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Vegetation of the Aorangirange,Southern WairarapaJ. Wardle aa Forest and Range Experiment Station ,Forest Research Institute , RangioraPublished online: 10 Feb 2012.

To cite this article: J. Wardle (1967) Vegetation of the Aorangirange,Southern Wairarapa, New Zealand Journal of Botany, 5:1, 22-48, DOI:10.1080/0028825X.1967.10428733

To link to this article: http://dx.doi.org/10.1080/0028825X.1967.10428733

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22 [MARCH

VEGETATION OF THE AORANGI RANGE, SOUTHERN WAIRARAPA

1. WAROJ.E

Forest and Range Experiment Station, Forest Research Institute. Rangiora

(Received for publiclIlion 19 luly 19M)

SUMMARY

In primeval times the Aorangi Range supported an almost continuous forest cover broken only by occasional slips and young stream terraces. Forest still forlT15 the mo.,t extensive cover in the area. It can be broadly classified into four main types: mahoe-hinau-rewarcwa forest en the more stable slopes and flats up to an altitude of 1,700-1,900 ft; black b(;ech forest on the drier, more exposed spur sites up to 1,600 ft; red beech forcst on the mid slopes between 1,400 ft and 1,900 ft; and silver beech forc.~t along the ridges between 2,000 ft and the highest points of the range.

In recent times the vegetation of the area has been modified by fire, browsing, and erosion. Erosion has favoured the extension of seral plant communities such as are found on slips, in gully heads, and on aggrading river terraces. Burning has given rise to tall kanuka scrub at lower altitudes and to subalpine scrub and tussock grassland at higher altitudes. Browsillg mammals have considerably altered the composition of cach forest type and have modified Succc.~sion on bared areas.

The first section of this paper hriefly describes the physical nature of the range; the second deals with the distribution, structure, and composition of the main forest types, and the third descrihes how erosion. fire. and hrowsing have altered these to produce the present seral and suhclimax communities.

I NTRODII('TION

The greater part of the Aorangi Range (commonly referred to as the "Haurangis") lies within permanent State forest. The total area of State forest is approximately 34.000 acres. divided into a northern and a southern section by a strip of hilly grazing country lying to the true left of Turanganui River (Fig. I).

The range is comparatively low. the two highest peaks being Mt Ross 0.226 ft) in the southern section. and Mt Te Munga (3.211 ft) in the northern section. It constitutes a major block moving between two trans­current fault zones which trend in a NE-SW direction. The present sharp ridges of the range resulted from extensive southward movement of the block during the Tertiary period (Wellman. 1956; Kingma. 1959). This

N.Z. JI BOI. 5: 22-4H.

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1967]

North ISland N Z.

J. I,U

WARDLE - AORANGI RANGE

AORANGI

3

Fig. 1

'LEGEND'

Forest Margin

Scrub

Trig Station

MOUNTAIN RANGE

Scal. of MIl ..

o 3 G

FIG. I-Locality map showing the main stream systems. The forest and scrub margins have been modified from 1he NSMI. N 165 and N 168. However. these margins arc not striclly realistic (See Fig. 3).

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24 NEW ZEALAND JOURNAL OF BOTANY [MARCH

movement caused the development of a series of minor faults and consequent large-scale internal deformation. Many of these minor faults are still active (Fig. 2). The drainage pattern of the range is largely influenced by the pattern of fault lines and shatter zones. and is of a somewhat trellised nature. As a result downcutting by the streams is rapid and the valleys are typically V -shaped and steep walled. with numerous gorges.

FIG. 2-Recent tectonic activity . Pararaki valley. Note tilted trees and drawn roots along line of fault.

The parent rock is predominantly grey wacke. dating from the Aptian and Neocomian stages of the Cretaceous (Grindley. Harrington. and Wood, 1959). Minor volcanics. probably of Jurassic age (Wellman, (956), border the south-western margin. and towards the north-east there are pockets of limestone resulting from movement during the Kaikoura diastrophism (Kingma. 1959). The Aorangi Range. and the mountain ranges to the north-east. are a continuation of the ranges of the eastern Marlborough region, and are composed of similar rocks (Wellman. 1956).

There are few climatic records available for the Aorangi Range. Climatic maps of New Zealand show the area as having between 40 and 60 in. mean annual rainfall. This is probably true for the lower elevations. but rainfall increases rapidly with gain in altitude and probably exceeds 100 in. at 3,000 ft (Wairarapa Catchment Board. pers. comm.).

The range is exposed to strong winds from all directions. The prevailing winds are from the north-west, but southerlies are also common. These southerly winds are frequently .of high velocity. and. in contrast with the drier north-westerlies, are heaVily laden with moisture.

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1967] WARDLE - AORANGl RANGE 25

Snow is infrequent and the little that falls does not lie for long enough to have much effect on the vegetation. The incidence of fog and cloud is low. Information on frosts and temperature patterns is not available.

The material presented in this paper was mainly gathered during a survey of the range carried out by the Napier branch of the Forest and Range Experiment Station in January and February 1963. During the survey 36 permanent vegetation plots were established. The composition of the main vegetation types has been determined from information yielded by these plots and from allied observations.

DISTRIBUTION, STRUCTURE, AND COMPOSITION OF THE MAIN VEGETATION TYPES·

Distribution and composition of the vegetation typ~s in the Aorangi Range were originally determined largely by natural site characteristics such as altitude, aspect, and slope. The area has, however, been subjected to modifying influences which have caused departure from the original condition and given rise to an extremely complex vegetation pattern (Fig. 3). The plant communities may be classified into two major groups -those that have not yet been modified beyond recognition, and those that are at a sera I or subclimax stage. The former are discussed in this section; the main modifying influences and the way in which they have affected the original vegetation types are described in the following section. .

The unit of vegetation employed in the descriptions is the "type", here defined as a plant community characterised by certain physiognomically prominent species. Consistency in classifying an area where the vegetation pattern is so complex requires that the types be broad and that many minor variations be ignored. A certain amount of detail will thus be lost. The following vegetation types are defined in terms of their physiognomic dominants. Each is described as it is at present; and although the canopy layer in each probably retains its original composition, it must be borne in mind that the subcanopy layers will usually have been modified by browsing mammals. Detailed data on the composition of the types are given in the appendix.

MAHOE-HINAlJ-REWAREWA FOREST

Mahoe (Melicytus ram;florus)-hinau (£Iaeocarpus dentatus)-rewa­rewa (Kn;ght;a excelsa) forest is the most complex and probably the

• Botanical names used in this paper are according to Zotov (1963) for the grass sub-family Arundinoidt'ae; Cheeseman (1925) for the remainder of the indigenous Monocotyledones; Philipson (1965) for the genera of the Araliaceae; Allan (1940) for the introdl:ced species and Allan (1961) for all remaining species.

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FIG. 3-Aerial photographs of portion of the Aorangi Range from the Waitutuma catchment eastwards to >the coast. These photographs give an indication of

most extensive forest type in the Aorangi Range. It occupies the more stable slopes and flats up to an altitude of 1.700-1,900 ft. Its composition varies with changes in site characteristics. On the younger soils and on moister soils mahoe assumes local dominance and may form a forest little more than 20-30 f1 high. but on the better drained and deeper soils hinau becomes abundant and usually forms an overstorey to the mahoe. Rewarewa is present throughout as a scattered emergent.

To the north of Otakaha Stream podocarp species occasionally OC{;ur. overtopping the mahoe. hinau. and rewarewa. They are more common

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the extent (0 which the original forest has been modified by fire and erosion. (Photo ref.: 266/8 and 266/10. North at top.)

on the shadier. moister, southerly and easterly aspects than on the more exposed, northerly and westerly aspects. These podocarps, which include rimu (Dacrydium cupressinum). matai (Podocarpus spicatus). and miro (Podocarpus ferrugineus), with rimu the most abundant species, may reach heights in excess of 80 ft on the lower. sheltered slopes and on old river terraces. but with increase in altitude and in exposure their height becomes much reduced.

The understorey of mahoe-hinau-rewarewa forest varies from site to s~te. but pigeon wood (Hedycarya arhorea) occurs regardless of site

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28 NEW ZEALAND JOlJRNAL OF BOTANY [MARCH

variation. Silver treefern (Cyathea dealhata) generally forms a dense understorey on the moister, shadier slopes. Kawakawa (Macropiper excelsum) often occurs as a band at about 1,000 ft, and Coprosma rhamnoides, Cyathodes fasciculata, and Helichrysum glomeratum are often present on the drier ridges. Toro (Myrsine salicina) is usually confined to altitudes above 1,500 ft.

The composition of the forest-floor cover varies with changes in canopy density. Where the canopy is fairly open and drainage is good, a dense cover of Uncinia spp., Pel/aea rotundifo/ia, and Microlaena avenacea is usually in evidence (Figs. 4 and 5). Water fern (Histiopteri.l' incisa) is found in moister sites and seepage spots. Where the canopy is denser the forest floor is often bare or only sparsely vegetated with small ferns and herbs, prominent among which are Asplenium hookerianum and Ranunculus hirtus.

The composition of the forest is not greatly influenced by the presence of podocarp species. Silver treefern and pigeonwood are still the main understorey species, but the variety of small trees, shrubs, and herbs is usually greater---especially the variety of herbs. Epiphytes such as Astelia solandri, f;arina mucronata, E. autumnalis, Dendrohium cunning­hamii, and Griselinia lucida are more common, and Blechnum filiforme is usually present on the forest floor and on tree trunks.

BLACK BEECH FOREST

In the north-west corner of the Aorangi Range, and particularly in the Waihora catchment, the bulk of the forest is dominated by moderately dense, 50-60-ft-high black beech (Nothofagus solandri). In this area the species occurs with equal prevalence on valley sides, river flats, and ridge tops. Throughout the rest of the range, however, black beech forms an open cover, 30-40-ft-high, and is limited to the drier, more exposed spurs.

From the headwaters of Tauanui River northwards hard beech (Notl/O­fagus truncata) may occur in association with the black beech. Hard beech reaches its maximum abundance in the Paharakeke catchment where, in limited areas, it may be dominant. Throughout the rest of the Aorangi Range black beech is usually not associated with any other tree species, except in the transition zones. where it may be found in associa­tion with the dominants of the surrounding mahoe-hinau-rewarewa. red beech (Nothofagu.l' fU.l'ca). and silver beech (Nothofagus menziesii) communities.

The understorey of black beech forest generally lacks complexity. Kamahi (Weinmannia racemosa) may form a scattered second storey and there is often a moderately dense shrub tier of Cyatho<ies fal'ciculata and small-leaved species of Copro.l'ma. Where the canopy is open Uncinia spp. and various small grasses. such as Notodanthonia gracilis. may form the forest-floor cover. Where the canopy is denser the main vegetation is

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1967] WARDLE - AORANGI RANGE

FIG. 4-Mahoe-hinau-rewarewa forest on river terrace, Pararaki valley. Browse-resistant Uncinia spp. and Micro/aena av('nacea have spread in response to opening up of canopy.

FIG. 5-Mahoe-hinau-rewarewa forest, Mangatoetoe valley. The partial1y open canopy has al10wed sufficien.t light to reach the forest floor to permit the development of a cover of browse-resistant Uncinia spp.

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30 NEW ZEAI.AND JOllRNAI. OF BOTANY [MARCH

FIG. 6-Black beech forest on spur, Pararaki valley. The trees form a moderately dense canopy at 30-40 ft. The forest floor is bare except for litter and occasional ephemeral seedlings.

of small ferns such as Grammiti.1 hillardieri, HymenOIJ/Jyl/um spp .. and Trichomanes renijorme. but the floor is predominantly litter-covered or bare (Fig. 6).

Black beech in the Aorangi Range does not usually grow above 1.600 ft. Occasionally it may be found as a pure stand at 1 ,SOO ft. and scattered trees Occur up to 2.6()() ft. e.g .. on the summit of Whawanui.

RED BEECH FOREST

Red beech forest is extensive from Otakaha Stream to the northern limits of the range. Generally it forms a band, the lower altitudinal limit of which varies with aspect and with the composition of the neighbouring communities but usually lies between 1.400 ft and 1.900 ft. The upper altitudinal limit is usually about 200- 300 ft below the crests of the higher ridges. i.e .. those between 2.0no ft and the highest point in the range.

Red beech generally forms an open forest dominated by large. well spaced trees, 70-S0 ft high and frequently more than 40 in. in diameter.

Changes in associated species are often related to the composition of the neighbouring communities. At the lower altitudes. where red beech abuts on mahoe-hinau-rewarewa forest. there may be a subcanopy of mahoe and perhaps. rewarewa. Podocarps. of which rimu is the most common. are sometimes emergent and may locally form pure stands to the exclusion of red beech. With increase in altitude they decrease in importance until their upper limit is reached at about 2.2<X) ft. Oil the other hand silver beech and kamahi. which occur in the upper p~rt of

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1967] WARDLE - AORANGI RANGE 31

FIG. 7-Blechnum discolor and pepperwood in red beech forest. Taua­nui catchment. Blechnum discolor often forms a very dense floor cover on the sheltered slopes between 2.000 ft and 2.200 ft.

the red beech zone. gradually increase in importance with increase in altitude.

In the lower to middle part of the red beech zone Olearia rani and the treefern Cyathea smithii (which replaces silver fern at these higher al,titudes) are often important subcanopy species. The shrub tier is usually dominated by Coprosm':l linariifolia. C. rhamnoides, and Cyathodes fasciculata. The canopy is usually open and this permits the development of a dense floor cover of water fern in seepage spots. and of Micro/aena avenacea and Uncinia spp. in the better drained sites. Between 2.000 ft and 2.200 ft. particularly on the sheltered aspects. there is often a very dense band of B/echnum discolor (Fig. 7) . Above 2.200 ft pepper­wood (Pseudowintera colora/a) becomes increasingly important as a subcanopy species. Silver beech and kamahi increase in abundance. and the density of ,the Blechnum discolor floor cover decreases. Mosses. lichens. and Blechnum minus also increase at these higher alti,tudes. but except for these and a few small ferns such as Hymenophyllum spp. and Grammitis billiardieri the forest floor is largely bare.

SILVER BEECH FOREST

Where red beech is important silver beech is restricted to a narrow band along the ridges between approxima'tely 2.000 ft and the highest points of the range. i.e .. Mt Te Munga at 3.211 fit and Mt Ross at 3.226 ft. Where red beech is not important. as in the sooth. silver beech forest extends downslope until it meets mahoe-hinau-rewarewa forest .

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32 NEW ZEAI.AND JOURNAl. OF BOTANY [MARCH

On the exposed north and west faces and on the highest ridge crests throughout the range. silver beech forms a dense. stunted forest. often less than 30 ft high. with a smooth wind-shorn canopy. Kamahi is frequently associated with silver beech in the tree layer. but except for occasional stunted pepperwood and Coprosma parvifiora. a shrub under­storey is lacking. Mosses. lichens. and filmy ferns often form a mat over much of the forest floor and extend up the trunks and branches of the trees.

Below the ridge crests and on the more sheltered slopes silver beech forest becomes taller and less dense. thus permitting the development of a more complete understorey. Pepperwood and Cyathea smithii may form dense thickets which are difficult to penetrate; Pseudopanax simplex. toro. fuchsia (Fuchsia excorticata), and broad leaf (Griselinia Iittoralis) are occasionally important species; and a floor cover of water fern. Blechnum minus, and B. discolor may also be present. Red beech is often associated with the silver beech on these more sheltered sites.

Hall's totara (Podocarpus h!JIlii) is sometimes found in association with silver beech. Mature trees of this species are usually confined to the sharper. more exposed ridge-top sites. and most show signs of senility. Few medium-sized trees are to be found. although seedlings and saplings are often fairly plentiful.

MAJOR MODIFYI NG FACTORS

EROSION

The bedrock of the Aorangi Range has been shattered and deformed to an extreme degree by the progressive movement of numerous faults and thrust planes--many still active. In consequence. downcutting by rivers. many of which follow fault lines and shatter zones. is rapid, and slipping is frequent. The seral communities which develop in these disturbed areas may be divided into three broad groups-those that occupy. respectively. the slips. the higher eroding gully heads. and the aggrading stream beds.

The pattern of succession on slips is a complex one. varying with slope. aspect. altitude. type of slip. position on slip. and composition of the vegetation surrounding the slip. Slips in black beech forest may be colonised by kanuka (Leptospermum ericoide.l'). Cyathodes fa,dculata. and C. juniperina. or sometimes by black beech seedlings. In the red beech and upper mahoe-hinau-rewarewa zones. water fern. Urtica ferox. and Uncinia spp. may be the pioneers. In the lower mahoe-hinau­rewa rewa zone the succession varies considerably. On moist slips and towards the shaded edges of drier slips. Blechnum spp .. water fern. or even treeferns may pioneer. Frequently there is direct colonisation by small tree species such as mahoe. wineberry (A ristotelia .,·errata). fuchsia. Carpodetus serratus, and Pennantia corymhosa. On the drier sites tauhinu (Cauinia leptophylla). kanuka, Helichrysum gfomeratum. and small-

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leaved species of Copmsma enter into the succession at an early stage. though they usually follow Raoulia glabra. Helichrysum filicaule. Noto­danthonia spp .. and in places Unicinia spp. and Microlaena avenacea. Along stream banks and on moist. well illuminated slump sites near streams. especially where the exposed rock is soft and much shattered. tootoo (Cvrtaderia fulvida) is sometimes an important coloniser.

Mahoe. fuchsia. and wineberry were probably the most important pioneer species in the actively eroding gully heads and along the sides of steep. down-grading tributary creeks. Browsing by introduced herbivorous mammals. however. has reduced fuchsia and wineberry in these sites. and in many places only dead sticks provide evidence of their former importance. Where these two species have disappeared mahoe forms a very open stand with a dense ground cover of water fern. Urtica femx. and U. incisa. Broadleaf is often associated with mahoe in the gully heads. but is more important on the slightly better drained. more stable sites. Scattered rewa rewa are usually present in the gully heads and Hoheria sexstylvsa may be locally common.

The succession on stabilising river terraces seems to follow a fairly consistent pattern. The two scabweeds. Raoulia glabra and Ravulia tenuicaulis. Epilobium spp .• and small grasses and other herbs are often the first pioneers on fresh gravelly alluvium. These rapidly give way to tauhinu scrub. of varying density. which provides protection for the development of a moderate ground cover of herbs. mosses. and lichens. Frequently tauhinu itself will colonise fresh alluvium.

If the river terraces remain stable for long enough. the tauhinu eventu­ally gives way to kanuka scrub and this in turn to tree species. Up to an altitude of 700-800 ft titoki (A lectryon excelsus) may replace the kanuka scrub. Titoki often forms pure stands 20-30 ft high. with a shrub under­storey of pigeonwood. Myrsine australis. Carpodetus serratus. Lopho­myrtus spp .• Pennantia corymbosa. Copmsma areolata. and C. rham­noides. and a floor cover of Uncinia spp. and Microlaena avenacea. Seedlings and saplings of mahoe. hinau. and rewarewa are common. and these species eventually replace the titoki. On some of the more mature terraces rimu. miro. kahikatea. and matai grow up through and overtop the mahoe. hinau. and rewarewa.

Ngaio (Myvporum laetum) and karaka (Corynocarpus laevigatus) are two tree species often associated with titoki on the river flats. Karaka may have been introduced by the former Maori inhabitants of the area (Adkin. 1955).

FIRE

The Aorangi Range. especially its southern section. has had a very complex fire history (Fig. 8). Before Europeans settled in the area there was a heavy Maori population concentrated in the coastal fringe (Adkin. 1955). The vegetation pattern of the range suggests that these Maoris were responsible for large-scale fires. More recently. land-clearing and

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34 NEW ZEALAND JOURNAL OF BOTANY [MARCH

burning-off fires started by European settlers have spread into the range. The unpublished Annual Report of the Wellington and Wairarapa District Acclimatisation Society for 1891 records that quite extensive areas were burnt during that year. Ring-count studies on kanuka suggest that other large-scale fires occurred between 1850 and 1880. In the early 1920s a fire burnt an area of forest in the north-west corner of the southern section of the range.

FlO. 8-Remnant stands of silver beech. Waitutuma catchment. The fire­induced scrub between these stands i~ dominated by kanuka.

Three main vegetation types have developed in response to this fire history. One of these. tall kanuka scrub. is seral and eventually replaced by adjacent climax communities. The other two. subalpine scrub and subalpine tussock grassland. are maintained in a subclimax condition by a harsh climate and by heavy animal use.

Tall Kanuka Scrub

Patches of fire-induced kanuka scrub occur throughout the Aorangi Range, but most of it ~s concentrated near agricultural land. This scrub seldom extends to altitudes above 1,500-2,000 ft. Kanuka seedlings are generally considered to be intolerant of the shade provided by the parent trees. and second-generation kanuka stands are rare. although there are areas where wind throw or death of some of the larger trees has allowed the species to regenerate. More frequently. however. dispersal of other species from the surrounding communities occurs during the first genera­tion. and eventually these species replace the kanuka. Here and there. both on slips and burns. an unusually vigorous kanuka may survive and

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form part of the new canopy. This, and the incidence of kanuka as a slip coloniser on the better drained sites, accounts for the scattered occurrence of the species in mahoe-hinau-rewarewa, red beech. and black beech forest.

Understorey and forest-floor composition varies greatly with the age of the kanuka stand. In the younger stands. which are frequently dense. the kanuka litter forms a thick layer over the mineral soil. At this stage there is little undergrowth. but as the stands become older they open up and other species establish. Lycopodium spp. frequently comprise the first ground cover. but Cyathodes fasciculata and C. juniperina are often found where the canopy allows sufficient light to penetrate.

Subalpine Scrub

Most of the subalpine scrub in the Aorangi Range is confined to a relatively small areaaround Mt Barton. although there are patches near the summits of Whawanui, Mangatoetoe. and Bull Hill. Charred remains of logs are numerous throughout. Most of these appear to be of beech­almost certainly silver beech. Occasional large. broadleaf trees are still to be found growing along moist drainage ways well within the scrub boundaries. and on the south side of Mt Barton a solitary silver beech was found 200-300 ft from the summit. It seems unlikely. too, that Mt Barton would have supported a climax subalpine scrub while Mt Ross and Mt Te Munga. approximately 300 ft higher, were and still are clothed to their summits in silver beech forest (Fig. 9). From the

FlO. 9-Silver beech regenerating freely in a trig station clearing on the summit of Mt Te Munga. 3.211 ft. The regeneration is vigorous enough to withstand the heavy browsing to which it is subjected.

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36 NEW ZEALAND JOlJRNAI. OF BOTANY [MARCH

evidence of the charred logs it can be assumed that fire destroyed the forest cover on Mt Barton and elsewhere and was thus responsible for the development of the subalpine scrub. The main factors perpetuating this scrub vegetation have probably been repeated burning. the low regenerative potential of forest species in the harsh climatic conditions found at higher altitudes. the slow rate of spread of silver beech. and the heavy browsing of tree seedlings by introduced mammals.

As would be expected with fire-induced vegetation. distribution patterns are complex and the species occurring on any particular site can be explained more readily in terms of which species got there first. than in terms of habitat preferences of the species concerned. Even so. most of this scrub can be classified into one or other of two broad types. The more extensive of these is dominated by Dracophyllutn fi/ito/iun!. manuka (Leptospermum sco{JariUln). Cyathodes fascicu/ata. and CO{Jrosma {Jarvi­fiora. This type is confined to the Mt Barton area. but is found on all aspects and on a wide range of sites. It is generally open and seldom exceeds 4 ft in height (Fig. 1 (). Both height and density vary with degree of exposure; on north-west slopes and at higher altitudes the scrub becomes short and very open. In the open spaces ground cover is usually provided by Ce/misia spectahi/is. B/echnum minus. Senecio /aRopus. Lycopodium spp .. and small grasses. Ce/misia spectahi/is is the com­monest of these species; it becomes increasingly important as the shrub component diminishes with increasing exposure of the site (Fig. II). The relative importance of the shrub species in this type varies considerably. In some areas manuka and Cyathodes fascicu/ata form a local community to the exclusion of the other two species. In others Dracophyllum filitolium may dominate. and in moist. sheltered sites CO(Jfosma {Jarvifiora may be the only shrub species present.

On Whawanui and Mangatoetoe, and to a minor extent around Mt Barton, scrub dominated by Hehe venustu/a and Senecio e/aeaRnifo/ius is to be found. Associated species again include manuka. Cyathodes fasciculata, and Coprosma {Jarvifiorr.J. This type may reach a height of 10-12 ft and is usually denser than the one described above. Floor cover is usually sparse or absent. On the more exposed sites Hehe venustu/a may be the only shrub species present. Where this is so. the height of the scrub is generally not more than 3-5 ft. A number of small tree species found in the upper forest types are also common in the Hehe venustu/a­Senecio e/aeaRnifo/ius scrub. Prominent among these is broad leaf.

Suhalpine Tussock Grassland

Tussock grassland is found mostly in a small area surrounding Mt Barton. This community .. like subalpine scrub, appears to have been induced by fire. The dommant species. Chionoch/oa cheesemanii, varies in density with the degree to which the site is exposed. On sheltered sites it forms a dense cover 3-:-5 ft. tall. with occasional plants of CO{Jrosma spp. and Phormium co/ensOl (Fig. 12). On more exposed sites the tussocks are smaller, and the spaces between them large and frequently bare of

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FIG. 100Induced scrub near Mt Bavton dominated by Dra(:ophyllum filifolillm, manuka, and Cyathodes fasciculata. Remnant trees of silver beech are scattered throughout. Numerous remains of logs testify that this area was formerly under forest.

FIG. II-Induced scrub near Mt Barton dominated by Dracophyllum fili/olium. With increase in al1titude the scrub cover becomes sparse and stunted, and Celmisia spectabilis becomes common.

37

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38 NEW ZEALAND JOURNAL OF BOTANY (MARCH

FIG. 12-lnduced Chiolloch/oa cheesemallii tussock grassland near Mt Barton . The tussock forms a dcnse cover 3-5 ft high, and has few associated species .

vegetation. A greater variety of herbaceous species is present, the most important being Celmisia spectahilis. This species becomes increasingly common as the degree of exposure increases.

INTRODUCED HERBIVOROUS MAM MALS

Deer and goats were reported to be present in high concentrations throughout the Aorangi Range by the year J 880 (L. T. Pracy, pers. comm.). The first reliably recorded opossum liberations took place in 1921 (Pracy, 1962), but it seems probable that there was an carlier liberation in 1903 (L. T. Pracy, pers. comm.). There has therefore been sufficient time for modification of the vegetation by introduced herbivor­ous mammals to proceed to an advanced stage.

To gauge the degree of modification that has taken place one must reconstruot a picture of the forest as it was before these mammals were introduced. For this purpose observations have been made of animal excIosures; of remnant patches of vegetation from which browsing animals are e.xc~uded by the nature of the topography; and of the com­position of Similar forest types in areas where browsing pressure has been less intense. Species of known high palatability have been observed as ephemeral seedhngs .. as epiphytes. as mature trees, or as bare skele­tons, 'but not as established seedlings, saplings, or as trees within reach of browsing animals. Verbal descriptions of the area by people who were familiar with it in the past have also been used. From sources such as these it has been pos~i~le to obtain an approximate idea of the former struoture and compoSItion of the main forest types.

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The mahoe-hinau-rewarewa forest previously had a very much denser understorey than it has now; in it the large-leaved species of Coprosma. C. australis, C. rohusta. and C. lucida. were important. These species are still very plentiful as seedlings. but rare as shrubs. Other shrub and tree species which were probably once abundant in this forest type but are now rare include hangehange (Geniostoma Iigustrifo/ium). Pittosporum eugenoides, P. tenuifolium, and Pseudopanax edgerleyi.

The black beech forest. on the other hand. has probably always had an understorey dominated by xerophytic species of low browse preference. although large-leaved species of Coprosma were probably. again. more abundant than they are now. The presence of logs shows that northern rata (Metrosideros robusta), a tree species associated with black beech on dry spur sites. was once fairly common in the Aorangi Range. but it is now nearly extinct. Opossums have probably been the main factor in its elimination.

The red beech and silver beech forests formerly had understoreys rich in palatable species such as Pseudopanax simplex, Coprosma foetidissima, fuchsia. and broad leaf. These species are still present. but only as old trees out of reach of browsing animals. or in areas that are difficult of access.

The near-elimination of many of these palatable shrub and small-tree species from the understoreys opened an ecological niche that could be successfully filled only by species of low browsing preference. At first. while the forest canopy remained intact. these species would have had to be tolerant of moderately shady conditions. The small-leaved species of Coprosma were virtually the only shrub species capable of spreading under these conditions.

The seedlings and saplings of most of the important tree species in the Aorangi Range are palatable to deer and goats. Browsing of seedlings and saplings by these animals eventually permitted increased amounts of light to reach the forest floor. since it eliminated the young trees that would have filled canopy gaps. The opening of the canopy was undoubtedly hastened by the activities of opossums. Better illumination of the forest floor encouraged the spread of the species with low palat­ability. In mahoe-hinau-rewarewa forest the large-leaved spp. of Uncinia were the most important of these. In red beech forest the fern Blechnum discolor became widespread. and in silver beech forest the more open conditions resulted in an increase of pepperwood. These species. especially the non-woody ones. form a dense floor cover that tends to prevent the establishment of tree seedlings. The ultimate stage in this forest modifica­tion may well be conversion to a turf. for many of the species which are now increasing. even though they thrive in 'ovell illuminated conditions. show signs of being unable to survive once the protection of the open forest cover is further reduced.

Perhaps the most spectacular modlification by animals is to be seen on the slips and in the gully heads. Many of the plant species that formerly colonised these sites were fast-growing. at least in the seedling and sapling

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40 NEW ZEALAND JOURNAL OF BOTANY [MARCH

stages: they included fuchsia, the large-leaved spp. of Coprosma, Pseudo­panax spp., SchefJIeru digitata, and wineberry. The early luxuriant growth of these plants made them particularly susceptible to browsing, so that they are now to be found on such sites only as seedlings. In consequence succession on slips and gully-head scars is much slower than it was formerly.

ACKNOWLEDGMENTS

I wish to thank for their assistance the personnel of the Napier branch of the Forest and Range Experiment Station, New Zealand Forest Service, and the students employed on the Aorangi Range survey. I also wish to thank Messrs J. T. Holloway and M. J. Wraight for their helpful suggestions.

REFERENCES

ADKIN, G. L. 1955: Archaelogocial evidence of former native occupation of east Palliser Bay. J. Polync.\". Soc. 64: 450-480.

ALLAN, H. H. 1940: "A Handbook of the Naturalized Flora of New Zealand." Government Printer, Wellington.

ALLAN, H. H. 1961: "Flora of New Zealand." Vol. I. Government Printer, Wellington.

CHEESEMAN, T. F. 1925: "Manual of the New Zealand Flora." 2nd ed. Govern­ment Prin'ler, Wellington.

COCKAYNE, L. 1958: "The Vegetation of New Zealand." 3rd cd. Engelmann, Leirzig.

GRINDLEY, G. W., HARRINGTON, 1. H., and WOOD, B. L. 1959: The geological mal' of New Zealand, t :2,000,000. BII/I. Gco/. Sun'. N.Z. n . .\". 66.

HOLLOWAY, 1. T. and WENDELKEN, W. 1.1957: Some unusual prcoblems in sample plot design. N.Z. JI For. 7 (4): 77-83.

KINGMA, 1. T. 1959: The tectonic history of New Zealand. N.Z. JI Gco/. Geophys. 2: I-55.

PHILIPSON, W. R. 1965: The New Zealand genera of the Araliaceae. N.Z. JI Bot. 3: 333-41.

PRACY, L. T. 1962: I~troduction and liberation of the opossum (TrichoSllrtls vlI/pcelllll) Into New Zealand. N.Z. For. Servo In/orm. Ser. 45.

WELl.MAN, H. W. 1956: Structural outline of New Zealand. BII/I. N.Z. Del'. sciellt. indo Res. 121.

WELl.MAN, H. W. and ~RODIE, 1. W. 1954: A note on the geology of Cape Palliser, New Zealand. N.Z. JI Sci. Technol. B35: 440-50.

ZOTOV, V. D. 1963: Synopsis of the grass subfamily ArundinoidCC/c in New Zea­lanJ. N.Z. JI Bot. J: 78-136.

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ApPENDIX I-COMPOSITION OF THE MAIN FOREST TYPES

Tables 1-4 give a quantitative summary of the composition of the main forest types. To conform with the plot measurement technique employed. tree in the following tables refers to any stem with a diameter of 1 in. or more at 4 ft 6 in. above ground level, and shrub to any woody plant whose total height exceeds I ft and whose stem diameter. 4 ft 6 in. above ground level. is less than 1 in.

The figures presented in Tables I and 2 are averages from a number of 1/10 acre cruciform plots. The cruciform plot consists of a belt transect measuring 200 X 25 links running up and down slope from the plot centre. with the addition of two wings. each 100 X 25 links. running across slope through the plot centre.

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TA

BL

E I-

Basa

l ar

ea (

sq f

t)

and

num

ber

of t

rees

, pe

r ac

re

.j:>.

IV

M

aho

e-h

inau

-B

lack

bee

ch

Red

B

eech

S

ilve

r be

ech

rew

a rew

a fo

rest

fo

rest

fo

rest

fo

rest

S

peci

es

(8

plot

s)

(6

plot

s)

(7

plot

s)

(7

plot

s)

Bas

al

No

. o

f B

asal

N

o. o

f B

asal

N

o. o

f B

asal

N

o. o

f ar

ea

tree

s ar

ea

tree

s ar

ea

tree

s ar

ea

tree

s ---

Mel

ieyt

us r

ami{

ioru

s 52

13

0 <

1

<5

<

1

<5

Z

tT

l E

laeo

earp

us

dent

atll

s 43

20

~

Kni

ghti

a ex

eels

a 41

20

6

\0

<1

<

5

6 \0

N

H

edye

arya

arb

orea

33

16

0 1

10

tTl >

G ri

seli

nia

Iitt

oral

is

19

\0

2 <

5

2 <

5

7 20

r >

P

odoc

arpu

s sp

ieat

lls

14

<5

z 0

Not

hola

glls

sol

andr

i 7

<5

12

1 14

0 2

<5

6

10

.... W

einm

anni

a ra

eem

osa

20

150

19

70

39

190

0 c:: N

otho

lagl

ls I

llsea

15

\0

15

7 12

0 38

20

;.:

l z N

otho

lagu

s m

enzi

esii

4

\0

35

30

144

290

>

r L

epto

sper

mll

m

eric

oide

s 6

\0

7 10

38

40

0

Pod

oear

plls

le

rrll

gine

lls

5 <

5

19

\0

'Tl

Cya

thea

sm

ithi

i 3

10

19

50

t:::O

Dae

rydi

llm

eu

pres

sinu

m

9 <

5

2 <

5

11

<5

~ >

D

ieks

onia

sq

uarr

osa

\0

20

z P

seud

owin

tera

eol

orat

a 7

100

30

2 70

-<

Myr

sine

au

stra

lis

5 50

1

10

<1

\0

N

otho

lagl

ls

trun

eata

8

30

Myr

sine

sa

/ici

na

2 <

5

8 <

5

\0

2 20

C

yath

ea d

ea/b

ata

<5

5

<5

1

<5

~

Pod

ocar

pus

hall

ii

<1

<

5

5 20

>

O

ther

sp

ecie

s·

11

7 3

9 ;.:

l ("

) ;x:

Tot

al

250

204

230

357

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*Tre

e sp

ecie

s pr

esen

t w

ith

basa

l ar

eas

not

exce

edin

g 5

sq f

t pe

r ac

re i

n an

y of

th

e m

ain

fore

st t

ypes

. T

hese

inc

lude

:

In M

ahoe

--H

inau

-Rew

arew

a F

ores

t

Ale

etry

on e

xeel

sus

Bra

ehyg

lott

is

repa

nda

Car

pode

tus

serr

atus

C

opro

sma

rotu

ndif

olia

C

ordy

line

au

stra

lis

In B

lack

Bee

ch F

ores

t

Car

pode

tus

serr

atus

C

yath

odes

fa

sdeu

lata

C

yath

odes

ju

nipe

rina

In R

ed B

eech

For

est

Ari

stot

elia

se

rrat

a C

opro

sma

aust

rali

s C

yath

odes

fas

deul

ata

In

Sil

ver

Bee

ch

For

est

Car

pode

tus

serr

atus

C

opro

sma

foet

idis

sim

a C

yath

odes

fas

deul

ata

Dra

coph

yllu

m f

ilif

oliu

m

Gau

lthe

ria

anti

poda

Lep

tosp

erm

um

seop

ariu

m

Lop

llO

myr

tus

obeo

rdat

a M

aero

pipe

r ex

eels

um

Pse

udop

anax

arb

oreu

m

Ole

aria

ran

i

Ole

aria

ran

i P

seud

opan

ax

cras

sifo

lium

S

ened

o e

laea

gnif

oliu

s

Myr

sine

di

vari

cata

O

lear

ia r

ani

Pse

udop

anax

era

ssif

oliu

m

Pse

udop

anax

sim

plex

O

lear

ia r

ani

Pse

udop

anax

era

ssif

oliu

m

Rub

us s

pp.

-\0 ~

~

~ [2 t"l1 I ~ >

Z 9 ~

>

Z

(;)

t"l1

~

w

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44 NEW ZEALAND JOURNAL OF BoTANY [MARCH

TABLE 2-Number of shrubs per acre

Mahoe-Black beech beech beech hinau- Red Silver

Species rewa rewa forest forest forest forest (6 plots) (7 plots) (7 plots)

(8 plots)

Coprosma rhamnoides 480 640 190 160 Brachyg/olTis repanda 220 Pellllantia corymbosa 190 Helichrysum g/omeratum Ito 70 <5 Pseudowintera c%rata Ito 150 740 Coprosma microcarpa 10 760 70 Cyathodes /ascicu/ata <5 640 570 90

Coprosma ballksii 520 110 <5 Cyathodes juniperilla 450 20 N otho/agus so/andri 300 20 Coprosma linarii/olia <5 40 410 20 Coprosma parviflora 180 Macropiper exce/sllm 80 Urtica /erox 70 Parsonsia heterophylla 50 Carpodetus serratlls 30 20 <5 Coprosma rotundi/olia 20 Rhipogonllm scandem to Knightia excelsa 10 <5 Lophomyrtlls obcordata to Pseudopanax crussi/oli/utl 2(:) 10 Leptospermum ericoides <5 to to Coprosma areo/ata to Nothofagus menziesii <5 80 70 Notho/agus /usca 50 <5 Podocarpus /errugineus 20 Myrsine divaricata 20 Coprosma /oetidissima <5 to <5 Podocarpus hallii <5 to 60 Coprosma australis to <5 Pseudopanax co/ensoi to Wl'inmannia racemosa <5 <5 10

Leptospermllm scoparium <5 <5 to Psclldopanax simplex to

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Other shrub species. present at densities of less than 10 stems per acre. include:

In Mahoe-Hinau-Rewarewa Forest

Beilschmiedia tawa Gymnelia lanceolata Melicope simplex Metrosideros diUusa

In Black Beech Forest

Gaultheria antipoda Gymnelaea lanceolata

In Red Beech Forest

Aristotelia serrata Coprosma lucida Dacrydium cupressinum Elaeocarpus denta/us Fuchsia excortica/a

In Silver Beech Forest

Cassinia leptophyl/a Dacrydillm cupressinum Dracophyl/um (ili/olium

Olearia paniculata Olearia rani Plagianthus betulinus Podocarpus spicatus

Notho/agus truncata

Griselinia littora/is Olearia rani Pseudopanax edgerleyi Rubus cissoides

Fuchsia excorticata Olearia paniculata

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46 NEW ZEALAND JOURNAL OF BOTANY [MARCH

Tables 3 and 4 list the herhaceous plants making up the forest-floor cover; these have been arbitrarily classified into major (more than 1 ft high) or minor (not exceeding 1 ft). The frequency of a species in any forest type is the percentage of milliacre plots within that type in which the species occurs, i.e., in which it is rooted. There are 20 miIIiacre plots in each cruciform plot-five spaced at 20-link intervals along the centre lines of each of the four sections.

TABLE 3-Frequcncies of major forcst-floor species

Mahoe-hinau- Black beech Red beech Silver bcech

Species rewa rewa forest forest forest forest (120 plots) (140 plots) (140 plots)

(160 plots)

Uncinia spp. (broad-leaved) 42 II 12 6 Cyathea dea/bUla JO 4 2 Po/ystichum silvaticum 8 Juncus spp. 5 Uncinia (narrow-leaved) 3 10 3 Micro/aena avencea 3 5 9 6 B/echnum disc%r 51 JO Dicksonia squarrosa 6 I Chionochloa sp. S Cyathea smithii 2 14 Histiopteris incisa 4 2 8

Other major species which are present but with I rcqucncics of Icss than 5, arc:

In Mahoe-Hinau-Rewarewa Forest B/echnum lanceo/atum Carex spp. Ctenitis decomposita DianeJ/a intermedia Echinopogon ovatus

In Black Beech Forest Deycuxia crinita Gahnia sp.

In Red Beech Forest Gahnia sp. Uhcrtia ixioides Po/ysticltum vestitum

In Silver Beech Forest Biecltnum fluviatile Libertia ixioides PeJ/aca rotundifolia

Hypo/epsis rugosu/a Libcrtia ixioidcs N %dan/honia sp. Pel/aea rotlllldifo/ia Ptcris maci/cnta

HymcllophyJ/um sp. Pteris macilellta

Tociea Itymcllopltyl/oidcs Todell su pcrha

I'o/ysticltllm vestitum Rumoltra lldillntiformis Todell superha

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TABLE 4-Frequencies of minor forest-floor species

, Mahoe-hinau- i Black beech! Red beech I Silver beech Species rewarewa forest, forest forest foreSit

(160 plots) , (120 plots) I (140 plots) , (140 plots)

Ullcillia spp. (broad-leaved) 42 27 41 34 Pel/aea rotundi/olia 39 3 6 Asplenium hookerianum 39 5 II Phymatodes diversi/olium 21 IO 9 15 Ranunculus hirtus 18 2 7 Polystichum silvaticum 15 Asplenium flaccidum 14 10 9 14 Pyrrosia serpens 14 II 4 11 Sonchus sp. 14 2 10 Oxalis lactea 14 IO Polystichum vestitum 13 7 Carex sp. 13 Asplenium bulbi/erum 11 6 4 Hypochoeris radicata 10 4 2 5 Hymenophyllum spp. 1 50 55 36 Grammitis billardieri 3 47 63 49 Dianella intermedia 5 14 4 Trichomanes reniforme \3 7 Blechnum minus 2 15 46 Blechnum discolor 14 5 Blechnum fluviatile \3 7 Chiloglottis cornllta 2 18 Nertera spp. 5 2 16 Cory bas triloba 6 3 13 Cerastium spp. 9 1 11 Polystichum richardii 9 6 Hydrocotyle spp. 9 3 9 Cardamine debilis 8 2 Gnaphai;um spp. 6 3 Oxalis corniculata 6 Viola spp. 6 4 Cirsium vulgare 5 Acaena anserinifolia 5 Notodanthonia gracilis 9 Pterostylis banksii 7 Grammitis heterophyl/a 6 2 f;arina alltumnaiis 5 Todea superba 6 1 Microlaena avenacea 3 3 1 8 Histiopteris incisa 1 4 7

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48 NEW ZEALAND JOURNAL OF BOTANY [MARCH

Other minor species which are present but with frequencies of less than 5, are:

In Mahoe-Hinau-Rewarewa Forest

Agropyrum scahrum Arthropodium candidum Asplenium falcatum Asplenium lucidum Blechnum filiforme Blechnum lanceolatum Craspedia un if/ora Crepis capillaris Ctentitis decomposila [)aclylis glomerala Echinopogon ovalus Epilohium spp. Galillm sp. Hypolepis rugosula

In Black Beech Forest

Asplenium falcotum Echinopogon ovalus Galillm sp. Luzula sp.

In Red Beech Forest

Asplenium faleatum Bleehnum eapense Bleehnum lanceolatum Chionochloa sp. EpiJohium sp. Luzula sp.

In Silver Beech Forest

Anthoxanthum odoralum Asplenium falealum Bleehnum capense Epilohium sp. Holcus lanatus luneus spp. Lagellophora peliolala

Luzula sp. N otodanlllOnia sp. Paesia scaherula Poa anceps Poa eaespilosa Poa imhecilla Pleris tremula Raoulia tenuicaulis Rumohra hispida Senecio jacohaea Solanum nigrum Taraxacum ofjicinale Wahlenhergia gracilis

Pleris Iremula Senecio I(/goplls Uncinia spp. (narrow-leaved)

Paesia scaherula Pleroslylis venosa Raoulia glahra Rumex acelosella Rumohra adianliformis Todea hymenophylloides

Libertia ixioides Luzula sp. Paesia seaherula Poa caespitosa Rumohra adianliformis Tmesipteris tannensis Wahlenhergia gracilis

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