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A study of modern pollen deposition, Southern Alps, South Island, New Zealand
Review of Palaeobotanyand Palynology, 64 (1990): 263-272
263
Elsevier Science Publishers B.V., Amsterdam
A study of modern pollen deposition, Southern Alps, South Island, New Zealand P.M. R a n d a l l Department of Plant and Microbial Sciences, University of Canterbury, Christchurch (New Zealand) (Received October 11, 1988; revised and accepted January 25, 1990)
ABSTRACT Randall, P.M., 1990. A study of modern pollen deposition, Southern Alps, South Island, New Zealand. Rev. Palaeobot. Palynol., 64: 263-272. The pollen and spores dispersing from natural and semi-natural vegetation in the South Island of New Zealand were studied from moss polsters collected from 35 sites along a transect from Westland across the Southern Alps to Canterbury. Sites in Westland were dominated by local broadleaved angiosperm species (Metrosideros,Quintinia, Weinmannia)and Poaceae pollen. Sites above the treeline on the Main Divide showed anomalous high counts of exogenous Podocarpaceae pollen. This seems to be washed out of strong NW winds by orograpbic precipitation. Peat cores taken from 3 sites confirm that exogenous podocarp pollen influx has continued over at least the last 500 years. Nothofaguspollen dominates within the beech forest areas, but drops to 10% a short distance from the forest edge. Poaceae pollen shows low frequencies in forested sites, but dominates in grassland areas.
Introduction
Study area
Prior to the w o r k of M o a r (1970, 1971), Pocknall (1978, 1980, 1982) and M c G l o n e (1982), pollen analysts in New Zealand lacked detailed information on the relationship between modern pollen deposition and the vegetation from which the pollen originated. A s u m m a r y of pollen deposition studies up to 1981 is outlined by Pocknall (1982). Most such work in New Zealand has attempted to discover information a b o u t relative dispersal and p r o p o r t i o n a t e representation of pollen o f the main taxa present in Quaternary sites. The aim o f the present study is to amplify information a b o u t the m o d e r n pollen deposition in order to assist interpretation of Quaternary pollen diagrams. M a n y o f the sites suitable for Quaternary pollen analysis are in mountain localities in the Southern Alps, where pollen accumulates and is preserved in Sphagnum bogs or other wet sites.
The study area contains a complex system of mountain ranges, valleys and basins with peaks rising to 2270 m and valley floors at 460 to 610 m. Arthur's Pass, on the Main Divide, which forms the boundary between Westland and Canterbury, is at an elevation of 920 m. The mixed forest of the Taramakau and Otira valleys in Westland contains substantial amounts of the podocarp tree species Dacrydium
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cupressinum, Podocarpus hallii, Prumnopitys ferruginea, P. taxifolia and Dacrycarpus dacrydioides but is dominated by angiosperm species Weinmannia racemosa, Metrosideros umbellata, smaller amounts of Quintinia acutifolia, and many other angiosperm species. Nothofagusfusca is present in the Otehake and Taramakau river catchment area with some Nothofagus solandri var. cliffortioides and N. menziesii. Scattered stands of N.fusca occur on young river terraces of the lower Otira river and a small stand of N. menziesii in the lower Otira gorge.
© 1990- Elsevier Science Publishers B.V.
264
There is a steep rise from the Westland valleys to the Main Divide where the vegetation is predominantly subalpine scrub, grassland and herbfield (Calder and Wardle, 1969). At the Canterbury side there is a gradual drop into the Bealey River valley in which Nothoj~tgus solandri var. cliffortioides dominates. It is accompanied by N.fusca and N. menziesii in the upper Poulter River and N. fusca near the Bealey river confluence with the Waimakariri and in the Hawdon and Andrews river catchment (Wardle, 1970). Further east, in the Waimakariri basin, the forest is discontinuous. Extensive grassland and scrubland dominates, with patches of Nothofagus solandri var. cliffortioides on the Craigieburn and Torlesse Ranges. The main vegetation in the Porter Pass area is mostly Dracophyllum scrub and grassland with only isolated small patches of Nothofagus solandri var.
cliffortioides. Precipitation is high in Westland ( > 5000 mm per annum), diminishing eastward (c. 1750 mm at the Bealey confluence, 1000 mm at Castle Hill). In the mountains strong westerly winds predominate with peaks in spring and summer (Coulter, 1973). Methods
Moss polsters were collected at 35 sites on a transect from Westland to Canterbury across the Main Divide (Fig. 1 and Appendix 1). At least four to six subsamples from each site were combined into one sample. To reduce the possibility of differential filtration of pollen by the moss polster, the whole thickness of living moss was removed (Crowder and Cuddy, 1973). Moss polsters are believed to yield pollen spectra representative of t h e pollen deposition for one to several years. Samples were collected where possible from open bogs and not from beneath forest canopy. The samples were transported to the laboratory in sealed plastic bags. The samples for pollen analysis were boiled in 5% sodium hydroxide, sieved and treated with hot
P.M RANDALL
40% hydrofluoric acid to remove silicates, followed by standard acetolysis (Faegri and Iversen, 1964), before mounting in glycerine jelly. The pollen and spore counts were carried out at a magnification of x 400 and 700 pollen grains and spores were counted per sample along regularly spaced traverses of the slide. To obtain additional information regarding the deposition of pollen on sites at Arthur's Pass on the Main Divide, a spore marker was added to five surface samples to determine the absolute numbers of pollen and spores present. The method of Stockmarr (1971) was used. Two Lycopodium tablets containing 10,850_+200 spores per tablet were added to one gram of dry material and processed as above. The analysis was done in the normal way, counting to 700 pollen and spores of all types, plus the number of marker Lycopodium. The pollen sum, on which the percentage calculations are based, consists of all terrestrial taxa excluding those, such as sedges, that are mainly restricted to bogs and wet sites. All fern spores were also excluded from the pollen sum. Three peat monoliths (38 to 42 cm deep) were sampled to establish the immediate past history of the vegetation in the region. The fossil pollen spectra were then compared with the surface samples paying particular attention to the pollen markers of modern times (Pinus and Salix) as well as prominent native species (Dacrydium cupressinum, Prumnopitys and Nothofagus fusca type). The record of European settlement in the area is well documented. Using the first occurrence of pollen of indicator species (Pinus and Salix) as well as charcoal, approximate dates can be fixed for the peat cores. Samples were collected using 50 cm PVC tubes of 20cm diameter, taken to the laboratory and sectioned at 2 c m intervals for pollen analysis. Subsamples were taken at the middle and bottom of the core for radiocarbon analysis. Radiocarbon samples were pretreated by the removal of roots which had grown down into the
Fig. 1. Location of the 35 sites for surfacesamplescollectedalong a transect from Westland to Canterbury across the Main Divide and the two main forest typesin the study area. Podoearp/broadleafforest (light shading) and beechforest (dark shading), and grassland or subalpine (clear areas). For details of sites see Appendix 1. Based on the NZFS Forest class maps series 6 No. 18, 19, 20 (1974).
265
STUDY OF MODERN POLLEN DEPOSITIONAL, NEW ZEALAND
SOUTH ISLAND
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at sites E-G. Nothofagus solandri var. cliffortioides forest is present nearby. (3) Nothofagus fusca type (which includes both N.fusca and N. solandri var. cliffortioides) dominates at sites H-L, all surrounded by beech forest. (4) At all other sites to the east, grasses are dominant. Grass pollen is also dominant at the westernmost site (site A) which is surrounded by pasture, and abundant at the Main Divide sites, but unimportant at sites closely surrounded by beech forest (e.g. sites H-L). Nothofagusfusca type values, high in forest sites, fall to less than 10% in sites mainly surrounded by grassland and scrub. Exceptions are minor Nothofagus pollen peaks where samples were taken near small pockets of beech forest.
peat. This was done to reduce the possibility of contamination by the presence of younger carbon. The samples were then dried and forwarded to New Zealand Radiocarbon Dating Laboratory, Institute of Nuclear Sciences, DSIR, Lower Hutt for age determination. Pollen diagrams
Surface samples The results of counts from the surface samples are shown in Fig.2. The outstanding features are: (1) At sites B-E, broadleaved angiosperm counts are high (predominantly Metrosideros 320/0, with Quintinia 8%, and Weinmannia 4%) and Nothofagusfusca type and Podocarpaceae are very low. (2) Pollen of Podocarpaceae predominates (Dacrydium cupressinum, Prumnopitys and Dacrycarpus dacrydioides) at sites E - G on the Main Divide, where these trees are absent (they are virtually confined to sites west of the Main Divide). Nothofagusfusca type pollen is also very abundant
Peat Cores Is the array of pollen spectra described above from modern surface samples reflected through time, or is it strictly a modern phenomenon due to either (i) the length of time represented by the pollen deposited in the moss polsters or (ii) unusual
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Fig.2. Pollen diagrams from sites of the West-East transect, from Westland (Site A) across the Main Divide (Sites E-G) to Porters Pass (Site AI). The Podocarpaceae pollen (Dacrycarpusdacrydioides, Dacrydium cupressinum, Prumnopitys) at the Main Divide (Sites E-G) is not local (see text). Only major pollen taxa shown.
267
STUDY OF MODERN POLLEN DEPOSITIONAL, NEW ZEALAND
Podocarpus totara type, Phyllocladus, Asteraceae type, Coprosma and Poaceae. Nothofagus forest is only 500 m to the east of the site today and contributes consistently about 20% of the pollen over the last 150 yr. Disturbance of the area by fire in 1890 is indicated by the presence of charcoal at the 26-28 cm level. A gradual increase in grass pollen after that time is due to the introduction of European grasses to the area (Calder and Wardle, 1969).
weather patterns over the past few seasons? Analyses of the pollen present in short peat cores should enable these questions to be answered.
( A ) Main Divide The peat core from the Main Divide (Fig.3) was taken from a bog dominated by Donatia novaezelandiae and Oreobolus pectinatus. It consisted of dark brown humified peat to 50 cm. The full depth of the bog was not determined. The core was too young to be radiocarbon dated, but using the occurrence of Pinus at 24 cm and Salix at 22 cm as indicators of European introduction of these species to the region (about 1870 A.D.), the bottom of the core was estimated at approximately 150 yr B.P. The results from the core show that there has been a steady influx of pollen of the various Westland tree species (Dacrydium cupressinum, Prumnopitys, Metrosideros and Quintinia) through the period of accumulation of the peat. Pollens which are mainly locally derived are
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(B) Bealey River Bealey River (Site H, Fig.l) is situated within the Nothofagus solandri var. cliffortioides forest. The core from this site (Fig.4) consists of uniform, dark-brown well humified, Donatia novae-zelandiae and Oreobolus pectinatus peat. As for the Main Divide core, the full depth of the bog was not determined. The 40cm level was dated at 482+60 yr B.P. (Libby T1/2 5568 yr) 496___61 yr B.P. (New TI/2 5730 yr) (NZ No 7534). Nothofagus
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Fig.3. Pollen diagram from Lower Bog (Site E) at Arthur's Pass, on the Main Divide of the Southern Alps. Only major pollen taxa shown.
268
P.M RANDALL
fusca type pollen dominates for the last 500 years, but there has been steady influx of Dacrydium cupressinum, Prumnopitys, Metrosideros and Quintinia pollen dispersed over the Main Divide from Westland. Both of these cores indicate that the European settlement (since the mid 1850s) has not greatly affected deposition of Dacrydium cupressinum and Prumnopitys pollen, although during this time there has been extensive clearance of forest to the west. The increased grass pollen percentage on the Main Divide site and the presence of Pinus, Salix and Rumex pollen are the only indicators of European influence.
(C) Cave Stream From Cave Stream (Site AD, Fig. 1) a 38 cm core was taken. The core is composed of light brown, humified unconsolidated peat with some sedge remains. The 102 cm of peat lies on alluvial gravel. The core was too young to be radiocarbon dated, but the presence of charcoal near the base
(22-38 cm) (Fig.5) and the occurrence of Pinus, Salix and Rumex pollen, suggests a maximum of about 80 yr B.P. The pollen profile (Fig.5) shows the removal of Nothofagus forest and scrubland by fire and their eventual replacement by grassland, which began with settlement of the area by European sheep farmers, beginning in 1857 (McLeod and Burrows, 1977).
Addition of exotic' spores The counts from samples with the Lycopodium marker were compared statistically with those without markers. The correlation coefficient for the two pollen counts was 0.97. This high correlation between the corresponding pairs of counts indicates a high level of repeatability in the samples. The results from the marker counts at five sites (Table I) indicate greater input of winddispersed podocarp pollen to the upland sites than to lowland sites in which the source trees grow. However, the other Westland species,
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Fig.4. Pollen diagram from Bealey River (Site H), Canterbury. Core taken from within the Nothoflagus solandri var. cliffortioides forest. Only major pollen taxa shown.
STUDY OF MODERN
269
POLLEN DEPOSITIONAL, NEW ZEALAND
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Fig.5. Pollen diagram from Cave Stream (Site AD), Canterbury. Core taken from a bog in an open grassland/scrubland habitat. Only major pollen taxa shown.
Metrosideros, Quintinia and Weinmannia, which are bird or insect pollinated and thus not well dispersed, do not show the same trend. Further interpretation of these absolute count values may be difficult as Boyd (1986) has shown in a similar study. Those sites on and near the Main Divide (C, E, F, G, H) show variability in the total pollen counts. Such differences seem likely to arise from differences in the effectiveness of various species of moss to act as pollen traps. At Otira, Racomitrium moss was collected, Sphagnum cristatum was collected at the Lower Bog and Dobson sites and Sphagnum falcatulum was collected at the Upper Bog and Bealey sites. The different length of time represented by each moss polster (Carroll, 1943) and the difference in the ratio of surface area to the weight or volume of the moss (Boyd, 1986) may be responsible for the variation shown in pollen counts (Table I). The percentage representation data (Table II) do not show any marked variability between the sites
although they do depict a general west to east trend in pollen dispersal. Discussion
In places of average wind flow (2-4 m/s), pollen and spores have little tendency to settle under the influence of gravity; they normally move in a horizontal direction as a result of wind currents (Tauber, 1965). Tauber proposed that pollen deposition consists of three components: pollen derived from (i) trunk space, (ii) above the canopy and from (iii) washout by rain. These three components give rise to a different pollen spectrum. In the Main Divide area, there is a strong westerly wind flow, with peaks in the spring and summer. The predominant westerlies are often accompanied by heavy precipitation (total annual average about 5000 mm; Burrows, 1968). Even when weather is fine to the east and west, cloud, fog and drizzle may prevail on the Main Divide
270
PM. RANDALL TABLE I The total number of pollen grains in one gram of dry weight of moss. Only major taxa s h o w n Main Divide Location Site Taxa
Otira C
Dacrydium cupressinum Podocarpus totara type Prumnopitys Halocarpus type Dacrvcarpus dacrydioides Lepidothamnus Pinus Phyllocladus Ascarina Metrosideros Nothofagusfusca type Quintinia Weinmannia Coprosma
Upper F
Dobson G
Bealey H
5,008 6,677 3,338 0 3,338 0 5,008 0 1,669 352,208 10,015 100,154 43,400 45,069 31,715 5,008 200,308 238,700
1,482 1,220 3,137 87 174 436 784 959 174 1,046 3,922 436 349 3,050 174 2,092 174 10,196
12,112 3,533 14,467 6,224 1,346 3,869 3,869 6,056 505 2,019 12,112 505 673 4,542 1,178 1,346 1,178 14,130
2,648 3,246 9,056 683 427 5,126 1.538 2,307 342 940 7,604 513 256 3,588 683 1,538 427 8.458
6,982 4,340 17,926 4,906 i, 132 9,246 1,698 4,717 189 1,510 47,740 943 566 3,774 189 755 566 4, ! 51
1,168,462
61,004
117,752
59,803
132,087
Apiaceae Asteraceae
Coriaria Poaceae Total pollen count
Lower E
T A B L E II Percentage representation data for the Main Divide sites. Only major taxa shown. Main Divide Location Site Taxa
Otira C
Lower E
Upper F
Dobson G
Bealey H
Dacrydium cupressinurn Podocarpus totara type Prumnopitys Halocarpus type Dacrycarpus dacrydioides Lepidothamnus Pinus Phyllocladus Ascarina Metrosideros Nothofagus fusca type Quintinia Weinmannia Coprosma
tr tr tr 0 tr
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3 tr
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1 1
1 3
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Coriaria
17
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Poaceae
21
33
15
16
4
1
271
STUDY OF MODERN POLLEN DEPOSITIONAL, NEW ZEALAND
(Coulter, 1973). The westerly wind tends to flow up the western valleys and is forced to rise at the Main Divide. Pollen and spores are completely washed out of the atmosphere with as little as 1 mm of rain, provided that the bulk of the raindrops are small in diameter (of the order of drizzle diameters, 0.2 mm) (McDonald, 1962). So podocarp pollen, which is released from trees in Westland during dry weather and blown eastward, will frequently encounter conditions suitable for washout over the Main Divide. The high values of Dacrydium cupressinum and Prumnopitys type pollen in the Main Divide sites corresponding to components (ii) and (iii) of Tauber (1965) may be accounted for in this way. The present study establishes some general principles about aerial pollen deposition across the Southern Alps. There is little evidence of podocarp pollen being deposited locally; instead it is carried by strong westerlies to the Main Divide where the high rainfall causes deposition o f large amounts of it. Is this feature peculiar to upper valleys such as Otira? The phenomenon of transport of podocarp pollen upslope has been noted by previous workers (Moar, 1970; McGlone, 1982; Pocknall, 1982). Pocknall (1982) presumed it to be due to low pollen production of the local sub-alpine vegetation but the present results suggest that a different interpretation is warranted. Low podocarp pollen count values are apparent for all four Westland valley sites. Other observations in Westland (Pock-
nail, 1980) show podocarp pollen being deposited locally, although low frequencies of Dacrydium cupressinum have also been observed at some sites. These apparently anomalous results were considered by Pocknall (1978) to be due to the effects of vegetation structure and dispersal of large amounts of pollen of other taxa in the vegetation, but it appears that wind carriage of pollen to distant sites is a normal feature for this species. The dispersal pattern of Nothofagus fusca type pollen is such that it is not possible to distinguish between a few trees near the sample site and many trees at a great distance. Nothofagus pollen appears to be dispersed from east to west but only in trace frequencies. This is probably due to the predominant north-west wind flow. Poaceae pollen dominates in the grassland sites, with only low frequencies in forested areas. This has been considered to be due to only small amounts of Poaceae pollen being transported into and through forested sites (Pocknall, 1978). This study suggests that Poaceae pollen may not be well dispersed.
Acknowledgements I am indebted to Colin Burrows and Dave Kelly for useful suggestions on the MS and to Matt McGlone and Neville Moar of Botany Division, DSIR for assistance in identification of pollen and compiling the pollen diagrams.
Appendix 1 Site
Location
Grid Ref.
Altitude (metres)
A B C D E F G H I J K L M
Aickens BarrackCreek Otira Gorge RollestonRiver Lower Bog Arthur's Pass Upper Bog Arthur's Pass DobsonMemorial Bealey River Quarry Knob Bog Quarry Knob WoolshedHill Bog Lower Woolshed Bog HorribleBog (N)
098470 455 056394 520 054392 520 043381 520 052327 915 053326 920 054323 920 050303 825 088217 730 090216 740 230235 1055 223234 670 205190 640
Generaldescription of site Grassland bordered by podocarp/broadleaf forest River terrace bordered by podocarp/broadleaf forest Grassland/scrubland bordered by podocarp/broadleaf forest Scrubland bordered by podocarp/broadleaf forest Subalpine scrub, grassland and herbfield Subalpine scrub, grassland and herbfield Subalpine scrub, grassland and herbfield Nothofagus solandrivat. cliffortioides forest Nothofagus solandrivar. cliffortioides forest Nothofagus solandrivat. cliffortioidesforest Nothofagus solandrivar. cliffortioidesforest Nothofagus solandrivar. cliffortioidesforest Grassland/scrubland
272
P.M. R A N D A L l
Appendix 1 (cont.) N O P Q R S T U V W X Y Z AA AB AC AD AE AF AG AH AI
Horrible Bog Waterfall Terrace Cass Misery Swamp Lake Sarah S.E. Lake Sarah Kettlehole Bog Waimakariri St Bernard Ribbonwood Stream Lake Hawdon Lake Hawdon Bog Slovens Stream (N) Vagabonds Inn Slovens Stream Craigieburn Cutting Cave Stream Broken River Castle Hill Willow Tree Bog Lake Lyndon Porters Pass
205184 208172 230171 208162 244157 251146 254147 271147 264137 244115 308091 311090 328051 319038 329032 213028 207022 208974 205959 185881 207853 216851
640 640 550 610 580 580 610 485 610 580 580 580 550 610 550 915 790 700 700 825 9l 5 930
Grasstand/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland Grassland/scrubland Scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland
Maps used: Sites A-L NZMS 1 $59 Otira 1:63350 4th Edition 1985 Sites M AF NZMS 1 $66 Broken River 1:63350 3rd Edition 1980 Sites AG AI NZMS t $74 Springfield 1:63350 6th Edition 1986
References Boyd, W.E., 1986. The roles of mosses in modern pollen analysis. The influence of moss morphology on pollen entrapment. Pollen Spores, 28: 243-256. Burrows, C.J,, 1968. The ecology of some alpine grassland. Thesis, Univ. Canterbury (unpubl.). Calder, J.D. and Wardle, P., 1969. Succession in subalpine vegetation at Arthurs Pass, New Zealand. Proc. N.Z. Ecol. Soc., 16: 36-47. Carroll, G., 1943. The use of bryophytic polsters and mats in the study of recent pollen deposition. Am. J. Bot., 30:361 366. Coulter, J.D., 1973. Ecological aspects of the climate. In G.R. Williams (Editor). The Natural History of New Zealand. Reed. Wellington, pp.28-60. Crowder A.A. and Cuddy, D.G., 1973. Pollen in a small river basin: Wilton Creek, Ontario. In: H.J.B. Birks and R.G. West (Editors), Quaternary Plant Ecology, Blackwell Scientific Publications, Oxford, pp.61 77. Faegri, K. and Iversen, J., 1964. Textbook of Pollen Analysis. 2nd ed. Munksgaard, Copenhagen, 237 pp. McDonald, J.E., 1962. Collection and washout of airborne pollen and spores by raindrops. Science, 135:435 436. McGlone, M.S., 1982. Modern pollen rain, Egmont National Park, New Zealand. N.Z.J. Bot., 20:253 262. McLeod, D. and Burrows, C.J., 1977. History of the Cass
District. In: C.J. Burrows (Editor), Cass, History and Science in the Cass District, Canterbury, New Zealand. Univ. Canterbury, 418 pp. Moar, N.T, 1970. Recent pollen spectra from three localities in the South Island, New Zealand. N.Z.J. Bot., 8:210 221. Moar, N.T., 1971. Contributions to the Quaternary history of the New Zealand flora 6. Aranuian pollen diagrams from Canterbury, Nelson, and North Westland, South Island. N.Z.J. Bot., 9:80 145. Pocknall, D.T., 1978. Relative pollen representation in relation to vegetation composition, Westland, New Zealand. N.Z.J. Bot., 16:379 386. Pocknall, D.T., 1980. Modern pollen rain and Aranuian vegetation from Lady Lake. north Westland, New Zealand. N.Z.J. Bot.. 18: 275-284. Pocknall, D.T., 1982. Modern pollen spectra from mountain localities, South Island, New Zealand. N . Z . J . Bot.. 20: 361 371. Stockmarr, J., 1971. Tablets with spores used in absolute pollen analysis. Pollen Spores, 13:615 621. Tauber, H., 1965. Differential pollen dispersion and the interpretation of pollen diagrams. Geol. Surv. Denmark II, ser. 89, 69 pp. Wardle, J., 1970. The ecology of Nothojagus solandri. 1. The distribution and relationship with other major forest and scrub species. N.Z.J. Bot., 8: 494-531.
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Elsevier Science Publishers B.V., Amsterdam
A study of modern pollen deposition, Southern Alps, South Island, New Zealand P.M. R a n d a l l Department of Plant and Microbial Sciences, University of Canterbury, Christchurch (New Zealand) (Received October 11, 1988; revised and accepted January 25, 1990)
ABSTRACT Randall, P.M., 1990. A study of modern pollen deposition, Southern Alps, South Island, New Zealand. Rev. Palaeobot. Palynol., 64: 263-272. The pollen and spores dispersing from natural and semi-natural vegetation in the South Island of New Zealand were studied from moss polsters collected from 35 sites along a transect from Westland across the Southern Alps to Canterbury. Sites in Westland were dominated by local broadleaved angiosperm species (Metrosideros,Quintinia, Weinmannia)and Poaceae pollen. Sites above the treeline on the Main Divide showed anomalous high counts of exogenous Podocarpaceae pollen. This seems to be washed out of strong NW winds by orograpbic precipitation. Peat cores taken from 3 sites confirm that exogenous podocarp pollen influx has continued over at least the last 500 years. Nothofaguspollen dominates within the beech forest areas, but drops to 10% a short distance from the forest edge. Poaceae pollen shows low frequencies in forested sites, but dominates in grassland areas.
Introduction
Study area
Prior to the w o r k of M o a r (1970, 1971), Pocknall (1978, 1980, 1982) and M c G l o n e (1982), pollen analysts in New Zealand lacked detailed information on the relationship between modern pollen deposition and the vegetation from which the pollen originated. A s u m m a r y of pollen deposition studies up to 1981 is outlined by Pocknall (1982). Most such work in New Zealand has attempted to discover information a b o u t relative dispersal and p r o p o r t i o n a t e representation of pollen o f the main taxa present in Quaternary sites. The aim o f the present study is to amplify information a b o u t the m o d e r n pollen deposition in order to assist interpretation of Quaternary pollen diagrams. M a n y o f the sites suitable for Quaternary pollen analysis are in mountain localities in the Southern Alps, where pollen accumulates and is preserved in Sphagnum bogs or other wet sites.
The study area contains a complex system of mountain ranges, valleys and basins with peaks rising to 2270 m and valley floors at 460 to 610 m. Arthur's Pass, on the Main Divide, which forms the boundary between Westland and Canterbury, is at an elevation of 920 m. The mixed forest of the Taramakau and Otira valleys in Westland contains substantial amounts of the podocarp tree species Dacrydium
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cupressinum, Podocarpus hallii, Prumnopitys ferruginea, P. taxifolia and Dacrycarpus dacrydioides but is dominated by angiosperm species Weinmannia racemosa, Metrosideros umbellata, smaller amounts of Quintinia acutifolia, and many other angiosperm species. Nothofagusfusca is present in the Otehake and Taramakau river catchment area with some Nothofagus solandri var. cliffortioides and N. menziesii. Scattered stands of N.fusca occur on young river terraces of the lower Otira river and a small stand of N. menziesii in the lower Otira gorge.
© 1990- Elsevier Science Publishers B.V.
264
There is a steep rise from the Westland valleys to the Main Divide where the vegetation is predominantly subalpine scrub, grassland and herbfield (Calder and Wardle, 1969). At the Canterbury side there is a gradual drop into the Bealey River valley in which Nothoj~tgus solandri var. cliffortioides dominates. It is accompanied by N.fusca and N. menziesii in the upper Poulter River and N. fusca near the Bealey river confluence with the Waimakariri and in the Hawdon and Andrews river catchment (Wardle, 1970). Further east, in the Waimakariri basin, the forest is discontinuous. Extensive grassland and scrubland dominates, with patches of Nothofagus solandri var. cliffortioides on the Craigieburn and Torlesse Ranges. The main vegetation in the Porter Pass area is mostly Dracophyllum scrub and grassland with only isolated small patches of Nothofagus solandri var.
cliffortioides. Precipitation is high in Westland ( > 5000 mm per annum), diminishing eastward (c. 1750 mm at the Bealey confluence, 1000 mm at Castle Hill). In the mountains strong westerly winds predominate with peaks in spring and summer (Coulter, 1973). Methods
Moss polsters were collected at 35 sites on a transect from Westland to Canterbury across the Main Divide (Fig. 1 and Appendix 1). At least four to six subsamples from each site were combined into one sample. To reduce the possibility of differential filtration of pollen by the moss polster, the whole thickness of living moss was removed (Crowder and Cuddy, 1973). Moss polsters are believed to yield pollen spectra representative of t h e pollen deposition for one to several years. Samples were collected where possible from open bogs and not from beneath forest canopy. The samples were transported to the laboratory in sealed plastic bags. The samples for pollen analysis were boiled in 5% sodium hydroxide, sieved and treated with hot
P.M RANDALL
40% hydrofluoric acid to remove silicates, followed by standard acetolysis (Faegri and Iversen, 1964), before mounting in glycerine jelly. The pollen and spore counts were carried out at a magnification of x 400 and 700 pollen grains and spores were counted per sample along regularly spaced traverses of the slide. To obtain additional information regarding the deposition of pollen on sites at Arthur's Pass on the Main Divide, a spore marker was added to five surface samples to determine the absolute numbers of pollen and spores present. The method of Stockmarr (1971) was used. Two Lycopodium tablets containing 10,850_+200 spores per tablet were added to one gram of dry material and processed as above. The analysis was done in the normal way, counting to 700 pollen and spores of all types, plus the number of marker Lycopodium. The pollen sum, on which the percentage calculations are based, consists of all terrestrial taxa excluding those, such as sedges, that are mainly restricted to bogs and wet sites. All fern spores were also excluded from the pollen sum. Three peat monoliths (38 to 42 cm deep) were sampled to establish the immediate past history of the vegetation in the region. The fossil pollen spectra were then compared with the surface samples paying particular attention to the pollen markers of modern times (Pinus and Salix) as well as prominent native species (Dacrydium cupressinum, Prumnopitys and Nothofagus fusca type). The record of European settlement in the area is well documented. Using the first occurrence of pollen of indicator species (Pinus and Salix) as well as charcoal, approximate dates can be fixed for the peat cores. Samples were collected using 50 cm PVC tubes of 20cm diameter, taken to the laboratory and sectioned at 2 c m intervals for pollen analysis. Subsamples were taken at the middle and bottom of the core for radiocarbon analysis. Radiocarbon samples were pretreated by the removal of roots which had grown down into the
Fig. 1. Location of the 35 sites for surfacesamplescollectedalong a transect from Westland to Canterbury across the Main Divide and the two main forest typesin the study area. Podoearp/broadleafforest (light shading) and beechforest (dark shading), and grassland or subalpine (clear areas). For details of sites see Appendix 1. Based on the NZFS Forest class maps series 6 No. 18, 19, 20 (1974).
265
STUDY OF MODERN POLLEN DEPOSITIONAL, NEW ZEALAND
SOUTH ISLAND
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at sites E-G. Nothofagus solandri var. cliffortioides forest is present nearby. (3) Nothofagus fusca type (which includes both N.fusca and N. solandri var. cliffortioides) dominates at sites H-L, all surrounded by beech forest. (4) At all other sites to the east, grasses are dominant. Grass pollen is also dominant at the westernmost site (site A) which is surrounded by pasture, and abundant at the Main Divide sites, but unimportant at sites closely surrounded by beech forest (e.g. sites H-L). Nothofagusfusca type values, high in forest sites, fall to less than 10% in sites mainly surrounded by grassland and scrub. Exceptions are minor Nothofagus pollen peaks where samples were taken near small pockets of beech forest.
peat. This was done to reduce the possibility of contamination by the presence of younger carbon. The samples were then dried and forwarded to New Zealand Radiocarbon Dating Laboratory, Institute of Nuclear Sciences, DSIR, Lower Hutt for age determination. Pollen diagrams
Surface samples The results of counts from the surface samples are shown in Fig.2. The outstanding features are: (1) At sites B-E, broadleaved angiosperm counts are high (predominantly Metrosideros 320/0, with Quintinia 8%, and Weinmannia 4%) and Nothofagusfusca type and Podocarpaceae are very low. (2) Pollen of Podocarpaceae predominates (Dacrydium cupressinum, Prumnopitys and Dacrycarpus dacrydioides) at sites E - G on the Main Divide, where these trees are absent (they are virtually confined to sites west of the Main Divide). Nothofagusfusca type pollen is also very abundant
Peat Cores Is the array of pollen spectra described above from modern surface samples reflected through time, or is it strictly a modern phenomenon due to either (i) the length of time represented by the pollen deposited in the moss polsters or (ii) unusual
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Fig.2. Pollen diagrams from sites of the West-East transect, from Westland (Site A) across the Main Divide (Sites E-G) to Porters Pass (Site AI). The Podocarpaceae pollen (Dacrycarpusdacrydioides, Dacrydium cupressinum, Prumnopitys) at the Main Divide (Sites E-G) is not local (see text). Only major pollen taxa shown.
267
STUDY OF MODERN POLLEN DEPOSITIONAL, NEW ZEALAND
Podocarpus totara type, Phyllocladus, Asteraceae type, Coprosma and Poaceae. Nothofagus forest is only 500 m to the east of the site today and contributes consistently about 20% of the pollen over the last 150 yr. Disturbance of the area by fire in 1890 is indicated by the presence of charcoal at the 26-28 cm level. A gradual increase in grass pollen after that time is due to the introduction of European grasses to the area (Calder and Wardle, 1969).
weather patterns over the past few seasons? Analyses of the pollen present in short peat cores should enable these questions to be answered.
( A ) Main Divide The peat core from the Main Divide (Fig.3) was taken from a bog dominated by Donatia novaezelandiae and Oreobolus pectinatus. It consisted of dark brown humified peat to 50 cm. The full depth of the bog was not determined. The core was too young to be radiocarbon dated, but using the occurrence of Pinus at 24 cm and Salix at 22 cm as indicators of European introduction of these species to the region (about 1870 A.D.), the bottom of the core was estimated at approximately 150 yr B.P. The results from the core show that there has been a steady influx of pollen of the various Westland tree species (Dacrydium cupressinum, Prumnopitys, Metrosideros and Quintinia) through the period of accumulation of the peat. Pollens which are mainly locally derived are
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268
P.M RANDALL
fusca type pollen dominates for the last 500 years, but there has been steady influx of Dacrydium cupressinum, Prumnopitys, Metrosideros and Quintinia pollen dispersed over the Main Divide from Westland. Both of these cores indicate that the European settlement (since the mid 1850s) has not greatly affected deposition of Dacrydium cupressinum and Prumnopitys pollen, although during this time there has been extensive clearance of forest to the west. The increased grass pollen percentage on the Main Divide site and the presence of Pinus, Salix and Rumex pollen are the only indicators of European influence.
(C) Cave Stream From Cave Stream (Site AD, Fig. 1) a 38 cm core was taken. The core is composed of light brown, humified unconsolidated peat with some sedge remains. The 102 cm of peat lies on alluvial gravel. The core was too young to be radiocarbon dated, but the presence of charcoal near the base
(22-38 cm) (Fig.5) and the occurrence of Pinus, Salix and Rumex pollen, suggests a maximum of about 80 yr B.P. The pollen profile (Fig.5) shows the removal of Nothofagus forest and scrubland by fire and their eventual replacement by grassland, which began with settlement of the area by European sheep farmers, beginning in 1857 (McLeod and Burrows, 1977).
Addition of exotic' spores The counts from samples with the Lycopodium marker were compared statistically with those without markers. The correlation coefficient for the two pollen counts was 0.97. This high correlation between the corresponding pairs of counts indicates a high level of repeatability in the samples. The results from the marker counts at five sites (Table I) indicate greater input of winddispersed podocarp pollen to the upland sites than to lowland sites in which the source trees grow. However, the other Westland species,
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Fig.4. Pollen diagram from Bealey River (Site H), Canterbury. Core taken from within the Nothoflagus solandri var. cliffortioides forest. Only major pollen taxa shown.
STUDY OF MODERN
269
POLLEN DEPOSITIONAL, NEW ZEALAND
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Metrosideros, Quintinia and Weinmannia, which are bird or insect pollinated and thus not well dispersed, do not show the same trend. Further interpretation of these absolute count values may be difficult as Boyd (1986) has shown in a similar study. Those sites on and near the Main Divide (C, E, F, G, H) show variability in the total pollen counts. Such differences seem likely to arise from differences in the effectiveness of various species of moss to act as pollen traps. At Otira, Racomitrium moss was collected, Sphagnum cristatum was collected at the Lower Bog and Dobson sites and Sphagnum falcatulum was collected at the Upper Bog and Bealey sites. The different length of time represented by each moss polster (Carroll, 1943) and the difference in the ratio of surface area to the weight or volume of the moss (Boyd, 1986) may be responsible for the variation shown in pollen counts (Table I). The percentage representation data (Table II) do not show any marked variability between the sites
although they do depict a general west to east trend in pollen dispersal. Discussion
In places of average wind flow (2-4 m/s), pollen and spores have little tendency to settle under the influence of gravity; they normally move in a horizontal direction as a result of wind currents (Tauber, 1965). Tauber proposed that pollen deposition consists of three components: pollen derived from (i) trunk space, (ii) above the canopy and from (iii) washout by rain. These three components give rise to a different pollen spectrum. In the Main Divide area, there is a strong westerly wind flow, with peaks in the spring and summer. The predominant westerlies are often accompanied by heavy precipitation (total annual average about 5000 mm; Burrows, 1968). Even when weather is fine to the east and west, cloud, fog and drizzle may prevail on the Main Divide
270
PM. RANDALL TABLE I The total number of pollen grains in one gram of dry weight of moss. Only major taxa s h o w n Main Divide Location Site Taxa
Otira C
Dacrydium cupressinum Podocarpus totara type Prumnopitys Halocarpus type Dacrvcarpus dacrydioides Lepidothamnus Pinus Phyllocladus Ascarina Metrosideros Nothofagusfusca type Quintinia Weinmannia Coprosma
Upper F
Dobson G
Bealey H
5,008 6,677 3,338 0 3,338 0 5,008 0 1,669 352,208 10,015 100,154 43,400 45,069 31,715 5,008 200,308 238,700
1,482 1,220 3,137 87 174 436 784 959 174 1,046 3,922 436 349 3,050 174 2,092 174 10,196
12,112 3,533 14,467 6,224 1,346 3,869 3,869 6,056 505 2,019 12,112 505 673 4,542 1,178 1,346 1,178 14,130
2,648 3,246 9,056 683 427 5,126 1.538 2,307 342 940 7,604 513 256 3,588 683 1,538 427 8.458
6,982 4,340 17,926 4,906 i, 132 9,246 1,698 4,717 189 1,510 47,740 943 566 3,774 189 755 566 4, ! 51
1,168,462
61,004
117,752
59,803
132,087
Apiaceae Asteraceae
Coriaria Poaceae Total pollen count
Lower E
T A B L E II Percentage representation data for the Main Divide sites. Only major taxa shown. Main Divide Location Site Taxa
Otira C
Lower E
Upper F
Dobson G
Bealey H
Dacrydium cupressinurn Podocarpus totara type Prumnopitys Halocarpus type Dacrycarpus dacrydioides Lepidothamnus Pinus Phyllocladus Ascarina Metrosideros Nothofagus fusca type Quintinia Weinmannia Coprosma
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1 1
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2
Coriaria
17
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21
33
15
16
4
1
271
STUDY OF MODERN POLLEN DEPOSITIONAL, NEW ZEALAND
(Coulter, 1973). The westerly wind tends to flow up the western valleys and is forced to rise at the Main Divide. Pollen and spores are completely washed out of the atmosphere with as little as 1 mm of rain, provided that the bulk of the raindrops are small in diameter (of the order of drizzle diameters, 0.2 mm) (McDonald, 1962). So podocarp pollen, which is released from trees in Westland during dry weather and blown eastward, will frequently encounter conditions suitable for washout over the Main Divide. The high values of Dacrydium cupressinum and Prumnopitys type pollen in the Main Divide sites corresponding to components (ii) and (iii) of Tauber (1965) may be accounted for in this way. The present study establishes some general principles about aerial pollen deposition across the Southern Alps. There is little evidence of podocarp pollen being deposited locally; instead it is carried by strong westerlies to the Main Divide where the high rainfall causes deposition o f large amounts of it. Is this feature peculiar to upper valleys such as Otira? The phenomenon of transport of podocarp pollen upslope has been noted by previous workers (Moar, 1970; McGlone, 1982; Pocknall, 1982). Pocknall (1982) presumed it to be due to low pollen production of the local sub-alpine vegetation but the present results suggest that a different interpretation is warranted. Low podocarp pollen count values are apparent for all four Westland valley sites. Other observations in Westland (Pock-
nail, 1980) show podocarp pollen being deposited locally, although low frequencies of Dacrydium cupressinum have also been observed at some sites. These apparently anomalous results were considered by Pocknall (1978) to be due to the effects of vegetation structure and dispersal of large amounts of pollen of other taxa in the vegetation, but it appears that wind carriage of pollen to distant sites is a normal feature for this species. The dispersal pattern of Nothofagus fusca type pollen is such that it is not possible to distinguish between a few trees near the sample site and many trees at a great distance. Nothofagus pollen appears to be dispersed from east to west but only in trace frequencies. This is probably due to the predominant north-west wind flow. Poaceae pollen dominates in the grassland sites, with only low frequencies in forested areas. This has been considered to be due to only small amounts of Poaceae pollen being transported into and through forested sites (Pocknall, 1978). This study suggests that Poaceae pollen may not be well dispersed.
Acknowledgements I am indebted to Colin Burrows and Dave Kelly for useful suggestions on the MS and to Matt McGlone and Neville Moar of Botany Division, DSIR for assistance in identification of pollen and compiling the pollen diagrams.
Appendix 1 Site
Location
Grid Ref.
Altitude (metres)
A B C D E F G H I J K L M
Aickens BarrackCreek Otira Gorge RollestonRiver Lower Bog Arthur's Pass Upper Bog Arthur's Pass DobsonMemorial Bealey River Quarry Knob Bog Quarry Knob WoolshedHill Bog Lower Woolshed Bog HorribleBog (N)
098470 455 056394 520 054392 520 043381 520 052327 915 053326 920 054323 920 050303 825 088217 730 090216 740 230235 1055 223234 670 205190 640
Generaldescription of site Grassland bordered by podocarp/broadleaf forest River terrace bordered by podocarp/broadleaf forest Grassland/scrubland bordered by podocarp/broadleaf forest Scrubland bordered by podocarp/broadleaf forest Subalpine scrub, grassland and herbfield Subalpine scrub, grassland and herbfield Subalpine scrub, grassland and herbfield Nothofagus solandrivat. cliffortioides forest Nothofagus solandrivar. cliffortioides forest Nothofagus solandrivat. cliffortioidesforest Nothofagus solandrivar. cliffortioidesforest Nothofagus solandrivar. cliffortioidesforest Grassland/scrubland
272
P.M. R A N D A L l
Appendix 1 (cont.) N O P Q R S T U V W X Y Z AA AB AC AD AE AF AG AH AI
Horrible Bog Waterfall Terrace Cass Misery Swamp Lake Sarah S.E. Lake Sarah Kettlehole Bog Waimakariri St Bernard Ribbonwood Stream Lake Hawdon Lake Hawdon Bog Slovens Stream (N) Vagabonds Inn Slovens Stream Craigieburn Cutting Cave Stream Broken River Castle Hill Willow Tree Bog Lake Lyndon Porters Pass
205184 208172 230171 208162 244157 251146 254147 271147 264137 244115 308091 311090 328051 319038 329032 213028 207022 208974 205959 185881 207853 216851
640 640 550 610 580 580 610 485 610 580 580 580 550 610 550 915 790 700 700 825 9l 5 930
Grasstand/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland Grassland/scrubland Scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland Grassland/scrubland
Maps used: Sites A-L NZMS 1 $59 Otira 1:63350 4th Edition 1985 Sites M AF NZMS 1 $66 Broken River 1:63350 3rd Edition 1980 Sites AG AI NZMS t $74 Springfield 1:63350 6th Edition 1986
References Boyd, W.E., 1986. The roles of mosses in modern pollen analysis. The influence of moss morphology on pollen entrapment. Pollen Spores, 28: 243-256. Burrows, C.J,, 1968. The ecology of some alpine grassland. Thesis, Univ. Canterbury (unpubl.). Calder, J.D. and Wardle, P., 1969. Succession in subalpine vegetation at Arthurs Pass, New Zealand. Proc. N.Z. Ecol. Soc., 16: 36-47. Carroll, G., 1943. The use of bryophytic polsters and mats in the study of recent pollen deposition. Am. J. Bot., 30:361 366. Coulter, J.D., 1973. Ecological aspects of the climate. In G.R. Williams (Editor). The Natural History of New Zealand. Reed. Wellington, pp.28-60. Crowder A.A. and Cuddy, D.G., 1973. Pollen in a small river basin: Wilton Creek, Ontario. In: H.J.B. Birks and R.G. West (Editors), Quaternary Plant Ecology, Blackwell Scientific Publications, Oxford, pp.61 77. Faegri, K. and Iversen, J., 1964. Textbook of Pollen Analysis. 2nd ed. Munksgaard, Copenhagen, 237 pp. McDonald, J.E., 1962. Collection and washout of airborne pollen and spores by raindrops. Science, 135:435 436. McGlone, M.S., 1982. Modern pollen rain, Egmont National Park, New Zealand. N.Z.J. Bot., 20:253 262. McLeod, D. and Burrows, C.J., 1977. History of the Cass
District. In: C.J. Burrows (Editor), Cass, History and Science in the Cass District, Canterbury, New Zealand. Univ. Canterbury, 418 pp. Moar, N.T, 1970. Recent pollen spectra from three localities in the South Island, New Zealand. N.Z.J. Bot., 8:210 221. Moar, N.T., 1971. Contributions to the Quaternary history of the New Zealand flora 6. Aranuian pollen diagrams from Canterbury, Nelson, and North Westland, South Island. N.Z.J. Bot., 9:80 145. Pocknall, D.T., 1978. Relative pollen representation in relation to vegetation composition, Westland, New Zealand. N.Z.J. Bot., 16:379 386. Pocknall, D.T., 1980. Modern pollen rain and Aranuian vegetation from Lady Lake. north Westland, New Zealand. N.Z.J. Bot.. 18: 275-284. Pocknall, D.T., 1982. Modern pollen spectra from mountain localities, South Island, New Zealand. N . Z . J . Bot.. 20: 361 371. Stockmarr, J., 1971. Tablets with spores used in absolute pollen analysis. Pollen Spores, 13:615 621. Tauber, H., 1965. Differential pollen dispersion and the interpretation of pollen diagrams. Geol. Surv. Denmark II, ser. 89, 69 pp. Wardle, J., 1970. The ecology of Nothojagus solandri. 1. The distribution and relationship with other major forest and scrub species. N.Z.J. Bot., 8: 494-531.