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Tussock grassland communitiesin the Mackenzie Country,South Canterbury, New ZealandH. E. Connor a & C. G. Vuceticha Botany Division , Department of Scientific andIndustrial Research , ChristchurchPublished online: 20 Feb 2012.

To cite this article: H. E. Connor & C. G. Vucetich (1964) Tussock grasslandcommunities in the Mackenzie Country, South Canterbury, New Zealand, New ZealandJournal of Botany, 2:4, 325-351, DOI: 10.1080/0028825X.1964.10428730

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NEW ZEALAND

JOURNAL OF BOTANY Department of Scientific and Industrial Research, Wellington.

Editor: Mabel Rhe

VOLUME 2 DECEMBER 1964 NUMBER 4

TUSSOCK GRASSLAND COMMUNITIES IN THE l\IACKENZIE COUNTRY, SOUTH CANTERBURY,

NEW ZEALAND

H. E. CoNNOR

llotany Division, Department of Scientific and Industrial Research, Christchurch

(With Appendix by C. G. VUCETIQI)

(Reu;I'ed lor I,,,b/iea/ioll, J July 1964)

SUMMARY

There lire three major tussock Itrassland communities in the MJckenzie Country: (i) fescue·tuss(l(k ItrasslJnd where FIS/llra lIo,'ae·ze/a'ldide is abundJnt, (ii) red. tunock grassland where Chiolloch/oa rllbra is the physiognomic dominant, and (iii) ,nnw.tussock grassland where Chio/lo,.h/od riRida is the rhysill~nllmic (Iuminant. Stlcioligical analyses of these communities in which severa pha~es ore reco~nised lire Iti"en in tables. The histocy of the granlands is traced from the destructIOn of the (orests thruugh tall.tusso(k stages to the current extensive shorl·tussock grassland of u'(('nt origin.

INTRODUCTION

111C~ Mackenzie Country has captured attention from the first days of the legend of Mackenzie the sheepstealer, through the days of settlement and the early ascents of Mt Cook in the National Park, to the present when new extensive plans have been drawn to utilise its vast water resources (or the production of electricity. The grasslands of the Mackenzie Country, and nearly all of the Mackenzie is grassland, have since 18~7 produced sheep and wool. Dut this part of the 13 million acres of Soutb Island tussock grassland has not been described in any botanical detail.

The Mackenzie Country is a basin bounded on the north and west by the Main Divide, to the cast by the Two TIlumb, Rollesby, D.1Igety, and

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Kirkliston Ranges and on the south by the Ohau and Waitaki Rivers. TIle Grampian Mountains swing into the Plain from the Kirkliston and Dalgcty Ranges. Three passes, llurkes, Mackenzie, and Hakataramea, separate the various eastern ranges (see Fig. 1).

Although the Mackenzie Country was known to the Maoris and its presence reported by Edward Short land to the alief Protector of Aborigines In 1844 (Gillespie, 19~8) and though C. O. Torlesse marked the Mackenzie Pass ("Mauahouna Pass") on his map of 1849 (Mating. 19~8) there is no record of entry into this basin until 18~'. In that year J. H, C. Sidebottom, after a very brief visit, informed his employer "There seems to be a tine plain just at the back of the Snowy Range, and a first-rate Pass through the mountains to it". Gillespie (oP, cit.) reported that late in 18~~ G. Dunnage explored the Mackenzie llasin and wrote. "The feed was very poor but would improve with burning", Dunnage found the plain strewn WIth large boulders among which grew quantities of sharp wild spaniards (Adph}'lItJ srp.). }laast (1879) traversed the Mackenzie Country in 1862, and though gIven to describing landscape makes no mention of the general vegetation of the area; he did, however. collect many plant specimens.

The geology is broadly outlined by Grindley, Harrington, and Wood (19~9), who show the rocks of the mountains as undilferentiated grey­wackes. Within the basin extendin~ out from the Main Divide to somewhat south of Lakes Tekapo and Pukakl are extensive moraines. and beyond the moraine lies the large outwash plain dissected by rivers and streams. The moraines are predominantly of till and gravel, and the outwash plains 0' well-graded grey wacke gravel. Tussock-covered fans of subangular gravel are present at the foot of the eastern ranges, e.g., the large fans (rom the Dalgety Range extending down to the swamps at the change in slope. Speight (1963) discussed the "landform associations" in the vicinity of take Pukaki.

Soil sets indicated in the tables are from data supplied by Mr C. G. Vucctich, Soil Uurcau, Department of Scientific and lndustrlal Research, Christchurch, see Appendix II,

The climate of the region is very varied, (rom the high rainfall Ilt Mt Cook to the low rainfall in the Uasin. Garnier (19~H) places the Mackenlie Country in the "Inland South Island (Climatic) Region", ". , , which is essentially microthc:rmal and either semi-arid or subhumid; ill contains the driest climates and the most extreme temperatures r('Corded in New Zealanll; anll it contains such variations over short distances thaI .1 map of its climatic types displays a mosaic of contrasting conditions". The following meteorological values arc from Garnier (01'. (it,), based on Lake Te:kapo readings; mean temperature 47°(0'; mean diurnal range of tem­perature: 20'40 ",; mean extreme maximum temperature H~'3°F; mean extreme minimum temperature B·~oF: mean annual rainfall 22'14 in. These values do not, however, represent the climate on the eastern ranges, which are especially subject to desiccating north-westerly winds. Nor do the values given indicate that mist pours over these eastern ridges. and particularly throu~h the passes and saddles, from the pile-up of cloud on the coastal side. Neither do they describe the dry southern area nor the wetter northern

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lands. Rainfall isohyets are presented in Fig. 1. Garnier also suggest~d. an ililand mOllntain dimaJe, ". . . a change to cooler and wetter condltlOns at an elevation between 3,000 and 3,500 ft above the sea". This altitude corresponds with the upper limits of the montane zone as indicated by Wardle (1963), and agrees with the present-day distribution of short- and tall-tussock grasslands.

All the land used for grazing sheep is held on lease from the Department of Lands and Survey except for one station and for the small freehold areas on the various runs.

Rabbits became a very serious problem in the Mackenzie and ~ere responsible for much damage to the fescue-tusock grasslands, more espeCially to those on the drier sites. Burnett (1927) reports "1878. Rabbit invasion lirst became acute. Regular rabbitcrs lirst employed at Haldan Station". Fol­lowing the formation of the Rabbit Destruction Council, and the adoption of a Killer Policy in 1947 control measures were effected. Rabbits were devalued in 1956. Recovery of the vegetation even at some of the most seriously depleted sites is now noticeable.

METIIODS

After a general inspection of tussock grassland in the area, sample sites considered representative of the various communities were selected. All samples were 100 sq. metres. At each selected sample area, altitude, aspect, and slope were measured, special features noted, and the sample plot carefully examined to determine all the plant species present. Each species was scored for the combined value abundance-dominance on a 5 to r scale, and {or sociability on a 5 to 1 scale; the scores used in these scales are presented adjacent to Table 1. Any species present in the stand though not in the sample plot were also recolded with appropriate notes.

In the tabulated data the live-figure number at the top of each column is the rei eve number; the lirst two digits correspond to the season of saml'ling. the remaining three to the sample number for the season, e.g., in Table I rcleve 63124 is {rom the one-hundred and twenty-fourth stand examined in the season 1962-63. A\I sample sites are indicated on Figure 1; details are given in Appendix I.

Grasslands were examined over a widc range of altitudes. aspects and slopes from the lower reaches of the Godley River to Hakataramea Pass on the eastern ranges, in the Dasin, and on the Den Ohau Range to the west. All major communities have been investigated, though the whole of the Mackenzie Basin has not been examined.

Variability within the tussock grasslands is of importance to botanists, to run·holders. to conservators. and to those who have the responsibility of administering these lands (or grazing. By taking a large number of samples. by sampling in places where the vegetation had not been disturbed {or some time. by sampling at different altitudes and on differing aspects. as well as by sampling in severely degraded communities. a range of variability has been retorded.

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SoCIOLOGICAL ANALYSIS

In the Mackenzie Country there are several distinct grassland communi· ties. Fescue·tussock grassland occurs on the plain, on much of the morainic country, and on some of the slopes of the boundary ranges. Red·tussocks are common at the sides of some streams and in the wetter places around them. but red-tussock grassland is also typical of much morainic country between Lakes Tekapo and Pukaki. Snow.tussock grassland occurs on the slopes of all the ranges. often right to the foot, and outliers of snow.tussock extend out on to the plain. e.g .• at Haldon Corner.

Ninety.one reI eves from altitudes between 1,4'0 and 4,800 ft are pre· sented. In general terms these lie within the montane and subalpine ~ones of Wardle (1963). Of these re1evcs

26 are from fescue·tussock srassland, Table 1 ; 4 are from weed commumties (sorrel or Broil/liS /etlorlllll). Table 2: 8 are from red·tusock grassland, Table 3; 9 are from grassland derived from red·tussock. Table 3;

36 are from snow·tussock grassland, Table' : 8 are from grassland derived from snow.tussock. Table 6.

Connor (1961) recognised four groups of species in snow·tussock grass. land in western North Otago, t'iz. SIlow·l/lJSod~ f,rass"lIId sprritS, uiJr. spread I/lssorl! f,rass/and spedtS, uiJrspreaJ aJ,'r1ll;,'r' spu;rs, and Irans­f,rtSsivr or duidenla/ speries. Analysis of about 300 species lists from various grassland communities has led to a better evaluation of the sociological rank of the numerous species. In this ,Paper six species groups are recog. nised. mainly because more communities have been examined here than in the North Otago area. The group u:iJesprrad tklrr'lIlit't sperits will not be used, though it has some merit. as most of its species are an integral part of the grasslands. The six groups are:

(i) Jl'/idespreaJ IllJJork f,raJs/alld speries. Those species found com­monly in all tussock grassland.

(ii) Shorl-llISso(k Krass/alld spuies. Those species which occur only or preferentially in fescue· or silver·tussock grassland.

(iii) U,(J·//lJJork ""au/and spuies. Those species which ocrur only or preferentially in red·tussock wassbnd.

(iv) Snow-II/ssork trass/and spefltJ. Those species which occur only or r.referentially an snow-tussock grassland.

(tI) Tai/·/llJJorl? "rdsslauJ spedes. Those species which occur in both red- and snow-tusso<k grassland and show no particular preference for either.

(Ii) AffiJmla/ Sptfiel alld sptfies 0/ III/rerfditl sial/IS. TIlose species which plainly belong in some other community but ocrur in tussock graSSland from time to time, and those infrequent species about which not enough is known.

These groups are based principally on the frequency and abundance/ dominance scores (or the species in tussock grasslands. even though the same species might occur in an adjacent habitat such as river bed. Some species listed were found no more than once or twice, and no importance

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is attached to them. Other species that occurred once or twice in ~ach of several community types are more difficult to deal with. No speCIes (or group of species) should be regarded as exclusive to any particular type of grassland. though some are almost so. and it is premature to designate any (haracter-species in the Ziirich-Montpellier sense however closely that cate­gory may appear to be approached. Detailed examination of other grasslands may well lead to reclassification of these groups.

FESCUE-TuSSOCK GRASSLAND

Table 1 presents data from 26 stands at altitudes between 1,500 and 2.800 ft, and in a variety of areas. It illustrates the wide variability within the fescue-tussock grasslands. Some subcommunities can be recognised though intergrading occurs.

Table 1 is arranged to show the gradient from iloristically simple to floristically complex re1evcs which follows increasing rainfall and the associ­ated soil set. It is divided into six phases. Some releves are difficult to place and may be the connecting links between phases; despite this, the separations indicate differences even though they cut across some lines of nearly continuous variation.

l:ach phase has a floristic complement that differs in some respect from the others. There is an increase in the number of typical mature phase I/;orl­IliJSod~ grassland species from Phase A to Phase F; this is most marked at Phase E. At the same time the small group of short-tlissock f.I'aJslanJ Il'uitl typical of Phase A (see below) decreases from Phase A to Phase E and is aimost absent in Phase F. Similarly the "Simons Hill Element" of Phases D and D (see below) does not enter the mature phases. The wide­sl'rea.i lussock grasslal1d species increase in number from Phase A through to Phase F.

TIle six phases are:

PUASE A. ltdevcs 63124, 63125. 63127: from gently sloping or flat wind-eroded areas on thin stony soil on fans lying into the north-west wind. FtSlllca nO"ae-ulafldide and Poa (o/mso; are the chief grasses, with abundant RII/IIIX au/osella and many plants of the small grass /'oa "Mllio-1010. I:cw other species are <juantitatlVely important. A group of shorl. IIl1Sork graulanJ Il'edtl is typical-Raollfia alllira/il, GYPlol'hila IIlblllosa. [;pilo/;i"111 ros/rallll". A1:roslis 111I1S(OSd. POd mani%lo, COnf'ohtlitll Vert­(1II1dtiS. and Pimt/ea p,d,·inariJ.

This 110ristically simple phase is a degraded fescue-tussock grassland con­taining some of the species typical of areas which have been severely ucpktro by rabbits (see also Table 2).

l'lIASr: U. RtlcvCs 6}087. 63091.63090, 63089: from flat and moderately stett' country Iring to the sun at "Simons Hill". Fes/llcd tl0t'alf-zllanJia(f is the chief native grass but the naturalised annual species Aira (drJopbyllta. VII/pia bromoidts. and DrolJl/lS lu/or/(/II are quantitatively important. Mat plants Raoll/ia allS/ra/is and R. hoo/uri are si~nificant. More widespread IIISJork .~assland spetiu are present than in Phase A. A small group of

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shorl-IIIJJork grdJs/and specieJ, the "Simons Hill Element", DrolllllS Itt/Ofl'III, Cirsilllll t'II'gart, Oxalis rorniCIIlaJa, Araena bll(hanallii. Mllthienbtrli.ia (om­pltxa, Dro11I11S mollis. Trifolillm bybridllll1, is typical.

PHASE C. Releves 63101, 6H02, 63104, 63105: from flat country on deeper soils than Phase A. This phase is characterised by F. tlot'ae-ztlandi.u and Poa rolensoi as the perennial grasses, and by much of the annual grass Aira raryophyllea. The "Simons Hill Element" of Phase II is almost absent. as too are the Phase A species Raolilia allslra/is to COIII'Oit'III,1S t'trerlmJlls. Phase C shows a strong increase in widesprtad /fISJork grasslalld sptrits. though few of them are quantitatively important. The shrubby species Pillle/a oreophila, Carll/khat/ia 1II0l1roi, and C. pttriti are prc:5ent, and among the species of lc:5ser stature Raoft/ia sllbserirea and Araena raesii­"Ia"ra. IVI/fle."( areloulla is abundant.

PHASE D. Releves 63088, 63092. 62133: maturer than Phase D, mostly from shaded slopes. The "Simons Hill Element" of Phase D is present, but fewer of the typical Phase A species occur. FtSlllra "ot'ae·ztlaIlJiolt hu still the highest abundance/dominance score. and to the U'ideJpreaa /I/Slock f,rassland speries are added lItlirhrysllln filiratlle. Sde"allrhllJ Il11iflorllS, and II ydrororylt no,'ae-zehmdiae.

PIIASE E. Relevcs 63137, 63126, 63149: from exposed sites, or on thin stony soils. The number of shorl Ifmork K'asslal1J sptties is similar to that in Phase F, but because of aspect or soil factors some species found on drier sites or on poorer soil remain. e.g., Raoldia allslralis and Dro11llls Ittlortol/. The list of tllidespre.ld IlIssoril f,rdJsland spuitJ is about the same as in Phase D, but less than in Phase F.

PHASE F. Relevcs 63150. 631'2, 63153, 631'4, 6309-1, 63122, 63139, 6313', 63136: from morainic downs and outwash terraces. TIlis is the most mature phase of fescue-tussock grasslands recognised in the Mackenzie Country. It is characterised by many sborl-IIIJSo(/i trdJslalld JPttitJ. by the absence of the Phase A s~C('ies and the "Simons Hill Element", and also by many tiJidtSprtdd IImodl "rassland sperits including the following which were either absent or present only at low frequency in other phases: Kotltri" Iwrlzii, Diuari.l 101ll1ldloll. Plantago spa/h,dala. Llizl/la (,dll/peSlris. Linlllll ralbarlirllll/, and Ctill/isia tratilmla. Among the Jhort-IIISJOr/i J',raJs/.md SpttitS is Coprosllla pelriti. a mat.forming woody species. Drdrb),(om, shulair;i is a fairly frequent mou··tllssork grass/and Sptt;es.

The six phases, although prc:5ented in sequence, do not represent the various stages in the evolutIOn of mature fescue·tussock grassland, but rather the condition of various stands, regardless of their ori~in or of the influence of man or animals. Nevertheless there is relationship among the phases. Phases E and F are more closely related than is Phase E to Phase D. Phases Band D arc closely related. though perhaps not as closely as are Phases D and F. Phases A and II show some similarity but the "Simons Hill Element" is an essential dilTerence. Further comparisons can readily be made from Table 1.

As was indicated above. the phases are closely associated with the soib and the rainfall gradient in the area. Despite this association there is tvi.

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dence that the stands for reI eves 63124 and 63125 (Phase A) were not always in their present condition. Re1eve 63126 (Phase E) is from a small area of grassland, not noticeably wind eroded, in the same area as 63124 and 63125 (Phase A). This sample may be representative of the tussock grassland before the severe depletion reflected in 63124 and 63125 had commenced. An opposite trend is seen from the stand for releve 631~9 (Phase E), which appears to be returning to the maturer stage seen. 10

rei eve 63150 (Phase F) in its recovery from previous heavy rabbit graziOg. Barker (1953) recognised a "Fesl/lca - Poa A lIiancc" of two associations;

in one Disraria 10/IIIIa/Oli was extensive, and the other, which had a larger species list, was from higher altitudes but still within the lowland zone. The fescue-tussock grasslands of the Mackenzie Country differ, not un­expectedly, from those described by Barker. Poa caespilosa at Hunters Hills was present often at high abundance/dominance values. In the Mackenzie it is found mostly on river beds and on slopes recovering from very serious depletion in cover. Anthoxan/hll1l1 odoraflllll was of importance in Barker's low-tussock alliance, but in the Mackenzie the abundance/dominance value never exceeded + in the releves presented, nor did the Presence Class (III) approach that in the Hunter Hills (V). Nor was Discaria /Ollllla/Oll in the Mackenzie such a feature, floristically or physiognomically, that a phyto­sociological unit would be recognised. The examination of the two sets could be extended by a comparison species for species. Many species (the most common of the widespread tllssock ~ra.rsland species excepted) would ditTer in one direction or another. The grasslands described here and those of Barker belong to the one community-type, though this may not be her rather restricted "Pesfllca-Poa Alliance".

IV' eed C01l1mtll1i/ies

l1lC~ short-tussock grassland is interrupted by patches, often quite large. dominated by naturalised weed species. Some of these stands are perhaps only one stage up from riverbed colonisation, though most seem to result from rabbit depredation. At all sites the soils were shallow. In Table 2 details are presented f rom four stands examined. Rllmex ace/oul/a was abundant in 3 relevcs (63084, 63086, and 63151). Species accompanying R au/osella w~re not numerous, an~ only oc~as.ional1y attained significant abundance/dommance scores-Raolllt.: hookerl m two rei eves, and Aira rJr'1opbrll~a in one. The species lists in the It are/osella·dominated stands differ from those of Moore (1954) from Molesworth. Droll111J /ec/ortllll. locally abundant in the Mackenzie Country, dominated the stand for rclcve r,;08~.

The stands detailed in Table 2 arc related to those listed in Phase A of Table I, particularly in that all the shor/.ftlJJork. Kr.usland species typical of Phase A are present. l1le step from a weed community to an incipient fescue-tussock grassland is not great. One area examined contained young vigorous plants of teJ/tlfa 'lOt·.u-%tlandia~. many growin.'t in mats of UaOll/i.l bookeri. The sample was composed of 27 species. but only .-lira (dryofJIJyllea :\.4, Villpia !Jr011loiJes 2.3. Raolllia hookeri 2.3, IlJllllex aCt!foul/a 2.3,

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Carex breviculmis 1.1, and Geranium !essiliflorl1l11 1.1 scored significantly. The remaining 21 species, including Fes/llca novae-zelandiae, were scored + or r. Poa eolensoi was not present. This sample represents a rabbit­induced weed community evolving again towards fescue-tussock grassland.

RED-TuSSOCK GRASSLAND

Red-tussock grassland, where Chionoch/oa rtlbra is physiognomically im­portant, is widespread in the Mackenzie Country and was even more wide­spread in the immediate past. In much of this grassland C. 'libra is reduced in density (depleted), and samples of dense red-tussock stands are not readily available for study. Table 3 presents the data collected for 17 reI eves, all from stands at elevations between 2,700 and 3,000 ft in the morainic country between Lakes Tekaro and Pukaki. Table 3 is arranged in order of decreasing quantities 0 red-tussock, from two dense stands to fescue-tussock grassland with isolated red-tussock scattered erratically through it. In both dense stands examined (releves 63130 and 63144) Chionoeh/oa rubra scored 5.5. Additional species never attained high abundance/domin­ance scores except for Agroslis lenllis in 63144. By comparison with red. tussock grassland in other parts of Canterbury red-I/I!sock grassland species were few, although a further five species considered characteristic of red­tussock grassland elsewhere were present in rei eve 63144, 1Iiz. Anis%me aroma/iea var. flabellifolia, Carex eoriaeea, Halo-ragis depreJIa, llln""s effllsus. and Schoenus pallciflorlls. Only a few widespread /Imock grass/and species occurred in these two stands.

Transformed Red-Tussock Grassland

A striking change in floristic composition occurs when the red-tussock grassland community is opened up. The number of species quickly increases, vide the group of rei eves 63131-61021 in Table 3, and shor/-//mock grau­land and snow-lllSSoek grassland species occur in the community, together with many widespread IImock grass/and species. Where red-tussock is even further reduced to an abundance/dominance of + or r, a fescue-tussock grassland is produced. This is seen in Table 3 in the group of reI eves 61022-63132. In three places, "llalmoral", near Irishman Creek, and near Mt Cox, the sequence was followed in detail. At "llalmoral" rcleves 63130, 63131, 63128, 63129, 63132, 63133, and 63134 are such a series. Releve 63130 is from a dense red-tussock stand. Releve 63131 was sited a few yards away and here C. ",bra was at 3.3. At this state the light-demanding Poa eolensoi-absent in 63130-scored 3.4, and the mat plant Raolllia slIbsericea. Fes/llra novae-zelandiae, and Geranirlm Jessiliflorll111 all reached a score of 1. Further reduction in abundance/dominance of C. rtlbra to 2.2 (rei eve

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63128), indicates additional changes in the community. Here Fesluc.a novae-zelandiae was more abundant than Poa co/enso;, and HypochoerlS radicala appeared to be reaching its highest abundance/dominance. The four remaining releves (63129, 63132, 63133, and 63134) are from fescue-tussock grassland, but traces of C. rubra still occur. I~ rei eve 63129 Poa colensoi and Fes/llca novae-zelandiae scored equally In abun~ance/ dominance, and RII1l1eX ace/oulla reached the highest abundance/dommance value accorded it in this stage of community development. Releve 63132 tends to follow on from releve 63131, for Poa colensoi exceeds F. novae­zelandiae in abundance/dominance. Two releves (63133 and 63134) were recorded from stands some distance away. Both are floristically similar despite aspect differences, and represent well developed fescue-tussock grassland.

Two further stands were examined at "Balmoral" on Old Man. Releve 63142 was from red-tussock grassland on a steep slo~e and. is one of ~he easternmost stands of red-tussock in the MackenZIe Basm. AlongsIde it releve 63143 was recorded from what is now a fescue-tussock grassland. The pattern of change here is the same as on other parts of "Balmoral", but the slope is steeper.

The sequence examined near Irishman Creek (rei eves 63144, 63146, 63145, 63147, 63148) was a series upslope from a basin in the moraines. Simi­larly near Mt Cox (releves 61021 and 61023), further inland from the "Dalmoral" and Irishman Creek stands, in red-tussock grassland where C. rllbra was 3'3 and 2'2, widespread IllSSock grassland species are numerous and abundant. Releve 61022, very close to 61021, was a fescue-tussock grassland containing more Dim tria IOtlmalou than any other stand examined in the area.

In other parts of Canterbury many red-tllSJock grassland species persist into the fescue-tussock grassland stage, but this is not so in the Mackenzie Country. Some fescue-tussock grassland sites listed in Table 1 may have developed from red-tussock grassland, but this cannot be determined on a floristic basis. Percentage presence of species found in depleted red-tussock grassland. in fescue-tussock grassland derived from a former red-tussock grassland. and in fescue-tussock grassland are given in Table 4 for those species showing significant differences. Species below Presence Class II have been excluded.

f!ichelachne crinita occurs most frequently in fescue-tussock grassland, while both Carex 11111e/leri and P'Irrhanlhera exiglla are at their highest in fescue-t~ssock grassla~d derived from a former red-tussock grassland, For 13 species the t~end IS t.owards .lower percentage presence in fescue-tussock grassland; for nme specIes the mverse holds; and for three species two are at their highest in fescue-tussock ~rassland formerly red-tussock and one is at its lowest in that same class. While these trends indicate the selectivity of these species, consideration must be given to their abundance/dominance in the three states listed above. Four species, A~rostis tenllis, AnthoxanlhllfJt odora/um, Acaena caesiiglallca, and Vitladinia al/stralis, had higher abundance/dominance scores in fescue-tussock grassland than in the other two communities.

Some few mow-llIJsock ~rassland species are present in the opened-up

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red·tussock grasslands. Three species, Brdrhyrollle sindairii. Adphylla allred, and Lagenophora rlmeala, attained Presence Oass IV, III, and III respec. tively, but the abundance/dominance scores were never high. CbiollorUoa riXida was present at four sites examined.

SNow·TuSSOCK GRASSLAND

Snow.tussock, Chionorhlod riXida, ,is present on all the mountain ranges studied. At high altitudes an alpine C. rigida/Poa colemoi grassland occurs. Below this a C. rigida/Feslllr(J novae·zeldl1dide grassland is found on the castern and southern ranges, predominantly in the drier regions where the desiccating north·west winds have their strongest efTect, while a C. r;g,ida/ F. mallht>Wsii grassland occurs on the western ranges in the higher rainfall lone on land of predominantly shaded aspect. A C. f'igida/F. l1ot·ae·zelandiae grassland is often found on the slopes below the C. ri!,ida/F. mal/hewsi; community of the western ranges. In the northern part of the Den Ohau Range this sequence of communities is illustrated by: (i) alpine C. rif.ida/ PO:l colemoi rcleves (63112, 63113); (ii) C. rif.ida/F. mal/hewsii grassland (relevi's 63114, 63111, 63119, 63120, 63121); (iii) C. ri!,ida grassland with both F. mallhelllJii and F. not·ae-zelandiae (releve 63118); (Ill) C. rif.ida/F. notlae-zelandiae (releve 63116); and (1/) a fescue-tussock grassland at the foot of the slope •.

Alpine Chiollorb/oa rigida/Poa (olemo; Grassland Two snow·tussock stands were examined in the alpine wne. These are reported in rei eves 63112 and 63113 (Table S). Both were on the Den Ohau Range above the head of Lake Pukaki, upslope from the C. ri.r:ida/ F. lIIallhewsii stands 63111 and 63114. POll colemol was the chief associate grass and Raoltlia species provided some cover. In reI eve 63112 Pmttl (bond-fa /7","ila and Celmisia Iyalli; both show significant cover, but not in the denser snow-tussock stand of rcleve 63113. Difference in aspect between these two rei eves may be sufficient to cause these diJferences, but rei eve 63113 was from an isolated stand of tussock between two I.ugc screes which descended far below it; it appeared to be little grazed. Stands such as these are common on mountain slopes. These two releves can be compared with 61017 and 61004 from stands at 4,(,00 ft on the eastern ranges. Although both 61017 and 61004 are from stands of southerly aspect there are some similarities, as here, too, Poa (o/msoi is the chief associate grass, and the mow-tllssock ~rassland and widespread tllSJor/! f.raJII(J11d spuies are much the same. In both places Feslllrtl species were of low abundance/domainance. This was also the case in inland North Otago (Connor, 1961).

·n1(~ main species here were: AI.'Ollis '",lIis ~.4. FrIlII(a nOl'a,·zelallJia, 2.3, C:raliJoJn f,au,i 3'~1 COI"O.ll1hl pe/,iei 2.3, PI',""II)'" n,lII.1 2.3. lIi,.,ar;lIlII /"dea/IIIIII 1.3, IIJ/'or/Jo"rh ,./lairala 1.'. G"tII;.lIIa (orymbi!"ra 1.1, SI.lckholtJid "';11;",.1 1.3 Cllulila "/1ilh)'1IlNIII 1.3: 35 othef 5J'('cies scored + or f wefe nlsn pre5ent. . ,

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Cbio/l0ch/oa ri"ida/Feslllca novae-zelandiae Grassland

The 20 rei eves from Chioltochloa rigida/Feslllca nova~-ze!a~diae gr~ssl~~d presented in Table 5 show marked variability, but agam It IS a vart~bJltty that can be ordered. Cbionocb/oa ri"ida and either P. novae-zel~nd~ae or Poa colemoi are the important species and make th~ largest ~on~rtbutlOn to the cover. From place to place Rao~"ia SIIbsericea tocreas~s. to Importance, especially when the abundance/domtoance scores for C. rtg/da are low.

Five phases are recognised. Within each there is some v.ariability and individual rei eves are often quite difficult to place, and, as to the fescue­tussock grassland, some rei eves are the connecting links between the phases. Thi~ grouping of the rei eves into phases is somewhat arbitrary, but the variation IS not altogether continuous and some phases are clearly recog­nisable. Phases A to 0 are from the eastern ranges and Phase E is from the western ones.

PHASE A. Releves 61017, 61004, 61005, 61007: from country of shaded aspect, primarily in the lower alpine and upper subalpine zones. Chionochloa ,if/da and Poa co/emoi are the main grasses. Acip!Jy/la 11I0nro; and Agroslis sldJII/aJa, and a relatively short list of widespread IIIssock grassland species (but one which includes the omnipresent plants LtlZlda campes/ris, Viola CIIntlin"bamii, If/ ab/enber"ia albolllar"inala, etc.), define this phase. Short­tussock grassland species are rarely recorded.

PHASE n. Releves 61011, 61014, 61015, 61020: from country of shaded aspect primarily of the upper subalpine zone. Similar to Phase A but lacking two or three high altitude species and with more widespread IlIssock grass­/and species includin~ Craspedia Imiflora, Genliana corymbifera, and An/hox­anl/JlIIII odorallllli. Peslllca novae-ze/andiae and Poa co/ensoi are extensive.

PHASE C. ReJevcs 61019, 61002, 61009, 61018, 61006, 61003: from country of sunny aspect, primarily in the upper subalpine and lower alpine zone where much bare ground and unstable subangular gravel are present. Abundance/dominance values for C. rigida are low; there are relatively few snoW-tlISSOC!'? J!,rassland species, while either Poa (olemoi or F. not'ae­rtlandiat is the main subsidiary grass. Although the altitudinal range is such that some high altitude species, e.g., CelmiJia lyalli or Aripbylla 1I10nroi could occur, these are not prescnt. Snow-Illssock r,rassland species are fewer than in the preceding phases. Senecio ,"Iallcophylllls ssp. discoidetls is present and is an indicator of the amount of detritus on the slopes, while fI iuadinid. allS/ra/is and Carmichaelia pelriei indicate sunny aspects though these specIes are higher upslope than usual. The distribution of widespread 1IIJS0ck "rass/dnd species is similar to that in Phase A which is from com­parable altitudes.

PHASE D. Relevcs 61013, 61010, 61001, 61016, 63140: from the montane lone. Snow-Illssork grassland spteitS are few at low altitudes. ShorJ.lIIssock "rass/and species are present, e.g., Cop,osllla ptlriei. Diche/admt crinila. and Aira rar),ophyllta, and there are more widespread ///Ssock "rass/and JPtcitJ. The allinities with fescue-tussock grassland are somewhat grcater than in the prcceding phases.

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PIIASE E. Releves 63095, 63097, 63116: from country of sunny aspect, primarily in the montane lone on the western mountains. Very sImilar to Phase D except that .rhorl-/llSIod! gras.rlal1d .rpede.r are poorly represented. ll1ese three releves are from grassland near a C. rigida/F. matthew.rii com­munity. The floristic interrelation among these five phases is reflected in different ways and is emphasised by the arrangement in Table 5. Differing altitudes and aspects accentuate the floristic differences. The lower alpine reI eves of Phase A are closely related to the upper subalpine releves of Phase 13. Aspect similarities here seem more important than differences in altitude. The relationship between the reI eves of Phases A and C, which are from much the same altitudinal lone, is not as strong as might have been expected. Aspect differences again are here more important, and have pro­duced a different phase of the snow-tussock grassland. Phase 0 from the montane zone differs radically from Phase A. and the change in species composition is associated with altitudinal differences. Further comparison ~mong the five C. rigiJa/F. not'ae-ze/alldiae phases reveals interplays of various kinds. most of them concerned with altitudinal differences. For example. the differences between Phases D and D and between Band E are of much the same order as when Phase A is compared with Phases D and E. Although these five phases can be viewed as distinct parts of the one community the differences among them are unequal. Releve 63118 is intermediate between the C. ri{ida/F. nOlJae-zelaI1Jiat' grassland and the C. rif.ida/F. mal/hew.rii grassland to be discussed below, Here at "Glentanner" on the Ben Ohau Range both F. not'ae-ze/a1Idiae and F. mal/hew.rii were present. At higher altitudes only F. lIIal/h~wsii was present. and downslope only F, not'ae-ze/al1diae (releve 63116). Relev6 63118 is clearly from an interphase between these two snow-tussock grass­land communities. Barker (l9H) described a "Danlhonia-Ce/misia Alliance (Tall Tus­sock)". which she con~idered consisted of associations, which (perhaps wisely) she declined to define. The community discussed contained Ce/lllisia spt(lahilis at high abundance/dominance scores, while Fes/llfa not'at­ze/andiat and Poa fo/emo; were the chief associated grasses. The naturalised Al1Ihoxal1/I1IlIIl odorallllll was sometimes quite important. The grassl.tnd of Barker is floristically different from the C. riKiJa/F, notlde-ulalldiae Jtrassland described here, even allowing for difference in altitude and climate. However it may be unwise to attempt a fuller comparison with 13arker's records as all but five of the 15 relevcs she presented were from an area that had been burned less than a year and a half before examination. The illustrations and tabulations of 13arker indicate a run-down phase of a Chionocbloa riKiJa grassland. and the abundance of Celmi.ria spurahilis is probably quite long-standing in terms of European occupation. Celmisia .rperlahilis does not entcr the Mackenzie Country. but because of its wide extent of the seaward side of the surrounding northern and eastern ranges. and because C. dellsif/ora may in time show a similar behaviour pattern, attention was paid to the distribution of other species of Celmi.ria. Only the ubiquitous C. Kraci/mla was recorded often. Celmisia fyallii and C. bre,-i/o/ia

·n.lfllho"ia {lal'f'IU"1 of her lext i, Chio"nrh/na ,ill.iJ".

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was observed at higher altitudes only. On rocks on the slopes of ~1t. Dalgety at 4,150 ft C. demijlora, C. angtuli/olia (and possible hybr.l.ds betwee~ them) were growing with Chionoch/oa rigida, Pimelea Iraverw, an~ Hell­chrjslll" selago. In the same vicinity at 4,500 ft on a deep fine s~11 on a southerly facmg slope in a localised area C. brevi/olia cx;curred m large patches as does C. spectabilis in other areas. The foHowmg were scored, here: Celmisia bret:i/olia 4.4, Chionochloa rigida 2.2, Festllca novae-zela~diae 1.1, Poa cO/e1Isoi 1.1. On the Grampian Mountains at 4,100 ft C. densylo~a was recorded on rock outcrops, whife at 4,800 ft on dry rocks C. brevl/olla had the same distribution pattern as on the slopes to Mt. ~algety. N?ne of these Ce/misia occurrences is comparable to the extensive C010O1es of C. speclabilis found outside the Mackenzie Basin.

Chiol1och/oa rigida/ Feslllca matthewsii Grasslal1d

Eleven releves are presented in Table 5, from C. rigida/F. matthewsii grasslands at altitudes ranging from 1,800 ft to 4,600 ft; all are from stands on the Ben Ohau Range. Three phases are recognised:

PUASE I~. Releves 63096, 63098: from subalpine stands of north-westerly aspect. Snow-tussock grassland with Fes/llca mal/hewsii and Poa colensoi. adjacent to C. rigida/F. not!ae-ze/andiae grassland. Species typical of C. rigiJa/F. mal/hewsii grassland, apart from F. 1J1allbewsii, are not represented. This phase shows some interaction with the C. rigida/F. novae­ulalldiae grassland close by, though it is not clearly as transitional as is releve 6.H 18.

PHASE G. RelevCs 63100, 63106, 63107, 63108: from the montane zone on country of shaded aspect. Low altitude C. rigida/F. mal/bewsii grassland in which Angelka deClpie1ls, Ranlllull/'IS Kracilipes, Anisolome aromalica var. ;'I(isa are additional snow-IIlSIock Krass/and species, and Blecbnll1l1 ftl1l1a-marina and Afyoso/is allslraiis of the lal/-IIlSIock grassland spt'des are Important. Some sborl-/llSSock Krass/and species. e.g., Carex breviCli/mis and C. co/msoi. are present. The widespread "lJSo(k grass/and species present are much the same as in low altitude C. riKidalF. nOllae-ze/andiae grassland.

PHASE H. Releves 63111, 63114, 63119, 63120, 63121: from the sub­alpine ~one in higher rainfall area than Phases F and G. Snow-tussock grass­land wIth F. mal/hewsii and Poa cO/ef1soi as the chief smaller grasses. The snow,-t,!ssoc.k grassland species present are comparable with Phase B of the C. "Kidall-. I~ot·ae-%e/an~ia.e grassland, but Celmisia demijlora and Penta­cbol1dra .PIII/llia are addItions. Among the fal/-tllssock Krassland species, Galli/herM depressd, Craspedia /allala, and Bledmll1l1 pel11la-lIIarina are often prescn.t i!l signi~cant quantities. The widespread IIISIock rrass/alld sft'(ies are dlstrtbuted In much the same way as in Phase Bothe C. "Kidal F. 1I0l'dt-u/alldiae community.

TIl ere is variability in the C. rigidalJl. lIIallbetusii grassland just as there is in the C. riKidd/l-'. flot'at-ulandiae community, both between and within the phases. Major floristic differences between Phases G and H lie in

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(i) the presence of a few shorl-Illssock Krass/and species in Phase G, (ii) the presence in Phase G of wideJpread I/mock Krass/dlld species common to low altitude stands elsewhere, (iii) more snow-I/lJJock Krass/and spefies in Phase H, and (ill) higher abundance/dominance values for IdI/-I/lJJock Krass/and species in Phase H. These differences are chiefly associated with altitude, and in general the influence of altitude here is parallel to that in C. rigida/F. llot1ae·zelandiae grassland on the eastern ranges.

Four samples were taken from stands on the western slopes of the Ben Ohau Range near Greta Stream. These releves (63095, 63096, 63097, and 63098) show the interaction of altitude and aspect with the occurrence of either P. l1ol'ae·ze/andide or Po lJIal/hewsii, as was recorded in North Otago (Connor, 19(1), In stands on slopes with northerly aspect F, tlOl'ae· zeldl1diae was present at 3,100 ft (releve 63095) and at 3,400 ft (rcleve 63097). In two stands, one at 3,300 ft on a westerly facing slope (releve 63096) the other at 3,MO ft also of westerly aspect (releve 63098), P. lIIallbewsii was the important species of Fes/llca, though Po "Ot'cle­zelalldiae was prescnt to a lesser extent. By contrast with these sunny western slopes, f1. lIIallhewsii was rresent at 1,800 ft on a slope of shaded (southerly) aspect on the eastern SIde of the range (re1eve 63100),

The C, rigida/F, 1IIa1lbewsii grassland in the Mackenzie agrees well with that from stands on the Barrier Range on the southern side of Lake Ohau, and those from further south around Lindis Pass and Ahuriri River.

Trtllls/ormed Snow-TllJJock Grasslands

Cockayne (1928) indicated that snow·tussock grassland may be trans· formed into short·tussock grassland by burning, This change is accelerated by grazing after the burn. In Table (, eight releves are presented from stands of two kinds of tall·tussock grassland which have become short· tussock grassland, Site descriptions for these relevcs are presented in detail in Appendix I.

All the rcleves disrussed here are from stands either well within the range of the C. rigiJa communities. or where C. ri!!,ida was at least sparselY' present (e.g., relevcs 63093 and (3138). No other criterion has been used.

As the community is opened up by the reduction in number and canopy spread of C, ri!!,iJa, Iight.demandlng species such as Peslllca not'ae·uland/a, and I'oa (o/ensoi increase appreciably 10 abundance/dominance, so too does Uaolilia SldJSericea, and at rower altitudes and in appropriate sites Diu.trid 101llllaloll, There is a general increase in the number of species. but the increases in abundance/dominance of many of them are often more import. ant. The eventual destruction of snow·tussock communities lead to grasslands of the types detailed in Table (I. Snow·II/HOck ![yass/,mJ spuies and the group of species characteristic of C. ri!!,ida/F, IIIdlliJeU!sii community arc still present. "al/.II/Hock !!,r,mland sprcies. which were often extensive in the C. ri!!,iJ.,/P. mlllll,ewsii grassland also still occur. In the grassland derived from a C. 'f';Xitla/ p, no".,t·zeldfldi.,t community, shorl.tllssock Xraulcllld spuies arc more numerous than in those that derived from C. rixidal

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F. mallhewsii grassland, primarily because most releves were from stands in the montane zone. The formation of short.tussock grassland from the C. rigidd/F. malthewsii community is nowhere as extensive as that formed from C. ri"ida/F. novae·zelandiae grassland.

Table 6 gives the percentage presence of the species there listed,. together with the percentage presence in montane C. rigida/F. novae·zelandtae grass· land (5 rei eves) and f escue·tussock gra~sland (15 releve~, the ~a,?e ~s us~d in the red·tussock grassland compaCisons). For speCIes orlgmatmg 10

C. rigidd/ Po mallbewsii grassland, percentage presence in 10 releves from Table 5 (releve 63114 is excluded) is given. Any trends are probably over· emphasised by the small number of samples presented.

HISTORY OF TIlE GRASSLANDS

TIle observations reported here permit a more precise outline of the history of the grasslanas of the Mackenzie than has been ~iven by earlier authors, though much in their theories is confirmed. In part~cular, emphasis is placed on the extent of former tall·tussock grasslands whICh have largely been destroyed.

Pre.Ellropean Grass/ands

In a recent paper (Molloy el al., 1963) it is stated that "Surface logs, buried wood and charcoal, wind·throw dim(?les and buried pod sols provide convincing evidence of a former widespreaa forest cover over now.treeless tracts of the South Island". The Mackenzie Basin is one of these tracts, now treeless except for beech (Nolbo/a8'Is) forest in some wetter western valleys. Most of the subfossil remams from the Mackenzie Country are podocarp charcoal, but beech charcoal and buried podsols are recorded near Lakes Pukaki and Ohau. Charcoal and uncharred seed cases of matai were found in soils on rolling moraine and terrace land near Lake Tekapo. A sample of totara wood from the Dalgety Range had a radiocarbon date of 1450 ± 70 years before 19~0 (Fergusson and Rafter, 19~7), which is 500 to 700 years older than the general run of dated old logs and charcoal from South Island tussock grassland.

TIle forest probably reached an altitude of 4,000 to 4,500 ft and there gave way to snow·tussock grassland dominated by C. rigida and followed at still higher ~ltitudcs br other alpine associations. Within the forest :lone one can envIsage smal enclaves of both red· and snow·tussock grassl.lOd, while rock ~utcrops, river. beds, and recently developed fans would all sup.port 5.pCCICS now found an tussock grassland. Near rIvers and particularly beSide tributary streams red·tussocks would flourish. A short.tussock grass· land would occur on some flood plains.

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eXPLANATIONS OF SYMBOLS FOR ALL TABLES

Estimates of ABUNDANCE/DoMINANCE for each species arc given as a

single value according to the following scale:

r = of erratic occurrence. + = sparsely or very sparsely present, cover very small.

1 = plentiful but of small cover value.

2 - very numerous or covering at least %0 of the area.

3 = any number of individuals covering 1 to i of the area.

4 = any number of individuals covering! to i of the area.

, = covering more than i of the area. D ::::: indicates occurrence outside plot but within the stand.

The SOCIABILlTY of each species is estimated-

1 ::::: growing one in a place, singly.

2 = grouped or tufted.

3 == in troops, small patches, or rushions. 4 ::::: in small colonies, in extensive patches, or forming carpets.

, - in great crowds.

In the records the abundance/dominance value is followed by the sociability figure.

PRESENCE CLASS where calculated is expresed in the followinB scale:

Class 1 species present in 1-20% of the rcleves.

II species present in 20-40% of the releves.

III species present in 40-60% of the releves.

IV species present in 60-80% of the releves.

V species present in 80-100% of the rcleves.

LIFE FORMS

N Nanophanerophyte Cil Chamaephyte H Hemicryptophyte

*Naturalised Species.

G Geophyte Th Therophyte D

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1964] CONNOR - TUSSOCK GRASSLANDS 341

The forest seems to have been destroyed mostly by fire, and probably one of the first new plant communities to evolve would have been a short­tussock grassland, derived from the discrete communities within the forest or from the alpine zone. This would contain, contributed from the alpine lone, POd (olmsoi, Feslllea mallhewsii in appropriate places, Deyellxia at'mo;des, D. lorsleri, and perhaps a high altituJe "race" of Feslllea not'ae­zelandiae (Connor, 1960b; Scott, 1961) with Bra(h),come sinclairii, LaJ:eI10-phora (linea/a, and some species of EpiiobilllJl. In the montane zone Pes/llca 1IOt'ae-zdandiae, AJ:ropyron scabr/lll/, Dichelaehne (fini/a, ErJlhranlbt·ra pIIIJlila. PJrrhanlhera exiJ:lla, Geranillm sessi/ifi()rtllII, and species of Carex. Uaoll/ia and Celmisia would be early colonisers. The succession here need not have passed through all the stages listed by Cockayne (1928) and Calder (1961) unless catastrophic landscape changes followed forest destruc­tion. The outwash plain appears to have remained in short-tussock grass­land (called Steppe by Zotov, 1938), now much deteriorated.

During the relatively quick formation of short-tussock grassland in the montane and subalpine zones, Chionoch/oa riJ:ida would migrate from the alpine zone down the mountain slopes into recently formed short-tussock grassland. Uecause of its slow growth and irregular seeding it would br entering a somewhat mature short-tussock grassland. Certain other specie' would spread downslope from the alpine zone, but the new communiti('9 would not be replicas of the alpine snow-tussock grassland. As the snow­tussocks increased in number and in the size of the canopies, li~ht-demanding short-tussock grassland species would be progressively eliminated, until a snow-tussock grassland had replaced the short-tussock grassland.

There was no barrier to the downslope migration of snow-tussock. Chiol1ochloa ri;,idd is there to the foot of the mountain slopes today, as it is in the remainder of the South Island, though often only in small groups or as isolated plants.

The time from forest destruction to a close-canopied snow-tussock com munity might have been about 200 years. This estimate is well within the time from forest destruction (a thousand years or more before 1950) to the arrival in the Mackenzie of European sheep-farming colonists in 1857. No pre-European dense-canopied stands remain and their structure a'ld com­position may only be inferred. Any dense stand found today haw without doubt been burnt, possibly many times.

Some features of C. riJ:ida plants need comment. Chiolloch/oa ri~ida does not flower abundantly every year, but in ~ood lIowering years larg'! numbers of seed heads are produced; as in other Indigenous grasses adequate seed is set; seed-set figures are given in Connor (196Oa ) from some fiela collections and from plants in cultivation. The seed from the alpine zone would, in good Ilowering years, be sufficient to permit C. ri,Jda to make an appreciable move­ment into any new zone. l:lorets with the small caryopses arc usily wind dispersed, so that migration need not be only by marginal advance. Further, C. ri~idd flowers heavily in otherwise poor /lowering years whenever the tussocks have been burned in a previous season (Uarker, 1953, O'Connor and Powell. 1963). and any pre-European grass fires would assist this.

Judged solely by the rate of growth in experimental gardens ChiollocMoa riJ:ida is a very slow growing plant. Five years from seed to the first

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342 NEW ZEALAND JOURNAL Of' BOTANY [Orc.

production of flowering stems is an average based on singte,sp.lrcd pl.anl5 grown in the Hotany Oivision experimental garJens at Linwln. and pJ.anb twice as old are of the size of mature plants in the field. In nature the rate of growth is very much slower. While Chiollorh/od rigiJd was the important tall·tusock on the mountain slopes, evidence points to the importance of CNOI/Orb/od r,",rd (red.IUS5(xk) on the moraines. Morainic country is extensive, and passes bark towards the glacial regions at the heads of the lakes. Near Lake Pukaki red·lussockl\ occur along the sides of streams but none was found in the morainic country between the Lake and Simons Pass. Today the easternmost nlcnsion of red·tussock in the "Halmoral - Irishman Crc:ck" area is the line of the f.:nd of the moraines. It is therefore suggested that most. though possibly not all, morainic country between the Two 111umb Range and Lake l'uk.lki; fasman River was dominateJ by C. r"brd. This tall red-tussock gra.ssland probably evolved in much the same: way as did lhe montane and subalpine snow·tussO<.k grassland, i.e., by invading a first formed short·tu~S()(k gr.lSS· land. Some dilferent species may have been involved here. and the procrss was perhaps faster than on the mountain slopes, aided by an annual seeding in red·tussock. The nuclei from whkh this extensive red·tu5S(xk grassland was formcJ would he found not only in small basins of imperfect drainage within the forest and beside streams, but also in stands of C. ,,,b," on shoulders and Hat places often above 3,000 ft, and possibly from above the timber·line. CI.Jiollo(/JIoa ,libra has a wide ecological amplitude and dccs not eschew steep slo~es or dry places that arc at least seasonally ..... et. It ..... ould become the dommant species on the rolling moraines because of the range of habitats available to it in this higher rainfall are" \\hi(h is also prone to long.lying snow. As was indicated above, some few snow-tussocks were: found among the moraines between Lakes Tekapo and Pukaki ..... here no other tall·tussocks were present, and a (ew snow-tussoc::ks were found in red·tussock grassland and the fescue·tussoc::k gr.lssl.lnd derived from it. These occurrences are not unknown in tussoc::k.gruslanJs throughout Soulh Island, and most appear to represent a recent migration by this species into sites now available to it through r(ccnt (hanges in the veget.1tion, though some are also obviously ecotonal. If snow·tussock had C\'er been widespread on the moraines in the Mackenzie Country it ..... ould have lltttl on the drier moraines. Evidence in support of uU·tusso(k grassland on the slopes and on the moraines is afforded by the present occurrence of these species there­C. rigiJa to the (oot of the slopes and C. rll/Jrd to the edge of the dumpell moraine. So far, Ihen. it IS possible to suggest that tall·tussock grassland would ~ the prc-European state of this pm of the Mackenzie Country. in keeping with the pattern of tall·tussoc:k distribution in most in!'lnd basins in South hlanlf. In the past C. ,,,/I'd could prob.lbly have extended out beyond the moraines on to the outwa!\h plains, (or rcd·tuS5oc:k can p~rsist on (rcc-draining soils as well as on those with impcr(e<l drain.1gc. Uut the stony shallow soils--thco majorily of the outwa~h plain-- arc o\'crdraincd and this would militate against mudl red.tussock grassland in :l subhumid dim.llc. No rcd·tussoc:ks, nor any r,·J-IIiSlod: r.rdss/,IIIJ spuiu (and the!\e

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1961] CONNOR - TUSSOCK GRASSLANDS 343

are few in the Mackenzie Country). have been found on the plains. Some snow.tussocks occur on the plains however, but only very close to existing snow.tussock stands. e.g., at Haldon Corner and on the terraces of the OllJU River close to the southern end of the Uen Ohau Range. It lIcems unlikely that they also were ever extensive on the outwash plains.

Indl/red Shor/·r/mork Grassland

The formation of fescue· tussock grassland from both red·tussock and ::now·tussock grasslands has been presented above (pp. 332 and 333). The precise nature of such changes over broad areas in the past may only be mferred, since no documentary records are available. Dunnage's HJ~~ comment that "The feed was poor but would improve with burning" (Gillespie op. (il.) would seem to apply to tall.tussock, for, entering the Mackenzie through Uurkes Pass, he must have traversed extensive areas of ~now.tussock grassland (still present today) before he arrived on to the plains and reached Lake Tekapo. The pastoralists were confronted with much grassland unsuited to sheep grazing, and following the experience and practices of earlier settlers nearer the coast the use of fire was natural. Burning of the grasslands here, as elsewhere, became the accepted pro· redure. They heavily stocked the burnt land.

Sheep were first introduced into the Mackenzie Basin in 1857 and by I H95 were estimated at 270,000. It is difficult to find a satisfactory record of sheer. numbers before 1895, but the sheep numbers for the Mackenzie County from 18R4-95 given in the Statistics for the Colony of New Zealand suggest Ihat there were never more than 300,000 sheep in the basin even in the early eighteen eighties. Sheep numbers given below were supplied by Mr R. D. Dick of the North Canterbury Catchment Uoard for the period IS95 to 1957, and for 1963 by Mr D. G. Reynolds of the Depart. ment of Agriculture.

No. of Sheep at 30.4.1895 - 270,000

30.4.1906 - 223,000

30.4.1936 - 199,000 30.4.1947 - 173,000

30.6.1957 - 169,000

30.6.1963 - 19·',000. Burning alone, except in severely hot out·of-season fires or where very

persistent, would probably not ploduce a grassland community with a new dominant species, but when burning is associated with heavy grazing many tall·tussocks must die, and then a grassland with new dominants can evolve. O'Connor and Powell (196.\) concluded from burning and cutting r.ludies conducted on the eastern side of Mackenzie Pass at 2,300 ft that Cb/ollo(bloa rit(ida plants would recover from burning to almost full size and vigour in about three to four years, providing there was no grazing. From

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344 NEW ZEALAND JOURNAL OF BOTANY [DEC.

further experiments at higher altitudes (4,000 ft and 5,100 ft) in the Craigieburn Mountains O'Connor (1963) concluded that restoration of the C. ri/!,ida canopy would be a slow process, and even slower at 5,100 rt if the tussocks were very severely defoliated.

Durning followed by heavy stocking has led to the replacement of much tall-tussock grassland by short-tussock grassland. There has been a rdurn to the state at the time of the migration downslope by ChiollO(hlod "i.~iJJ and across the moraines by C. ,.lIbra, with the difference that this tame naturalised species-RIIIJJex ace/Olel/a, Anlhoxalllhlllll oJora/lIlII, lIypo(houiI radiea/a for example-have entered most of the grassland and are now integrated in it, and that subsequent management has maintained it as !l

short-tussock grassland. The era of highest sheep numbers possibly coincided with the final period of the major destruction of montane snow-tussock and red-tussock grasslands and their transformation into fescue-tussock grass­land. Snow·tussocks are heavily grazed if kept short or if burnt; they are preferred to red-tussock or fescue·tussock, and it seems reasonable to suggest that much sheep and wool production came from the grazing of the burnt· snow·tussocks during the redevelopment of the fescue-tussock grasslands.

The change from tall-tussock grassland to a short-tussock grassland more suited to free-ranging sheep was achieved with considerable success on l.trge parts of the Mackenzie Country, but not always without the loss of pro­tective cover in the more critical upper montane and subalpine zones, or in those areas subject to the full blast of the desiccating north· west winds. Not all the subsequent burning to promote succulent new shoots of less palatable grasses has met with the same success, and much depleted vegetation and eroded country has resulted. Short·tussock grassland has been produced on the morames from Chiofloehloa ",bra grassland. On the mountain slopes short· tussock grassland has been produced from C. rigida grassland on al\ extensive scale. On the whole, short-tussock grassland has b:en produced (through the combined etTorts of burning and grazing) up to about 3.000 ± ~oo ft on the eastern ranges and to a somewhat higher elevation on the western ones.

On the eastern ranges, persistent efforts to extend short·tussock grass­land dominated by Pes/lied lIol·ae·zrlandiae further upslope of this altitude have frequently been disastrous. Continual burning foHowed by grazing, and fires running upslop~ to ~arious altitudes, have led in many places to Brassland of the type detailed III releves 61006, (,1003, 61018, and 61009 in fable 5. Here there is much unstable subangular gravel and bare ground and 1'. lIol'a,-ulafldia, does not increase, nor does C. rigiJa. From this zone of weakness, often just above mid·slope and at the boundary bc:twttn the montane and subalpine zones, much erosion both upslope and down has occurred, som.e mergin~ with t.alus from .erosion. higher upslope. Molloy (19(2) has discussed tIllS accretmg zone With speCial rderence to the debris (rom higher altitudes. One furth.er result fro~ t~ese attempts to increase the area of short·tussock grassland IS a new altltudrnal :zonatIon for C. ,.ir.ida. In the presence of forest C. rigida was in the alpine zone-above 4,000--4.500 ft; with the destruction of forest a new lower limit at less than 2,000 (t was established; subsequent to the destruction of much C. rigida a new intermediate limit. about 1,000 ft below that which obtained in the presence

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19641 CoNNOR - TUSSOCK GRASSLANDS

of forest, has been produced. Expressed alternatively, C. ril{ida was formed,. restricted to the alpine zone, today it occupies the subalpIne zone as wei , and in the very recent past it also occupied the montane zone.

On the wetter western ranges where Fes/llt'a not'ae·zelal1diae and F. mal/heU'sii both occur, the results from attempts to extend short-tussock grassland further upslope ditfer from those already described for the eastern ranges. On these ranges P. novae·zelal1diae occurs at lower elevations and P. mal/hewsi; higher up although on shaded aspects the latter may be present at quite low altitudes. Pes/lira mal/hewsi; is not subject to the same limitations on expansion above 3,'00 ± '00 ft as is F. l1ot'ae-zelal1diae. It forms a short·tussock grassland (with Poa rolenJoi in particular) higher upslope (see also Wraight, 1963). Nor is there the strikIng zone of weak. ness described from the eastern ranges of westerly aspect, though it may be there in an incipient stage.

Short·tussock grasslands produced from the destruction of red-tussock communities have been discussed above. Red-tussock plants are more difficult to destroy than are snow·tussocks, and this has led to a mosaic of (i) patches of dense red-tussock grassland at the wetter sites, (ii) stands of thinned out red-tussock grassland with many uidespread IIlJJot'k graJJland speries on some of the slopes and flat terraces, and (iii) stands of fescue-tussock grass­land with traces of red·tussock. In the Mackenzie the formation of short­tussock grassland from red·tussock communities has not been as success­ful as that from snow-tussock grassland on the slopes, although towards the terminal moraines, in a somewhat drier area, the transformation has been more successful.

Much short·tussock grassland in the Mackenzie Country is therefore of fairly recent origin, as the operations responsible for these changes began little more than a century ago.

ACKNOWLEDGM ENTS

I wish to acknowledge with thanks the assistance in the field in 1961 of Mr E. D. Penny, and the particular assistance both in the field and during the preparation of the tables of Mr A. H. MacRae, Botany Division. Mr C. G. Vucetich, Soil Bureau, Department of Scientific and Industrial Research. Christchurch, kindly provided Appendix II and determined the soil :;cts. Many collea~es have helped with critical discussion in the many phases of the preparatIOn of this text and to them latTer grateful thanks. Mr K. R. West, Botany Division, prepared Figure 1.

REFERENCES

UAa"FR. A. P. 1953: An f:colo~ical Study of Tussock Gnml.lnll. Buntcn lIilk N.Z. /)tl'. 1(;. illdNJIr. ReI. nNII. un.

DU.NETt'. T. D. 1927: "Natural History of Cantcrbury." Simpson and William5, Christchurch.

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346 NEW ZEALAND JOt1RNAL OF llOTANY

c'uou, D. M. 19(,1: Plant Ecolo~y of Suball"ine Shin~le Ri\'er-hcd~ in untrrhury. New Zealand. I. Ero'. 49: ~HI-9·f.

COCKAYNIl, L. 192M: "The Ves:etation of New ZealJnll." 2nd t'll. En,l!t'lm.lOn. Lc-iNil'

CoNNOR, II. E. 19(,Oa: Bret'liin,l! Systems in New Zeal.lOtI GraS5e~ III. N .7.. I "It.",. Rei. J: 12H-H.

19Mb: Variation in Leaf Anatomy in reJ/llfd 1101·""-:t'I.,"J,.,I' (I hrk.) Cocka)'ne and f. mallhfu's;; (1I.tck.) Cheesem.ln. N.7. J. S..,. J: .1(.)1 •. '\0" --- 1961: A Tall·tuss(l{k Grassloln,1 Community in New 7.eal.IO,1. Ib,J. oJ' 1125-3'5.

FrR(;l!SSON, G. ).; RAI'TF.JI. T. A. 19~7: New Zeal.IOlI "c A,I!e Me;nuremenh--' N.l. J. Sc;. Tuh. n.J8: 732-49.

GARI'IIlR, n. J. 19~R: The Climate of New ZeaIJnd." Edward Arnold. u,ndnn

GII.I.r.SPIIl, 0 A. 19~R: "South Car.tcrbur)', a Record of Settlement." Suuth Canter· bury Centennial History Committee. Timaru.

GRINI)t.F.Y, G. W.; HARRINGTON. H. ).; \'\'001>, B. L. 19S9: TIlt' Gelllo~ic." M"I' ,., New Zeahlnd. N.Z. 1.1'01. SII',.. nllll. II.r. M.

MAt.tNG. P. B. 195H: "The Torlesse Parers." The Pe~asus Preu. Christchurch. 2\M 1"1'.

Mou.oy. n. P. ).; BITRROWS, C. J.; Cox. J. E.; JOHNSTON, J. A.; \'('AROI.I', P. 19{,\' Distribution of Subfossil Forest RemJim. E:utern South hl.tnll. 1\:ew Zealand. N.l. J. 1101 I: Mi-71.

MonoY, n. P. J. 19(12: Recent Chan,l!es in Mountain Ve~etJtion anll r/T«h lin Ptll~le St.lbility. PrOf. N.? Soc. Soil Sri. ,: 22-(,

MOOR 1', I.. B. 19)4: Some Rllllle)l: (lcf'/oJl'lIoi Communities in New 7.t';lIJnd ,. '~d"',o '-6: 2(,R-711.

O·CONNOR. K. F.; POWFII.. A. ). 19M: Stu,lie~ (In the MJn.l,lo!ement e,f Snllw·tu,,,,,k Gra551,IOII. r. N.l. J. axr;c. R,J. (i: ~H-(,7.

O'CONNOR, K. F. 1%3: Studies (In tltl' MJna~l'ment IIf Sn\lw·lu'~\I(k Gra,dJIltI II Ibid. 6: l('H-n.

N:nTT. D. 19M: iemrcrature Resronses of Nati\'e Short Tuu(l(k, N.?. 1:,',,1. S",. P,oe. 8: 27-211.

S"fH.ll'r. J G. 19M: I..ate Pldstocene historical Gcmnnrrhnlo,lt}, of the 1.llke I\,bki Arca, New Zealand. N.?. /. GI'f1I. G·,.ol,h,·,. (i: IftO-HM.

VON liMn. J. 11197: "TIle Genlo.ItY of the Prll\'inccs IIf ClOterhury nn,1 \X'('~t'.IO.I. New ZeaIJnll." T/;, TIIIII'" Christchurch. 4H(, rr W' ARIlI.r.. P. t 96\: Evolution lind Distribution of Ihl' New Zeal;IO,1 rIot;! II. A/Tcct('\1 h)· Quaternary Climates. N.?. /. n(Jl. I: l-l?

\'('ItAle.IIT, M.) 19(,\: 111(' Alpine IInll Urrcr Montane Gr.ml.IO,I) of the \'('llitJU R,ver Clt(hment. M.ulhorou~h. Il.iJ. I: H 1-7f,.

ZUTOV. V D. 19\11: Survey (If the TU~~(l(k.~r.mIJnJs e,f the &lulh hl.tnd. New Zeal.lnd. N.?. f. Sri. Tnh. }o: 212A-t1A

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Relev~ Nn.

CoNNOR - TUSSOCK GRASSLANDS

ApPENDIX I. SAMPLE SITE LoCAUTIES

(Order of rei eves as in tables)

TAlILE I-Fescue·tussock Grassland

63124 Plains below Mackenzie Pass. lJ} 12 ~ Plains below Mackenzie Pass. 63127 Plains below Hakataramea Pass. lJ}OS7 Outwash plain near Simons Hill. lJ}091 Outwash plain near Simons Hill. 6}09O Slopes on Simons Hill. Oversown and topdressed. 63089 Slopes on Simons Hill. Oversown and topdressed. lJ}101 Plams near Twizel River. lJ} 102 Plains near Twizel River. 63104 Terraces above Ohau River.

347

631O~ Terraces above Ohau River. Set1uio haaJtii here is the form typical of rock outcrops nad snow-tussock grassland in the region.

('3088 Slopes on Simons Hill. Aspect contrast with 63089, 63090.

63092 Slopes on Mary Range. 631 n Morainic downs near Lake Pukaki. Carlllichaelia petrie; important

in this sample from sunny aspect. 631 H Morainic downs near Lake Alexandrina. 63126 Plains below Mackenzie Pass. Compare with releves 63124, 6312~. M 149 Terrace above Tekapo - Forks Rivers junction. MI '0 Terrace above Tekapo - Forks Rivers junction. 63151 Terrace above Tekapo - Forks Rivers junction. (,31 H Terrace above Tekapo - Forks Rivers junction. The 4 rei eves 631·j9

to 63153 were sampled on the series of terraces leading back to the morainic downs.

63154 Morainic terrace "The Wolds". (,J09·~ Morainic downs near Lake Pukaki. Compare with releve 63123 for

efTect of aspect difTerences. M 122 Morainic downs near Simons Pass. (,3139 Morainic terrace near Mt John. 63135 Slopes on Old Man. ('3136 Morainic downs near Lake Alexandrina. Compare with 63137 for

clTcet of aspcet difTerences.

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348 NEW ZEALAND JOURNAL OF llOTANY

TABLE 2-Weed Communities Relev~

Nil. (,3084 Terrace near Tekapo River. 63086 Terrace near Tekapo River. 631' 1 Terrace above Tekapo - Forks junction. near releve 631'2. (,308' Terrace ncar Tekapo River.

TAIILE 3-Red-tussock Grassland Relcvl'

Nu. 63130 Morainic downs "Ualmoral" near Forks Stream. 63144 llasin near Irishman Creek. (,3131 Morainic downs "llalmoral" ncar Forks Stream. (,314(, Morainic clowns ncar Irishman Creek. 63142 Slopes on Old Man. (,t 02} Morainic downs ncar Mt Cox. (,312H Morainic downs "llalmoral" ncar Forks Stream. (,1021 Morainic downs ncar Mt Cox. 61022 Morainic downs ncar Mt Cox. (13129 Morainic downs "llalmoral" ncar Forks Stream. 63133 Morainic downs "llalmoral" ncar Forks Stream. (,31 H Morainic downs "llalmoral" ncar Forks Stream. (,3143 Slopes on Old Man. 6314' Basin ncar Irishman Creek. (,3147 Morainic downs near Irishman Creck. M 14H Morainic downs near Irishman Creck.

rOrC.

M 132 Morainic downs "llalmoral" ncar Forks Stream. Relc:v~ (1}14·I-631.h~ arc a series from the bottom of a basin up to a lIat-torped ridge.

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19641

Releve Nil

61017 61 DO·!

6100~

61007

61011 61014 61015 lilO20 lilO19 61002 61009 61018 61006

61003 61013 61010 61001

61016 li3140 lilO9~

li}097 MI16 MIIR li}096 li309R 63100 63106 6.\107 63108 (,J1I1 l!3114 Mll9 M120 63121 MI12 (,3113

CoNNOR - TUSSOCK GRASSLANDS

TABLE 5-Snow·tussock Grassland

Slopes on MI Edwards. Aspect contrast with 61018. Slopes on Mt Dalgety. Slopes on Mt Dalgety. Aspect contrast with 61003. Grampian Mountains above Hakataramea Pass. Aspect contrast with

61006. Boundary Stream, Richmond Range. Slopes above Godley River.1Not burnt since 1941. Slopes above Godley River(Not grazed since 1950. Slopes on Mt Edwards. Slopes on Mt Edwards. Dalgety Range above Mackenzie Pass. Grampian Mountains above Hakataramea Pass. Slopes on Mt Edwards. Grampian Mountains above Hakataramea Pass. Slopes on Mt Dalgety. Macauley River. Hakataramea Pass. Burkes Pass. Burnt since examination. Slopes on Mt Edwards. Haldon Corner. Burnt since examination. Greta Stream, slopes on Den Ohau. Greta Stream, slopes on Den Ohau. Slopes above Jacks Stream. Slopes above Twin Stream. Greta Stream, slopes to Den Ohau. Greta Stream, slopes to Ben Ohau. Slope above edge of Lake Ohau. Slopes on Den Ohau. Slopes on Den Ohau. Slopes on Ben Ohau. Slopes above Jacks Stream. Slopes above Jacks Stream. Slopes above Twin Stream. Slopes above Twin Stream. Slopes above Twin Stream. Slopes above Jacks Stream. Slopes above Jacks Stream.

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NEW ZEALAND JOURNAL OF UOTANY

TABl.E 6-Fesrue-tussock Grasslands, Derived from Snow-tussock Grassland

Rcleve No.

(DEC.

61012 Uoundary Stream, Richmond Range. CbionorlJ/oa ril!,iJa both above and below stand. 63011 is from an immediately adjacent shaded Slope.

63099 Greta Stream, slopes to Uen Ohau. Immediately below the stand for 63097. Oversown many years ago.

63141 Haldon Corner. Immediately below C. ril!,iJa(P. "o,·ar-:rI.l1Ididr grassland. 63140 on adjacent shaded slope.

63093 Morainic downs near Lake Pukaki. An isolated group of C. ril!,iJ.1 present in the stand.

63138 Morainic downs near Mt John. An isolated group of seven C. ril!,iJa plants occurred within the stand.

63109 Lower slopes to Uen Ohau. Sparse C. 1'if.iJa present. Uelow 63lOli. 63110 Slopes above Jacks Stream. A few C. rif.ida plants present. 6311 ~ Slopes above Jacks Stream.

ApPENDIX II. SOILS

By C. G. VUCllTIC.ll, Soil Uureau, Department of Scientific and Industrial Research, Ouistchurch.

l:ive main groups of soils all formed on grey wacke materials are repre­sented in sample r.lots described in this account. Table 7 shows the soil set names, their claSSification, and moisture classes (based on the 1110rnthwaite method of water balance), as identified with the aid of site notes from the Soil Map of the South Island, New Zealand (in press). Approximate overseas classification equivalents are given.

Most soils have been m<xHfied by processes of accelerated erosion when vegetative cover has been inadequate, doubtless both before as well as dter European settlement. Today the ccmmon process of frost action followed by wind and/or sheet erosion continues in some areas, while in other!> stabilised by plants. topsoils are darkened with humus and show little apparent evidence of an erosion history. DilTerences between north and south facing slopes have sharpened soil differences in some localities, e.g .. Simons Pass (respectively Waltaki and Omarama soils). and on low eastern slopes of 1.1en Ohau Range (Iikewi~e Omarama and Tekoa soils). Ibther lIilrcrent CIrcumstances apply at mldslope levels on the ranges 0.000--4.~OO ft), where stccpland soils once developed under and protC'Cted by (oresl have been modified to resemble the Kaikoura and Benmore soils normally occurring upslope of them, and arc mapped therewith on the 4 ml - 1 in. soil map . .1 t follows that the generalised s?il ,:"a~. ca.nnot ~o .~ull justice to locl1l situatIons, and there may be anomalies 10 SOil typmg of sample rIots. c.g .• several rIots on the steep lower slopes of the Ben Ohau Range I1re identified as Tekoa soil, commonly mapped in midslope positions in some other mountain districts.

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1964J CoNNOR - TUSSOCK GRASSLANDS 3'51

Brolt'II-grey farthI (Waitaki steepland soils) are droughty soils (subxeruus) receiving Insufficient water for efTective leaching, and are chemically fertile. Associated very poreus shallow and stony Mackenzie soils on neighbouring outwash plains are. however. leached and infertile as well as being droughty.

)' e/low-trey ear/hs are represented by the Omarama soils on steep slopes, as near Simons Pass and Hakataramea Pass. These soils do not have the well developed subsoil fragipan as on the Hakataramea Downs, and are considered to be related stcepland soils but with comparable moisture, "/%.

subhygrous. lIiKh.(o/m/ry ),ellow-brou1Il ear/hs are friable and free.draining soils

widely represented throughout the district. Those with subhygrous moisture status are: on the plains the Pukaki soils (moderate depth to gravels) and the more droughty Acheron soils (shallow to gravels). On the morainic downs. as near Balmoral, Tekapo soils on loess capping bouldery till are more retentive of moisture. Those receiving higher rainfall (hygrous) are more strongly leached and with brighter subsoil colours. They are repre· sented by <.ass soils on moraines. and by Tekoa steepland soils on stable midslope sites of glacial detritus as near Twin Stream and Jacks Stream, and by Kaikoura steepland soils at higher levels.

Midslope sites on the Two Thumb Range are classed as the very friable ~nd stony Kaikcura steepland soils, in contrast to the drier Grampians Range where U,nmore soils occur on mid and upper steep slopes and deeper, less siony Kirkliston soils onlhe rolling tops.

N.Z. Snil CI.l"ification

Uruwn·,:rcy carlh~ A"I1(i;IICd ~tony 'nih

lIilo!h counlry yellow. brown Citrth!

TABI.F 7

Soil Sets (in this account)

AIS, Waitaki strrpland Mes. Mdckrnzie

OrS, Omar:lma steepland

Pi, Pukaki; Ac. Achcron

Mr, Mesorotami'l; C~. Cas~

Kill, Kirkliston hill; Rt. Pliketeraki

DmS, Benmore strcpland TkS. Tckoa strcpland KrS. Kaikollra sl('cpland

Moisture O\·ersC.':ls CIJs~ifiealion Rel:ime Equivalrnh

Subxerolls

"

SubhYl:rllus

Jly,!:rous

Non·cnlcic brown soih Not correlated

Grey.brown f'O<,lsolic soils

Acid brown ('arths and brown (';trtlu

Arill brown ("lftll'

---- ----- ---------- _._--- -- - ---------- -

For reference to classification and moisture regime see Taylor and Pohlen (1%2) Soil Bureau Bull. 25, and for soil sct names see "Soils of the South Island. New Zealand," Soil Bureau Bull. 27 (in press).

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