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The hypothesis that there is a large number of specific adaptations in New Zealand flora to browsing by the moa (large flightless birds) is critically discussed. While there clearly can be no definitive test of the hypothesis as the moa are extinct, the circumstantial evidence previously proposed in its favour is mostly ambiguous or irrelevant. Tests of the hypothesis based on the current ecological preferences of plants with putative anti-moa browsing adaptations are likewise inadequate or inconclusive. Experimental tests with browsing mammals are unhelpful. Of the many browsing adaptations that have been suggested, only spines and mottling of understorey leaves merit further investigation. Nevertheless, while specific adaptations to avian browsing may be rare in the New Zealand flora, more generalised anti-herbivore adaptations are common.
Before European settlement, New Zealand had no browsing or grazing mammals. Their role was filled by large flightless birds, collectively called the moa (Order Dinornithiformes). Polynesians, arriving about 1000 years ago, had hunted the moa to extinction by about 400 years ago (Anderson 1989b). New Zealand was therefore free of terrestrial browsing vertebrates from then until the European introduction of a wide range of browsing and grazing mammals, beginning with goats in the late eighteenth century.
Early this century it became apparent that introduced mammals were rapidly degrading the vegetation, primarily by killing browse-sensitive trees and shrubs and preventing regeneration (Cockayne 1928). It was widely assumed that moa had been primarily grazing birds of open grasslands and had therefore exerted insignificant browsing pressure, resulting in a low level of defences against browsers in the indigenous flora.
Greenwood and Atkinson (1977) presented a detailed argument for widespread adaptation to moa browsing in the New Zealand vegetation, greatly extending previous suggestions that moa browsing may have been responsible for the evolution of the distinctive divaricating shrubs (Carlquist 1974:242, Livingstone 1974, Taylor 1975, Melville 1978). Since 1977 the moa browsing hypothesis, although meeting some opposition (McGlone and Webb 1981), has been strongly supported in both in the scientific literature (Burrows 1980, Lowry 1980, Mitchell 1980, Caughley 1983, Lee and Johnson 1984) and in the popular media. Atkinson
Critics of introduced mammal control policy have pointed out that New Zealand forests and grasslands had evolved in the presence of browsing birds, and that the vegetation immediately before European settlement — from which moa had been absent for several centuries — could not have been in a natural state (Caughley 1983, Butchler 1989). Introduced mammals, they claim, are restoring a more natural vegetation. The question of whether or not the moa and other browsing birds exerted a strong or weak influence on the plants of New Zealand is therefore not only important for a complete understanding of the ecology of the present vegetation, but also has management implications.
Jared Diamond (1990) in a vivid metaphor described the New Zealand vegetation as still being influenced by the ‘biological ghosts’ of long extinct birds. Can these ghosts be laid to rest or does a complete understanding of our vegetation depend on our acknowledging their intangible but influential presence? Our aim in this paper is to subject the now reformulated moa-browsing hypothesis to critical scrutiny.
There were many herbivorous birds in the pre-Maori New Zealand fauna. Flighted folivores included the extant New Zealand pigeon (Hemiphaga novaeseelandiae), kokako ( Callaeas cinerea), and the red-crowned and yellow-crowned parakeets (
However, moa were doubtlessly the dominant vertebrate browsers from ground level to a height of about 3 m. The 11 species had a range in body weight from 22 to 230 kg, and a range of head-up stature from 93 to 252 cm (Anderson 1989a). Moa ranged through the entire landscape: they occurred in subalpine grassland, shrubland, scrub, open dry forest and dense, wet, lowland forest (Worthy 1990). However, their populations were unlikely to have been large: the rapid elimination of the moa by the Maori, the number of fossil sites, and arguments based on the biomass of ratite birds in other places, all tend to suggest low densities. Moa probably had a rather slow rate of increase (Anderson 1989b). Anderson (1989a: 87) concluded that “… the total population of moas in each island was measured in tens of thousands, the South Island population was about twice that of the North, and the greatest concentration of moas occurred in the eastern half of the South Island.”
Burrows (1980a, 1980b) has shown from his analyses of fossil gizzards that some moa could cut through twigs at least 6 mm thick, and that twigs formed a normal and substantial part of the diet. He later concluded: “The Dinornis moa seem to have behaved like deer, cattle or goats, in taking a variety of food items, including browse.” (Burrows 1989). Anderson (1989a) postulated that had a large food
Because sensory perception and mouth structure in birds differ substantially from those in browsing mammals, Greenwood and Atkinson (1977) asserted that anti-herbivore adaptations in New Zealand must also differ markedly from those typical of areas where mammals are the main vertebrate herbivores. Three features, they suggested, contrast moa with herbivorous mammals.
• Moa lacked a prehensile tongue and teeth, which would have made them more reliant on a clamping, pulling, and breaking feeding action, than on the cutting and chewing action of ungulates.
• In common with many other birds, moa had good colour vision but a poor olfactory sense. Hence food location and selection depended more on sight than smell.
• The beak and associated horny covering of the face would have protected the head nearly up to the full depth of the bite. Spines, which are effective against the soft noses of mammals, would have been less effective against moa.
These assumptions are only partly valid. The difference between moa and ungulate browsing is probably less than Greenwood and Atkinson (1977) initially envisaged. Both groups could cut and ingest substantial twigs; the significant difference may have been the nature of the two digestive systems. Moa probably could not debark trees and shrubs. While moa lacked soft, vulnerable noses, their eyes and tongues were certainly exposed to spines so that the presumed different reaction to spines may be relative rather than absolute.
Greenwood and Atkinson (1977) reversed the relative sensory capabilities of the moa. Moa probably had poorish vision and a well-developed offactory sense (Anderson 1989, Worthy 1990). The larger relative size of the factory area of the moa brain suggests that moa probably had a better developed sense of smell than most other bird groups (Worthy 1990). Atkinson and Greenwood (1989) suggested that as perception of the indigo-violet-red part of the light spectrum is restricted in some palaeognathous birds, these colours would also have been less apparent to moa.
In this section we discuss the many features of the New Zealand flora that have been interpreted as anti-browsing adpatations (Table 1).
Batcheler (1989) suggested that because a number of low-statuted, fast-growing plants normally found on seral sites have toxins (e.g., Sophora, Coriaria, Brachyglottis repunda, Solanum have alkaloid defences), this is evidence for plant defence against moa browsing. His hypothesis, following from a similar line of reasoning by Greenwood and Atkinson (1977), is that moa browsed preferentially on nutrient-rich sites and on the lower strata of forest, and therefore high levels of defence could be expected in plants that habitually grew there. Low nutrient
Pseudopanax crassifolius (Mitchell 1980).
First of all, there is no reason to assume that toxins, carbon-based chemical defences, and low nutrient and fibrous leaves are specifically anti-moa browsing adaptations; they are effective against a wide range of herbivores, including invertebrates. All the indigenous species of flowering plants in New Zealand toxic to livestock are related to toxic species elsewhere, and no toxin restricted to New Zealand has yet been recognised (Connor 1977). The New Zealand situation appears to be a particular example of a universal trend.
Secondly, toxins may be a preferred means of defence for plants growing on nutrient-rich sites, and the concentration of toxic plants on such sites must be seen in terms of general plant defence strategies. The faster-growing a plant, the less profitable it is for it in terms of absolute growth rate to invest in defence (Coley et al. 1985). Pioneers, for example, accumulate nutrients more quickly than mature forest species, and allocate a great proportion of their resources to growth, but little to defence (Dirzo 1984). As fast-growing plants on nutrient-rich sites have more nitrogen to spare for defence than do slow-growing plants on poor sites, successional plants often possess nitrogen-based toxins such as alkaloids, which have a high turnover within the plant and can be produced in variable quantities according to need (Rhoades 1983). On the other hand, slow-growing plants in low-light, nutrient-poor, or stressed environments rarely have nitrogen-based toxins for defence, but rely on carbon-based chemicals such as polyphenolic compounds and lignins, which are less energy-demanding and use mimimal amounts of scarce nutrients. Nutritive qulity of leaves is usually low in species growing on poor soils, and such leaves tend to be sclerophyllous and are retained longer, possibly to minimise nutrient loss through herbivory, environmental stress, and nutrient leaching (Chabot and Hicks 1982). However, although low nutrient status leads to less browsing through herbivore avoidance, the long life of the leaves means the browsing that does occur has a great impact.
In general, therefore, plants growing in low-nutrient and environmentally stressed sites have well-defended leaves that are not browsed to the same extent as those from plants growing on richer sites. Fast-growing plants invest more in growth than defence, and rely on growing through their vulnerable phases, or deploy facultative defences such as highly effective low-bulk toxins. These defence strategies in the New Zealand flora are not necessarily a consequence of moa-browsing but a generalised adaptation to all herbivory.
The risk of attack on a given plant by a herbivore is referred to as its “apparency”. Plants can reduce their apparency by modifications that variously make the plant look dead, as if it has already been attacked, like a non-plant object such as a stone, like another distasteful plant, or render it less visible. To a greater or lesser extent all plants employ an apparency strategy, as nearly every change in their external appearance or life history must alter the ease with which they are located and attacked by herbivores.
Mottled leaves. Atkinson and Greenwood (1989) suggested that mottled leaves in seedling plants may have reduced discovery by browsing moa (e.g., Parsonsia capsularis, and
Some insects use colour vision and leaf patterns to locate food plants, although not as commonly as they use smell and taste (Prokopy and Owens 1983). Smith (1986) has suggested that mottling of the leaves of Byttneria aculeata (a subcanopy vine in Panama) mimics effects of leaf miners, and thus discourages ovipositing and leaf damage. He showed that mottling was more common in clearings than in shady sites, possibly because of the trade-off between reduced photosynthesis and decreased insect damage in mottled leaves, and the reverse in unmottled leaves. Givinish (1990) has shown that in northeastern North America mottled leaves are more common in herbs of shaded forest understoreys than in any other growth form, and that they are essentially absent in trees, shrubs, herbs or vines of sunny sites. He suggested that mottling camouflages from vertebrate herbivores the foliage of particularly vulnerable phenological groups (evergreens; spring ephemerals) in light-dappled understoreys. There is therefore a strong possibility that some leaf mottling is directed against herbivores, but investigation of mottled plants in New Zealand would be needed to establish whether this is so here.
Dark coloration. Dark bronze and purple leaves are common in the New Zealand flora, in particular in new foliage and the exposed foliage of high-altitude plants. Blanc (1989: quoted in Givinish 1990) suggested that dark-coloured foliage held close to the ground could be camouflaged against herbivores. In nearly all plants he examined, dark-coloured understorey leaves were within 20 cm of the ground surface. If the foliage is held higher the apparency advantage is largely lost as the leaves are viewed against a contrasting background rather than the dark soil. However, most New Zealand examples of dark-leaved understorey plants (Table 1) I hold their foliage well over 20 cm above the ground, and furthermore, the coloration of the leaves varies seasonally.
It is also possible that herbivory is not involved in the New Zealand situation. Other possible adaptive explanations for dark leaves include modification of the heat balance of the leaf, protection from ultra-violet radiation, response to frost damage, and modification of the photosynthetic capacity of the leaf. Without more investigation we cannot choose between these explanations.
Mimicry. When a plant strongly resembles another object, living or non-living, less palatable than itself, it is often regarded as a mimic. Many cases of mimicry have been alleged but are exceptionally difficult to prove, as assessment of the similarity is usually subjective (see Edmunds 1990 for a lucid discussion). Mimicry by living leaves and stems of unpalatable objects such as stones and dead twigs is essentially different from the better known examples of camouflage mimicry in the insect world. When an insect mimics a leaf, twig or stone, only in a very superficial way does it take on the characteristics of that object. It remains an insect, heterotrophic and mobile, and its energy exchanges and life-style are but marginally affected. However, when a plant mimics an inanimate object, it is changed totally. A leaf or stem that resembles a dead twig, for instance, is very different from a green leafy stem. Its changed shape, ridigity, and pigmentation will substantially alter its heat absorption, gas exchange, light energy capture, construction costs, and competitive relationships. We should therefore be cautious in assuming that a given plant is mimicking an inanimate object, however close the resemblance; its resemblance may be a consequence of environmental factors. Atkinson and Greenwood cited several examples of the various types of mimicry. None of them is convincing.
Some New Zealand plants resemble dead twigs (Table, 1). Muehlenbeckia ephedroides resembles plants from arid regions that have a similar grey pubescence and leafless habit. It is typical of exposed unstable substrates such as river beds
Muehlenbeckia's leafless grey stems are an adaptation to the harshness of the local environment and its prostrate habit a response to frequent flooding and abrasion.
The reduced twig-like leaves of the juveniles of forest-dwelling Pittosporum obcordatum, are assumed to increase its resemblance to a dead or partially browsed plant (Atkinson and Greenwood 1989:83). However, there are plausible reasons why a juvenile plant may restrict its photosynthetic surfaces, the most likely of which is the necessary diversion of scarce resources to roots (Wright 1992). The prominence of the whitish mark along the midrib is a consequence of the reduction of the reaf blade; it may or may not be significant in apparency.
When one plant is allegedly mimicking another plant, the problem is more complex. It first has to be established that the model is both less palatable to herbivores and more widespread and abundant than the mimic plant, and that the two taxa frequently occur together, Without these conditions, selective pressure would not be sufficient to result in mimicry. A second complexity is that in adopting the form, structure, and colour of the unpalatable model, the leaves of the mimic also begin to resemble the model in essential life functions such as light interception, heat balance, and water relations. It then becomes debatable whether it is environmental or habitat convergence, or true mimicry. Atkinson and Greenwood (1989) mentioned a number of possible examples of plant-plant mimicry. For none of them do they establish a convincing case for mimicry, although it remains a possible interpretation.
Alseuosmia pusilla is a small, usually unbranched shrub, the palatable leaves of which are strikingly similar to the red-mottled, distasteful leaves of Pseudowintera colorata. However, although A. pusilla fulfils the requirement of a mimic, in that it is less common than its model but grows with it, this is not sufficient to establish mimicry. Aside from the red mottling, which is only well developed in sun-exposed leaves, there is nothing distinctive about P. colorata leaves. Red, brown, or yellow mottling of leaves is common in New Zealand, and seems to be a response to leaf damage, especially in the presence of bright light. Other species of Alseuosmia, which have dissimilar leaves to Pseudowintera colorata, also have red and brown mottles. Moa, as they possessed an excellent sense of smell, are unlikely to have been confused by the visual similarity between the two species. Therefore, the most likely explanation of the visual similarity between the two species is chance.
The moa's excellent sense of smell makes the proposed mimicry between spiny-leaved Aciphylla species and some similar appearing, but non-spiny tussock-forming plants improbable. Atkinson and Greenwood (1989) link the spiny-leaved Aciphylla subflabellata and unpalatable
Although numerous plants in the New Zealand flora have tough fibrous leaves, Atkinson and Greenwood (1989) singled out Pseudopanax crassifolius, P.ferox, P. lineare, Cordyline australis, and
Tough, fibrous stems and leaves do deter herbivores (Choong et al. 1992). However, it is likely that in many plants the anti-herbivory effects are a secondary consequence of their structural function. In the suggested New Zealand examples, the long leaves are held at angles ranging from horizontal to near vertical. The single-stemmed juveniles of Cordyline and the lancewoods gain fully adult leaves only when branching begins. Their leaves alone form the juvenile canopy and must provide the necessary extension; hence they have the strength and toughness characteristic of branches. Phormium has the form of a giant tussock, with individual leaves up to 3 m in length, thick at the base and often quite stiffly erect except for the top few centimetres, and therefore clearly acting as stems. These species have developed these leaves primarily in relation to their particular requirements for canopy formation.
The claim that plants reduce the size of their leaves or the total area per plant, or dispense with them either entirely or seasonally primarily because of herbivory (Atkinson and Greenwood 1989, Batcheler 1989) has little merit.
Batcheler (1989) suggested that deciduousness in the New Zealand flora is related to herbivory. However, deciduousness in New Zealand species is linked to cool winters. For example: Fuchsia excorticata and
The New Zealand brooms (Carmichaelia, Chordospartium, Corallospartium) follow a typical broom pattern of reducing or dispensing with leaves in favour of photosynthetic stems, an adaptation therefore unlikely to be specifically directed at moa. It is, moreover, an adaptation that probably developed to reduce water-loss rather than herbivory. However, as with many such adaptations, it undoubtedly also reduces the palatability of the plant to a range of herbivores.
Reduction in total leaf area, or loss of leaves for a period, we believe therefore is best explained by the plant being unable to afford the cost of leaves if their income in form of photosynthate is low versus outgoings such as respiration, increased root area, or stem mass. Reduction in leaf area because of an elevated risk of herbivory seems inexplicable, given that there are other ways to minimise herbivory that do not involve sacrificing photosynthetic potential.
Spines are usually a defence against animals. Spininess is in particular a characteristic of arid-land plants, and documented evidence of avoidance or lower usage of such plants by vertebrate herbivores suggests that it evolved as a defence against them (see discussion in Janzen 1986:616–620). A strong point in Greenwood and Atkinson's (1977) hypothesis is that the New Zealand flora has few spiny plants, and they made a convincing case for this being because birds have horny beaks and therefore are undeterred by spines. However, they made an exception for Aciphylla, regarding their stiff sharp pointed leaves as effective anti-browsing devices, presumably because the spines threatened the eyes of browsing birds.
Strong stiff leaves with thick cuticles will tend to be spine-like. Hence, adaptation to drying windy climates by a tussock species could bring with it spininess as a mere byproduct. As previously pointed out by McGlone and Webb (1981),
Moa should have been attracted to plants of high nutrient status. Therefore, if a plant with a putative anti-browse feature is also nutritionally attractive to birds, it will have passed this test.
Lee and Johnson (1984) measured the concentration of several nutrients in leaves of six divaricate and four non-divaricate Coprosma species. Nitrogen, phosphorus, calcium, and sodium were all present in higher concentration in divaricating species, and only potassium was more abundant in non-divaricating species. A canonical discriminant analysis of the data showed that the leaves fell into two groups, largely on the basis of nitrogen, phosphorus and sodium, in which small-leaved divaricating species were separated from large and small-leaved non-divaricating species.
However, if we take the two most important nutrients, nitrogen and phosphorus, and plot the data (Fig 1), a less clear-cut picture emerges. The two most weakly divaricating species of the group (C. rotundifolia and C. rubra) are separated from the rest by their high levels of nitrogen and phosphorus. The other group of divaricate and non-divaricate species have substantially lower nitrogen and phosphorus levels. The least nutritious leaves among the divaricating plants, belong to the most tightly divaricating species (C. propinqua and C. crassifolia). It would be hard to argue, despite their lower nutrient levels, that any of these Coprosma spp. are unattractive to vertebrates. Coprosma lucida and
We argue that the high nutritional status of small thin leaves is probably related to their high ratio of photosynthetic tissue to supporting tissue, and possibly to higher photosynthetic activity than in the larger leaves. Therefore, acquisition of a small thin leaf or of a large, thick leaf could alter the nutritional balance regardless of any other factor. Highly active leaves may be more attractive, and therefore browsed more often, but this is off-set by higher growth-rates and turn-over, as discussed above.
As they often occur as successional plants on nutrient-rich sites, but often grow slowly, and allegedly devote a high proportion of their resources to defence, divaricating plants must be seen as highly anomalous. If browsing was so intense on nutrient-rich sites in New Zealand, it would seem that high levels of defensive chemicals, in particular toxins, would be a more optimal solution, as it would demand minimal change in the structure of the plants, be more flexible, and permit faster growth.
Is this test a good one? What if nearly all divaricating plants had leaves of exceptionally low nutrient content? Would this lead to the abandonment of the browsing hypothesis? We think not. For instance, Atkinson and Greenwood (1989:88) include among adaptations to moa browsing “…linear and fibrous juvenile leaves of low nutrient value of Pseudopanax crassifolius.” If, as in the thin-leaved divaricating shrubs, the leaves of
The anti-browsing adaptation should be best developed at that part of the plants life-cycle when it is most exposed to moa browsing. The anti-browsing feature should not be retained when the plant is out of reach of moa.
This test is based on the assumption that terrestrial vertebrate browsing necessitates a greater defence investment in foliage borne low on the plant. This is a reasonable assumption with some plants and in some habitats. For instance, in Europe, holly (Ilex aquifolium) has more abundant spines on leaves close to the ground and leaves on heavily browsed shoots have longer spines (Peterken and Lloyd 1967), and in Africa thorny plants are intrinsically thornier below 3–5 m from the ground (Milewski et al. 1991) and browsed shoots tend to become more densely spiny. However, in some situations juvenile spininess is reduced and lost in the adult form without the presence of large vertebrate herbivores. For example, in the Hawaiian Islands certain Cyanea species have juveniles that are densely armed with stout, sharp prickles, while the adults are smoother, and less prickly (Lammers 1990). The prickles are regarded as a defence against phytophagous land snails and insects. Presumably, either attacks by animals are more intense close to the ground or the effect of the browsing is more severe on the juveniles. Regardless, the greater defence of the juveniles is not in any way related to the size per se of the browsing animal. At a more general level, Lowman (1985) found that leaves closer to the ground in tropical Australian forests were grazed more heavily by insects. It would seem reasonable to suppose that similar sorts of mollusc and insect attack occurred in New Zealand, and that for whatever reason, it was more intense closer to the ground. There was a large array of flighted and climbing browsing birds and insects in pre-historic New Zealand. The prevalence or better development of a putative browsing defence when the plant is small, which is then lost or reduced at greater heights, cannot be automatically attributed to the browsing height range of moa.
Even if it is accepted that moa browsing was a decisive factor in the evolution of plants, it is by no means clear that many allegedly browse-resistant plants pass the ‘life-cycle test’. Although most divaricating plants are of low stature, nearly 20% regularly attain heights of more than 4 m, and some can grow to 12 m, thus beyond the reach of even the tallest moa. Unless the divaricating plant form is a juvenile phase only, the degree of alleged defence does not change, no matter how tall the plant. For instance, in tall divaricating plants, such as Coprosma crassifolia. Melicope simplex and
The degree of divarication often alters markedly with the aspect of the plant. Those parts of the plant exposed to the sun or the prevailing wind are usually strongly divaricate, and leafy shoots are often protected by one or more layers of near leafless branches. Sheltered, or shaded parts are mostly less strongly divaricate, are usually leafy to the exterior, and the leaves are larger. If a divaricating plant is growing completely in shade, leaf size and leafiness approximate more closely to those of the interior of an open-grown plant, and it is rare for divarication to be as well-developed as in the open. Many (for example, Coprosma rhamnoides) develop spreading, sparsely leaved plagiotropic arrays of thin branches at intervals up the stem.
In windshorn plants degree of divarication is primarily related to exposure not to ease of access by herbivores. For example, in exposed plants of Hoheria angustifolia the small-leaved divaricating juvenile foliage will extend further up the windward side of the plant, with adult foliage at the same level on the relatively protected lee side. In extremely windy sites, adult foliage often does not extend above the level of juvenile foliage, but the stem with adult leaves grows parallel to the ground on the lee side (McGlone, pers. obs.). Similar cases can be seen when divaricating plants grow in the shelter of rocks. Why divarication should be so intense a few centimetres above the shelter of a rock, and relaxed in the lee of the
Most divaricating plants therefore fail the life cycle test, in that they are heavily defended when apparently there is no need, such in the uppermost layers out of the reach of moa, and weakly defended, if at all, underneath a canopy well within the reach of any moa. In most open-grown plants, the lower parts of the plant and the side away from environmental exposure are inexplicably left weakly guarded.
The height at which the tough-leaved single-stemmed juvenile of Pseudopanax crassifolius begins to change into the less coriaceous and branched adult tree certainly is at about the level where moa browsing would have been greatly reduced or absent. However, measurement in a range of habitats on the height to the beginning of canopy branching (which is a conservative indication of the height at which the tough juvenile foliage is replaced by adult leaves) show a variable range of heights for changeover (Table 2). Changeover occurs at a height of 1–2 m lower in the open when the competing vegetation is low-growing than under tall dark forest canopies. Under very dark, high canopies juveniles can reach as high as 7 m, with only a small cluster of leaves at the top of a stem often less than 5 cm in diameter. Juveniles in the open tend to have abundant leaves down the length of relatively thick stems. If moa-browsing is the main reason for the juvenile form, why should it be retained at heights well above the risk of attack? We suggest that the single-stemmed juvenile is an adaptation designed to carry the stem apex to a height where branching will spread the canopy above competing foliage. There fore, the lower the light level, the longer the juvenile form is retained.
Plants with anti-browsing features should reach their greatest abundance on fertile sites where moa are certain to have browsed most intensively, and be unimportant on sites inaccessible to moa.
Divaricating plants are more abundant on landforms characterised by soils of moderate to high fertility. They are often abundant on alluvial lowland soils, but are not so common on soils derived from mafic rocks (Lee 1992), or on acid or leached soils. However, it is possible to exaggerate the relationship. For instance, on Stewart Island, the variety and abundance of divaricating plants are greatest on alluvial flats, and shrubs such as Pseudopanax anomalus and trees with divaricating juveniles such as Prumnopity taxifolia and Plagianthus regius are virtually confined to these sites (Wilson 1987). However, despite the acid soils of low fertility that characterise the island, some divaricating shrubs are both widespread and common (e.g., Myrsine divaricata, Coprosma propinqua, Coprosma rhamnoides, and
Greenwood and Atkinson (1977), Batcheler (1989) and Atkinson and Greenwood (1989) pointed out that few if any epiphytes are divaricating, despite the their dry branch and rock environment. They saw this as contradicting claims that the divaricating habit is related to drought avoidance (see McGlone and Webb 1981), and also as demonstrating that the divaricating syndrome was rare when the risk of moa browsing was low. Some divaricating plants have developed an ability to grow in dry areas, and the small expendable leaves, near leafless exterior, and self-shading habit of many divaricating plants could help explain this. Keen (1970) concluded on the basis of experimental studies that the small-leaved divaricating
Astelia solandri and
Greenwood and Atkinson (1977) claimed that divaricating plants are rare on steep cliffs, where moa would have found it impossible to browse. Like tree brunches, cliffs present a relatively specialised environment. Many cliffs are unstable, and seem to have a large opportunistic rather than cliff-specific element in their vegetative cover. Furthermore, cliffs vary greatly in all sorts of characteristics, including accessibility. Sheltered moist cool and relatively stable cliff sites are not favourable to divaricating plants, but sunny dry windy and exposed cliffs on Banks Peninsula have an abundance of divaricating plants (Hugh Wilson, pers comm).
We have discussed above why toxic foliage in seral vegetation is not necessarily a defence against moa browsing, but a more general adaptation to herbivore attacks on palatable, fast-growing plants. The argument could be reversed. Fast-growing but browse-sensitive plants characteristic of fertile soils, for example, Aristotelia serrata and
Areas known not to have had moa should have reduced numbers of species and smaller populations of plants with strongly developed anti-herbivore defences, than in areas that had moa.
We are certain that moa did not occur on most offshore islands more than a few kilometres from the mainland, although Stewart Island had moa (Anderson 1989). This test therefore contrasts islands with mainland sites.
Greenwood and Atkinson (1977) showed that the percentage of divaricating plants (in relation to the woody flora) on moa-free islands was lower than on comparable areas of coastal mainland (4.7% versus 10%). In general, the coastal mainland sites had more woody species (average 47) than the island sites (32). However, the Chatham Islands have a percentage of divaricating plants (8%) not much lower than the New Zealand flora as a whole, and higher than two of the mainland coastal areas chosen as comparisons. The low numbers of species involved and differences in climatic conditions between island and mainland sites (which are only rough proxies for islands) make this comparison of doubtful significance.
Of more significance are species or closely related species that occur on islands and the mainland, but differ in degree of divarication. Greenwood and Atkinson (1977) noted that a non-divaricating relative of Myrsine divaricata and a non-divaricating form of
Sophora microphylla also has non-divaricating juvenile populations on the mainland, and the divaricating form is best expressed only in the far south and south-east of the South Island (Godley 1979) and in the east of the North Island.
In general, we are dubious the geographic distribution test is satisfactory. Islands differ so much from mainland situations the only convincing test would be similar islands with and without moa. What evidence there is appears to support the view that it is not prehistoric moa presence or absence but current climatic conditions which control the numbers and abundance of divaricating plants on islands. We would suggest that the other putative anti-moa adaptations would likewise show no consistent geographic relationship to the past distribution of moa.
Where vertebrate terrestrial herbivores are or were absent, heavily defended plants should be at a competitive disadvantage in relation to less well defended plants.
This is not a test proposed by Atkinson and Greenwood (1989) but one that should be considered. A major weakness of the moa-browsing hypothesis is the way that divaricating plants and other plants with supposed anti-browsing defences remained abundant in the absence of substantial browse pressure and in the presence of fast-growing competitors. A fair assumption is that anti-browsing strategies come with some cost to the plant. Nevertheless, divaricating plants, and other plants with supposed anti-browsing features, can be successful in the absence of any browsing pressure, and in the presence of ‘normal’ large-leaved, fast-growing competitors. With the extenction of the moa some 400 years ago, and major depletion starting about 600 years ago (Anderson 1989), non-defended plants have had between 300 and 500 years without significant browsing to overwhelm their defended competitors. That this did not happen is clear from the abundance of divaricating plants, spiny plants, plants with tough leaves, and dark
A possible counter to this argument is that although defended against browsing, these plants are better adapted to their characteristic environments than other species, and therefore will not be outcompeted, even in the absence of browsing. If this is granted, it leaves the problem of what it is about plants with these defence strategies that makes them so suitable for these environments.
The argument can be reversed: if browsing pressure was strong enough to induce in the vegetation such a wide range of anti-browsing strategies, it follows that before the extinction of the moa, many palatable and weakly defended plant species should have been restricted by browsing. This contention is not supported by the fossil evidence. Pollen of highly palatable species of Griselinia, Pseudopanax, Aristotelia, and Fuchsia are not often abundant in the fossil record because of poorly-distributed grains, but as far as can be told, they neither increase or decrease with the elimination of the moa. At some pre-human sites they are at times abundant (McGlone, Neall and Clarkson 1988), and this suggests that moa-browsing pressure was certainly not sufficient to restrict them.
As far as we are aware, there has been only one attempt to experimentally investigate the nature of browsing on putatively moa-defended plants. However, this investigation, reported in Atkinson and Greenwood (1989:87), is badly flawed.
‘The main restriction to mammalian browsing of divaricating plants appears to be mechanical. In a feeding trial of 3 replicates I.A.E.A. offered fresh leafy branches of Psedopanax anomalous, to 3 cattle and 3 goats, all pasture fed animals. In all cases the branches were briefly sniffed and then either nibbled sparingly or ignored. After removing all the leaves from these branches, and offering them again to the goats, they were rapidly consumed (leaving none for the trials with the cattle!).’
Livestock are not closely related to moa, and moa browsed in a different manner and had different sensory capabilities. Furthermore, the livestock were pasture-fed animals unaccustomed to browsing. Therefore, we cannot be sure that their reaction to the branches was not one of unfamiliarity, whereas the stripped leaves resembled their normal diet. This experiment therefore adds nothing to our knowledge of how any animals may have fed on a divaricating plant. But why use mammalian browsing to test an hypothesis centred on moa? For instance, many divaricating plants can resist browsing by deer, goats, hares, and possums in the wild (see, for example, Clarkson 1986:46–47). However, we believe that browsing resistance in these plants is but a side effect of an adaptation to a set of environmental stresses and is a consequence of the unique climatic and biological history of New Zealand. Studies of mammal browsing are largely irrelevant, as most of the putative anti-moa strategies are not common in other regions with large populations of indigenous mammalian herbivores.
Of all the many suggestions put forward for specific anti-moa-browsing strategies in the New Zealand flora, we believe that only spines in Aciphylla and dark or mottled leaves in juvenile plants are worth investigation as possible anti-moa adaptations. Nevertheless, while we largely dismiss the notion of widespread specific adaptations to moa-browsing, we are convinced by the evidence for generalised adaptations against browsers. As elsewhere in the world, tough,
It would be difficult enough to achieve resolution of the question of evolution of anti-browsing adaptations if the moa and other extinct browsing birds were extant. In their absence, and therefore the absence of any but the most scanty evidence as to their sensory capabilities and diet preferences, the task is formidable. We stress that demonstration of the resistance of some elements of the New Zealand flora to present day mammalian browsing is not convincing evidence that such plants were adapted to vertebrate browsing.
There is therefore little hope of settling the browsing question by this sort of debate, in which the evidence is indirect and disputed. We need a more sophisticated approach towards understanding the various adaptations and peculiarities of the New Zealand flora. Only by exploring a much wider range of ecological hypotheses than those currently proposed will we be able to realistically appraise the likelihood that some of those forms and structures can be explained only by herbivory. It is primarily for this reason that we suggest that the most fruitful way forward is the study of our plants in relation to the primary influences of the physical environment.
We thank our many colleagues both in New Zealand and elsewhere who have either endured with good grace our persistent attempts to destroy their belief in the presence of specific adaptations to moa browsing in the New Zealand flora, or have assisted us with debate and information. In particular, Ian Atkinson, Tony Druce, Eric Godley, John Lawton, Bill Lee, David Lloyd, Peter Wardle, Colin Webb, and Hugh Wilson have been generous with their time and insights. We thank Peter Wardle, Beverley Clarkson and Joanna Orwin for their critical reading of the manuscript. We also thank Mike and Rowan Glynn, and Joan Esterle for assistance with the field measurement of Pseudopanax branching heights under trying conditions.
Spores and pollen obtained from a condensed stratigraphic sequence of carbonaceous silts and muds from Porirua, near Wellington, indicate that the sequence straddles the Last Glaciation/Postglacial boundary. The lower part of the sequence contains dispersed rhyolitic glass shards identified as belonging to the Kawakawa Tephra. This, and a radiocarbon date on a basal Phyllocladus log of 21100 +/- 300 yrs BP, gives a maximum age for the base of the sequence. The spores and pollen indicate a change from subalpine Phyllocladus scrubland and/or grassland to lowland podocarp forest dominated by Podocarpus totara and Dacrydium cupressinum and a change from a cold, moist environment to a warm, humid one. The sharp junction between these two assemblages indicates an unconformity, but none is apparent in the monotonous sequence of silts and muds. Sequences straddling the Last Glaciation/Postglacial boundary without major and distinctly recognisable discontinuities have not been recognised from the Wellington are, but at Porirua it is suggested that much less time is missing.
Pollen analysis, pollen diagram, paleoenvironments, radiocarbon dates, Kawakawa Tephra, Last Glaciation, Postglacial, Porirua, Wellington.
In late 1988 an excavation, beneath what is now the Porirua K-Mart shopping complex, exposed a 1.7 m sequence of carbonaceous mud infilling a fossil gully. The site is about 50 m above sea level on the east side of the main road to Titahi Bay, Porirua, near Wellington, at grid reference R27/644 061 (grid reference for sheet R27 based on the national thousand metre grid of the 1:50 000 topographical map series INFOMAP 260). The fossil record number assigned is R27/f163 (New Zealand Fossil Record File number based on the metric INFOMAP260 series). A more general picture of the locality, which is about 1 km from the head of Porirua Harbour, is given by Mildenhall (1980) and Pillans et al. (figure 1; 1993). The sequence is a non-bedded, graded mud with secondary granules dispersed through it. The colour when wet is medium to dark grey.
A sequence of fifteen samples was collected in December 1988 by Brad Pillans and Brent Alloway (Victoria University of Wellington) for palynological analysis. Also noted at the time were pieces of wood, leaves and cladodes of Phyllocladus. Subsequently, a radiocarbon date on Phyllocladus wood from the base of the section, close to rhyolitic glass shards, which form about 5% of the fine sand fraction in the basal 40 cm and identified as belonging to the Kawakawa Tephra (Pillans et al. 1993), gave an age of 21 100 +/- 300 yrs BP (NZ7707). Since the glass shards are dispersed throughout much of the base of the sequence, and not in a discrete horizon, the tephra is almost certainly recycled.
The sequence overlies weathered Mesozoic greywacke basement, downthrown to the east of the Ohariu (Owhariu) Fault (Grant-Taylor 1976; Mildenhall 1980), and forming part of a depressed basin modified by a Last Glaciation river valley system which formed when sealevel was much lower (Stevens 1974). The subsequent rise in sealevel, which began about 14 000 years ago, was the catalyst allowing for Postglacial sedimentation to occur.
All fifteen palynological samples were examined and counted 500 grains were counted from each sample and the percentages of the various taxa calculated. In the summary pollen diagram (Fig. 1) the percentages of spores are based on the full count while those of the pollen grains are based on total pollen only. Only the most significant taxa (in terms of numbers) are presented, while the distribution of climatically and environmentally significant taxa are indicated. A brief summary of the palynological results, including the same pollen diagram, was originally published in Pillans et al. (1993).
The pollen percentages determined from the samples from 1.3–1.7 m are less reliable than the others, as they are completely dominated by Cyathea and smooth monolete spores (Blechnum?) leaving total pollen counts at less than 100 grains.
The sequence (Fig. 1; Zone A) records the drying and silting up of an acid, woody swamp, dominated by Phyllocladus, probably P. alpinus, and Halocarpus, through which fresh water flowed, and occurring in a moist, cool, subalpine grassland/shrubland environment. In the basal 1.2 m, pollen from Dracophyllum, Asteraceae and Poaceae is also common. The acidity of the depositional environment is indicated by the presence of numerous Gleichenia circinata spores. The lack of pollen from beeches and podocarps, other than Phyllocladus and Halocarpus, indicate that no substantial forests, or possibly even refugia, existed in the area at this time. The existence of fresh water is indicated by the presence of colonies of the shallow fresh water, lacustrine, alga Botryococcus. Since this alga requires some degree of warmth to exist, it suggests that the summers, at least, were warm enough to allow continual survival of this taxon.
This swamp changed into a full forest environment (Fig. 1; Zone B), dominated by tree podocarps, initially Podocarpus totara and Prumnopitys taxifolia, and subsequently
Thus, the top of the sequence at about 1.6 m above basement, where Dacrydium cupressinum first dominates, would be no younger than 9 500 yrs BP. If the exposed sequence represents sedimentation from the time immediately after the deposition of the Kawakawa Tephra, at 22 590 +/- 230 yrs BP (Wilson
Therefore the section is either condensed or the disconformity represented by the sudden change in the vegetation represents more time than the sediments themselves. A gap in sedimentation after deposition of the Kawakawa Tephra, representing much of the late Last Glaciation and a small part of the early Postglacial, is widespread in the Wellington area (Lewis & Mildenhall 1985; Mildenhall 1992a; 1992b), and elsewhere (McGlone et al. 1984), where it is recorded as being the result of landscape instability marking the onset of the maximum of the Last Glaciation.
The sequence also shows the following palynological features:
1. Unlike several other sequences of the same age in the Wellington area, for example Lindale, Paraparaumu (McIntyre 1970) and Wallaceville, near Upper Hutt (Harris & Mildenhall 1980; 1984), Nothofagus menziesii and to a lesser extent N. fusca type (pollen of Nothofagus fusca, N. solandri and
2. Most of the samples are dominated by spores which mask the character of the surrounding vegetation, especially in the upper part of the sequence. It is quite clear that the spores are local and come from plants growing in a damp valley or stream-side environment because they are generally well preserved, while the pollen grains are often poorly preserved and show signs of having been transported to the site of deposition by water.
3. The dominance of Phyllocladus (Zone A; Fig. 1) at the base of the sequence may prove to be useful in correlating other undated Phyllocladus dominant sequences in the Wellington area. However, Phyllocladus is not known from dated sequences to be as dominant at this time, although it is known to occur at some localities within this age range (Harris & Mildenhall 1980; 1984; Mildenhall 1987; 1992a), as well as much older localities (e.g. Mildenhall 1983). Other taxa, notably Nothofagus menziesii and N. fusca, are the more prominent pollen types from woody vegetation (Mildenhall 1992b).
4. The presence of some taxa gives additional information on the environment.
Epilobium grows in open spaces; its presence in the base of the sequence in tetrads shows that the genus grew at the site of deposition.
The presence of abundant Botryococcus, Haloragis, Myriophyllum, Plantago and Potamogeton in the basal 1.2 m of the sequence all indicate flowing, probably shallow, fresh water. The presence of numerous liverwort and hornwort spores (Anthoceros, Megaceros, Phaeoceros and the family Ricciaceae) suggests sheltered, moist, stream-side environments, with bare ground constantly available for these colonising plants. The frequency and abundance of Sphagnum spores indicates that much of the sedimentation occurred in a sphagnum swamp.
Fuchsia, found in the top 0.5 m of the sequence, is often found growing along banks of streams; it needs semi-shade for germination and seedling development. The presence of Pseudowintera often indicates disturbance in marginal forests; it also needs light for germination and seedling development.
The relative scarcity of Prumnopitys taxifolia and
The presence of pollen from the root parasite Dactylanthus taylori, towards the base of the sequence, is interesting for several reasons. Firstly, it is restricted to the North Island, where it is most abundant on the Volcanic Plateau (Macphail & Mildenhall 1980) and occurs no further south than Kaitoke, near Wellington (Aston 1909). Secondly, it is more tropical or sub-tropical in origin, and, finally, it is not common in the fossil record. Its occurrence at the base of the sequence in sediments deposited during a period of cool to cold climate to the south of its current range is unexpected. It may be recycled.
Most of the other taxa identified are wide-ranging in ecological tolerance.
There are several other sedimentary sequences in the Wellington region, containing the Kawakawa Tephra, that show similar characteristics.
At Wainuiomata the Kawakawa Tephra occurs near the top of a 61.6 m sequence (Begg & Brown 1991) which goes from fusca beech at the base, through menziesii beech into Phyllocladus. The topmost Last Glaciation samples are dominated by Poaceae (Mildenhall 1992b).
Similarly, at Taita Phyllocladus, with Poaceae and Nothofagus menziesii, occurs immediately below the Kawakawa Tephra with N. menziesii and Podocarpus and Prumnopitys species dominant above (Harris & Mildenhall 1980; 1984). The sharp changes in the vegetation above and below the Kawakawa Tephra indicate an unconformity and that considerable time may be missing from the condensed 2 m section at this locality.
At the site of the proposed new Pacific Cultural Centre and National Museum, Lambton Harbour, Wellington, 28 m of blue-grey to brown-grey silt occurs below 12 m of fill. At about 16–17 m glass shards, presumed to be from the Kawakawa Tephra, were identified in the field by R.D. Beetham (J.G. Begg pers. comm. August 1992). Here the entire silt sequence consists of grass-dominant assemblages, with Phyllocladus most common between 18–16 m (Mildenhall 1987).
Clearly, therefore, parts of the Wellington region, at the peak and towards the end of the Last Glaciation, contained a montane grassland/shrubland with sparse, probably stunted Phyllocladus as the dominant tree species (Mildenhall 1992b). Conditions were cold, constantly damp and probably windy, with temperatures the equivalent of the upper part of the Tararuas at the present day. The occasional pollen of more temperate taxa, like Dacrydium cupressinum, Dacrycarpus dacrydioides and
Brad Pillans and Brent Alloway, Victoria University of Wellington, are thanked for collecting the samples and for the use of some of their site information. Geoff Gregory, Institute of Geological and Nuclear Sciences Limited, and Bruce Sampson. Victoria University of Wellington, reviewed the manuscript and made useful suggestions for improvement. The pollen diagram was drafted at Victoria University under the guidance of Brad Pillans, who also reviewed a draft of the manuscript. The samples were processed by Roger Tremain to whom thanks are also extended.
Minute Landsnail eggs were collected in the field and incubated indoors. Successful batching of eggs was recorded for Serpho kivi (Gray), Phenacohelix giveni (Cumber), Therasia traversi (E A Smith). Omphalorissa purchasi (Pfeiffer), Flamulina perdita (Hutton), and Paralaoma species. Apart from Serpho kivi which was observed ovipositing, the actual date of laying was not known. Some notes on the life span of sub-adult snails collected in the litter are included.
There is an almost total lack of basic life history data on New Zealand landsnails. “For the numerically dominant Punctidae and Charopidae we do not know feeding specialisations, length of life, breeding seasons, annual or seasonal fluctuations in numbers.” to quote Solem et al. (1981) This paper records some opportunistic observations on the hatching times, growth rates, and life span of 22 species of minute landsnails from four North Island localities.
Hakirimata Range, Tainui Biological Region, Raglan District. Infomap 260-S14 973896 150 m.
Eggs were collected on 28 March 1982. The initial group of eggs was taken because it was possible to date the laying. A mature Serpho kivi was seen just completing laying a cluster of pellucid eggs in deep wet litter on the ground. The eggs were transferred carefully with Tawa ( Beilschmiedia tawa) leaves from adjacent litter. Extra litter was taken to prevent desiccation. The litter was housed in a 1.2 litre glass jar with lid, and daily examination and records were kept. The jar was kept in a room with good overhead light but no direct sunshine. Successful hatching occurred, so further collections of eggs were studied in the same fashion. A light mist spray was used spasmodically to maintain suitable moisture levels. If no snails were visible on inspection of the jar it was put briefly into sunlight.
This proved to be a good method of making the snails more active. Occasionally a check was made at night but active snails were seen infrequently. After eight months when there was no longer any sign of activity, the litter was checked again and then dried out. (Test ended 16 November 1982.)
Okareka. Northern Volcanic Plateau, Rotorua lakes. Infomap 260-U16 026304 400 m.
Small transparent eggs were collected on 10 April 1984 in a mamaku fern stem ( Cyathea medullaris) 30 cm off the ground. Extra litter was collected but care was taken to leave reasonable visibility in the jar. Many slime trails formed on the glass. The upper trails were removed with damp tissue on two occasions.
Minginui, East Volcanic Plateau, Whirinaki District. Infomap 260-V18 321725 400 m. A raft of flattened bluish eggs were collected 20 April 1984 - on bark. Small oval eggs were collected from the ground at the same time. Loose litter was also added to maintain moisture levels and provide food.
Kaimai, Roberts Farm, Northern Volcanic Plateau, Otanewainuku District. Infomap 260-U15 731679 220m. A collection of gelatinous, transparent eggs was
Serpho kivi. Eggs of this species from Hakirimata provided the most complete data on development, since their time of laying was known. These eggs darkened after 21 days and hatched after 58 days. When the litter was dried out only three shells were found. The maximum number of animals seen moving around the jar was 3, although a minimum number of 12 eggs had been collected.
Five weeks after hatching the snails gathered on the lid of the jar. They continued to move to a lesser extent but returned to the lid to rest. One snail died after 106 days. Two lived 132 days. The small shell was broken, the others measured 1.5 mm, 1.4 mm. Adult diameter 8.5–10.0 mm. (Powell)
Seven S. kivi from Kaimai hatched after only 30 days after collection. They moved around in the litter and one was recorded dead on the lid after 31 days. The remaining 6 were alive and on or near the lid at 38 days, but were not checked again.
Omphalorissa purchasi. Four hatchlings of this species from Okareka were seen after 61 days. The eggs had not been seen in the litter, and the snails lived in the litter for 280 days. Omphalorissa purchasi were not measured, the shells were missing when the litter was dried out.
Phenacohlix giveni. Twenty three hatchlings appeared after 25 days on 4 May 1984. Three died after 114 days, and one lived 171 days. The maximum measurement was 3.9 mm — one complete body whorl plus 2 riblets. Powell gives width as 5.00–6.00 mm. This indicates a growth-rate from hatchling to half-grown in less than six months in captivity. The colour pattern appeared after 39 days, before the first riblet formed. It was fortuitous that a succession of the fungus Mycena miniata, a toadstool, grew in the jar and was eaten by two species of snail. The Phenacohelix juveniles were frequently seen resting close by the fungus. The stipe of the fungus was eaten more than the cap. One Therasiella neozelanica was also seen adjacent to the Mycena. This food was not available for the second batch of Phenacohelix juveniles which hatched on 1 August 1984. This was 107 days since collection (time of laying unknown). These snails lived a minimum of 35 days and were not seen after this. Measurements were 0.9 mm, no riblets.
Therasiella neozelanica (Cumber). Two snails inadvertently collected in the litter were seen moving for 244 days.
At no time did the membranous plaits on the periostracum collect litter or dirt. For illustration see Cumber (1967). Growth was almost imperceptible during the period of captivity. Some plaits were damaged or missing when measured: 2.5 mm–3.0 mm. Maximum diameter of adults is 4.0 mm including plaits, (Cumber 1967).
Paralaoma sp. three snails were collected in the litter and seen for 143 days, but no trace of shells was found in the litter when dried.
Also present in the litter when it was collected were — Laoma marina (Hutton) lived 212 days, diameter 3.1 mm. Adult diameter 3.5 mm.
“Mocella” maculata lived 294 days, diameter 2.6 mm. Adult diameter 3.2 mm.
The following observations are all from the Minginui sample.
Therasia traversi. Thirty five eggs hatched after 38 days. They were all dead 63 days later, and lack of food was the most likely cause. The bark took up much space in the jar and therefore less litter was available to hatchlings. Colour pattern was showing after 27 days. Measurements were diameter 1.5–2.0 mm. Adult diameter 12.0 mm.
Paralaoma sericata. Three eggs hatched after 113 days in the jar. These were seen up to 28 days later but not again. On checking dried litter they appeared to have grown on for a further 42 days. Collected in the litter by chance were the following species, all were dead 36 weeks later.
Live snails seen during the observation period, but shells not found at the count included:
This would indicate that shell breakdown occurs in moist litter in a terrarium.
The final group of snails appeared in the Kaimai sample:
Flamulina perdita. 105 hatchlings emerged from the smaller eggs in the litter after 39 days (17 June 1989). Two weeks later 38 had migrated to the lid of the jar, but 30 were dead within two days. Five remained alive 39 days after hatching and were still growing. The smallest measured were 0.7 mm and the largest were 1.2 mm in diameter. Maximum diameter of adults 5.75 mm.
The following group were alive after 71 days:
It is possible to incubate minute landsnail eggs indoors without elaborate equipment and ultimately build up data on behaviour and feeding habits of particular snails. As no carnivorous snails were kept in captivity, the snails surviving longer periods found sufficient food on the leaf litter enclosed.
The small red-coloured fungus Mycena minata was a good food source for Phenacohelix giveni.
Eggs which showed blue colouration when collected hatched more quickly and it was observed that further colour changes occurred near hatching. For this reason only transparent eggs should be collected in order to gather more accurate data on hatching times. In this study the egg stage varied from a minimum of 25 days to a maximum of 113 days. Eggs of two arboreal species were included in these observations. Although laid at ground level, the newly hatched snails migrate upwards at 14 days in F. perdita, and 35 days in S. kivi.
Juveniles of Flamulina perdita may be more food specific than S. kivi with the result that they migrate earlier. Very small F. perdita have been observed at about 1 m and higher on the bark of Podocarpus totara and Dacrycrpus dacrydioides. Small snails included in the litter by chance are able to live for several months in captivity. Since they do not aestivate they can be assumed to be getting sufficient food and moisture. Seven to nine months was recorded for several species.
The growth-rate of Therasiella neozelanica in captivity was imperceptible
I would like to thank Dr. F. Climo for assistance, identification and encouragement over a number of years and Dr. G. Gibbs for editorial advice and comments.
The ecology of common woody plants naturalised in New Zealand is reviewed from the point of view of successional interactions between these plants and the indigenous flora. The reduction of fires, often associated with plantation forestry, has allowed pioneer stands of naturalised woody plants to grow to closed stands and to open up enough for indigenous plants to establish, at least in higher rainfall areas.
The naturalised plants are trees, shrubs and vines, grouped below in accordance with their ability to encourage indigenous trees and shrubs.
Groups 1–5, in higher precipitation areas, and often close to indigenous forest remnants are:
1: Pinus radiata, Ulex europaeus, Buddleja davidii,
Cytisus scoparius, Crataegus monogyna.
These plants, all light-demanding, initiate successions containing mesophyll (broadleaf) shrubs and small trees. The earlier indigenous nanophyll(small-leaved) shrub and small tree stages of Leptospermum scoparium and
2: Berberis darwinii, being relatively shade tolerant, but still capable of establishing in open sites, initiates an indigenous mesophyll scrub succession but persists under the canopy of the resulting low forest.
3: Calluna vulgaris. The density and youth of stands of this shrub do not yet allow a positive opinion on its role in succession. However, more open stands contain small Leptospermum scoparium, growing higher than the
4: Erica lusitanica. This low shrub can give way to the taller Leptospermum but because of the reproductive vigour of E. lusitanica its stands may remain monodominant. Their youth does not yet allow an assessment of the nature of succeeding vegetation.
5: Hakea salicifolia, Hakea sericea, Hakea gibbosa. These three species invade very leached soils. Succession has been recorded in stands of H. salicifolia and H. sericea, usually containing nanophyll indigenous shrubs that would have been present in the absence of these two Hakea.
6: Pinus contorta, Rubus fruticosus, Clematis vitalba. No succession has been seen in stands of this group of one tree and two vines.
Groups 5 and 6 form dense stands in areas of low soil moisture:
7: Rosa rubiginosa. Thymus vulgaris: these shrubs form dense stands in semi-arid areas, distant from any indigenous tree seed source; the only invaders are grasses and herbs, some indigenous.
8: Lupinus arboreus, Lycium ferocissimum. Found on edaphically dry and exposed coastal sites.
New Zealand, naturalised woody plants, succession to indigenous vegetation.
Plant names used here are those of Allan (1961), Brownsey et al. (1985), Connor and Edgar (1987) and Webb et al. (1988). Clmate records are from New Zealand Meteorological Service (1980)
New Zealand's relatively equable climate has allowed the vigorous growth of many plants naturalised from parts of the world where climates are more extreme. Fortunately almost all the woody plants naturalised in new Zealand are light-demanding and there are few cases of invasion of intact forest, such as is the case in Hawaii (Gerrish and Mueller-Dombois, 1980). However, much indigenous seral woody vegetation has been displaced, in its early stages, by naturalised shrubs and
Cockayne, (1928) describes communities of naturalised shrubs, such as Ulex europaeus, but does not describe any succession beyond the dense stands he found. It is probable that fires, both intentional and accidental, were more frequent than today and that there had not yet been sufficient opening up with age to allow entry of indigenous plants.
The first detailed account of succession beyond Ulex europaeus is by Druce (1957), where he details change at c. 40 years from
Since then an increased appreciation of the role played by naturalised woody plants in succession to indigenous vegetation has resulted in far more information being available on the interaction between naturalised and indigenous woody plants. This increased attention to naturalised plants also has two other causes:
Firstly, the increased areas of distribution of many of these plants has made them more obvious, and a matter of concern in protected natural areas.
Secondly, the stricter control of fire, particularly in areas of expansion of plantation forestry since 1925, has allowed development of naturalised shrub stands to the stage where successful establishment of other plants is possible in these stands.
Most of the woody plants whose ecology is described below were introduced intentionally in the earlier years of European colonisation in the 19th century. A full account of dates of first records in this country is given in Webb et al. (1988). Some common examples are: Pinus radiata for timber,
New Zealand is now a country with an array of domestic and feral mammal browsers. None of these mammals were here before the late 18th century. Only two indigenous shrubs ( Discaria toumatou and
The naturalised plants to be dealt with here are only those which have spread over considerable areas and densely enough to give their own physiognomy to the vegetation. Almost all the naturalised shrubs and vines are found almost exclusively in the warm temperate or montane /cool temperate zones. The bioclimatic zones are those of Wardle (1991). Only Calluna vulgaris and Pinus contorta are found in quantity in the subalpine and lower alpine zones, having been introduced in areas where open land is continuous to the alpine areas. In higher rainfall areas forested at the time of European arrival, restriction of the rest of the shrubs to lower altitudes is partly a function of very limited forest clearance in the subalpine zone, and the consequent existence of forest as buffer belts between lowland and mountain environments. East of the South Island mountains, largely deforested before Europeans arrived, it is probable that the more continental climate there imposes limitations on the upward migration into the subalpine zone of most naturalised shrubs.
The list below is arranged in approximate climatic order, from coolest and wettest, to driest climates. Distributions and common names come from Webb et al.(1988), with the distributions in countries of origin in italics. The botanical
(1) Upper montane to subalpine, high to medium precipitation.
Calluna vulgaris L. (Hull). (Ericaceae) Europe, Scandinavia, Asia minor, N. Africa. North, South, Campbell Islands. Heather
Pinus contorta Loudon (Pinaceae) North America. North and South Islands.
Contorta pine.
(2) Warm and cool temperate/montane
Pinus radiata D.Don. (Pinaceae)California,
Ulex europaeus L. (Leguminoseae)
Rubus fruticosus L. (Rosaceae) northern hemisphere temperate North, South, Stewart, Chatham Islands. Blackberry.
Hakea gibbosa Cav., Proteaceae. E. Australia. Northern North Island. Downy hakea
Hakea salicifolia (Vent.) Burtt, Proteaceae. E. Australia northern North Island and northwestern South Island. Willow leaved hakea.
Hakea saricea Schrader et Wendl. E. Australia N.Z. distribution similar to Hakea salicifolia. Prickly leaved hakea
Buddleja davidii Franchet,
Berberis darwinii Hook. (Berberidaceae) southern South America. North (localised), South and Stewart Islands. Darwin's barberry.
Clematis vitalba L. (Ranunculaceae)
Erica lusitanica Rudolphi (Ericaceae) southwestern Europe North and South Islands. Spanish heath.
Cytisus scoparius L. (Leguminoseae) Europe. Asia Minor, Russia. North, South, Stewart, Chatham and Campbell Islands. Broom.
Crataegus monogyna Jacq. (Rosaceae) Europe. North and South Islands. Hawthorn.
(3) Cool temperate, low precipitation.
Rosa rubiginosa L. (Rosaceae) Europe, N. Africa. North, South, Stewart and Chatham Islands, but more common in drier climates. Sweet brier
Thymus vulgaris L. (Lamiaceae) Mediterranean. South Island, dry inland areas. Culinary thyme
(4) Coastal, high to medium precipitation.
Lupinus arboreus Sims. (Leguminoseae)
Lycium ferocissimum Miers. (Solanaceae) Southern Africa. North and South Islands. Boxthorn.
The most extensive and obvious invasion by Calluna vulgaris is within Tongariro National Park, which surrounds and includes New Zealand's active mainland volcanoes. Much of the landscape here, from 600 m up to c. 1600 m is dominated by Chionochloa rubra, a tussock up to 1m high and with orange leaf ends, which impart a golden glow to the landscape. This tussockland spread from naturally poorly drained sites and alpine areas because of forest destruction by pre-European fires. The soils, of low pH, and often compact and moderately drained, were ideal for Chionochloa rubra, whose dense stands appear to slow down invasion by shrubs. The tussocklands were, when not reburnt for early farming, being invaded only slowly by indigenous plants. Species included Leptospermum
scoparium at lower altitudes and
Calluna vulgaris was introduced intentionally in 1912 in the northwest sector of the Park, as food and shelter for the introduced game bird, the grouse (Lagopus lagopus scoticus). The grouse did not survive, but Calluna vulgaris did and is now expanding its area at the expense of the tussock lands and other more open subalpine zone vegetation. In an area along the Mangatepopo Rd., tussock with subalpine scrub, at higher altitudes, was burnt in 1948. The tussock had regenerated all over by the late 1950s, and was invaded by Calluna vulgaris in the 1960s. The cover of Calluna now is almost complete with only sparse suppressed tussocks surviving. Away from the area of the 1948 fire, in most of the north and west of Tongariro National Park, Calluna vulgaris has also spread extensively into tussock and indigenous heath communities which had not been burnt in the mid-20th century. Up to about 1200m, where the Calluna vulgaris stands are not completely closed, Leptospermum scoparium grows through more open stands of
Chapman and Bannister (1990) did not find rapid invasion by Calluna vulgaris of Dracophyllum subulatum heath in the study area. Dracophyllum subulatum has a similar physiognomy and stature to Calluna vulgaris. The least spread of Calluna recorded by these authors has been in boggy ground. Although Calluna vulgaris produces seed only below 1200m altitude (Chapman and Bannister, 1990), the lightness and quantity of this seed has enabled it to establish at 1600m, above the treeline, either blown or adhering to mountaineers' muddy boots. The capacity of Calluna vulgaris for vegetative reproduction ensures its spread there.
In a similar way the distribution of Calluna vulgaris along highways and railways near Tongariro National Park has been aided by man. A plant, found in the Tararua Conservation Park (and dug out !) 250 km south of Tongariro National Park could easily have travelled there on the boots of a mountaineer.
Calluna vulgaris is also locally distributed at The Wilderness Scientific Reserve, near Lake Te Anau. Although only small plants remain in the reserve, after control operations there are more mature stands nearby (P.Bannister, pers. comm.) There is still a risk that Calluna, uncontrolled, may suppress the ground flora beneath the tall bushes of Halocarpus bidwillii in the reserve.
Calluna vulgaris is one of our most unfortunate introductions. In Tongariro National Park Calluna was introduced into a mountain area of great natural beauty and with a unique indigenous vegetation. Calluna is easily dispersed naturally, and its dispersal is assisted in an area of high recreational use. The same trampers often go to other, distant mountain areas, and seed can easily go with them.
This plant matures in relative shelter to a small tree and many of its invasive attributes are shared with naturalised shrubs. It seeds precociously, as early as seven years of age, in New Zealand. Its early growth is often as a multibranched shrub and it resprouts when cut back, even at ground level.
The first infestation of Pinus contorta to spoil a natural landscape was on the southeast slopes of Tongariro National Park. In contrast to the formerly tussock covered northwest area of this Park, much of which is now covered with Calluna vulgaris, the southeast side of Mt. Ruapehu, from 1000–1800 m altitude was of a desert aspect with sparse vegetation. There is a series of alluvial fans of loose sandy tephra, where substrate instability is maintained by the strong fohn winds in the rainshadow of Mt. Ruapehu, by frost at any time of the year and, near the Whangaehu River, by episodic extensive lahar floods. Rain is adequate for forest growth, 1100mm annually at 823m altitude, and wetter with altitude. Substrate
On the edge of this “desert” landscape a plantation of P. contorta was established in the 1930s. By the 1960s saplings were establishing on non-vegetated surfaces. By the 1970s there was a dense scrub of P. contorta advancing into intact tussock grassland 10–15km north of the original plantation. By this time also there were isolated plants established in the lower alpine zone open vegetation, often well above the altitude of the nearby treeline. Control measures today in the National Park include hand pulling and grubbing out, much done by volunteers from mountain clubs. To the southeast of the National Park lies the Waiouru military reserve, an extensive area of tussock grassland, of Chionochloa rubra and, at lower altitudes Festuca novaezelandiae, the only extensive area of the latter tussock in the North Island. There are interspersed forest patches, mostly of Nothofagus. Over the last 10 years the Army has undertaken an extensive cutting and burning programme, hoping to eliminate Pinus contorta. The native tussocks respond to burning by good new regrowth.
The second phase of P. contorta introduction started in the 1950s, when foresters decided that, as the South Island's tussock covered mountains east of the Southern Alps had been forested until fires of the 13th and 14th centuries, they should reforest these mountains. This planting was done with the aim of reducing erosion, much of which was then believed to have resulted from pastoral activity in the time since European arrival. While the vegetatively reproduced Salix spp. were used for torrent stabilisation, and Alnus viridis for scree planting, the most commonly used plant on open slopes was Pinus contorta. Hand planted at times, but also directly seeded from the air, Pinus contorta has become established not only on unstable slopes, but also in intact tussock grassland and indigenous open scrub. Allen and Lee (1989) found that, of the three conifers spontaneously establishing in tall tussock grassland, (Larix decidua, Pinus nigra, P. contorta), the latter showed greatest tolerance of the proximity of tussock bases of Chionochloa rigida. This result was ascribed to a possible greater shade tolerance of Pinus contorta, a supposition that could explain its ability to invade indigenous vegetation with a range of canopy densities, from semidesert to open scrub. None of the plants of these communities can survive in the pine's shade.
At about 40–50 years age the Pinus contorta plantations on the south side of Mt. Ruapehu are dense and dark beneath with no growth under them, indigenous or naturalised. They were felled around 1980. In its native territory, of inland North America, Pinus contorta stands can remain at a dense stage for over a century, with little or no self-thinning. This character may be related to the slight shade tolerance noted by Allen and Lee (1989). It seems likely then that this conifer will not provide a starter for indigenous successions for a very long time
In some South Island areas of conservation and tourist interest, attempts are being made to remove P. contorta, but the job is even more vast than in the earlier invasion of Tongariro National Park.
Extensively planted in large blocks, for timber and as windbreaks, this tree, which can reach 40m height in 30 years is now the most noticeable naturalised plant in New Zealand.
From planted trees the wind dispersed seed travels into more open indigenous communities, and trees may establish and ultimately suppress the local vegetation. Julian (1983) described P. radiata establishment, from seed from nearby trees on to the site of Leptospermum scrub, burnt 10 years previously. Although there was vigorous scrub regrowth, this was accompanied by the establishment in the first 3 years after the fire of clumped stands of P. radiata saplings, whose lateral growth
From this stage an estimated 30–40 years has to elapse before the establishment of a dense indigenous understorey, generally of mesophyll (Specht, 1979) shrubs.
Many plantations of Pinus radiata were established, after burning, on the sites of nanophyll (Specht, 1979) scrub or heath. After 25–30 years of pine growth, a tall mesophyll and tree fern scrub develops, very similar in appearance to the understorey of an indigenous forest. The understorey which developed under
Under 25–30 year old Pinus radiata, near Tokoroa, with 1600 mm of annual rain, a dense indigenous understorey developed. The soils are sand to silts from the Taupo Pumice eruption of the second century A.D., overlying deep, clayey, fine tephra of earlier Holocene age, Previous to planting, the vegetation burnt was a mosaic of
Under the changed light and soil conditions, the first plants of the understorey were tree ferns. By 30 years of pine age, on slopes where the pumice was shallow, older moister tephra were in root range, and a dense understorey developed. This understorey was of broadleaf shrubs together with tree ferns, up to 10m high Together with most of the other shrub species in this understorey, the tree ferns were completely absent from the scrub and heath that was burnt before planting. On deeper pumice soils of infilled valleys, the tree fern understorey persisted, with little invasion by shrubs.
Close to an area of intact indigenous podocarp forest a few seedlings of Dacrycarpus dacrydioides and
Following winter clear-felling of the pines, with lack of heat for pine seed release ther was little establishment of pine seedlings. Many of the species of the crushed understorey resprouted, and in the first years the light-demanding nitrogen fixer Coriaria arborea increased from a few scattered plants to form a locally dense cover.
When the pine was felled in summer, its cones opened and its dense seedling growth soon suppressed any indigenous regrowth.
As the aim of these plantations is the continuing production of wood, all indigenous and pine regrowth is now defoliated and burnt before replanting with selected stock.
The species of the Tokoroa understoreys are all mull forming, and associated with fertile seral sites (Druce, 1957), would be expected on the youthful Taupo Pumice.
In the Rai-Whangamoa forests of the northern South Island, Jelinek (1977) reported vigorous invasion by indigenous species of a Pinus radiata forest after thinning at 30 years of age. The deep humus under
On former Nothofagus sites, on deep weathered loess near Upper Hutt, with rainfall of 1298 mm annually, Vella (1984) decribed an open understory under Pinus radiata about 45 years old.
From Wellington, Smith (1979) described understorey composition in pine plantations of 45 years, with rainfall of 1240 mm. These plantations are on steep slopes with skeletal soils. On east to south slopes, the following species were abundant, reaching to about 3m height, but not forming a dense scrub: Coprosma
grandifolia, C. robusta, Brachyglottis repanda, Melicytus ramiflorus, with the ferns
The Wellington shrub assemblage under P. radiata reflects the fertility of a skeletal young soil. The site, on analogy with indigenous forest remnants in Wellington probably carried a mixed forest, with some podocarps, in a Beilschmiedia tawa matrix. The Wellington stands have a completely different shrub understorey from the Upper Hutt stand, described above, by Vella(1984), where the older soil, having previously supported
In areas of reasonable moisture supply, where Pinus radiata has been felled after the development of an indigenous understorey and no attempt is made to regenerate the pine, a mesophyll scrub of indigenous plants results. For example such scrub is very visible from the interisland ferry, on the east of Picton Harbour, the pines there having been felled in the last 10 years. Dependent on the proximity of seed trees, such scrub and its counterparts in other areas, could eventually lead to tall indigenous forest, where commercial forestry does not go beyond a first rotation.
This fearsomely spiny shrub, with bright yellow flowers is one of the most widespread and visible of New Zealand's naturalised shrub flora. Newsome (1987) records 254 000ha of land occupied by Ulex europaeus although only on 20 000 ha is
As scattered clumps in pasture, Ulex europaeus is common throughout New Zealand, from high precipitation areas of the South Island west coast, down to c.600 mm of annual rain in the east. Introduced last century as a hedge plant, its spread on to pastures is only checked by browsing of domestic stock when pasture growth is inadequate, mostly in drier areas. It is in these areas that gorse hedges are still successfully maintained.
In higher rainfall areas, as stock numbers decrease with weaker grass growth on unfertilised pastures, Ulex europaeus is able to expand and often, after repeated fires, displaces indigenous seral plants such as
Ulex europaeus is a plant with strong reproductive capacity. It bears seed young, and the seed is catapulted from the pod by sun or fire heat, and can last up to 30 years in the ground. Fire stimulates its germination (Zabkiewicz, 1976). Plants of all ages can reproduce from root-shoots after fire and can coppice from cut stems. Its growth rate is high, Lee et al.(1986) describe rates of 20cm/yr. for height and 0.5cm/yr for diameter growth. The maximum height recorded by them was 7m with diameter (at 10cm height) of 21.7cm. This is certainly more vigourous growth than the 1–2m high plants of Ireland or Portugal.
Ulex europaeus is relatively short-lived in New Zealand. Druce (1957) recorded 46 years as a maximum near Wellington. Usually by 40 years
Druce (1957), Healey (1969) and Oates(1988) all record, from central New
Ulex europaeus scrub by indigenous species, generally broadleaf shrubs within time spans from 15 to 30 years of
Ulex europaeus stands were studied by Oates (1988) on former
All three studies cited above are in areas of moderate rainfall of 1000–1200mm annually, and have mild climates. Ulex europaeus litter breaks down under these conditions, allowing seeds of indigenous plants to germinate.
Ulex europaeus enriches the soil with more nitrogen than under an equivalent indigenous sere (Egunjobi, 1971). The addition of this nutrient can result in a richer soil, with consequent changes towards more fertility-demanding species than were present in the original forest.
Under the cooler conditions of the southern South Island, near Dunedin, Lee et al. (1986) reported that Ulex europaeus litter breaks down slowly and can reach 75cm in depth. Under these conditions there are few plants, indigenous or naturalised, in the
1. Melicytus ramiflorus is present in both areas.
2. Griselinia littoralis occurs only in stands near Dunedin, reflecting the cooler climate of that study area.
3. Kunzea ericoides occurs only in stands near Dunedin, no doubt reflecting the openness which allows decomposition of the deep
Apart from its role in succession on old forest sites Ulex europaeus also invades open ground in many circumstances. On an intact shoreline forest edge south of Haast,
Although Ulex europaeus grows well on leached, former forest soils, it will not invade the poorly drained pakihi heathlands of the South Island west coast, growing only along artificial drainage channels through these wet areas.
The high reproductive capacity of Ulex europaeus has enabled it to displace indigenous, early stage woody seral vegetation. This displacement has always been aided by fire in the indigenous woody vegetation, as
These two similar needle-leaved shrubs to small trees are widespread on the North Auckland peninsula in the very leached soils of gumland scrub, on old Agathis sites. Hakea sericea is, as well, abundant on leached soils of northwest Nelson.
Enright (1989) found that the two species rarely co-occur in the gumland heath of North Auckland, Hakea sericea being characteristic of very leached sandstone soils and in young vegetation on clays, while Hakea gibbosa was found on less leached sandstone soils. Neither species was found on more fertile volcanic soils.
Fire opens their persistent woody seed capsules and accelerates the invasion of the resulting clear ground. Beever (1988) reported, from a site burnt five years previously, young plants of Hakea gibbosa and Hakea sericea, with mean densities of 1.6/m2 for Hakea sericea and 0.7/m2 for Hakea gibbosa.
From northwest Nelson, in the Abel Tasman National Park, Esler, (1961) reported that Hakea sericea excludes all other plants of seral scrub and that it and Hakea salicifolia (below) invade unburnt scrub. Williams, (1992, in press), from Northwest Nelson reported that Hakea sericea lives to about 15–20 years before stand collapse by windthrow. When Hakea sericea plants are growing with Leptospermum and Kunzea shrubs these species continue a succession towards indigenous forest. When, however, Hakea sericea is in dense pure stands, stand collapse is followed by invasion of Pteridium esculentum, which can in turn initiate a sere back towards forest.
This shrub to small tree, up to 5m high, has flat leaves, to 11 cm long. It is widespread and often dense in Leptospermum and Kunzea scrub in northwest Nelson, including a serious infestation of “several thousand acres” (Esler 1961) within the Abel Tasman National Park. Here the soils are leached, although well-drained, deep granitic clays. Indigenous scrub succession on them is often very slow or even apparently static (Gabites, 1979).
Williams (1992), in this National Park, found evidence of succession on to indigenous vegetation, following invasion of scrub by Hakea salicifolia. As Hakea salicifolia does not produce annual rings he aged stands from Kunzea ericoides associated in the same stands.
On the shallowest soils, however, Hakea salicifolia is still invading the mixed Leptospermum-Kunzea scrub, a scrub which has shown very little change since first described by Dumont d'Urville in 1827.
R. fruticosus grows as a scrambling shrub, capable of layering. It has relatively soft leaves, up to 16 cm long, and intensely barbed stems. Introduced as a fruit plant last century its spread by seed has been accomplished by defecation by birds, pigs and humans. As well, its vegetative reproduction allows broken fragments to establish new plants on stream beds after floods.
Fortunately R. fruticosus is not tolerant of the shade of New Zealand's evergreen forests, so it has not invaded intact forest. Its main distribution is in areas of higher rainfall, throughout the Auckland province and western side of both
Rubus, are now being used again, as free ranging animals. They present a hazard to indigenous vegetation when not controlled themselves. Rubus fruticosus grows well on leached poorly drained soils. As such it can be a serious hindrance to the regeneration of Nothofagus forests on the west coast of the South Island.
The density and tendency to collapse, of the low canopy (2m) of R. fruticosus prevents invasion by indigenous shrubs.
Although limited in distribution at present in its invasion of indigenous communities, this garden species' strong colonising ability in protected natural areas is a matter of concern. Smale(1990) reports on its occurrence on gravel river beds in Urewera National Park, and Brown (1990), in the Rimutaka Conservation Park, near Wellington. Dobson (1979) also reported its spread on gravel riverbeds near Kaikoura. There is a range of precipitation in these three sites, in the same order, from 2400 mm down to 888 mm annually. Erosion is active on shattered grey-wackes of the three areas, and the resultant depositional surfaces are colonised by Buddleja davidii. It is a short lived plant, from 15 to a maximum of 20 years. Within this time its initially dense stocking declines and seedlings of indigenous, broadleaf woody plants establish. Among these Smale (1990) recorded
This spiny leafed shrub, which can grow to 5 m tall, is a common forest understorey and seral scrub plant in Nothofagus forests of southern South America, in climates very similar to New Zealand. Eskuche (1968, 1969) records Berberis darwinii as common in both evergreen tall forest and in shorter deciduous forests of Nothofagus. As well, Berberis darwinii can dominate scrub following fire in these forests(Eskuche, 1969; Hildebrand, 1983). The spread in New Zealand of Berberis darwinii, from garden plants to localised dense patches has only been recorded since 1946 (Webb et al. 1988). It has small fleshy fruit and is easily transported by both naturalised and indigenous birds (Allen, 1991). As well as by seed establishment, Berberis darwinii spreads vegetatitively by suckering. So far it is present as an invader of open land from the southern end of the North Island down to Stewart Island in areas of regular rainfall. A study by Keller (1983) near Wellington showed that Berberis darwinii colonises low fertility pastures, resists browsing by stock, and forms a tight divaricating form in wind exposed areas. As the Berberis darwinii scrub closes canopy and this canopy gets taller, indigenous plants will establish. In Wellington Keller (1983) recorded, beneath Berberis scrub about 2–3m high, seedlings and saplings of the small trees Carpodetus serratus,
Near Dunedin Allen (1991) recorded, as principal associated indigenous plants: the light-demanding Kunzea ericoides, (whose quantity has an inverse relation to that of the
As the Berberis canopy becomes taller and begins to open the indigenous plants emerged and form a low secondary forest. In this forest Berberis darwinii persists as small lianoid trees beneath the canopy. Allen and Wilson (1992) found that these forest conditions are better suited for germination of Berberis darwinii seeds and establishment than open condition. Therefore Berberis darwinii is likely to remain as a constituent of the forest, as it does in its natural distribution in South America.
There has not yet been a study of the chronology of successions involving Berberis darwinii. It is impossible to say whether Berberis darwinii simply persists in an indigenous forest resulting from succession through Berberis darwinii or whether Berberis darwinii in fact actively invades an intact indigenous forest.
There are few shade tolerant naturalised plants in New Zealand, but the presence of Berberis darwinii, considering its spread within half a century, is a danger to the indigenous purity of forests, particularly to remnant and reserved patches of lowland forests in farmland.
This liane, originally a garden plant, is normally of modest growth in its Northern Hemisphere climates, which have more severe limitations of temperature and precipitation than New Zealand. Here the seeds are easily carried by wind, so Clematis vitalba has spread far from settled areas into open forest remnants and on to the edges of closed forest. The growth habit of
Clematis vitalba thus presents a very real threat not only to the smaller forest patches already scarce in the New Zealand lowlands, but even more to those farm areas where smaller trees form an attractive woodland and summer shade for stock. Control of
This nanophyll shrub, which can grow to 3m high, is common in milder climates of both main islands. Mather and Williams (1990) from whom this account is drawn, reported that it cannot tolerate frost at above 400m altitude in Canterbury, nor can it grow in areas with low rainfall. Erica lusitanica can tolerate leached and degraded soils in a similar way to Leptospermum scoparium but has reproductive
Cytisus scopurius is widely distributed throughout New Zealand, but reaches its greatest densities on freely drained alluvial or tephra-derived soils, (Hunter and Blaschke, 1986). In this respect its edaphic tolerance parallels its distribution in Europe, where it is regarded as an indicator of deep friable soils. (author's observation.) The distribution of Cytisus scoparius in New Zealand excludes the driest areas, and those areas considered to have a marginally Mediterranean type climate (McQueen, 1984).
This distribution accords with its European distribution in climates without a pronounced summer drought; Cytisus scoparius is not, as Williams (1981) believes, a plant of truly Mediterranean climates (Braun-Blanquet 1951, Fournier 1961).
Cytisus scoparius is a fastigiate, almost leafless shrub, which grows to 2.5 m high. It spreads easily, by seeds catapulted from the pod in high natural temperatures, and the seedlings will grow in as little as 10% of full light (Willams, 1981). Growing with a single leader Cytisus scoparius has a competitive advantage over the frequently associated Ulex europaeus.
Where there are forest remnants close by, in areas above c.1000mm of annual rain, indigenous broadleaf shrubs and small trees can establish under Cytisus scoparius. Williams (1983) describes one such succession where mixed stands of Ulex europaeus and
Cytisus scoparius has a wide range of soil fertility tolerance, particularly to phosphate (Williams, 1981), which enables it to grow equally as well on a leached former forest soil as on a young volcanic deposit. It also actively colonises the broad gravel beds of aggrading rivers, and the old gold dredge tailings of the southern and western South Island.
A shrub or small tree, Crataegus monogyna has spines on its stems, but younger plants are browsed back by stock (Williams and Buxton, 1986.) These authors descibed its establishment and spread in the eastern foothills of the Southern Alps, in about 1500mm of annual rain. Here Crataegus monogyna establishes in the protection of the indigenous spiny shrub, Discaria toumatou, and can grow in this protection until above browse height. In pastures where
A spiny shrub, to 4 m tall, this deciduous shrub of medium height is is widely distributed in New Zealand pastoral land. Rosa rubiginosa is mapped as dominant (Hunter and Blaschke, 1986) in the northeast of the South Island, with c.700mm of annual precipitation and in the driest parts of the intermontane basins of the central South Island, with down to 340 mm annual precipitation.
Rosa rubiginosa reproduces by seed in a fleshy fruit capable of bird transport and favoured by wild pig as an autumn food. The shrub also reproduces vegetatively, by rhizomes and root suckers. Consequently Rosa rubiginosa, with its curved spines, forms thickets impenetrable by man, stock or the dogs used to move the stock.
Molloy (1976), describes its ecology on the northern area of dominance on Molesworth Station, with precipitation around 600–800mm annually. He shows that Rubus rubiginosa is mostly distributed on actively eroding soils, up to the upper limit of Festuca novaezelandiae low tussock grassland (750–1200m). Plants aged from their rootstocks show that their first establishment was in about 1937. The virtual elimination of rabbits in the 1950s had little apparent effect on recruitment of Rosa rubiginosa, and the absence of new plants after the early 1960s was attributed to competition from the vigorous, aerially sown pasture plants sown for the reclamation of depleted rangeland.
Eastwards of the area described by Molloy (1976) in higher rainfall Williams (1989) describes dense Rosa rubiginosa scrub, in an area where there has been little land management, except the rabbit control of the 1950s. Rosa rubiginosa has been in this area since earlier than 1953, when it was mapped as dominant by McQueen (1954). Here Rosa rubiginosa grows in an impenetrable scrub, 4m high, sometimes with Discaria toumatou and the divaricating
Further south, in the intermontane basin of Central Otago, Partridge et al. (1991), in a study of vegetation on the slopes above a major gorge, found that Rubus rubiginosa was most commonly in a species-poor community of steep rocky faces, with the indigenous low-growing divaricate shrub Melicytus alpinus. Away from this detailed study area, the more gentle slopes and flat areas of Central Otago have become increasingly covered with open scrub of
A low (to 60cm high), multibranched shrub, aromatic and with small grey leaves, this naturalised plant has a restricted but dense occurence in Central Otago, ia region of 340mm of annual precipitation. The altitudinal range of Thymus vulgaris there is from 250–750m. The texture of the schist-derived soils beneath Thymus vulgaris varies from gravels of old gold workings to remnants of the silty loess of the remaning topsoil. These soils are usually near neutral acidity, because of the aridity of the area.
Wilkinson et al. (1979) give the following account of its biology. Originally introduced in the 1860s as a culinary herb, Thymus vulgaris has spread to occupy some 2 000 ha densely, and a much larger area as scattered plants. It flowers and seeds prolifically and the seeds, within calyces, are easily wind-carried as well as
Much of the land which Thymus vulgaris has occupied is that which Cockayne (1928) described as man-made desert (see above, Rosa rubiginosa). The main cover was of the indigenous mat plants Raoulia spp. After the rabbit populations had been severely reduced in the 1950s, the vegetation surveyed in 1967 (McQueen, 1981) included breaking up mats of Raoulia, prostrate species of the indigenous grass Rytidosperma, and an ephemeral cover of naturalised spring green herbs and grasses. By 1990 Thymus vulgaris had spread on to many of the sample points of 1967. In all areas invaded by Thymus vulgaris there is little trace of the previous vegetation. Plants commonly found with Thymus vulgaris are the indigenous grass Elymus tenuis, and the naturalised
Thymus vulgaris is one of New Zealand's more agreeable invaders. It is not spiny, it smells nice, makes good honey and its low greyish growth harmonises with an arid landscape.
There are two shrubs which have modified the landscapes of New Zealand sand or gravel coasts. These are Lupinus arboreus and
This low shrub, up to 2 m high is widespread on coastal dunes, introduced intentionally to stabilise the backs of the foredunes on aggrading coasts, or where pastoral attempts had caused renewed sand movement. Lupinus arboreus has suppressed most indigenous plants of the inland sides of foredunes.
Lupinus arboreus is also common on steeper scree slopes in some coastal areas Fuller (1985), and on gravel and sand riverbeds stretching to some scores of kilometres inland.
A tall shrub at times 5m high, this spiny, tightly branched plant was introduced for hedging in coastal areas. Its fleshy fruit and probable bird transport have resulted in its spread along many New Zealand coasts, both on sand dunes and on gravel storm beaches. Once established it layers from lower branches and can form large dense clumps. Here the indigenous pioneer vegetation is dominated by the mat forming Muehlenbeckia complexa. Lycium ferocissimum is usually scattered in this simple community, and may spread further inland into coastal pasture.
Both Lupinus arboreus and
Naturalised woody plants in New Zealand occupy a wide range of habitats, but the majority are light-demanding pioneer plants and thus prone to be eventually part of succession back to indigenous forest. This succession is governed by availability of indigenous seed, and it is only in areas of adequate rainfall and closeness of forest remnants that the naturalised plants can give way to indigenous vegetation. Such succession has been documented for Pinus radiata, Ulex europaeus, Berberis darwinii, Cytisus scoparius, Erica lusitanica, Buddleja davidii. In most cases succession through this group of trees and shrubs results in the following indigenous vegetation being of mesophyll shrubs. The presence of the naturalised plants usually leads to the omission of initial nanophyll indigenous successors, such as
In two cases of widely distributed woody plants there is evidence of local differences in the composition of the successional vegetation:
1. The understorey of Pinus radiata varies with soil conditions; on rich, young soils the understorey plants are large-leafed and mull-forming, on older leached soils many species of the pine understorey are smaller-leafed and mor-forming plants, often those of indigenous forests on older leached soils.
2. Under stands of Ulex europaeus and
In higher rainfall environments, in forest remnants are two shade-tolerant species: Berberis darwinii which can survive within a secondary forest, and Clematis vitalba which can cause canopy dieback by physical suppression of large trees in forest remnants.
The naturalised shrubs of semi-arid areas, Rosa rubiginosa and Thymus vulgaris, show that much of the grassland areas found by Europeans last century are capable of supporting woody growth. All except the driest of these areas have evidence of pre-13th century forest, and in fact still have remnant areas of indigenous shrubs. The distance from indigenous forest seed sources, in these oftburned lanscapes, so far prohibits any succession back to indigenous forest.
The coastal spread, on rocky beaches and sand dunes of the erect shrub Lycium ferocissinum as a primary coloniser, and Lupinus arboreus on sand dunes are examples of the introduction to New Zealand of life forms not previously indigenous in these environments. Neither of these species appear yet to be initiating successions towards indigenous vegetation.
I am very grateful to the post-graduate students, cited in the text, who have provided much of the detail of dynamics of naturalised woody plant successions in the Wellington area. I should also like to thank co-workers who have provided photographs and the School of Biological Sciences, Victoria University of Wellington for assistance in photographic processing. Pamela Searell, Drs. Mary McEwen and Bruce Sampson provided excellent editorial criticism. Much of the travel involved was financed by the Internal Research Grant Commitee of Victoria University of Wellington, and the Miss E. L. Hellaby Indigenous Grasslands Trust.
Three of the following articles are based on papers presented at a symposium in 1989 entitled “Contributions to the Scientific Knowledge of New Zealand by the Early French Voyages”. This was sponsored jointly by the Alliance Française de Wellington and the Wellington Branch of The Royal Society of New Zealand. The fourth article by J Bartle was partly based on his contribution to the general discussion.
The background to the early French and other voyages of exploration was the intense interest in Europe in the expansion of knowledge in a scientific sense during the “Age of Enlightenment”. This applied particularly to the little known Pacific Ocean, both in terms of natural history and also as a means of testing out the concept of “Noble Savage” uncorrupted by civilisation.
Let me state at the outset that the term “philosophical” is to be used in this talk in its appropriate eighteenth century context, at a time and place when a philosopher was first and foremost a lover of knowledge, committed, as the full title of the Royal Society so aptly put it, to the improvement of knowledge in every field. The philosopher was not a specialist, but still Plato's synoptic man who took a comprehensive view of the universe and consequently a researcher to whom no field of enquiry was closed.
The most striking aspect of eighteenth century research is the appearance of encyclopaedic works of all kinds. It was as though the man — and the woman — of the Age of Enlightenment had said “Let us first gather all that we know, all that we believe, all that we suspect, so that we may then go on to discover what lies beyond the clearly defined frontiers of knowledge”.
And so the eighteenth century opens with Pierre Bayle's great Dictionnaire historique et critique, over 3000 pages long, while in England John Harris publishes his Lexicon technicum; in Italy Coronelli struggles with his Biblioteca universale, although after seven volumes he had only reached the letter C; in Germany Hübner supervises the writing of the Reales Staats Zeitungs und Conversations Lexicon and Zedler gets to work on the massive Grosses vollstandiges Universal Lexicon.
But it was in France that an encyclopaedia became a manifesto of the philosophical movement as well as a massive compendium of human knowledge. It had been conceived originally as a straightforward translation of Ephraim Chambers' Cyclopaedia, published in two volumes, of 1728. It became a 21 volume collection of articles and essays by the leading writers and philosophers of the day:
More specialised encyclopaedias were also being written, which often were more than works of reference: in keeping with the enquiring and above all challenging spirit of the time, they were also calls to action.
The driving force behind many of these compilations was a desire for knowledge which made itself felt outside the world of science. It spread into and through the educated middle class and, in France particularly, reflected a trend away from the literary salons and the fashionable passion for writing poetry, letters and plays which had turned the seventeenth century into the great age of European classicism. A fascination for a knowledge of nature and the physical world, soon transformed into a devotion to scientific enquiry, was combined with a critical examination of society.
Social analysis, eventually associated with a re-assessment of political and religious structures and of how they fitted into broader philosophical concepts, was brought about by the rigidity of French society and the resulting weakness of the French social system, but it also encompassed comparative analyses of institutions outside France. The political systems of England, Austria, Russian and the German states, were examined and envied or rejected, while correspondents throughout Europe added their contribution to the ferment of analysis. Curiosity spread beyond the boundaries of European society. China and some other eastern countries were known, very imperfectly, for their products and their art, and their despotic forms of government; the Near East held a lesser fascination while what was known of the African continent was unappealing; but North and South America had produced a rich harvest for students of natural history. There could not be boundaries to a world which had so much to reveal to an age bent on unveiling and analysing all that Man and Nature had to offer in such overwhelming diversity. The philosophes were driven by an Aristotelian desire for completeness, for a grasp of the entire panorama of human knowledge and of the physical world.
This curiosity, this thirst for new discovery and to some extent for sheer novelty and the exotic, lay behind many of the activities and the reactions of those who entered the Pacific during the second half of the eighteenth century. They certainly formed part of the mental make-up of the readers who, in large numbers, seized enthusiastically upon the accounts of Pacific voyages and eagerly discussed what was reported about an ocean and places that were both fascinating and frightening.
Expectations and responses depended on this European vision of the Pacific. It was a part of the world whose immensity was appreciated, now that the miscalculations of sixteenth-century Spanish navigators had been swept aside.
It was a blank space on the world charts that was still cluttered with imaginary islands, mysterious nations, fabulous animals left over from the imaginings of earlier cartographers. With enthusiasm and expectancy, the eighteenth century appeared to clear up this now accessible but still only vaguely perceived region of the globe. Exploration and scientific observation would as the philosophes expected, “improve human knowledge,” They would also offer possibilities for trade and for the consolidation of imperialist aspirations — but those are other matters best, at this juncture, overlooked.
In keeping with the approach of the savants, the first step was an encyclopaedic compilation, a broad but complete survey which would also show what needed now to be done.
The most influential such work is Charles de Brosses' Histoire des navigations aux terres australes of 1756. It was a historical survey of what had been achieved in the southern seas and the Pacific and, probably more important, what remained to be done.
It was not a compilation for its own sake, as was the Abbé Prevost's Histoire générale des voyages which also appeared, in many volumes, in the 1750s, itself inspired by John Greens A General Collection of Voyages, and Travels of 1746, but a systematic analysis of Pacific exploration encased between a lengthy essay on the
philosophes were eager to promote actual exploration.
De Brosses' background is of interest because, although he belonged to the philosophe movement, he was not a parisian, but a resident of Dijon in Burgundy. His work and his subsequent influence were in no way lessened by his provincial background. Indeed, he provides evidence of the movement being not exclusively centred on Paris, but being active throughout France and throughout Europe.
He was a typical man of the Enlightenment, wide ranging in his interests and tireless in his studies. He had already written on Roman history, on archeology and on the origins of the languages. When Pierre de Maupertuis' open letter to King Frederick of Prussia, On the Progress of the Sciences was read at the Dijon academy in 1752, De Brosses had expanded his views on the importance of exploration to the advancement of science in a Memoir which he presented to the academy. This led the great naturalist, Buffon who was a patron of the Dijon academy and the author of the great multi-volume Histoire naturelle, to suggest to De Brosses that he expand his memoir into a full volume (eventually there were two), and to underline the likelihood that voyages of exploration would result in the discovery of new plants and new animals.
Letter recognised the thoroughness of de Brosses' work and was, so far as he could be, sympathetic. philosophes over social progress and the real benefits of civilisation.
For scientists, knowledge and discovery were goals to be pursued without the need for philosophical justification. The world was there to be discovered, its secrets waiting to be revealed. The good that would result depended, not on what was discovered, but on the use which man made of it.
Others — and they were numerous — who speculated on the evolution of social and legal systems — were less definite. Laws seemed to exist to protect property rather than the individual, the status quo rather than innovation. And as society became more complex, man became less free, more exploited, less happy. Montesquieu's L'Esprit des lois went back to the origins of laws to discover what principles guided their emergence and their evolution. But
Viewed from the standpoint of the French political and legal system which had become rigid, ineffectual and corrupt, earlier societies, especially small pastoral or agricultural social units, aroused a nostalgia which gave rise to the simplistic myth of the Noble Savage. Civilisation seemed to have corrupted man, oppressed him, entangled him in a web of clumsy laws. The ownership of property, the accumulation of personal wealth were to a large extent the causes of this corruption.
Why therefore travel to distant seas to bring the chains of civilisation to simple people who, noble savages living in isolation, were undoubtedly happier than the civilised poor of Europe?
Reverse the proposition and you get the scientist's eagerness to test a theory, for if on faraway islands there could be found some unspoilt savages, the theory would have been proved right. Some aspects of their behaviour could be identified and copied in order to reform European civilisation. And as a by-product, newly discovered plants and animals could be brought back to Europe and used for the
One cannot either overlook the religious impetus: Christianity could be brought to the newly discovered lands. Unacceptable moral practices could be eliminated and salvation assured to nations hitherto deprived of the message of the Gospels, even though, to many philosophes, the spreading of the power of the established church, or churches, was in no way a benefit for which any Noble Savage would be likely to be grateful.
The exploration of the Pacific was not envisaged as what we now know it to be: sailing across vast empty seas, making occasional landfalls on small islands and atolls. The great blank on the charts of the eighteenth century was confidently expected to be filled by a continent, or at the very least by extensive tracts of land.
For those who kept an open mind, there was simply no reason why the south Pacific ocean should be void of lands at least as large as Japan or the Dutch East Indies, of places as populated and developed as Ceylon, or even as vast as South America. For a number of eighteenth century geographers and cartographers, the dictates of geophysics actually required the presence of large land masses to preserve the equilibrium of the globe. The theory had an elegant simplicity look at any terrestrial globe in any learned geographer's study, remember the relatively newly discovered laws of gravity, and it would soon become evident that, if the earth had not already toppled over, it was because the continental masses of the northern hemisphere are counterbalanced by roughly equal land masses in the southern hemisphere, some of which clearly remained to be discovered.
The expectation that new countries and in all likelihood new civilisations would be found was a significant component in several voyages. Surville sailed, rather hopefully — ál'aventure, as was stated — in search of an undiscovered land, spurred on by garbled reports that the English had come upon the fabled Land of Davis. Bougainville, who openly admitted his debt to De Brosses was more openminded about the existence of a southern continent, but he nevertheless hoped that if new lands were there to be discovered, they would fall to France through his endeavours, rather than to the rival English who were undeniably planning a world strategy which included in it the control of the Pacific. And Alexander Dalrymple, an influential man in his day and an admirer and friend of De Brosses, had no doubt at all about the existence of a southern continent.
Although there was no continent, if one excepts the Antarctic which is not really what these geographers were speculating about, there were islands still waiting to be added to the map. The Solomons, for instance, which Surville and Bougainville were to visit and report on, had been discovered by the Spanish but, in the absence of reasonably precise longitudes, had been lost again: the French were fairly confident they did exist and that exploration would solve what had unaccountably become a geographical riddle.
“Discovery”, of course, is a term which is now open to some criticism, as it disparages the achievements of the real first discoverers of the Pacific islands who were Polynesian or Melanesian wanderers: but its true meaning is real enough, namely to find something which was not known to the rest of the world. The explorer's task was twofold: to determine as precisely as possible the geographical location of the place newly discovered, and to describe its appearance, its climate, its flora and fauna and the way of life of its inhabitants, if any.
Most eighteenth century navigators were adequately equipped to carry out this descriptive function. They were sufficiently educated to be accurate and methodical: they often had some training and, if not the captain, at least some of the officers had studied botany, zoology, and sometimes geology. Linnaeus had introduced a rational methodology for the naming of species, and in France men
Conditioned to be systematic and responsible in their observations, they were also preconditioned when it came to assessing the primitive societies they encountered. Bougainville, who followed Samuel Wallis to Tahiti, found a peaceful and friendly people. He was not to know that Wallis had cowed them into submission by the use of firearms, and consequently he put it down to the naturally good and uncorrupted nature of primitive people. He was disabused in due course, but not so the philosopher-naturalist Philibert Commerson who hailed the island as the true abode of the Noble Savage. Out of his enthusiasm, shared by others on board and spread by his writings among the French and European public, the lasting myth of an idyllic Tahiti was born. Commerson had found what he expected to find, an expectation fulfilled strengthened the theory on which it was built, so that a number of later voyagers would sail in the belief that the South Seas harboured island paradises.
The first Frenchmen to come to New Zealand were different. Seemingly unaware of—or untouched by — the writings of social philosophers, their expectation of New Zealand was built on the reports of the only Europeans to have preceded them more than a century before: the Dutch. Abel Tasman had been attacked without provocation, so that the natives of New Zealand were expected to be fierce and bloodthirsty. They were not: no indication that the Maoris were particularly warlike was apparent when Surville and his men put into Doubtless Bay. The French were pleasantly surprised and for most of the fortnight's stay relations were highly amicable.
A quite different expectation was carried to New Zealand by the next visitor, Marion de Fresne. Although an experienced seaman with a fine wartime record, he was something of an optimist and a dreamer. He had accepted the idea of the Noble Savage with few of Bougainville's reservations and in fact he had undertaken to return to Tahiti the islander, Ahu-toru, whom Bougainville had taken to France.
Marion du Fresne first called at Tasmania where he endeavoured, with little success, to befriend the aborigines. The Maoris of the Bay of Islands, however, rapidly lived up to his expectations and to the reports which James Cook (down-playing the loss of life which had marred his first contacts) was giving of them. His overconfidence led him to overstay his welcome; he had no inkling of the laws of tapu and little suspicion of the extent of intertribal warfare — and with a number of his officers and men he paid the price with his life in June 1772.
The optimistic theories of
The role of the learned societies should not be overlooked. Their function was to assist with the instructions that were issued to the various captains. La Pérouse received pages of them — from the Academy of Science, the Society of Medicine, the Dean of the Paris Faculty of Medicine, from the Royal Geographer Buache, from Buffon, and from the King's Gardener. D'Entrecasteaux, although his primary duty was to search for La Pérouse, received lengthy detailed instructions from the Academy of Science and the Society for Natural History. There were also experiments to be carried out, such as recording sea temperatures in specified locations, testing new chronometers, new methods of preserving food, desalinisation and the role of insects in the corruption of drinking water.
In the nineteenth century, instructions became centralised through officials of the Navy and, although the influence of the learned societies did not lessen, their independence was restricted, while gentlemen who wished to have their scientific theories tested during a naval expedition to the Pacific were requested to go through the proper channels.
The background against which nineteenth century navigators set out, moreover, had changed. The theory of the Noble Savage had faded. Attacks by natives who feared and resented the arrival and presence of strangers destroyed the idea of peaceful societies of island dwellers; a growing knowledge of class distinctions, feudal island structures, slavery, warfare and cannibalism destroyed the notion that civilisation undermined the essential and natural goodness of man. Science itself was becoming increasingly specialised, the amateur gentleman being elbowed out in the process. Experiments were carried out under far more rigorous conditions. The eighteenth century had prepared the ground, surveyed the horizons, and now it was time to get down to precise analysis and detailed, disciplined study. The writings of Dumont d'Urville are evidence of the new trends: whereas previous generations had painted broad all-embracing canvasses, the accounts of his voyages are loaded with details, quotations, references to sources to an obsessional degree, which makes the massive volumes untranslatable in their entirety. Even his attempt at writing a novel set in New Zealand, lengthy enough in itself, is further burdened by a massive appendix of explanatory notes.
The degree of co-operation which existed between philosophers and scientists of different nations during the Age of Enlightenment and into the nineteenth century was quite remarkable when one remembers how often the great powers were at war with each other during these years. The attitude of these research workers, amateur and professional, was well expressed in the title of a study by Gavin de Beer: The Sciences were never at war (1960). Sir Joseph Banks, as the most influential Englishman of his day in the field of science and exploration, played a significant part. He helped La Pérouse and D'Entrecasteaux, and willingly provided information and advice to would-be navigators, supplying scientific equipment and helping to release French collections of natural history specimens impounded during the wars. Documents were issued to explorers by all European rulers — the kings of England, France and Spain, in particular — so that all might sail without let or hindrance and with no fear of capture should war break out while they were on the high seas bereft of news from Europe and with no hope of rescue.
James Cook, La Pérouse, D'Entrecasteaux, Baudin, were protected in this way, but even without a specific passport genuine scientific expeditions had little to fear. It was understood that the search for new knowledge was a task that transcended national rivalries. Admittedly, local governors sometimes had doubts or were afraid of what their home government might say if they allowed a blockade runner through under pretext of exploration or research. But it was indeed the Age of
The friendship and mutual help one encounters in the relations between navigators and scientists from different countries are some of the most gratifying and impressive features of the age of the philosophes. The age of merchants and colonisers which followed it cannot lay claim to the same fidelity to the ideals of science and learning.
The contributions to botanical knowledge of New Zealand resulting from French voyages from 1769 to the settlement at Akaroa in the 1840′s are evaluated. Translations of general comments by the French botanists on aspects of the flora and vegetation of New Zealand are provided.
It is clear that a certain amount, perhaps a considerable amount of national rivalry was involved in the voyages of discovery. Scientists, however, like to believe that science is international with the shared common purpose of advancing knowledge. This is not a new idea. Montagne in his introduction to the botanical account of D'Urvilles last voyage says
“When the noble emulation which reigns between great nations such as France and England, both placed at the forefront of civilisation, has for its purpose only to expand knowledge and does not degenerate into a jealous and deplorable rivalry, it can only be eminently useful to science and the whole of humanity.”
Both the earliest French and British voyages visiting New Zealand in the 1760′s and 1770′s were not as productive botanically as they might have been. Extensive and very valuable plant collections were made on Cook's voyages but the accounts of these from the first voyage were prepared, but never published and the published botanical results of the later voyages were mostly very inadequate. No plant collections appear to have been made during the two French visits in the same period. Crozet in his journal from Marion du Fresne's ill fated voyage did provide some general comments on the plant cover of the Bay of Islands as well as some geological speculation which has turned out to be remarkably far sighted.
“Might not the subterranean fire, which formerly burned and vitrified so much matter in New Zealand, have also by several shocks detached this island from new Holland or from the Austral lands or from some other continent?”
He comments on the widespread occurrence of bracken fern which we now know would have largely followed destruction of forest by fire. He also remarks on some of the larger forest trees, possibly pohutukawa and northern rata and certainly kauri which he terms the “olive-leaved cedar”. He was very impressed by the size of these trees, their abundant resin and the quality of the wood which he judged “very suitable for making ships' masts”.
He also commented on the fact that even in winter he did not see “a single tree shed its leaves”.
It was not until 1824 that another French expedition visited New Zealand. This was the first of 3 voyages involving Dumont d'Urville, although on this occasion the commander was Duperrey.
By now there was a strong emphasis on scientific discovery. Unlike British voyages no scientists as such were included but some of the officers and particularly the surgeons were also reputable scientists. For instance D'Urville would have regarded himself as much a botanist as a navigator.
As far as Botany was concerned observations were made of the vegetation and plant specimens were collected. The latter are mostly preserved for later study by drying and pressing to make what are known as herbarium specimens. The plants
Some of the collectors had some artistic ability and sketched living material, but most of the often handsome illustrations in the published accounts were made by artists at the Paris Museum aided by the sketches and notes of the collectors.
There was little about New Zealand plants in the publication on Duperrey's voyage, perhaps because many of the species at the much visited Bay of Islands had already been described. In his journal René Lesson was rather unimpressed by New Zealand vegetation partly because he was seeing it at a place where the forests had been greatly reduced by fire.
“The season of our visit was not a good one for collecting botanical specimens. The flowering season was over, and although there was vegetation, it was verdant only in ravines and damp places; on the mountain sides it took on a reddish appearance from the closely packed mass of a fern with edible roots. The forms of vegetation are few and monotonous; very different from the splendour and profusion of tropical plants. In their uniformity and dreariness they are nothing like the plants of New South Wales, resembling rather the vegetation of Chile. Some hills are covered with trees of medium height, with dull grey foliage like an olive tree (probably pohutukawa J.D.). Large trees, birches and pepper plants grow in the sandy bays. I found no edible fruit, apart from a kind of small bluish plum (tawa or taraire) which the plump pigeons swallow whole. The korarou and the Phormium grow in damp places, while an Oxalis and a small daisy were the only plants flowering on the turf at the time. Trees which because of their hardness and their great size, are most suitable for maritime construction, are found in the interior of New Zealand. This timber and the linen plant (Phormium) are the most commercially desirable products”. “There seems to be little variety in the vegetation of the bay region. I was there in autumn of these parts at a time when the flowering season was partly over; my herbarium was enriched by only 5 or 6 plants with their flowers. There are scarcely any trees, except in the gullies, but as the soil is composed of a deep layer of humus, the trees there attain the most majestic proportions. The hill tops are bare of shrubs, and from a distance they seem to be covered by bright green turf, owing to a thick growth of fern 2 or 3 feet high. As one goes inland, vegetation increases, the bush is thicker, trees of very hard red and black wood rise on the slopes, and the soil is kept damp by a carpet of the pretty kidney-shaped trichomane.” “Vegetables planted by Marion du Fresne grow abundantly.”
The second voyage of D'Urville, this time as commander, visited more localities in New Zealand in 1827 — the northern part of the South Island and points on the eastern North Island coast and in Northland, including the present site of Auckland, and finally again the Bay of Islands. Extensive plant collections were made mostly by A. Lesson (younger brother of R. Lesson), and on their return A. Lesson and A. Richard, a notable botanist at the Paris Museum, prepared and published in the account of the voyage a detailed account of the vegetation of New Zealand as well as descriptions of 380 species of which 41 were illustrated. The account was entitled “Essai d'une Flore de la Nouvelle-Zelande”. About this Cheeseman later commented “This is the first publication dealing with the flora of New Zealand as a whole, and possesses considerable merit, so much so that it is regretted that so little use has been made of it by New Zealand botanists”.
As the authors had seen several localities in New Zealand including some where the forests were in their original state they were more favourably impressed by the flora and vegetation than was R. Lesson.
“Among the many countries visited by the Astrolabe, during its voyage of circumnavigation, there is none more interesting than New Zealand, a region little known up to now from the point of view of geography and natural history. The researches of the officers and naturalists, under the command of Captain D'Urville, have given results of great importance. Confining ourselves to that which specially concerns Botany, the plant collections made by M. Lesson the younger, pharmacist of the expedition, added to those Captain D'Urville himself managed to collect amongst more important activities which occupied his time have enabled us to gain a much better understanding of the original vegetation of New Zealand.
If one reflects on the few localities that have been explored by naturalists, and above all the short time that their excursions have lasted, one will easily understand that there still remain many species to be discovered. Furthermore, we do not pretend to have made a complete Flora of New Zealand, but simply an essay in which the plants that have been observed up to now have been brought together.
Independently of the species we have observed ourselves and which belong to the collections made by Captain D'Urville and Lesson the younger during the last voyage of the Astrolabe we have had at our disposal a great number of type specimens, collected by Forster himself, which form part of the rich herbarium of the Muséum d'Histoire Naturelle de Paris. These original specimens have been very precious for us, in giving us the means of naming with certainty, by direct comparison, a part of the species collected on the voyage of the Astrolabe. But an advantage no less important which will be appreciated and shared by all botanists is that we have had access to Forster's manuscripts. It is known that a large number of the species mentioned by this botanist are often known only by the simple diagnostic phrase, which he gives in his Prodromus. Furthermore several of them are so uncertain that they have been omitted in the lists of some authors. We have been able by this means to remove the doubts concerning several of these species, either by ourselves making detailed descriptions from the original Forster specimens, when we had them in front of us, or by publishing the manuscript descriptions for the species we do not hold or of which we have only specimens that are too incomplete or in too poor a state. We have been careful to acknowledge these borrowings; either by citing Forster's manuscripts or by indicating by the symbol † the species described by the voyager.
Before giving an overview of the relations of the vegetation of New Zealand with those of various regions of the globe we are going to sketch, from the notes of M. Lesson and verbal information furnished by Captain D'Urville, the general appearance of the country. It is traversed by mountains which are progressively higher the further they are removed from the coasts. At Astrolabe Harbour (Tasman Bay) these mountains attain a great height and several are covered with snow. Near the anchorage they rise almost vertically from the little sandy beaches of the shore. Once one has attained the summits one can easily go from one to the other, and one can enjoy from the heights of these peaks an entrancing view. Most of them are covered with ferns, several with trees and shrubs. The humus layer is often very thick; this soil is blackish, light, still filled with plant debris and watered by a large number of torrents and streams.”
“We said above that the humus layer was generally thick: this probably applies only to the littoral explored by the navigators, as the crests of some mountains they climbed had a thinner layer which could only support ferns and lichens; some small trees form sparse tufts on the eminences of Tasman Bay. In Hauraki Bay and its various arms, where several islands were discovered by Captain D'Urville, only
Dacrydium cupressinum, Podocarpus dacrydioides and several other green trees, whose descriptions we will give, probably form the basis of these aggregations. Astrolabe Harbour, in Tasman Ban is one of the places where MM. D'Urville and Lesson made the most abundant collections. What we are about to say about the vegetation of this locality will suffice to give a general idea of that of the whole of New Zealand, because these diverse regions are similar to each other, with the exception of some plants which seem to characterise some regions, but which might well occur in other parts of the archipelago which is still not completely explored. The vegetation of Astrolabe Harbour is very beautiful, although the number of cryptogamic plants almost equal the seed plants. The European is surprised to discover here some plants of his own country or at least closely related species. For example one finds here wild celery (
Ferns are remarkable for their number and diversity. One sees them above all in the shade of forests where a spongy soil, composed of plant debris, favours their growth. They reign there almost alone as their great abundance chokes the few phanerogam herbs which try to establish there. One finds a great number of them on the stems of trees, where they develop in the manner of parasites, in ravines, and as far as the almost bare rocks of the mountains. Lichens are, after ferns, the most abundant cryptogams. They grow on soil, rocks and trees. The crawling stems of certain polypodiums and other ferns are often garnished with these lichens, above all in moist places; for example, in the vicinity of cascades.”
Following D'Urville's second voyage there were two further visits to the Bay of Islands by French expeditions — La Place in 1831 and Dupetit-Thouars in 1838, but little of botanical significance seems to have resulted.
Decaisne in Dupetit-Thouars described and illustrated a few New Zealand species. One of these is something of a mystery as it is an attractive daisy shrub from the Chathams—Olearia semidentata and so far as I can discover the venus did not visit the Chathams. Perhaps the specimen was given by someone on a whaling ship at the Bay of Islands that had come from the Chathams although the name of the collector — Pfihl — is certainly not French.
D'Urville's third and last voyage reached New Zealand in 1840. This time the Auckland Islands to the south of New Zealand were visited, as well as Otago Peninsula, Akaroa and inevitably the Bay of Islands. It is of interest that the Auckland Islands were also visited that year by the eminent English botanist
The published botanical results of this voyage were not so extensive as those of the second partly due to the death of Hombron, one of the botanical collectors, before the account had been finished. However, one important contribution was the publication of several new species of seaweeds and mosses from the Auckland Islands.
This was the last French voyage of discovery to visit New Zealand but this was not the end of the early French contribution to New Zealand Botany. The French colony was established in Akaroa in 1840 and the surgeon with the support vessels, Etienne Raoul, was also a keen botanist.
“The corvette L'Aube, on which I embarked as surgeon major, left Brest under the orders of M. Lavaud, captain of vessel, the 19th February 1840, and after two stops, one at Sainte-Croix de Ténériffe, the other in Senegal, arrived at the Bay of Islands on the 11 July. The very advanced state of the season and the abundant rain only permitted me to collect at that time some well known species in the environs of Kororareka, Pahia, Waitungui. L'Aube left the Bay of Islands the 31 July and on the 15 August lowered anchor at Akaroa, Banks Peninsula, from where it did not depart until 21 November 1841. The first stay, encompassing a complete series of seasons enabled me to collect many botanical specimens in their diverse states. During a second appearance of the Aube at the Bay of Islands, from 2–13 December 1841, I augmented with a good number of species the limited collections made in 1840. After returning to Akaroa on the 26 January 1842, after a short voyage to New Holland, L'Aube was replaced by L'Allier, onto which Lavaud carried his flag and of which the health service was confided to me. L'Allier did not leave Akaroa until the 11 January, 1843 and made its return to Europe, stopping for a third time at the Bay of Islands. During these different excursions to the Bay of Islands and especially during the long sojourn at Banks Peninsula I was able to gather most of the species, to describe and draw in fresh conditions most of them, which make up the Flora of this part of the South Island, up till now little explored.
On my arrival in France, in October 1843, my collections were deposited in the Museum of National History of Paris, and at the request of MM. Brongniart and de Jussieu, the Minister of Marine authorised the work which I prepared under the supervision of M. Decaisne who oversaw the classification of the herbarium specimens and their analysis and willingly corrected the deficiencies in my notes on a subject of which I had not made in advance a sufficient study.
My plants have been compared with Forster's, contained in the museum, and with some of those used by M. Richard for his work on the Vegetation of New Zealand.”
As an evidence of the international character of science he warmly acknowledges the assistance and advice of Joseph Hooker during a visit he made to the Paris Museum and who later sent specimens from Kew for comparison with Raoul's specimens.
“J.D Hooker, to whom the flora of the antarctic regions is so well known, pointed out to me, during his visit to Paris, several new species in my herbarium, and later sent me a series of specimens from New Zealand compared by him with Forster species kept at the British Museum. The result of this double comparison is that under the same name there exist in the Museums of Paris and London different species collected by Forster himself. These identification errors have resulted in some errors on the part of botanists who have consulted one or other of these herbaria; thanks to the goodwill of M. Hooker I have corrected them. 1 am equally obliged to MM. J.D. Hooker and Taylor for the determination of my lichens, liverworts and mosses, which M. Léveillé has kindly described.”
The result of Raoul's work was a handsomely illustrated book — “Choix de
After completing his book Raoul was sent to tropical Africa for medical work, but he later returned to his home town of Brest where he died at the very early age of 37.
Belligny was another botanist in the Akaroa colony. He had been on the staff of the Botanical Gardens in Paris and although he did not produce any publications he sent back specimens of plants (including seeds), birds and insects.
The early French contribution to New Zealand Botany was clearly considerable. Consulting Allan's more recent account of the New Zealand Flora I have determined that although only about 10 early French botanists actually visited and collected in New Zealand a total of 55 French botanists were subsequently involved in the describing and/or naming of New Zealand native plants.
Three phases of French involvement in New Zealand zoological discovery, dating from the late 18th century are identified and described.
The zoological discovery of New Zealand, that process by which specimens of its fauna were collected and described, was made by individuals representative of several different nations. It was a collective effort and it seems almost quibbling to single out or compare the contributions made by individual countries. Nonetheless, if an occasion or a need arises to evaluate the contribution made by one nationality or another, then such evaluations can sometimes be revealing in terms of national character, the way in which scientific problems were approached, and the basis on which approaches were made. Some evaluations of this kind have already been attempted (Andrews, 1987) and the present paper takes some of that work a step further. Appropriately, in this case, we look at the French on the occasion of the bicentennial of the French Revolution, itself a major influence on science in that country.
The topic under discussion will be examined in comparison with the British contribution in another paper in this collection (Bartle). Meanwhile the present paper sets out to describe some aspects of the early French discovery and collection of New Zealand zoological specimens.
Three distinct phases characterise French involvement with New Zealand zoology. They are:
1. The late 18th century, in which the French, along with other Europeans, took advantage of (a) the specimens brought back from Cook's voyages and (b) the ensuing hiatus that resulted from the English naturalists failing to make the most of their opportunities in terms of description and publication.
2. A period when the French undertook their own voyages of exploration in the Pacific with specific scientific objectives.
3. A colonial phase to which the small French community in New Zealand, and its supporters, made a modest contribution.
In each case there were elements of imperialist science — a response to the demands of scientists in France, based in an institution — in this case the Muséum d'Histoire Naturelle or its antecedents.
First what was its context? Did the initial exploration of New Zealand by the French arise out of a thirst for scientific knowledge; was it a simple desire for conquest; or was it based on commercial or trade considerations? Or was it simply, as seems more likely to be the case, a combination of these things.
To understand the reasons for these early explorations of New Zealand we have to go back to the Seven Years War and the intense rivalry that existed between Britain and France (Dunmore, 1965). This rivalry was to continue beyond the
In the event the first French voyage to reach New Zealand was that of Jean de Surville in command of the St Jean Baptiste, in 1769. In contrast to Cooks voyage which, fortuitously, reached New Zealand at the same time, there was no scientific motive attributed to the voyage. It was privately organised and New Zealand was encountered more or less by accident. De Surville was looking for Tasmania and refreshment for his crew, many of whom were suffering from scurvy, when he changed course and encountered New Zealand instead.
Their stay in New Zealand was a short one, and their scientific observations limited by time and the absence of a natural historian. In spite of these deficiencies the First Officer, Guillaume Labé, recorded the presence of lizards which the English naturalists on the Endeavour (then not far distant) had failed to record or collect, although they must have seen them (De Surville, 1981).
Labé also reported some fish and several species of birds, but again none of these were collected or given any form of scientific description. They released two pigs which thus featured among the very first introduction of European mammals to New Zealand. However in all probability the pigs did not last very long, and were eaten by the Maoris. De Surville himself did not survive the voyage back to France, drowning off Peru, leaving Labé to report that the first French contact with New Zealand and its fauna, flora and indigenous people had been made.
Following de Surville several decades passed before the French again set foot in New Zealand, but French naturalists, like others from Europe were able to acquire specimens brought back from the South pacific by the three voyages of James Cook. Although Cook's voyages by a number of measures were very successful, there were considerable problems describing the countless plant and animal specimens they brought back with them. Into this vacuum came a number of European scientists who were either correspondents of Joseph Banks or who visited him in London. Banks held on to voyage collections, but he was generous in lending or donating specimens to fellow naturalists (Andrews, 1987).
In this way some of the species in Banks' collections were able to be given proper descriptions and scientific names according to the Linnaean system of classification which had achieved a fair degree of support by that time. Regrettably not all were treated like this and substantial parts of the collections were originally described using non-Linnaean names. Why the English naturalists and their Danish colleague Daniel Solander, who was one of Linnaeus' pupils, were unable to get their collections published in the scientific literature is a matter of debate and speculation, the detail of which need not concern us here. The overall consequence of the free-for-all that followed was that many of the specimens went to Europe and acquired scientific names that had German, Danish and, of course, French authors.
The French were particularly well positioned to gain access to the English collections. They were geographically close and could visit Banks relatively easily or, later, attend the auctions of collections that included Cook voyage material. Civil turmoil in France in the late 18th century positively encouraged this traffic.
Molluscan shells found early favour with the French, as they were universally sought after by naturalists and collectors alike. Jacques de Favanne and his son were responsible for publishing a work called La Conchliologie in 1780, which is one of the earliest formal records of French interest in New Zealand zoology. Being collectible items many shells might have found their way from Banks' and
One French link with New Zealand zoology that lasted well into the 19th century was initiated by Guillaume Olivier, a founder of the Linnaean Society of Paris (Andrews, 1987). Olivier came to London in 1789 to examine the Banks collection of insects, the year of the French Revolution. The animals he described were rather non-descript Hemipterans which in itself is surprising as the tendency in those days was to favour large and more colourful species. Although his visit and subsequent study took place at the time of the Revolution, he managed to get his work published. He was assisted in his work by Pierre Latreille, a conservative priest and entomologist who had endured a spell in prison at the hands of the Revolutionary authorities. He did not suffer unduly from his incarceration and lived to arrange the entomological collections of the Muséum d'Histoire Naturelle and achieve other scientific distinctions.
Another Frenchman to visit the Banks collection was Pierre Broussonet, between 1780–82, who worked on the fish under Solander's supervision. Broussonet described the carpet shark from New Zealand using non-Linnaean nomenclature, calling it “Isabella”. The scientific name of this shark was later provided by another Frenchman, Bonnaterre, in 1788. The carpet shark thus became the first properly described and named New Zealand fish. In 1789, like so many others, Broussonet was forced to flee France, becoming physician to the United States Embassy in Morocco.
Before moving on from the fish some mention must be made of Georges Cuvier the famous French anatomist who was to dominate French zoology in the early 19th century. He was supplied with tracings and copies of fish painted or drawn by the artists on Cooks voyages. The copyist was Sarah Bowditch who with her husband, became closely associated with Cuvier in Paris. One New Zealand fish which was described in this way was the common yellow eyed mullet, Aldrichetta forsteri (Andrews, 1987).
Apart from the almost inexplicable appearance of the tui in a French book on African birds by Le Vaillant, at the end of the 19th century (Stresemann, 1975) there is little else to report on this first phase of zoological contact between France and New Zealand. Although it relied heavily on the Cook collections it was meritorious in view of the fact that a surprising amount of study and publication was achieved in one of the most turbulent periods of French history, and almost entirely on the basis of gifted or borrowed specimens.
A revival of French interest in the Pacific in the early decades of the 19th century saw some scientifically well equipped expeditions come in the direction of Australia and New Zealand. Not all of these were outstandingly successful, but the three voyages that reached New Zealand were. It was these voyages and the post-revolution development of the Muséum d'Histoire Naturelle, as well as the distinguished group of naturalists based at the museum, that formed a solid basis for the second phase of discovery by the French of the New Zealand fauna and flora.
The first voyage of significance was that of Duperrey and the Coquille in 1824. Among the naturalists, were R P Lesson and Prosper Garnot.
They did not spend long in New Zealand, and all their collecting was carried out in the vicinity of the Bay of Islands, which was reached in April 1824. They
One of the interesting features of this voyage was the better balance struck between invertebrate and vertebrate animals. It was clear that there was increasing interest in invertebrates as the 19th century wore on — whereas earlier naturalists were preoccupied with vertebrates — particularly birds and to a lesser extent fish. From this voyage Lesson was able to describe the rare flax snail — the first terrestrial snail to be described from this country and the common pipi. Duperrey's voyage might also have been significant for its acquisition of some part of a kiwi, possible little more than a few feathers, donated by the missionary Thomas Kendall. This interesting bird was discovered barely a decade earlier and was so unusual that its very existence was doubted by European naturalists at the time.
The scientific accounts of Duperrey's voyage were published in grand style and most volumes had appeared before 1830. This was the start of an outstanding series of zoological works arising out of the French voyages to New Zealand. They were comprehensively and richly illustrated with fine hand coloured engravings which were incorporated as an ‘Atlas’ to accompany the voyage reports.
It spite of these considerable achievements, it was the next voyage that stood out as the most significant of the three conducted during this period. Under Dumont d'Urville's captaincy the Astrolabe reached New Zealand in January 1827. They collected in Astrolabe Bight and in the vicinity of French Pass before heading for Auckland and the Bay of Islands. They discovered some new bird species including the South Island fantail, the wrybill, the grey warbler and the quail (Wright, 1950, Andrews, 1987). But again it was the invertebrate collection that predominated, and became associated with the two naturalists, Quoy and Gaimard. They recorded many new molluscan shells and the magpie moth, a cicada and other insects that were later described by the French entomologist Boisduval.
This voyage too resulted in an elegant publication with a fine atlas of engravings. Quoy himself was very good at executing watercolours of the numerous shells and other marine life that they discovered. These paintings are still in the Muséum d'Historic Naturelle.
There was a mystery associated with this voyage that is still unresolved. In the museum at Marseille is a specimen of the world's largest gecko (Bauer, pers. comm.). It is believed to have come from New Zealand, and there are fairly reliable accounts of large reptiles of this description having been present in the late 19th century. How it got to Marseille no one knows and it is not mentioned in any of the voyage accounts, although the vessels from both expeditions called there on their return.
D'Urville's last voyage with Astrolabe, and the Zelée under Jacquinot, reached New Zealand in March 1840 calling at the Auckland Islands and there making the first significant collections from one of these interesting localities that lie well to the South of New Zealand. A new species of penguin was found here as well as a variety of insects and shells (Wright, 1955; Andrews, 1987).
They called at Otago, Akaroa, Poverty Bay, and the Bay of Islands. Their collections were well received back in France but perhaps were not as substantial as the previous voyages. Also, by now voyages returning from the South Pacific were relatively commonplace. However, most significant in the collection was a pair of kiwis that were illustrated in the atlas that accompanied descriptions of the voyage zoology. Ever since the kiwi was first described the French zoologists had been keen to lay their hands on a specimen, and for many years remained unconvinced of the authenticity of earlier accounts. Their doubts were now put at rest.
The third and final phase of the French contribution to New Zealand zoology was that represented by the French whale fishery and the protecting vessels of the French Navy, as well as the small French community at Akaroa. Amongst their leaders was St. Croix de Belligny who was a naturalist from the Jardin des Plantes in Paris. He was apponited a travelling correspondent to that institution and asked to send specimens back to France when time permitted.
De Belligny collected more than 130 birds that were sent back to France, probably on one of the naval vessels. The officers of these vessels themselves collected specimens, for example Lavaud and Raoul on L’Allier, Jaeger Schmidt on the Heroïne and Arnoux on LeRhin. Some material went to the Maison Verreaux, a firm that deal in natural history specimens, one of whose members, Jules Verreaux, was known to have visited New Zealand (Andrews, 1987).
This last phase was diffuse and less organised than the voyages that preceded it, and its output cannot be said to have surpassed the volume of scientific work of the voyage naturalists. The activity in the third phase took place into the 19th century when most of the large animals at least were well known and represented in the major museums.
Taken as a whole the French contribution to early New Zealand Zoology was a major one and somewhat tidier and more complete than the British efforts that preceded and followed it. Many of the natural history findings of the voyages of Cook failed to reach publication - or at least did so in a piecemeal way and then not always adhering to Linnaean nomenclature. The later voyage of the Erebus and Terror which included New Zealand was also a substantial affair, but it took many years for its findings to appear in print. Speed of publication is all important for the scientific record - specimens can soon rot or dry out in their bottles, and publication delays can often lead to nomenclatural problems.
The French contribution to New Zealand science began around the same time as the French Revolution. In spite of the political and social turmoil that followed, and the upheaval that resulted from wars with Britain, the French found time to throw substantial light on the natural history of a little known part of the world. Their contributions in many cases have stood the test of time and still remain a valuable part of the permanent scientific record.
A comprehensive approach to the cataloguing of natural diversity was developed by the natural philosophes in France and became part of French government policy following the revolution of 1789. This resulted in the establishment of the professionally-based Muséum National d'Histoire Naturelle in Paris which, through a system of voyageurs-naturalistes, was able to ensure that well-documented natural history collections were made during pacific voyages and preserved in state institutions. A similar commitment to museum-based natural history research and exhibition or even to scientific collecting was not developed by the British until after 1840, by which time many of the new wonders of the Pacific had been described. Thus most early British name-bearing zoological specimens from the Pacific were lost.
I am neither a historian nor an expert on voyages of discovery, but rather a museum curator concerned with tracking down and verifying early type specimens. With very old collections this can be a complex and difficult task as types were mostly not labelled as such. Sometimes the identification of type specimens revolves around the identification of particular labels, or handwriting. Thus it is often necessary to become familiar with the personnel and organization of scientific expeditions if one wishes to identify the actual specimens described. From such work the idea for this paper developed, constrasting approaches to the organization and documentation of zoological collections in England and France. It arose following discussions on French contributions to scientific knowledge of New Zealand at a symposium organized by the Alliance Francaise de Wellington and the Royal Society of New Zealand on 28 October 1989, and formed the basis of a public lecture at the National Library of New Zealand on 10 October 1991, one of a series on New Zealand as seen by the French.
My interest in the scientific organization of voyages of exploration and the fate of their collections was stimulated by a study of early specimens of bellbirds Anthornis melanura (Bartle and Sagar 1987). The selective synonymy that follows illustrates something of this history and in particular highlights the involvement of both British and French expeditions in their collection and subsequent species description:
“Mocking Creeper” Latham 1782
- missing type, ex-Leverian Museum
Certhia melanura Sparrman 1786
- “Cape of Good Hope.” Museum Carlsonianum
Philedon dumerlii Lesson & Garnot 1828
- 5 collected La Coquille Bay of Islands April 1824
- figured in Atlas (1828)
- full description in text (1830), recognised as the same as C. melanura Sparrman.
The first two names had their source in the British expeditions of James Cook, familiar to most New Zealand readers, whereas the last name arose out of the first of the major French expeditions to New Zealand begun by Duperrey and continued by D'Urville, which are less well known.
Many of us have been led to believe that French interest in this part of the world was largely concerned with territorial aspirations, including a rather limited
I was brought up to believe that the perfect models for conduct of voyages of exploration and discovery were those of Cook, and that the best way for a government to organize the scientific programme for such voyages was to delegate it to a gifted amateur like Sir Joseph Banks. Banks had a key role in all British voyages of discovery from 1766 until his death in 1820. He became a general repository for all collections and scientific knowledge which Cook and his followers accumulated (Mackay 1985). The influence of Banks can be traced right up to the Challenger expedition of the 1870s — perhaps the first completely professional British voyage of scientific discovery. And what happened to Banks' great repository of knowledge and collections? Where are the published journals and monographs, and what happened to the collections?
Perhaps I was influenced in my youth in favour of an inspirational rather than pragmatic approach to scientific expeditions and the production of research results by Sir Charles Fleming, himself a great admirer of Banks. Scepticism really only set in when, a few years ago, I started to try and track down bird specimens from British voyages … Only then I learnt of the delays, dissention and intrigues that beset publication of the results of Cook's voyages in England (Hoare 1982). In my opinion these private jealousies set back scientific knowledge of this part of the world by fully 50 years — from the 1770s to the 1820s, until after the end of the Napoleonic Wars.
English voyages of scientific exploration actually depended on private patronage for over 70 years — from the time of Cook to that of James Clark Ross, in 1840. Sir Joseph Banks, England's greatest promoter of science during the period (Mackay 1985), was a conservative figure, favouring slavery and believing that the benefits of science and mercantilism should be reserved for the landowning class (Mackay 1985). This went with a paternalistic attitude toward indigenous peoples. Other key amateurs involved in promoting scientific exploration in the eighteenth century included Lord Sandwich, Daines Barrington and Thomas Pennant.
Museums were the main government agencies involved with collections and publications resulting from expeditions of scientific discovery in the nineteenth century. Again, the famous English institutions are held up as models of their kind, but this was certainly not the case until the 19th century was almost over.
Montague House was purchased in 1754 by the British government to house the collections of Sir Hans Sloane. The natural history cabinets on the first floor were reached by a magnificent staircase guarded by a stuffed rhinoceros and three giraffes. Early keepers owed their position to patronage, and the stipend was insufficient to offer a livelihood. Montague House was maintained as a nobleman's cabinet of curiosities, rather than as a national musuem for the benefit of the masses (Gunther 1975). The trustees seemed to wish to make it hard for the right people, and absolutely impossible for the wrong people, to get into the place at all. The museum was open only three days a week (daily opening would not arrive for 120 years), and visitors had to apply in writing for admission to the Principal Librarian several weeks in advance (Panter-Downes 1980). Staff worried that the
It was not British government policy to encourage organized collecting during this period (Gunther 1975). So the advance of natural history depended on the study of collections in private hands. In eighteenth century England the most important of these was that of Sir Ashton Lever, who built a museum in Leicester Square, open to the public for a small fee. The Leverian Museum was much more popular with both the public and scholars than the British Museum, and the latter was described in 1799 by a visiting professor of natural history as containing “amidst a vast quantity of insignificant trifles, few important specimens”. It was “no longer instructive” whereas, in contrast, “the Leverian museum may be seen for a trifle, and the (well arranged) collection exceeds everything of the kind I have seen” (H.F. Link, in Stresemann 1975). This was partly because so many of the ethnological and zoological items from Cook's voyages were shown there. Pennant, Latham and Shaw found their richest quarry there, and by 1784 the Leverian Museum had 28,000 items and was world famous. But its owner had been too prodigal, and to make money, he held a lottery with 36,000 tickets at a guinea a head — all blanks except one, which entitled the holder to the entire museum! On the day of the draw Lever had sold only 8,000 tickets but, as luck would have it, the winning ticket was amongst these and was bought by a dentist, James Parkinson. The unhappy loser took to drink and died soon after, in 1788 (Stresemann 1975). James Parkinson, despite re-housing and expanding the museum, auctioned the entire collection, in May and June 1806.
The British government turned down the chance to acquire this, the greatest of all eighteenth century museums (Gunther 1975), and the lion's share was bought by the showman, William Bullock, who had also built a private museum in London. Later, in 1819 the government likewise refused to buy Bullock's collections, on the grounds of lack of space at Montague House (Farber 1982). By this time it included more than 3,000 birds, including all those from Cook's second and third voyages, presented by Sir Joseph Banks. So this priceless collection was sold at auction and dispersed throughout Europe with many specimens and associated data lost forever (Medway 1981).
Meanwhile, the state of the British Museum's collections was not good:
“… mouldering or blackening in the crypts of Montague House, the tomb or charnel-house of unknown treasures [where] …
moths, ptini, dermetesare busily employed amid the splendours of exotic plumage, or roaring through the fur of animals, we do not know a single insect visible to the public, of all that have been deposited in the British Museum” (Traill 1823).
Neglect of the British Museum's natural history collections by the poorly-paid staff became a well-recognised scandal by the 1830s. Konig (1836) wrote that:
“… Most objects of the Sloane collection were in an advanced state of decomposition and they were buried or committed to the flames one after another. Dr Shaw had a burning every year; he called them his cremations … Some persons in the neighbourhood complained and threatened with an action, because they thought the moths were introduced into their houses by the cremations in the Musuem garden.”
And, later, in the time of Dr Leach, when:
“… the gardens of the Museum were still the favourite resort of the Blooms-burians, but the attraction of the terraces and the fragrance of the shrubberies were sadly lessened when a pungent odour of burning snakes was their accompaniment. The stronger the complaints, however, the more apparent
became Dr. Leach's attachment to his favourite cremations” (Edwards 1870).
Collections made by government and private expeditions prior to 1840 were not offered to the British Museum, but instead to the museums established by societies like the Zoological Society of London and the Royal College of Surgeons, and by organizations such as the East India Company. Treasury, the Foreign Office and the Admiralty felt that the British Museum was unworthy of collections from overseas expeditions, and instead gave these to the Zoological Society and to private collectors (Gunther 1975). By the time John Gould retired as keeper of the Zoological Society's museum in 1836, this collection alone already outstripped that of the British Museum (Stresemann 1975).
By 1835 the situation had deteriorated to the extent that a Parliamentary Select Committee was set up to look into the condition and management of the British Museum. The need for a properly-paid professional staff was identified, as well as the advantage of placing all government-collected specimens in a national collection; and the value of employing “travelling naturalists” (Farbor 1982). During this review the Muséum National d'Histoire Naturelle, Paris, was repeatedly cited as an example of how a major national museum should be organized. Over the next decade these recommendations were slowly put into effect, although G. R. Gray, in charge of the bird collections at the British Museum since 1831, did not employ collectors abroad, like the French. It was not until Bowdler Sharpe took over the bird collection in 1872 that really substantial advances in ornithology were made (Stresemann 1975).
The origin and establishment of the Muséum National d'Histoire Naturelle, Paris played a key role in encouraging scientific exploration in the Pacific during the early nineteenth century.
The idea of cataloguing nature to reduce its diversity to a comprehensible system was developed by natural philosophes in France before the revolution. After the revolution such knowledge was felt to belong to the masses, and not just to an educated élite. This, from 1793 to 1840 French voyages of exploration resulted in government-funded publication of detailed accounts of voyages and discoveries and the journals and objects recovered became the property of the people through the new national institutions.
One of the four major philosophes of the French Enlightenment was Georges-Louis Leclerc, Comte du Buffon, superintendent of the Jardin du Roi since 1739. He was responsible for the aggrandizement of that institution from a minor botanical garden for the study of pharmaceutical plants to a major research institution. His Histoire naturelle, générale et particuliere was the third most popular book in France in the late 18th C (Farber 1982). Buffon's writings were one of the important factors in popularizing science in the period of the Enlightenment. However, Buffon really needed thousands of bird specimens for this undertaking. Although 700–800 had been accumulated by 1770, many of these were destroyed by insect pests, and no one knew how to protect them. The big breakthrough was made by Jean-Baptiste Bécoeur, an apothecary in Metz, who developed an arsenical soap that protected skins without destroying them. Although Bécoeur kept his recipe a secret during his lifetime, in the hope of benefiting financially from it, it somehow passed to the Muséum and became the accepted method of preparation (Farber 1982). Louis Dufresne, taxidermist at the Musěum since 1793, popularized arsenical soap in an article on taxidermy that he wrote for the Nouveau dictionnaire d'histoire naturelle (1803–1804). This single development enabled the Muséum in Paris to build up the greatest collection of birds that the world had ever seen in the early 19th C.
In the same year as the appointment of Dufresne, the revólutionary government of France took the bold decision not to dismantle the Jardin du Roi, but rather reorganize it as a national museum of natural history that would encourge the solution of practical problems, would provide lectures to the public, and would reflect the glory of France as the leader in the world of ideas (Farber 1982). Under the energetic direction of Etienne Geoffroy Saint-Hilaire, the Paris Muséum rapidly became the centre for European zoology. The Revolutionary armies had brought in many remarkable and important rarities when they carried off the Prince of Orange's famous collection as booty from The Hague to Paris in 1775. Its condition in 1797 was described by H.F. Link:
“The museum of natural history in the botanic garden of Paris is far more interesting than the British Museum, and contains a great number of specimens, and very extraordinary productions. London possesses nothing that can be compared with it.”
In 1817 William Kirby wrote:
“Every part of the Museum is in beautiful order, systematically arranged, so that every student may in a moment find every object that he wants, and every facility is afforded to him that he can desire. I wish the zoological department of the British Musuem was in similar order.”
During the following years the French government took a series of far-sighted decisions which effectively integrated the work of the Paris Muséum with expeditions and voyages of scientific discovery. On February 3, 1819, at a meeting of the staff of the Muséum, the Secrétaire général of the Ministre de l'Intérieur informed the professors that the budget contained twenty thousand francs to create “a school for young naturalists destined to make voyages to various parts of the world”. The sum soon grew to twenty-five thousand francs per year and was used to train, equip, and cover the expenses of collecting for about ten voyageurs-naturalistes. With the aid of government support, the Muséum was able to send people into the field in localities where it wished to strengthen its collection. Missions were sent to the Cape of Good Hope, South America, Australia, West Africa, Madagascar, North America and India (Farber 1982). Of course not only specimens were returned to the museum, but accurate accounts were kept and preserved in the Muséum, where they are today. Foreigners and colonials were encouraged to send material by designating them correspondants (Farber 1982). The Muséum enjoyed enormous official support, for it drew on the rising public taste for natural history and promoted that interest by lecture series and by its popular exhibitions. This at the time when the few enterprising souls who actually obtained entry to the British Museum commented most unfavourably on the state of the collections and on the lack of enterprise of its staff.
D'Entrecasteaux's expedition in search of the ill-fated La Pérouse was the first post-Revolutionary expedition to the Pacific. Unlike earlier French voyages, it was well-equipped to make observations and collections, although unfortunately everything was seized in Java, and it was years before the naturalists could return to France, bereft of notes and material.
Nicolas Baudin's voyage to Australia in the Géographie and the Naturaliste (1800–1804) was well-staffed with scientists, artists and instruments, and one of the largest natural history collections to date, including nearly 1,000 birds, was brought safely back to France. Baudin's expedition, although highly successful from a scientific point of view, was torn by internal strife. The bitter disputes between the scientific staff and the naval personnel led to the decision that henceforth the collection and observation of natural history on navy ships would be entrusted to medical officers.
This change in policy reflected a growing professionalization of the French navy as well as a shift in emphasis in the purpose of major naval expeditions (Farber
Muséum. To the museum came hundreds of rare species from the government expeditions of the Coquille (1821–1825), the Astrolabe (1826–1829), and the Astrolabe and Zélée together (1837–1841), which amassed a wealth of new birds from the coasts and islands of the world's oceans. Added to these were the astonishing novelties collected by the daring “travelling naturalist” Goudot and the Ship's doctor Bernier during the 1830s on the mysterious island of Madagascar. Beginning with Alcide d'Orbigny (1826–1834), Justin Goudot (1827–1843), and Claude Gay (1830–1842), explorers of the Andean countries were mostly French (Stresemann 1975).
Publication of the scientific results of these expeditions was prompt and comprehensive, thanks to the availability of government funds. Familiar New Zealand animals such as the sea-horse and red gurnard were first described in Duperrey's volumes. On D'Urville's later voyages the collecting became quite systematic, with particular attention paid to specific groups. D'Urville also had firm instructions regarding disposal of the collections to the Muséum from the Secretary of the Navy (Andrews 1986), in contrast to the contemporary British voyages in which amateurs participated at their own expense. The result was that the Muséum in Paris was often overwhelmed with specimens and drawings. In 1828–30 1263 animals were illustrated, 500 species of insects collected, and 520 birds accessioned, together with many botanical and geological specimens (Andrews 1986). Ten volumes of publications were approved containing the finest hand-coloured copper plate engravings.
Thus the Muséum went from strength to strength, enjoying a status and financial support unequalled by any comparable institution in Europe. In 1840 Swainson remarked that the Museum was “… the most celebrated in the world …” The arrangement and accessibility of its collections, combined with their scope, made it the centre for ornithology in the first half of the nineteenth century, and the standard by which other museums were judged. Because it was a permanent, government-funded institution, workers could take advantage of having type specimens in a known, fixed location. In days when travel was difficult, expensive and sometimes dangerous the importance of having a large and accessible permanent collection in a central location was essential for anyone writing a general treatise or monograph.
Throughout this intensive period of cataloguing nature, discoverers of new species increasingly designated particular specimens as ‘types’. Original descriptions were often inadequate, and later authors needed to check exactly which species was being discussed. Fortunately type specimens were generally labelled as such at the Paris Muséum, and can be easily located even today. In England, however, the frequent breakup of important collections and the formation of new private collections throughout the 19th C has greatly impeded zoology and, in particular, ornithology.
Worse still, as the British Museum was part of the Civil Service, all correspondence, including that relating to the acquisition of specimens, was systematically destroyed up till 1965. This means that the only documentation for specimens is usually the brief register entry, or the label, which could be removed or amended. In addition, when the Trustees of the Natural History Museum ordered the demolition of Walter Rothschild's “insect room” laboratory at Tring, in 1970, all documents relating to the Rothschild collections, including the journals of collectors like Henry Palmer in Hawaii and the Chatham Is, were systematically burnt (Rothschild 1983, pp 185, 299, 331). The full implications of this great crime are yet to be recognized.
The funding of a public natural history museum in 1793 by the revolutionary government of France was a turning-point in the history of science. It provided a proper foundation for the collection and description of natural history specimens by French expeditions and naturalists with the result that, for many years, their collections, displays and publications were superior to, and the envy of, their British counterparts.
Lessons on the effects of the different levels of state involvement in zoological research in the Pacific and on the contrasting organization of British and French natural history museums in these far-off times still have relevance today. State responsibility for funding of biological collections is essential for progress in scientific knowledge.
Ideas for this paper were accumulated during periods of work at the Museum of Zoology, University of Cambridge; Sub-department of Ornithology, British Museum (Natural History), Tring; and at the Muséum National d'Histoire Naturelle, Paris, in 1980 and subsequently. I am grateful to the following curators and staff for their hospitality and many helpful discussions: the late C.W. Benson; I.C.J. Galbraith and P.R. Colston; C. Erard and C. Jouanin. The work of J. Dunmore has been an inspiration. The following kindly commented on this paper: J.R.H. Andrews, J.W. Dawson, J. Dunmore, and J.C. Stahl.