Plant Ecology 148: 81103, 2000. 2000 Kluwer Academic Publishers. Printed in the Netherlands.

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Classication and ordination of plant communities along an altitudinal gradient on the Presidential Range, New Hampshire, USA
Santiago Sardinero
Instituto Nacional de Investigaciones Agrarias (INIA), Centro de Investigaci n Forestal (CIFOR), carretera de La o Corua km 7, E-28040 Madrid, Spain. (e-mail: sardiner@inia.es)
Received 11 December 1998; accepted in revised form 9 December 1999

Key words: Alpine tundra, Bioclimatology, Krummholz vegetation, New England, Northern Hardwoods, Spruce-Fir Forests, White Mountains

Abstract An analysis of vegetation along an altitudinal gradient on the Presidential Range, New Hampshire, USA, using the BraunBlanquet approach followed by multivariate data analysis is presented. Twelve main plant communities have been distinguished. Floristic information is presented in twelve tables and one appendix. The relationships of the communities to complex environmental gradients are analyzed using Correspondence Analysis. Floristic composition and community structure are controlled primarily by the altitudinal gradient (temperature, precipitation), and by mesotopographic conditions (snow accumulation, exposure and cryoturbation, slope position, and soil moisture). Abbreviations: CLC complete linkage clustering, MNVAR minimization of variance in new clusters, MISSQ minimization of the increase of error sum of squares, SAHN sequential, agglomerative, hierarchical and nonoverlapping clustering techniques, CA correspondence analysis, ss saplings and seedlings, kr krummholz. Nomenclature: Gleason & Cronquist (1991) for vascular plants, except for Betula cordifolia Regel, Coptis groenlandica (Oeder) Fernald, Empetrum nigrum ssp. hermaphroditum (Lange ex Hagerup) Bcher, Huperzia selago ssp. arctica (Grossh. ex Tolm.) Sipliv., Minuartia groenlandica (Retz) Ostenfeld, Scirpus caespitosus var. callosus Bigelow ex Torr., Vaccinium vitis-idaea ssp. minus (G. Llodd.) Hult n. Nomenclature for lichens follows Esslinger & Egan e (1995). Introduction The Presidential Range (New Hampshire, USA; Figure 1) is a classical locality, frequently visited by botanists and plant ecologists since the earlier nineteenth-century, including Antevs (1932), Baldwin, Bigelow, Bliss (1963), Booth, Cutler, Griggs, Monahan, Oakes, Peck, Robbins, Thoreau, and Tuckerman, among many others. Fenneman (1938) included the Presidential Range in the Appalachian Highlands, New England Province, White Mountains Section. Braun (1950) included it in the HemlockWhite Pine-Northern Hardwoods Region, New England Section. Rivas-Martnez et al. (1999) included it in the North American Atlantic Region, Appalachian Province, Appalachian Sector, North Appalachian Subsector. Kchler (1964) identied the northern hardwoods (Acer-Betula-Fagus-Tsuga), the northern hardwoods-spruce forest (Acer-Betula-Fagus-PiceaTsuga), and the northeastern spruce-r forest (PiceaAbies) as the potential natural vegetation within the territory (see also modied version from U.S. Dep. Agric. 1978, and Barnes 1991). Greller (1988) identied two major deciduous forests within this area: hemlock-white pine-northern hardwoods, and sprucenorthern hardwoods. Bliss (1963) presented quantita-

82 tive data on the principal alpine plant communities and their ecological relationships with soils and meteorological factors. In this paper we present a classication and ordination analysis of the vegetation along an altitudinal gradient on the Presidential Range according to the Braun-Blanquet approach (Braun-Blanquet 1964; Mueller-Dombois & Ellenberg 1974; Westhoff & van der Maarel 1978; Ghu & Rivas-Martnez 1981; Rivas-Martnez 1987), followed by multivariate data analysis. Our purpose is to develop a comprehensive, oristically based, vegetation classication in the study area. The Presidential Range constitutes the territory with largest alpine belt in the northern Appalachians. For this reason, it is the best place to develop a basic vegetation model which could be extended along the mountains of northeastern US with alpine summits: Presidential Range, Mts. Lafayette and Lincoln on Franconia Ridge (NH), Mt. Manseld and Camels Hump in the Green Mountains (VT), Katahdin (ME), and Adirondacks (NY; Figure 1). This classication might be used as a basis for the development of syntaxonomic vegetation units in northern Appalachians, in vegetation mapping, and in climatic change studies. Furthermore, it would be possible to compare these plant communities to other temperate, arctic, and alpine communities along the Northern Hemisphere: Alaska (Walker et al. 1994), Eastern North America (Bergeron et al. 1984; Grandtner 198990; Miyawaki et al. 1994), Europe (Mucina 1997), Greenland (Danils 1982, 1994), North Japan (Miyawaki 1988; Nakamura & Grandtner 1994; Nakamura et al. 1994), etc., and to better understanding their global composition and distribution. Study area The Presidential Range comprises the highest peaks in the northern Appalachians, which along with the Franconia Range to the west, contain the highest peaks in the White Mountains. These mountains are composed of Lower Devonian mica schists and gneises of the Littleton Formation, intruded by small dikes of quartzite. Glaciers eroded these mountains, leaving cirques, Ushaped valleys, glacial grooves in the rocks, erratic boulders, etc. (Billings et al. 1946). The climate Davis (1976; 1981) has documented multiple glacialinterglacial cycles within the 2 million-years of Pleistocene time, possibly as many as sixteen, with an average of 125 000 years for the complete cycle. Relatively short interglacial periods of 10 000 to 20 000 years were followed by long glacial periods up to 80 000 to 100 000 years. The Pleistocene glaciers reached their last maximum extension about 18 00020 000 years ago during the late Wisconsinan glaciation, and retreat began about 16 500 years ago (Delcourt & Delcourt 1979). The Holocene, the present warm climatic interval, began 10 000 years ago and reached a temperature maximum 5 000 to 8 000 years ago (Mitchell 1977; Houghton et al. 1996). Present conditions appear to represent those near the end of a typical interglacial interval (Mitchell 1977). Meanwhile the Earth seems to be involved in a process of global warming, specially the Northern Hemisphere (Mann et al. 1998, 1999), correlated with a substantial and sustained rice in the carbon dioxide atmospheric concentration over the last two centuries (Keeling & Whorf 1998; Etheridge et al. 1998). Present climatic data were obtained for two meteorological stations within the study area (National Oceanic and Atmospheric Administration, NOAA), at the top of Mt. Washington (Figure 2b), and at Pinkham Notch (Figure 2c). The increments of the mean annual temperature, and mean annual precipitation are 0.57 C/100 m, and 63.27 mm/100 m, respectively, assuming a regular increment for them along the altitudinal gradient. Bioclimatic nomenclature follows Walter & Lieth (19601967), Walter (1985), RivasMartnez (1997) and Rivas-Martnez et al. (1999). Methods Field sampling The study was based on a data matrix composed of 183 samples and 93 taxa. The sampling was done during 1995 and 1996, using the centralized replicate sampling procedure (Mueller-Dombois & Ellenberg 1974). Plot locations were subjectively chosen between the two meteorological stations, in areas of homogeneous vegetation, according to plant-forms and physiognomic-ecological plant formations (Rbel 1930; Raunkiaer 1937; Braun-Blanquet 1964; Ellenberg & Mueller-Dombois 1967a,b; Whittaker 1975; Box 1981; see Appendix 1). The moss layer was not recorded.

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Figure 1. Location map of the Presidential Range showing also other locations having alpine summits in the northern Appalachians.

Data analysis The characterization of homogeneous plant communities is one of the principal aims in phytosociology, phytogeography, phytoclimatology and landscape ecology (see Legendre 1990). Clustering methods represent a useful approach to this problem. Syntaxonomic table sorting (Braun-Blanquet 1964; Mueller-Dombois & Ellenberg 1974; Westhoff & van der Maarel 1978) was used to detect and characterize preliminary vegetation types in the data matrix with 183 samples and 93 taxa. The abundance/dominance values of the 6-grade scale of BraunBlanquet in this compiled raw table were transformed into a 09 ordi-

nal scale according to van der Maarel (1979). Several Sequential, Agglomerative, Hierarchical and Nonoverlapping clustering techniques (SAHN techniques; Sneath & Sokal 1973; Podani 1989a) were performed in order to classify the data. Similarity ratio, chord distance, and euclidean distance were adopted to compute the distance between every pair of samples in the resemblance matrix. Distance-optimizing clustering methods like Complete Linkage clustering (CL), as well as homogeneity-optimizing clustering methods like Minimum Variance of newly formed clusters (MNVAR), and Minimum Increase of Error Sum of Squares (MISSQ), were carried out with the programs NCLAS and HMCL (Podani 1993, 1994). Then the

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85 alternative results were compared looking for similarities in the composition of clusters, so we could have a good reason to assume that there is some real tendency in the data towards grouping (Podani & Dickinson 1984; Podani 1989b, 1994). Ordination analyses were performed by Correspondence Analysis (CA), carried out with the program PRINCOMP (Podani 1993, 1994). The units dened by cluster analysis were superimposed on the ordination diagrams in order to provide a more complete analysis of the grouping, and to analyze relationships between vegetation and ora in terms of unimodal response models (ter Braak 1985), by drawing sample plots and taxa plots. A good explanation on the interpretation of CA biplots may be found in ter Braak (1995). Since very different life forms, plant formations, and sample plot sizes were involved in the data analysis, a progressive dimensionality reduction was carried out, splitting the data in data subsets with the help of classication and ordination results. Further analyses of the data subsets structure were carried out by comparison of alternative clustering algorithms and CA ordinations (Podani 1994). For a nal result, computer program ELLIPSE (Podani 1993, 1994) superimposed classication results on ordination diagrams by drawing ellipses of equal concentration around the centroids of clusters (Mardia et al. 1979; Lagonegro & Feoli 1985) at 95% probability level. The mean and standard deviation of each ordination dimension for every community were calculated. Differences between means among the twelve communities were tested by Students t-test, giving the probabilities that the null hypothesis is true, i.e., the probabilities among every pair of plant communities of being the same community. plant communities, and giving good reason to assume that there is some real tendency in the data towards grouping. Nevertheless, several differences can be observed between both clustering methods: Figure 3a separates groups 5a and 5b at the highest dendrogram level, emphasizing their oristic differences, whereas Figure 3b gathers both groups in the Picea mariana Abies balsamea krummholz vegetation, following a physiognomic criterion. This last criterion has been followed in this paper. Also, Figure 3a gathers groups 10, 11, and 12, whereas group D in Figure 3b gathers groups 812 in the same branch; groups 8, 9, and 11 are located in one side, and groups 10 and 12 in the other side. Figure 3b suggested two main groups, the rst one comprising the forests (group A in Figure 3), and the second one comprising dwarf trees (krummholz) and alpine vegetation (group B in Figure 3). Classication using euclidean distance and MISSQ (dendrogram not shown) suggested the same two main groups (A and B), and also the same twelve plant communities. Preliminary ordination The results obtained by preliminary classication were conrmed by the ordination analyses, and we distinguished the same two main groups (A and B) along the rst CA axis, and groups C and D along the second axis (Figure 3c); plant community numbers correspond to those in Appendix 2. A dimensionality reduction was carried out, splitting the original data matrix into two data matrices, one of them comprising the forests data (group A), and the other one comprising the krummholz and alpine vegetation (group B). Forests data matrix classication Two additional clustering algorithms were tested with the forest samples, euclidean distance and MISSQ (Figure 4a), and chord distance and CL (Figure 4b). The plant communities yielded by these methods were the same that those obtained in Figures 3b and 3a. Euclidean distance and MISSQ (Figure 4a) provided an easy interpretation of the forests data, so they were sorted according to this clustering method. Thus, four main forest types were distinguished (Tables 14). Fagus grandifoliaAcer saccharum community Table 1: samples 116, Group 1 in Figure 4 Upper-montane northern hardwood forest. Fagus grandifolia combines with Acer saccharum, Acer pen-

Results and discussion Preliminary classication The samples were sorted according to similarity ratio and CL (Figure 3a) which have been demonstrated to be effective algorithms in syntaxonomical studies (Mucina & van der Maarel 1989). Plant communities in this dendrogram correspond to those in Appendix 2. Classication using chord distance and MNVAR (Figure 3b) have also been demonstrated to be useful algorithms in syntaxonomical studies (Mucina & van der Maarel 1989), yielding the same composition of

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Figure 3. Preliminary analysis. (a) Clustering using similarity ratio and CL. (b) Clustering using chord distance and MNVAR. (c) CA ordination for samples. (d) CA ordination for taxa. Vegetation types: (A) Forests. (B) Krummholz and alpine vegetation. (C) Upper-subalpine krummholz, and alpine dwarf shrub ericaceous vegetation. (D) Remaining alpine vegetation. Plant community numbers correspond to those of the twelve tables and Appendix 2.

sylvanicum, Betula alleghaniensis, Acer rubrum, Betula cordifolia, and Acer spicatum in the tree layer. Acer spicatum, Acer rubrum, Acer pensylvanicum, Acer saccharum, Betula alleghaniensis, Fagus grandifolia, Viburnum alnifolium, Abies balsamea, Sorbus americana, Picea rubens, Tsuga canadensis occur in the subtree layers. Abies balsamea, Picea rubens, and Tsuga canadensis occur in the tree layer with low cover. Oxalis montana, Dryopteris campyloptera, Maianthemum canadense, Trientalis borealis, Aralia nudicaulis, Clintonia borealis, Lycopodium annotinum, Cypripedium acaule, Thelypteris phegopteris, Trillium undulatum, Medeola virginiana, Viola selkirkii, Uvularia sessilifolia, and Smilacina racemosa occur in the herb layer. Two variants can be detected: Betula alleghaniensis Acer rubrumAcer pensylvanicum dominated stands

(Group 1a in Figure 4, and in Table 1), and Acer saccharumFagus grandifolia dominated stands (Group 1b in Figure 4, and in Table 1), where the latter over time seems to replace the former, according to the models shown by Pacala et al. (1996). This forest is replaced by Abies balsameaBetula alleghaniensis community (number 3) on xeric soils (see Figure 8). This community bears close resemblance to Betulo alleghaniensisAceretum saccharii Grandtner 1989 1990, and Oxalis montanaDryopteris campyloptera Fagus grandifolia community Okuda 1994. Betula cordifoliaSorbus americana community Table 2: samples 1724, Group 2 in Figure 4 (Montane)-subalpine softwood forest where Betula cordifolia and Sorbus americana dominate the tree

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Figure 4. Analysis of the forests data matrix. (a) Clustering using euclidean distance and MISSQ. (b) Clustering using chord distance and CL. (c) CA ordination for samples. (d) CA ordination for taxa. Plant communities: (1) Fagus grandifoliaAcer saccharum (1a. var. dominated by Betula alleghaniensis. 1b. var. dominated by Fagus grandifolia and Acer saccharum. (2) Betula cordifoliaSorbus americana. (3) Abies balsameaBetula alleghaniensis. (4) Abies balsameaBetula cordifolia (4a. var. with more abundance of Dryopteris campyloptera, Cornus canadensis, Trientalis borealis, Clintonia borealis, and Lycopodium annotinum. 4b. var. with more abundance of Picea rubens).

layer, occurring with Abies balsamea and Betula alleghaniensis. Abies balsamea, Sorbus americana, Acer spicatum, Sambucus canadensis, Viburnum alnifolium, Picea rubens, and Acer pensylvanicum occur in the subtree layers. Oxalis montana, Dryopteris campyloptera, Aster acuminatus, Maianthemum canadense, Trientalis borealis, and Aralia nudicaulis occur in the herb layer. Since it is a softwood and rapid-growth community, it seems to be the secondary forest of Abies balsameaBetula cordifolia community (Table 4), and the Abies balsamea Betula alleghaniensis community (Table 3) which grows close to the montanesubalpine boundary (Figure 8). This community has been described according to the Code of Phytosociological Nomenclature (CNP,

Barkman et al. 1986) as Sorbo americanaeBetuletum cordifoliae Sardinero in Rivas-Martnez et al. (1999). Abies balsameaBetula alleghaniensis community Table 3: samples 2535, Group 3 in Figure 4 Upper-montane-(subalpine) spruce-r forest dominated by Abies balsamea and Picea rubens, combined with Betula cordifolia, Betula alleghaniensis, Acer pensylvanicum, Acer rubrum, Acer saccharum, Fagus grandifolia, Tsuga canadensis, and Sorbus americana. This forest grows on xeric soils, below Abies balsameaBetula cordifolia community. Towards deeper soils it contacts with Fagus grandifolia Acer saccharum community (Figure 8). Structurally, it is an Abies balsameaPicea rubens dominated forest, but oristically it has some transgressive species from the deciduous forests such as Betula alleghaniensis,

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Table 1. Fagus grandifoliaAcer saccharum community (1a. var. dominated by Betula alleghaniensis. 1b. var. dominated by Fagus grandifolia and Acer saccharum). Reference no.: 001. Pinkham Notch, Tuckerman Ravine Trail, 09/14/95/01. 002. Tuckerman Ravine Trail, 06/14/95/02. 003. Gulf of Slides, 09/27/96/08. 004. Tuckerman Ravine Trail, 06/14/95/03. 005. Gulf of Slides, 09/27/96/05. 006. John Sherburne Ski Trial, 09/28/96/01. 007, 008, 009, 010. Old Jackson Road, 09/28/96/08,06,05,09. 011. Tuckerman Ravine Trail, 09/14/95/03. 012. Gulf of Slides Ski Trail, 09/27/96/03. 013, 014. Old Jackson Road, 09/28/96/02,04. 015, 016. Gulf of Slides Ski Trail, 09/27/96/01,02. Plant community No. Plant community variant Relev No. e 000000 111111 aaaaaa 000000 000000 123456 000000 667666 573972 000300 242522 1+12+2 1.1111 1121+3 .+1.+. ...1.+ .2.... 1..1+. .1+... ...+.+ .1.1.1 ...1.+ .....1 243345 421111 222221 1212+. 322211 221233 222121 231212 232222 322222 12.22+ 11+111 1+2121 ++.+1. 22.212 3332.+ 12.111 .1.112 0000000000 1111111111 bbbbbbbbbb 0000000000 0001111111 7890123456 0000000000 7668766666 8750062425 3002300000 2222222232 4545433331 2132343445 2233222231 2231222232 .1211.+..1 +.1+1...1. ....1...1+ .+1.....++ .+....+++. ...1....+. .1....+.11 1.11....1. 2211133211 .l+..++1.+ ....2+11+1 ....1...12 2322333333 +211212111 22213122.. 1221222122 2211113122 3222323122 1.12++++1+ 11..1+++11 11.+1++.1+ 1.+11+.++1 1+..2.2211 +...333133 11..111.11 ++++1..+1. Table 1. Continued Sorbus americana ss Lycopodium annotinum Trillium undulatum Other taxa: Thelypteris phegopteris Picea rubens Aster acuminatus Monotropa uniora Acer spicatum Carex arctata Amelanchier bartramiana Mitchella repens Fragaria virginiana 21111+ 31.... .1+.++ ..+2+1 12.11. ....11 ...... .1.1.. ...+.+ ...+.. ....+. .....1 .++.1.1... ...3321112 .+..++.... ..1.1...+. ....++.... ...1.1.... ...+..+.++ .....2.... ......+... ..+......+ ...1.....+ ...+....1.

Altitude (x 10 m)

Slope (x 10%) Area (x 100 square m) Faithful and differential taxa: Acer saccharum Fagus grandifolia Acer saccharum ss Fagus grandifolia ss Viola selkirkii Uvularia sessilifolia Cypripedium acaule Smilacina racemosa Sambucus pubens Carex grayii Medeola virginiana Brachyelytrum erectum Constants: Betula alleghaniensis Acer rubrum Acer rubrum ss Acer pensylvanicum Viburnum alnifolium Aralia nudicaulis Acer pensylvanicum ss Clintonia borealis Maianthemum canadense Dryopteris campyloptera Abies balsamea Abies balsamea ss Betula cordifolia Picea rubens ss Acer spicatum ss Oxalis montana Trientalis borealis Betula alleghaniensis ss

Additional taxa with one or two occurrences: Tsuga canadensis ss: 1 in rel. 1, and + in rel. 2. Polygonatum biorum: + in rels. 5 and 15. Prunus serotina: 1 in rel. 6, and + in rel. 15. Thalictrum polygamum: + in rels. 6 and 15. Thelypteris noveboracensis: + in rels. 7 and 12. Cornus canadensis: + in rels. 13 and 16. Arisaema triphyllum: + in rels. 13 and 16. Tsuga canadensis: 1 in rel. 1. Sorbus americana: 1 in rel. 4. Betula cordifolia ss: 1 in rel. 6. Lycopodium obscurum: 1 in rel. 9.Cornus alternifolia: + in rel. 11.

Acer pensylvanicum, Acer rubrum, Acer saccharum, Fagus grandifolia, Tsuga canadensis, Viburnum alnifolium, Aralia nudicaulis, Trillium undulatum, and Cypripedium acaule. The tree species of this transgressive ora never dominate the tree-layer. This community bears close resemblance to Betulo alleghaniensisAbietetum balsameae Grandtner 19891990. Abies balsameaBetula cordifolia community. Table 4: samples 3656, Group 4 in Figure 4 Subalpine balsam r forest where Abies balsamea dominates the tree layer, accompanied by Betula cordifolia and Picea rubens. The presence of Picea rubens is lower than in Abies balsameaBetula alleghaniensis community. Abies balsamea, Sorbus americana, Betula cordifolia, and Picea rubens occur in the subtree layers. Oxalis montana, Dryopteris campyloptera, Maianthemum canadense, Trientalis borealis, Cornus canadensis, Coptis groenlandica, Lycopodium annotinum and Clintonia borealis occur in the herb layer. This community bears close resemblance to Abieti balsameae-Piceetum rubentis Marcotte & Grandtner 1974, and Betulo papyriferaeAbietetum balsameae Grandtner 19891990.

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Table 2. Betula cordifoliaSorbus americana community. Reference no.: 017. Old Jackson Road, 09/28/96/07. 018. Boott Spur Trail, 09/30/96/04. 019, 020, 021, 022. John Sherburne Ski Trail, 09/15/95/05,01,04,02. 023, 024. Lion Head Trail, 10/02/96/08,09. Plant community No. Plant community variant Relev No. e 00000000 22222222 -------00000000 11122222 78901234 00101011 78091922 40000305 00444400 22212122 54534344 22221222 2123222+ .2.33323 11211.22 +++.+1.. 23323332 22333332 2.+11221 111111.+ 2212+... .22221.. 12++...+ 1122.2.. 1.1113.. 1.1+.+.+ 1....+22 +1....+1 +..1.1.. .+....21 ....12.2 .....212

Table 3. Abies balsameaBetula alleghaniensis community. Reference no.: 025. Dolly Copp Road, 09/29/96/03. 026, 027. Boott Spur Trail, 09/30/96/02,03. 028. Gulf of Slides, 09/27/96/04. 029. Boott Spur Trail, 09/30/96/01. 030. Old Jackson Road, 09/28/96/03. 031. Tuckerman Ravine Trail, 06/14/95/04. 032. Pinkham Notch, Tuckerman Ravine Trail, 09/14/95/02. 033. Gulf of Slides, 09/27/96/07. 034. John Sherburne Ski Trail, 09/14/95/04. 035. Gulf of Slides, 09/27/96/06. Plant community No. Plant community variant Relev No. e 00000000000 33333333333 ----------00000000000 22222333333 56789012345 00000000000 57865666787 06169385100 40404000000 22222221222 44442444353 22233232313 23233332334 33333322312 21222121222 11+11111+2+ 12+2111111+ 11+1.22211. 11+.21.++11 .++1+++111. 11..2221121 +11+..2112+ +1+1.111+++ ++.1..+1++. +1.1+1.++.. .+.+1+2..++ +...+21++.. 122.1311221 11..+3+22+1 .11..211333 ++..1.11.11 11.+1.....+ 1+...2...11 ....1.111.. 121........ +..1.1..... +..1.+..... .1+.1......

Altitude (x 10 m)

Slope (x 10%) Area (x 100 square m) Faithful and differential taxa: Betula cordifolia Sorbus americana ss Acer spicatum ss Sorbus americana Aster acuminatus Sambucus canadensis Constants: Abies balsamea ss Dryopteris campyloptera Viburnum alnifolium Trientalis borealis Abies balsamea Oxalis montana Picea rubens ss Betula alleghaniensis Maianthemum canadense Aralia nudicaulis Other taxa: Betula cordifolia ss Prunus serotina Acer pensylvanicum ss Dennstaedtia punctilobula Cornus canadensis Clintonia borealis

Altitude (x 10 m)

Additional taxa with one or two occurrences: Betula alleghaniensis ss: 1 in rels. 17 and 22. Trillium undulatum: + in rels. 18 and 23. Carex arctata: 1 in rels. 23 and 24. Brachyelytrum erectum: 1 in rel. 17. Cornus alternifolia: + in rel. 17. Sambucus pubens: + in rel. 17. Viola selkirkii: + in rel. 17. Acer saccharum ss: + in rel. 17. Acer saccharum: + in rel 17. Picea rubens: + in rel. 18. Lycopodium annotinum: 2 in rel. 22. Fragaria virginiana: 1 in rel. 23. Thelypteris phegopteris: 1 in rel. 23. Amelanchier bartramiana: 2 in rel. 24.

Slope (x 10%) Area (x 100 square m) Faithful and differential taxa: Abies balsamea Picea rubens Abies balsamea ss Picea rubens ss Betula cordifolia Acer spicatum ss Acer pensylvanicum ss Viburnum alnifolium Aralia nudicaulis Betula alleghaniensis Clintonia borealis Sorbus americana ss Acer rubrum ss Betula alleghaniensis ss Acer rubrum Trillium undulatum Acer pensylvanicum Constants: Dryopteris campyloptera Maianthemum canadense Oxalis montana Trientalis borealis Other taxa: Aster acuminatus Lycopodium annotinum Betula cordifolia ss Lycopodium obscurum Fagus grandifolia Acer saccharum Coptis groenlandica

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Table 3. Continued Dennstaedtia punctilobula Tsuga canadensis Cornus canadensis Sorbus americana .++......1. ....2.11... ........+++ .1.......11 Table 4. Abies balsameaBetula cordifolia community (4a. var. with more abundance of Dryopteris campyloptera, Cornus canadensis, Trientalis borealis, Clintonia borealis, and Lycopodium annotinum. 4b. var. with more abundance of Picea rubens). Reference no.: 036. Mt. Washington Road, 06/14/95/05. 037. John Sherburne Ski Trail, 09/15/95/06. 038, 039, 040, 041, 042. Boott Spur Trail, 09/30/96/10,11,12,14,05. 043. Lion Head Trail, 10/02/96/06. 044. John Sherburne Ski Trail, 09/14/95/05. 045. Boott Spur Trail, 09/30/96/08. 046, 047, 048. John Sherburne Ski Trail, 09/15/95/08,03,07. 049, 050. Lion Head Trail, 10/02/96/04,03. 051. Tuckerman Ravine Trail, 10/02/96/02. 052. Lion Head Trail, 10/02/96/05. 053. Tuckerman Ravine Trail, 10/02/96/01. 054. Lion Head Trail, 10/02/96/07. 055, 056. Boott Spur Trail, 09/30/96/07,09.

Additional taxa with one or two occurrences: Cypripedium acaule: + in rel. 25, and 1 in rel. 29. Fagus grandifolia ss: 1 in rel. 28, and + in rel. 30. Tsuga canadensis ss: 2 in rels. 29 and 32. Acer spicatum: 1 in rel. 25. Monotropa uniora: + in rel. 25. Vaccinium angustifolium: + in rel. 26. Smilacina racemosa: + in rel. 31.

Forests data matrix ordination The ordination diagrams obtained using CA (Figures 4c, 4d) illustrate two main sources of variability. Axis 1 describes the altitudinal sequence, starting by Fagus grandifoliaAcer saccharum community (group 1) at the right side, following by Abies balsameaBetula alleghaniensis community (group 3) in the middle, and nishing with Abies balsamea Betula cordifolia community (group 4). Axis 2 seems to be related to the successional process between Betula cordifoliaSorbus americana community (group 2), and group 4 and the upper part of group 3 (Figure 8); it also expresses internal variability within group 4. Classication of krummholz and alpine vegetation Classication using similarity ratio and MNVAR (Figure 5a), euclidean distance and MISSQ (Figure 5b), and chord distance and MNVAR (group B in Figure 3b), yielded the same composition of plant communities, showing that a solid data structure was underlying. All of these classications suggested also two main groups, the rst one comprising the uppersubalpine krummholz and alpine dwarf shrub ericaceous dominated vegetation (group C in Figures 3b and 5), and the second one comprising the remaining alpine vegetation (group D in Figures 3b and 5). Ordination of krummholz and alpine vegetation The ordination diagrams obtained using CA (Figures 5c, 5d) illustrate two main sources of variability. Axis 1 describes the altitudinal sequence, starting by Picea marianaAbies balsamea community (group 5) at the right side, following by Vaccinium uliginosum Cetraria islandica community (group 6) in the middle, and nishing with groups 8, 9, 10, and 11 at the left side. Axis 2 mainly isolates Empetrum

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Figure 5. Analysis of krummholz and alpine vegetation. (a) Clustering using similarity ratio and MNVAR. (b) Clustering using euclidean distance and MISSQ. (c) CA ordination for samples. (d) CA ordination for taxa. Vegetation types: (C) Upper-subalpine krummholz, and alpine dwarf shrub ericaceous vegetation. (D) Remaining alpine vegetation. Plant community numbers correspond to those of Tables 5-12, and Appendix 2.

hermaphroditumVaccinium cespitosum community (group 7), and Deschampsia exuosaSolidago cutleri community (group 12). This axis seems to be related to the snow cover gradient. Axis 1 also distinguishes groups C and D obtained by clustering. For this reason, a new dimensionality reduction was carried out splitting the data matrix into two data matrices. The rst one comprising the upper-subalpine krummholz and alpine dwarf shrub ericaceous dominated vegetation (group C in Figures 3 and 5), and the second one comprising the remaining alpine vegetation (group D in Figures 3 and 5). Classication of krummholz and alpine dwarf shrub ericaceous dominated vegetation Classication using chord distance and MNVAR (Group C in Figure 3b), similarity ratio and MN-

VAR (Group C in Figure 5a), euclidean distance and MISSQ (Group C in Figure 5b), euclidean distance and CL (Figure 6a), and euclidean distance and MNVAR (Figure 6b) yielded the same composition of plant communities. Euclidean distance and CL (Figure 6a) provided an easy interpretation of this data set, so it was sorted according to this clustering method. Thus, three main plant communities were distinguished (Tables 57). Picea marianaAbies balsamea community. Table 5: samples 5786, Group 5 in Figures 3, 5, and 6 Upper-subalpine black sprucebalsam r krummholz vegetation in which two main variants have been distinguished: Closed krummholz constituted fundamentally by Abies balsamea with lower amounts of Picea mariana. Plants such as Cornus canadensis, Coptis groen-

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Figure 6. Upper-subalpine krummholz, and alpine dwarf shrub ericaceous vegetation analysis. (a) Clustering using euclidean distance and CL. (b) Clustering using euclidean distance and MNVAR. (c) CA ordination for samples. (d) CA ordination for taxa. Plant communities: (5) Picea marianaAbies balsamea. 5a. closed krummholz = var. with Cornus canadensis. 5b. scattered krummholz = var. with Vaccinium vitis-idaea ssp. minus. (6) Vaccinium uliginosumCetraria islandica. 6a. var. with Ledum groenlandicum and Betula cordifolia. 6b. var. without Ledum groenlandicum nor Betula cordifolia. (7) Empetrum hermaphroditumVaccinium cespitosum.

landica, Maianthemum canadense, Oxalis montana, and Gaulteria hispidula may be found inside of this vegetation (group 5a in Figures 3, 5, and 6; Table 5, samples 5765). Scattered krummholz, in which Abies balsamea has progressively less signicance at the same time that Picea mariana is increasing in abundance. The inuence of the alpine belt starts here with the presence of Vaccinium vitis-idaea, Cetraria islandica, Cladina rangiferina, Juncus tridus, and Carex bigelowii (group 5b in Figures 3, 5, and 6; Table 5, samples 66 86).

Vaccinium uliginosumCetraria islandica community. Table 6: samples 87115, Group 6 in Figures 3, 5, and 6 Lower-alpine dwarf shrub heath community dominated by Ericaceae, basically Vaccinium vitis-idaea and Vaccinium uliginosum. Cetraria islandica and Cladina rangiferina are the most important lichens, although other species such as Cladonia pleurota, Cetraria nivalis, Cladina stellaris, etc. also occur. Two variants have been distinguished: dwarf shrub heath variant characterized by the presence of Ledum groenlandicum, and Betula cordifolia (Table 6: samples 87101, Group 6a in Figures 3, 5 and 6). This variant is typically located just above

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Table 5. Picea marianaAbies balsamea community (5a. closed krummholz = var. with Cornus canadensis. 5b. scattered krummholz = var. with Vaccinium vitis-idaea subsp. minus). Reference no.: 57. Lion Head Trail, 09/13/95/02. 58, 59. John Sherburne Ski Trail, 09/15/95/10,09. 60, 61, 62. Lion Head Trail, 10/02/96/35,36,28. 63, 64, 65, Lion Head Trail, 09/18/95/38,37,39. 66. Davis Path, 09/18/95/30. 67. Lion Head Trail, 09/13/95/01. 68. Lion Head Trail, 10/02/96/22. 69. Davis Path, 09/18/95/28. 70, 71. Boott Spur Trail, 09/30/96/15,13. 72, 73. Lion Head Trail, 10/02/96/24,30. 74. Lion Head Trail, 09/18/95/04. 75. Lion Head Trail, 10/02/96/12. 76. Lawn Cutoff, 09/18/95/33. 77, 78, 79, 80, 81, 82, 83, 84. Lion Head Trail, 10/02/96/11,10,14,18,16,20,26,32. 85. Davis Path, 09/18/95/31. 86. Lion Head Trail, 09/18/95/02. Table 6. Vaccinium uliginosumCetraria islandica community (6a. var. with Ledum groenlandicum and Betula cordifolia. 6b. var. with Rhododendron lapponicum, Potentilla tridentata, and Diapensia lapponica). Reference no.: 87, 88, 89, 90, 91. Lion Head Trail, 10/02/96/13,17,15,19,25. 92. Boot Spur Trail, 09/30/96/16. 93. Lion Head Trail, 10/02/96/23. 94. Davis Path, 09/18/95/18. 95, 96, 97. Lion Head Trail, 10/02/96/21,29,31. 98. Lion Head Trail 09/13/95/08. 99. Lion Head Trail, 10/02/96/27. 100, 101. Lion Head Trail, 09/13/95/03,04. 102. Mt. Washington east face 10/01/96/26. 103, 104, 105. Mt. Washington, south face, 09/16/95/02,06,08. 106, 107. Davis Path, 09/18/95/21,27. 108, 109. Mt. Washington, south face, 09/16/95/04,09. 110. Between Alpine Garden Trail and Tuckerman Ravine Trail, 10/01/96/09. 111. Alpine Garden Trail, 10/01/96/12. 112. Lion Head Trail, 10/02/96/33. 113. Lion Head Trail, 09/18/95/03. 114. Between Alpine Garden Trail and Tuckerman Ravine Trail, 10/01/96/10. 115. The Cow Pasture, 10/01/96/34.

krummholz (Bliss 1963), at the beginning of the alpine belt. dwarf shrub heath variant characterized by the higher frequency of Rhododendron lapponicum, Diapensia lapponica, Potentilla tridentata, and Juncus tridus, and by the very low frequency of Ledum groenlandicum and Betula cordifolia (Table 6: sam-

ples 102-115, Group 6b in Figures 3, 5 and 6). This variant is located just above variant 6a. This community bears close resemblance to Cetraria cucullataVaccinium uliginosum community (Nakamura & Grandtner 1994).

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Table 7. Empetrum hermaphroditumVaccinium cespitosum community. Reference no.: 116. Mt. Washington south face, 09/16/95/11. 117. Between Alpine Garden Trail and Tuckerman Ravine Trail, 10/01/96/11. 118. Lion Head Trail, 09/18/95/06. 119. Mt. Washington, south face, 09/16/95/07. 120. Davis Path, 09/18/95/29. 121. Lion Head Trail to Tuckerman Ravine Trail, 09/18/95/07. 122. Tuckerman Crossover, 09/18/95/11. Plant community No. Plant community variant Relev No. e 0000000 7777777 ------1111111 1111222 6789012 1111111 7758656 5152252 6436434 0000000 0200000 6524444 3343324 2213143 2.1..1. ..2.122 ..1..1. 323232. 21.211. 2..1221 1..1.11 .+.2121 Table 8. Minuartia groenlandicaAgrostis mertensii Reference no.: 123, 124, 125, 126, 127.Lion 09/13/95/06,05,09,07,10. 128. Mt. Washington, east face, 129. Nelson Crag Trail, 10/01/96/02. 130. Lawn Cutoff, 131. Mt. Washington, east face, 10/01/96/24. 132. Mt south face, 09/16/95/01. 133. Davis Path, 09/18/95/20. Plant community No. Plant community variant Relev No. e community. Head Trail, 10/01/96/25. 09/18/95/32. Washington,

Altitude (x 10 m)

Altitude (x 10 m)

Slope (x 10%) Area (square m)

Slope (x 10%) Area (square m)

00000000000 88888888888 ----------11111111111 22222223333 34567890123 11111111111 54555875866 08204629762 52050000000 00000000000 00000000000 22424641621 23333133323 323223213.. 332322+1.3. 22221.+.... .....+111++ ......+.... 111........ 11......... 1.+.+...... 1.......... ..+........ .....1..... .....1..... .........+.

Faithful and differential taxa: Empetrum hermaphroditum Vaccinium cespitosum Coptis groenlandica Lycopodium annotinum Cornus canadensis Constants: Vaccinium uliginosum Carex bigelowii Cetraria islandica Vaccinium vitis-idaea ssp. minus Juncus tridus

Additional taxa with one or two occurrences: Scirpus caespitosus var. callosus: 2 in rels. 116 and 117. Potentilla tridentata: 1 in rels. 116 and 117. Ledum groenlandicum: + in rels. 116 and 119. Cladina rangiferina: 1 in rel. 119, and 2 in rel. 122. Loiseleuria procumbens: 1 in rel. 117. Sphagnum sp.: 1 in rel. 118. Deschampsia exuosa: + in rel. 121.

Faithful and differential taxa: Minuartia groenlandica Agrostis mertensii Juncus tridus Diapensia lapponica Carex bigelowii Potentilla tridentata Other taxa: Betula cordifolia kr Vaccinium uliginosum Vaccinium vitis-idaea ssp. minus Ledum groenlandicum Huperzia selago ssp. arctica Carex capillaris Cetraria islandica Cetraria nivalis

Empetrum hermaphroditumVaccinium cespitosum community. Table 7: samples 116122, Group 7 in Figures 3, 5 and 6 Lower-alpine dwarf shrub heath dominated by Empetrum hermaphroditum, Vaccinium cespitosum and Vaccinium uliginosum. It is located on gentle slopes and little depressions, where there is larger snow accumulation than in Vaccinium uliginosumCetraria islandica community. Due to this higher moisture condition, Lycopodium annotinum, Coptis groenlandica, Cornus canadensis, and Scirpus caespitosus var. callosus also occur here, and the frequency of lichens decreases.

Ordination of krummholz and alpine dwarf shrub ericaceous dominated vegetation The ordination diagrams obtained using CA (Figures 6c, 6d) illustrate two main gradients. Axis 1 describes the altitudinal sequence, starting by the closed krummholz variety of Picea marianaAbies balsamea community (group 5a), located at the right side, following with the scattered krummholz variety of the same community (group 5b). Next comes Vaccinium uliginosumCetraria islandica community var. with Ledum groenlandicum and Betula cordifolia (group 6a), nishing with the Vaccinium uliginosum Cetraria islandica community var. without Ledum groenlandicum nor Betula cordifolia (group 6b), lo-

95

Figure 7. Analysis of the remaining alpine vegetation. (a) Clustering using similarity ratio and CL. (b) Clustering using chord distance and MNVAR. (c) CA ordination for samples. (d) CA ordination for taxa. Plant communities: (8) Minuartia groenlandicaAgrostis mertensii. (9) Diapensia lapponicaRhododendron lapponicum. (10) Carex bigelowiiSolidago cutleri. (11) Salix uva-ursiSolidago cutleri. (12) Deschampsia exuosaSolidago cutleri.

cated at the left side. Axis 2 mainly isolates Empetrum hermaphroditumVaccinium cespitosum community (Group 7), due to its different oristic composition supported on larger snow accumulation. Classication of the remaining alpine vegetation Classication using chord distance and MNVAR (Group D in Figure 3b), similarity ratio and MNVAR (Group D in Figure 5a), euclidean distance and MISSQ (Group D in Figure 5b), similarity ratio and CL (Figure 7a), and chord distance and MNVAR (Figure 7b) yielded the same composition of plant communities. Similarity ratio and CL (Figure 7a) provided an easy interpretation of this vegetation, so it was sorted according to this clustering method. Thus, ve main plant communities were distinguished (Tables 812).

Minuartia groenlandicaAgrostis mertensii community. Table 8: samples 123133, Group 8 in Figures 3, 5 and 7 Alpine community growing in rock cracks at the beginning of the alpine belt consisting of Minuartia groenlandica, Agrostis mertensii, Juncus tridus, and Diapensia lapponica. Also can be located on moister sites, along alpine trails and other disturbed locations, as primary succession communities on sandy soils, characterized by Minuartia groenlandica, Agrostis mertensii, Juncus tridus, little amounts of Carex bigelowii, and very low amounts of Diapensia lapponica. Data belonging to this community have been reported by Bergeron et al. (1984), and Nakamura & Grandtner (1994) from Mount du Lac des Cygnes.

96

Table 9. Diapensia lapponicaRhododendron lapponicum community. Reference no.: 134. Nelson Crag Trail, 10/01/96/05. 135. Alpine Garden Trail, 10/01/96/21. 136. The Cow Pasture, 10/01/96/17. 137, 138, 139. Davis Path, 09/18/95/25,24,17. 140. Tuckerman Crossover, 09/18/95/14. 141. Lion Head Trail, 10/02/96/34. 142. Davis Path, 09/18/95/26. 143. Nelson Crag Trail, 10/01/96/03. 144. The Cow Pasture, 10/01/96/13. 145. Lion Head Trail, 09/18/95/01. 146, 147. Davis Path, 09/18/95/22,23. 148. Tuckerman Crossover, 09/18/95/12. 149. Lawn Cutoff, 09/18/95/34. 150. Davis Path, 09/18/95/15. 151. Tuckerman Crossover, 09/18/95/13. 152. Davis Path, 09/18/95/16. Plant community No. Plant community variant Relev No. e 0000000000000000000 9999999999999999999 ------------------1111111111111111111 3333334444444444555 4567890123456789012 1111111111111111111 7776666667756665666 1021133212221139333 0000000000000000000 0000000000000000000 0000000000010000000 4962222422902222211 4342322221222222311 2..4434445434444334 1..21122212222112.. 1...12222..11122.1. 11+......++......2. .+2.1..+.+.2.......

Table 10. Carex bigelowiiSolidago cutleri community. Reference no.: 153. Ball Crag, 10/01/96/28. 154, 155, 156, 157, 158. The Cow Pasture, 10/01/96/15,14,18,16,29. 159. Alpine Garden Trail, 10/01/96/08. 160. Tuckerman Ravine Trail, 09/18/95/08. 161. Nelson Crag Trail, 10/01/96/07. 162. Alpine Garden Trail, 10/01/96/06. 163. Ball Crag, 10/01/96/27. 164. Mt. Washington, east face, 10/01/96/23. 165, 166. Nelson Crag Trail, 10/01/96/04,01. 167. The Cow Pasture, 10/01/96/20. 168. Mt. Washington, south face, 09/16/95/03. 169. Davis Path, 09/18/95/19. Plant community No. Plant community variant Relev No. e 11111111111111111 00000000000000000 ----------------11111111111111111 55555556666666666 34567890123456789 11111111111111111 77777776778877786 22222220324712022 00000000000000040 00000000000000000 11111210102200020 52602004260068954 54454554555555555 222332221....1... 2221.2+.+........ +....11.22112211. 21122...112...... .+.11.1.12....11.

Altitude (x 10 m)

Altitude (x 10 m)

Slope (x 10%) Area (square m)

Slope (x 10%) Area (square m)

Faithful and differential taxa: Juncus tridus Diapensia lapponica Rhododendron lapponicum Loiseleuria procumbens Minuartia groenlandica Agrostis mertensii Constants: Potentilla tridentata Carex bigelowii Other taxa: Vaccinium uliginosum Cetraria islandica Vaccinium vitis-idaea ssp. minus Solidago cutleri Cladina rangiferina Cetraria nivalis Salix uva-ursi Ledum groenlandicum Huperzia selago ssp. arctica

Faithful and differential taxa: Carex bigelowii Potentilla tridentata Solidago cutleri Constants: Minuartia groenlandica Agrostis mertensii Juncus tridus Other taxa: Vaccinium uliginosum Cetraria nivalis Diapensia lapponica Cetraria islandica Rhododendron lapponicum Stellaria borealis Vaccinium cespitosum Salix uva-ursi Geum peckii Sphagnum sp. Vaccinium vitis-idaea ssp. minus Carex capillaris Poa glauca

22222211111122..... .11112211211121...2

...2.+++.+.+12+.1.. 2321..++.....+..... ........1..1+1..... 1.+........1....... .1.1.........+..... ...1..+......+..... ...2............... .....1............. ............+......

1....1.1+........ ..1.++........... .++.............. ..1.+............ ..+...+.......... ..........+....l. ..1.............. .....1........... .......2......... .......2......... .......1......... ..........+...... ...............+.

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Table 11. Salix uva-ursiSolidago cutleri community. Reference no.: 170. The Cow Pasture, 10/01/96/30. 171. Nelson Crag Trail, 10/01/96/31. 172. Alpine Garden Trail, 10/01/96/32. 173, 174. The Cow Pasture, 10/01/96/33,22. Plant community No. Plant community variant Relev No. e 11111 11111 ----11111 77777 01234 11111 77777 22220 00000 00000 00110 69229 44444 21+1+ 22212 22111 21112 222.1 2112. 1222. 11.1. Table 12. Deschampsia exuosaSolidago cutleri community. Reference no.: 175, 176, 177, 178. Mt. Washington, south face, 09/16/95/05,10,13,12. 179, 180, 181. Close to Tuckerman Junction, 09/18/95/09,37,10. 182, 183. Tuckerman Ravine Trail, 09/18/95/35,36. Plant community No. Plant community variant Relev No. e 111111111 222222222 --------111111111 777778888 567890123 111111111 877766655 383300155 300000000 000000000 001111112 680000000 323242323 112222333 3332..11. .+1211..1 .11.211.1 ...123414 ....22222 .....2121 .....+111 ..1.11... ....112..

Altitude (x 10 m)

Altitude (x 10 m)

Slope (x 10%) Area (square m)

Slope (x 10%) Area (square m)

Faithful and differential taxa: Salix uva-ursi Solidago cutleri Constants: Cetraria islandica Cetraria nivalis Carex bigelowii Agrostis mertensii Juncus tridus Diapensia lapponica Rhododendron lapponicum Other taxa: Minuartia groenlandica Cladina rangiferina Vaccinium cespitosum Vaccinium uliginosum

Faithful and differential taxa: Deschampsia exuosa Solidago cutleri Vaccinium cespitosum Cornus canadensis Coptis groenlandica Veratrum viride Geum peckii Dryopteris campyloptera Scirpus caespitosus var. callosus Lycopodium annotinum Sphagnum sp. Constants: Carex bigelowii Other taxa: Carex brunnescens Juncus tridus Cetraria islandica Vaccinium uliginosum Agrostis mertensii Calamagrostis canadensis Thelypteris phegopteris Dennstaedtia punctilobula Luzula parviora var. melanocarpa

1.... .2... .+... ..1..

223332123

Diapensia lapponicaRhododendron lapponicum community. Table 9: samples 134152, Group 9 in Figures 3, 5 and 7 Upper-alpine wind-exposed, snow-free community dominated by Diapensia lapponica, Juncus tridus, Rhododendron lapponicum, and Loiseleuria procumbens, accompanied by Carex bigelowii, Potentilla tridentata, and Vaccinium uliginosum. Carex bigelowiiSolidago cutleri community. Table 10: samples 153169, Group 10 in Figures 3, 5 and 7 Alpine sedge meadows dominated by Carex bigelowii accompanied by Potentilla tridentata, Solidago cutleri, Minuartia groenlandica, Agrostis mertensii, and Juncus tridus.

111...... .1+...... .1....... ...1..... ...1..... ......1.1 .......12 .......1+ ......1..

Salix uva-ursiSolidago cutleri community. Table 11: samples 170174, Group 11 in Figures 3, 5 and 7 Alpine dwarf shrub willow community dominated by Salix uva-ursi, accompanied by Solidago cutleri, Cetraria nivalis, Cetraria islandica, Carex bigelowii, Agrostis mertensii, Juncus tridus, Diapensia lapponica, and Rhododendron lapponicum.

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Table 13. Summary information of CA ordination analyses. Axis 1 Overall data matrix: Eigenvalues Cumulative percentage variance Forests: Eigenvalues Cumulative percentage variance Krummholz and alpine vegetation: Eigenvalues Cumulative percentage variance Krummholz and alpine ericaceous vegetation: Eigenvalues Cumulative percentage variance Remaining alpine vegetation: Eigenvalues Cumulative percentage variance Axis 2 Axis 3 Axis 4 Sum of eigenvalues

0.900 14.1

0.586 23.3

0.538 31.7

0.347 37.2

6.379

0.364 21.8

0.167 31.8

0.134 39.9

0.091 45.4

1.667

0.672 13.4

0.632 25.9

0.382 33.6

0.326 40.0

5.028

0.529 17.2

0.391 29.9

0.285 39.2

0.221 46.3

3.077

0.738 21.8

0.372 32.7

0.337 42.7

0.285 51.1

3.392

Geum peckii, Vaccinium cespitosum, Carex bigelowii, Coptis groenlandica, and Cornus canadensis. Ordination of the remaining alpine vegetation The ordination diagrams obtained using CA (Figures 7c, 7d) illustrate two main gradients. Axis 1 represents the snow-cover gradient, starting by the snowfree and wind-exposed communities of Diapensia lapponicaRhododendron lapponicum (group 9) and Minuartia groenlandicaAgrostis mertensii (group 8) at the left side, following by Salix uva-ursiSolidago cutleri (group 11) and Carex bigelowiiSolidago cutleri (group 10). At the end of the sequence are located the snow-banks communities of Deschampsia exuosaSolidago cutleri (group 12). Axis 2 isolates DiapensiaRhododendron lapponicum (group 9) and Salix uva-ursiSolidago cutleri (group 11) from Minuartia groenlandicaAgrostis mertensii (group 8) and Carex bigelowiiSolidago cutleri (group 10). Figure 8 plus CA ordinations suggest that distribution of plant communities within the alpine belt has no correlation with altitude, but with mesotopographic conditions (snow accumulation, exposure and cryoturbation, slope position, and soil moisture). Probability ellipses and Students t-test After the plant communities classication, computer program ELLIPSE (Podani 1993, 1994) superimposed

Figure 8. Box plots showing the altitudinal distribution of the twelve plant communities described, and the estimated boundaries between montane, subalpine, and alpine bioclimatic belts. Plant community numbers correspond to those of the twelve tables and Appendix 2.

Deschampsia exuosaSolidago cutleri community. Table 12: samples 175183, Group 12 in Figures 3, 5 and 7 Alpine snowbank community characterized by Deschampsia exuosa, Solidago cutleri, Veratrum viride,

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Table 14. Probabilities that the null hypothesis is true among the twelve groups, i.e., the probabilities among every pair of plant communities of being the same community (axis 1 and 2 of CA ordinations, respectively, at 95% signicance level). Bold symbols indicate probabilities P > 0.40 103 .
CA 1 2 3 4 5 6 7 8 9 10 11 12 1 1 0.28E 07 0.78E 06 0.79E 06 0.15E 06 0.80E 06 0.80E 06 0.79E 06 0.80E 06 0.80E 06 0.80E 06 0.80E 06 2 1 0.046 0.98E 07 0.40E 07 0.78E 06 0.80E 06 0.81E 06 0.81E 06 0.81E 06 0.80E 06 0.80E 06 3 4 5 6 7 8 9 10 11 12

1 0.83E 07 0.79E 06 0.75E 07 0.40E 07 0.93E 07 0.14E 06 0.82E 08 0.29E 07 0.90E 07

1 0.78E 061 0.36E 08 0.62E 07 0.76E 08 0.45E 07 0.55E 07 0.13E 07 0.33E 07

0.77E 06 0.82E 06 0.80E 06 0.79E 06 0.80E 06 0.81E 06 0.79E 06

1 0.35E 04 0.17E 06 0.15E 06 0.19E 06 0.69E 07 0.11E 07

1 0.24E 06 0.39E 07 0.45E 07 0.77E 05 0.52E 07

1 0.44E 03 0.75E 02 0.130 0.62E 06

1 0.13E 06 1 0.150 0.13E 05 1 0.14E 06 0.68E 08 0.38E 05 1

classication results on Figure 3c by drawing ellipses of equal concentration around the centroids of clusters (Mardia et al. 1979; Lagonegro & Feoli 1985) at 95% probability level (Figure not shown). The mean and standard deviation of each ordination dimension for every community were calculated. Differences between means among the twelve communities were tested by Students t-test, giving the probabilities that the null hypothesis is true, i.e., the probabilities among every pair of plant communities of being the same community. Results of the Students t-test are shown in Table 14. The highest probability of the null hypothesis to be true, i.e., the probability of being the same plant community, based on axes 1 and 2 of CA ordination (P = 0.150) is that between Diapensia lapponica Rhododendron lapponicum (group 9) and Salix uvaursiSolidago cutleri (group 11). This result seems to agree with Danils (1994) who considers Salix uvaursi characteristic to LoiseleurioDiapension communities. Next lower probability (P = 0.130) is between Minuartia groenlandicaAgrostis mertensii (group 8) and group 11, also belonging to Loiseleurio Diapension. Next lower probability (P = 0.046) is between Betula cordifoliaSorbus americana (group 2) and Abies balsameaBetula alleghaniensis (group 3). This seems to be related to the similarity in their ora (see Tables 2 and 3, and Appendix 2), but the rst one is a soft-wood secondary forest of Abies balsameaBetula cordifolia community (group 4), whereas the second one is an Abies balsameaPicea rubens dominated community growing on xeric soils in contact with Fagus grandifoliaAcer saccharum community (group 1). At the beginning of the sub-

alpine belt group 2 seems to be the secondary forest of group 3 (see Figure 8). Next lower probability (P = 0.75 102) is between group 8 and Carex bigelowii Solidago cutleri (group 10). Next lower probability is between groups 8 and 9 (P = 0.44 103 ), both of them belonging again to LoiseleurioDiapension. The remaining probabilities are lower than 104 .

Acknowledgements This study was possible thanks to a postdoctoral fellowship awarded to the author by the Ministry of Education and Science of Spain for staying at Queens College of the City University of New York, and The New York Botanical Garden during 19951996. This paper improved with the help of Dr Andrew M. Greller (QC of CUNY), Dr Leslie F. Marcus (QC of CUNY), Dr D. S. A. Wijesundara (then of the QC of CUNY), Dr Enrique Forero (then of the NYBG), Dr Salvador Rivas-Martnez (Dep. Biologa Vegetal II, Universidad Complutense de Madrid), and two anonymous referees. Dr Vctor J. Rico (DBV II of UCM) revised the lichenic material. The author is grateful to all of them.

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Appendix 1. Field sampling design. The sampling was done using the centralized replicate sampling procedure (Mueller-Dombois & Ellenberg 1974). Plot locations were subjectively chosen between the two meteorological stations, in areas of homogeneous vegetation, according to the following plant-forms and physiognomic-ecological plant formations (Rbel 1930; Raunkiaer 1937; Braun-Blanquet 1964; Ellenberg & Mueller-Dombois 1967a,b; Whittaker 1975; Box 1981). Plant forms 1. Deciduous summergreen broadleaved hardwood trees 2. Deciduous summergreen broadleaved softwood trees 3. Evergreen boreal/subalpine needleleaved trees 4. Evergreen boreal/subalpine needleleaved treeline krummholz 5. Arctic/Alpine dwarf-shrubs Plant formations Hardwood forests Softwood secondary forests Boreal/Subalpine needle forests Krummholz Arctic/alpine heath, prostrate willow vegetation Arctic/alpine dwarf cushion shrub vegetation Sedge meadows Megaforb snowbanks Examples Fagus grandifolia, Acer sp. pl. Betula cordifolia, Betula alleghaniensis Abies balsamea, Picea rubens Abies balsamea, Picea mariana Vaccinium uliginosum, Ledum groenlandicum, Empetrum hermaphroditum, Salix uva-ursi Rhododendron lapponicum, Loiseleuria procumbens, Diapensia lapponica Carex bigelowii Veratrum viride

6. Arctic/Alpine dwarf cushion shrubs 7. Grasses 8. Summergreen megaforbs

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Appendix 2. Synoptic table of all vegetation communities. Bold symbols indicate faithful and differential taxa for one group. Regular boxes indicate taxa preferential to multiple groups. 1.Fagus grandifoliaAcer saccharum. 2.Betula cordifoliaSorbus americana. 3.Abies balsameaBetula alleghaniensis. 4.Abies balsameaBetula cordifolia. 5.Picea marianaAbies balsamea. 6.Vaccinium uliginosumCetraria islandica. 7.Empetrum hermaphroditumVaccinium cespitosum. 8.Minuartia groenlandicaAgrostis mertensii. 9.Diapensia lapponicaRhododendron lapponicum. 10.Carex bigelowiiSolidago cutleri. 11.Salix uva-ursiSolidago cutleri. 12.Deschampsia exuosaSolidago cutleri.

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Appendix 2. Continued.