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Tuatara aims to stimulate and widen interest in the natural sciences in New Zealand, by publishing articles which (a), review recent advances of broad interest; or (b), give clear, illustrated, and readily understood keys to the identification of New Zealand plants and animals; or (c), relate New Zealand biological problems to a broader Pacific or Southern Hemisphere context. Authors are asked to explain any special terminology required by their topic. Address for contributions: Editor of Tuatara, c/o. Victoria University of Wellington, Box 196, Wellington, New Zealand. Enquiries about subscriptions or advertising should be sent to: Business Manager of Tuatara, c/o. Victoria University of Wellington, Box 196, Wellington, New Zealand.
is the journal of the Biological Society, Victoria University of Wellington, New Zealand, and is published three times a year. Editor: H. B. Fell. Assistant Editor:
‘The term “plankton”, from the Greek planktos meaning drifting, was proposed by Hensen as a collective term for the small organisms which occur suspended or free-swimming in water and whose powers of movement are so restricted that they drift at the mercy of currents.’ It may seem paradoxical to refer to the ‘terrestrial plankton’ but the phrase ‘dry land’ is a misnomer, not even a desert is wholly dry. All terrestrial habitats retain moisture and the total volume of water held in these environments is probably almost equal to that of the volume of the freshwater lakes and rivers, 0.25 Gg. (Hutchinson, 1957, p. 223). Further, the freshwater lakes and rivers are being constantly replenished chiefly by water that has passed over or through terrestrial environments. This annual precipitation on land surfaces is of the order of a geogram (1020g.) or four times the total volume of all inland waters. The annual evaporation from land surfaces is equal to about two-thirds of this figure and the runoff to the sea is equal to about a fifth (Hutchinson, 1957, pp. 221-230). ‘Dry land’ is, therefore, deeply involved in the hydrological cycle and there is ample water to provide a rich variety of niches for aquatic organisms. The peculiarity of these niches is that the moisture is held in small discrete surface films or pore spaces, greatly restricted in volume, and contrasting sharply with the ocean, lakes, and rivers. Secondly, the currents to which this kind of water is subject are those of wetting and drying, of precipitation, drainage and evaporation. Thirdly, these waters tend to be much richer in mineral and organic nutrients than other natural bodies. Indeed the blooms of freshwater and marine plankton are often associated with the influx of fresh runoff carrying not only the major nutrients,
12 (Hunter et. al., 1956). Fourthly, and perhaps most significantly, there is the close association with higher plants, an organic cycle different to that of most freshwater and marine environments.
In terrestrial environments the variety of the plankton is not as great as that of the marine and freshwater habitats but the structure of the population is very similar. Exceptionally the organic cycle is dependent upon algal photosynthesis (e.g. Antarctica, Flint and Stout, 1960) but generally the vegetation is dominated by the higher plants. Romer has pointed out that the evolution of the higher plants was a necessary prerequisite for the evolution of terrestrial animals and it has also provided a wealth of niches for aquatic microbial life, not only in such bizarre reservoirs as those of the pitcher plant, the bladder wort, or the axils of Astelia but more typically on leaf. stem, and root surface and particularly in the decaying vegetable matter of the forest floor. Here at its richest and in its most typical form may be found the terrestrial plankton.
Algae and phytoflagellates, most commonly unicellular, bacteria, yeasts and protozoa form the nannoplankton. The three latter generally closely associated with fungal mycelium, fine roots, and decaying vegetation. The smaller metazoa include rotifers, nematodes, tardigrades, gastrotrichs. turbellarians, enchytraeids and other aquatic olgichaetes. The Crustacea are represented by copepods, cladocera, and ostracods. There are also insect larvae and minute gastropods. These organisms may attain very great populations. Bacteria may number 109 per ml. of free water, yeasts perhaps 1% of the bacterial total, protozoa several thousands, and so on. Some figures of populations estimated by a dilution technique are given below:—
No attempt has been made to estimate the mass of these organisms but it is clear, from the numbers alone, that they form an appreciable part of the terrestrial population.
The predominance of higher plants has displaced the importance of the algae in the food chain of the terrestrial plankton. Few of the zooplankton are obligate feeders on diatoms or other algae. The majority depend on bacteria, yeasts, or perhaps fungal mycelium. Some are phytophagous and others are micropredators. Ingestion of plant remains is rare among the smallest plankton although some of the nematodes and the larger animals, such as the copepods and aquatic oligochaetes, pass large quantities of plant detritus through their gut. However the greater part of
feeding on the plant debris but on its associated fungi and other micro-organisms. Because the restricted distribution of the free water in which the plankton live prohibits the use of such devices as filter feeding terrestrial predators are driven to the only alternative — a complete ingestion of the plankton and its substrate and the separation, by digestive processes, of the edible plankton from its inedible plant substrate. This may well explain the enormous activity of soil animals in plant litter and the failure in most cases to show the presence of significant amounts of cellulolytic enzymes. On the other hand the constant comminution of the dead plant material passing through the gut of soil animals renders it more accessible to microbial attack, and the animal droppings become active centres of microbial proliferation. The relationship may be seen therefore as one of symbiosis. Where larger animals tend to be absent, such as in peats, the process of degradation is slowed down and incompletely decomposed organic material tends to accumulate.
Some figures are available of the activity of the fauna both in the physical comminuation of plant litter and of its chemical degradation. Litter animals such as oribatid mites or amphipods may consume between 25 and 40% of their own weight of litter per day. Of this only about 20% is digested. In the case of mites it is estimated (Engelmann, 1961) that of the digested material by far the greater part is metabolised in the respiration of the animals but Clarke, in his study on the amphipods in a New South Wales podocarp-broadleaf forest, estimated that only about a sixth of the digested material was respired the rest being lost as dead animal tissue in the non-predatory mortality of the amphipod population. Such dead animal tissue, however, represents a far more readily accessible source of nutrients for other soil organisms than the plant tissue and one suspects that it would be rapidly metabolised by other organisms.
Physical comminution by soil animals is illustrated by Nef (1957). ‘If a pine needle 60 mm. long. 1 mm. broad and 0.5 mm. deep
2 is fed to an earthworm which reduces it to fragments of 1 mm. diameter the surface area will be increased to 240 mm.2 but if it is attacked by mites which reduce it to fragments of 1003BC diameter the surface area becomes 1.8 m.2 or 10,000 times the original.’
In Figure 1 an attempt is made to present the relationships of the terrestrial plankton to the larger animals in soil and particularly forest litter. The principal sources of available nutrients are:— (I) soluble nutrients, such as carbohydrates, amino-acids, and soluble proteins initially derived from leaf drip, dead leaves and twigs; and (II) insoluble nutrients, such as cellulose and lignin. The insoluble nutrients are rendered accessible to soil animals only by bacterial and fungal action and they become accessible in one of two ways, either in the form of microbial cell tissue or in solution as autolysed tissue. In the first case they may be eaten either by the microfauna or the macrofauna and in either case contribute to increase of animal tissue and respiratory activity. In the second case they contribute to the sum of soluble nutrients available in the system.
Similarly animals feeding on the microflora suffer either predatory or non-predatory mortality. In the first case their tissues are used in respiration or synthesis by their predators and in the second soluble nutrients may be released by autolysis or they may become centres of microbial proliferation. The relationships are complex but the pattern is uniform. The sum of synthesis and respiration, predatory and non-predatory mortality tends to release at least part of the available energy of the system in the form of soluble nutrients and these soluble nutrients augment the initial source from leaf drip; from frass, honey dew or other products of phytophagous arthropods; and from dead plant litter. It is this pool of soluble nutrients which constitutes the main substrate of microbial proliferation, supports the terrestrial plankton, and which directly or indirectly provides the basic nutrients of the great majority of soil animals. It also seems likely that it is the concentration of these nutrients in the soil profile which determines the rate of respiration rather than the size of population. Measurements on beech leaves have shown the highest rates tend to be in the more freshly fallen leaves — provided they are wet. Here the populations are lowest but it is likely that soluble nutrients, not yet leached away, are at their highest concentration. In beech leaves, which are not readily eaten by soil animals, the greater part of loss of weight of the litter can be accounted for by respiratory metabolism in situ, at least in the early stages of decomposition. It seems very likely that this reflects the predominant role of microbial growth and metabolism, which may well provide the immediate substrate of the associated animal population.
All planktonic organisms are minute but the terrestrial representatives are remarkable even within this range for their relatively small size. Thus the most common of terrestrial copepods are harpacticids, cyclopoids are rare, and the most common of the harpacticid copepods, Epactophanes richardi is only about 0.5 mm. in length. Similarly terrestrial ostracods are relatively small, the New Zealand species, Mesocypris audax, Chapman, 1961, being about 1.1 mm. in length. Of the aquatic oligochaetes the representatives of the Aeolosomatidae occurring in terrestrial habitats are all only about 1 mm. in length, the Naidid worms are larger, up to several mm. long. Phreodrilids and enchytraeids are larger again but in this case the worms possess a thicker body wall, a morphological character shared by the earthworms, which makes them less susceptible to variation in hydrostatic pressure.
The terrestrial nannoplankton, such as the protozoa, is similarly remarkable for its small size. Thus terrestrial species are almost invariably smaller than freshwater or marine representatives of the same genus and where more than one species is present it is typically the smallest which is the most common. Thus while a catch of terrestrial plankton has many similarities to a catch of fresh water or marine plankton the most striking difference is that it will need to be studied under a higher magnification and it is reasonable to associate this distinction with the confined character of their aquatic habitat which restricts moisture to thin films and narrow pore spaces.
A second feature of the terrestrial plankton is the relative simplicity of form and life history. Most groups are capable of asexual reproduction and the life cycle tends to be short and direct. Again it seems reasonable to associate these characters with the necessity of accommodating an aquatic mode of life to an environment capable of frequent desiccation. The formation of cysts or, in the case of copepods, eggs capable of enduring periods of drought is also typical.
Apart from the nannoplankton — the bacteria, the yeasts, and protozoa — little is known of the physiology of the terrestrial plankton. Many of the larger forms, such as the copepods, appear to be sensitive to high carbon dioxide concentrations or low oxygen tensions such as are associated with rapid decomposition under conditions of poor aeration. Most seem tolerant of acid pH, being common in sphagnum and forest litter with a pH below 5.0 and sometimes below 4.0. However from the few observations that have been made there is a suggestion that mull forest soils, such as those under puriri, may have a different copepod fauna to mor forest soils, such as those under beech, and this may imply physiological and ecological distinctions at the species or genus level. Little is known of the rate of reproduction or of any
It is this aspect, the successful colonisation of ‘dry land’ that has aroused speculative interest in the soil fauna generally (Ghilarov, 1956). How have animals evolved from strictly aquatic to strictly non-aquatic organisms? The fauna of the terrestrial environment, part rock, part water, part air, offers the most suggestive evidence on this problem. Three evolutionary pathways have been suggested. One, proposed by Hurley (1959) as the evolutionary pathway of terrestrial amphipods, suggests that a supralittoral fauna penetrated directly into the forest floor. A second possibility is the slow evolutionary adaptation of a fresh water pond fauna to increasing periods of desiccation until finally a true soil fauna enjoying brief periods of aquatic life is attained. The Tubificidate have been suggested as an example of this process. A third possibility is the evolution of a fauna originating in fresh water streams, extending through moist mossy banks and sphagnum bogs to the forest floor and so to mineral soil. This seems in the majority of cases to be the most convincing explanation. Thus with the naidid worms the closest relatives of one of the New Zealand species were recorded from shallow streams and adjoining moss carpets. The copepods and ostracods tend to occur in sphagnum as well as in forest litter (Harding, 1953, 1955), and this seems also the most convincing pattern of protozoan evolution. On the other hand the evolutionary trend is not solely towards greater emancipation from a permanently aquatic environment. The reverse trend is also evident. The occurrence of earthworms and pulmonate gastropods in fresh water lakes, and perhaps more dramatically the suggested evolution of hydracarines from terrestrial mites are cases in point.
The mobility of the fauna varies greatly. Some, such as protozoa, although very strictly aquatic are able to exploit the smallest and most transitory pockets of moisture; others, such as enchytraeids and ostracods, although requiring a high relative humidity, are able to move freely within such an atmosphere. These latter have close affinities both taxonomically and ecologically with earthworms and terrestrial amphipods and isopods. Their respiration is dependent upon relatively large moist exposed surfaces and they lack the cuticular waxes which are of such great value to the insects in their emancipation from water to air (Beament, 1961). Consequently they are susceptible to excessive transpiration losses. Such animals are always happier in a wholly aquatic medium than
Still the best and most readily accessible account of the forest fauna is that of Birch and Clark (1953). A more recent work is the new English translation by Norman Walker of Kuhnelt's Soil Biology (Faber). A number of papers have been published in recent years on the microfauna of New Zealand soils and these are also listed below.
Tuatara has over 700 subscribers, including 176 libraries. These, coupled with casual sales, now require a regular edition of of 1100 copies. The journal is distributed to Great Britain, United States of America, Union of Soviet Socialist Republics, Australia, Sweden, Belgium, Sudan, Fiji and Malta.
The increased size of the current issue (the largest yet) reflects the present financial situation resulting from the increased demand. No subsidy has been needed since the close of volume eight, the journal being now able to support itself from revenue. Provided certain technical problems are overcome, it is anticipated that one colour plate will be included in each annual volume. No change in the subscription rate is likely in the immediate future.
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‘To the memory of Daniel Carl Solander, F.R.S.
1733-1782’
So runs the dedication to Allan's 1961 Flora of New Zealand. But why Solander? Why not Forster, Manual (p.v.) ‘Every botanist who prepares a Flora starts from the standpoint reached by his predecessors in the same field’.
Solander had no predecessors — his was a virgin field. He had it is true the Species Plantarum at his hand. He had the advantage of personal tuition by Linnaeus. He had wide experience of botanical matters in Europe and in England, and he had, during 1769, botanised briefly in South America and Tierra del Fuego, and extensively in Tahiti. He was in fact a first class professional botanist. He had been offered the Chair of Botany in the Petersburg Academy of Sciences, was a fellow of the Royal Society and on the staff of the British Museum. He had also two very able colleagues — Banks, himself no mean botanist, with his vigorous intelligence, his youthful enthusiasm and his very necessary money — and Parkinson, with his technical skill and devotion. And in the background the master mariner Cook, who had brought them there and would take them home. The presence of Tupaia too was providential. To be able to discuss the native names and uses of the plants collected, with the people who had lived among them for centuries was invaluable. Solander's Pohutukawa and Kowhai — his modus praeparandi, are echoed unaltered 200 years later in the Floras of today. These things were in his favour. But against him was the very limited time ashore and the almost total strangeness of the vegetation he was studying.
This paper is based upon the typed copy of the Solander manuscript in the Auckland Museum. I am indebted to Dr.
Dendrobium cunninghamii Lindl. (Epidendrum pendulum Sol. ined.) Mscr. p. 1349, typescript p. 167. Engraved from an original unpublished drawing by
Banks said of it (Beaglehole 2 : 1962, p. 9) ‘Sow thistle, garden nightshade, and perhaps one or two kinds of grasses were exactly the same as in England, three or four kinds of fern the same as those of the West Indies, and a plant or two that are common to almost all the world; these were all that had before been described by any botanist out of about 400 species, except five or six which we ourselves had before seen in Terra del Fuego’.
The Primitiae Florae is of necessity a coastal Flora and the shadow of the Endeavour hangs over it. Much of the botanising was done during wooding and watering, fishing, shooting and surveying trips, and was subject to the exigencies of service. The Maori comes into it too. His clothes and cultivations, his ornaments and his children, and many of his plant names. The few errors are more interesting than otherwise — one in particular, Avicennia resinijera. Cook (Beaglehole 1 : 1955, p. 204) says —‘ … in speaking of Mercury Bay I forgot to mention that the Mangrove trees found there produce a resinous substance … we found it at first in small lumps upon the sea beach, but afterwards found it sticking to the Mangrove trees and by that means found out from whence it came.’ The resin was the familiar Kauri gum and it was not produced by the Mangrove. The name though misleading, was perpetuated by Geo. Forster and having priority has stuck — like the gum it commemorates.
This first Flora was a major event in the botanical history of New Zealand, although it was never published. Its importance is emphasised by the number of Solander's names which were taken up by later botanists and his skill by the fact that 123 plants still belong in the genera in which he originally placed them. The near misses are revelant too. Smilax, Pandanus, Piper, Fagus, Panax, Aralia, Veronica, Myrtus, Mesembryanthemum, Passiflora, etc., are but further proof of his knowledge.
To read Solander for the first time is to experience a feeling of familiarity — one has trodden this path before. In spite of the Latin and the odd quirks of the Linnean system, this collibus et campis is home sweet home. His descriptions have a modern ring about them and his locality-habitat notes a photographic clarity.
For example :—
Avicennia resinifera — ad latera limosa fluviorum lacuum solsorum. Disphyma australe — copiose juxta littora marina praecipue, in fissuris rupium.
Lycopodium billardieri — in sylvis … in arboribus parasitica dependens.
Entelea arborescens — ad latera vallium et ubi sylvae incipiunt. and of Leptospermum simply — ubique et copiose.
Odd words stand out here and there — Paesia scaberula — elegantissima and Phormium tenax — utilitissima, which tell more than pages of prose.
That the work had so profound an influence on subsequent Floras of New Zealand was due partly to Banks' place in the scientific world and to his generosity in allowing the collections to be studied by those interested, and partly to his choice of secretaries. First Solander, then Dryander and finally Students' Flora of 1899 and were handed over to Cheeseman who used them while preparing the 1906 Manual. They are still preserved in the Auckland Museum and are the basis of these notes. Cheeseman says ‘of their scientific value I cannot speak too highly’. I believe there is a photostat copy of the original manuscript and another set of drawings in the Dominion Museum and in the Turnbull Library. The whole of the ‘Banks and Solander’ material was studied by Flora. All these botanists quote Solander freely and with respect.
205 Fig. Pict. are listed and there exist several others. One which interests me is an unfinished but recognisable sketch of Earina autumnalis which is not otherwise mentioned in the typescript. Another plant left out by Solander is the Taro (Colocasia), which is however recorded several times in Banks' journal as a food plant cultivated by the Maori. The plants he does list as cultivated are :—
Sonchus oleraceus — in graminosis et cultis.
Broussonetia papyrifera — culta in septentrionali parte Nov. Zel. sed rara.
Lagenaria vulgaris — forte culta utensilaria varia e hupis fructo formant incolae.
Discorea sativa — culta. Ipornoea batata — culta.
Solander's Maori names are sometimes amazingly accurate. His Kawakawa, Kowhai, Mahoe, Manawa, Manuka, Ngaio, Piripiri, Pohutukawa, Poroporo, Tawa, Ti, Tutu, appear as they do today. Several others are recognisable though mis-spelt by modern standards — Karaka (Chalacha), Karamu (Charamugh), Kiekie (Geagea), Kumara (Kumala), Ramarama (Lamalama),
Rangiora (Rangiola), Mangeo (Tangeo), and Kowhai ngutu kaka (Kowhai no tugaga). There are as many again that are meaningless to me. One name is worth comment. He uses Kowhai for Sophora tetraptera, and Kowhai maori for S. microphylla. The word ‘maori’ I believe, originally meant normal or ordinary. The Kowhai maori was then the common or garden variety of Kowhai found along the coast as distinct from the larger S. tetraptera which is less frequent.
There are 349 names in the typescript. Of these 15 were not identified by Cheeseman — 12 ferns and 3 Cyperaceae. The remainder fall conveniently into four groups :—
(i) In the current Floras (Cheeseman, 1925, for the Monocotyledones, Allan, 1961. for the remainder) 49 names read exactly as Solander wrote them. Of these specific names 45 originated with Solander, 4 with Linnaeus. Of the 35 genera, 28 are Linnean and 7 by Solander. The 7 are Astelia, Dacrydium, Metrosideros, Nertera, Pimelea and Pittosporum. Familiar plants in this group are — Arundo conspicua, Asplenium lucidum, Astelia nervosa, Avicennia resinifera, Clianthus puniceus, Dacrydium cupressinum, Linum monogynum, Metrosideros excelsa, Nertera depressa, Pimelea longifolia, Pittosporum crassifolium, P. tenuifolium, Pteris tremula, Ranunculus hirtus, Rubus australis, Salicornia australis, Scirpus frondosus, Senecio lautus, Sophora microphylla, S. tetraptera, Tetragonia trigyna, Trichomanes reniforme and Weinmannia sylvicola.
(ii) 40 plants bear the specific or varietal name given by Solander or derived therefrom, but the genus has been changed. Included are — Alectryon excelsum, Ascarina lucida, Blechnum discolor, Bulbophyllum pygmaeum, Carpodetus serratus, Celmisia gracilenta, Coprosma acerosa, Cordyline australis, Disphyma australe, Dracophyllum longifolium, Earina mucronata, Elatostema rugosa, Griselinia lucida, Haloragis erecta, H. procumbens, Hebe macrocarpa, H. pubescens, Hymenophyllum dilatatum, H. sanguinolentum, Litsaea calicaris, Lophomyrtus bullata, Luzula campestris, Neopanax arboreum, Nothofagus fusca, Pellaea rotundifolia, Phormium tenax, Pittosporum umbellatum, Pseudopanax crassifolium, Rhopalostylis sapida, Sarcochilus adversus, Schoenus tendo, and Tetrapathaea tetrandra.
(iii) 74 plants were placed by Solander in the genera to which they still belong, although the specific names have changed. 45 genera, 41 of them Linnean, 4 by Solander. They include species in genera like — Adiantum, Arundo, Asplenium, Astelia, Carex, Clematis, Coriaria, Drosera, Elaeocarpus, Epilobium, Gaultheria, Gnaphalium, Hydrocotyle, Juncus, Lepidium, Lobelia, Lycopodium, Myoporum, Myosotis, Plantago, Samolus, Scirpus, Senecio, Solanum, Tillaea, Trichomanes and Urtica.
(iv) A further 186 plants were described but the names were never taken up. The group includes Acaena, Aciphylla, Aristotelia, Arthropodium, Beilschmedia, Brachyglottis, Carmichaelia, Corynocarpus, Cyathea, Dendrobium, Discaria, Dysoxylum, Entelea, Freycinetia, Fuchsia, Geniostoma, Hedycarya, Hoheria, Ipomoea, Knightia, Lagenaria, Leptospermum, Macropiper, Melicope, Melicytus, Microtis, Myrsine, Olearia, Orthoceras, Paesia, Plagianthus, Podocarpus, Pomaderris, Pterostylis, Rhabdothamnus, Rhipogonum, Schefflera, Thelymitra, Uncinia, Vitex, Wahlenbergia and many more besides.
Two names show transposition — Senecio perdicioides was written Perdicium senecioides by Solander, and Lagenaria vulgaris was called Cucurbita lagenaria.
Cheeseman considered 6 species to be duplicated —
Calystegia tuguriorum (Convolvulus lacteus — C. versatilis).
Carex forsteri (C. debilis — C. latifolia).
Geranium pilosum (G. pilosum — G. patulum).
Gnaphalium involucratum (G. involucratum — G. collinum).
Penantia corymbosa (Meristoides paniculata — Fagoides triloba).
Podocarpus dacrydioides (Dacrydium thujoides — Lycopodium arboreum).
A number of Parkinson's drawings have been published from time to time. I am aware of eight books and papers (there may be others) containing them. Twenty-nine plants are illustrated.
The 349 species collected and described by Solander in New Zealand are contained in 206 genera and (counting the Filicopsida as a single family) 86 families. Of the genera 130 belong to the Dicotyledones, 48 to the Monocotyledones, 25 to the Filicopsida, 2 to the Coniferae and 1 to the Lycopodiaceae.
When it is remembered that manuscript Floras were prepared also for Tierra del Fuego, Tahiti and the eastern coast of Australia, and that each of them was as coherent as the New Zealand section, and that he wrote in addition on the marine and bird life encountered on the voyage, the genius of the man shines forth, and we begin to understand why the present Flora of New Zealand is dedicated to the Swedish Doctor from the University of Uppsala. Well might Linnaeus write of the ‘immortal Banks and Solander’.
The ticks are the largest of all Acarina. They may be distinguished by the presence of a movable capitulum, distinct from the fused thorax and abdomen and visible dorsally in most genera, and the prominent barbed hypostome. Unfed ticks in all stages are dorso-ventrally flattened but the bodies of engorged females may be spherical and up to 10 mm. in diameter, when the sclerotized parts which provide the main taxonomic characters become relatively inconspicuous. The basis capituli may bear cornuae postero-laterally or auriculae ventrally and in the females has two dorsal porose areas. The mouth parts are borne in the median line anteriorly and consist of a dorsal cheliceral sheath and a ventral toothed hypostome which enclose the chelicerae and are apposed to form a sucking tube. This structure is flanked on each side by a labial palp which is four-segmented, though the fourth segment is small and often not visible dorsally. The scutum is a flat shield-shaped plate on the dorsum of the thorax immediately behind the capitulum. The legs are six-segmented and bear a sensory organ (Haller's organ) on the distal tarsal segment of the first pair of legs. The spiracular plates are sub-circular and are situated ventro-laterally behind the fourth coxae. The integument of the remainder of the body, except in the males, has a thin unsclerotized integument which is capable of great extension during engorgement. The eyes, when present, are situated on the sides of the scutum but they are not present in New Zealand species.
The females are oviparous and the development of individuals of both sexes includes a larval and a nymphal stage. The larvae have only three pairs of legs and the nymphs lack the genital opening of the males and females. The males and females of Ornithodoros differ only in the form of the genital opening, but in the Ixodidae the males have the scutum covering the dorsum and more numerous ventral plates, while the females have porose areas on the dorsum of the basis capituli.
All stages except the males are blood-sucking. The skin of the host is pierced by the cutting chelicerae and the recurved teeth of the hypostome, which is inserted into the wound, act as a holdfast. The argasid ticks such as Ornithodoros feed quickly and leave the host at once, and all stages are commonly found in the nest, where they shelter, and not on the host. All stages
This habit is of significance in the transmission of the diseases of man and animals, especially in tropical countries, which are caused by the pathogenic viruses, rickettsiae, and protozoans which the ticks may carry. In some cases the pathogenic agent is transmitted to the progeny of the infected female tick through the eggs, and ticks which have never fed on an infective host may thus carry infection. Apart from their role as vectors of disease the ticks inject salivary secretions when feeding and in some species these have a neurotoxic constituent which causes ‘tick paralysis’ in man and animals, especially when the feeding site is near the brain or spinal cord.
The eggs, which are spherical and about half a millimeter in diameter, are extruded in large masses containing as many as 1000 eggs. They are found in the nest of the host, or on the ground where the female has dropped from the host. Host finding is easy for the larval ticks hatching from eggs laid in a nest, burrow, or lair which is constantly used or periodically reoccupied. The tick species infesting free-ranging mammals may drop from the host anywhere, though they would tend to be concentrated in favoured camping places. The larvae or newly moulted individuals of such species ascend grasses or other vegetation and wait for the passing hosts to brush against them.
Ticks have been known to survive unfed for very long periods but there is no doubt that a large part of the mortality amongst larvae and nymphs awaiting a host is due to adverse climatic factors. One small hymenopterous (chalcid) parasite of ticks is known elsewhere, but invertebrate predators may take a greater toll of ticks on the ground. Birds such as the starling may take the ticks not only from the ground but from the host also. Two unusual cases are known of a sea-bird tick taken from the faeces of a tuatara and a kiwi tick from the faeces of a cat. On the hosts the ticks aggregate in positions where they are not readily dislodged by self-cleaning or scratching.
Because of their medical and veterinary importance considerable work has been done on the physiology, ecology, and control, of ticks. In New Zealand the cattle tick (Haemaphysalis) is the only species which has been studied (Myers, 1924).
Recent taxonomic work on New Zealand ticks is accessible in previous papers (Dumbleton 1943, 1953, 1958, 1961), and detailed host lists are given in two of these (1953, 1961), A generalised host list is combined with the list of the species in the New
Haemaphysalis leachi Audouin, Hyalomma aegyptium Linnaeus, and Ixodes ricinus Linnaeus, have been recorded as present in New Zealand. They were considered (1953) as doubtfully established and are now omitted from the list as they have not been collected since that date.
Little can be said regarding the origin of the tick fauna. The single species of Aponomma, a genus largely restricted to reptiles, would appear to have been contemporaneous with Sphenodon.
The same may be true of Ixodes anatis Chilton the only New Zealand species confined to land birds, more especially the kiwi. This appears to belong to a group differing from that of the other New Zealand species. While there are few or no Australian species confined to land birds I. anatis has some affinities with the species of Ixodes (Sternalixodes) which occur there on land animals.
The species occurring on sea birds are either cosmopolitan or belong to species-groups which are widespread in the Subantarctic or in Australia.
Keys to the families, genera, and the males and females of the genus Ixodes are given below. Nymphs, and usually larvae, are identifiable by association with adults and similarities in morphology.
Scutum absent in all stages; capitulum ventral, visible dorsally in larva but not in nymphs and adults; integument mamillated, with symetrical pattern of smooth plaques on dorsal surface; sexes differing only in form of genital orifice (figs. 1 and 2) . . Argasidae.
Scutum always present, covering all (males) or not more than anterior half (females, nymphs, larvae) of dorsum; capitulum anterior, visible dorsally and with prominent basis capituli; integument smooth; females with porose areas on basis capituli, males with median and ad-anal plates (figs 3, 4, 5 and 6). Ixodidae.
Without a marginal sutural line, separating the dorsal and ventral surfaces and differing in sculpture from both …. Ornithodoros.
(One species. Body distinctly conical anteriorly; hood and cheeks, enclosing capitulum, distinctly separated [figs. 1 and 2] O. capensis).
Ornithodoros capensis, dorsal. Fig. 2: Ornithodoros capensis, ventral; capitulum, hood (h), cheeks (c), and genital opening (go). Fig. 3: Ixodid male, dorsal; scutum (s). Fig. 4: Ixodid female, dorsal. Fig. 5: Ixodid male, ventral; coxae (ex), spiracle (sp). Fig. 6: Ixodid female, ventral; anus (a), anal groove (ag). Fig. 7: Aponomma sphenodonti ventral; festoons (f). Fig. 8: Haemaphysalis bispinosa, female capitulum, ventral (L) and dorsal (R); chelicera (ch), hypostome (hy), sheath (sh), palpal segments (2, 3). Fig. 9: Aponomma sphenodonti, female capitulum. Fig. 10: Ixodes uriae male, posterior margin, ventral. Fig. 11: Ixodes anatis male, genital opening, pre-genital plate (pg), coxal spur (cs). Fig. 12: Ixodes auritulus zealandicus male; pre-genital and jugal (j) plates. Fig. 13: Ixodid male, ventral; diverging anal groove. Fig. 14: Ixodid male, ventral; converging anal groove. Fig. 15: Ixodes uriae, female capitulum. Fig. 16: Ixodes anatis, female capitulum. Fig. 17: Ixodes auritulus zealandicus female capitulum; auricua (au), cornua (co), palpal spur (ps). Fig. 18: Ixodes eudyptidis, female capitulum; porose area (po).
Little is known about the root habit of many New Zealand plants. One of the few published works is a paper by McIndoe (1932).
One feature of at least some species of forest tree is their ability to form root grafts. This phenomenon seems to be common in the Nothofagus species and is probably found in others. There are several interesting ecological implications arising from root grafting as it has been shown overseas (see Fraser and Gaertner, 1961), that materials are transmitted from individual to individual through the root systems. Weak plants may survive in a dense stand in this way and this raises the question of the nature of competition in such a situation.
A set of observations on a peculiarity shown by some New Zealand gymnosperms was made by Foweraker (1929) and bears repeating. Foweraker found that Podocarpus totara could survive burial by quite deep deposits of river silt by sending out adventitious roots near the new ground level. Photographs show trees with as many as three series of roots girdling the stem and each separated by several feet. This ability could be important to species inhabiting the flood plains of rivers. Christensen (1923) showed that a number of other New Zealand trees and shrubs survived burial by riverbed gravel in the same way, while others succumbed.
Some of the most aggressive colonisers of bare ground in the mountains are able to produce adventitious roots from stems buried by silt or gravel, or shoots from erosion-exposed parts which are apparently root tissue in nature. These include Muehlenbeckia axillaris, Raoulia spp., Gaultheria rupestris and Cyathodes fraseri. This capacity is shared with the aggressive adventive plant Rumex acetosella.
It has been noted that some New Zealand trees and larger shrubs produce adventitious shoots readily from root systems which lie near the ground surface. Moar (1955) recorded this for Dacrydium colensoi and the author has seen it in species such as Hoheria glabrata. Griselinia littoralis, Dacrydium bidwillii and Olearia ilicifolia.
Amongst the alpine vegetation in New Zealand there are many shrubs, many woody small plants which could be called shrublets and numbers of semi-woody species. There are, in fact, very few soft-stemmed herbs proper. Most of the herb-like species are partially woody or fully woody perennials which creep along at ground level, sending down adventitious roots. This applies, of course, to dicotyledonous plants such as Celmisia discolor, C. lyallii, C. laricifolia, Forstera tenella, Gaultheria depressa, Drapetes dieffenbachii, Coprosma pumila, Ourisia sessilifolia and Geum uniflorum, but even the long-lived monocotyledons like the snowgrasses (Chionochloa spp.) and the Astelias have tough, fibrous, wood-like tissue in their main stems and have the usual monocotyledon capacity for sending out adventitious roots along their leaders. The carpet grass (C. australis) is an extreme example of this. Most of the subalpine scrub species also have the capacity for production of adventitious roots on branches lying along the ground. Gaultheria rupestris, Hebe macrantha, Coprosma serrulata and Podocarpus nivalis are some of these and Wardle (1960) recorded others in a description of downhill layering in subalpine scrub. Mr B. Fineran (pers. comm.) has similarly noted this characteristic in shrub species on the Snares Islands.
On a recent expedition to Western Fiordland it was noticed that under the thick growth of mosses and liverworts on the trunks of Nothofagus menziesii trees, as much a 8-10 feet above ground level, there were numerous adventitious roots. An investigation of other species in the same neighbourhood—Griselinia littoralis, Coprosma linariifolia, C. ciliata. C. astonii, Neopanax anomalum and Myrsine divaricata — showed that all the species examined exhibited the same characteristics. Presumably the plants extract moisture and nutrients from the humus which is present under the moss cushion. This is not so surprising when it is realised that most of the soils of Western Fiordland are peats so that the substrate in the moss cushion is similar to that on the ground. There was no evidence for this phenomenon in the same species in eastern Fiordland. It seems that very high rainfall is necessary for it. On the Snares Islands it was found that Olearia lyallii sometimes produced roots in the humus collected on branches. (B. Fineran, pers. comm,).
The ‘feeding roots’ of forest species both in beech forest and mixed broadleaved podocarp forest in New Zealand are found in the humus layer of the soil. Similarly in the alpine grasslands — the actively growing root systems proliferate in the humus layer
Chionochloa, as in many other species, there are normally masses of old dead and rotting leaf bases which surround the living tillers. ‘Feeding roots’ are found in these rotting tissues which remain wet even in the driest weather. It is probable that both water and nutrient requirements are largely supplied in this way.
Some New Zealand forest species are obligate epiphytes but many other species are able to exist as facultative epiphytes if the atmosphere is moist enough. This phenomenon is at its best in places of very high rainfall such as Western Fiordland or the Upper Hokitika River. In the latter area the forest is dominated by Metrosideros umbellata and Weinmannia racemosa, with an admixture of other broadleaved and gymnospermous species. Almost every tree in the forest has started life as an epiphyte and there are huge ratas, as much as 20 feet in diameter, formed from the coalescence of several trees (natural shoot grafts) and covered with a great luxuriance of filmy ferns, mosses and liverworts. In such situations, at least in the early stages of their life, these plants must obtain nutrients entirely from humus formed in the crotch of their host tree. It is interesting that so many species of shrubs, trees and herbs, in addition to the obligate epiphytes, are adapted to take advantage of the epiphytic habitat if other environmental conditions permit.
The facility with which New Zealand plants produce adventitious roots and the ability to utilise humus or peaty material with little mineral matter in it for much of their nutrient supply is a striking feature of our vegetation. There is scope for much future research on the root systems of New Zealand plants.
Additional Note: Since this paper was written a paper has been published (
Present address: C/o, Canterbury Museum, Christchurch, New Zealand.
As many enquiries are made each season, and as early records are somewhat confusing, it is considered as well to clarify the position concerning food-plants of ‘monarch’ butterfly larvae in New Zealand. In various early records the ‘monarch’, Danaus plexippus Linnaeus, 1758. was recorded under the names archippus, erippus, and berenice.
Fereday (1874b BNZE) recorded that a Mr. Meinertzhagen was told by the Maoris that ‘The caterpillar … feeds upon the pollen of the gourd which they grow in that part of the country (Hawke Bay).’ This record is not reliable and is not accepted. Fereday also recorded caterpillars and pupae found by a Mr. Nairn who ‘… had been feeding some new kind of caterpillar …’ and had described ‘… the shrub on which he found the caterpillar as the Gomphocarpus ovata, one of the milk producing plants, and a native of the Cape of Good Hope.’ It is considered by the present author that this is a valid record of Asclepias fruticosa (= Gomphocarpus fruticosus) as a food-plant of Danaus plexippus. A. fruticosa, the common ‘swan plant’, was recorded as a horticultural escape in New Zealand, under the name of Asclepias nivea, by Kirk (1870), and this species is a native of the Cape of Good Hope.
Asclepias curassavica but, as it is not known if this plant was at that time present in New Zealand, it seems best to consider the record as Asclepias sp. Colenso (1878 BNZE) was alos obviously referring to an Asclepias when he described ‘cotton plants’ and ‘… green capsule having the remains of soft spines …’. Records by Butler (1878 BNZE) and Asclepias sp. was given by Danais archippus became abundant in Wanganui
Gomphocarpus, …’. J. J. Walker (1914 BNZE) mentioned Gomphocarpus fruticosus R.Br. as a food-plant at Sydney (Australia) but did not record it as such in New Zealand. Tillyard (1926b BNZE) recorded two plant species, as ‘… Asclepias physocarpa (Gomphocarpus fruticosus) and A. curassiva.’, but as they were not specifically for New Zealand they are here considered to be Australian records. Gomphocarpus fruticosus,’, while Cottier (1956) gave the first record of Araujia sericofera as a food-plant in this country.
The known food-plants in New Zealand are all of the family Asclepiadaceae, members of which are sometimes known as ‘milkweeds’ or ‘cotton plants’. The species concerned are as follows :-
Asclepias curassavica L. Commonly known as the ‘blood flower’ this plant has also been called ‘red-head cotton’ in Australia. It is not common in New Zealand but is grown in some gardens and is known to be a food-plant here. This is the first undoubted record for this country.
Asclepias fruticosa L. (= Gomphocarpus fruticosus R. Br.), This, the ‘swan plant’, is generally known as a food-plant in New Zealand. In Australia it has been known as ‘bladder cotton’, ‘white cotton’, etc. Asclepias physocarpa Schlect. (= Gomphocarpus fruticosus Sims, not R. Br.) is a species very close to A. fruticosa L. and. although apparently not known in New Zealand (apart from one mention in a commercial catalogue), may well be grown here under the name of ‘swan plant’. It is a potential food-plant.
Araujia sericofera Brot. (= Araujia sericifera auct.), Although ‘monarch’ caterpillars are not known to occur naturally on the ‘moth-catching plant’ they will readily feed on leaves of this plant when supplies of the ‘swan plant’ have been eaten out.
The author wishes to acknowledge the assistance of Dr.
All references included in the ‘Bibliography of New Zealand Entomology’ (Miller, 1956) are referred to that work, in the above text, by the appropriate date and letter, followd by the letters BNZE. e.g. Butler (1878 BNZE), Fereday (1874b BNZE). Other references are given below.
This paper is not only an attempt to reconstruct something of the vegetation pattern but also to draw the attention of botanists and others to the need for archival research required by a topic of this nature. Thanks are due to MissA. Simpson for checking the botanical nomenclature.
The effects of the introduction into New Zealand of ‘European’ plants and animals have been described by a number of specialists in several fields notably Thomson, 1922, and Wodzicki, Others (Cumberland 1941 1961. 1962b; Holloway 1959; Johnston 1961) have attempted to reconstruct the pre-European vegetation on a New Zealand-wide basis. Clark (1949) notably, has shown in some detail, the successive impact of man and his plant and animal domesticates on a land that lacked indigenous grazing mammals. Clark, discussing the initiation of large-scale settlement in the South Island during the 1850's, rightly draws attention to the damage done to the vegetation by cutting, burning and the proliferating sheep. However, it is often not realised that a similar process was initiated in the Wairarapa district, almost a decade before settlement in the South Island began.
This paper is an attempt to describe the vegetation pattern of the Wairarapa on the eve of settlement in 1843, the manner in which the indigenous vegetation was attacked and the way in which new plants and animals were introduced.
The research on which the paper is based was predominantly archival and no attempt will be made to link present-day survivals with past conditions.
The vegetation of the Wairarapa in early European times was characterised by variety. The whole area was a patchwork of grass, swamp, scrub and forest mingled in varying proportions. This is well illustrated by Bidwill's description of the area adjoining Bidwill's Ridge, which is located between Featherston and Martinborough, about two miles from the latter. The land along the Ruamahanga River was in dense bush and fringing the Ridge was a swamp containing Phormium tenax. ‘About a mile to the north were low-lying ridges on which grew manuka (Leptospermum scoparium) and a small variety of flax, interspersed with open
To the west of Lake Wairarapa, the mixed podocarp/broadleaf forest extended down from the Rimutaka Range to reach the lake margin and similar salients of bush extended into the valley at several points, notably in a 20,000 acre block between the Waingawa and Waiohine Rivers. At its northern end the valley was closed off by an area of bush-clad hills and down-land that extended with little break to a clearing in the vicinity of the Manawatu Gorge. Bush then continued as far as the margins of the tussock lands of the Ruataniwha/Takapau basin. There is no reason to suspect that the podocarp/broadleaf forest was in any way unlike that covering much of the remainder of the North Island, although Colenso considered the North Wairarapa forest the most primeval of any he had seen in New Zealand. ‘The soil for many feet was composed of vegetable matter … and the trees were of immense size. The birds were very few … and a death-like silence reigned’ (Colenso, Journal), The last was probably a localised phenomenon since another observer claimed that ‘the woods are alive with kakas and pigeons’ (Weld in Lovat. 1914. p. 50). Colenso also noted some of the forest-dwelling grasses such as Oplismenus undulatifolius, Poa imbecilla, P. anceps, Microlaena spp., herbs such as Craspedia sp., and several species of Cotula, Oxalis and Epilobium (Hooker, 1867, p. 320-327).
In the lowland short tussock,