
Part I.—Quadrat Investigations of Natural Regeneration
In the Waitakere Ranges, all degrees of inter-mingling of the forest are to be found from a pure kauri stand to a type of forest dominated mainly by species of Podocarpus and Dacrydium, the broad leaf, Beilschmiedia tawa not being common. These are some of the members of the ultimate climax, but the forests in this region do not appear to have reached a great degree of stability.
It is obviously impossible to follow a single succession from its initiation to the ultimate climax, and resource has therefore to be made to the study of a number of examples of the same succession at different developmental stages contemporaneously. Similarly it is not possible to see kauri forest initiation to-day. The commencement of secondary forest can, however, be observed in the natural regeneration which usually occurs after the forest has been destroyed or damaged. It is realised that the sequence of events which we see to-day giving rise to a new kauri forest may be, and probably are, vastly different from the succession which gave rise to the kauri forest which we now see and which is dominated by trees that are many hundreds of years old.

A very brief description of the subsere and the kauri forest will suffice, as these various communities have been adequately described by Cockayne (1928). The succession of events is as follows. After the short-lived initial communities of grasses, and herbs there develops a tea-tree or manuka scrub. It is a typical shrub community with Leptospermum scoparium—the dominant—present in considerable numbers. In the early stages the ground is covered with a mass of Gleichenia circinata. Beneath this fern, which is often one metre and more deep, are to be found many of the forest tree seedlings, including Agathis. With the passage of time the fern dies out, Leptospermum scoparium grows larger, the allied Leptospermum ericoides enters the succession and becomes a co-dominant. Many individuals of the numerous species which go to make up the kauri forest are now seen as small saplings growing beneath the tea-tree canopy. L. ericoides is usually by now completely dominant, but it is eventually over-topped by individuals of Agathis which at this stage have a characteristic conical form. (Plate 43, fig. 2.) The Leptospermum ericoides persists for a considerable period, first as a co-dominant but later as a subdominant, and there comes into being what may be called “a forest of kauri rickers”. These rickers, which often form dense stands, increase in both diameter and height until eventually there is developed a community, little different from the mature forest. The main difference is the smaller size and very much larger number of individuals of kauri in the former type. The ground is by now covered with a heavy tussock-like growth of “kauri grass”, Gahnia xanthocarpa and Astelia trinervia, which frequently reaches a height of two to two and a-half metres. (Plate 44, fig. 2) By the time the mature kauri tree has developed, it has become the only dominant. Even in a dense stand, the trees are not close together but may be separated by distances of fifteen to twenty metres. There are also a number of smaller trees of other species but only one, Knightia excelsa, reaches into the sub-canopy layer. The canopy in the pure stand is quite irregular, being formed of the large rounded heads of the mature kauri tree. (Plate 43, fig. 1.)
The occurrence and distribution of natural regeneration was investigated by a series of quadrats laid down in the various successional stages, five types being recognised.
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(1) Young Leptospermum.
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(2) Older Leptospermum (usually lacks the ground layer of fern and rush found in the first type). (Plate 43, figs. 3, 4.)
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(3) Leptospermum–Agathis community.
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(4) Kauri ricker community. (Plate 44, fig. 1.)
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(5) Kauri forest. (Plate 44, fig. 2.)
A quadrat size of 36 sq. m. was decided upon from the species-area curve. Myers (1948) in a more comprehensive study of minimal area in Leptospermum shrubland also found that the minimal area for this community was 36 sq. m.
On every quadrat a list of all the species encountered was made. In addition, counts of all the individuals of Agathis were made. These individuals were classified into one of the six size-age groups listed below in order that a better picture of regeneration could be obtained. (Plate 45.)
Class I
Small seedlings with cotyledonary leaves only. The majority of these individuals are from the current year's seed fall, but retarded individuals are occasionally found which have come from a previous fall.
Class II
Seedlings with up to about twelve leaves and not more than ten centimetres in height; branching of the stem not apparent.
Class III
Seedlings between ten and fifty centimetres high, with a number of branches both lateral and terminal.

Class IV
Slender saplings with a stem not more than five centimetres in diameter. The height varies from fifty centimetres to four metres or more.
Class V
Saplings and small rickers five to fifteen centimetres in diameter.
Class VI
All individuals whose stem is more than fifteen centimetres in diameter.
A total of eighty-four quadrats were laid down, sixty of which were thirty-six square metres in area, the remaining twenty-four being one hundred square metres. The number of species occurring in the sixty smaller quadrats are eighty-six, with the addition of the twenty-four larger quadrats a further ten species were added, making a total of ninety-six.
Natural regeneration is considered first as the total number of individuals of Agathis in each of the five successional types, and secondly as the distribution of the size-age classes in these communities. In both cases, the data is taken from twelve 36 sq. m. quadrats in each of the five community types.
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| Community | Size-Age Classes | No. of quadrats on which individuals of Agathis were found (Total = 12) | |||||||
|---|---|---|---|---|---|---|---|---|---|
| I | II | III | IV | V | VI | Total | |||
| Young Lepospermum | 6 | 63 | 6 | 5 | – | – | 80 | 7 | |
| Proportion | 7.5 | 78.8 | 7.5 | 6.2 | – | – | |||
| Older Leptospermum | 22 | 213 | 45 | 6 | 1 | – | 287 | 12 | |
| Proportion | 7.7 | 74.2 | 15.7 | 2.1 | 0.3 | – | |||
| Leptospermum-Agathis | 34 | 233 | 49 | 97 | 54 | 26 | 493 | 12 | |
| Proportion | 6.9 | 47.2 | 10.0 | 19.7 | 10.9 | 5.3 | |||
| Ricker Agathis | 10 | 262 | 66 | 32 | 28 | 27 | 425 | 12 | |
| Proportion | 2.3 | 61.6 | 15.5 | 7.5 | 6.6 | 6.6 | |||
| Kauri forest | 23 | 88 | 18 | 19 | – | 3 | 142 | 11 | |
| Proportion | 16.2 | 62 | 12.7 | 7.0 | —2.1 |
A second set of quadrats 10m. × 10m. provided the following results, which are very similar to those obtained from the smaller quadrats, apart from the larger numbers of individuals.
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| Size-Age Classes | No. of quadrats on which individuals of Agathis were found. | |||||||||
|---|---|---|---|---|---|---|---|---|---|---|
| Community. | I | II | III | IV | V | VI | Total | No. of quadrats | No. of quadrats of occurrence | |
| Leptospermum-Agathis | 243 | 1472 | 224 | 558 | 181 | 36 | 2714 | 12 | 12 | |
| Proportion | 8.19 | 54.3 | 8.2 | 20.6 | 6.7 | 1.3 | ||||
| Ricker | 3 | 85 | 26 | 15 | 8 | 20 | 157 | 2 | 2 | |
| Proportion | 1.9 | 54.1 | 16.6 | 9.5 | 5.1 | 12.8 | ||||
| Kauri Forest | 191 | 341 | 104 | 39 | – | 16 | 691 | 10 | 10 | |
| Proportion | 27.6 | 49.4 | 15.1 | 5.6 | – | 2.3 |

It can be seen from the above table that there is considerable variation in the total number of individuals of all size-age classes in the various communities examined. The smallest number of individuals is found in the young Leptospermum, the highest at the middle stage of the succession. The number of individuals in the kauri forest is also relatively low, much lower than would be expected even for the class I individuals. There is adequate seed available for the germination of many more seedlings.
In the young Leptospermum it is only to be expected that the density of Agathis will be low, since invasion by this and the other forest species has only just commenced. This fact was reflected in the absence of kauri from almost half of the quadrats and the small numbers on several of the remainder. The absence of individuals in classes V and VI is also due to the early age of the community at this stage. By the time an older Leptospermum community has developed, there has been a considerable increase in the number of individuals of all age classes, especially classes II and III. There has been a slight increase in the numbers in class IV, and class V is now represented by a solitary specimen. The increase in total numbers is probably due as much to the “accumulation” of seedlings as to the operation of any factor more favourable to their growth. In spite of the increase in numbers there is evidence to show that there is a considerable mortality of young kauri seedlings; these results only show the “end point” of their survival at the time of counting, they do not show what the trends in survival are.
The next stage of the sere is an ecotonal community in which Agathis becomes codominant with the giant tea-tree Leptospermum ericoides. The most noticeable feature of this stage is the presence for the first time of individuals of the oldest size-age class. It must not be thought that the individuals of this class in this community are at all comparable with those of the same class in the kauri forest. Many of them will not yet be producing seed and those that are, only small quantities. This added seed source much nearer to hand is probably partially responsible for the large increase in the number of class II seedlings. It is possible, too, that conditions in the community are more favourable for the germination and survival of small seedlings. On the whole, it can be said that natural regeneration is at its greatest in the Leptospermum-Agathis community.
There is a change from the sub- or co-dominance of Agathis by the time the ricker community develops. There is apparently a slight decline in the regenerative capacity of the sere as shown by the reduction in the number of class IV and V seedlings. Further evidence on this point is the absence of one or other of the various size-age groups on almost all of the quadrats. The ricker community gradually changes into a mature forest. In the kauri forest, there is a decrease in the number of individuals of Agathis and a steady increase in the size of those remaining. The diameter of any of the adult specimens examined was not less than sixty centimetres, and many were considerably larger than this. The first point to be noticed about this community is the small number of individuals of all age classes and more importantly the complete absence of individuals of class V in all probability showing that domination by Agathis is now on the downgrade because it is apparently not replacing itself. The largest individuals of class IV were not more than two or three centimetres in diameter and can scarcely be considered as potential replacements for some of the dominant individuals. If a community is to maintain itself as a stable climax, the dominant species must be able to reproduce themselves so that there is formed a more or less even aged stand. If the seedlings of tree species cannot meet the competition of lesser species, whether it be in the herb or shrub stratum, such trees must eventually disappear from the community. The small number of individuals of all age classes and the complete absence of class V indicates that, at least in some of the strata, Agathis is not competing successfully. Permanent or true dominance involves the ability to compete successfully in all strata of the

community and Agathis is certainly not doing this. It might be argued that there are sufficient individuals in class IV allowing for a considerable mortality, to maintain the stocking of class VI at its present level. However, the forest is surely old enough for individuals of Agathis intermediate between class IV and class VI to have developed and the fact that they have not already done so indicates that they may not under the present environmental conditions. At the same time, it does not seem possible to regard the members of class IV as potential replacements of the present mature individuals. The greatly reduced occurrence of seedlings of Agathis on quadrats within the mature forest is harder to explain for this community than for any of the others. Collections of seed by means of traps have shown that there is more than sufficient seed falling to the ground in one season to produce the number of seedlings of all ages recorded, let alone those which are only in the cotyledonary stage (size-age class I). The activities of at least one insect are responsible for the destruction of a large proportion of the seed; this does not explain, however, the complete absence of one of the size-age classes and the small number of individuals in the remainder.
