The Alpine Tussock Grasslands
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12 MARK AND BURRELL: THE ALPINE TUSSOCK GRASSLANDS V E GET A T ION S T U DIE SON TH E HUM B 0 L D T M 0 U NT A INS FIORD LAND PART 1: THE ALPINE TUSSOCK GRASSLANDS A. F. MAHKo AND JULIET BUHHELLt Botany Department, University 0/ Otago, Dunedin INTRODUCTION NOMENCLATUHE Nomenclature follows Allan (1961) for Tussock grassland vegetation on the western pteridophytes, gymnosperms, and dicotyledons, slope of the Humboldt Mountains overlooking and Cheeseman (1925) for the monocotyledons, the Hollyford Valley jn northern Fiordland except for the grass genera Chionochloa and was analysed quantitatively with a series of Notodanthonia which follow Zolov (1963), or sample plots at six sites. These plots were \vhere authorities are cited. located on the prominent spur which runs southwest from the summit of Ocean Peak THE AREA (1,848 m.-Fig. 1). The study was made to determine whether changes in composition and Geologically the area consists of partly cover of the grassland vegetation bet\veen the schistose grey wacke except for a band of silver beech (Nothofagus menziesii) timber line undifferentiated b3sic volcanic rock, cOlumonly and the upper limits of closed vegetation at schistose and highly altered, along the summit about 1~640 ill., conform to a gradient or of the range (VVood 1962). Both belong to the whether discrete communities are recognisable. Caples Group. Since the transect began within 3 krn. of the Topography is moderatcly to steeply undu- Deadman's-Harris Saddle track, it could be lating whh the slope generally between 20° considered as an upward extension of the alti- and 35°, but steepening appreciably with a tudiu::ll gradient already described for the forest marked increase in bare rock over the last 300 (Mark and Sandcrson 1962). Ill. of Ocean Peak. The westerly aspect exposes all sites to prevailing winds. Precipitation data The work was carried out by a party of are available for Homer Tunnel (650 Ill.) 16 botany students and staff during a University km. to the west in the Upper Hollyford Valley of Otago Science Students' Association expedi- where mean annual rainfall is 711 em., and tion in May, 1963. for Marian Camp (344 m.) 10 km. to the southwest on the valley floor where it is 450 em. An annual precipitation of over 500 em., evenly distributed throughout the year, can thus be expected on the area. Information on tempera- ture and snow is not available, aJthough snow may fall durjng any month :md usually covers the ground for considerable perjods during winter. There was no evidence of recent fires nor of heavy grazing by deer, although several deer '"ere sighted. METHODS Sampling sitcs \vere selected to sbow changes in vegetfltion which seemed to be due to alti- tude. There \Vas no fixed interval between them; indeed at the highest n1titude (1,640 m.) FIGUHE 1. View northeast from ahout (220 m. Miss E. L. Hellaby Grasslands Research Fellow. to the summit of Ocean Peak at 1,848 m., t Present address: Botllny School, University of Me}. covering the location of four of the six sites. honrne. Australi<l. MARK AND BURRELL: THE ALPINE TUSSOCK GRASSLANDS 13 two were contiguous, one being in an area (site 6 in Table 1) where snow accumulates. where snow accumulates. The greatest interval At this site the upper stratum is virtually in elevation was 200 m. absent and the luiddle stratum is also of minor At each site, measurements were made of importance. The smaller snow tussock C. plant cover and frequency of species. Cover crassiuscula (no. 51) is absent from the timber was measured with point intercepts at 30 em. line area and of minor importance above 1,500 intervals along 10 lines each 15 m. long, m_, being rare where snow accumulates (site 6). making a total of 500 points per site. The Between 1,100 m. and 1,500 m., however, it is lines were parallel, about five m. apart, and at an important co-dominant. Celmisia petriei right angles to the contour. Three strata were (18) is common as a sub-dominant below 1,500 recognised. The upper stratum (> 30 em. m., and Astelia cockaynei (41) and Schoenus high) included tall subalpine shrubs and herbs pauciflorus (42) occur throughout. which occur close to timber line, and the large Among the ground layer species Dacrydium snow tussock Chionochloa flavescens. The laxifolium (58) and Astelia linearis (32) are middle stratum (15-30 em. high) included important at site 2 (1,140 m.) where the medium herbs and shrubs such as Celmisia reduced slope somewhat impedes drainage. petriei and Hebe hectori, and the smaller snow Chionochloa oreophila (101) is confined to the tussock Chionochloa crassiuscula. The ground two high-altitude sites but is important and stratum « 15 cm. high) included dwarf shrubs dominant only where snow accumulates (site and herbs such as Myrsine llummularia and 6). Poa colensoi (37) occurs throughout but Astelia linearis. Tall plants were also included increases in importance with altitude to become in lower strata if they were intercepted there. co-dominant with C. oreophila and Celmisia Since only the first hits were recorded for hectori (98) at site 6. plants in each stratum, the term dominance Distributions of certain species of Celmisia may be more appropriate than cover, to express are of interest. C. bonplandii (35) occurs at the results (Hanson 1962) . Values for per- all sites but is of minor importance. C. petriei centage dominance for each species in each (18) is confined to the lower four sites, whereas stratum were based on the number of inter- C. walkeri (61) and C. glandulosa (62) are ceptions. In addition, species lists were com- common between 1,120 m. and 1,500 m. C. piled for a rectangular plot 2 m. wide, centred petiolata (76) and C. sessiliflora (77), on the on each line. Frequency values for each species other hand, were seen above about 1,150 m. and were based on occurrence in the 10 plots. C. hectori (98) occurs only above 1,500 m. Voucher specimens of most of the species Bare earth is unconunon at all sites but recorded are deposited in the Otago University Herb3rium. litter, mostly leaves of snow tussock and, near timber line, of Dracophyllum, covers up to 48% of the ground. RESULTS Index of Similarity Between Sites Values for frequency and percentage domin- Sorensen's Quotient of Similarity-K (Soren- ance in each stratum are shown in Table 1, sen 1948)-was used to assess the degree of "vhere species are arranged by strata and then relationship between the flora at the various according to altitudinal distribution. The habit sites. The method involves expressing the of each species is also indicated as follows, tall species common to each pair of sites as a shrubs and herbs, > 30 cm. high; medium percentage of the total species present at both, shrubs and herbs, 15-30 cm. high; and dwarf thu" shrubs and herbs, < 15 cm. high. The many 2C patterns of species distribution shown in Table Kab = X 100 1 suggest an altitudinal gradient. Tall shrubs A + B [lrc almost confined to the vicinity of the timber where A and B arc tbe number of species at line where they contribute about half of the sites a and b, and C is tl'c number s'harcd. dominance in the upper stratum. Several other Results for the six sites are shown in Table 2. woody and herbaceous species (nos. 1, 2, 4, 5, Each coefficient was then tested for significance 10, 11 and 12 in Table 1) are similarly by the method reccmmended by Looman and restricted in distribution. The tall snow Campbell (1960). tussock Chionochloa flavescens (no. 44) is The use of this test of floristic affinity might important at all except the high-altitude site be questicned en the basis that sites were 14 MARK AND BURRELL: THE ALPINE TUSSOCK GUASSLANDS selected rather than chosen at random. How- present above about 1,100 m., is a tussock ever, selection was based largely on topography grassland dominated by C. f/avescens and the and dominant species, and these species, being shorter C. crassiuscula. In snow pockets above a small proportion of the total flora at a site, about 1,500 m. this is replaced by the third make only a minor contribution to the results. community, in which the dwarf tussocks C. Similarities in the vegetation between each oreophila and Poa colensoi share dominance of the six sites were determined by a similar with Celmisia hectori. Both grass species method but using different units (Oosting characterise snow hollows in other parts of the 1956). The cover provided by species com- South Island (Burrows 1962). Tall shrubs mon to a pair of sites is expressed as a per- are unimportant in the grassland except near centage of the total cover at the two sites, thus: timber line, but several species of Celmisia 2W are prominent. The grassland dominated by Cab = X 100 tall evergreen species of Chionochloa at sites A + B 1 to 5 would come within the "low(er) alpine" where A and B are the totals of percentage zone, whereas the dwarf grassland occupying plant cover at sites a and b, and W the sum the site of snow accumulation (site 6) would of the lowest of each pair of percentage cover be designated "high alpine" (Wardle 1964). values of species present at both sites. The A consideration of floristic affinities between dominance values for the three strata were sites substantiates separation of the snow combined for these analyses. No test of tussock-scrub as a distinct community within significance is available for these coefficients.