Publicly accessible
URL: http://nzetc.victoria.ac.nz/collections.html
copyright 2006, by Victoria University of Wellington
All unambiguous end-of-line hyphens have been removed and the trailing part of a word has been joined to the preceding line, except in the case of those words that break over a page. Every effort has been made to preserve the Māori macron using unicode.
Some keywords in the header are a local Electronic Text Center scheme to aid in establishing analytical groupings.
is the journal of the Departments of Botany and Zoology, Victoria University of Wellington, and Victoria University Press, New Zealand, and is published twice a year.
Joint Editors:
This article is modified from a chapter in a forthcoming book — “Forest Vines to Snow Tussocks. The Story of New Zealand Plants.”
Keywords: Climbing plant, epiphyte, forest, liane, New Zealand, parasite, saprophyte.
The specialised forest growth forms are defined.
The vines are divided into subcanopy herbaceous species (ferns) climbing by roots and woody canopy lianes climbing by roots, twining stems, twining leaf petioles, tendrils, hooks or scrambling stems.
For epiphytes the primary categories are shade epiphytes growing low on trunks and sun epiphytes in tree crowns. The latter are subdivided according to their forms or locations — mat, nest, pendent, small shrub, large shrub, tree or “strangler”, on tree ferns, on leaves (epiphyllae).
Parasites attached to roots or branches are reviewed. Our few saprophytes are described and their life style compared with that of parasites.
Woody and herbaceous vines and vascular epiphytes or “perching plants” are abundant and distinctive features of tropical rain forests. Surprisingly, in view of our temperate latitudes, a comparable range of vines and epiphytes is equally conspicuous in the New Zealand conifer broadleaf forest By contrast the larger vines and epiphytes are uncommon in or absent from beech (Nothofagus) forests. The mistletoe parasites, however, are more conspicuous in beech than in conifer broadleaf forest.
This partial neglect is perhaps understandable, as it is not easy to get close to the foliage, flowers and fruits of many of these plants, particularly where they occur high in the branches of tall trees. Binoculars help, but few botanists are as intrepid as J. L. Harrison-Smith, who spent many hours “wandering about” among the branches of giant kauris recording epiphytes. In his own words …“In order to catalogue the plants the trees were climbed. Ordinary gum collectors' climbing gear (spiked boots and climbing hooks) were used. Descent was accomplished with a “bo'sun's chair” suspended from a rope. It was then possible to examine the trunk and any isolated branches on the way down.” Fortunately it is possible to reach most vines and epiphytes without taking such risks. Some species grow low on the trunks of trees and so are readily accessible, but even those normally restricted to tree crowns may descend to lower levels in well-lit situations. Also, in the normal course of events, trees are blown or fall over and then a full range of high epiphytes and vines, albeit somewhat damaged, can be examined.
Not infrequently the categories of vines, epiphytes and parasites are confused, so that if it is stated that a plant “grows epiphytically on trees” further enquiry is often necessary to determine the precise mode of growth of the species in question. It is true that the word “epiphyte”, meaning “plant that grows on other plants,” fits all three categories, but as in other respects their life styles are quite distinct, this term should be restricted to plants that germinate and establish on the trunk and branch surfaces of trees. Vines by contrast establish on the ground, then grow up the trees, and, although some parasites are similar to epiphytes, in that their seeds germinate on trunks and branches, they differ markedly in having special root-like organs (haustoria) that penetrate the living tissues of their host and draw water and nutrients from them.
Vines in this category are herbaceous and attach themselves to tree trunks by special roots arising from the stems. They ascend for varying distances up the trunks, but mostly do not enter the tree crowns. They are all able to reproduce in the reduced light of the forest interior.
In New Zealand all vines in this category are ferns and in some, as in the climbing members of the arum lily family in the tropics, there is a remarkable increase in size and complexity of the leaves with increasing height above the ground. The best known example of this is Blechnum filiforme a common plant in lowland forest as far south as the northern South Island. (Fig. 1).
Arthropteris tenella also has relatively thin, dark leaves and slender vertically ascending stems, but these generally reach to only a few metres above the ground. This species also reaches the northern South Island, but is less common than
The light green, thin-leaved Phymatosorus scandens also shows a trend from juvenile to adult leaves with increasing height, but in this case the juveniles are about as long as the adults, up to 35 cm, but are narrow, undivided and usually sterile. There is often a fairly abrupt change to the adult leaves, which are much wider and are deeply incised into a number of narrow, lateral segments bearing sporangia. This species often occurs with
The thick-leaved Phymatosorus diversifolius, with its stout, grey-green, black-flecked stems, is better known to most people. It ascends to higher altitudes than the species so far considered and also reaches the Auckland Islands to the south of New Zealand. It also differs in often preferring to climb inclined trunks and inclined to
Erratum
Legend for Fig. 1 should read:
Upper right: Leaf from near ground on a tree trunk. Left: Leaf 3 metres above the ground.
Below right: Fertile leaf.
As the name of the species indicates it has a similar range of leaf forms to P. scandens. On a tree with a heavy growth of the fern the large, shiny, bright-green leaves are rather widely spaced and are deeply incised into narrow segments with an abundance of sporangia beneath aggregated into distinctive orange spots or sori. Young plants establishing in moss on trunk bases have narrow, undivided sterile leaves. Phymatosorus diversifolius also grows on the ground, most abundantly on rocky slopes. In rocky exposed places the narrow, undivided leaves may persist, but in these circumstances they bear sporangia.
A third species of Phymatosorus—P. novae zelandiae—is found in montane forests throughout the North Island, but is absent from the South Island. With its thick, leaves and stout stem it is similar to P. diversifolius, but the stems are densely covered with straw-coloured scales and the leaves are generally larger with more numerous, narrower and longer lateral segments. It appears that there are no marked variations in leaf form in the species.
Rumohra adiantiformis ranges throughout New Zealand in lowland to montane forests and is most common as a climber on tree fern trunks. The much divided leaves have a leathery texture and bear conspicuous sori. There is a modest increase in size of leaves with increasing height.
Particularly in high rainfall areas filmy ferns may be common as climbers. In some cases the fronds are very small and delicate and grow intermingled with mosses, but some species — Hymenophyllum dilatatum, H. scabrum, H. sanguinolentum — have relatively large leaves which, Holloway notes, increase in size with increasing height above the ground. Holloway suggests that the increased leaf areas in this case enable more effective absorption of water by the thin leaves in the drier tree trunk habitat. The kidney fern (
Six of the climbing ferns considered here are restricted to New Zealand. The ranges of the others are:
Phymatosorus diversifolius (Australia, Tasmania, Tropical Polynesia)
P. scandens (Australia, Norfolk Island)
Arthropteris tenella (Australia, Norfolk Island, New Caledonia)
Rumohra adiantiformis (South temperate zone, tropical Polynesia, tropical America)
Vines whose foliage eventually spreads into the canopy of the forest are usually woody and are often referred to as lianes. At the adult stage they are light-demanding and generally produce flowers, or spores, only in well-lit situations. Young plants on the forest floor are more shade tolerant, but nevertheless establish most abundantly in the better-lit earlier stages in forest development or in canopy gaps in mature forest. Unlike the sub-canopy climbers the lianes climb by a variety of means — attaching roots, twining stems, hooks and tendrils.
Most prominent here are the climbing ratas, currently included in the genus Metrosideros. They are able to grow up quite large trunks and are perhaps most abundant on conifers and northern rata. The climbing stems are usually quite slender and the leaves, forming a close mosaic on the tree trunk, are generally smaller, thinner and more rounded than those of the adult stage. When the stems reach full light high in the tree crown, or at lower levels where light is adequate, they form a bushy growth of branches which extend away from the support and eventually bear flowers. At this stage the stems extending up from the ground enlarge considerably and swing away from the host trunk as woody cables (Fig. 2). Metrosideros fulgens and
Metrosideros albiflora and
It has been said that New Zealand is the only place where climbing species of Metrosideros occur, but in fact there are climbers related to ours in New Guinea and the Philippines.
The only other root climbing liane in New Zealand is the kiekie (Freycinetia baueriana var. banksii). It belongs to a distinctly tropical family, the Pandanaceae, which is represented by many species of Freycinetia and Pandanus in tropical rain forests. One might expect that the out-lying New Zealand species would be of reduced form and perhaps rare, but in fact it compares with the largest and most robust tropical species and is abundant in lowland, especially swampy forests as far south as the south west of the South Island. Tree trunks are often completely obscured by the foliage of kiekie, which can extend into the highest crowns 30 or more metres above the ground. The leaves are dark green, narrow and a metre or more long with finely toothed cutting edges. The male and female inflorescences, on separate plants, are cone-like and surrounded by leaf-like white or purplish bracts, which are sweet and edible. The stems are a few centimetres in diameter and distinctively ringed with leaf scars. They give rise to slender, attaching roots, which branch freely towards their ends and attach themselves firmly to the trunk. Other roots are stouter and grow down the trunk to the ground, often building up into quite thick and rather untidy masses.
Twining lianes have climbing stems which wind around supports in a clockwise or anticlockwise direction, depending on the species, until they reach the full light of the forest canopy. Unlike root climbers many twiners are not able to climb large tree trunks —their turning circle is too small for that — so they either have to climb young slender
Undoubtedly, supplejack (Ripogonum scandens), which ranges throughout the country, is the most familiar twiner in New Zealand forests particularly on alluvial and swampy sites. It belongs to the lily family, taken in a wide sense, and its almost black, jointed, bamboo-like climbing stems often form entanglements that greatly impede progress (Fig. 3). Fortunately, unlike several of its relatives in Australia, supplejack does not have prickles.
Aggregations of woody, tuber-like rhizomes below ground give rise to the climbing stems which are dark-brown to black, 1-2 cm in diameter and bear pairs of long, narrow, often twisted scales in the place of leaves. The stem tips are reminiscent of Asparagus and are soft and easily broken. They can elongate at an average rate of 5 cm per day in summer and while growing upwards the upper part of the shoot revolves slowly in an anticlockwise direction. If it does not encounter a support it bends down to the ground and grows up again from the tip.
Supplejack mostly climbs fairly slender supports, but can also twine around quite large trunks, the record being a 1.5 m diameter kohe-kohe. When a climbing stem reaches the forest canopy, lateral climbing stems arise from its upper parts and eventually these bear relatively slender, leafy stems of limited growth, which are unable to twine. The leaves are broad and distinctively veined with two strong lateral veins more or less parallel to the midrib. The leafy stems bear small flowers followed by bright red berries.
When lateral climbing stems are formed near the ground they are often swollen and tuber-like at the base and produce roots which may descend more than a metre to the ground. Supplejack is restricted to New Zealand, but other species of Ripogonum are found in eastern Australia and New Guinea.
The two species of Parsonsia sometimes known as native Jasmine, are found in lowland forest and shrubland throughout the country. Parsonsia heterophylla is the larger of the two with stems up to 10 cm in diameter which attain heights up to 20 m above the ground. It is commonest near forest margins, but reaches the crowns of taller trees deeper in the forest by spreading from lower to higher levels in the forest canopy.
In this species there is a very marked difference in size and shape between juvenile and adult leaves. The seedlings establish in sometimes quite shady places on the forest floor and the first few leaves produced are small and almost circular. These are followed by leaves tending towards the second type of juvenile leaf, which is long and narrow with smooth or wavy margins. Intermediates may be narrow at the base and round at the tip, or narrow at base and tip and rotund in the middle, and, to make matters even more complicated, lateral branches on the seedlings usually repeat the same sequence. The result is a bewildering and apparently random arrangement of leaf forms. Some of the seedlings are completely green, others are largely brown and some leaves of the latter are unusually attractive with a mosaic of green, dark brown and pale brown patches. This is an interesting phenomenon also to be found in the equally variable leaves of juvenile pokaka (Elaeocarpus hookerianus), juvenile Pittosporum obcordatumand and seedling lancewood (
Seedlings, rotating in an anticlockwise direction at their tips as they grow upwards, reach about 45 cm in height without support and somewhat higher if two or more seedlings twine about each other. If no support is encountered then the stems bend down to the ground and grow along it until they find something to climb. The supporting stems are usually slender, although Parsonsia heterophylla has been observed climbing tree trunks up to 25 cm in diameter.
Parsonsia capsularis has small flowers of a different form to those of
The small, fragrant flowers of these two species are borne in clusters and are white, yellow or red (P. capsularis only). The fruits are pod-like, hang downwards and split open to release numerous seeds, each with a dense tuft of hairs for wind dispersal.
It has been suggested that the distinctive juvenile forms of our Parsonsias are peculiar to New Zealand, but in fact the phenomenon is also found in species in eastern Australia and New Caledonia. The genus ranges from tropical Asia to the Pacific.
Two of the New Zealand species of Muehlenbeckia are common twining lianes throughout New Zealand in lowland to montane forests. Both are deciduous, M. australis is the larger species and its seedlings are often abundant on the forest floor in both shady and well-lit places, but mature plants are most commonly found at forest margins or in regenerating forest. Young stems bend to the ground if they don't find support, branch, and spread on the forest floor. Unlike most other twiners the erect stems rotate in either direction and in some cases change direction when they begin to climb. The supports are always slender and often become very deformed as they expand within the coils of the vine. Sometimes they die, and this may be caused by the vine, but at other times it is the latter that dies leaving as evidence on the supporting stem a pronounced helical groove.
By climbing up smaller trees Muehlenbeckia australis may extend into the crowns of tall trees 30 metres or more above the ground. A distinctive feature of this vine is the formation of firm cane-like “searcher shoots” during the autumn from any part of the stem sysfem. Where these arise on stems coiled on the forest floor they grow erect for several metres beginning to rotate only after the first metre; where they develop on stems in tree crowns they extend more or less horizontally often from one tree crown to another and in this way the vines become extremely widespread through the forest canopy. Particularly in second growth forest there sometimes seems to be more of the draping foliage of the
The adult leaves are several centimetres long, broad, thin and pale green, sometimes with a drawn-out tip. Juvenile leaves are much smaller, round, or oval and sometimes fiddle-shaped.
Muehlenbeckia complexa is similar in its growth habit to
Muehlenbeckia complexa is endemic and
Tecomanthe speciosa is undoubtedly New Zealand's rarest vine in nature, only one plant having been discovered on the Three Kings Islands. It has robust twining stems which, in cultivation, extend high into supporting trees. The leaves are pinnately compound with quite large leathery leaflets and the tubular flowers although large, are of an inconspicuous cream colour. Most of the other species of Tecomanthe are found in New Guinea.
Lygodium articulatum (Mangemange) belongs to a largely tropical genus of ferns unique for their ability to climb by twining, although in this case it is not the stems that twine, but the axes of compound leaves that have indefinite growth and sometimes extend from the ground to the tops of high trees. The New Zealand species is restricted to the north of the North Island, but is common there particularly in kauri forests. The true stems spread over the forest floor. The axes of the leaves arising from these are slender and wiry and often twine about each other as well as their supports to form springy masses, near the ground in well-lit places or on the forest roof. Compound
It might be wondered why the fern Lygodium with its twining leaves is not included here. Leaf climbers are defined as plants whose stems are supported as they grow upwards by the sensitive petioles (stalks) of their otherwise unmodified leaves, which wind round any slender supports with which they make contact. In Lygodium the true stems remain on the ground, but the primary axis of the leaf is like a twining stem in its indefinite growth and indefinite ability to twine. Indeed, it takes a botanist to appreciate that with Lygodium we are dealing with twining leaves rather than twining stems, so it seems more realistic to treat it as a twiner rather than a leaf climber.
New Zealand representatives in this category are all species of Clematis, a genus widespread in temperate regions and also found in the montane tropics. The best known and largest New Zealand species is Clematis paniculata, which is found in lowland forests throughout, particularly marginally, and is greatly appreciated in the spring when its sprays of large, pure-white flowers stand out against the dark foliage of the forest. The adult leaves are divided into three leaflets, which are broad, dark-green, and smooth-margined. Leaves on young plants on the forest floor are very different. The first leaves are long, narrow, membranous in texture and undivided. These are succeeded by compound leaves with three narrow leaflets which in subsequent leaves become broader and deeply lobed and gradually trend to the adult form where lighting is adequate. Similar juvenile leaves have been reported for Clematis species in eastern Australia. Seedlings rotate in anticlockwise direction and are able to twine around slender supports. Similar twining ability, at least when young, has been recorded for some other leaf climbers outside New Zealand, nevertheless the primary mechanism for climbing is the clasping leaf petiole.
When a leaf is formed at a stem apex it is first erect then gradually bends downwards until it projects at right angles to the stem. The petioles of the leaf as a whole and of the leaflets are well developed at this stage and if any of them touch a suitable support they are stimulated, by a process not yet understood, to wind round it. The portion of a petiole in contact with the support enlarges and becomes strengthened. Clearly the leaves cannot attach to large supports, so where a large stem, up to 10 cm in diameter, of Clematis paniculata ascends to a tree crown 10 metres or more above the ground, it must have attained that position via smaller trees and shrubs.
The four or five other climbing New Zealand species of Clematis are smaller than C. paniculata, are incompletely known and in some cases not yet clearly defined. Several grow at forest margins, including C. forsteri and C. foetida, and some also in shrubland.
Tendrils are similar to the petioles of leaf climbers in that they are sensitive to touch and respond by twining round a support. They differ in that they are derived from plant organs — branches, inflorescences, leaves or leaflets — that have completely lost their original function and are used solely for climbing. Further, once a tendril has attached to a support, it coils into two opposed helices in its free part, which increases its elasticity and also draws the stem closer to the support.
In New Zealand we have only one forest liane that climbs by tendrils. This is the native passion vine — Passiflora tetrandra, which ranges through the North Island and to Banks Peninsula on the east of the South Island. The leaves are dark green and shiny and drawn out to a point at the tip. The flowers are much smaller and less colourful than those of cultivated species and less elaborate in their form, but the fruit compensates for this, being bright orange, 2-3 cm in diameter and it is greatly sought after by birds.
The tendrils arise in leaf axils and are considered to be modified inflorescences. They are at first erect then bend downwards and if they encounter a slender support they wind round it. The part in contact gradually becomes thickened, until it is about twice the diameter of the free part of the tendril. The native passion vine is most common in the lower marginal parts of forests, but it spreads so effectively over the forest roof that it frequently reaches the tops of taller trees. The woody stems can be up to 12 cm in diameter and in their lower parts often form tortuous coils on the forest floor.
The New Zealand hook climbers are all species of Rubus, a widespread genus in temperate regions and the montane tropics, which includes the familiar blackberry and raspberry. In north temperate regions, the species of Rubus do not climb and are shrubs or scramblers in open habitats, but most of the New Zealand species and a number of Australian and tropical species are low to high climbing forest lianes. The adult leaves of most species are palmately compound with three or more coriaceous leaflets and the undersides of the petioles and leaflet midribs, as well as the stems at times, are beset with backwardly curving hooks or prickles, which effectively prevent the stems from slipping back from any position attained. This tenacity earned for the New Zealand plants the name of “bush lawyer” in colonial times, a perhaps unwarranted slur on the legal profession of the day. In fact the “lawyers” are the only plants in New Zealand forests that are prickly, which contrasts with the rain forests of Queensland and south east Asia where many spiny climbers are unpleasantly in evidence. Probably the original lack of browsing mammals in New Zealand is the explanation for this and the same would apply for the lack of spiny plants in New Caledonian forests.
The bush lawyers are sometimes included in the “scrambler” category of vines. Scramblers are small, unspecialised climbers whose weak, drawn out stems grow up between the branches of shrubs and trail over them. The lawyers begin their ascents in a similar way, but the presence of hooks enables them to reach great heights equal to those attained by more specialised vines. For this reason I think they warrant a special category. The liane species of New Zealand Rubus occur throughout the country, including Stewart Island, in lowland to montane forest.
Young plants, the first leaves of which are simple, may have quite stout stems and so are able to stand erect without support for 60 cm or more. If nothing is available to climb, the young plant bends to the ground and branches and spreads widely over the forest floor until some of the branches find supports and make their way into the forest canopy via shrubs and smaller trees. On woody stems looped on the forest floor and those extending to the forest roof “searcher shoots” are frequently produced, those arising near the ground being able to stand without support for one to several metres. Thus the searcher shoots are very effective in expanding sites of Rubus foliage in the canopy and in establishing new sites.
The commonest species is Rubus cissoides which has long and narrow, sharply toothed leaflets. The adult stems may be up to 17 cm in diameter and the foliage can reach to 15 or more metres above the ground.
Rubus schmidelioides has stems up to 10 cm in diameter. Its leaflets are similar in shape to those of R. cissoides but generally smaller with blunt teeth and a dense covering of whitish hairs beneath.
Rubus australis is most common in swamp forest and is sometimes referred to as “swamp lawyer”. The leaflets are short and fairly broad and sometimes almost circular. In this species there is a distinct juvenile form, which spreads and roots widely over the forest floor and bears leaves with small, membranous, more or less round leaflets with reddish coloured veins. At the adult stage this species can reach for 10 metres or more into tree crowns with stems several centimetres in diameter.
Rubus squarrosus is perhaps the most remarkable, as in open situations and on shrubs the leaflet blades are not developed and the leaves consist of rather elongated
R. australis.
Some low climbers of more open habitats — Parsonsia capsularis, Muehlenbeckia complexa, Clematis (several species) — have already been considered in association with their higher climbing relatives. Other small but specialised non-endemic vines of northern coasts are:
The remaining species are mostly unspecialised “scramblers” as defined in the last section in the discussion of hook climbers. They grow through shrubs and small trees at forest margins or in shrub communities in drier eastern localities. The species in this group are: Fuchsia perscandens; Brachyglottis (Senecio) sciadophilus and Helichrysum dimorphum of the Compositae or daisy family;
One of our species of Lycopodium, the attractive trailing L. volubile, can also be a scrambling climber over shrubs. This species ranges to the tropical Pacific and south east Asia and in the latter region is recorded as sometimes extending into tree crowns.
A number of the vines in the absence of support can grow as shrubs in open situations. This is particularly the case with the small, unspecialised scrambling species, as well as the small twiners Muehlenbeckia complexa and
Some of the taller, more specialised forest vines may also grow as shrubs. Rubus squarrosus is particularly striking in this role as it is entirely leafless and presents a forbidding mass of slender stems and petioles beset with yellow prickles.
Several of the climbing ratas (Metrosideros) too, can be encountered as shrubs, M. perforata in particular having a dense, billowy form, which gives no hint of any ability to climb.
Such dual roles are not peculiar to New Zealand vines, but have also been observed in a number of tropical lianes.
Sharing a need for much brighter light than is available on the floor of a closed forest the majority of species of epiphytes, vines and parasites grow high in tree crowns. In this situation epiphytes alone face special problems. Vines are able to obtain soil water and mineral nutrients via their stems; parasites can tap the supplies drawn up by their host trees; but epiphytes either have no connection with the ground throughout their lives or send roots down to it only after a period of some years. Soil does not form easily on trunks and branches and as they are sunnier, windier and better drained than the ground, they suffer both more frequent and more severe droughts. In response to these stressful conditions many epiphytes have evolved modifications enabling them to store water and to reduce its loss by evaporation. Water can be stored internally in special cells, whose presence confers fleshiness on the organs concerned, or externally in cavities formed by appropriately shaped and arranged leaves. Some epiphytes can get by with a minimum of mineral nutrients and need little or no soil, others build up considerable quantities of dark humus largely from the decay of their own old leaves and roots with a varying contribution of bark flakes and leaves from surrounding trees. Many other epiphytes unable to form soil themselves take advantage of those that can.
Now that we have defined the epiphyte category how rigorously do we interpret the definition in deciding whether or not a particular species should be included? Certainly not so rigorously that we exclude those species which, although normally epiphytic, are sometimes to be found on sunny, rock outcrops that provide conditions similar to those of tree tops. In fact, it might well be that there are no epiphytes, even those of tropical forests, which are unable to grow on the ground in suitable circumstances.
Going to the other extreme, should we include species that are normally terrestrial, but can occasionally grow on trees? In this case the answer is “no” as apart from reservations about stretching the definition so far, the number of species involved would be inconveniently large. In certain circumstances almost any plant is able to grow as an epiphyte. For example, in forests of high rainfall regions, particularly where frequent mists maintain high atmospheric humidity, seeds germinate just as readily on moist, moss and lichen covered trunks and branches as on the ground. A notable example of chance epiphytism in these circumstances in New Zealand, is an occasional silver beech (Nothofagus menziesii) growing on a tree of the same species. Even in forests of average rainfall and atmospheric humidity the branch systems of large long lived trees, such as the kauri, are available as habitats for so many centuries that quite unlikely species can sometimes be found as epiphytes on them. The intrepid Harrison-Smith found a 3 metre kauri growing on a kauri, as well as a few to several examples of other conifers—rimu, totara, kahikatea—and also several angiosperm trees. In most cases the occasional epiphytic plants of otherwise terrestrial trees and shrubs are small and do not grow on to reproductive maturity.
Lichens, often followed by mosses, are generally the first epiphytes on trees in both temperate and tropical regions. In New Zealand small filmy ferns are often associated with the mosses. The thin layer of soil that these small epiphytes form is important for the establishment of most of the vascular epiphytes, which are the concern of this section.
These do not require a very high level of light and as epiphytes they mostly grow low on tree trunks where they escape the shading of the larger plants of the forest floor.
Five small species of fern and two flowering plants occupy this station in New Zealand although they are also to be found on rocks. Of the ferns three are species of Grammitis characterised by tufts of narrow simple leaves arising from short stems. G. pseudociliata differs from G. billardieri and G. magellanica ssp. nothofageti in having an abundance of reddish hairs on its leaves. G. pseudociliata is concentrated in the North Island, G. billardieri and G. magellanica extend throughout the country and the last also occurs in southeast Australia.
Ctenopteris heterophylla occurs throughout the country as well as in the subantarctic islands and south east Australia. Its habit is similar to that of the
Anarthropteris lanceolata is found in the North Island and near the northern shores of the South Island, as an epiphyte or on rocks. It is also said to occur in Vanuatu. The leaves are very similar in shape to those of
These four epiphytic ferns are related to the climbing Phymatosorus species and have similar prominent, rounded, brown to orange sori.
The two flowering low epiphytes are both species of Peperomia, a large genus of small succulent plants in tropical and subtropical regions. As epiphytes they grow mostly near the coast in the northern half of the North Island.
P. tetraphylla ranges from the East Cape district through to the Bay of Plenty. It has leaves in whorls of four as its name indicates. The same species is also found in Australia and Polynesia. P. urvilleana, with leaves borne singly, is found throughout
These are more numerous and diverse than the shade epiphytes and can be grouped into several growth forms.
These form mats or patches mostly on inclined to horizontal branches, and comprise three orchids and one fern which range throughout the country. These mat epiphytes may establish directly on bare bark, particularly if it is rough and fissured, but may also avail themselves of moss cushions. The one fern is Pyrrosia serpens belonging to a genus of epiphytes centred in tropical Asia. Our species is also found in Australia and the islands of Polynesia.
The epiphytic orchids belong or are closely related to large tropical genera and can be regarded as outliers reduced in both leaf and flower size. They all have specialised roots, which as well as serving for attachment, also efficiently absorb and store water in a special outer layer of dead cells known as the velamen.
Drymoanthus adversus, often attached to quite smooth bark, is unlike the other species in that it has a short stem, which does not grow along the bark surface. The roots arising at the base of the tuft of leaves are particularly conspicuous as they spread out “like the rays of a spider's web” for a considerable distance, often encountering the roots of other plants of the same species.
Much more evident to the casual observer are the massive nest epiphytes perched high in tree crowns (Fig. 4). In New Zealand there are three long and narrow-leaved species belonging to two closely related genera of the lily family — Collospermum hastatum, C. microspermum and
All three nest epiphytes usually establish among mosses and lichens in branch forks or on inclined to horizontal branches. As their stems, completely hidden by the leaf clusters, are short and more or less erect the plants are fixed in position, although branching to form additional leaf clusters often results in massive clumps of foliage
Astelia solandri is more shade tolerant than the Collospermums and so is often found below them in the lower branches and on the upper trunks of trees. The silvery green leaves are in three ranks and are one or two metres long, but only 2-3 cm wide. Their bases are tightly folded forming a narrow ridge at the back.
Collospermum hastatum accompanies
Collospermum microspermum is equally specialised but its leaves are as narrow as those of
The collospermums are the only known tank epiphytes outside the family Bromeliaceae, but as they very efficiently form soil, they are also nest epiphytes.
Four New Zealand-wide pteridophyte species often grow as epiphytes with their roots or rhizomes embedded in the soil of epiphyte nests. Though they occur elsewhere as well it is in such sites that their growth is most vigorous and their pendulous stems or leaves attain their maximum length. Of the four, Lycopodium varium is the most impressive, its slender stems sometimes forming huge masses up to 1½ metres long below collospermum nests. The stems branch repeatedly by equal forkings or dichotomies, so that a dense, but well balanced mass is formed. It is almost constantly in motion as even the lightest breeze can set the tassels swaying. In their upper parts the stems are clothed with small spreading leaves, which grade into small, close set scales enclosing the sporangia towards the branch tips.
The ferns Asplenium polyodon (A. falcatum) and
Tmesipteris, a genus restricted to the south west Pacific, is sometimes referred to as a “living fossil” as it is considered to be one of the most primitive genera of land plants. One of the highlights for botanical visitors to New Zealand is to see a living plant of this genus.
Tmesipteris elongata subspecies robusta has been observed growing from Collospermum clumps at a number of localities through the North Island, but not yet in the South Island. Its stems, with their small, simple leaves, are unusually long for a Tmesipteris and dichotomize freely. Other species of Tmesipteris rarely branch. The smaller Tmesipteris tannesis may also hang from tree trunks or branches.
Three orchid species can also be included as pendent epiphytes. The two Earinas belong to a small genus with other species in New Caledonia and Polynesia, but is considered to be closely related to the large genus Epidendrum of tropical America. Both species have spreading stems and can sometimes extend for several metres along branches. The stems bearing the leaves droop downwards and can be 30 or more centimetres long. The leaves are in two rows and more or less in one plane; those of E. mucronata are narrow, thin and quite grasslike while those of E. autumnalis are broader and thicker in keeping with the more robust nature of the plant as a whole. Both species form terminal sprays of small flowers, E. amucronata in the spring and E. autumnalis in the autumn. The flower clusters of the former hang down and are yellowish orange, those of the latter turn upwards and are waxy white with a strong spicy perfume.
Our sole species of the large tropical genus Dendrobium — D. cunninghamii — is the largest of our epiphytic orchids, its freely branching stems and narrow leaves forming feathery drooping masses. The stems are polished, often bright yellow and very bamboo-like in appearance. The white, reddish centred flowers are scattered and while modest by tropical standards, are at 2 to 2½ centimetres diameter, the largest among our epiphytic orchids.
The two species of Pittosporum and one species each of Senecio and Coprosma in this category are usually not more than a metre high when growing as epiphytes, but may attain small-tree size on the ground. All are endemic to New Zealand.
Pittosporum cornifolium is found throughout the North Island and although it is quite a common plant, many people are unaware of its existence perched as it is inconspicuously in tree crowns. The stems of this plant are spindly and they often hang down below the branches. The leaves are thin but firm with prominent veins and the flowers are small and yellowish red. The round, woody seed capsules are a surprise. When they open they reveal a bright red lining and shiny black seed embedded in a sticky bright yellow fluid.
Pittosporum kirkii has a more restricted range, not being found further south than the central North Island. It has a more erect growth habit with thicker stems and longer, thicker almost fleshy leaves with obscure veins. The flowers are bright yellow and the capsules are unusually large, up to 4 cm long, and are flattened and pod-like. Kirk, after whom the species is named, states that the “valves contract in a curious manner when the capsule bursts”. The capsule is apparently not so colourful as that of
Senecio kirkii is found in lowland forests throughout the North Island but has not been recorded from the South Island. Its growth form has been described as “candelabralike”. The leaves are soft and somewhat fleshy and the flowers, up to 5 cm diameter, are pure white and crowded into dense heads.
The thick and shiny-leaved karamu ( Coprosma lucida) is best known as a ground plant in shrubby early forest regrowth on drier sites, but is also reasonably common as an epiphyte in asteliad nests. The species is found throughout the coountry but presumably is common as an epiphyte only within the range of nest epiphytes.
As well as being larger than those of the preceding category, these also eventually send a root to the ground and so overcome the water supply and soil nutrient problem.
Puka ( Griselinia lucida) is the most notable in this category, its large, dark-green shining leaves usually contrasting so strongly with the foliage of the supporting tree that it stands out even to the casual observer (Fig. 5).
Puka is distributed in lowland forests throughout the North and South Islands, but is more common in the north. Its seedlings generally establish in asteliad nests situated at branch forks and its roots ramify through the humic soil. After a few years a strong root begins to grow down the trunk of the supporting tree towards the ground. This root and its branches are closely appressed to the bark of the trunk, frequently growing into crevices and behind bark flakes. The root tips are white and smooth, but from a short distance behind them, the root surfaces are densely clothed with short root hairs. These apparently persist until cork formation commences, as they are undiminished up to a metre from the growing points. Where the roots are in contact with the trunk, they are anchored by the root hairs and the union is sometimes so complete that when the roots are pulled away they either remove portions of bark or leave strips of their own tissue behind.
When the root tips reach the ground generally one main vertical root enlarges greatly until it attains a diameter of ten or more centimetres. This main root usually has a few major branches near the ground and the whole system has a very distinctive appearance resulting from the more or less continuous and pronounced longitudinal grooves and ridges of the bark. In its upper parts the main root gives rise to slender horizontal, girdling roots, which often encircle the trunk or the supporting tree many times and so ensure that the puka will not be dislodged even by the strongest gale.
Two other species, Griselinia littoralis and
Griselinia littoralis (broadleaf) is the only other species of its genus in New Zealand. Its leaves are smaller than those of puka, yellowish green, and symmetrical or only slightly asymmetrical at the base. In puka the leaf base is very asymmetrical as the two parts of the leaf divided by the midrib are of quite different lengths. Broadleaf has been observed as an epiphyte on a variety of trees. Its descending roots are often more massive than those of puka, but they are not grooved. The species ranges throughout New Zealand including Stewart Island. Beyond New Zealand
Pseudopanax colensoi (mountain five finger) also has a wide range, but is absent north of 36 degrees S and from Stewart Island. I have observed it growing as an epiphyte on
The descending roots of puka and mountain five finger seem too slender in relation to their height to stand alone when the supporting trees die, but this may be possible for the more massive roots of the broadleaf.
Northern rata (Metrosideros robusta) is the most notable and common example here. It is found in lowland forest throughout the North Island and near the north west coast of the South Island. It is much more frequent as an epiphyte than a ground plant and prefers the tall emergent conifers as supporting trees. The earlier stages of its life cycle are very similar to those of the puka. Establishment is usually in asteliad nests, although young plants have been observed attached directly to rough bark. A distinctive feature of some small plants of northern rata is the development of tuberlike
With the development of such a massive root system, when the supporting tree eventually dies the northern rata is able to stand alone on its “pseudo-trunk” (Fig. 6). If the support was an emergent then the rata now replaces it in that role.
The northern rata and tropical epiphytic trees of similar habit, are often referred to as “stranglers”. This implies that these epiphytes kill the supporting trees by compressing their trunks with a complete or partial network of roots. Popular writers on New Zealand plants have taken enthusiastically to this idea describing the northern rata variously as a “predatory gangster”, “forest bandit” or “notorious strangler” which “crushes”, “smothers”, “stifles”, or “squeezes” the supporting tree in an “iron” “deadly” or “fatal embrace”.
Partly as a reaction to these verbal flights, some botanists in recent times have tended to take a contrary view. They point out that the light demanding northern rata generally establishes in the well-lit crowns of mature trees so that, by the time the former is large enough to stand alone, the supporting tree might well have died of old age. It does seem, however, that the northern rata must have some deleterious effect on the supporting tree by partial overshading, root competition, and perhaps in cases where the supporting trunk enlarges within a well developed rata root cage, by some restriction in the movement of water and dissolved nutrients.
Recently a distinctive new tree species of Metrosideros has been discovered. It is restricted to a few forest patches near North Cape and has a similar epiphytic habit to northern rata.
Southern rata ( Metrosideros umbellata) is rare and localised in the North Island, but quite common in montane and higher latitude lowland forests in the west of the South Island. It is mostly terrestrial, but has been observed growing as a “strangling” epiphyte at several places.
Similar Metrosideros epiphytes are known in New Caledonia, Fiji and Hawaii.
These are considered separately, as tree ferns provide a substrate rather different from the bark of ordinary trees. To begin with tree ferns do not branch, so only the trunk is available for colonisation. Secondly, the trunks are built up from persistent leaf bases, which provide a variety of surfaces according to the species, but none is quite like bark. Also the large crowns of leaves cast considerable shade so any epiphytes need to be shade tolerant at least when young.
The different species of tree fern vary in their suitability for epiphytes. In our largest and most handsome species, the mamaku ( Cyathea medullaris) the leaf bases decay down to hard leafscars which collectively form an armour-like surface unsuitable for epiphytes. In the lower parts of the trunk masses of slender roots grow out adding considerably to the diameter of the trunk, but these too form a hard dry surface.
In the ponga or silver tree fern ( Cyathea dealbata) the leaf bases decay more gradually and do not form well-defined scars. As a consequence soil forms readily in the interstices and a variety of epiphytes are able to establish. Wheki (
In moist situations lichens, mosses, liverworts and smaller and larger ferns may be abundant on tree fern trunks. Climbing ratas and other climbers may also be present as well as a range of seedlings of trees and shrubs which die before reaching maturity. Here we will consider only the consistent and specialised tree fern epiphytes. It is worth noting that the species concerned are mostly different from those occurring on ordinary trees.
A Lycopodium is quite commonly encountered. It is smaller than Lycopodium varium on trees, and the leaves associated with the sporangia towards branch tips are often not reduced to scales. Some treat it as a distinct species,
Asplenium flaccidum, common on trees, also inhabits tree fern trunks. Several species of
One shrub and one tree frequently, and several other species less commonly, play this role on tree ferns.
Five finger ( Pseudopanax arboreus) is common as a terrestial plant in shrubby forest regrowth, but in more mature forest it can be surprisingly frequent as a tree fern epiphyte, mostly on the ponga, but also on wheki. The seedlings establish at the top of the trunk and being fairly light demanding, their leaves soon push between and above the fern fronds. The primary root begins to grow down to the ground, but soon gives off a branch root which grows horizontally around the trunk sometimes returning to and fusing with the vertical root. It is thus comparable with the girdling roots of the puka and northern rata. The vertical root eventually reaches the ground and sometimes branches to enclose the tree fern trunk in a network of roots in its lower part. In the meantime, the crown of the five finger has continued to branch and grow upward with the tree fern crown following behind it. (Fig. 7).
Less frequently Pseudopanax edgerleyi (raukawa) and Coprosma grandifolia adopt a similar life style.
Kamahi ( Weinmannia racemosa), the main canopy dominant in many montane and higher latitude forests, also frequently begins its life as a tree fern epiphyte, particularly on wheki (
As had already been mentioned, epiphytes sometimes grow on the ground and in some circumstances may be important components of terrestrial communities. On Rangitoto Island for example, a volcanic cone only a few centuries old in Auckland harbour, the epiphytes northern rata (Metrosideros robusta)), puka ( Griselinia lucida), Senecio kirkii,
Epiphyllae as they are termed, are not so evident on the generally smaller leaves of the New Zealand rain forest as they are on the large leaves of the tropical rain forest. As in the tropics the plants concerned are filamentous algae, leafy liverworts and lichens.
One epiphyllous alga in New Zealand is commonly observed on the leaves of mahoe ( Melicytus ramiflorus.) This is a species of
Five epiphyllous species of liverwort each from a different genus, have been recorded in New Zealand. They are related to epiphyllae of the tropics and in New Zealand have been found mostly on fern leaves, but also on leaves of trees and shrubs including Pseudowintera(horopito).
Epiphyllous lichens were the subject of a detailed study by Allan at Kitchener Park, Feilding. They were found to be abundant on the leaves of the conifers totara (Podocarpus totara), matai P. spicatus and kahi katea (P. dacrydioides), on tawa ( Beilschmiedia tawa), titoki (
The leaves of supplejack (Ripogonum scandens), the species of Coprosma, Pittosporum, Hoheria and puka ( Griselinia lucida) were free of lichens.
Clearly much still remains to be learnt about leaf epiphytes in New Zealand.
Although all flowering plant parasites agree in having structures known as haustoria, which penetrate into the living tissues of the host, they differ quite widely in a number of other respects. Some are complete parasites as they lack chlorophyll and so are unable to utilise light energy to manufacture sugars. Others, having green leaves, make their own organic nutrients and derive from the host mostly water and inorganic nutrients.
Some parasites are attached to roots, others to trunks and branches.
Dactylanthus taylori is a complete parasite attached to the roots of a range of mostly small tree species in lowland to montane forest throughout the North Island. It is not readily observed as only the reddish-brown scaly inflorescences appear above the ground. The strange appearance of the flower heads apparently arising directly from the ground, led the Maoris to give the name Pua reinga (Flower of the Underworld) to this species.
Apparently the embryo root of a Dactylanthus seed penetrates the slender root of a suitable host, then gradually expands into a tuber-like structure which eventually surrounds the host root. The terminal portion of the latter then dies away. The “tuber” continues to enlarge and the end of the host root enlarges with it into a more or less disc-like form. Both can attain a diameter up to 30cm. The tuber has a flattened balllike shape and is covered with hard warty protuberances. Inflorescence buds originate between these, bearing male flowers on some plants, females on others. The flowers have a strong sweet perfume which is attractive to flies.
The junction between host and parasite is not flat, but formed into radiating grooves, v-shaped in section. It has been found that if the host/parasite mass is boiled the parasite can be removed exposing the expanded, fluted ends of the host roots. These “wooden roses”, as they are called, are prized as curios.
Dactylanthus is restricted to New Zealand but belongs to a largely tropical and subtropical family.
Although Mida salicifolia is a root parasite on a wide range of trees including kauri, unlike
These all contain chlorophyll so are only partly dependent on their hosts for organic nutrients. All but one of the New Zealand parasites in this category belong to two largely tropical families — Viscaceae and Loranthaceae, collectively referred to as mistletoes. In the first the flowers are small and inconspicuous, in the second they are much larger and often brilliantly coloured.
Three small species of Korthalsella represent the Viscaceae. All have vestigial leaves and strongly jointed stems, which in K. lindsayi and K. clavata are strongly flattened and in K. salicornioides cylindrical. The latter is found throughout the country, the former two from the central North Island southwards and both parasitise a wide range of shrubs and small trees.
The New Zealand species in the family Loranthaceae are all green-leaved, freely branching shrubs up to one metre in diameter. Currently all these species are referred to a number of small genera, with one exception endemic to New Zealand, although formerly some were included in the tropical genera Elytranthe and Loranthus.
Tupeia antarctica is the only species of a genus restricted to New Zealand. Each plant is attached to a ball-like mass, which is a combination of the parasite haustorium and host tissues.
Peraxilla colensoi and
Ileostylus micranthus has small green flowers and yellow berries. It is widespread in New Zealand and Norfolk Island and has as hosts a range of shrubs, small trees and sometimes conifers both native and introduced.
The species of all these genera except Tupeia send out roots over the bark surface which form secondary haustoria at intervals. Ileostylus alone can form new leafy shoots from its roots.
The branch parasites have berries eaten by birds and the seeds are deposited on tree branches. The seeds are attached to bark by a sticky secretion.
Some branch parasites elsewhere have explosive fruits, which shoot the seeds for several metres. This has been observed in the New Zealand species of Korthalsella.
The remaining parasite in this category is Cassytha paniculata and it is quite different from the rest as it is a twining vine as well as a parasite. Its seeds germinate in the ground and the slender yellow green primary stem with rudimentary leaves rotates in anti-clockwise direction winding tightly around any stems it encounters. At frequent intervals haustoria penetrate the host. The stems of the Cassytha branch freely, but remain slender, festooning the shrub hosts with tangled stringlike masses. This species is restricted to the northern half of the Northland peninsula where it grows on shrubs and particularly manuka ( Leptospermum scoparium). It is surprising to find that
Vascular saprophytes are completely without chlorophyll and are often small, pale plants growing in leaf litter in very shady places in rain forests. It is thought that they gain their organic nutrients from decaying plant material. Their underground parts are penetrated by fungal threads and recent studies, some in New Zealand, have shown that in some cases the fungal threads are also attached to the roots of nearby trees. It is suggested that such saprophytes and probably others, may be secondary parasites drawing nutriment from tree roots via fungal threads.
The New Zealand saprophytes are all orchids, except for one belonging to the Burmanniaceae a family closely related to the Orchidaceae.
The one non-orchid is Thismia rodwayi. It has only been found in the northern half of the North Island and there mostly on the volcanic plateau. The pinkish scale-leaved stems, arising from a branching root system, each end in a relatively large delicate flower, which has been likened to a red lantern. Our species is also found in Tasmania and Victoria and there are other species in Australia, tropical Asia and America.
Corybas cryptanthus is the only saprophyte among the eight New Zealand species of the genus. It has been collected at scattered localities throughout the country. Only the flower appears above the leaf mold, the stem then elongating to carry the capsule to about 15 cm above the ground. The genus ranges from south-east Asia through Australia to New Zealand.
The 15 species of Gastrodia ranging from India and Japan to Australasia, are all saprophytes. The branching underground rhizomes are tuberous and filled with starch and those of our species were eaten by the Maoris. The stems are tall, up to one metre, and can be attractively if strangely coloured. They often appear as if polished, with flecks of white and brown giving a resemblance to wood grain.
C. cunninghamii, mostly in beech forest, and G. minor mostly under Leptospermum are found throughout. G. sesamoides has not been discovered further south than 42 degrees S in the South Island and is found in open forest and shrubland.
Yoania australis was only discovered in recent times, but is now known from several localities in Beilschmiedia tarairi forest on the Northland peninsula. The stems bearing the small flowers are a pale rose colour and up to 20 cm tall. The genus is entirely saprophytic and is known at several localities in Asia and north Africa.
In view of its wide range of specialised growth forms it is not difficult to conclude that New Zealand conifer broadleaf forest comes closer to tropical rain forest in this aspect than to any other type of vegetation despite New Zealand's temperate latitudes. In the light of certain fossil evidence the most likely explanation for this is that, before the Ice Age, forest of the general type now largely confined to tropical latitudes was also widespread in middle latitudes of both hemispheres. Plant fossils from the vicinity of London dating back to early Tertiary times (80 million years ago) belong to genera, including some of palms, now largely restricted to the tropics. Fossil floras with similar relationships have also been discovered in Oregon, U.S.A. The Ice Age, whose effects would have been more severe in the largely continental northern hemisphere, virtually eliminated such forests from middle northern latitudes while limited examples persisted in middle southern latitudes and in New Zealand best of all.
Remnants of such middle latitude rain forests, with fewer species than those of New Zealand and in particular fewer vines and epiphytes, can be found in parts of New South Wales and Victoria in Australia, along a portion of the south-east coast of South Africa and in Central Chile. The rain forests of these areas may have been more reduced than those of New Zealand by the development of arid continental climates and by their longer history of human and natural fires as well as by Ice Age coldness. New Zealand's narrow oceanic land mass would have ameliorated the two climatic factors and enabled the survival of our fascinating array of vines, epiphytes, parasites and saprophytes.
Keywords: Creationism, evolution, fossil, history, philosophy, religion.
The history of the debate between evolutionists and creationists in New Zealand is examined. The history of this debate is compared to that in the United States and the activities of The Institute for Creation Research in the United States and New Zealand is discussed. A general discussion of belief systems forms a basis for discussion of the Institute's activity. The nature of the scientific evidence presented by the Institute is examined and found to be inadequate by normal scientific standards. An appeal is made for New Zealand scientists to recognise the interest people have in their origins and to provide information for the general public.
In New Zealand, debate on the merits of evolution and creationist world views goes back to the 1870's. Arguments occurred between Darwin's supporters, such as
The arguments were fierce. However, as Stenhouse (1984) strongly points out, the arguments were between scientists who were, and remained, orthodox religious believers. There were no New Zealand equivalents to the militant English agnostics like Huxley and Tynfall. It was not a conflict between science and religion. Attempts to define the conflict in so simple a fashion stemmed largely from the United States.
Over the years local groups, such as the New Zealand branch of the Evolution Protest Movement, continued to produce some creationism literature for local fundamentalist groups (e.g. Milne, undated) or to distribute literature from overseas publishers writing on behalf of groups such as the Jehovah's witnesses (anon., 1967). However support for creationism by scientists had virtually disappeared by the end of the 19th Century (see e.g. Parsonson, 1984).
Why then do so many of the general public get the impression that the occurrence of evolution is still a live scientific issue? I believe there are two main reasons for this in New Zealand.
First, biologists themselves have been involved in heated debate recently about the mechanisms of evolution. For example it has been argued that Darwin's natural selection is of secondary importance (Grehan, 1984), poorly understood (Cherfas, 1984), well proved (Charlesworth, 1984) and a universal biological law (Reed, 1981). It has been argued that evolution is gradual (Smith, 1981), rapid (Iltis, 1983) or a mixture of the two (Lister, 1984). It has been argued that evolution has come about by subtle molecular mechanisms (Dover, 1982) and as a result of mass extinctions brought about by the impact of an asteroid (Hallam, 1984) or even regular cosmic catastrophies (Maddox, 1984). In this welter of argument the layman could well be excused for thinking that the whole idea of evolution was under attack.
To most biologists, on the other hand, the argument is steadily strengthening our understanding of evolution. Whether that mixture of evolution and genetics often called neo-Darwinism will survive in its classical form with a new lease of life or be so transformed that it will need another title remains to be seen. However that argument resolves itself, there have been no recent suggestions in major scientific journals that evolution has not occurred. A brief argument about whether evolution qualifies as a genuinely scientific theory (“theory” is used here to mean a model for which sufficient evidence exists for it to be used as a base for further predictions, as opposed to a “hypothesis” which must be further tested before it is used in this way) seems to have been firmly answered, and the answer is yes (Sparkes, 1981).
The second reason for public doubt comes from the activities of an interconnected San Diego based group of organisations. These are:
the Institute for Creation Research (I.C.R.)
Christian Heritage Colleges
Creation Research Society.
Membership of the Creation Research Society stands at about 550, all with M.Sc. or Ph.D. qualifications, and all having signed a statement of faith as fundamentalist Christians. The Christian Heritage Colleges exist to train fundamentalist Christian teachers, with a strong emphasis on creationist teachings. The I.C.R., which I visited in July of 1983, is physically adjacent to a Christian Heritage College, shares staff with it, and works under contract to it. Thirty students have graduated over the past three years although only one has been in biology. This student's thesis was entitled “A rationale for the Christian College Biology Curriculum” (Anon, 1983). Research generally has been “literature based” or “field based” (Dr Morris, pers. comm. 1983). The I.C.R. is only now in the process of building laboratories, although it has accepted graduate students for training for several years.
I.C.R. members are seldom represented at scientific conferences, their publications tend to be published by the organisation itself and therefore are not subject to the usual peer scrutiny, but they are very active politically. They are, for example, putting pressure on boards that select schoolbooks in the United States not to buy books that devote time to evolution unless they also give space to creationism. In general, publishers have not been willing to promulgate creationism, but they have tended to seek to increase their sales by cutting back sections dealing with evolution. Boards are also being pressured to include “creation science” in their syllabi (Budiansky, 1983). This pressure would be even more effective if it were not for the activities of scientists who, with the backing of the American Association for the Advancement of Science, attend school board meetings and put the case for science. Pressure on legislatures has produced bills for equal time for creation and evolution in schools in Arkansas (where one such bill was recently declared by a Federal Court to be a constitutional violation of the separation of church and state), and in Alabama, Mississippi, West Virginia and 21 other states where the bills were defeated in the legislature (Budiansky, 1983).
In the States this crusade dates back to William Jennings Bryan and his concern that people with a university education were caught up in a flood of disbelief sweeping campuses and which he attributed to the teaching of evolution (Numbers, 1982; Marsden 1983). Support for the cause is widespread in the United States to this day. One Judge Braswell Dean was quoted in Life magazine for January, 1982, as saying: “This monkey mythology of Darwin is the cause of permissiveness, promiscuity, pills, prophylactics, perversions, pregnancies, abortions, pornography, pollution, poisoning and proliferation of crimes of all types.”
Many scientists reading this may well have the reaction that this is the kind of thing one expects to encounter in the United States but it couldn't happen in New Zealand. In
I believe that there are a number of important issues that need to be raised in this context. The first is the intensity of belief involved and people's rights to their own beliefs.
Everyone has a unique way of explaining the mass of data they take in through their senses. We perceive the sights, textures, odours, tastes, sounds, and words of our world and then seek to make sense of these by making pictures, saying words or using our imagination to reproduce inside our heads, patterns which allow us to give an imaginary past, imaginary future or an imaginary internal working to what we see (Lankton, 1980). Since we are receiving far more information that we can possibly pay attention to, the way we perceive the world, i.e. what we pay attention to, is itself influenced by the patterns we use inside our head to explain things. None of us likes to change these patterns we have developed. Most of us do change them from time to time, either to make them more internally consistent or to bring them more into line with other people's. This applies to all belief systems. Scientists are no happier to give up their belief systems than anyone else. Thus as Sparkes (1981) points out, there are “core theories” in science which are heavily protected, i.e. we would rather explain away exceptions than change the “core theory”. The core theory with its subsidiary explanations is what Kuhn (1970) refers to as a paradigm. There are paradigms in the study of history and religion. What is different about science is that “core theories” and all other scientific hypotheses and theories are constantly producing predictions which can be tested in the real world in a repeatable way. If too many of these predictions fail, then even “core theories” will be modified, as Newton's theories were by Einstein's.
The views of individual scientists will of course be coloured by the theories they know and their own philosophical background. Thus even the experimental process itself, as well as its results, will be modified by the attitudes the experimenter already has (see Fig. 1). In this figure (modified from Sparkes, 1981) I have referred to a scientist's existing set of paradigms, scientific or non-scientific as “authority filters”.
People are naturally very interested in their origins and every group seems to have its own paradigm of how we got here. Sproul (1979) gives 134 versions, including seven biblical versions and four Maori versions. This in itself, by the way, makes nonsense of the I.C.R.'s claim for “equal time for teaching creation and evolution in science courses”, since they only want “equal time” given to their own particular version of creation. The scientific equivalent of these creation myths is the theory of evolution. That scientists have as much emotional investment as anyone else in their favourite version of “how we got here” can be measured by the heat that is sometimes generated in discussions of the mechanism by which evolution operates. Because of the large
For example, my own background is that I was a fundamentalist Christian as a teenager. I was unusual in that I accepted most of the doctrine, e.g. need for salvation through a conversion experience, but had no problem in also accepting evolutionary theory when I came across it. Later I read widely in other religious writing and became convinced that it was unlikely that any one group has a monopoly on religious truth. My present position is that I am a theist in the sense that I believe that we are all part of some thing/being, and that sacred writings are an attempt to come to terms with this. From this point of view creation and evolution are equally satisfactory models. My prejudice towards evolution comes from 25 years experience of finding it a useful thinking tool in looking at biological systems as they exist today. Creationism does not make sense of the data.
Members of I.C.R. on the other hand are all committed fundamentalist Christians. They believe that the bible is literally true and they are convinced that the results of their “research” will be “even greater confidence in the Word of God” (Vardiman, 1984). Jukes (1984) accuses Gish of deliberate lying about the nature of a chemical reaction (Gish continues to claim the bombardier beetle uses an impossibly explosive mix of hydrogen peroxide and hydroquinone “Although he knows that — the mixture slowly and quietly turns brown” instead of exploding). From my contact with members of the Institute I doubt that they would lie deliberately to further their cause. I was invited by them to visit their Institute and at no time had any impression that they wished to mislead me or hide anything from me. Rather, I have the impression of very sincere men with a paradigm so inflexible that even correcting obviously wrong detail is very difficult for them.
With this in mind the we can look at I.C.R.'s claim to be a scientific organisation. I.C.R. has also been criticised by the British Biblical Creation Society as being ascriptural and anti-scriptural (Professor Edgar Andrews, reported in Howgate and Lewis, 1984) and on numerous grounds by a wide spectrum of theologians (Frye, 1983) but these are theological arguments and beyond the scope of this paper. I wish to suggest that from a scientific point of view the work of the Institute can be broadly divided into two categories or parts. The first part is not scientific, and in some cases is anti-science, in that it uses unscientific methods to attack scientific work, and a second category which is science but simply poor quality science.
Each month the I.C.R. publishes an article in its Impact series. In the last 12 months (to September, 1984) the following have appeared: one article on politics (getting “God-centered” theories into schools), one on (?) archaeology (the search for the Ark on Mt Ararat), one on the finances of I.C.R., one on philosophy, and four that are pure theology, using arguments based solely or mainly on the Christian Bible. All eight articles I would rank as non-science. This leaves three that could be called scientific, to which I will refer below, and one that is a good example of what I have called anti-science. In this last article, Morris (1984) uses quotations out of context from cosmologists to attack the assumed views of evolutionists. This is a common method of argument by this group. It is a good debating tactic, used in many of their publications. In Morris (1982) we find scientists such as Stephen Jay Gould, Colin Patterson, Kimura Motoo, Pierre Grasse, Roger Lewin, and Sewell Wright, who have made major
Those arguments that I.C.R. members use against evolution involve misunderstanding, often of very basic science. I will take three examples.
Creationists' argue that the second law of thermodynamics makes evolution impossible since it requires that all systems increase in entropy (i.e. become less and less ordered with time) (e.g. Wysong, 1976; Morris, 1984). As Hull (1982) and others have pointed out, this law applies only to closed systems, with or without a source of energy. It therefore does not apply to Earth since Earth is an open system (see Fig. 2). In such systems with a heat source (the Sun), a receiver and transmitter of energy (the Earth) and a heat sink to receive the energy (space), entropy either remains constant or decreases. Bernstein et al (1983, p. 190) describe a simple experiment to demonstrate this principle. A frying pan (preferably with a copper bottom and about 20-30cm in diameter) is filled with enough oil to just cover the bottom (about 1 mm deep) and placed on an electric element of about the same size as the bottom of the pan. At high stove settings the surface will appear dimpled as a result of developing a convection pattern. What is happening is that a heat source (the stove) is supplying heat to the oil (the receiver and transmitter) which then radiates it into the room (a heat sink). The parallels to our solar system are obvious and the decrease in entropy (production of pattern) is both obvious and repeatable.
This situation, called a steady-state system, is stable for long periods of time so long as: deS = diS ≤ O [Where deS is the entropy flow between the system and its environment and diS is the entropy production of irreversible processes within the system (diffusion, chemical reactions, heat conduction, etc.)(Csanyi, 1982, p.17)].
The three issues of Impact referred to above as scientific, include that of Robbins (1984) asking if the redwoods of California can be used to date the flood. Since he only raises the question but does not suggest how it could be answered, I will take it no further. Vardiman's (1984) paper speculates on the possible atmosphere before and during the Biblical flood. It is highly speculative and depends on the prior assumption that a world-wide flood has taken place. Geologists have no evidence that this has occurred. The third article (Cumming, 1984) uses examples from life cycles of several organisms and from biochemistry (his later reference to this material as “ecology” is puzzling) to support Paley's (1802) argument that nature is so well designed that it requires the direct intervention of a designer. It seems appropriate to answer such an ancient argument with classic answers. As Darwin and Hutton pointed out over 100 years ago, the following suggest that if a designer had directly intervened it must have been on a bad day:
the New Zealand native bird, the kakapo, has wings that are too long for its weakened muscles to operate.
young calves have canine teeth they never use.
whales have a pelvis so tiny that it is hard to find.
the Patagonian goose has webbed feet but never goes near water.
Certainly there are many examples of fine adaptation of organisms to their environment. Darwin's idea of natural selection following natural variation was based in part on this observation. Darwin's theory also quite happily explains relict structures, retained to be used for some new purpose, or being gradually phased out as
Finally let us consider the work by Wysong (1976). Wysong, more than most creationists, attempts to be scientific (or if one is uncharitable, goes to great lengths to hide his creationist views until late in his book). Let us then examine one of the real scientific tests of creationism as set forth by Wysong. If the creationists are correct and the earth is only about 10,000 years old, then we should find the remains of what we regard as modern organisms mixed in with those which we believe to be from the geological past. Wysong (1976, pp. 370-383) produces 15 items as supporting such “disordering”. Let us consider them in turn.
Item 1 refers to the finding of pollen from Angiosperms in supposedly pre-Cambrian rocks. Stainforth (1966) in reporting this find admits it poses a problem but suggests several possible mechanisms. The item then goes on to quote Axelrod (1959) on woody plant remains from Cambrian rock. Wysong says: “Woody plants supposedly did not arrive on the evolutionary scene until over 200 million years after the Cambrian”, implying that this find represents an anomaly. Axlerod on the other hand actually states on the first page that the finding of these ancient plants solves many problems concerning the early evolution of land plants, and goes on to quote other records of early Cambrian land plants.
Item 2 refers to the finding of pre-Cambrian arthropod fossils by the U.S.G.S. but gives no references.
Item 3 quotes the speculative writer Velikovsky as saying that a figurine made by humans was found under “tertiary” larva in the course of drilling for water.
Item 4 quotes a newspaper report that a gold chain was found embedded in a “chunk of coal” and item 5 deals with an iron pot also encased in coal reported in a Creation Research Society publication.
Item 6 brings up the suggestion of human foot prints found among dinosaur footprints. The only reference given is to Ingalls (1940). Ingalls states “unless 2 and 2 are 7 … these prints were not made by Carboniferous man”. The possibility of mistaking dinosaur footprints for human ones, given their lack of clarity and the possiblity of fraud by local guides or other persons, has all been pointed out before. Milne and Schafersman (1983) give examples from the Paluxy River-Glen Rose site which seems to prove fraud by someone.
Item 7 is an old (1860, 1886) record of human remains in “pliocene strata” quoted from the British Evolutionary Protest Movement.
Items 8 and 14 refer to human remains found in coal or rock. Unfortunately the references are unavailable to me.
Item 9 is a picture that purports to be a trilobite squashed into the impression of a shoe. The trilobite is not clear and no evidence is given to suggest that the two shallow symmetrical impressions are made by a shoe rather than some natural agency.
Items 10—13 concern pottery designs and pictures made by primitive peoples and said to be of 5-toed llamas, dinosaurs and mastodons. This is then used as an argument that the animals concerned must have been contemporaneous with the peoples who made the representation. The one example figured (fig. 123) does not reveal anything like the detail which would be needed to back up such a claim.
Item 15 lists several similar items from one of the Society's own publications.
Clearly the standard of “proof” given here is below that which would be accepted from a junior university student, much less than the standard required by a science journal. Until such time as creation scientists can produce evidence that shows significant fossils clearly misplaced in the time frame that evolution would predict, we are entitled to be very sceptical. The obvious place to look for such inconsistencies is among those organisms that leave the best fossils, such as ammonites and foraminifera.
It is not surprising that some people like the creationist approach. Modern science has reached a stage where we are forced to recognise that the world is complex beyond our power of understanding. It is much harder to imagine a beginning for our universe 20,000,000,000 years ago than to go with Bishop Usher, who in the 17th Century worked back through the genealogies in the Bible and decided that creation occurred at 9a.m. on the 23rd October, 4004B.C. People prefer simple answers. So do scientists, for after all we often use Occam's razor to choose between hypotheses. As good scientists we will choose the simplest hypothesis that fits the facts. That people can accept that the creationist argument, as put foward at present, fits the known facts of Biology, Astronomy, Chemistry, Physics, etc. is a sign of the organisation and commitment of I.C.R. supporters and an indictment of the job we have done as scientists in communicating our knowledge to the general public. Perhaps the best way to rectify the situation is for scientists to make themselves available to schools, churches and other interested bodies. I hope that those responsible for education and for informing the public will feel free to approach our universities and research centres for assistance.
I wish to thank Professors
Key words: Anthocerotae, Dendroceros allisonii, Dendroceros endiviaefolius, Dendroceros giganteus, Dendroccros granulatus, Dendroceros megasporus, Dendroceros nodulosus, Dendroceros validus.
The taxonomy, morphology and distribution of Dendroceros giganteus (Lehm. & Lindenb.) Prosk., D. granulatus Mitt, and D. validus Steph. are reported. D. allisonii Herz., D. endiviaefolius (Mont.) Prosk., D. megasporus Herz. and D. nodulosus Steph. are reduced to synonymy.
Key to New Zealand species.
(Synonymous with Dendroceros endiviaefolius (Mont.) Prosk. and Anthoceros colensoi Mitt.)
In New Zealand this very beautiful species is found growing as a caespitose carpet overlying mosses, liverworts or water-retentive litter in swampy ground in regions of high rainfall, as in Fiordland, Arthur's Pass and the Ruahine Range. It occurs also on the Auckland Islands (collected P. D. endiviaefolius — a name regarded here as a synonym.
At the British Museum I was able to examine the type material by courtesy of the Curator of Bryophytes. Other specimens that I examined came from the herbaria of the British Museum, the New York Botanical Garden, National Museum at Wellington, Botany Division at Christchurch and E. A. Hodgson at Massey University. Living plants were collected at Arthur's Pass, Milford Sound, Wilmot Pass and Mt. Cargill and grown in culture at Massey University. The species was first named Anthoceros giganteus (Lehmann, 1832), then transferred to Megaccros (Stephani, 1916) and later to Dendroceros (Proskauer, 1953). Proskauer remarked on its similarity to D. endiviaefolius of Chile and Argentina, as had Schiffner earlier (1890) using the name Anthoceros. Detailed notes and illustrations of D. endiviaefolius are given by Proskauer (1953) and by Hassel de Menendez (1962). Mitten (1855) described Anthoceros colensoi from specimens collected on the summit of the Ruahine mountains and now housed at the New York Botanical Garden. Already Proskauer (1953) has used the name Dendroceros giganteus for these plants. Goebel used living New Zealand plants in his studies (1905, 1906) but Khanna (1944) unfortunately, as Proskauer (1953) noted, did not have correct material of this species.
The bright-green, strap-shaped thallus, up to 5 cm long and 1.5-5(−8)mm wide, may branch a few times (Fig. 1). It consists of a flat or slightly convex midrib region and wide unistratose wings, which are deeply cut into rounded fimbriate lobes, that at times are so greatly folded that they completely obscure the midrib. However, in young parts of the thallus the wings are absent. The tip itself is slightly coiled (Fig. 3). Rhizoids are
The midrib region is up to 27 cells high. Above and below are small cells each with 1-4 disc-shaped chloroplasts. The central cells are larger with up to 8 chloroplasts except for scattered cells containing mucilage. They are somewhat elongated in an axial direction and often their walls appear pitted due to enlarged primary pit fields. Pores are present on the underside of the thallus leading into cavities. These may become
Nostoc colonies which usually grow so large that they protrude from the surface.
The wings develop late from a marginal meristem, resulting in a regular cell arrangement similar to that derived from a cambium. They are unistratose and except in the scattered mucilage cells have 1-4 disc-shaped chloroplasts per cell.
The slender brown capsule lacks stomata and is up to 8cm long and 0.3mm wide (Fig. 2). It is surrounded at the base by an involucre, 2cm high, which gradually tapers upwards from a basal width of 2mm. When ripe the capsule splits some distance below the apex into 2 valves which remain united at the tip and twist spirally. The spores when shed are green, unicellular and up to 58 m in diameter. The inner face is finely dotted (punctate) and has an indistinct triradiate marking (Fig. 4a.); the spherical face is punctate and verrucose, with irregular knob-like protuberances 2.5um high (Fig. 4b). The elaters are 180-250um long and have a maximum width of 8-12 m narrowing somewhat towards the tips; they are made of 2-4 cells with a pale-brown helical band.
(Synonymous with Dendroceros allisonii Herz.)
This species was collected by Schiffner in 1894 on Mt Singalang. Bonner (1965) gives the distribution as Sumatra and Philippines. Plants from New Zealand, which are now considered to belong to this species, were first collected by Lophomyrtus bullata near a stream in Pelorus Bridge Scenic Reserve, Marlborough on 30/9/1983 and independently by B. V.
Sneddon on 9/12/1984 and by T. Moss on 10/12/1984 in the Akatarawa Range.
Probably it has been overlooked in other New Zealand localities for it grows intermixed with other epiphytes and is obvious only when sporophytes are present. (Fig. 5).
Dendroceros validus was first described by Stephani (1917) from Schiffner's specimen which is now housed in the Herbarium at Geneva. A more complete description in English, accompanied by several illustrations, was provided by Hasegawa (1980). Herzog (1935) gave a Latin description of what he named D. allisonii, using a New Zealand specimen which had been collected by
The thallus is green in colour and about 1cm long and 5mm broad. It may branch a few times. There is an ill-defined midrib region, about 0.3mm wide, which is flat above and convex below and thin, broad, crispate wings which often become so strongly folded and curved upwards that they obscure the midrib (Fig. 6). Short rhizoids arising ventrally from the midrib attach the plant to twigs or bark. Plants are monoecious. Antheridia occur singly in antheridial cavities on the midrib, in which position archegonia and later sporophytes are also located.
The midrib region, as seen in transverse section, is compact (Figs 7 & 8). Medianly it is 6 cells (0.12mm) deep, but becomes thinner as it merges into the wings which are only one cell deep. The internal cells show conspicuous simple pits on the walls and usually contain a single chloroplast. Those of the upper surface each have a somewhat thickened outer wall and one large chloroplast while some of those on the lower surface produce rhizoids. On both surfaces near the apex, pores are present and further back Nostoc colonies bulge strongly from the lower side of the thallus. In the wings each cell, except for occasional mucilage cells, has a single large chloroplast of irregular shape.
Often at the angles where the cells meet there are small corner thickenings (trigones) or sometimes these are perforated so leaving air-filled regions (Fig. 9). The outer walls of the cells are somewhat thickened.
The sporophyte, 8-15mm high, is at first green and is surrounded at the base by a brown cylindrical involucre 3-5mm high. The capsule lacks stomata and on the outside has a trabeculate appearance due to the outermost cells having thickened lateral walls (Fig. 10). When ripe it splits from the apex downwards into 2 valves which turn chestnut-brown and undergo slight twisting. The spores, 60-65 um in diameter, are multicellular when shed. Their surface is finely punctate (Figs. 11a & 11b) and has a weak triradiate marking (Fig. 11a). The elaters consisting usually of 3 cells, are 200 um long and show a helical band of thickening.
(Synonymous with Dendroceros nodulosus Steph. and Dendroceros megasporus Herz.)
Dendroceros granulatus is widely distributed in the islands of tropical Asia and the South Pacific (Hasegawa, 1984). Plants considered to belong to this species are found in northern New Zealand as far south as Wairoa. They grow as an epiphyte intermixed with other bryophytes on the trunks of nikau palm, Rhopalostylis sapida Wendl. & Drude, and on the stems of trees and shrubs such as manuka,
Dendroceros granulatus was first described by Mitten (1871) from plants collected in Samoa by Powell. Later Stephani (1917) described it separately and also listed it as a synonym of D. javanicus Nees. Unfortunately Stephani's outline drawing (sheet 2248) shows little detail. A more complete description accompanied by illustrations is given by Hasegawa (1982), who kindly supplied a duplicate specimen collected in 1982 in New Caledonia (N. Kitagawa 22259). The name D. nodulosus was given by Stephani (1909; 1917) when he described a specimen collected by Cheeseman in Auckland in 1895 and now housed in the Stephani Herbarium, Geneva. Notes and drawings made by Stephani (sheet 2299) were available for study by courtesy of the curator of the Stephani Herbarium. The species has been collected frequently in New Zealand and is represented in the herbaria of E. A. Hodgson, National Museum and Botany Division. A specimen collected last century by Anthoceros colensoi in the Mitten Herbarium, has been annotated by Proskauer as presumably D. nodulosus. D. megasporus was established by Herzog (1935) who provided a Latin description based on a specimen which had been forwarded by E. A. Hodgson and had been collected by D. nodulosus by D. granulatus. The description given below is based on fresh material collected from Piha, Auckland, New Zealand and maintained in culture at Massey University.
The thallus is green in colour and up to 3cm long and 2.5mm wide. It may branch a few times (Figs. 91 & 13). There is a midrib region, up to 0.8mm wide, which is flat above and convex below, and broad wings which are deeply lobed and under moist conditions a little crispate. However, on drying the wings curl upwards strongly. A few short rhizoids may be present on the under side of the midrib. Plants are monoecious.
As seen in a transverse section the midrib region is compact (Figs. 14 & 15). Medianly it is up to 11 cells (0.26mm) deep but towards the wings it gradually becomes thinner. The upper and lower cells have a single chloroplast of irregular shape while the latter may show a somewhat thickened exterior wall. Large blue-green algal colonies, forming within pores, produce prominent projections on the lower surface and occasionally on the upper surface also. The central cells each have 1 or 2 chloroplasts and often show trigone thickenings at the angles and a reticulate appearance on the lateral walls due to large primary pit fields. (Fig. 15). The wings are unistratose and, except in the scattered mucilage cells, have one large chloroplast per cell. At times there are triangular or rectangular trigones at the angles and these may be perforated (Fig. 16). As well, in older parts of the thallus, there are a few larger perforations where one or more mucilage cells have broken down (Fig. 17).
The sporophyte, usually only 10mm high, is surrounded at the base by a tuberculate involucre 4-5mm high. The capsule lacks stomata and is at first green but later turns chestnut brown. It is described as nodulose because of prominent thickenings on the lateral walls of the epidermal cells (Fig. 18). When ripe it opens at the top by a slit on one side, then flattens and twists slightly as the slit lengthens. The spores are green and multicellular when shed, with a diameter of 55-80 m. The surface (Figs 19a & b) is punctate (dotted) except at the weak triradiate marking (Fig. 19a). Elaters consisting of 3-5 cells are up to 400 m long and have a pale brown, helical band.
Note: Once the spores are shed regeneration takes place by new shoots developing at the margin of the thallus and the old plant tends to die off.
The writer is deeply indebted to J. Hasegawa for information and helpful discussion regarding Dendroceros species found in the islands of the South Pacific and of tropical Asia; to the curators of the herbaria at Botany Division, National Museum, British
Key to the genera of Anthocerotae in New Zealand.
From Tuatara 25(1), 1981.