evolutoides is assigned the most aggregated position in both samples. Parmelia omphalodes exhibited a distribution on the transect that can be correlated with the presence of bare earth. This lichen was found on the bare areas at the base of the transect, irl and around the frost-boils found on the steeper portion (25 to A5 m.), and on the hummocks at the upper end. Evidently Parmelia Qmphalodes had a frequency which was high enough to lower the aggregation index.and the localized distribution of this species is not indicated. The value of descriptive data like the histogram is realized in analyz- ing such distributions. A repeated pattern of multimodal densities is evident on the histo- grams for the following lichen Species: Cetraria islandica, Q. nivalis, Dactylina arctica, Cladonia Sp., Sphaerophorus globosus, Thamnolia germicularis, and Cornicularia divergens. At each point where these species decreased or ceased to occur, a moist area was present. The line representing elevation shows a slight depression at each of these sites (20 m. and 75 mt). These lichen species were xerOphilic in their 24 distribution in the entire Barrow area, being found in their highest densities on polygon and hummock teps, and on the beach ridges, both recent and old. It is believed that Cetraria richardsonii was limited to the lower portion of the transect (See Appendix Figure AAXI) by the predominantly northeasterly winds. The wind struck the beach ridge and the transect at the higher point and a slight lee effect was exerted by the 2.7 meter difference in elevation between the lower and upper ends of the transect. The large, strap-like thalli of this lichen were found loosely tangled among the grass stems and offered the most wind resistance of any of the lichens. The lichens are the most difficult of the plants to analyze accord- ing to ecological and phytosociological methods. The methods used in this study could hardly be used in temperate or even in boreal areas. The lush growths of lichens occurring in central Alaska and in some north temperate forests could not be analyzed using actual densities. But because of the stunted or reduced condition of the lichen flora in the Barrow area, and eSpecially on this beach ridge, it was possible to use a sampling method such as list-quadrats. Higgins (1951) remarked that large areas densely covered with either Sphaggum or lichens were uncommon in the immediate vicinity of Point Barrow. Even under these conditions, it is realized that an arbitrary method has been used but the correlations that have been observed seem to warrant the use of this method. Density counts of such life forms, although they could not represent the actual numbers of individuals present, serve as an index or gauge of abundance which otherwise might not be measureable. The application of photographic techniques may solve part of the problem of evaluating life forms such as lichens and mosses. Summary. The effect of the number of quadrats upon the values of density, frequency, and measures of aggregation was described in this section. It was found that density and frequency were represented reason- ably well in the smaller sample. however, when a species was aggregated in reapect to some micro-habitat feature, the smaller number of quadrats was liable to inaccurately represent the relative status of the species. In phytosociologial studies, concerned with these micro-distri- butional relationships, the larger sample of the total area is desirable. If sampling merely to record the dominants of an area, the smaller number of quadrats appears to produce essentially the same results as the larger, more detailed sample. The 2l-quadrat transect was a 1.25 percent sample of the total transect area while the loo-quadrat transect was a 5 percent sample. In these comparisons, it can be seen that the effectiveness of the measures of aggregation is dependent upon the number of quadrats and the distributional factors present in the habitat. The measures when applied to the grasses, rushes, and sedges gave comparable results for both series of quadrats. In the miscellaneous flowering plants and the lichens, there was a tendency for the order of the Species to be 26 significantly changed when the smaller number of quadrats was employed. The Species most commonly misrepresented in degree of non-randomness were those exhibiting relationships to micro-distributional features. Therefore, if a study is concerned with the phytosociological Vcharacteristic of aggregation or contagion, the level of sampling should be high enough to compensate for micro-distributional effects. It appears from this study of tundra vegetation, where habitat differences occur in relatively short distances, that a 1.25 percent sample of the total area was not detailed enough to describe accurately the distri- butional aSpects of the vegetation. The dominant species, however, could be determined from the smaller number of quadrats. Comparison of duadrat Size Appendix Tables 10 to 14 are concerned with the comparison of quadrat sizes. As described previously, the quadrat size was halved and the resulting data were used in exactly the same manner as the original transect data. Mean density, frequency, and the four measures of aggre- gation were computed. Appendix Table 9 merely lists the data for the IOU-quadrat transect, using the quadrat size of 0.025 square-meter. This transect is designated as Transect IIIb in the Appendix Tables. Density and frequengy. ln Appendix Table 10, the density and frequency values for the two quadrat sizes were compared. Among the grasses, rushes, and sedges there was no disagreement at all. The means (i) seemed to be approximately halved in the smaller quadrat data. when density is considered as the number of individuals per quadrat, it becomes directly prOportional to quadrat size. The high mean, low frequency condition of EriOphorum scheuchzeri was repeated in the smaller quadrat but the difference between density and frequency was not as large as in the larger quadrat. Curtis (l950) and hNans (1952) both stated that estimates of rela- tive frequency are affected by the size of the quadrat employed while density values are proportional. Apparently EriOphorum scheuchzeri represents this condition since the density value was almost halved, yet the frequency drOpped only one percent in the smaller quadrat data. The resulting discrepancy in the measures of aggregation for this Species illustrates the significance of this condition and will be discussed in the analysis of the measures of aggregation. Saxifraga cernua, in the comparison in Appendix Table 10, drOpped to fourth most abundant species. Of the miscellaneous flowering plants, é. EEEEEE had the most discontinu— ous distribution on the transect. The histogram (Appendix Figure V111) for this species showed three density modes coinciding with relatively moister soil conditions. This species was also poorly represented in Transect Illa, in which the number of quadrats was reduced. It was evi- dent that the smaller quadrat size did not record the actual density conditions for this Species as well as it did for the other Species which did not have such a discontinuous distribution. Curtis (1950) found in his studies on artificial pepulations that there is some indication that highly contagious species are over-repre- sented by relatively small quadrats. The unusually high frequencies L3 indicated for daxifraga cernua and Eriophorum scheuchzeri in the smaller quadrat data appear to illustrate this condition since these species were contagiously distributed. The rest of the density values for the miscellaneous flowering species showed a very good agreement in the order of the species, especial- 1y among the less dense species. The lichen densities showed little Variation between the two quadrat sizes so that the Species maintained their same relative positions in each transect. The proportional relationship of quadrat size and density seemed to apply to the lichens as well as the flowering plants. The frequency relationships of the flowering plants were also repeated in the lichen species, with high frequencies indicated in the smaller quadrats. Cetrari§_islandica had a frequency of 86 percent in the 0.05 squaresmeter quadrats and 83 percent in the 0.025 square-meter quadrats. Measures of aggregation. Appendix Tables ll to 14 coxpare the measures of aggregation for the species in the two transects, using the two different quadrat sizes. The index D-d/d2 showed the most sensitivi- ty in Appendix Table 11 where the grasses, rushes, and sedges are listed. It was noticed in the comparison of the densities and frequencies that Eriophorum scheuchzeri had an unusually high frequency in the smaller quadrat data. In Appendix Table 11, according to D—d/d2, the relative position of g, scheuchzeri drops to fifth most aggregated species. In the 0.05 square-meter quadrat data, g, scheuchzeri had been considered as the most non-random, or the second most non-random. 29 A consideration of the histOgram (Appendix Figure III) for this species indicates that Eriophorum scheuchzeri does exhibit a highly contagious distribution. It is evident that the high frequency observed in the smaller quadrat data results in a more random condition being a indicated by the measure, D-d/d2, than actually exists. The three other measures in Appendix Table ll apparently are not sensitive enough to ~g record this condition and show little disagreement between the two quadrat sizes. The various measures of aggregation for the miscellaneous flowering Species are listed in Appendix Table 12. Saxifraga cernua, as noted previously, shows an unusually high frequency in the smaller quadrat. Fracker and Brischle's index, D-d/dz, indicates a more random condition for é, cernua in the smaller quadrat data. The unusually high frequency and the resulting discrepancy in the measures of aggregation revealed in the values for Eriophorum scheuchzeri seem to be repeated in the values for Saxifraga cernua. On the basis of the effect of quadrat size upon the measures of aggregation for these two species, the assumption of Curtis (1950) that contagious Species are over-represented in relatively small quadrats appears to be substantiated. Appendix Tables 13 and IA are concerned with the measures of aggre- gation computed for the lichen Species, using the two quadrat sizes. The relative variance (Sz/f) in this comparison is affected the most by the smaller quadrat size while D-d/dz, the most sensitive of the measures when applied to the flowering plants, showed little disagreement between 30 the two quadrat sizes. Nhitford's measure, &’ also shows close agree~ ment between quadrat sizes in this comparison. The measure used by thinnies, Q/d, does not appear to be affected by the change in quadrat size but the order of the species in degree of non-randomness is actually not as indicated by this index. The high frequency for Cetraria islandica in the 0.025 Square—meter quadrats has not resulted in the usual change in the relative randomness. If a change were to occur, it would be expected in the D-d/d2 values, which indicated this change in the flowering plants. It is possible that lichen distributions cannot be measured by these indices, even though individuals can be arbitrarily delimited and density values obtained. The asexual methods of reproduction (especially in the Arctic Species), the means of dissemination, and many other characteristics of the lichens, may produce distributional effects that render this type of data ineffect- ive. Summary. From these various comparisons, from which the most conSpicuous examples have been drawn, it can be concluded that density, when it is defined as the number of individuals per quadrat, is affected by change in quadrat size in 8 directly prOportional manner. Frequency exhibits more complex relationships, especially when the Species is contagiously distributed. Variations in frequency are related to the effect of quadrat size upon the measures of aggregation. The misrepresen- tation of the degree of aggregation shown by a Species is often the result of relatively small quadrats being used. Whenever randomness is indicated in plant pOpulations,while at the same time it is apparent that micro- distributional influences are present, more than one quadrat size should be used. In this case, an increase in quadrat Size would be in order because of the tendency of smaller ouadrats to misrepresent the degree of aggregation. The use of nested quadrats, or counting the quadrats in sections and keeping the count separate for each section, are methods which can be used to record data for several quadrat sizes simultaneously. Since populations have been Shown to conmonly occur in aggregated distributions, and random dispersions are seldom found, it would seem that investigators in phytosociology should consider the employment of several quadrat Sizes an indeSpensible part of their technique. The- nature of contagious distributions has been studied by several bio- metricians and phytosociologists. heyman (1939): Cole (l9h6) Archibald (1948, 1950), Thomas (l9h9), Thomson (1952) and Beall and Rescia (1953) have deve10ped or modified statistical distributions which can be fitted to the observed frequency distributions found in biolOgical pOpulations. Comparison of Transects Transect I was established on a lower or more mesic area on the beach ridge to provide a basis of comparison with the higher portion sampled by Transect Ill. Transect I was located 1/8 mile north of Transect III on the same beach ridge. The same sampling procedure was used to study both transects. Only 21 quadrats were counted on Transect I, but they were Spaced every fourth meter. The area sampled in this transect was the same so that in Transect IIIa which comprised 21 quadrats selected as previously mentioned from Transect Ill. Comparable data from these two transects are summarized in Appendix Table 8. Several differences are observable in this comparison. The relative densities of several grasses and sedges vary significantly. Luzula nivalis and p, confusa occurred in reduced densities on the moister area (I). These two Species were found in the more xeric sites in the entire Barrow area. Alopecurus alpina was present in moderately high densities on Transect 1, sharing co-dominance with 223 arctica and Arctagrostis latifolia. Gelting (l93h) observed that A10pecurus alpina occurred eSpecially in bogs and other damp places in East Greenland, where it is often dominant with Arctagrostis latifolia. Alepecurus alpine was absent from Transect llla which was evidently too xeric for this species. The comparatively higher density of Carex aggatilis on Transect I, was indicative of the higher moisture requirements of this Species. Carex aguatilis occurred in highest densities in the polygonal ditches and trough and was decidedly hydrOphilic in its distribution in the Barrow area. The absence of several Species of miscellaneous flowering Species from Transect 1 also indicated the influence ofia more hydric habitat. Vaccinium vitis-idaea, CassiOQe tetragona, Saxifragagflagellaris, g, gppositifolia (not in quadrats), were all characteristic of the higher area (Illa) but were absent on Transect 1. Certain lichen Species such as Cetraria nivalis, Cornicularia divergens, §phaer0phorus globosus, and Cladonia gracilis occurred in considerably lower densities in Transect 1 than in TFlLSCCt Illa. Theac same Species showed density maxima in the more xeric sites on Transect Illa. Further evidence of their xeric tendency was provided by their distribution in the Barrow area on polygon tOps, hummccks, and other beach ridges. On the basis of the Species present in Transect III, of which Transect Illa was a part, this higher area can be considered to represent a more xeric habitat. The soil data further confirm the condition that was noticed at the beginning of tne study on tne basis of vegetational characteristics. Discussion of Photographic Sampling As noted previously, Salix rotundifolia and Petasites frigidus were sampled using a photographic method. Text Figures I and II are based on the resulting data. The cover values of Salix rotundifolia indicate that this Species has a rather wide tolerance of moisture conditions. Reduced cover values, however, were noted in extrenely wet areas. The low cover Values in the 20 - 25 m. area (Text Figure l) were related to the seepage occurring at the 20 meter station and the soil—frost activity on the steeper portion of the slope (25 m.). The absence of this species in the upper portion of the transect (80 to 100 m.) is esplained partially by the soil-frost activity in the hummock area. The polygonal trough at 75 m. was evidently too moist and, in addition, competition with grasses and 3A sedges was greater there. The quadrat frequency of Salix rotundifolia was at percent which agrees reasonably well with the 08 percent obtained in the photographic sample. The density-cover relationship of Petasites frigidus is graphically represented in Text Figure 11. It was possible to obtain both density and percentage cover values for this Species. The relationship of density and percentage cover varies as indicated by the text figure. The discrepancies, where they occur, are believed to be related to the micro-habitats present in the area where Petasites frigidus occurred. The pattern of hummocks in this area presented a mosaic of soil and relief conditions upon which this species was distributed. The varying conditions of the micro—habitat were eXpressed in the vigor of the plant and hence its vegetative growth. Plants on hunmocks that were not affected by soil-frost activity as much as others had larger vegetative parts (leaves) and hence possessed a greater cover value. BeCause of these and similar factors in the environment, densities will not agree at all times with cover values. .X/ EQ (J o > o h OOH «0 Gal ON; 0m. 0.: Qmi 00. can 00. OOH. 00 mm om b Ampvpofivaoomcmne as AK. mo 00 mm Om m4 oq mm on mm om ’ ’ \yoég I. «I I./ .>pzaue maaomwvcsaou Kwamn madmnmouonn =m & :d 50pm nocfisuouou mm uo>oo ommpcoouom .HH massed same ma OH m l O 5 WHObd-P-HOG O.m m.m O.m IO 3 OH ON om o: On 05 s1 (3 o > m h OOH 00:161-th mm Ampmuoevpoomcmha 382. R msosmmommsos «mom mmow l 0 «.7360 R lill I hufimcoa .mmwga A.qv assawwtn moswmmumm dflnmcowpmaom uo>ooahpwmcoa HHH poomcmha .HHH musmaa sway ma pa .D .m. #04” LTJH0>€U+3-HOQ E. V \x) \j Discussion of Soil-Vegetation Relationships he beach ridge has been described previously as representing a more xeric site in comparison with the polygonal and marsh habitats. The most evident reasons for this condition were the gravel substratum and the slight elevation. Active layer measurements were made'at ten stations on Transect III and I several thnes during the summer (Text Figure III and IV). These measurements were made with a metal probe. It was possi- ble to feel the presence of graVel with the probe and as each depth read- ing was made, the presence or absence of gravel was noted. At the base of the beach ridge, the lowest point on the transect, the active layer was the deepest with a definite stratum of gravel, six inches thick, occurring just above the permafrost. The gravel was felt at all the stations up to the sixty meter area. In the trench at 60 meters, the gravel was present in a 2 to 3 inch stratwa directly over the permafrost table. Approximately 17 inches of weakly stratified soils were superimposed upon the gravel layer at this station. At 05 meters, however, the gravel disappeared and could not be felt with the probe in the remaining upper portion of the ridge. A trench dug at the 100 meter station failed to disclose any gravel on the permafrost table. it is possible that the gravel layer in the upper 30 meters of the ridge was absorbed into the permafrost. The average active layer depth at the 100 meter station was slightly over 30 inches. From the active layer readings made with the probe and the trenches dug at various points on the transect, it is known that a gravel 9 substratum existed under a najor part of the ridge and that it occurred just above the permafrost table. huller (1947) stated that percolation of ground water takes place upon permafrost gradients. From Text Figure III, the gradient of the permafrost table on Transect 111 can be seen to be quite steep, especially from 70 meters to the base of he transect. From field notes, taken while the trenches were being dug, a phenomenon can be described which illus~ trates the percolation of ground water on the permafrost table. As the trench at the 30—meter station was being dug, an upwelling of water occurred as soon as the gravel layer was entered. A similar upwelling took place in the trench at the 2C-meter station. The water in both trenches appeared to be under a slight hydrostatic pressure. The trench at 20 meters soon filled and remained filled for the entire summer. The 30-meter trench event‘ally drained and did not fill again. water was not encountered in any of the other trenches above the 60— meter trench or below the 20-meter trench. In the trench dug at the O-meter station, 1.3., the base of the transect, an active layer of 22 to 31.5 inches was noted with approximately six inches of loose gravel upon the permafrost table. The gravel was slightly damp and plant roots had penetrated into this layer. There was no water in this trench at the time of digging nor did it fill with ground water at any- time during the summer. The reason for the lack of water in the lowest trench at 0 meters was very likely the "hump" in the permafrost table that occurred between .20 meters and the base of the transect. In every series of active layer readings, this area (10 m.) had a shallower active layer than the one either above or below it. This bulee upward in the permafrost table exerted a damming effect upon the percolation of ground water. The area from 18 to 21 meters was a very moist site and this saturation could be reasonably attributed to the damning of ground water just below the 20- meter station. The comparatively shallower active layer beneath the lO-meter area could be easily correlated with the thicker vegetative mat occurring at this zone. Wiggins (1951) found that areas of bare ground nad deeper active layers than areas with a thick vegetative mat. Salix rotundifolia shows high cover values on tnis area; loa arctica, Arctagrostis latifolia Luzula nivalis, and L, confuse are present in moderate to high densities here. The histograms for these Species in the Appendix Figures from I to VII show the actual densities on this area. The mosses also con- stituted a very important part of the vegetative mat on this area. Ceratadon purpureus, Tomenthypnum nitans, Drepanocladus uncinatus, Distichwm flexicaule, and Oncophorus pplycarpus were the most abundant Species of mosses. Thalli of the alga, hostoc commune, were also found in this moist area from 10 to 20 meters. The diagram of active layer depths for Transect 111 (Text Figure III) shows a pronounced deeper layer during the whole summer on the area from 20 to 50 meters. The steepest gradient and also the most bare ground on the whole transect occurred in the area between 20 and 50 m.. 1,0 The bare ground was in the form of frost ”boils" or ”scars". Text Figure V shows ten-meter cumulative values for the frost "boils" and hummocks in the transect. The associated vegetation of this area was typical of bare earth habitats where pioneer Species are apt to be found. Continued frost action on this area very likely kept the Species from ever establishing a vegetative mat and hence this area is always in a colonizing stage. h0pkins and Sigafoos (1950) concluded that the concept of climax vegetation must be modified when applied to tundra vegetation. They stated that the tundra differs from climax assemblages in that bare areas, and those covered by pioneer plants, are intimately mixed among areas covered by vegetation representing the highest stage in the succession. Most of the flowering Species either drOpped in density on this area (20 to AC m.) or they did not occur here at all. Such Species as Luzula nivalis, L, confusa, Potentilla emarginata, Papaver radicatum, Draba glabella, Saxifraga foliolosa, é. punctata, s. caespitosa, Stellaria laeta, and CassiOpe tetragona are found on this area. Poa arctica and Arctagrostis latifolia occurred here in reduced density. Saxifraga cernua was virtually absent from this area and appeared to occur in highest densities when in moister conditions. The histograms indicate the distributions of these species in regard to this area (Appendix Figures I to XV). CassiOpe tetraggna and Papaver radicatum were present in the quadrats from 25 to 35 meters. They were represented Ll by only one or two individuals. Several lichen Species exhibit density maxima on this bare earth area. Most of the fruticose Species occurred densely here, while the foliose Species such as gobaria linita and Peltigera canine were more hydrOphilic. As indicated on the diagram of the active layer depths for Transect III (Text Figure III) a consistently deeper active layer was also present at the upper end of the transect. Again this feature can be related to the presence of bare ground in the hummock area which occurred from 80 to 100 meters. This site was a poorly deveIOped polygonal area with many small hummocks. The tops of most of the hummocks were bare except for a few plants of Arctagrostis latifolia, Poa arctica, vaccinium vitis- ig§g§_and lichens which did not form a continuous vegetative cover. The lack of turf, the exposure to winds, and the absence of snow cover all favored the formation of hummocks in this area. The low site at 75 meters represented a polygonal trough with possibly a melting ice wedge supplying the moisture which made it more hydric. The distribution of Eriophorum angustifolium on the histogram (Appendix Figure VII) is to be noted with reSpect to this feature. This species was commonly found in areas much moister than the beach ridge. The pH values recorded for ten stations each on Transect III and Transect I are diagrammed in Text Figure VII. The more acidic condition consistently found on the upper portion of Transect III was very likely related to the increase in elevation and the leaching associated with the local drainage conditions. The soil in the hummocks, which were present in the area from 80 to 100 meters, was a light yellow-brown color and the pH range was usually from A.6 to A.9. The troughs between the hummocks, with slightly darker soil, did not have pH values of much over 5.0 so that the entire micro—relief of the Site was decidedly acid in reaction. The distribution of Vaccinium vitis-idaea on the transect is an example of the acidOphilic tendency of 3 Species which was related to the micro-relief present on the hummocks previously described. This Species was entirely restricted to the upper portion of the transect (III) where it occurred only on the hummock tops. In Text Figure VII, the pH values for Transect III and I are com- pared. Although these transects were located at least 1/8 mile apart, a repetition of the high pH values is apparent at the same points on both transects. The active layer depths for Transect I were not as deep as on the higher area (Transect III) but this would be expected since the vegetative cover was more continuous and denser on Transect I. At the 20 meter station, an unusually high pH of 7.h was recorded twice during the summer. In the trench that was dug at this station, a well— defined stratum of fine, blue—gray, clayey-sand was discovered just above the permafrost table. This layer was approximately two to three inches thick. when tested with HCl, this soil effervesced indicating free lime. The chemical analysis of this soil (Sample #h, Text Table I) shows a comparatively high potash and calcium content. The soil sample was examined for Foraminiferan tests but only unidentifible fragments were found. It is possible that this stratum of fine, compacted clayey-sand was an organic deposit laid down beneath a pool or pond. Pools of this type commonly form on the backslopes of beach ridges and were seen on the present beach ridges of the Arctic Ocean in the vicinity of Point Barrow. Whatever its formation, this stratum evidently created a rich soil habitat in the lower portion of the transect.) Saxifraga flagellaris occurred on the lower or alkaline portion of the transect between 0 and L5 meters. This species was consistently found on the lower part of the beach ridge for several hundred yards on either side of the transect. Gelting (193A),in his studies on the vascular plants of East Greenland, considered this Species to be calciphilic. He found this plant eSpecially numerous on the raised marine terraces in company with Saxifraga caespitosa. Six Specimens of S. caeSpitosa were present in the quadrat at as meters. The soil samples from the trenches were analyzed chemically by active tests and mechanically for sand, Silt, and clay. The results of these tests are given in Text Table I. It was not possible to analyze every stratum that was found; therefore characteristic strata were analyzed from several trenches. For example, a light brown sandy loam was present on most of the transect just beneath the dark humic layer associated with the vegetation. Samples #1 and #10 in Text Table l were from this layer and Showed little difference except in pH. Sample #10, from the lOO-meter station, was lighter in color which was possibly an indication of the degree of Ah leaching. In the 20 to AS meter area, this sandy loam was mixed with the gravel stratum which appeared here at the surface. The soil in this area was very homogeneous and did not show the stratification apgarent in the other portions of the transect. This was eXpected, since the area from 20 to 50 meters had the largest number of frost "boils" of any portion of the transect, and these features are an indication of the churning (congeliturbation) occurring in a soil. Sigafoos (1951) stated that the formation of frost scars or "mud Spots" was the initial result of congeliturbation on most sites. Sample fit was from the blue-gray clayey-sand in the trench at 20 meters. The chemical analysis of this stratum correlated with the high pH values obtained at this point. Sample #3 was a sample of the dark—brown humic layer associated with the vegetative mat. It was present, either in an unbroken or broken form, on most of the transect.' This layer varies from pH 6.6 at the base of the transect to pH 4.8 at the lOO—meter station. Com- pared to a sample eight inches below in the sandy loam, this dark-brown humic layer had twice as much potash and calcium. The possibility that some of hese soil conditions were present over a large area on the beach ridge was indicated by the similarity of the pH patterns for both transects, I and III. The trench at the loo-meter station yielded several interesting samples which are possibly related to the history of the beach ridge. L5 The gravel layer was not found in the trench at 100 meters but, as stated previously, it was possible for this layer to have been absorbed into the permafrost. A narrow stratum of dark-brown peat, one to two inches thick, was found on tOp of the permafrost table in this trench. Sample #9, from a 15 inch depth in the trench at 100 meters, showed a noticeable increase in phosphorus, potash, calcium, and manganese. The color of the soil at this depth was gray. The light brown layer, previ- ously described, and this deeper gray stratum very likely represent two periods in the development of the beach ridge. Rex (1953) considered these beach ridges in the Point Barrow area as being loess-covered. Evidently the upper layer, light brown in color, represented the eolian deposition while the deeper, gray layer was associated with the earlier, marine history of the beach ridge. Eolian deposition of similar material was common in Alaska where deep deposits have been laid down in relative- ly short periods. The peat layer (ph - 5.2) present on the permafrost table at the lOO-meter station represented the remains of the initial vegetation of beach ridge. The 0.5 ppm of iron recorded in sample #8 was associated with the peat layer. In connection with certain ecological studies conducted in the Point Barrow area by Thompson (1953): active layer depth transects were established in three distinct habitats (Text Table 11). One of the mm pauoxo .mowaafiE\mpth :fi mosam> HH< * m. 0 O O o om ms.m o.« o o o o a: m.mo o.m> 0.0: o.m>H o.mu n.5m mo O.m 0.4 o.m mN.o mN.o mN.o wz N.OH m.oa o.m 0.9m 0.0H m.4 zmmpom 4\H 4 H m\._H J\H m\H m\H .904 .monm N.m H.m m.q m.> 0.0 m.o ma pmoa *mamsaaqa assesses n o.HN m.mm m.mm u N.mm hmao u o.qm 4.mm N.4 I m.5m paflm I 4.4a o.mq no I an vcmm salsa zma spawn =ma spams =oa spaoe=ma spate zm spams =m mumpmalooa myopoaaooa mpopmsuooa mummeIom mummeIO mumposao AmvmpeoSpfipmcoo we as as aw m» as mmamsom aflom mwmhamc< HwOficmnomz cheese same .m. .04 m4 . O.N . o.m a m.m .moe II IIIIII mega poomcmup .a OOH A .s A an.” excesses: mo songs gang I I moss poowcmnp .s OOH M .E H S.” .333 $0.3: no gonna: 330.4 HHH sameness 8292580 30m anon...“ use xoosfii .wo gumopmflm A mHo>co+=-HOE 5' V pH pH pH Values Transect I 700‘ 605‘ /.\ \\ / \\ ~\\\:::,// 000‘ I/‘\:\\\ ><:/ \ ’/O \\ ’/’ p,,"" 5.5‘ 5.0‘ o——-——-—o30 July 1953 u-——————*29 August 1953 14-5‘ 1'0 2'0 3'0 #0 5E) 605 76 86 at E0 pH Values 705‘ O /\ Transect III 7.0. ° ° \ 4‘ \O ”\ 6'5 % //\\’\ “ /\ \\ ””’ T \ X 6-01 " \fA " \ ' .’<.\.4’\’‘1'\ 5-5 \\\ . 500' ‘5}:VR-53“ '\ 4.5 o . o 29 July 1953 h 0 ------- 16 August 1953 x——— 19 August 1953 3.5 ‘ 1'0 2'0 3'0 LIE 5b 66 7b 81) 96 E0 Text Figure VII. Transect (meters) 53 SUhhARY This phytosociological study concerns the vegetation of an uplifted marine beach ridge on the Arctic Coastal Plain, near Point Barrow, Alaska. The vegetational complex on the beach ridge was sampled quantitatively by list-quadrats placed randomly in a belt—transect. These quantitative data were evaluated by three statistical methods: (1) Comparisons of the observed frequency distribution with the Poisson series. (2) Density histograms -- which diagrammed the actual number of individuals occurring in each quadrat. (3) Measures of aggregation -- statistical indices which evaluate the amount of aggregation in the distribution of a Species. Few species were found to have distributions that approximated the Poisson distribution. The density histograms revealed relationships existing between the contour of the beach ridge and soil ph. The histo- grams also furnished a basis of comparison for evaluating the measures of aggregation. The measures of aggregation were found to agree with the distri— butions of the species as indicated by the histograms. The effect of the number of quadrats and the size of the quadrat upon density, frequency, and the measures of aggregation was analyzed. 55 Twenty-one quadrats were found to be as effective a sample as 100 if the object of the sampling was merely to determine the dominant Species. If phytosociological relationships were to be investigated, a larger sample was recommended. The relative order of the Species in regard to the degree of aggregation was affected by the number of quadrats used, eSpecially when micro-distributional factors were present. It was shown that density was prOportional to quadrat size, when density was defined as the number of individuals per quadrat. Frequency, however, did not react to quadrat size in the same manier. The degree of aggregation apparently influenced the change of frequency that occurred when the quadrat size was reduced. The measures of aggregation were also affected by quadrat size and tended to misrepresent the degree of aggregation when the smaller quadrats were used. This was eSpecially evident when the Species were distributed very contagiously. A conparison was made of two transects, which represented distinct areas on the beach ridge. 0n the basis of Species alone, the higher area could be identified as a more xeric site. The measurement of percentage cover was accomplished by a photo- graphic method for two Species, Salix rotundifolia and Petasites frigidus. §§lix rotundifolia was shown to exhibit low cover values on extremely moist sites and on areas affected by soil-frost activities. The frequen- cy of Salix rotundifolia in the photographic sample was approximately 55 Twenty-one quadrats were found to be as effective a sample as 100 if the object of the sampling was merely to determine the dominant Species. If phytosociological relationships were to be investigated, a larger sample was recommended. The relative order of the Species in regard to the degree of aggregation was affected by the number of quadrats used, eSpecially when micro-distributional factors were present. It was shown that density was prOportional to quadrat size, when density was defined as the number of individuals per quadrat. Frequency, however, did not react to quadrat size in the same manner. The degree of aggregation apparently influenced the change of frequency that occurred when the quadrat Size was reduced. The measures of aggregation were also affected by quadrat size and tended to misrepresent the degree of aggregation when the smaller quadrats were used. This was eSpecially evident when the Species were distributed very contagiously. A comparison was made of two transects, which represented distinct areas on the beach ridge. On the basis of Species alone, the higher area Could be identified as a more xeric site. The measurement of percentage cover was accomplished by a photo- graphic method for two Species, Salix rotundifolia and Petasites frigidus. Salix rotundifolia was shown to exhibit low cover values on extremely moist sites and on areas affected by soil-frost activities. The frequen- cy of Salix rotundifolia in the photographic sample was approximately 5c the same as the quadrat frequency of this Species. The relationship of percentage cover to density was determined for Petasites frigidus. The effects of micro-habitat were analyzed in relation to percentage cover and density and found to be related. Variations in the vegetational pattern were found to be correlated primarily with soil-frost phenomena, structure of the ridge, and chemistry of the soil. Climatic factors Such as exposure to wind, and snow cover were also effective. An analysis of the structure of the beach ridge and the component soils revealed several correlations with vegetative characteristics. Soil-frost phenomena were shown to create a habitat which possibly excluded a Species but which, more often, merely caused a reduction in density. The particular conditions of active zone depth and snow cover were discussed briefly. The beach ridge was found to have a deeper active layer than either the marsh or polygonal habitats. The early withdrawal of snow cover was also indicated by the series of readings made in several sites. 57 LIST OF REFERENCES Acock, A. M. 1940. Vegetation of a calcareous inner fjord region in Spitzbergen. Jour. Ecol. 28: 81-106. Archibald, E. E. A. 1948. Plant pOpulations I: A new application of Neyman's contagious distribution. Ann. Bot. 12: 221--235. . 1950. Plant pOpulations II: Estimation of number of individuals per unit area in heterogeneous plant pOpulations. Ann. Bot. 1A: 7--21. Ashby, Eric. 1935. The quantitative analysis of vegetation. Ann. Bot. A9: 779-802. . 1936. Statistical ecology. Bot. Rev. 2: 221--235. . 19b8. Statistical ecology. Bot. Rev. 1h: 222--23h. Bauer, H. L. 1943. The statistical analysis of Chaparral and other plant communities by means of transect samples. Ecology 2h: A5-60. Beall, G., and R. R. Rescia. 1953. A generalization of Neyman's con~ tagious distributions. Biometrics 9: 35h-—386. Benninghoff, W. S. 1952. Interaction of vegetation and soil frost phe- nomena. Arctic 5: 34--L4. Black, R. F. 1951. Eolian deposits of Alaska. Arctic A: 89-112. Blackman, G. E. 19L2. Statistical and ecological studies on the distri— bution of species in plant communities. Ann. Bot. 6: 351-—366. Bbcher, Tyge N. 1951. Distributions of plants in the circumpolar area in relation to ecological and historical factors. Jour. Ecol. 39: 376-395- . 1952. Lichen-heaths and plant successions at ¢Sterby on the Isle of Laesgi in the Kattegat. K. Danske Videnskab. Selskab. Biol. Skrift. 7: l--2A. Bouyoucos, G. J. 1936. Directions for making mechanical analysis of soils by the hydrometer method. Soil Sci. 42: 225-229. ‘Jv O) Clapham, A. 1. 1936. Over-diapersion in grassland communities and the use of statistical methods in plant ecolOgy. Jour. Ecol. 2L: ’3'2'.‘ ’)£’1 4/~--lv).~ 0 Cole, L. C. 19Lb. a theory for analyzing contagiously distributed pOpu- lations. bcolOgy 27: 329--3h1. Curtis, J. T. 1950. The interractions of certain analytic and synthetic phytosociological characters. Ecology 31: h3h--A55. Evans, F. C. 1952. The influence of size of quadrat on the distributional patterns of plant pOpulations. Contrib. Lab. Vert. Biol., Univ. of Mich. 5A: 1--1L. Fracker, S. 8., and H. A. Brischle. 19hh. Measuring the local distribution of Ribes. Ecology 25: 283-303. Gelting, Paul. 193A. Studies on the vascular plants of East Greenland between Franz Joseph Fjord and Dove Bay. Meddel. om Grénland 101: Gimingham, C. H. 1951. The use of life form and growth form in the analysis of community structure, as illustrated by a comparison of two dune coxmmnities. Jour. Ecol. 39: 396-—406. Goodall, D. N. 1952. Quantitative aSpects of plant distribution. Biol. ReV- 27: l9h—-2A5- Greig-Smith, F. 1952. The use of random and contiguous quadrats in the study of the structure of plant communities. Ann. Bot. 16: 293-- 316. Hale, h. E. Jr. 1952. Vertical distribution of cryptogams in a virgin forest in Wisconsin. Ecology 33: 398-—h06. Hanson, H. C. 1951. Characteristics of some grassland, marsh, and other communities in 4. Alaska. Ecol. hono. 21: 317--378. . 1953. Vegetation types in N.N. Alaska and comparisons with communities in other Arctic regions. Ecology 34: 111-140. HOpkins, D. M., and R. S. Sigafoos. 1950. Frost action and vegetation patterns on Seward Peninsula, Alaska. Geol. Surv. Bull. 97h-C. Hulten, Eric. l9hl-l9h9. Flora of Alaska and Yukon. Parts I-IX. Lunds Universitats Arsskrift, N.F. Vols. 37-h5. 59 Johnson, D. N. 1919. Shore processes and shoreline development. Ailey and Sons, hew York. Lindsey, A. A. 1952. Vegetation of the ancient beaches above Great Bear and Great Slave Lakes. Ecology 33: 53A--5A9. LcGinnies, N. G. 193A. The relation between frequency index and abundance as applied to plant pOpulations in a semi-arid region. Ecology 15: 263-—282. Molina, E. C. 19h2. Poisson‘s EXponential Binomial Limit. D. Van Nostrand 00., New York. pps. l-—A7. Miller, S. w. 19h7. Permafrost. Ann Arbor Press, J.w. Edwards, Ann Arbor, Mich. Neyman, J. 1939. On a new class of "contagious" distributions, applic- able in entomology and bacteriology. Ann. Math. Statis. 10: 35- 57. Preston, F. W. l9h8. The commonness, and rarity, of Species. Ecology Rex, R. 1953. Uplifted beach ridges and first generation lakes in the .Barrow, Alaska area. Unpublished manuscript. Russell, R. Scott, and P. 3. Wellington. 19A0. Physiological and ecologi- cal studies on an Arctic vegetation. I. - The vegetation of Jan Mayen ISlando Jouro E0010 28: 1.53"].790 Sigafoos, R. S. 1951. Soil instability in tundra vegetation. Ohio Jour. Sci. 51: 281-298. . 1952. .Frost action as a primary physical factor in tundra plant communities. Ecology 33: 480-487. Taber, S. 19A3. Perennially frozen ground in Alaska, its origin and history. Bull. Geol. Soc. Amer. 5A: 1433--1548. Thomas, M. 19A9. A generalization of Poisson's binomial limit for use in ecology. Biometrika 36: 18-25. Thomson, G. w. 1952. Measures of plant aggregation based on contagious distribution. Contr. Lab. Vert. Biol., Univ. of Mich. 53: l-16. Thompson, D. Q. 1953. Unpublished data. 60 Whitford, P. B. 19A9. Distribution of woodland plants in relation to succession and clonal growth. Ecology 30: l99--208. Wiggins, Ira L. 1951. The distribution of vascular plants on polygonal ground near Point Barrow, Alaska. Contr. Dudley Herb. A: Al-56. APPENDIX Histograms Frequency Distribution Figures Tables 61 Amaouosvpoomzdue _ .H ousmfim xfiocoam< OOH no 00 mm Om mm 05 no 00 mm Om m: 0: mm on mm ON ma 0H m , _ _ =5: 2.: a om. . . m. 0.: 3. . .oA «.4 Col. . .m.a o.m Om __ V. ¢w39470M9 .O.N m.m x I\ / OOH; I lyx/ mom 0.0 . / N l .4 O. Me O H / \\\ ///= m 0 x 04H. m.m 0.5 a a so? has mm m c c o Oma .o a a CON. .u a cam; . 40H I cflflaoca no .> mm n aonosoemm :oflpmcowpmxo Hsoflpuo> o 21? n M a CNN . .m 0amL HHH noomqmue .um.: weapons «om mo EduMOpmfi: hufimcoa 63 Dogma-opt». OOH mm 3 am l RII:II x 0/0 ._ . % Oa ON on es. Om. Om; Om. OOH. OHH. ONH. Ampopoevpoomcmua am Oh no Ob mm L am am °|||C Oma. .eomaeu A.em.ma eaaomaeea HHH seeeeeee mwpnoummuou< mo EmLMOpmwm hvfimcoa .HH ousmwm xfiocmaa< Om a4 as am am MN Om ma OH m E==_________ . . .m. . O.H m.H ¢OflQLK OoN \\/ / //4III //// \\h/llll 00m /x\ /: .m.m Tag a o . d 9 «OH - eaeaoea co , a cowpdpomwmxo Hmoflupm> > . 0 Ram u hocosdoum H o~.eH -.m a m 614 I. m DOCUI'HQ OOH mm Om Amuoposv poomceua .HHH ousmHm fiocoag nm 0m on ON. 00 0.0 Mm On 3 0.4 mm O.m mm Om «H OH m. 0/ 3 .\\\\.\ .m. Om. . .\ ,O.H 81/! .nA / OXO . / . 9% 34. II//O\ \l/ CON . , NIIIX\\ /// Om. .. I [mix . n.~ .\ a // 00. / \x/III Com /_ /.\\ . Ob. m.m Om. . TB Om. . a o OOH. H e. OHH. . M o ONH. xOH I oHHMOAQ no . H COHpmnmmmmxo H0305; m OMH. . RON .. honoswoum oi. 9m - m OmH. N.0 0>H HHH poomcmus canon Huousosonom Suonaomml mo :BuwoumH: 033.com 0.4 «.4 O.m 0.0 «.0 O; :1: n 65 QQme-‘I‘Ph 0H. ON mml on. on. as. 3. On. mm. O0. O L K, I o: .03 am: .3] u? L: .1. AI 1 sea I eaaaeea co COHpmpmmmmxo HmoHpuo> MMm I mocozoopm 3.0 I M HHH poomcmhe .Hnmx mHHHumzom Negev Mo swamOpmHm thmcoO . O.H .m.H . O.N m.m 5.5 HHO>CG+>°HOC O.m m.m 0.0 0.0 o.> ma 66 QQCUJ-Hpbz OH. 0H1 :3 (Y es. as. mm. O0. sou I eufieoea we COHQmummmmxm HmoHpum> mHm I Aocoscomm 3.x. I N HHH seemeeee .Ewm A.EHHonxv mHHOHHQMH .hm> .thom A pmomqv mHHm>Hc «Hausa mo smAMOpmHm.huHmco0 Cm o.m .m.m A WH°>¢GPHOC é V m.e o.“ m.m 0.0 “.0 0.5 Saigon-Hp» 0H ma. om mm 0m mm 04 «a. 0m. mm. oel as am am as “WW «0 aoH I easeoea we COHpmuowmme Hmoprm> R00 I mocvswomh asIM so me on ma sea 0 o $0Mgd7mg x / IK\\ / / me . HHH seemeeee .FN; Ger—rt my.” ’vthcc< ON mH OH m an om mu _ /. he 0.H A .och mmsmcoo mHsNSH mo SwamoumH: thmcoQ Ia.H o > m 9-H o 0' é V 0.0 m.0 0.5 :1: Q. ngmflah om. mm. 00. m0. 0m. mm. mml\ \‘I on I eaaaoee co COHpmuommmxo HmOHpuo> &OH I hocmsvocm mmH I M HHH peeeeeee .hcczoco: ESHHomHemmem automaOHum mo EmpmopmH: mpHmcoO .O.N .m.m \X///¢ 00m MHO>0QHOG é v m.m 0.0 m.0 0;. ma QGGm-Hpbe, mm. 3. m0. Om. mm. 00. IC‘ 08H I 03.093 we GOHpmpommmxo HmoHauo> mum I hoconwoum m0.m I M HHH poomcmue .A macuoo ammuMwam no swamOpme thmcoO _ A HH0>G+J~40S3 S. v 0:0 “:0 o.“ 2 0.0 as O.> 00:03-th coHpmuommmXo HmoHpuo> . . $00 I hocosvoum OOH n )0 mm 0m mm. 0A.. mo )3 mm. 0m m.0 0.0 mm )0 nu. 0v mH 0H m 52 . ___ ml . .m. 0: .3 mH.x//lx// o .m.H /X/ 0 / DNA \/0 o .OoN /R, \ / O O [IK\ // ,. a x/ . “m..;flaeea z x/ .m m // 0m. / \\u// / .0.m / \\ d— R. .. .3 00. . Aév m0. . c o Om. . H p I a 0 mm. MOH oHHmoc 00 > o H m 4m.0 I M HHH poomcdhe .mUpszHm mpomH mHamHHopm mo swamOQmHn thmcoa 0:0 “:0 0.m m.m 0.0 m.0 0.0 :1: D. 71 QGCM-Hpb; OH mH. ON. AN; 00H 0m 00 mm 00 mm 00 m0 00 mm Om m0 0.0 mm 0.0 mw Ow n.H OH m 9\ .IIIIIIII 00. \Olf. _. . . /l O, .Q\\\ // O O /IIII meI.// // \.I /I\\ mm. 04. m4. 0m. mm. 001 m0. sod I eaaeoae co GOHHMhomMon HmoHuuo> mum I hocosvoum Sam I M HHH eeeueeee .noHum A.an:0HmHam mopHmcpoa no swamOpmHm huHmcoO .O.H .m.H 0.N A43 IflIH 0 > m.p-d o a m.m 0.0 m.0 0.0 72 0000)?ka 0H mH 0N. mml mm. 3. m4. 0m. A A ’ P p xoa I eaaeoee 00 GOHpmuowmmxo Hmotho> R04 I mocmswoum OH.m I m HHH peeeeeee .:= =. 2.. .nompuom H.0onoHv msousmusaoppm 0Hoocom no smumopmHm thmcoO . OoH A [fir-10>fl-o-3HOC3 8' V 0:0 3 0a 3 0.0 73 00:03-th 0H. 0m. em. a: 0.: 3. 03 rI\ KN > n~.\ aoa I eaaueee 00 :OHpmuommmxo HmoHauo> now I mucosvoum mHH I M HHH accesses .pHafi 71000.3 053003: .3020 2H 00.3055 000.5080 mo 52000me 00350 .m. .0.H .m.H .0.N 05V LflHO><flQ¢HOCI . 0.4 3 0a m.m m.0 0; ma 7A (Duane-40>, III. :.. .. . I I I I I _ _ \0/0\ m. o\. . m. 00. .\\\\\\\\ . 0.H m.m.aliix / O/ . moH Ix/ 0/ o\flflfim>0dm 0 ON. \/‘ 0\\\\\ \n/ . . O N I] \\ / OIIIIIII I». /n// mma\\\\\\\\ :mIIx/ .m.~ m // m mm. a . m.m 04. .A m4 0m. Noa I caeaoea co ccHumuvmmmxo Hmopro> aHm I mosescoum 00. I.m MH°>03+>HOG é v HHH accesses .zmusm mgmckumEu mHHHpcopmm mo EwumopmH: muHmcoa 0.0 m.4 0.m 0.0 m.0 ma 75 QOCm-I—Ipia D r 0H. WHJK!’ 8. fl - . III| .- 1/ ./ .53 \ /X' .\\ \X/ I I IKI‘. / 40H I 3820 .3 COHpmuowmmxo H00pr0> mmm I mocoswopm 00. Im HHH seemeeee .um.m mmOHoHHom mmmumesm mo smumopmHm apHmcoO II p o\\\/ . A mHo>pros é v 0.. m. 76 m mm. 3. 3. 00:03-va R. a. xoa I eaeaoee no QOHumsommmxo H00th0> fimH I mocosvopm 40. I.M .uHsmA.0coqv mscHE .00090 .4 momuHImH0H> adHcHoomm r V I? Emumoamfim haHmcoO .m. .0.H .0.N m& O mHo>ddeG 3 an I hocosvoum dem .eeeesm 2. .0000? meeeflaomeau emetuaxem 0.4 m.4 0.m 0.0 m0 9A :3 o. 77 Oman-Hub. aoa I eaaaeea 0e 00Hpmumwmmxo HmoHpuo> fiom I mucosdoum ON.MH I.m HHH poomzmue .000 A.3nv mHudeoHsuo> mHHoHEmze mo EmamoamH: mpHmcoO me O .H m .H 0.m m.m A mHo>auHoc S. V 0.4 m.4 0.m m.m 0.0 m.0 0.0 :1: GI Chm ccnmIp A OH + I _ y 1 . E, E E; _...__ md .mod Om. .0.N mm. m.m om \I/I/LOOM an. . «a 04. .H.av : m4. . o H R. . . s aoa I eaaeote no 2 mm. coHpmuommmxo Honnuo> A > - 0 R00 I mucosvoam H 3.2 I m e HHH seemeeee .Hmz A.xoozv wOHuoua aCHthowO mo smumopmflm haHmcoQ ‘2 .0 a... 0.m 0.0 m.0 0.0 In 79 QQCflw-ih “whoposv poomcmpe OOH mm 00 mm om m0 00 no 00 OH mH mm. Om. mm- 04. m4. 0m, j l: \fl//I.I\\\ aoa I eaaaeea an mm. COHpammmbmxo HmoHpuo) m0m I accosvomm mH.OH I M D (m.. 0: mm \u/ // IQ x! HHH seeeeeee .:o¢ A.qv mowvcmHmH mHHmwuoO mo SeymoumHm ApHmcoQ 0m . g, a m .HHH>H oustm Xchmaq< N 0m mH 0H m ______ m.H 0.m m.m .O.m A 0 LflH0>d+JfiOC3 5 0.0 an l l Inn-443.5: (DZ C2 mH 0N mm. Om mm 00 3. 0m. OOH mm 00 L Amuopoev poomcmue 00 mm 30 mo 00 mm Om m0 0: mm 0 :OHpm>mHu X :a /./ I . <00 I maaaeea 00 COHpmuommmxo HmoHpmm> flmm I hocosvopm 8.0 I m HHH seoeeeee Om mm .HHN mohsmHm Nchoda< ON mH OH a z i. .ch> A.mvsmv nanonon msuozaouomnqm no amumoanm haanon .A mHo>mpfiOC é V m o 0.H 1 8 (30:01-th 00H mm 00 mm m OH mH. mm. 0m. mm. 0.0. m0. Om. Ampoposv poomcmum. HHH aoomcmus .:o< acomAe>H0 mHumHsochoo Ho smumopmH: haHmcoO .HM mousmHm 03.0009? 00 m0 00 m0 00 mm 0m m4 04 mm 0.m mw 0N mH OH m . I: __ o\\\ no . o.H ul/IX 0 “0H ///n g/ / o 0 $7 .. \\\\HVXA/MHH///I.\\\\\\\\ cowm\./ . 0.m \ l/IQ/x/ .m.N . / / \\x/l/ .OOm /.\ .. m.m .05 a o H x2 I .3080 .8 w COHumuommmxo HmoHpuo> > 0 R40 I hocosvoum H N0.ml.m m 0:. m.. 82 f3 0 : m-H.p >. OH. mH ON. mm. 0m. mm. Amumpmsv commemEB .HHH oMSMHm KHucmaad OOH mm 00 mm Om m0 00 mo 00 mm On 0.0 04 mm 0;” mm 0N nH afiH m _ H1 . _ __ .IIIIIIII. m. 0.0 O.H m.4 t/l/X// 0 - “0‘” 00m I/If// . . \A//.\\ x/ OoN mom sIII/aII\II\ / o e /K// x3093. $0 / / m.m 0.0 // . . //// \\\\\\\./II//IIIJ.0 m m 0 .. m.m 0.0 .A.EV :0 c o H a . a0H I oHaeoca we a COHpmuommmxo HaoHuuo> > o mm0 I hosezvonm H mo.m I.m. m HHH seemeeee .004 A.qv eaae>ae.mmweupoo we eeemoseam Asameea 83 QOQm.r{+)>z 00H OH. mH. ON. mm. on. mm. 00. m0 Om. Amuouosv poomcmue 38 $8383 mm 00 m0 O0 04 mm 0n NOH I OHHHOhQ Ho :OHaanemmaxo HaoHuuo> N00I hocoscohm «m.m I m HHH cooncaua nmm chovaHO no saumoumH: hancoQ .HHdH 00930 030503 mm 0N mH 0I\|II|II\\IIO — . . 0.m m.m . 0.m m.m . A £fl,H o > d-p-H o G é v m.0 0.m m.m 0.0 m.0 0.0 :1: O. 8h QOCw-Hfih 00H 0H. mH. mm. 0m. m.m. 00. m4. m0 Amumpmav poomcmae 00 a 0... as 0... 3 8 R on 3 a. m.m 0m mm _ _ xoa I oaamoea 00 COHQmuommmxo HeoHpuo> m0: I mocozcoum 004 I m HHH eeeeeeae .0HHH2 A.Hv mchmo whomHuHom Ho EmumopmH: thmcoO .HHHHH mpsmHm Kchmugd om m0 0H m +.- O\. .m. .0.H .m.H mHo>deoc é \J 0.4 0.m m.m 0.0 m.0 0.0 :1: D. 85 30:200-th Annaposv pommcmps .0HxH wustm xHucoaa< 02 a. 0.. a... 8 3 00 we 00 a 0a a... 3 3 0m mm 0m 3 0a a _ a _ =— m. 0 .m. OH 0 .OoH m.m.xlllx \ol \ .m.H /./ . . 0033.3. 00. \fl.\ \I\\./ / 0.0 all“. le\ /KIII mm.\\\\\\\\ mm2// .m.~ I / K. om // \\\ ///..00M 3. . m.m 04 A ”ca I oaamoee we :OHpmquMQXo H00pr0> &0m I mucoscoum mmH IM HHH 0.0.0.00 .0eoeem A.aea0 eeHeaH MHo>deoc é \J «thnoH mo adumoamHz szmcoO 0.0 m.0 0.0 :1: D. 6 8 chm-th OOH mm 00 “4 OH. ma. ON‘ mmI om. mm. of 3. amhwpmav pommcdpe .>MA mhdmfim KHUCmQQd £8 MBRSBMmRSBRREONMHSm .co< A.AV mouoadgqao maawfiuam Mo adhmOpmfl: haamcwa _I a I\. .% .oA \GOAI l/x \\oI/O .moH / O/\\ /h 0 \\u/ .OON . IIII\ // xiii/u c an / m m // m \u// 6. /¥\\\ /: m.m .73 T. o H Moa I oafiuopg mo w :oaumpmwmnxo Hmowago> > mm I hoaosdmum M +34 I M m HHH aoomsane o; 0.m m.m 0.0 m.0 0.5. :1: D. (Du-443% CDC Q OOH mm Om mm Om mu 0% mo A mH.T ON. mW\\\\ 04. mq. Om4 o//o\\ xoa I maflmogg mo coflpmnwmmmxo HwOfippm> wma I mozmsvmum mu. I_M Amummpvav pommcmue Om m4 OJ \/ HHH poomcwge .902 A.Av mwamfiocz mflcoumao no swumopmw: hpfimcma / I, mm .H>%£ mpzmflm xaucmma< ON ma 0!. .qqmmmmJMm OH m .m. .o.H .m.a . 0.m A mHo>prOG é V 0.4 0.: 0.m m.m 0.0 0.5 mm 88 :3 m c m-H.p >3 OOH OH mH.II.II. ON mm. on. mm. 04‘ mm ’ '1 0» mm Aum mm Oh we 00 Amgmpmsv pommcmpe mm Om «q. O4 mm Om mm .HH>HH mhdem chchQ< ON mH OH m .I\\\\\\I Q\\\\\ \ \Gfima o xoa I oaflmopa we COprnvmmmxo Honppo> Rmm I mocosvoum oo. I_m HHH pooncxpa .UHHH} A.Hv «nonpnaa anomHuHom mo squOpmHm hpfimsva .m- .O.H m.H .O.N .m.m .A mH'o>dp«-Ioc: E. v 0.4 3 0.m m.n 0.0 (Ow-149‘?) JG) ”mumpmsv pommcmpe .HHH>AM mpdem XHocoaa< 03 .3 3 mm 8 SI 0.» 3 oo 3 an 3 on mm on 3 om 3 B m \\\\\ MO 0\0 “0 OH. .\ o.H mIH .IIII‘I/l of “OH I/ \a o 0 9.670 ON. \A//x/o\%\\.m.\\a/ CON 0 0 [III\ // MN .-\ IIIQu/ . m.m / om // \\\\\r/IIII 0.m //I\I\I I mm. .3 OJ. .A 3 . _ xoa I oflfimoua mo COHpmnowwwxo Hdowpno> ROH I hocoswohm mm. IM MHo>deoc é \J HHH puomcwus hopm H.2mmzv mouHoqum>o QOHSaooououm mo Ewpmounflm manCoO o; 3 m.m 0.0 m.0 0;. ma O 01 ‘2 O c m-H.p b. SAmpomev pommcmpe .MHHH mnSMHm vachQ< OOH mm Om m5 Oh no mm Om m4 0d m.m Om mN OW m.H OH m \0 ML O\\O .m. OH 0\ 00H mH.I:II m.H . lul/ o/O lt/ O \\o\§.¥d?.a .OoN ON \fflo’l.\\\x/// I O O IIIIR imam mm \ IQ // u/ Cam //!\\I mmI m.m 04. N «4. a “ca I oafiuopa Ho :oHuwuommmxo Hwowauo> mmH I hocosvopm Om. I.m HHH aoomcana 'mHo>a»Hon é V omchq A.um.nav mooa>HHondn moqumwdo Ho EnumOpmH: thmcoO 0.4 m.q .Oom m.m 0.0 m.0 o.u a: O. 1 Q/ QOCID°H+3>> Ampmpmav pommcwue .xxd whdem KHUcmaad 03. no om mm om SI 2 3 8 a 0.m 3 3 mm on mm 8 2 B m 0 OH. mH. ON. mm. Om. «mg 04. 3 TI Oml D afiII .—qIIIIIIII—,uql . AHII uqxxxwx o\ .m. o\ 00H IR/ 0 .MOH l/x \ f0 //.////\\\\é3%%a .Qm /IIOI \\\l// A xoa I mflflmoga go a cngmhomwwxv HMOHauo> am I hoGoddogm NH.O I M [fir-IObdi-D'HOS E. v HHH poumcmua .xoo: HHnomuumonu wwhmhpoo no Emumopmflr mpflmzoa o.¢ «.4 0.0 m.@ 0.5 3: D. 2 01 M OOH IWNIJ/ m.m. 0.m Viiilu/ .\ 'P I .Hxxx whammm xaccoag< @3qu OOH 969m 33:982.“ cmmfisom x. mm 0.m ON. mo Ow mm \Om m4 0: m.m. 0.m mm Ow MH OH m .../ / lat I /...\../.I.> ......)k/Nfi... 1.1:... ./\......./§W\.W.X.\\<1....v.\..A.>..../..\..\.:./...-.1 \(kaNi/I/ m .0. . m H .om I cogsnwunofiu aonuHom I cofipangupudu m hogan—won.“ 3:030 3.me I m .um.m onuouw mom #:OHpanuanO honoSUoum HHH uoomcmpe 93 .HHHHH whsmHm XHccmaad .UmppHso OMH pa hosmddmpw mco * On mo 00 mm Om m4 04 mm on «N ON «H OH m x I II . - . ...- ...-.- .. . . . . I . . I A... . .< z.../\ /I\ / \zlwI./.\.<./ ...”... .I .\{./®K/KxpounO aéHIm .oemfiuu A..m.mv aaaomfimmw naauommwgmmw *GOHpanppmHO h0263doum HHH QOQnGth. 70~ 60 50‘ 40‘ 30* 20‘ 10‘ 9h Transect 111 Frequency uistribution Eriophorum scheuchzeri HOppe 2-8.9 Observed frequency distribution - x x Poisson distribution - o—————o -/:-"“\- .éh’x d/V‘IAK I l"\‘ I’V‘fin ' . __ X gr» . 5 10 15 20 25 30 35 1.0 45 Appendix Figure XXXIII. 50* [40‘ 10‘ Transect 111 95 Frequency Distribution Carex aquatilis Wahl. i’ — 8.1+5 Observed frequency distribution - x Poisson distribution - . 23 Appendix Figure XXXIV. OX 50‘ 30: 26 10‘ Transect III 96 Frequency Distribution Luzula nivslig (Laest.) Beurl. var. latifolia (Kjellm.) Sam. 3(- - 7.15 Observed frequency distribution - x——- Poisson distribution - o ./ \ A._‘ -.- --. “K /‘ 4 f / \xk / ‘/ V 0:\/\J44\g \_ LOO ,\,/-.\,- :- 26-441 .- 3 s 34‘1/59/5': . . ' HUL'O'H'I'S'H'Z'O' 25 3 Appendix.Figure XXXV. 55 50 45 #0 35‘ 30‘ 25 20‘ 15‘ 10‘ /. \'\ 1 10 Appendix.Figure XXXVI. L/'. p . \ J/Jx;/‘\u/R\V/fl\-“/fi\:\:‘ng\c/’\./R\u/”'”"‘. 4/A\L,x\k 5 ' 113' i0 25 3 Transect 111 97 Frequency Distribution Luzula confusa Lindb. a? - 6.91 Observed frequency distribution - I Poisson distribution - o o 35 to AS n—x t " "‘ ‘x/ \xl’\ui/\L1If¥/ \ I. 1 50 X Transect III 98 Frequency Distribution F Eriophorum angustifolium Honekeny. i - 1.95 As Observed frequency distribution - x x Poisson distribution — ._____. #0. 35 30. 25« 20‘ 15‘ I" 10- 5. O «V» A -".‘~. .-”.'\- .«r» . 5 i0 i5 2b 25 30 35 no as 50 Appendix Figure XXXVII- 99 .HHH>NHH oudwwm Navcodg< 3 23 mm om mm om 3. .3 m P P D r b P 1...“... - 11-..}, . , . , 3 /0/0 x/nIn \( \ / J. I ./ I. . . ./ \ \/. . /.\. . O o I COHpanhumHv commHom X x I QOHudnHuumHu hocoswmhm vo>pomnO 34. I M .a assume owmnMmem QOHuannamHO accosvoum HHH accesses Transect III 100 Frequency Distribution Stellaria laeta Richards. 2? - 6.51. F Observed frequency distribution -x. x 40‘ . Poisson distribution -»0 o 35 ll 30 25‘ 20 15 10‘ 5‘ ‘\\7Z>/\\/\ /f I ._/,,/1\\[\/ + .\ _‘ K/ H 5...\./.'..'\../..\./..\:..x fl ' " 5 """"" 20 25 30 Appendix Figure XXXIX. h5‘ #0- 35‘ 3O 25 20 15 ' 10 ‘ / 5 Transect III 101 Frequency Distribution Petasites frigigg§_(L.) Fries. i - 5.75 Observed frequency distribution — x——__¢ Poisson distribution - . 1o 15 . 20 25 30 Appendix Figure XL- ...l—R/l /\ ”-K \ {/H/X/H ' \- \f""- “7"“ ’.'\-"\-= rué—u—v—h. \ " x 35 55‘ 50 1&5 hO‘ 35' 30- 25* 20‘ 15‘ 10‘ ,O\ /. Transect III 102 Frequency Distribution Senecio atropurpureus (Ledeb.) Fedtsch. i - 5.]. Observed frequency distribution - u—I Poisson distribution - o \\\..l[\/_‘ \f-“-'\ -\/"""\f K/’\ . X 15 \20 25 30 Appendix Figure XLI. Transect III 103 Frequency Distribution Saxifragg punctata L. ssp. nelsoniana (D.Dcn.) Hult. i-IJB F Observed frequency 80 75 70‘ 65‘ 55 50‘ AS‘ 40‘ 35 30* 25 20 15‘ 10‘ distribution — x——x Poisson distribution — 0 J O J / \xfl \\/ \~?”*v . . . X 5 10 15 20 Appendix.Figure XLII. lC‘ 10h Transect III Frequency Distribution Vaccinium vitis-idsea L. subSp. minus (Lodo.) fiult. i-.»3L. Observed frequency distribution -'x X 0 Poisson distribution -0 Appendix.Figure XLIII. BOT 75' 70. 65 60 55' 35‘ 30‘ 25. 20‘ 15 10‘ Transect III Frequency Distribution Potentilla emarginata Pursh. 76-.80 Observed frequency distribution - Poisson distribution - I \\ /x\{l\_ ./‘_'\n/'\x . ‘ '5 10 15 20 Appendix Figure XLIV. I °-—-—° 2'5 85 80. 75‘ 7O 55‘ 5O 1+5. #01 35‘ 25. 20 ‘ 15 10 ." 5 Transect III 106 Frequency Distribution Saxifraga foliolosa R.Br. i-.67 Observed frequency distribution — I————J Poisson distribution - ° - °\g/f\,/x\/\, , . . x 10 15 20 25 Appendix Figure XLV. 25¢ 20‘ 15' 10 Transect III 107 Frequency Distribution Thamnolia vermicularis (Sw.) Ach. i - 13.70. Observed frequency distribution - x X Poisson distribution — o ./\(:3/;¥:(\: A/:,,:I\>§W‘"\/\ , /\.._.'*\.._..../\../\,..frm - . X 20 25 30 35 1+0 155 50 55 60 05 Appendix Figure XLVI. 30‘ 25‘ 20‘ 151 10 Transect 111 108 Frequency Distribution Dactylina arctica (HOOko) Nyl. SE - 12.76 Observed frequency distribution - x x Poisson distribution - o——___. /" \ " " 2L” \\F\/\H)<\XZ \ /" /"\ ‘- I'c-U—nx 144‘ W 5 ,, 1 x 1o 15 2'0 25 3o 35 no Appendix Figure XLVII. 25‘ 20‘ 15¢ 10‘ / -\/>