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The following is entirely preliminary and is aimed at the problem of the origin of the New Zealand flora and vegetation solely for the purpose of drawing attention to different lines of evidence for transoceanic dispersal of plants in order to stimulate more effort to assemble information about this neglected subject. There has been much speculation and some debate among plant geographers and ecologists about the modes of origin of the New Zealand flora and vegetation. From studies of the distribution of the various floras of Pacific regions, particularly the islands, a number of schools of thought regarding the causes of the present distribution have developed. One of these schools has emphasized the long distance migration of plants across bodies of water by such natural agencies as ocean currents, winds, and some animals, particularly birds. Another school has put emphasis on formerly continuous, or nearly continuous land areas over which plants migrated in the past, or parts of the land areas themselves migrated by drifting apart. This land migration school postulates two main types of land area changes (1) that a series of land bridges formerly existed with narrow water gaps between areas that are now far apart, and (2) that there has been some drifting apart of continental areas or fragments of such areas. The first, or land-bridge theory, was formerly held by more persons than at present. The second, or continental-drift theory, is now more popular and advocated by such plant geographers as Good (1947).
The relative merits of these three possible explanations of the present distribution of plants can not be fully discussed in this short paper but some of relative values of evidences for these will be briefly considered. A number of writers, Cain (1944), Oliver (1925), Gordon (1949) and Fleming (1950) have discussed these three theories. In general, the exponents of either the land-bridges or the continentaldrift theories have heavily discounted the effectiveness of dispersal of various elements of the floras across oceans and seas. They have, as a rule, used the patterns of the present and some of the past distribution of species, genera, families and other categories of plants to show how former land areas were arranged or of different size. This mode of postulating former land areas has much less geologic evidence to support it than is desirable and many conclusions based on such hypotheses are open to question until the evidences from geophysical investigations, paleobotany, and paleogeography more conclusively support these theories. Some fragile constructions of logic have resulted from attempts using distribution to prove land bridges and continental drift without more direct evidences.
It is probable that the most satisfactory explanations of present plant distribution will eventually involve both land area changes and trans-oceanic dispersal, and until a better consistency of theories is evolved from the slowly accumulating evidences for each it is very appropriate to examine some of the sources of evidences of transoceanic dispersal. New Zealand is a group of islands particularly well suited by the nature of its flora and its geographical position to studies of the dispersal of plants, their establishment in new localities, and their incorporation into different types of vegetation. The ecology of vegetation, plant communities, needs to be considered in the evaluation of modes of origin of the floras.
The normal circumstances involved when a particular plant species expands its distribution into new areas not previously or recently occupied by it are usually very definitely associated with the nature of the vegetation this species in invading. There is usually much competition unless the vegetation is of the open type of a primary succession. In general, a region with well established climax or relatively fixed types of successional plant communities tends to keep its flora fixed or consistent as regards the abundant species. Therefore the migration of plants is not the end of the process of establishment and a whole complex of factors concerned with successful establishment (Weaver and Clements, 1938) is involved.
Long geologic periods may provide for species, genera, families and even larger category changes in the flora on a given area. However, for rapid changes of the flora numerous additions from outside the existing vegetational complex are most effective. This has been the case through the agency of man in modern times, particularly in New Zealand (Clark, 1949). During the long geologic past additions from the outside have had a great cumulative effect on many floras and vegetational complexes even though the rate and consistency of long distance dispersal may have been slow and hazardous.
Activities of many animals, birds particularly as regards New Zealand because mammals were almost non-existent, help disperse plants. Ridley (1930) in his monograph on the dispersal of plants, Darwin (1845 and 1859) and Guppy (1906 and 1917) considered these agents and gave many examples of plants dispersed by them, showing in numerous cases a very high degree of efficiency with which some plants are dispersed. Much, but not perhaps enough evidence has been accumulated to show how many plants are dispersed by them. Oliver (1925) states the problem thus as regards New Zealand and South America: “The conclusion seems to be inevitable that plants have been carried from New Zealand to South America by agencies comparable
Among the plants many parts are dispersed, such as spore, seed, fruit or vegetative part, or even the whole plant. The edible, fleshy fruits are often bird-carried, the pappus fruited species of the Compositae and the small spores of ferns are often wind-carried, and the floating fruits of some shore plants, and the floating seedlings of some mangrove swamp plants are transported by water. Hundreds of special adaptations of plants that aid their dispersal are considered by Ridley (1930) and others, and many of these adaptations indicate that no doubt they have depended on certain agents to increase their range of distribution.
The reliable factual data of long distance dispersion are very few because any complete observation of disseminules migrating long distances requires elaborate observations. Davis (1940) experimentally showed how mangrove, Rhizophora mangle, seedlings were efficiently aquivectant for nearly a hundred miles, and Ridley gives some instances of similarly accurate observations. Future research to establish direction, rate and distance of dispersal should be done and probably can be done with more modern techniques such as the airplane, better observations of bird migrations, and more data about hurricanes and other violent winds.
In spite of this lack of direct evidence, however, there are many indirect evidences among which are (1) the observed establishment of floras, and some inferred establishment, on new land areas, especially new islands; (2) the invasion of species into a vegetational complex, which species were not observed previously; and (3) related geological and paleobotanical evidences, some of which may be used to postulate certain floristic changes on islands.
There are also some usually neglected ecological evidences which can be considered. Among these are (1) an abundance of epiphytes in the vegetation; and (2) the occurrence of the same species or similar species near or on the shore areas and far inland on the high mountains with few or no such species in areas between these localities. These two will be stressed in this brief discussion because they apply particularly well to New Zealand.
The volcanic islands Krakatau, near Java, and Rangitoto, near Auckland, New Zealand, and some of the islands of the Florida Keys west of Key West (Davis, 1942) furnish examples of the establishment of floras on new islands. These islands are not far way from land masses and there are few if any good records from islands distant from
In relation to the long spans of time man has had very little time to accumulate information about migrations of plants. Doubtless many records of flora changes lie buried in accounts of explorations and other documents. Exact information will accumulate slowly and for the present much reliance must be placed on indirect evidence. Later experiments may be undertaken to measure rates and distances of dispersal, especially by winds and water currents.
Many of the common plant successions are the results of the successful invasion of new species into the sere communities. The primary successions on coastal and other physiographically young areas follow a course of development that is frequently related to the relative ability of the new species to migrate. At times bare mountain areas have received plants from a great distance by wind, and coastal areas often receive plants that came in by water or wind currents. Secondary successions, or successions after fire or other devastation, are partly the results of new plants invading the areas concerned. Some seeds, or other reproductive parts, of species of the former flora may remain after fire but in many cases the fire followers are species that may have migrated a great distance.
These are but a few of the examples of the ability of plants to migrate into non-climax vegetation. On islands such successional and sub-climax communities are often in greater proportion to the total vegetation than on continents. This seems to be particularly the case in New Zealand where so many of the lowland, non-Nothofagus types of forest vegetation are in a state of flux and very few true climax communities exist.
The invasion of a climax vegetation by new species is much more difficult to accomplish because, in general, the survival and establishment of the new species are relatively less than in successional communities due to the facts that most space is occupied and the species of the climax are so well established that few new species can displace them.
Only a very sketchy outline can be made of this indirect line of evidence because little is known of the past floras of New Zealand and almost nothing of the past vegetation. Most of the interpretations are speculations and inferred from possible analogies with other areas
The affinities of many genera and families lead to the assumption that some plants migrated from Malaya, some from Australia, some from other Pacific island regions, and possibly some from the Antarctic areas, these latter being assumed to have been larger and warmer than at present.
Perhaps early in the Cretaceous the genus Nothofagus was established and some of the climax beech forests developed. These forests probably have varied in size and composition since then but there is little doubt that the main elements, species of Nothofagus, have persisted and because of the climax nature of the community few species of modern floras have invaded these forests.
The other dicotyledonous forest trees and herbs and grasses now constituting the flora of the mixed Podocarp and Dicotylous forests, and the various grasslands or scrub vegetation probably did not reach their maximum number until mid-Tertiary times and maybe now are increasing in number by both additions from the outside and the development of new species on these islands, endemism and hybridization now being relatively abundant.
Cockayne (1928) and others have emphasized the endemic nature of the New Zealand flora and the extensive hybridization. These two conditions may be interpreted to indicate a state of flux in the vegetation types as well as the flora, perhaps because of the relatively late arrival of additions to the flora, intensive competition, selection, and other factors. Much of the endemism is of the initial type, rather than the relic, which indicates that some plants are migrating into new habitats, physical and social. A number of the new flora developed since the Tertiary began, and probably after mid-Tertiary times, seem
Olearia, Senecio, Epacris, and Cyathodes.
Land connections of the past may explain some of the Tertiary additions to the flora of New Zealand but care has to be exercised in making such interpretations from maps of the present distribution of species, genera, and families, or from the scanty fossil members of similar groups. A notable case in point is the genus Nothofagus which has been assigned a subantarctic origin because of its then known distribution. However Nothofagus forests have been observed recently in New Guinea (Archbold, 1942, and Landon, 1947) within a few degrees latitude of the equator, and on this island it is not probably related to any antarctic land mass. It is now just as plausible to consider Nothofagus a genus of Malayan origin as antarctic origin.
Migrations of plants across wide oceans may not have occurred consistently during the long geologic periods but it is extremely likely that even more intensive and extensive migration occurred during certain times. With the advent of flying animals, possibly reptiles and certainly birds, which migrate in large numbers animals may have moved countless disseminules in nearly all directions. Birds are now very effective in carrying fleshy fruits and hard indigestible seeds or fruits, or fruits or seeds that become attached to them. Some New Zealand plants now readily carried by birds are Rhopalostylis, eaten by parrots, Astelia and Hedycarya, the latter eaten by pigeons, and probably many species of Coprosma and Podocarpus. Perhaps over 150 of the present species are consumed by birds and transported.
Ocean currents of the past may have been even more effective than at present. The present ones are fairly well known (Davies, 1947) and from their direction of movement some correlations with dispersal of plants is sufficiently exact to indicate transportation by these currents.
In general insular areas are converging points for many surface ocean currents, some air currents, and migratory routes of birds. It seems probable that islands during a long period of time have therefore received more plants from afar than would continental areas of similar size and location. Even a small amount of observed migration of plants during the present has significance because the interval of observation has been very small compared to geologic ages and the long past has multiplied enormously the effectiveness of the occasional migration.
These three lines of evidence are greater than presented here in so brief a paper, and they are not enlarged upon because they have been more fully considered in other publications, especially about other regions. Each evidence is significant but the concerted value of all
Cocos nucifera might well be studied and experimental migration tests made. Oceantographers, ornithologists and botanists with such training can do much to contribute more information.
The above lines of evidence have been based to some extent on principles of plant ecology, the two following points of evidence are particularly related to ecological concepts and observations. They involve mainly (1) the significance of epiphytes as these plants imply certain modes of dispersal, and (2) the singular manner of distribution of certain genera, and species either along the coasts or on the high mountains, or at both places. In both instances disseminules are “precipitated” from the air; epiphytes often being perched in trees because the winds or some bird deposited them there, and coastal or high mountain plants occurring where they are mainly because winds are frequently reduced in speed or otherwise changed by coastal features, or the mass of the high mountains. Thus the two agents, birds and winds, are in a sense precipitating from the air many disseminules onto the forest canopy, or onto the relatively unpopulated areas of the coasts or high mountains. The fruit eating birds of the past and present have helped lodge disseminules above the ground. The winds have acted to carry many light weight disseminules to the coasts and to the high mountains. In New Zealand at present the most constant winds imoinge upon much of the western coasts from the northwest thus breaking their force against the high mountains of the Southern Alps and seasonally the Fohn wind blows off these mountains onto the Otago and Canterbury plains.
Fortunately Oliver (1930) has very thoroughly considered the kinds and nature of the epiphytes of New Zealand, His description of them also stresses their possible mode of origin as regards affinities to species of other land areas. He did, in some cases, particularly stress their means of dispersal by winds and birds, which may be considered the sole means of dispersal in numerous instances. Classifying the epiphytes as to modes of dispersal he noted: 35 species of Pteridophytes dispersed by minute spores, winds; 7 species of orchids by minute seeds, winds; one common Senecio species by pappus fruits, winds; 2 species
Pittosporum, by viscid coated seeds, birds mostly; and 5 species with fleshy fruits, birds. These are but the consistently epiphytic species. If the occasionally epiphytic or ephemeral epiphytic species are also considered the list is much longer and includes some of the important forest trees as Dacrydium cupressinum, Podocarpus spp. and many species of
The epiphytes of all types, including the lichens, mosses, and other non-vascular plants are very numerous in some of the New Zealand forests. In terms of number of species these epiphytes may not be many more than found on some other areas of similar size and conditions elsewhere, but in terms of number of individuals they are probably more numerous than in many other similar areas. This abundance of perched plants, many of which were precipitated from air currents or were dropped by birds signifies an efficient mode of dispersal by these agents. Cockayne (1928) and others have related the abundance of the epiphytes in general to the “rain-forest” condition and the subtropical nature of the vegetation. There is little doubt that the abundant precipitation over large areas and the high humidity account for some of the epiphytes, but there is some doubt as to the subtropical nature of the forests. The relatively great abundance of epiphytes may be a reflection of the ease of dispersal of the species involved, their ability to survive and other ecological considerations, as well as the climatic conditions.
Possibly the habit of epiphytism is a partial result of their mode of dispersal. Some species normally not epiphytic, or of genera that are non-epiphytic, upon migrating into a dense type of vegetation depend upon their only survival by their ability to remain perched above the soil and away from the intense competition of the forest floor. The genus Pittosporum illustrates this possible mode of development of epiphytism. The genus is distributed over Malaya and, with the exception of 2 species that are epiphytic in New Zealand, all so far found are non-epiphytic. Griselinia may be another example. It is a genus of 8 known species with all but 2 species in Chile, and some in Chile and in New Zealand are epiphytic. Similarly the genus Astelia furnishes some evidence of trans-oceanic dispersal. It is represented by about 19 species, some occur in dense lowland forests and some as subalpine mat plants. Significantly, species in New Zealand are both epiphytic and terrestrial. Epiphytes of this genus also occur in Fiji, Samoa, Society Islands and New Caledonia. Winds and birds may have dispersed these widely. It is interesting to note that the most fleshy fruited
Astelia are the most widely distributed indicating that birds may have aided in their dispersal.
Species and genera normally epiphytic wherever found are well represented in the flora and many are important vegetational components, particularly the many ferns and a few orchids. These autoepiphytes do not suggest special adaptability to the New Zealand conditions but their great abundance may indicate that they were easily dispersed, and precipitated.
The numerous facultative or ephemeral epiphytes such as Podocarpus, Dacrydium, Olea, Metrosideros and Coprosma are suggestive by the adaptability of their young that they may be readily dispersed by winds and birds. Metrosideros with many species in New Zealand is of very wide distribution, as far west as South Africa, but only in New Zealand are the species ephemeral epiphytes or climbers. The climbing or vine habit is in some ways much like the epiphytic habit and the abundance of this growth form of plants is also a notable feature of many New Zealand forests.
Oliver concludes from his study of epiphytes thus: “The evidence from geographical distribution thus points to the derivation of most of the epiphytes of New Zealand from the element of Malayan tropical facies in the flora. The method of dispersal is important, for obviously for the dissemination of epiphytes from tree to tree it is necessary for the seeds or fruits to be carried to suitable spots by such agencies as birds and air currents.”
As pointed out above the winds often converge upon island areas and these are frequently in two main strata, the low winds and the high winds. Both these may be in the same direction or may be in different directions. They normally are reduced in speed by obstructions provided by the physiographic features of the land mass. In many cases the winds are reduced at the coasts; in some other cases, especially the high elevation winds, the high mountains act as a barrier to them. At both localities there probably would be more deposition or precipitation of objects from the wind currents than at places between the coasts and the high mountains. Birds are affected by winds and may also drop more disseminules at these localities.
If, therefore, there are species similar or the same along the coasts and in the mountains there presence might indicate these modes of origin of the species. Frequency of such species is difficult to determine from the floristic literature and many more observations are needed to get enough data to give proper weight to this possible line of evidence for trans-oceanic dispersal. However, some data has been accumulated and a few samples of it will be considered.
There are a number of genera of the Compositae with pappus fruits which are distributed over the two main islands and some of the islands off the New Zealand group in the manner we are considering. Olearia, as an example has many endemic species such as O.
semidentata, O. traversii, O. allomii and O. traillii on the small offshore islands and along the coasts of the main islands are species such as O. virgata, O. paniculata, O. operina and O. angustifolia. In the mountains, often on the high subalpine parts, are species of Olearia such as O. macrondonta, O. avicenniaefolia, O. moschata and O. ilicifolia. Some species of Olearia, as O. colensoi, occur at the coasts and in the mountains.
Another genus with similar pappus fruits and coastal and mountain distribution is Celmisia. On islands or the coastal areas are C. rigida, C. major, C. lindsayi, C. vericosa and C. graminifolia, and in the subalpine areas are C. alpina, C. gracilenta, C. spectabilis, C. coriacea, C. argentea, C. lyalii and many others.
The genus Senecio has S. lyallii, S. bellidioides and S. eleagnifolius in the mountains and S. rotundifolius, S. glaucophyllus and S. greyii on islands and along the coasts. Other pappus fruited Com-positae genera as Helichrysum and Gnaphalium are similarly coastal and montane. Some genera, notably Haastia, are entirely montane and Raoulia is chiefly montane but has some species along the coasts.
A number of genera of the Epacridaceae, particularly Dracophyllum are common near the sea and on the high mountains. Of Dracophyllum the species D. rosmarinifolium and D. menziesii are subalpine and scoparium is typically coastal, but D. longifolium is both coastal and subalpine. Some of the other species spread over areas from the coast into the mountains.
Other families have genera of similar distribution, as Ourisia of the Scrophulariaceae, which are herbs with small seeds in capsules. The species O. colensoi and O. sessilifolia are high montane and O. macrocarpa and O. macrophylla are distributed from the coastal sounds to the mountains.
Astelia, a genus of the Liliaceae has berry fruits and may be transported by birds. Some of the coastal species of this genus are A. banksii and A. sublata, and some of the high mountain species are A. linearis, A. petriei and A. cockaynei.
Many more genera could be cited to illustrate this coastal and high mountain distribution which may reflect the mode of dispersal and indicate something of the effectiveness of such dispersal. Some species with members both along the coasts and on the high mountains similarly, and perhaps more effectively, also indicate the wind and bird types of dispersal. A few have been cited above, and some more are Raoulia australis, Celmisia cordata, Olearia avicenniaefolia, Senecio bellidioides, and some grasses as
Persons more familiar with New Zealand vegetation and flora can no doubt cite many more examples of this disjunct type of distribution than given here, and possibly some of the species cited are not good examples as the writer has relied mainly upon distribution notes from Cheeseman (1925) and a few observations and conversations.
Relative values of different modes of origin of the New Zealand flora and types of vegetation are briefly discussed and the possibly important mode of dispersal across oceans and seas is emphasized. Among the different lines of evidence supporting the importance of this trans-oceanic dispersal of plants the evidences that may be obtained from the abundant epiphytic flora and the ecology of the condition of epiphytism are stressed. Similarly the disjunct distribution of species of some genera and of some species are considered as significant evidence of dispersal, possibly across oceans, the disjunct distribution being near the coasts and on the high mountains.
Last year, specimens of live peripatus, Peripatoides novae-zealandiae, were sent from Victoria to the Chicago Museum of Natural History. Peripatus is an animal which most New Zealanders have probably seern at some time or another when they have kicked open a piece of rotting wood in the bush, but have not recognised as such, imagining it to be merely “another caterpillar.” The interest it holds for zoologists is reflected in this article written on the arrival of the New Zealand specimens at the Chicago Museum by Rupert L. Wenzel, assistant curator of insects, and published in the Chicago Natural History Museum Bulletin (July, 1949).
These remarkable creatures belong to the Phylum Onychophora, an extremely ancient group that has existed for at least one-half billion years. A fossil form, Ashyeaia pedunculata (see figure 2), has been described from Middle Cambrian deposits of that age. The illustration shown is of a hypothetical reconstruction, but the actual fossil specimens show most of the details of the external structure rather clearly.
If the fossil described as Xenusion auerswaldi (see figure 2) actually represents an onychophoran or onychophoran-like animal, as it seems to, then the beginnings of the group might be extended back another one-half billion years, for Xenusion is from Proterozoic Algonkian rocks that are approximately a billion years old. Judging from the deposits in which these fossil forms were found, the early Onychophora were marine.
The living species are terrestrial. They are few in number—about eighty species—and occur in the West Indies, Central and South America, South Africa, and the Indo-Australian region. Although they are terrestrial and have special breathing tubes known as “tracheae,” they are restricted to very moist environments; their thin skin makes them subject to very rapid dessication. They generally avoid light and live in rotten logs, under stones, under loose bark, etc., where they feed on small insects and other micro-organisms. Most of the species are small—about 2 to 3 inches long—but at least one attains a length of 5 inches.
One of the interesting protective adaptations of these animals is the ability to squirt a sticky, slime-like secretion that effectively tangles their enemies. The slime is secreted by long internal slime glands that extend almost the entire length of the body and is ejected from a pair of “oral papillae” (see figure 2), of which one is on each side of the head. Certain species can squirt the slime as far as twelve inches. Some Onychophora lay eggs. Others give birth to living young; in such species, special placenta-like structures may develop to facilitate the diffusion of nutrient materials through the uterine wall of the mother to the developing embryo, in much the same fashion as in mammals.
It is not only because of their great antiquity that the Onychophora excite the curiosity of the zoologist. An even greater fascination lies in the fact that these animals possess anatomical features both of the segmented worms (Annelida), of which the common earthworm is a familiar example, and the Arthropoda, the great phylum that contains the crabs, lobsters, shrimps, spiders, scorpions, millipedes, centipedes, and insects. The first peripatus described was considered to be a mollusc, and it was not until careful anatomical studies had been made that its position in the animal kingdom was appreciated. There can be little doubt that the arthropods evolved from worms or worm-like ancestors, and many zoologists consider that the Onychophora are an intermediate or linking group between the worms and the Arthropods. It is more probable, though, that the Onychophora represent an offshoot from the main line of evolution between the two.
Modern soil textbooks make a point of impressing the reader with the need to bear in mind that “the soil is a living body.” Such warning is hardly intended for the biologist who, no matter what his special interest, cannot fail to be aware of soil as an integral part of the plant and animal environment, and further he will know that many organisms have it in their power to modify their immediate habitat, making it conform more nearly to their needs. Soil, therefore, has many of the attributes of a living body solely because it supports so many living organisms. These range in size from bacteria to kauri trees. The by-products of their life in the soil and the residues left in the soil as a result of their occupation together constitute the soil organic cycle.
The earliest members of the organic cycle are those micro-organisms, mainly soil bacteria, which derive their energy from the multi-valent elements like sulphur, iron or manganese, and build up carbon compounds from carbon dioxide which may be present in either gaseous or dissolved form. Organic compounds built up in the bodies of these pioneer organisms then become a source of energy for more specialised micro-organisms which have the chemical tools required for breaking down complex carbohydrates.
The organic cycle commences at a very early stage in soil formation, often simultaneously with normal soil weathering processes. In many cases it can be shown that organic acids released by soil microorganisms are an important factor in the weathering of mineral particles.
The biotics division of the Soil Bureau is a research team recently set up to enquire into the nature of the organic cycle in New Zealand soils. So far it has done mainly exploratory work to see whether our soil classification (based upon the concept that soil types can be grouped by considering the action of a common set of soil forming factors upon the various kinds of parent materials) means anything to the organisms that live in the soil. N. solandri and N. cliffortioides in the Taupo region was examined.
can modify the soil environment, thereby incidentally altering soil fertility—a property of the soil that is closely regarded by the farmer. R. H. Thornton has commenced a survey of fungi and actinomycetes in New Zealand soils but the problems of identification are too difficult to be resolved by a solitary worker in this field and he is at present studying under Professor Chesters at Nottingham University. P. J. Culliford has commenced a survey of the conditioning influence of the different native tree species upon soil development. This is going to be an important line of research, for the great majority of our soil types are developing mainly through the process of leaching, and differences in the organic cycle (of which the tree is usually the strongest component) can readily cause major differences in the rate and kind of leaching and so greatly modify the kind of soil formed. This is well illustrated by the mosaic pattern which is often noticeable in freshly ploughed land that has formerly been covered with forest. The lighter gray patches of soil are areas once occupied by those trees whose litter was, for various specific reasons, prone to accumulate, forming a layer of peat in which slow decomposition was carried out by a predominantly fungal micro-population. The resulting increase in acidity has accelerated normal leaching and caused the “bleached” appearance of the soil.
As we learn more of the activities of the higher plants and animals, and microorganisms, we find that their metabolism is often controlled by specific soil conditions. Out of this grows the technique of bio-assay, whereby the growth of an organism can be used as an indicator for a particular soil condition. The soil fungus, Aspergillus niger, requires a small amount of copper to complete its life cycle. The ramifying hyphae can extract copper from a mineral soil in about the same degree as the roots of higher plants. Thus the fungus can be grown in agar cultures with a small volume of soil added to supply copper, and the colour of the spores at maturity can be used as an index of the amount of copper in the soil which is available to higher plants. This technique is being employed by D. Breen to make a survey of the available copper in New Zealand soil types. Trichoderma sp. for boron bio-assay work, and Cunninghamella sp. for zinc, while Aspergillus niger can also be used to estimate magnesium, manganese and phosphate.
The higher plants are themselves employed for evaluating availability of certain trace elements by E. J. S. Gridley of the plant indicator research unit. This section grows indicator plants, such as mustard, virginia stock, lettuce, etc., in pots and studies the effect of a standard range of trace elements on plant growth in different soil types. We have been able to demonstrate that hitherto unsuspected cases of boron, copper, zinc and manganese deficiencies are present in a number of important soil types.
Considering their importance for our well-being, it is remarkable how little is known about the working parts of the organic cycle in soil. The biotic surveys were expected to find many new and unexpected facts about soil life but so far it has been the bio-assay work (originally started as a service to supplement the work of the chemistry division of the bureau) which has stirred up the more thought-provoking problems. While using Aspergillus niger for phosphate bio-assay, Breen discovered that on some phosphate deficient soils the fungus is quite unable to grow unless supplied with either citric acid or a small amount of soluble phosphate as a “starter” in the medium. It is well known that A. niger and many other soil fungi secrete organic acids but Johnston has shown that many of these organic acids readily dissolve calcium and magnesium phosphates and even attack iron and aluminium phosphates. Johnston considers that there perhaps lies the explanation of such problems as the phosphate nutrition of lichens growing on exposed fresh rock surfaces, the beneficial results of mycorrhizal association in the roots of higher plants, and the tooth-destroying powers of the organisms responsible for dental caries. Theoretically it should be possible to culture powerful phosphate solubilizing micro-organisms in phosphate deficient soils and, as a result, make available to the higher plants phosphate from sources which are normally not available to plant roots. Pot trials with sour milk (nourishing lactic acid bacteria) and Aspergillus broth have gone some way to confirming the hypothesis.
Another unexpected idea came from our trace element experiments. It is customary to add most trace elements as sulphates, and both copper and zinc sulphates have been fairly consistent in promoting increased plant growth. It would be unwise, without further experiment, to conclude that this indicates a general copper or zinc deficiency in the New Zealand soils, for the increased plant growth strongly resembles that which normally occurs when nitrogen is added to the soil. It may yet be shown that some trace elements, when added to the soil, have their main effect upon the growth and metabolism of the soil micro-organisms rather than the higher plant, and that at least a part
Naturally, an editor of an Encyclopedia cannot have a sub-editor for every animal, but that is what the zoologist apparently expected of me. Matters are far worse than in the days of Dr. Holme's naturalist who flew into a rage because someone called him a Coleopterist. He was no smatterer, he said, trying to spread himself over the Coleoptera; he was a Scarabaeist. Nowadays a zoologist seeks out his animal in early life and henceforth stays with it. Often the intimacy between them is so great that it seems indelicate to intrude. I have known a bivalve and a man to develop interests in common so exclusively molluscous or bivalvular that no human dared to break in.
Collecting, to a zoologist, is only a means to an end. His real work starts when the collection is fully documented, and is complete enough to enable him to recognise the animals of a region. A student interested in species as such may follow up the taxonomy of a group, leading to a study of its evolution, and the relationships of its members. Or his main interest may lie in the adaptations of the animals to their environment, leading to a study of the biology of one species, and later to that of a group of animals-ecology or animal sociology. From either point of view, the forming of a collection will serve as a tool to discover and a tack further problems. And to the New Zealand zoologist, problems will soon present themselves.
The following notes are designed for the student beginning serious collecting. They outline the steps to be taken from the time of locating and securing the specimen, through the stages of fixing, preservation and recording.
Later articles will cover methods of collecting and preserving the Insects and Arachnids, the terrestrial Arthropods (Millipedes and Centipedes), Vertebrate animals, and the various groups of endoparasites.—Editor.
Organisation of field work is best left to the collector. Most marine work will be done between tide marks and full advantage should be taken of the predictions and times of lowest tides in the Nautical Almanac. On rocky coasts especially, the area should be reached an hour or two before low tide. The upper portions of the reef can then be thoroughly combed, and will not be neglected in favour of the richer substratum. A general recennaisance of the ground may be made before collecting; if the locality is a new one, a simple sketch plan together with a photograph often helps. Rapid sketches and measurements should be made of zonation patterns of sessile plants and animals.
In summer, or on North Island coasts, shorts or swimming trunks are the most serviceable dress for moving about in at low tide mark. Strong, open sandals are the best footwear, allowing the water to run out, but protecting the feet. During winter, rubber knee-boots are useful and comfortable. They should never be worn, however, on mudflat or soft sandy bottoms, where they render the wearer helpless if he gets bogged. A strong-bladed pocket knife should be carried, together
Formalin, where its use will not harm the specimen, is a better preservative than alcohol. It penetrates more rapidly and internal organs remain in better condition. Commercial formalin (40%) should be diluted with ten volumes of water to a 4% solution. 2% formalin with seawater is an excellent quick preservative for small specimens.
Alcohol as used in zoology is generally 95% ethyl alcohol (white spirits) which may be diluted with distilled water to strengths of 70% and 80%. At least 70% is required for safe storage of material. Alcohol is a valuable preservative for crustacea, polychaeta and echinoderms with corrodable bristles or hard-parts. A teaspoonful of glycerine in a quart of alcohol helps to preserve natural colours and to keep integuments flexible. Alcohol in jars containing preserved specimens should be changed at intervals—once or twice a year—and evaporation should be guarded against. One of the most important fixatives for the general collector of invertebrates is Bouin's Fluid, which is excellent for general structural and histological work, and for preservation of animals for dissection. Fixation time is at least 12 hours for a specimen of lcc. bulk, correspondingly longer for larger material. There is not much danger of over-fixing. Afterwards specimens should be washed in 70% alcohol, to remove excess picric acid, and stored in 70-80% alcohol. Formula for Bouin's: 75 parts saturated aqueous picric acid; 25 parts 40% formalin; 5 parts glacial acetic acid.
The illustration shows a suitable type of field record. Several hundred of these should be cyclostyled on quarto sheets at the beginning of a collecting season. The paper should take ink, and it may be perforated along the margin for filing. During field work, the collector will use his note-book for “on the spot” observations. On returning home, he should—even at some inconvenience—sort over and roughly classify the specimens, discarding unwanted material at once. Sketches and colour records must now be completed while the animals are still alive. The field sheets should then be filled in, before exact details
reference number corresponding to the serial sheet number. Specimens are thus numbered simply in order of collecting and writing up. An abbreviation for the name of the class can be usefully added to the sheet below the serial number, e.g. (CTEN — GASTR — CRUST —). The sheets may then be filed in serial order, and, if necessary, as the collection grows, cross-indexed in taxonomic and ecological categories. Details of locality, type of environment, ecological association should be specified, with date and collector's name. Space is left for details of novelty or special interest, but with well-investigated species, this will not necessarily be filled. Reference should be given to any life drawing, colour record, slide or other preparation made from living material. After identification the specific name can be added in the space provided, together with a note referring to taxonomic or general literature. A brief summary of diagnostic features can often be added with advantage.
As the collection grows, the greatest need will be for an orderly storage system, with facilities for quick reference to any container, to filed field notes and any other information. It may be desired to have the collection visible for permanent display, and with mollusc shells or dried material a set of flat cabinet drawers is best for storage. Most invertebrates, however, will be kept in bottles, and sets of tubes or jars can be massed together in a small space by placing them in deep cardboard boxes, conspicuously labelled with serial box numbers, and either the ecological to taxonomic classification of the material contained, e.g., either Low Tide — Wharf Piles — Devonport, or Echinodermata (Ophiuroidea and Holothuroidea). At the same time, the field record sheets should be kept in readily accessible files, again either in ecological or taxonomic order.
A tidy-minded collector will probably prefer several uniform sizes of tubes and jars. These pack more economically into limited space and are fairly cheap in quantity. Jars should be wide-mouthed, and of clear glass, thick enough to withstand knocks. Bakelite or other noncorrodable tops are preferable to metal caps. Alternatively, widemouthed bottles may be stoppered with firm, good quality corks.
Preserving jars with rubber washer lids are ideal for larger specimens, while for the bulkiest material, metal drums of two or four
The bulk of the information about a specimen should be entered on the record sheet. It is not usually convenient to label material exhaustively in situ. Labels attached to the container should include the habitat, locality and date, and the collector's initial, with perhaps a condensation of any other information to which it is desired to refer without turning up the field sheet. Most important, there should be a reference number to the written field record. Alternatively specimen labels can be placed in the preservative within the container. They should then be written on stiffish, non-absorbent white card, either in pencil (not “indelible” pencil) or in indian ink. Ink labels should be allowed to dry, then steeped for a few minutes in a 3% solution of acetic acid, which effectively sets the ink and prevents “running” when placed in preservative. An example of a completed label is shown. Note that the accurate identification of the specimen is not important at this stage. Often it may not be possible immediately; and the name will have to be entered on the field sheet at a later date. Sometimes identification by a specialist will be needed, while in a few cases the collector may have the thrill of bringing home a yet undescribed species.
Low Tidal Sand-Flat (L.W.S.T.)
Cheltenham Beach 5.11.49. W.J.B.
Commensal in burrows of Trochodota
105 (LAM) (Scintillona zelandica)
1. Coelenterates are difficult to preserve adequately. The pelagic
2. In the Platyhelminthes (Flatworms) and
3. Polychaeta or larger bristle worms present fewer problems. Their colours are often brilliant and characteristic, and should be recorded in life. Formalin—unless rendered neutral in reaction—damages the setae, on which systematic work so much depends. Use, therefore, 70% alcohol, bringing up to 80% with a drop of glycerine to prevent shrinkage and undue fading. Worm tubes — calcareous, parchment or sandy, should be kept. Habit sketches should not be neglected, especially with worms having a specialised relation to the substratum. Feeding, respiratory and cleansing mechanisms can be properly studied only with living material, and the field biologist will get useful “hunches” from watching the behaviour of his specimen before collecting, and observing the action of tentacles, mucous glands, parapodia and ciliary fields while the worm is still alive.
4. In the Echinodermata identifications are based to a major extent on skeletal parts. Formalin should therefore be avoided in favour of 70-80% alcohol, with glycerine to keep soft articular membranes flexible. Much more difficult to handle than the starfish or urchins are the delicate
The Holothurians or sea-cucumbers are equally temperamental—eviscerating themselves when irritated. Before preservation the animals should be narcotised by the addition of magnesium sulphate or menthol to the sea water in which they are contained. When completely
5. Ascidians and Enteropneusta. The tunic of simple ascidians should be slit to allow penetration of formalin when the specimen is detached from the substratum. Colour notes, both of test and contained animal, are of great importance. Shrinkage invariably takes place, so that accurate measurements must be taken in life. Compound ascidians, after the all-important colour note, should be fixed directly, some left in the test, others pressed out on slides. Glacial acetic acid is recommended, with transfer after 5 minutes to 95% alcohol. In the
6. Crustacea. With large crayfish, anomurans and crabs, as well as barnacles, material may be dried, but fluid preservation is better, using 70-80% alcohol with glycerine, not formalin. Good preservation should keep setae intact and joint membranes flexible. Penetration of alcohol to viscera is poor, and where specimens are wanted for dissection, it may be necessary to use form-alcohol, even with detriment to the exoskeleton. Crustaceans are especially interesting in the modifications of mouthparts and other appendages, which should be observed in action in living material. Notes should be made of the dates at which specimens are in berry, and in the case of parasites, host relationships should be recorded. In the
7. Most of the Mollusca are very easily preserved, and while perfect cabinet specimens are not necessary for the general zoologist, shells may be carefully scrubbed with a fine nail-brush to remove encrusting growths. Chitons should be firmly bound to wooden splints
Bouin's fixative is recommended, and it is often advisable to sacrifice one or more shells, lest the animal be damaged by extraction. Small gastropods should be gently cracked and dropped into Bouin's intact. The shell is perfectly dissolved, and the fixed animal remains, with chitinous operculum and radula intact. Bivalves should be wedged open slightly to allow entry of fixative. Colour notes and habit sketches of the extended animal should not be forgotten. In bivalves, note the size and shape of foot and siphons, in gastropods, locomotor, feeding and protective mechanisms, as well as details of spawning and copulation.
The soft-bodied Opisthobranchs are of unrivalled interest in their colours, and in diversity of structure and habits. They are, however, the most troublesome of invertebrates to preserve. Colour sketches and records of external form in life are indispensable. Narcotisation is difficult: a few methods may be suggested, but to a large extent the collector must be guided by his own mistakes. Slow, gradual tincturing of seawater with formalin, alcohol or coccaine is often successful. Alternatively, crystals of menthol, naphthalene or magnesium sulphate may be sprinkled upon the top of the water. For the study of internal structure it is best to pin freshly collected living material in extended condition, flood with Bouin's fluid and incise the dorsal integuments. Internal structure will otherwise probably deteriorate while the half-narcotised animal is still responsive to stimuli.
Most zoologists will have a working familiarity with one or more groups, and in many cases the collector himself will have specialist knowledge of his material. A field zoologist can usefully aspire to an acquaintance with a whole fauna as far as the family or generic level. In practice, however, there will always remain a large number of animals which must be sent to a specialist, usually a professional zoologist, for identification. Sponges, amphipods, polychaetes, and planktonic crustacea are likely to remain specialist's groups. Taxonomists are busy people. This should be remembered in sending specimens, which should be carefully sorted, separated as far as possible into distinct forms, securely packed and sufficiently stamped. Good specimens
Zoological collecting does not entail “sitting on” rare material to the envy of less fortunate competitors. Unused specimens are of little value lying on shelves. If you know of a man interested in—for example—barnacles, nudibranchs or crabs, let him have your material to work it up, even if he is not at once able to give quid pro quo.
Systematic coverage of New Zealand marine and freshwater invertebrates is very uneven. The following list of references is not exhaustive, but introduces the principal authorities and indicates the gaps still to be filled.
Mr. N. Hornibrook, Geological Survey, The Terrace Wgton.Foraminifera
Miss Gwyneth Parry, Canterbury University College, Christchurch (at present at Glasgow Univ.). Papers by F. G. A. Stuckey, T.N.Z.I. 1908, vol. 41. Miss Miss Beryl Brewin, Otago University, Dunedin.Coelenterata
Anemones:
Corals, Ctenophores, Hydrozoans:
Alcyonarians:
Miss S. Jonathan, Canterbury University College.Porifera
Prof. E. Percival, Canterbury University College. (esp. Rhabdocoelida.) Miss M. L. Fyfe, Otago University. Miss F. R. Nurse, Canterbury University College.Platyhelminthes
Turbellaria:
Land Planarians:
Freshwater Planarians:
Prof. E. Percival, Canterbury University College.Brachiopoda
Mr. C. R. Russell, 108 Knowles Street, Christchurch.Rotifera
Dr. G. H. Uttley, Canterbury Museum, Christchurch. Dr. Brown, Geol. Dept., Otago University.Polyzoa (Fossil)
Mr. Papers by Sir Mr. George Knox, Canterbury University College. Papers by Sir Prof. Annelida
Oligochaeta:
Polychaeta:
Hirudinea:
Prof. Dr. Miss Mr. A. S. Fuller, Auckland University College. Mr. B. M. Bary, Victoria University College. Mr. N. Hornibrook, Geological Survey, Wellington. Mr. A. W. E. Hurley, Victoria University College.Crustacea
Brachyura:
Barnacles:
Planktonic Crustacea:
Ostracoda:
Amphipoda, Isopod:
Mr. Mr. Dr. Marwick, Geol. Survey, Wellington. Mr. Miss Ralph, Victoria University College. Mr. Mollusca
Opisthobranchs:
Gastropoda, Pelecypoda:
Dr. H. B. Fell, Victoria University College. Mr. Echinodermata
Asteroidea, Echinoidea, Ophiuroidea, Crinoidea:
Holothuroide:
Miss Beryl M. Brewin, Otago University, Dunedin.Ascidians
“But so crafty have crustaceans been in keeping dark and not courting unnecessary observation, that very few persons know how numerous are the species or how astonishingly multitudinous are the individuals of some of those species. Still less known is the great variety of situations in which they may be successfully sought for. It is not only the shores of the sea and its various depths, down to about three thousand fathoms, that yield them. Out of the many miscellaneous dwellings which they favour, a few may be mentioned as samples, some of them being themselves crustaceans since the list includes the gills of a lobster, the cheek of a prawn, the abdomen of a hermit-crab, the viscera of a shore-crab, as well as the eye of a sprat, the mouth of a cod, the back of a turtle, the floating gulf weed, the slates of a roof, the fruit in a garden, the leaves of a tree, a deep well, a horse-pond, a cart rut after a shower of rain, or a packet of dried mud. In all these stations different kinds may at times be found. Some are parasites pure and simple, degraded in shape or almost without any definiteness of form; some are semi-parasitic and proportionately lethargic; others are free-living and extremely vivacious; nor are there wanting some so framed as to extort admiration even from the ignorant, a success of which nature has every reason to be proud, since even the best work of art or the most finished essay cannot always command it.”—T. R. R. Stebbing (1894).
To the average person the term “Crustacea” has the somewhat limited application of the familiar reddish-hued animal frequently encountered in trams. In reality, it covers a multitudinous range of species, rivalling, and only excelled in numbers and variety by, the orders of insects. Yet these species are to be found almost everywhere, including the back garden, for two of the orders of the group have invaded the land so successfully as in some cases to become garden pests. These two are the Orders Isopoda, or, more colloquially, the “slaters, sow-bugs, pill-bugs, woodlice,” and the Order Amphipoda, or “hoppers.” Both are found in garden litter, in rotting grass, under rotting wood, and in particular under the ground covering of leaves and decaying undergrowth in the bush. Both groups require a fairly high degree of moisture, and both groups form a comparatively large proportion of the leaf-mould fauna of New Zealand.
There is one very easy method of telling an amphipod from an isopod—the amphipod bounds away in leaps out of all proportion to
Most people know what an isopod looks like, under one name or another. “Woodlice” or “slaters” are the most common New Zealand terms for them. There is only one other kind of animal which may be mistaken for an isopod—a millipede found in New Zealand, mainly in high country. To tell them apart, turn the specimen on its back, and if the number of legs is obviously more than seven pairs, the animal is a millipede.
History: The first scientist to describe any of the native isopod species from this country was J. D. Dana, who led the United States Exploring Expedition of 1851-52. Most of his specimens were collected from the Bay of Islands, from such exotic—sounding places as Wykare and Taiammai. The only copy of the Crustacean section of his report which I know to be in New Zealand is in the Turnbull Library. Since Dana's expedition, isopod collections have been made by one French and at least three German expeditions. In New Zealand,
Of the forty-eight species known from New Zealand, four (Porcellio scaber, Armadillidium vulgare, Porcellionides pruinosus and Ligia exotica), are cosmopolitan. The last-named is mentioned as being found in this country by Jackson (1941) in his “Checklist of the Terrestrial and Fresh-water Isopoda of Oceania,” but I have seen no verification of this record. Amongst those of Pacific distribution are Actaecia euchroa, found in Tasmania, and Trichoniscus thomsoni which is recorded from Austral Island in the Polynesian Group.
Habitat: Although the term “terrestrial isopoda” has been used in this article to cover all forms of the Sub-order Oniscoidea, a number of the members of the sub-order are not truly terrestrial but littoral. These include
This habitat has had some influence on their distribution as is shown by specimens which Chilton reported to have been carried far down country on logs during a flood. The occurrence of Trichoniscus magellanicus and of species of Deto in South America has also been attributed
Perhaps the most interesting of the New Zealand species is Trichoniscus commensalis, which is found in ants' nests. This curious cohabitation is not uncommon in other parts of the world; in England and Europe Platyarthrus sp, and in South Africa Phylloniscus sp., Schoblia, Titana, and Kogmania sp. are commonly discovered commensal with ants or termites.
Prevention: It has already been mentioned that one of the best places to find isopods is in the back garden. In some cases they may become so numerous as to attack vegetables, plants and cultivated mushroms, their chief food being vegetable matter. They also show a liking for decaying wood or leaf mould. For those gardeners troubled by them, one recommended procedure is as follows: Trap a dozen or so, bottle them in alcohol of strength 75-95 per cent., and post with all details to a zoologist for identification. (N.B.—Do not expect an immediate answer—some of the material of the 1872-6 “Challenger” expedition is still being examined.) Then commence an all-out attack with paris green. One method is “to mix 4 ozs. paris green with 6d. bran and a little sugar. Moisten with treacle mixed with hot water and form into little balls. Place balls all round plants or under light boards where slaters gather. This is poison.”
A recent gardening column recommended D.D.T. In spite of the proven affinity of D.D.T. for chitin, it does not seem to be very effective on isopods. Paris green is the time-tried prescription, and can be bought in numerous proprietary mixtures.
Identification: To a certain extent, identification is made easy by the varied sculpturing on the back of animals. However, as in most crustacean groups, many of the species differ in points which are comparatively insignificant, and resort must be had to the examination of mouth-parts and appendages for specific identification. Again, some species differ in a number of very small points, each in itself not particularly obvious, but forming a complex of a quite distinctive nature. As a result, it is hard, and in fact impossible, to construct a key which will include all recorded species, and satisfactorily separate them without using these characters as distinguishing ones. For that reason it is necessary to explain some of the terms used.
Porcellio scaber, for instance, is an isopod which is unable to roll up into a ball (i.e., to “conglobate”). The head, or cephalon, carries two pairs of antennae. The first antennae are extremely small and lie between the second antennae of seven segments. The first five segments of the second antennae comprise the “penduncle,” the remaining two the flagellum. (In Ligia novaezealandiae there are 20 flagellar segments). The eyes are compound. On the under-surface of the
terminal branches (rami). Each of the seven thoracic segments bears one pair of legs on the ventral surface, whilst the under-surface of the first five abdominal segments is furnished with a pair of over-lapping platelike appendages (pleopoda) (fig. 15). The first two, or in some species the second pair only, are narrowed and modified in the male. The females lack this modification. Instead the thoracic segments are modified ventrally to form a broodpouch or “marsupium” in which the eggs and young are carried. The external plates of the pleopoda overlap other soft, vascular, leaf-like plates which are part of the pleopoda and are used for respiration.
The paired mouth-parts, as found in the Oniscoidea, named from the lowest and outermost pair first are:
Maxillipeds: A pair of somewhat oblong, flattened appendages, the median borders straight and fitting close together, and serving as a covering for the other mouth-parts. They have near the end a short, flattened, segmented palp (e.g., figs. 26, 27).
Maxillae: Two pairs, the second maxillae outermost. These latter are broadly plate-like, without teeth, and only partly divided into inner (median) and outer lobes. The first maxillae (e.g., figs. 21, 22, 23) have two (outer and inner) branches or lobes arising from a slender transverse basal segment. The outer lobe may have setae or it may have eight or more curved teeth terminally; and the inner, which is more slender and weaker, has two or more brush-like tufts of setae.
Mandibles: Pear-shaped appendages (e.g., figs. 13, 14) ending in a curved process bearing a few strong teeth. The inner margin may carry a ridged upraised area (“molar process”).
Above these paired appendages is the unpaired upper lip.
The first thoracic segment is often provided with an outwardly bent or rolled up border along its lateral margin. In forms which conglobate, this border has a cleft at the rear corner. When the animal rolls up, the leading edge of the epimera of the second segment fits into the cleft which may be continued forward as a furrow along the lower aspect of the border of the first segment. The furrow is known as the “coxopodite sulcus” or “sulcus,” and the segment is spoken of as being “sulcate.” This character is a very important aid in distinguishing species.
Identification in most cases requires dissection of mouth-parts. This can best be done with finely sharpened mounting needles under a dissecting microscope. The parts may be removed together, stained
References: The latest and most complete check-list is Jackson's “Check-list of the Terrestrial and Freshwater Isopoda of Oceania,” in the Smithsonian Miscellaneous Collections, Vol. 99, No. 8, 1941. Bowley (1935) has further separated the species of
Two species are insufficiently described to be keyed out: Spherillo brevis and Oniscus cookii. Oniscus cookii is probably identical with P. kenepurensis.
The group, Spherillo setaceus, S. bipunctatus and S. squamatus, has not been figured or described in sufficient detail to determine whether or not the pleon is sculptured. Accordingly, the key has been arranged to separate them in either dichotomy.
Acknowledgment: I wish to thank Miss Eileen A. Bowley for her kind permission to reproduce figures 14, 16, 17, 18 from her paper on
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Cephaloscyllium, Inflation of the Abdomen in, By
China, Fishing Industry in, By
Climate Relations of Fossil and Recent Floras. By
Botanical Specimens, A. Guide to the Collection of, By Zoological Specimens, Collection and Preservation of (Marine Invertebrates) by Collection:
Conservation, Soil, The Place of the Botanist in, By A.
Crabs—See Brachyura and Crustacea.
Crinoids, N.Z., By H. B. Fell. 3. (2): 78-85.
A Guide to the Brachyrhynchous Crabs. By A Guide to the Oxyrhyncha, Oxystoma and Lesser Crabs. By Lyreidus australiensis Ward (Brachyura, Gymnopleura) from Cook Strait. By Corrections and Additions for the Guides to the Brachyura. By Isopods, N.Z. Terrestrial, By Crustacea:
Cytology and Genetics and their Application to N.Z. Plants. By
Disease;
Mosquito Borne, and the War in the Pacific, By
Dispersal of Plants to New Zealand, Evidences of Trans-Oceanic, By
Asteroids of N.Z., Key to the Littoral, By H. B. Fell. 1. (1): 20-23. Echinoids—Key to the Sea Urchins of N.Z., By H. B. Fell. 1. (3): 6-13. Ophiuroids, N.Z. Littoral, By H. B. Fell. 2. (3): 121-129. Holothurians of N.Z., A Guide to, By Echinoids—Key to Sea Urchins to New Zealand, Additional Species, By H. B. Fell. 3. (1): 42. Crinoids, N.Z., By H. B. Fell. 3. (2): 78-85.Echinoderms:
Editorial. By
Editorial—By
Eels, N.Z. Freshwater, By
Epiphytes—Evidences of Trans-Oceanic Dispersal of Plants to N.Z., By
Entomology, Shakespearean, By
Feather-Stars—N.Z. Crinoids, By H. B. Fell. 3. (2): 78-85.
Eels, N.Z. Freshwater, By Grayling, The N.Z., A Vanishing Species, By K. Radway Allen. 2. (1): 22-27.Fish:
Fisheries: Problems of Marine and Freshwater Fisheries Biology in N.Z., By
Fishing Industry in China. By
Fossil Flora of N.Z. By Climate Relation of Fossil and Recent Floras. By Evidences of Trans-Oceanic Dispersal of Plants to N.Z. By Flora:
Geological History of N.Z. By Climate Relation of Fossil and Recent Floras. By Geology (see also Palaeontology):
Genetics and Cytology, Application to N.Z. Plants, By
Genetics Discussion Group. By A.
Grasslands in N.Z., The Conversion of Rain Forest to, By E. Bruce Levy. 2. (1): 37-51.
Hepaticae, Classification of N.Z., By E. Amy Hodgson. 3. (1): 20-32.
Hepaticae, Classification of N.Z., Corrections to, By E. Amy Hodgson. 3. (2): 86.
Isopods N.Z. Terrestrial, By
Holothurians, A Guide to the N.Z., By
Kapiti Island. By
How to Use Keys for Identifying Organisms. 1. (2): 6. Botanical:
Lichens, Note on with a Key to the Commoner N.Z. Genera, By Lichens, Crustaceous, A Note on the N.Z., By Lichens, Stictaceae of N.Z., Bv Mosses, Some Note on, with Key to Commoner N.Z. Genera, By Mosses, Correction to Key. By Hepaticae, N.Z. Classification of, By E. Amy Hodgson. 3. (1): 20-32. Hepaticae, Classification of N.Z., Corrections to, By E. Amy Hodgson. 3. (2): 86. Zoological:
Asteroids of N.Z., Littoral, By H. B. Fell. 1. (1): 20-33. Spiders, Common, of the Wellington District, By Sea Urchins of N.Z. By H. B. Fell. 1. (3): 6-13. Crabs. Brachyrhynchous, A Guide to the, By Crabs, Oxyrhyncha, Oxystoma and Lesser, By Crabs, Brachyura, Corrections and Additions for the Guides to, By Ophiuroids, N.Z. Littoral, By H. B. Fell. 2. (3): 121-129. Shags, Identification of N.Z. Mainland, By Sea Urchins of N.Z., Key to, Additional Species, By H. B. Fell. 3. (1): 42. Holothurians of N.Z., A Guide to, By Crinoids, N.Z., By H. B. Fell. 3. (2): 78-85. Isopods, New Zealand Terrestrial, By Keys:
Kirk, Harry Borrer, In Memoriam, By F. A. de la Mare. 1. (3): 1-4.
Lancelet, N.Z., The Generic Status of, By
A Note on with a Key to the Commoner N.Z. Genera, By N.Z. Crustaceous, A Note on By Key to the Stictaceae of N.Z., By Lichens:
Classifications of N.Z. Hepaticae. By E. Amy Hodgson. 3. (1): 20-32. Classification of N.Z. Hepaticate, Corrections to, By E. Amy Hodgson. 3. (2): 86.Liverworts:
Marchant Ridge, Tararuas. By
Mosquito Borne Disease and the War in the Pacific. By
Note on Some N.Z., with a Key to the Commoner N.Z. Genera, By Correction to Key. By Mosses:
Museums, N.Z., Biology in, By
Nothafagus, Requests for Observation on Flowering of, By
Notornis, Correction, By J. H. S. 2. (3): 115.
Obituary—In Memoriam—
Onychophora—See Peripatus,
Ophiuroids, N. Z. Littoral, By H. B. Fell. 2. (3): 121-129.
Oxyrhyncha, Oxystoma and Lesser Crabs, By
Geological History of the N.Z. with Reference to the Origin and History of the Flora and Fauna. By Vertebrate, in N.Z., By Flora of N.Z., Fossil, By Flora, Climate Relations of Fossil and Recent, By Palaeontology:
Peripatus “Living Fossil” and “Missing Link.” By Rupert L. Wenzel. 3. (3): 98-99.
Phytoplankton. By D. Alleyne Crawford. 1. (1): 15-20.
Plant Quarantines, Changing Conditions and, By
Plant Research Institutions, N.Z., Scope of the Biologist in, By
Plymouth Marine Station Life at, By
Botanical Specimens, A Guide to the Collections of, By Zoological Specimens, Collection and Preservation of (Marine Invertebrates), By Preservation:
Rain Forest to Grassland in N.Z., The Conversion of, By E. Bruce Levy. 2. (1): 37-51.
Rata the Killer. Bv
Plant Research, Scope of the Biologist in N.Z. Institutions, By Plymouth Marine Station, Life at, By Museums, N.Z., Biology in, By Cawthron Institute. By Sir Research Insitutions:
Biology for Australian Students (W. M. Curtis). By P. M. R. 2. (1): 52.Reviews:
Salamanders at Metamorphosis, Loss of Memory in, By B. M. B. 2. (1): 28.
Sea Cucumbers—A Guide to the Holothurians of N.Z., By
Sea Lilies—N.Z. Crinoids, By H. B. Fell. 3. (2): 78-85.
Sea Urchins, Key to the N.Z., By H., B. Fell. 1. (3): 6-13.
Sea Urchins. Key to the N.Z., Additional Species, By H. B. Fell. 3. (1): 42.
Shags, Identification of N.Z. Mainland, By
Shakespearean Entomology. By
Soil Conservation, The Place of the Botanist in, By A.
Soil Organic Cycle, Recent Researches on the, By
Sphenodon punctatus—The Tuatara. By
Spiders of the Wellington District, A Key to the Common, By
Starfish—Key to the Littoral Asteroids of N.Z. By H. B. Fell. 1. (1): 20-23.
Statistical Method, Some Basic Ideas in, By
Swell Sharks—Inflation of the Abdomen in Cephaloscyllium, By
Timber, Seasoned, Biology and Control of Beetles attacking, By
Tuatara, The, By
Tuatara—“To A Tuatara Alive in My Hand.” By
Whaling Station. Biological Interests at a, By
Woodlice—N.Z. Terrestrial Isopods. By
Vertebrate Palaeontology in N.Z. By
Zoological Specimens (Marine Invertebrates), Collection and Preservation of, by