DESCRIPTIONS OF SOME MATURE KAURI FORESTS OF

by Moinuddin Ahmed and John Ogden Department of Botany, University of Auckland, Private Bag 92019, Auckland

ABSTRACT

A quantitative sampling of 25 mature kauri stands, throughout the natural limits, was carried out. Each kauri stand is described in terms of its phytosociological attributes (frequency, density and basal area) for kauri and associated canopy and subcanopy species. A species list of under 10cm dbh is also given with their relative frequencies in each stand. In all stands kauri comprises most of the basal area. It is associated with 10 different co-dominant species. However, most of the forests have a similar species composition. It is suggested that all these kauri forest samples belong to one overall association.

INTRODUCTION The vegetation of various kauri forests has been described by Adam (1889), Cockayne (1908, 1928), Cranwell and Moore (1936), Sexton (1941), Anon (1949) and more recently Barton (1972), Anon (1980) and Ecroyd (1982). A quantitative description of some kauri forests was given by Palmer (1982), Ogden (1983) and Wardle (1984). However, due to extensive past disturbance and milling, most of the above accounts do not describe the natural forest state. Observations on the population dynamics of mature kauri forests were presented by Ahmed and Ogden (1987) and Ogden et al. (1987) while multivariate analyses were performed by Ahmed (1988). However, no comprehensive attempt has yet been made to analyse mature undisturbed kauri forest stands in relation to their species composition. Kauri forests have a restricted distribution in the North Island. Among these remnants, there are few, if any, truly untouched sites. Change to, or removal of, surrounding vegetation has changed not only the surrounding microclimatic conditions but also the denser central parts of most of the virgin stands of kauri. Bearing these points in mind, the paper presents details of 25 study areas mostly sited in undisturbed kauri forests.

MATERIALS AND METHODS Mirams (1957) regarded mature kauri as being at least 60cm diameter breast height (dbh). However, our criteria of mature kauri forest were:

89 Tane, Vol. 33,1991 1) that it should contain greater than 60cm dbh; 2) that there should be no sign of obvious disturbance (in some cases where signs of disturbance were found after sampling had commenced, this criterion was relaxed); and 3) that it covered at least 2ha in area. Quantitative samplings were carried out from 1980 to 1983. Point Centred Quarter Method (Cottom and Curtis, 1956) with the modification of Ogden and Powell (1979) was used to sample 25 kauri stands. In each stand 20 points were taken at 25m intervals along four 100m transects. However, in some cases due to landslides (slips) or the small area of the stand this procedure was not possible and shorter transects were taken. Trees > 10cm dbh only were included in the PCQ sampling. A species list with a frequency table for understorey plants (< 10cm dbh) was made using a circular plot (5m diameter) at each point. Lower plants were ignored but some widely distributed Pteridophyta were included. Species in the canopy, subcanopy and canopy gaps were noted separately. Slope, aspect and phytosociological attributes (density, frequency and basal area) were recorded. Ogden and Powell (1979) and Palmer (1982) used the term 'PCQ frequency' (PCQ = Point Centred Quarter Method) because according to Curtis and Mcintosh (1951) frequency values from plotless methods are not strictly comparable with those obtained using plot methods. Importance values (IV = the sum of relative values of frequency, density and basal area) give more information about the species than any other single attribute alone and reflect the realistic ecological importance of the species in a stand (Brown and Curtis, 1951). Every species was ranked according to its importance value and the species with the highest importance value in the stand was considered the dominant species. The data for each stand was summarised following Mueller- Dombois (1974). Nomenclature generally follows Allan (1961), or Moore and Edgar (1970). In some cases the two species of Weinmannia were not separated, although W. silvicola predominates. Dracophyllum was recorded to the if not identified. "Sprouting seedlings" were those with usually three or four while the term "seedlings" describes juveniles under 1cm dbh. All species recorded in the circular plots (<10cm dbh) are defined as understorey or ground flora.

RESULTS AND DISCUSSION Locations of the various kauri forests studied are shown in Fig. 1 while details are given in Table 1. Relative attributes with importance values of the first four dominant species of each stand are presented in Table 2. Absolute values for each stand and species (> 10cm dbh) are shown in Tables 3 and 4

90 respectively. A complete species list (< 10cm dbh) with relative frequencies in plots in each stand is given in Ahmed (1984), where more topographical, geological and meteorological details of each stand are also given. Species occurring in less than 4 stands have been excluded from Table 4, but are listed in the footnote.

Fig. 1 Map of northern position of North Island of New Zealand showing location of kauri forests studied. Numbers are stand number. Details are given in Table 1.

91 Table 1: Characteristics of 25 kauri sampling sites

Stand Locations Latitude Longitude Altitude Slope Aspect No S E (m) (degrees)

1. Te Paki Coastal Park 34° 28 172° 46 220 24 W 2. Omahuta Sanctuary 35° 15 173° 37 150 12 Flat Puketi State Forest 3. Te Harua Stream 35° 16 173° 44 305 35 N 4. Onekura Bluff 35° 11 173° 45 305 30 N 5. Loop Track 35° 15 173° 44 274 31 NE 6. Takapau Track 35° 13 173° 45 252 19 Flat 7. Manginangina Scenic 35° 12 173° 48 274 24 E Reserve 8. Russell State Forest 35° 23 174° 15 221 26 SW Warawara State Forest 9. Ridge Site 35° 22 173° 17 468 32 N 10. Plateau Site 35° 23 173° 17 358 28 N Waipoua State Forest 11. Waipoua Sanctuary 35° 29 173° 34 145 19 SW 12. Waipoua Coastal 35° 37 173° 29 244 21 SW 13. Trounson Kauri Park 35° 43 173° 38 175 19 N 14. Mount Manaia East facing slope 35° 49 174° 31 320 31 E West facing slope 35° 49 174° 31 320 26 W Little Barrier Island 15. Upper Thumb Track 36° 12 175° 04 335 32 S 16. Lower Thumb Track 36° 13 175° 04 213 26 NE 17. Valley Track 36° 13 175° 04 243 34 S High altitude Kauri 36° 12 175° 04 426 38 S 18. Te-Moehau 36° 31 175° 24 450 29 NW 19. Manaia Sanctuary 36° 52 175° 32 350 30 NW Waitakere Ranges 20. Cascade Kauri Park 36° 53 174° 31 240 35 EW 21. Piha 36° 58 174° 30 274 29 NW 22. 36° 58 174° 34 274 31 NW Hunua Range 23. Konini Forks 37° 04 175° 08 335 30 NE 24. Mount William 37° 13 175° 02 350 26 N 25. Kaimai-Mamaku State 37° 36 175° 52 350 30 NW Forest

92 1. Te Paki Coastal Park The Te Paki coastal park is located on the east coast of the Cape Reinga district and the study area is one of the ridges to the east of the Cape Reinga road a few miles south of the lighthouse. Fifteen species were recorded in the study area (Table 3). Kauri comprised 67% of the total basal area and 39% of the total density (Table 2). Leptospermum ericoides was abundant having 9 % basal area. kirkii, , Libocedrus plumosa, Phyllocladus trichomanoides and excelsa were other associates of kauri with low importance values. The canopy was irregular and high stand density (464 ha"*) with low stand basal area (34.5m2 ha-1) was the characteristic feature of this stand. Forty one species were recorded during the circular plot study. The vegetation was mainly composed of juvenile of Myrsine salicina, Gahnia xanthocarpa, Geniostoma ligustrifolium and kauri seedlings. These species were recorded in 65 to 95% of quadrats. Blechnum capense, Coprosma areolata, Cyathea smithii, Cyathea dealbata, Cyathodes fasciculata, Dodonea viscosa, Freycinetia banksii, Leptospermum ericoides, Nestegis lanceolata and Phebalium nudum were also found in 25 to 55% of quadrats. The study area was dry and steep. In many places the ground surface was exposed to direct sunlight either with no vegetation cover or with a few scattered plants.

2. Omahuta Sanctuary Omahuta kauri forest stands to the west of Puketi State Forest and 1km south of Mangamuka Bridge on State Highway 1 between Whangarei and Kaitaia. The sampling site was in a gully in Omahuta Sanctuary. This area was protected from the logging and now contains one of the finest stands of kauri in New Zealand (Sale 1978). Kauri is the dominant species of the area, although 28 stems in the sample comprised only 23m2 ha basal area (Table 3). Ackama rosaefolia is the co-dominant species. A few Dacrydium cupressinum, Podocarpus hallii and Podocarpus ferrugineus are present with kauri in the canopy layer. In the subcanopy Ixerba brexioides, tawa, Elaeocarpus dent at us and Pittosporum tenuifolium are abundant. These species are also frequent and widespread in the canopy gaps which are created by the fall of kauri or other big trees. Nineteen species > 10cm dbh are found in the area. The understorey contained 35 species < 10cm dbh, but it was mainly composed of dense Gahnia xanthocarpa, Dicksonia lanata and Freycinetia banksii. These species comprised 65 to 80% of the plots. Geniostoma ligustrifolium, and juvenile Ackama rosaefolia were found in

93 over 45% of the plots. Ricker sized kauri individuals were rare and only observed along the track and outside the sampling area in open places. However, 25 % of plots contained sprouting kauri seedlings.

Puketi State Forest Four stands were sampled in Puketi State Forest, which is positioned between the Hokianga and Whangaroa Harbour near the East Coast.

3. Te Harua Stream The study area is located on top of a ridge near Te Harua Stream at the Southern end of Puketi State Forest. Due to a slip area and the very steep slopes only 10 points were taken in this area. Kauri was the dominant species of the area and associated with the co- dominant species Phyllocladus trichomanoides and Dacrydium cupressinum. These two species had similar importance values. The former species was more abundant while the latter had a larger basal area (Table 2). The canopy and subcanopy was irregular. A few Podocarpus hallii, and Ackama rosaefolia were also present. The understorey flora was comparatively poor and scattered. Twenty species (< 10cm dbh) were found during the plot sampling, with Astelia trinervia, Cyathodes fasciculata, Gahnia xanthocarpa and juvenile kauri most abundant and recorded in over 50% of the plots. Freycinetia banksii was found in patches.

4. Onekura Bluff The area studied is situated on a steep slope in Onekura Ecological area, Puketi State Forest. Eighteen tree species were found but the dominant species was kauri, with 26% relative density and 83 % of the relative basal area of the stand (Table 2). Ackama rosaefolia was recorded as a co-dominant species and often found in a canopy layer with Podocarpus ferrugineus, and Podocarpus hallii, occasional Dacrydium cupressinum and were also present as canopy trees. The subcanopy was dominated by Pittosporum tenuifolium, Phyllocladus glaucus and Elaeocarpus dentatus. In a few places the canopy was broken by large standing dead trees. With the exception of Beilschmiedia tawa, present in 60% of the circular plots, no tree seedlings were abundant on the forest floor. Although 23 species were recorded from the quadrats, most were uncommon. Gahnia xanthocarpa, Astelia trinervia and Dicksonia lanata occupied over 80% of the sample plots. Freycinetia banksii and Coprosma australis were recorded in 45% of the plots. Kauri seedlings were found in 35 % of the quadrats.

94 5. Loop Track The study site was close to the Te Harua Stream. Kauri was the most abundant species on the site. Its 102 stem ha"* (> 10cm dbh) occupied 76.3m2 ha"* basal area (Table 3). Weinmannia sp. (presumably VV silvicola) was another numerous tree in area and attained the highest importance value after kauri (Table 2). Twenty one tree species were recorded in which Beilschmiedia tawa, Podocarpus ferrugineus, robusta and Podocarpus totara were the canopy associates of kauri. In a few places due to large standing dead trees the canopy was relatively open. Beilschmiedia tarairi, Dacrydium kirkii and Elaeocarpus dentatus were present as subcanopy species but subcanopy was not distinct. Gahnia xanthocarpa, Blechnum capense, Astelia trinervia and Freycinetia banksii were most abundant in the understorey, forming an extremely dense and impenetrable mass. Juvenile Beilschmiedia tawa and Dicksonia squarrosa were found in 50 to 60 % of the plots while sprouting kauri seedlings were recorded in 40% of the circular plots. Forty two species were found in the understorey (Ahmed, 1984).

6. Takapau Track The sample site lies on a flat part of the Pukatea Ridge in Puketi State Forest. The dominant species, kauri, formed a closed canopy while Ixerba brexiodes was the most numerous species in the subcanopy. These two species comprised 86 % relative density and 93 % relative basal area of the stand (Table 2). Only eight tree species were recorded in the area studied. Dacrydium cupressinum and Podocarpus hallii were the associates of kauri in the canopy. A lot of individual trees and groups of trees were dead or rotten and badly damaged. Many trees were found to be swollen and misshaped near the base. The understorey and ground flora comprised 33 species under 10cm dbh but juveniles of upper strata species were not abundant with the exception of kauri which was found in 85 % of the plots. Freycinetia banksii was relatively rare and the forest floor was covered by Gahnia xanthocarpa and Astelia trinervia occupying 95% of the plots. Dicksonia lanata and juvenile were recorded in over 50% of the circular plots.

7. Manginangina Scenic Reserve The sampling site stands on the eastern edge of the Puketi State Forest, about 13km west of the State Highway No 10 between Kaeo and Kerikeri. The study area was dominated by kauri though 22 tree species were found during the sampling. Kauri was present at a density of 51 stems per hectare and 64%

95 relative basal area. Ackama rosaefolia was co-dominant, being the most abundant species but with a lower basal area than kauri (Table 2). Ackama rosaefolia, Dacrydium cupressinum, Beilschmiedia tawa, Podocarpus ferrugineus and a few Podocarpus totara were the associates of kauri in the canopy layer. Though in general the canopy was irregular and strata were indistinct, Beilschmiedia tarairi, Elaeocarpus dentatus and Weinmannia sp. formed a subcanopy layer in a few places. Coprosma australis and Dicksonia squarrosa were the most abundant species of the understorey and recorded in 60 to 70% of the sample plots. Gahnia xanthocarpa, Astelia trinervia, Podocarpus ferrugineus and Dysoxylum spectabile were recorded in 55 % of the plots. Freycinetia banksii was relatively rare while Rhopalostylis sapida was found in patches. Kauri seedlings were recorded in 30% of the plots.

8. Russell State Forest Russell is situated between the Bay of Islands and Whangaruru Harbour on the eastern coast. The area sampled lies in the north eastern corner of the Waikare Ecological area in Russell State Forest. Kauri was the most numerous and dominant canopy species of the area, encompassing 81% of the relative basal area of the stand. Its density was 213 stems > 10cm dbh per ha. (Table 3). On the basis of relative basal area Podocarpus totara was greater than Phyllocladus trichomanoides but the latter species was more abundant (Table 2). These two species with Podocarpus ferrugineus and a few tall Ackama rosaefolia were also present as canopy trees. Phyllocladus glaucus, Pseudopanax arboreus, Myrsine australis, Knightia excelsa and Leptospermum ericoides were growing in the understorey but the subcanopy was not distinct while the latter two species were more frequent in canopy gaps. The ground flora consisted of 40 species including juveniles of the above mentioned tree species. Cyathea dealbata, Pseudopanax arboreus, macrophylla, Corokia buddleioides and Olearia rani were most abundant and frequent. These species occurred in 55 to 70% of the circular plots. Though no Beilschmiedia tarairi was observed in the upper stratum, juveniles were found in 40 % of the sample plots on the forest floor.

Warawara State Forest Warawara State Forest lies approximately 24km north of Hokianga Head along the west coast. Two sites were sampled in this forest. Beside Boscawen's (1912) pioneering work, no quantitative botanical work has been published for the state forest.

96 9. Ridge Site A thick stand of kauri is located on a north facing ridge. Twenty tree species were recorded but kauri was the most numerous and dominant species. Its density was 185 stems > 10cm dbh per ha (Table 3). On the basis of relative basal area, Podocarpus hallii was greater than Knightia excelsa but the latter species was more abundant (Table 2). The canopy was predominantly composed of kauri and Podocarpus hallii but a few tall Phyllocladus trichomanoides, Podocarpus ferrugineus, and Podocarpus totara were also intermixed, lxerba brexioides, Quintinia serrata and Pseudopanax arboreus were abundant in the subcanopy. The understorey was composed of dense Astelia trinervia, Coprosma lucida, Geniostoma ligustrifolium and Dracophyllum sinclairii occupying over 75 % of the plots. Juvenile Beilschmiedia tawa and Beilschmiedia tarairi were present in 56% while kauri seedlings were found in 62% of the plots. The ground flora was also rich in lower plants.

10. Plateau Area The sampling site stands on a north facing but relatively flat area in Warawara State Forest. In this stand lxerba brexioides (co-dominant species) was the most abundant subcanopy species, but kauri was the dominant on the basis of relative basal area and importance value (Table 2). Several dead kauri and large podocarps were observed in the sampling site, and the canopy was irregular. Podocarpus hallii, Podocarpus totara and Ackama rosaefolia had similar importance values (Table 2). They were present in the canopy stratum with Dacrydium cupressinum, Podocarpus ferrugineus and kauri. lxerba brexioides was the main canopy-former in the gap between the kauri. Twenty one tree species were recorded. The ground surface had many boggy patches and a dense carpet of ferns and lower plants. The forest floor was dominated by Astelia trinervia and Gahnia xanthocarpa which were present in 90% of the plots. Forty one species < 10cm dbh were recorded from the ground layer. Juveniles of Beilschmiedia tarairi were recorded in 50% of the plot though no tree was observed in the upper storey. Kauri seedlings were found in 40% of the circular plots.

Waipoua State Forest Waipoua State Forest is situated in the Hokianga country, approximately 15km from Hokianga Harbour and a few kilometres north of Trounson Kauri Park. Waipoua State Forest is the largest remnant of the once extensive kauri forest of northern New Zealand. However, parts of the forest have been considerably

97 modified by milling, gum digging, burning, wild pigs, and grazing animals. Two mature stands were sampled in the forest.

11. Waipoua Forest Sanctuary The study area lies in the eastern part of the Waipoua State Forest. Twenty four tree species were recorded, in which kauri was the dominant forming a closed canopy. In terms of density and also relative frequency, Beilschmiedia tarairi was the most abundant tree species (Table 2) of the area, forming the second stratum with Ackama rosaefolia, Knightia excelsa and Olearia rani. A few individuals of Podocarpus ferrugineus, Podocarpus hallii and Podocarpus totara reached the canopy. Near a group of dead large kauri and Podocarpus hallii the canopy was open with Dysoxylum spectabile, Elaeocarpus dentatus and Pseudopanax crassifolius. Individuals of kauri and Podocarpus hallii comprised 27% of the relative density and 77% of the relative basal area of the stand (Table 2). In the sample plot study 37 species were recorded, but the understorey was scattered and composed of Freycinetia banksii, juvenile Beilschmiedia tarairi, Blechnum fraseri, Cyathea dealbata and Cyathodes fasciculata. These species were recorded in 60 to 70% of the sample plots. Astelia trinervia, kauri seedlings, Ripogonum scandens and juvenile Ackama rosaefolia were also present in 30 to 50% of the plots. In a few places Rhopalostylis sapida were found growing in patches.

12. Coastal Area The study area is located 1.6km from the start of Kawerua Road near a water collection site. On the basis of relative basal area kauri was the dominant species. Although only 9 % of the relative density it occupied 66m2 ha.~* basal area. Twenty one tree species were found in the area studied (Table 3) but the canopy was mainly closed by kauri and the subcanopy was predominantly Beilschmiedia tarairi, Dysoxylum spectabile, Myrsine australis, Ackama rosaefolia and Melicytus ramiflorus. A few Knightia excelsa, Vitex lucens and Beilschmiedia tawa were also found in canopy gaps. Freycinetia banksii, Geniostoma ligustrifolium and juvenile Dysoxylum spectabile occurred in 65 to 75% of the sample plot on the ground surface. Though 32 species were recorded only Cyathea dealbata, Coprosma australis, Myrsine australis and Knightia excelsa were found in over 50% of the plots.

13. Trounson Kauri Park Trounson Kauri Park is located east of state highway 12 between Kaihu and Donnelly's Crossing. The sampling site is on a ridge-crest area facing a valley

98 system that extends to Maunganui Bluff on the west coast. In terms of basal area (81 %) the vegetation was strongly dominated by kauri. Beilschmiedia tarairi was the most numerous species (31 % relative density) associated with kauri but a much smaller relative basal area (Table 2). The canopy was usually closed by kauri but in many places Podocarpus hallii and Podocarpus ferrugineus were also present as canopy trees. Around the dead stumps of old kauri trees aggregations of Beilschmiedia tarairi were observed. Knightia excelsa, Coprosma arborea and Myrsine australis were relatively common in the undergrowth. Twenty one different tree species were recorded in the forest (Table 3). Thirty six species were found in the circular plot sampling and the ground flora was dominated by Beilschmiedia tarairi (75% plots) and (65% plots). Ripogonum scandens and Cyathea dealbata were found in 49 and 55% of plots respectively. Rhopalostylis sapida, Gahnia xanthocarpa and Astelia trinervia were recorded in 20 to 35 % of the plots.

14. Mount Manaia Mount Manaia is located on Whangarei Head approximately 13km east of Whangarei city. On the western slopes of the area studied, surrounding foothills and valleys have been cleared for pasture. Extensive milling, gum digging and burning is reported by Sale (1978) from the Whangarei district. Beside some general botanical descriptions by Kirk (1884) and Carse (1920) no quantitative work has been published from the area. The sample included points on both eastern and western slopes of Mount Manaia. Most of the 27 tree species were represented by only a few individuals. Kauri was the dominant species of the area (227 stems ha."*, > 10cm dbh). Leptospermum ericoides, the most numerous and widespread species, was present as a co-dominant species with Coprosma arborea and Dysoxylum spectabile. The canopy was irregular and a few large Vitex lucens, Sophora microphylla and emergent Knightia excelsa were also present. Although the forest floor had no vegetation cover in many places, 44 species < 10cm dbh were recorded. Except for a few species, all were represented by only a few scattered individuals. Juvenile Hoheria populnea and Geniostoma ligustrifolium, the only widespread species of the understorey, were found in 70 to 90% of the plots. Cyathea dealbata and Coprosma rhamnoides were recorded in 40 to 50% of the sample plots. Rhopalostylis sapida was growing in patches or as individual trees. Kauri seedlings were frequent and pole and ricker sized individuals were recorded in the study area.

99 Little Barrier Island Little Barrier Island is situated approximately 80km northeast of Auckland City in the Hauraki Gulf. Before it was protected, nearly one third of the Island was milled and burnt. One stand of the Valley Track and two stands along the Thumb Track were sampled.

15. Upper Thumb Track Rennison (1964) described the area as a kauri-tawa stand but the present quantitative sampling indicated that it might be better designated as a kauri¬ Leptospermum community. In this area of the Lower Thumb Track graded in Leptospermum ericoides and became rare. The area was dominated by kauri, comprising 62% relative basal area of the stand. Co- dominant species Leptospermum ericoides and Olearia rani were similar in terms of density but the former had greater basal area (Table 2). These two species were found in the subcanopy with Knightia excelsa, Weinmannia sp., Pseudopanax arboreus and Nestegis lanceolata. A few tall Metrosideros robusta emerged and were present with the canopy trees. Nineteen species were recorded in this stand (Table 3). Thirty five species (< 10cm dbh), including juveniles or upper storey trees, were recorded. The understorey was mainly dominated by Senecio kirkii, Olearia rani, Coprosma australis and Alseuosmia macrophylla, occurring in 70 to 90% of the plots. Juvenile Myrsine australis, Pseudopanax arboreus and Beilschmiedia tawa were recorded in over 50% of the plots. Freycinetia banksii was absent while Gahnia xanthocarpa and Astelia trinervia were found rarely. Kauri seedlings were recorded in 25 % of the sample plots.

16. Lower Thumb Track and 17. Valley Track These two stands had similar vegetation composition and dominant species and are treated together. The main difference was in the relative proportions of species and in rare species. Valley Track has relatively young kauri, though the largest kauri (230cm dbh) was observed near the junction of the Valley and Summit Track. In both tracks, kauri had similar relative density while the co- dominant, Nothofagus truncata had similar relative basal area (Table 2). The canopy was relatively closed at the Thumb Track. Leptospermum ericoides was also the most widespread species of the stands and present not only in canopy gaps, but also in the subcanopy. The subcanopy was dominated by Olearia rani, Knightia excelsa, Pittosporum umbellatum and Schefflera digitata. A few large Metrosideros robusta emerged through the subcanopy and were present with the canopy trees.

100 The ground flora of the two areas was similar. Thirty seven species were recorded from the Thumb Track and 35 species from the Valley Track. Freycinetia banksii and Gahnia xanthocarpa were rare while Astelia trinervia was abundant and widespread. Coprosma arborea, Myrsine australis, Senecio kirkii and Alseuosmia macrophylla were also frequent. Coprosma rhamnoides, Olearia rani and Pseudopanax arboreus were more common on the Thumb Track, while juvenile kauri were more abundant on the Valley Track and recorded in 70% of the plots. Nothofagus truncata was observed in 40 to 50% plots of both areas.

18. Mount Te-Moehau Mount Te-Moehau is located on the northern tip of the , about 60km east of Auckland. The study area was situated along the track leading from Stony Bay to the main peak. Due to the steep slope and a slip area only 10 points were taken in this area. The three kauri individuals were of sufficient magnitude to occupy 45% of the relative basal area of the stand. Widespread Weinmannia silvicola and Leptospermum ericoides show similar relative density and basal area but the first species leads in relative frequency and becomes the co-dominant species (Table 2). A few Podocarpus hallii and Beilschmiedia tawa were also present with the canopy trees while other species like Olearia furfuracea, Nestegis montana, lxerba brexioides and Weinmannia silvicola were growing under the canopy trees. Leptospermum ericoides and Knightia excelsa were present in the canopy gaps as well as in the undergrowth. Thirty four species (< 10cm dbh) were recorded from the ground surface plots but due to the dense and impenetrable carpet of Freycinetia banksii and Astelia trinervia only a few species were abundant. Juvenile Corokia buddleioides, Quintinia serrata, Myrsine salicina and Weinmannia silvicola were intermixed with these two species and found in 70 to 90% of the plots. Dracophyllum sp. and Blechnum capense were also recorded in 55% of the plots. Kauri seedlings were rare.

19. Manaia Sanctuary Manaia Sanctuary lies 24km north of Thames on the Coromandel Peninsula while the study area is located in a remote area near Kahatarahae Stream on a northwest ridge. Beside the timber and gum industries, mining associated with the gold rush has also had an effect on the vegetation of the area. No quantitative botanical work has been published from the area. The canopy was closed by tall, overmature kauri trees. Its 47 individuals ha."* occupied 127 m2 basal area ha."* (Table 3). Co-dominant lxerba brexioides and Weinmannia silvicola were the most numerous and widespread subcanopy species. Among 23

101 species recorded, Beilschmiedia tawa, Podocarpus ferrugineus, Podocarpus totara, Hedycarya arborea and Quintinia serrata were also common. The understorey was thickly covered by Freycinetia banksii with Lygodium articulata and juvenile Beilschmiedia tawa occupying 60 to 70% of the plots. Cyathea dealbata and juvenile Podocarpus hallii were also found in 45 to 55% of the plots. Astelia trinervia and Gahnia xanthocarpa were growing in patches.

Waitakere Ranges The Waitakere Range is situated 30km west of Auckland City. The forest cover of the area is highly disturbed but there are still a few areas in a relatively undamaged state, probably saved from milling by the steep slopes. Three mature kauri stands were studied in the Waitakere Ranges.

20. Cascade Kauri Park Cascade Kauri Park lies in the northern end of the Waitakere Ranges to the northeast of Waitakere reservoir. The study area was near the Upper Kauri Track. Large sized kauri were found on the steep slope while on the top of the ridge and near the track relatively young kauri were growing. It was the sole canopy tree with 188 individuals ha"*. 'Second growth' of kauri was observed just outside the sampling area. Seventeen species were recorded in the area (Table 3) and the Knightia excelsa was found as a co-dominant species, having a close resemblance to Phyllocladus trichomanoides in importance value (Table 2). Dacrydium cupressinum, Ixerba brexioides, Podocarpus ferrugineus and Hedycarya arbor ea formed the second stratum. A few Metrosideros robusta also occurred as canopy trees. Thirty eight species under 10cm dbh were counted. Astelia trinervia, Coprosma lucida, Coprosma arborea, Geniostoma ligustrifolium were abundant and found in 60 to 70% of quadrats. Kauri seedlings were recorded in 35% of plots. In many places, the ground surface was covered with Astelia trinervia and Gahnia xanthocarpa while Freycinetia banksii was found in 25 % of the sample plots.

21. Piha Piha is situated on the western side of the Waitakere Range. The study area is located just before the Piha road, around Cowan and Kauri Grove Tracks. Among 21 tree species recorded, kauri occupied 82% of the basal area (Table 2). The canopy was irregular due to death of large kauri and podocarp species. In a few places Podocarpus ferrugineus, Podocarpus totara and

102 Metrosideros robusta were also present in the canopy layer, lxerba brexioides, the most numerous associate of kauri, formed a second stratum and was absent in canopy gaps. This species was similar to Metrosideros robusta in importance values. Olearia rani, Dacrydium cupressinum, Hedycarya arborea, Knightia excelsa, Myrsine australis and Quintinia serrata were relatively common in the canopy gaps. Thirty seven species (< 10cm dbh) were listed during plot sampling but only five species were abundant. Freycinetia banksii was found in 90% of plots, forming thick and dense masses on the ground surface with Geniostoma ligustrifolium, Melicytus macrophyllus and Astelia trinervia (65-80 % ). Coprosma lucida (50%) and Rhopalostylis sapida (20% of plots) were also recorded.

22. Huia Huia is situated at the southern end of the ranges. The northwest facing sample area was near Hamilton Saddle just above the Lower Huia reservoir. The study area was dominated by kauri, which had only 48 individuals ha."* but comprised 70 % to the total basal area. Dacrydium cupressinum was the co- dominant. This species was similar to Knightia excelsa in importance value (Table 2). A few large Metrosideros robusta, Podocarpus totara and Dacrydium cupressinum were also associated with kauri as canopy trees. Podocarpus ferrugineus, Phyllocladus trichomanoides, Knightia excelsa, Hedycarya arborea and Beilschmiedia tarairi formed the second stratum of the community. Where the canopy was open Olearia rani, Hoheria populnea, lxerba brexioides, Pseudopanax crassifolius and Myrsine australis were growing. Thirty nine species were observed in the circular plot study. Where a canopy tree had fallen the ground was densely covered particularly with Freycinetia banksii intermixed with Geniostoma ligustrifolium and Cyathea dealbata. These species occurred in 75 to 95% of the plots. Gahnia xanthocarpa was also recorded with Astelia trinervia in 49 % of the plots.

Hunua Range Hunua Range is situated about 40km southeast of Auckland City, under the administration of Auckland Regional Water Supply Reserve. Two sites are studied in this area. Extensive cutting, burning, presence of wild animals and cattle are reported from this area.

23. Konini Forks The study site is located on a northeast facing slope about 3km from the water reserve. The sample area does not fall in any of Barton's (1972) types. Fifteen tree species were recorded but the canopy was closed by kauri and co-

103 dominant Beilschmiedia tawa. These two species occupied 68 % of the relative area (Table 2). A few large Dacrydium cupressinum and Podocarpus ferrugineus were also present with the canopy trees. Knightia excelsa, Pseudopanax crassifolius, Olearia rani, Olearia furfuracea and Myrsine australis were in the subcanopy. Thirty one species were recorded but the ground flora was irregular and the ground surface was often exposed with no vegetation cover. Cyathea dealbata and Paratrophis microphylla were most abundant and found in 65% of the circular plots. Kauri seedlings, Pseudopanax crassifolius and Freycinetia banksii were recorded in 40 to 50% of the plots. Gahnia xanthocarpa though present, did not fall within the plots sampled.

24. Mount William Mount William is located between Bombay and Pokeno approximately 50km to the southeast of Auckland city at the western edge of the Hunua Range. The area has a relatively young and dense stand of kauri. There were 416 stems ha."* with a diameter of greater than 10cm dbh (Table 3). Eleven tree species were found in the study area. With the exception of Dacrydium cupressinum, kauri was the only canopy tree. Phyllocladus trichomanoides was the co-dominant subcanopy species. These two species contributed 65 % density and 89% of the relative basal area of the stand (Table 2). Nothofagus truncata, Olearia rani and Knightia excelsa were also abundant and associated with co- dominant species. The understorey was scattered. During the plot sampling 28 species < 10cm dbh were found in which Coprosma spathulata and Geniostoma ligustrifolium were most abundant and recorded in 70 to 90% of the plots. Juvenile Phyllocladus trichomanoides, Cyathodes fasciculata, Cyathodes juniperina and Olearia rani were present in over 55 % of the circular plots. Juvenile kauri were also abundant on the forest floor and recorded in 65 % of the sample plots (Table 3).

25. Kaimai-Mamaku State Forest The sampling area stands in state forest park near the end of Hot Spring Road and 11km from Katikati. Extensive milling, gum digging, gum bleeding and burning have occurred in the state forest. Kauri grows in many places and further south, usually in small groups, as individual trees or regenerating stands. The sampling site represents a mature kauri stand close to the southernmost limit of the species. Kauri was the most abundant and dominant canopy species of the sampling area. In terms of relative frequency and relative density Ixerba brexioides and Weinmannia sp.

104 were close to kauri and found in subcanopy layers. These species comprised approximately 60 % of the relative density and about 84 % of the relative basal area (Table 2). All other species (18 tree species), with the exception of Knightia excelsa, were rare. Among the 40 species > 10cm dbh the most abundant and widespread species on the forest floor was kauri, juveniles and seedlings being found in 85% of the circular plots. Coprosma australis, juvenile Knightia excelsa, Geniostoma ligustrifolium, Dysoxylum spectabile and Hedycarya arborea were recorded in 50 to 60% of the plots. Astelia trinervia and Gahnia xanthocarpa were rare while Freycinetia banksii was not recorded.

CONCLUSIONS In the present study kauri stands were studied from 150 to 820m elevation, with relatively flat to extremely steep slopes and on various aspects. However, altitude and species composition did not clearly correlate (Ahmed 1988). In most cases kauri was not the dominant species in terms of density but it was in terms of basal area and importance value. An average of 129 individual ha."* of kauri (28 to 227 ha."*) with 57m2 ha. basal area (range 22.6 to 127.3 m2 ha."*) was obtained. Sixty nine tree species (> 10cm dbh) and 112 understorey species (including seedlings and saplings < 10cm dbh) of canopy and subcanopy trees) were recorded from the entire study (Ahmed, 1984, lists all). Ten species were recorded as co-dominant species: knightia excelsa, Dacrydium cupressinum, Phyllocladus trichomanoides, Beilschmiedia tawa, lxerba brexioides, Weinmannia sp., Ackama rosaefolia, Beilschmiedia tarairi, Leptospermum ericoides and Nothofagus truncata. In most cases the co-dominant species of any particular stand was also prominent in other kauri stands. Other prominent species were Podocarpus ferrugineus, Podocarpus hallii, Podocarpus totara, Elaeocarpus dentatus, Hedycarya arborea, Myrsine australis, Pseudopanax crassifolius, Olearia rani and Gymelaea (= Nestegis) lanceolata. Beside the seedlings of canopy and subcanopy species, understorey vegetation was characterised by Freycinetia banksii, Geniostoma ligustrifolium, Gahnia xanthocarpa, Cyathea dealbata, Astelia trinervia, Lygodium articulata and Alseuosmia macrophylla. Seedlings or saplings of kauri were recorded in all the stands studied and were abundant in some (Ogden, Wardle and Ahmed 1987). The present study shows much overlapping and similarities in species composition in widely separated kauri forests, implying that all kauri forests are relatively homogenous in nature.

105 Table 2. Site summary of PCQ sampling Relative phytosociological attributes and importance values of the first four leading dominant species of 25 kauri forests. Notes: 1. relative frequency; 2. relative density; 3. relative basal area; 4. importance value. (Percentages have been rounded to nearest whole number).

Stand Name of Species RF1 RD2 RBA3

1 australis 29 39 67 131 Leptospermum ericoides 20 20 9 49 Dacrydium cupressinum 16 13 8 37 Libocedrus plumosa 9 8 5 21 Other 11 species 31 21 10 63

2 8 13 62 83 Ackama rosaefolia 16 24 5 45 Dacrydium cupressinum 12 10 15 37 Ixerba brexioides 14 13 5 32 Other 15 species 50 40 13 103

3 Agathis australis 21 32 73 126 Phyllocladus trichomanoides 19 21 5 45 Dacrydium cupressinum 15 15 13 43 Podocarpus hallii 3 2 5 10 Other 13 species 42 29 4 75

4 Agathis australis 19 26 83 128 Ackama rosaefolia 20 22 2 45 Podocarpus ferrugineus 11 8 3 22 Beilschmiedia tawa 10 9 2 21 Other 14 species 39 34 10 84

5 Agathis australis 14 25 61 100 Weinmannia sp. 10 14 2 26 Beilschmiedia tawa 7 6 7 20 Podocarpus ferrugineus 7 7 5 19 Other 17 species 62 49 25 136

6 Agathis australis 30 25 74 129 Ixerba brexioides 35 61 19 115 Dacrydium cupressinum 12 5 3 20 Podocarpus hallii 11 4 3 18 Other 4 species 12 4 1 18

7 Agathis australis 15 17 64 95 Ackama rosaefolia 15 23 5 43 Dacrydium cupressinum 9 9 9 27 Beilschmiedia tawa 6 7 9 22 Other 18 species 55 44 13 112

106 8 Agathis australis 22 36 81 139 Phyllocladus trichomanoides 18 18 7 43 Podocarpus totara 12 11 8 31 Phyllocladus glaucus 10 8 2 19 Other 14 species 37 28 3 68

9 Agathis australis 15 27 64 106 Knightia excelsa 11 13 3 27 Podocarpus hallii 5 3 12 19 Phyllocladus trichomanoides 5 5 6 16 Other 16 species 64 52 15 131

10 Agathis australis 11 13 41 64 lxerba brexioides 17 23 10 50 Podocarpus hallii 8 9 11 28 Podocarpus totara 8 8 12 27 Other 17 species 57 48 26 130

11 Agathis australis 11 14 69 94 Beilschmiedia tarairi 17 26 9 52 Ackama rosaefolia 12 13 5 30 Knightia excelsa 12 9 2 23 Other 20 species 47 38 15 100

12 Agathis australis 9 9 68 86 Beilschmiedia tarairi 17 26 10 54 Dysoxylum spectabile 17 18 10 46 Knightia excelsa 17 18 3 39 Other 17 species 39 29 9 76

13 Agathis australis 15 20 81 117 Beilschmiedia tarairi 17 31 8 56 Knightia excelsa 10 11 3 23 Podocarpus hallii 10 8 2 19 Other 17 species 49 31 5 85

14 Agathis australis 14 18 56 87 Leptospermum ericoides 15 20 6 41 Vitex Iik:ens 5 4 27 36 Coprosma arborea 11 17 3 31 Other 23 species 56 41 9 105

15 Agathis australis 17 13 62 92 Leptospermum ericoides 14 23 17 53 Olearia rani 16 21 4 41 Knightia excelsa 15 13 3 30 Other 15 species 40 31 14 84

16 Agathis australis 26 26 63 115 Nothofagus truncata 27 44 26 97 Leptospermum ericoides 12 11 4 26 Olearia rani 8 5 1 14 Other 10 species 27 14 7 48

107 17 Agathis australis 23 26 51 100 Nothofagus truncata 23 38 27 87 Leptospermum ericoides 20 17 6 43 Metrosideros robusta 8 4 13 26 Other 8 species 27 14 4 44

18 Agathis australis 7 5 45 57 Weinmannia silvicola 20 23 10 53 Leptospermum ericoides 13 23 10 46 Podocarpus hallii 10 8 24 41 Other 11 species 50 43 11 103

19 Agathis australis 11 8 72 90 Ixerba brexioides 23 36 13 72 Weinmannia silvicola 17 20 4 41 Beilschmiedia tawa 8 8 4 20 Other 16 species 42 29 7 77

20 Agathis australis 21 41 90 151 Knightia excelsa 15 13 1 29 Phyllocladus trichomanoides 14 13 2 29 Dacrydium cupressinum 8 5 3 16 Other 13 species 43 29 4 76

21 Agathis australis 15 26 83 124 Ixerba brexioides 12 12 1 25 Metrosideros robusta 7 7 8 22 Olearia rani 9 9 1 19 Other 17 species 56 46 8 110

22 Agathis australis 14 20 70 104 Dacrydium cupressinum 7 9 11 27 Knightia excelsa 12 10 4 26 Olearia rani 6 8 1 16 Other 21 species 60 53 14 128

23 Agathis australis 14 24 45 84 Beilschmiedia tawa 13 24 23 59 Knightia excelsa 17 14 7 39 Pseudopanax crassifolius 12 8 1 21 Other 12 species 44 29 24 97

24 Agathis australis 27 48 78 153 Phyllocladus trichomanoides 19 18 11 47 Nothofagus truncata 14 11 3 27 Olearia rani 9 7 3 19 Other 7 species 31 18 5 53

25 Agathis australis 17 21 64 101 Ixerba brexioides 16 20 13 49 Weinmannia sp. 16 19 7 42 Knightia excelsa 12 8 2 22 Other 15 species 39 33 14 85

108 Table 3. Site summary of PCQ sampling. Absolute values of 25 kauri forests. Notes: 1, Numbers of species (NS); 2, stand density (SD); 3, kauri density (KD); 4, stand basal area (SBA); 5, kauri basal area (KBA).

71 n 7]—- Stand No Density ha 1 Basal area m ha NS1 SD2 KD3 SBA4 KBA5

1 15 464 180 34.5 23.1 2 19 216 28 36.7 22.6 3 17 355 114 54.4 39.5 4 18 345 90 79.3 65.6 5 21 407 102 125.4 76.3 6 8 408 103 88.1 65.5 7 22 308 51 75.3 48.0 8 18 586 213 80.8 65.1 9 20 696 185 65.6 42.3 10 21 673 84 85.2 34.1 11 24 494 71 69.5 47.8 12 21 776 68 97.0 65.7 13 21 671 134 145.9 118.4 14 27 1299 227 122.3 66.7 15 19 824 108 70.8 44.0 16 14 673 177 73.4 46.0 17 12 764 201 78.5 40.0 18 15 758 40 75.0 34.6 19 20 621 47 177.1 127.3 20 17 464 188 126.4 113.2 21 21 239 63 90.7 74.8 22 25 240 48 48.1 33.7 23 18 589 144 66.8 30.0 24 11 874 416 66.2 51.8 25 19 724 149 59.6 38.0

average 578.72 129.24 83.70 56.56 Standard Deviation 242.19 82.69 32.75 27.73

109 Table 4. Summary of species (PCQ sampling). Absolute values and importance values of species in 25 kauri forests.

Notes: 1 = Number of stands in which a species is recorded. Species found in less than 4 stands have been excluded (see Ahmed, 1984); 2 = Average density ha"'; 3 = Maximum density ha"'; 4. Average basal area m2ha"'; 5 = Maximum basal area m2ha"'; 6 = Average importance value; 7 = Maximum importance value.

Density Basal Area Importance ha"' m^ha"' Value Name of Species PR1 AD2 MD3 AB4 MB5 AIV6 Miv"

Ackama rosaefolia 11 41 77 2.01 8.60 22.42 45.72 Agathis australis 25 129 416 56.57 127.29 105.41 152.77 Beilschmiedia tawa 15 27 140 3.67 15.18 17.16 59.46 Beilschmiedia tarairi 11 58 200 4.47 11.10 30.10 56.00 Coprosma arborea 8 44 219 0.80 3.09 11.63 30.50 Cyathodes fasciculata 11 4 9 0.07 0.13 2.74 5.08 Dacrydium cupressinum 17 28 58 3.59 8.13 19.26 43.15 Dracophyllum latifolium 6 11 38 0.17 0.40 6.12 11.82 Dysoxylum spectabile 6 52 143 2.20 9.52 15.37 45.54 Elaeocarpus dentatus 18 14 31 0.47 1.23 8.36 19.04 Hedycarya arborea 13 13 31 0.32 0.76 7.57 15.22 Ixerba brexioides 12 76 250 6.24 23.43 37.51 72.10 Knightia excelsa 22 47 143 1.86 5.01 19.52 38.96 Leptospermum ericoides 10 97 260 3.90 11.74 28.64 52.62 Libocedrus plumosa 5 11 35 0.56 0.22 7.98 5.48 Melicytus macrophyllus 6 4 8 0.06 0.09 3.98 6.38 Melicytus ramiflorus 6 12 24 0.52 1.28 5.05 9.51 Metrosideros robusta 10 12 33 4.90 10.01 11.95 25.56 Mida salicifolia 4 5 8 0.29 0.47 2.80 3.72 Myrsine australis 17 12 40 0.24 1.10 5.39 12.33 Myrsine salicina 6 5 11 0.19 0.48 4.03 6.48 Gymelaea (= Nestegis) lanceolata 11 9 24 0.19 0.79 4.38 9.91 Gymnelaea (= Nestegis) montana 9 10 38 0.13 0.33 4.78 8.70 Nothofagus truncata 4 172 297 10.78 21.03 54.02 96.66 Olearia furfuracea 7 19 57 0.56 1.46 9.28 19.40 Olearia rani 13 32 175 0.73 3.10 12.98 41.24 Persoonia toru 6 4 6 0.10 0.16 3.06 3.67 Phyllocladus trichomanoides 16 33 153 2.50 7.09 20.52 47.12 Pittosporum tenuifolium 8 9 24 0.33 0.72 5.83 7.03 Podocarpus ferrugineus 20 18 43 2.21 4.74 12.54 22.47 Podocarpus hallii 15 20 63 4.07 18.09 14.53 41.10 Podocarpus totara 15 16 62 2.93 10.32 11.06 30.63

110 Pseudopanax arboreus 10 12 38 0.16 0.51 5.31 12.88 Pseudopanax crassifolius 13 18 48 0.27 0.72 9.20 20.89 Quintina serrata 9 18 63 0.48 2.07 7.50 22.38 Weinmannia sp. 11 52 171 2.32 7.94 18.03 52.72

Species occurring as trees > 10cm dbh in 3 or fewer stands (in some cases this is because the trees rarely reach this diameter, in others because they are not common): Brachyglottis repanda, Carmichaelia spp., Carpodetus serratus, Coprosma areaolata, C. australis, C. lucida, Corynocarpus laevigatus, Cyathodes juniperina, Dacrydium kirkii, Dodonaea viscosa, Elaeocarpus hookerianus, Eugenia maire, Geniostoma ligustrifolium, , G. lucida, Hoheria populnea, Laurelia novaezelandiae, Libocedrus bidwillii, Litsea calicaris, Melicope simplex, Metrosideros umbellata, Macropiper excelsum, Gymnelaea (= Nestegis) cunninghamii, Phebalium nudum, Phyllocladus glaucus, Pittosporum umbellatum, P. ellipticum, P. eugenioides, Podocarpus dacrydioides, P. spicatus, Pseudowintera axillaris, Schefflera digitata, Sophora microphylla, Vitex lucens.

ACKNOWLEDGEMENTS

Personnel of the New Zealand Forest Service, in the head office of the Auckland Conservancy, the district office at Thames, and the kauri Management Unit at Kaikohe helped with site location and logistics. Mr Richard Serra, Jonathan Palmer, and other students helped in various aspects of the field work. We extend our thanks to all concerned. The manuscript was typed by Miss Raewyn Arthur and Miss Debra Speed.

REFERENCES

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Ill Cockayne, L., 1908. Report on botanical survey of Waipoua kauri forest. Appendix to the Journal of the House of Representatives of New Zealand C-14. Cockayne, L., 1928. Vegetation of New Zealand 2nd Edition. Verlag Von Wilhem. Engelmann. Cottam, G. and Curtis, J.T., 1956. The use of distance measures in phytosociological sampling. Ecology 37 (3): 451-460. Cranwell, L.M. and Moore, L.B. 1936. The occurrence of kauri in montane forest on Te Moehau. New Zealand Journal of Science and Technology, 18: 531-543. Curtis, J.T. and Mcintosh, R.P., 1951. An upland forest continuum in the prairie-forest border region of Wisconson. Ecology 32: 476-496. Ecroyd, C.E., 1982. Biological flora of New Zealand 8 Agathis australis (D.Don) Lindl. (Araucariaceae) Kauri. New Zealand Journal of Botany 20: 17-36. Kirk, T., 1884. Botanical notes. Transactions and Proceeding of the New Zealand Institute 16:361- 368. Mirams, R.V., 1957. Aspects of natural regeneration of the kauri (Agathis australis Salisb). Transactions of the Royal Society of New Zealand 48 (4,1:661-680. Moore, L.B. and Edgar, E., 1970. Flora of New Zealand. Vol U. Government Printer, Wellington. Mueller-Dombios, D. and Ellenberg, H., 1974. Aims and Methods of vegetation Ecology. John Wiley and Sons. Inc. New York. Ogden, J., 1983. The scientific reserves of Auckland University. Quantitative vegetation studies. Tane 29:163-180. Ogden, J. and Powell, J.A., 1979. A quantitative description of the forest vegetation on altitudinal gradient in Mount Field National Park, Tasmania and a discussion of its history and dynamics. Australian Journal of Ecology 4:293-325. Ogden, J., Wardle, G.M. and Ahmed, M. 1987. Population dynamics of the emergent conifer (Agathis australis (D.Don) Lindl. Kauri) in New Zealand. II. Seeding population sizes and gap- phase regeneration. New Zealand Journal of Botany 25: 231-242. Palmer, J., 1982. A dendrochronological study of kauri (Agathis australis Salisb.). MSc thesis, University of Auckland. Rennison, G. 1964. An investigation of the kauri/hard community of South-West Little Barrier Island. Tane 70:84-87. Sale, E.V. 1978. Quest of Kauri. A.H. and A.W. Reed, Wellington, Sydney, London. Sexton, A.N. 1941. Note on the kauri-beech (Nothofagus truncata) association in Omahuta State Forest. New Zealand Journal of Forestry 4:308-310. Wardle, G.M., 1984. Observation on the structure dynamics of kauri (Agathis australis) forest. MSc thesis, University of Auckland, New Zealand.

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