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In the following brief survey of vertebrate palaeontology in New Zealand no account will be given of the Pleistocene specimens. It is possible only to give a general survey as in many cases more work is required before the group is properly known, but an attempt is made to indicate the principal literature of the subject and to give an idea of the vertebrate groups represented by fossils in this country.
The fossil fishes were dealt with as a whole by Chapman in 1918, and many teleosts later by Frost. These papers list the species and their localities and it is unnecessary to do so here. Species appear to have been described only from the Cretaceous and Tertiary. Of the Selachii Chapman lists 7 species from the Cretaceous, 23 from the Tertiary and 5 common to both. These are represented by isolated teeth, which are not uncommon in rocks of Upper Cretaceous and Tertiary age. The centra of vertebrae and fin spines also occur occasionally. There are remains of 2 Holocephali, one a species of Callorhynchus, both from the Upper Cretaceous of Amuri Bluff.
Most of the Teleosts are described by Frost from otoliths, and come from beds of Early Tertiary age. He describes 44 species, 29 of which are new. Only two Teleosts seem to have been recorded from rocks older than the Tertiary. Chapman mentions scales from the Upper Cretaceous of Amuri Bluff and the Clarence Valley, Marlborough, which he attributes to ? Thrissopater sp., and a specimen of Diplomystus coverhamensis, also from the Clarence Valley. Two fossil fishes from the Abbotsford Mudstone and the Burnside Marl have been described by Chapman, and several specimens have been collected from later rocks in Frazer's Gully, also in the Dunedin district. The tail of a large Tunny from the Oliogocene limestone at Duntroon, N. Otago, and the bones of another large fish from the greensand of the Hakataramea Valley are in the Otago Museum. They have not yet been described.
Quite a number of reptilian fossils have been collected in the Mesozoic rocks in various parts of the South Island, mostly from the Upper Cretaceous. There are some tantalisingly brief references in the literature to “saurian bones,” sometimes in older rocks, but specimens are not available, and ideas as to the age of the rocks from which they came may have changed with subsequent stratigraphical research. Park refers to Ichthyosaurus bones from Mt. St. Mary, near Kurow in N. Otago, and from Mt. Potts in Canterbury. These were said to be from Triassic beds, which would be an early date for an Ichthyo-
saurusMyopterygius from the Upper Cretaceous of New Zealand, but I have not found his authority for this statement.
The majority of the reptiles come from the Upper Cretaceous of the Amuri, Cheviot and Waipara districts in North Canterbury. First discovered by Hood in 1861, specimens were sent to Owen who described 3 species of Plesiosaurus. In 1868 Hood made another collection which was lost by shipwreck on the way to England. Haast examined the specimens before they were sent and mentions that one block contained the major portion of a skull at least three feet long. He also mentions that there were specimens, from the small seams of brown coal in the district, which consisted of a procoelous vertebra and part of a femur which he thought to be of terrestrial type and compared to that of Iguanodon! Further collections were made in the same locality, and also at Amuri Bluff, and several tons of rock containing bones were taken to the Colonial (now the Dominion) Museum and to the Canterbury Museum. These were described by Hector, who states that “portions of 43 individual reptiles, mostly of gigantic size and all of aquatic habits, and belonging to at least 13 distinct species, have been discovered.” He lists 6 species of Plesiosaurus, 1 Polycotylus and 2 Mauisaurus, all Sauropterygians, while there were 2 Mosasaurs, Liodon and Taniwhasaurus. There were also two vertebrae which he thought might be Crocodilus.
As no collecting seems to have been done for some 70 years, the writer recently visited the Waipara locality to see if more material had weathered out. The rivers here have cut deep canyons through the soft deposits, the steep sides of which are overgrown with beech trees. In the soft rock are numerous spherical concretions, up to five or six feet in diameter. They project from the cliffs and lie scattered along the bed of the streams and form a most extraordinary sight. Bones were observed in some of them, and in one case a concretion which had split in half exposed a small bone at its centre apparently forming the nucleus around which the concretion had grown. The concretions are extremely hard, and vigorous work with a pick produced little result other than a few sparks. Only one humerus of Mauisaurus, a bone some 13 inches long with a head 5 inches in diameter, and a few fragments were collected after several hours of effort.
All these reptiles are from Mesozoic rocks. The only Tertiary fossil reptiles appear to be some undescribed turtle bones in the Canterbury Museum.
The first fossil penguin bone to be discovered, the tarsus of Palaeeudyptes was found near Oamaru in 1857, and New Zealand is one of the only four localities where fossil penguins are found. One specimen is known from Australia, and many from Patagonia and from Seymour Island off the east coast of Graham Land.
The original tarsus was described by Huxley. In 1868 a number of bones, but without a tarsus, were collected near Brighton, Nelson, and attributed by Hector to the same species. About the same time, bones of two other penguins were collected near Oamaru and later described by Oliver as species of a new genus, Pachydyptes. Recently a considerable number of specimens have been collected, mostly in the greensand at Duntroon, N. Otago, but also in the Burnside Marl, Dunedin, and elsewhere. They have been described, and all the fossil penguins of New Zealand reviewed, by Marples (in M.S.S.).
So far fossil penguins have been discovered only in the South Island. One comes from Nelson, some from North Canterbury, South Canterbury near Waima'e and the Hakataramea Valley, North Otago, especially Duntroon and the Oamaru and Kakanui regions, and at Burnside near Dunedin. The oldest specimen is a fragmentary femur from Cheviot, which is at least as old as the Heretaungan stage of the Lower Eocene, while Pachydyptes comes from the Runangan stage of the Upper Eocene. Most of the specimens, assigned to the genera Palaeeudyptes, Platydyptes, Archaeospheniscus and Duntroonornis, come from the Duntroonian and Waitakian beds of the Lower and Middle Oligocene.
The fossil penguins of other countries are all assigned to the Miocene. It is interesting to notice that there are a number of differences between the fossil penguins of New Zealand and the Recent species, and that the Patagonian specimens are intermediate in structure. The Seymour Island and Australian specimens on the other hand are similar to the New Zealand ones. Even these Early Tertiary forms are fully specialised penguins, and the origin of this group from their flying ancestors must have taken place a very long time before the Oligocene. This is, of course, what would be expected from a consideration of the evolutionary history of birds in general, which does not show any important changes during Tertiary times.
Most of the species were large birds and some must have stood about five feet high. At least five genera and six species seem to be represented. Many of the specimens include a number of bones belonging to the same individual, which is important as most of the specimens found in other countries consist of isolated bones.
Flying birds as well as penguins existed in the Early Tertiary of New Zealand, but only fragments have so far been discovered. A clavicle from Duntroon, resembling that of an albatross, was described by Marples as Manu antiquus.
With the exception of the bones of a seal, said by Park to have been collected in the Burnside Marl, Dunedin, but which were never described and have been lost, the mammalian fossils are all cetacean. All the three subdivisions of the order are represented.
In 1880 McKay collected at Wharekuri, near Kurow in North Otago, a number of whale bones among which were some very large teeth of the Archaeocete Kakenodon. Other bones were attributed to
Kakenodon bones exist. Benham has published several papers recently on fossil whales, but there are still specimens which require re-examination, and some in the museums seem to have been lost. A natural endocranial cast in the Otago Museum appears to belong to an Archaeocete (Marples) but its locality is unknown.
Bones of Squalodonts have been found in the Oligocene rocks in various parts of the South Island, and were described by Benham as belonging to the genera Prosqualodon, Microcetus and Tangaroasaurus. The last, a fragment of a jaw, was originally mistaken for that of a reptile. Some well preserved skulls and other bones are in the Otago Museum, also a natural endocranial cast probably belonging to a young Squalodont. Fragments of several individuals have been found recently at Duntroon.
Benham also described two fragments of skulls, preserved in Dunedin, and several other bones all from the Oligocene of Otago, under the name Mauicetus parki. This is one of the Cetotheres, the most primitive group of the whalebone whales. Previously they have only been known from Miocene rocks, so that Mauicetus from the Lower Oligocene is much the oldest known member of this group. Recently a more complete skull and several bones of the same individual, as well as portions of other individuals have been discovered at Duntroon in the same Lower Oligocene beds as the fossil penguins. They have not yet been developed and described but they appear to resemble Mauicetus and may represent more than one species.
Before discussing some of the lines of cytological and genetical research which might be carried out in New Zealand, I will describe very briefly some of the types of studies which are now being carried out overseas.
In 1926, twenty-six years after the re-discovery of Mendel's paper, T. H. Morgan summed up knowledge at that time, of heredity and its physical basis in his book “The Theory of the Gene.” He noted that the characters of an individual are referable to genes which occur in a linear order on the chromosomes, and that the genes on each chromosome are held together in a linkage group. In diploids, homologous* chromosomes and their genes are paired, and each gamete contains only one of a pair, thus satisfying Mendel's first law. If genes are on different chromosomes they are assorted independently at game-togenesis, upholding Mendel's second law. Interchange of genes may occur between homologous chromosomes, and as the frequency of this interchange is roughly proportional to the distance between genes, a method was available of making linkage maps.
Research has proceeded in several directions since 1926. Genetic research on diploids has been extended to polyploids*. The statisticians have developed techniques for obtaining the maximum amount of information from a set of figures, as for example, in a linkage experiment, or for estimating the frequency of a gene in a population and the extent of its fluctuation due to selective influences. Refinements of technique continue to appear, but the development of new methods is usually beyond the power of the biologist, though he will naturally keep acquainted with the tools that the mathematician produces for his use.
R. A. Fisher in 1928 suggested that modifying genes were of great importance in the evolution of dominance and this stimulated research into the reaction of the same gene in different genetic environments. A great deal of research in this field has been done on cotton. The recognition that the phenotypic* effect of a main gene can change gradually this way or that under the influence of different modifying complexes, has given valuable support to the Darwinian theory that species evolved gradually by the accumulation of slight changes which have a selective advantage.
Further support for the Darwinian theory has been given by Mather from his recent researches on polygenes in Drosophila melanogaster. The polygenes occur in large numbers on all the chromosomes, even on the so-called “inert” heterochromatic Y chromosome. They
A full knowledge of the theory of polygenes is of great importance to plant and animal breeders, as most characters of commercial value are under the control of large numbers of genes each with a small effect.
The Ascomycete genus, Neurospora, has come to the fore in genetical research in recent years. It has been used to study the way in which genes control the synthesis of chemical substances. Its merit is that immediately after fertilisation an “ordered” ascus is formed consisting of 8 spores in a row. These may be dissected out in order and on germination tested for the mutant under consideration. The relative position of mutant and normal spores in the ascus show whether or not crossing over has occurred in the meiotic division. Irradiation with ultra-violet light induces biochemical mutants in Neurospora. They will grow only on a complete medium containing all the elaborated substances they could possibly need, and will not grow on a basic medium as they lack the power to synthesise some substance. This substance is identified by adding, in turn, all known vitamins and all known amino-acids to the basic medium until the spores germinate. Thus the mutant “lysineless” will not grow unless lysine is added to the basic medium (showing that the normal allele* of lysineless takes part in lysine synthesis). In the United States it has been shown that at least 7 genes control the synthesis of the amino-acid arginine, the first 4 building up ornithine, the next 2 changing that to citrulline and the last changing citrulline to arginine.
Genetical studies in other simple organisms are appearing in increasing numbers in the journals. Mutants in bacteria are identified by their resistance to various types of bacteriophage, while in America there are several workers who specialise in the genetics of Paramoecium.
Genetical research on micro-organisms is a specialised branch of a specialist subject and would be difficult to start as yet in New Zealand Universities. Such research can be better carried out elsewhere.
Before talking about work in New Zealand just a word about developments in cytology.
In 1926 cytology was not in as good a position as genetics. What happened during prophase of meiosis was still rather a mystery. Such a fundamental point as to whether the chromosomes paired end to end (telosynapsis) or along their whole length (parasynapsis) was not yet settled. However, during the next ten years the position improved rapidly, due mainly to the brilliant work of C. D. Darlington who
The first step in cytological and evolutionary studies in the N.Z. flora would be a chromosome survey, particularly of the grasses and herbs, one of the aims being to find easily cultivatible genera with large chromosomes in small numbers. At the moment the chromosome numbers of only about 60 species of N Z. plants are known, due mainly to the work of Dr. O. H. Frankel and Mr. J. B. Hair, of the Wheat Research Institute, Christchurch. Species (numbers in brackets) have been studied in the following genera:
Of these, two would be well worth further study. Chrysobactron Hookerii has 14 large chromosomes ranging from 10 to 18mu long; several have distinctive morphology due to their length, non-staining constrictions and position of the centromere. If plants from widely separated populations were crossed, studies of meiosis in the hybrid might reveal the beginning of structural differentiation between chromosomes from different parts of the species area. Clematis indivisa has 16 quite large chromosomes, and I have recently found that C. colensoi has the same number. A chromosome survey of the rest of the
Clematis may aid taxonomists, and it would be interesting to search for an XY sex-determining mechanism as the species are dioecious.
Chromosome surveys are of great use to the taxonomist, and the cytologist should keep pressed specimens of the plants which he examines. Chromosome surveys may support groupings that the taxonomist has already made. Thus Cockayne and Allan (1926) created two varieties of Hebe vernicosa, var.
On the other hand evidence from chromosome numbers may lead the taxonomist to revise his work. A type example comes from work done in N.Z. on the closely related genera Hebe and Veronica (Frankel, 1941). The majority of N.Z. species were placed by Cockayne and Allan in Hebe, while a few were included in Veronica. Frankel and Hair (1937) found that both N.Z. Hebes and Veronicas had basic numbers of 20 or 21, where as the Veronicas of the Northern Hemisphere had basic numbers of 4, 8, 7, 9, or 17. This showed that the separation of Hebe from Veronica by Pennel in 1921 had been justified, and it also suggested that N.Z. Veronicas were closer to Hebe than to the Northern Hemisphere Veronicas. Allan (1939), on re-examining the capsule dehiscence in N.Z. Veronicas found that it was essentially the same as in Hebe and therefore transferred them to the latter genus.
Again Calder's study of chromosome numbers in the N.Z. Danthonias showed that Danthonia semiannularis had 48 chromosomes, but that both D. semiannularis var. setifolia and D. semiannularis var. nigricans had only 24. The difference in chromosome number implies that species and variety are effectively isolated as a hybrid between them would probably be sterile. This fact, together with definite morphological differences, justified him in making two new species, D. setifolia and D. nigricans.
Chromosome numbers also indicate whether an inter-specific cross is likely to be fertile or sterile. Mr. Hair tells me that one of our Agropyron species has 42 chromosomes, and another has 28. A hybrid between them will have 35 chromosomes and probably be sterile.
Enough has been said to show the importance of a knowledge of chromosome numbers, and it is to be hoped that this gap in our knowledge of the N.Z. flora will be filled within the next few years. Of course, many other problems are often unearthed during a chromosome survey. Thus S. Smith-White (1948) in Sydney, while carrying
Leucopogon juniperinus was a triploid, and that contrary to most of the rules it was stable and reasonably fertile. The reason discovered for this is of great general interest.
The New Zealand flora should provide ideal material for studies in the evolution of species. The pattern of distribution in most of the genera has not been appreciably modified by man.
The beginnings of species differentiation might be sought for among the numerous cases of discontinuous distribution known within the country (see Wall, 1927). Individuals from two widely separated populations could be brought together in an experimental garden and studied for signs of cytological, taxonomical and, if possible, genetic divergence.
Ecotypes*, considered by many to be the first marked stage in species differentiation, have not been studied much in this country, possibly because of lack of experimental gardens. That such studies can produce interesting results has been shown in some work by Mr. H. Conner of the Botany Division, D.S.I.R., who has collected a number of distinct forms of Agropyron scabrum from both Islands. These have retained their distinctive characters when grown together under uniform conditions in Wellington.
On the species level, the hybrid swarms so characteristic of our flora show that many species, though morphologically distinct, are not yet differentiated enough to be inter-sterile. Cytological studies of F1 hybrids would show the nature of the chromosome differences between the species. Genetical studies would be difficult in some of the most interesting hybrids. The theory is that an estimation of the number of genes controlling, for example, the leaf shape difference between two species can be obtained by selfing the F1 hybrid and noting the proportion of original parent leaf types obtained in the progeny. Leaf shape and most other characters would doubtless be under the control of several genes, and to re-obtain the parent types many hundreds of F2 plants would need to be cultivated. In the case of shrubs, this would require much garden space for several years, until the plants matured. Other difficulties arise. In attempting to self-pollinate Corokia buddleoides X cotoneaster by bagging last season, no seed was set, though on open pollination it is usually prolific. It may be self sterile, or else bagging does not agree with it. The only chance of getting an F1 intercross would be to find two hybrids growing close together in an isolated garden, and let them cross-pollinate naturally. However, there is much scope for work on the synthesis of hybrids. Many of our puzzling plants have been called natural hybrids because of very strong circumstantial evidence. The forms are intermediate between two good species, and only occur when these species grow in proximity. The final experiment to clinch the argument is to synthesise the hybrid from the supposed parents.
A genus with its main representation in New Zealand could be the subject of an evolutionary study in the style of that of Babcock and others on the genus Crepis (Babcock, 1947). This genus has been under study for about thirty years. Some 113 species have been brought into cultivation at Berkley, California, and have been studied by taxonomists and cytologists, and the pattern of its evolutionary history is now fairly complete. Crepis has evolved from primitive perennial rhizomatous types with large flowers, leaves and achenes to more advanced tap-rooted annuals with small leaves and flowers and with small beaked achenes. The centre of origin was probably the northern part of central Asia, whence it has migrated over the Northern Hemisphere. Chromosome complements have evolved by decrease in basic numbers from 6 to 5, 4 and 3; by polyploidy; by increase in asymmetry*, and by decrease in size.
I had thought that the genus Celmisia might be studied in this manner. There is a large number of species, all but one of them confined to New Zealand; there is a good range of form, and the distribution of species is interesting, for example, a number of endemic species confined to the Nelson mountains. However it may prove very difficult to determine chromosome numbers accurately in this genus. In three species that I have studied, two had chromosome numbers between 80-90, and one about 120. This is very different from, say, Crepis capillaris with 6 chromosomes.
The breeding systems of plants is another important topic. Are some of our polymorphic species variable because they are obligately cross-fertile, thus continually reshuffling genetic material, or do they consist of a number of apomictic* strains each of which will inevitably breed true?
It should also be remembered that N.Z. Universities have quite a good tradition of research in plant anatomy. This could be used in collaboration with a geneticist in studying the control of plant form by genes. This branch of genetics is developing, and reference should be made to the second Symposium of the Society for Experimental Biology which deals with Growth and Differentiation (1948).
Cytological and genetical work on natural populations of Drosophila could be carried out in New Zealand, but first the taxonomy of the species would need to be clarified. Dr. Frankel has pointed out to me the good field for cytological research in insects here, and I have recently found that a male weta (species not yet determined) has 19 pairs of quite large chromosomes which are easy to study.
A prominent broadcast speaker on wild life made a statement recently that many of the details of the life history of Notornis were still unknown. This is, perhaps true of some aspects but certainly not true as the speaker claimed in respect to details of egg, nest and young chick, all of which were seen by the official party in January last. These findings have been published by Falla in the “Emu,” Vol. 48, 1949 to which journal the reader is referred for a full account should he desire further information.
Another point mentioned by Dr. Falla is on the nomenclature. Today the bird seems to be known as Notornis hochstetteri. The description of the living bird included a few small differences from the sub-fossil remains which have been named Notornis mantelli. Whether the species are really separable remains to be decided.
As the usual form of identification key would be of value only to those who are concerned with handling museum specimens and dead birds, the accompanying sketches have been prepared as an aid to distinguishing in life seven of the eight species of shags or cormorants to be found in New Zealand, including Stewart Island. A further six not figured here occur in the Chathams and the subantarctic.
The superficial similarity of shags disguises many differences of structure and habit; while age and seasonal plumage phases may lead to further confusion. These are mentioned in the following notes. Distribution and habitat are also important guides. As a group, those species frequenting inland waters and extending their range to estuaries and bays all have black feet. The oceanic shags, which never venture into fresh water, all have pink or yellow feet.
1. Specifically separated by most authors as Little Shag (Phalacrocorax brevirostris), variously called “White-throated,” “Little Pied,” or “Frilled.” The diagnostic features are short yellow bill and long tail. The plumage pattern is not uniform: the commonest form when adult is black with a white throat (1 c), and at the other extreme is the “little pied” plumage (1 a). Most young in their first plumage are entirely black (1 b), but the extreme “little pied” formP. melanoleucus.
Distribution: Lakes, rivers and sheltered coasts throughout New Zealand.
2. Little Black Shag. (P. sulcirostris). Distinguished from the young of the last by its longer bill, which is also dark, and its shorter tail. Feet black.
Distribution: Lakes, rivers, and estuaries of Auckland isthmus south to Hawke's Bay.
3. Pied Shag. (P. varius.) Larger than the last two, the male being nearly as large as a black shag. The bill is pale, and facial skin yellow, blue, and dull purple. Immature birds have white underparts slightly mottled with dark. Feet black.
Distribution: Warmer coastal waters and estuaries of Auckland, Bay of Plenty, East Coast, Marlborough, Nelson, North Canterbury, and Stewart Island.
4. Black Shag. (P. carbo). The largest inland shag, with a pale bill, and yellowish about the face. White chin, crests, a whitish mane, and white thighs are breeding ornaments only and are not always present. Immature birds have the underparts mottled with white. Feet black.
Distribution: Lakes, rivers, estuaries, and coastline throughout New Zealand.
5. Spotted Shag. (P. punctatus). A slenderly-built sea shag with pale-grey plumage except for oil-green back and thighs. Bright facial colours are developed in the breeding season; double crests, and pure-white feathers form a neck stripe and sprout from the darker areas of the plumage. The young are pale-grey below, and darker above. Feet, cream to yellow.
Distribution: Rocky coasts of both islands. The Stewart Island form, with less white, is known as the Blue Shag (P. steadi).
6. Rough-faced or King Shag. (P. carunculatus). The largest of the subantarctic shags, which are alike in having steel-blue necks and backs and oil-green wings. They have a cluster of orange pimples at the base of the bill, and blue eyelids. Young have the dark plumage dull brown. Feet pink.
Distribution: Western Cook Strait only, and not common.
7. Stewart Island Shag (P. chalconotus). Two distinct plumage forms (7a and 7b) occur, and they interbreed. The glossy dark bird is unmistakeable; and the white-breasted one is not unlike the King Shag. They differ in being smaller, having red throat pouches, and developing crests in the breeding season. Feet pink.
Distribution: Coasts of Stewart Island and Ruapuke. A larger form of the same bird inhabits Otago Peninsula. These birds are entirely marine. Superficially, the pied form could be confused with the Pied Shag (P. varius), and the dark form with the Black Shag (P. carbo).
Identification of shags is a matter of practical importance because while many are on the schedule of absolutely protected birds they are somewhat indiscriminately subject to shooting and nest raiding because they are fish-eaters. The basic falsity of the assumption that any predator effects serious reduction of its own food supply has long been revealed for what it is worth, but it still influences the views of gamekeepers, fish culturists, and some commercial fishermen.
The official attitude, maintaining a reasonable balance on available evidence, accords protection to all the purely marine cormorants; although prosecutions are infrequent except when heads are submitted for bounty payable on some river shags. With the exception of the Little Black Shag (P. sulcirostris), the estuarine and river species are not protected, and a good deal of lively controversy centres round the relationship of the Black Shag to stocks of introduced trout. Although the late
A good deal more investigation is required, backed by a good knowledge of birds, fish, and stream ecology. Some attempt at presenting evidence that can be checked has been made by Falla and Stokell (T.R.S.N.Z. vol. 74, 1945); but the New Zealand standard of investigation has not yet reached the level at which similar work has been carried out in the U.S.A., Canada, and Australia.
Australian investigations, because they deal with shag species also found in New Zealand, are perhaps of more immediate interest. Dealing with conditions on an estuary—the Swan River—D. L. Serventy (The Feeding habits of Cormorants and South-western Australia. Emu, 1938, p. 293) examined a total of 441 shags of four species in one year. His results showed the effect of the Pied Shag on commercially important fishes is negligible, the Little Pied is still more harmless, the Little Black Shag likewise harmless; the Black Shag alone being shown to include an appreciable percentage of marketable fishes in its diet. In Victoria, G. Mack (Cormorants and the Gippsland Lakes Fishery. Mem. Nat. Mus, Vic. LX, 1941, p. 95) found no evidence from stomach analyses that shags were detrimental to stocks of commercial or sporting fishes in that area. In New South Wales, K. McKeown (The Food of Cormorants and other Fish-eating Birds. Emu. 1944, p. 259) has published data which, as far as they go, confirm the findings from other States.
It may be concluded, on the basis of available evidence that no native marine or estuarine fisheries in New Zealand are adversely affected by the presence of a population of Cormorants. The problem seems to be reducible to the single issue of the black shag in relation to introduced trout. The whole of the fresh water system of the country has been stocked, or there have been attempts to stock it, with introduced salmonids. To those whose idea of adequate trout population really requires an artificial surplus to satisfy it, shag depredations can never be regarded with equanimity. But it should be possible to contemplate without alarm a state of affairs, and surely a desirable one, in which the growth of trout food keeps pace with the needs of a trout population, and the trout keep pace with their predators. The predatory relationship is not necessarily a destructive one and it has yet to be shown whether the level at which black shags affect trout has an adverse or a beneficial effect on the stock.
Approximately 180 species of echinoderms are now known from New Zealand, of which 71 belong to the Class Ophiuroidea, or brittle-stars. These are among the most numerous of all the bottom-dwelling animals of the seas. Most are scavengers, and as such play an important part in the bionomics of the ocean. Near the outlet of the fish-oil factory at Island Bay and where the local fishermen deposit offal, the number of individual ophiuroids must run into many thousands, even within the limits of “Fisherman's Creek” alone. Some 41 species are known to inhabit the littoral zone around New Zealand; the others occurring in deep water are not included in the following key. Colour notes are still required for some species, and in this field local students could do useful work. Some of the colour details given here have not previously been published. The relatively high proportion of species of Amphiuridae is a characteristic of the New Zealand fauna, not paralleled elsewhere.
No glossary is given for the technical terms which have had to be used in the key. The meaning of specialized terms should be sufficiently clear from the diagrams which follow. Before attempting identification, allow specimens to dry out, as this renders the skeletal plates easier to see. When counting oral papillae, include any infradental papillae in the total.
(The figures are not drawn to scale—where size is of diagnostic significance, the dimensions are given in the key.)
Abbreviations: AD. adoral plate; O.P. oral papilla; OR. oral plate; O.S. oral shield; R.S. radial shield.
Abbreviations: A.D., adoral plate; D.P., dental papillae; I.D.P., infradental papillae; LP., lateral armplate; O.P., oral papilla; OR., oral plate; O.S., oral shield; SC.SP., scale-like spines; SP, spine; SP.P., supplementary plate; T., tooth, teeth; T.F., tube-foot; T.S., tentacle-scale; U.P., upper armplate; V.P., lower armplate.
As stated in the “Guide to the Brachyrhynchous Crabs” the systematic standing of many species of our Brachyura is uncertain. The publication of the “guides” has revived interest in the group brought additional specimens, information and literature to hand so that already corrections can be made which promise systematic stability for some of our species. The following changes should be noted in their proper place in the “guides”:
F. Portunidae. Portunus borradailei (p.31) is now being accepted as a member of the genus Liocarcinus and should be named Liocarcinus borradailei.
F. Cancridae. Balss has shown that Heterozius rotundifrons (p.32) is not essentially a member of this family, but is more properly included in the Xanthidae. Mr. J. Morton has kindly supplied specimens from Auckland which prove to be Pilumnopeus serratifrons on comparison with named specimens sent me from the Australian Museum through the courtesy of Mr. F. McNeill. This is the first positive record since 1876. P. serratifrons has the palate ridged, the fronto-orbital border just more than half the greatest width of the carapace and so fits in the key alongside O. truncatus (p.32) from which it is distinct in having three well-developed sharp-edged anterolateral lobes, a sharp spine on the wrist, and a strong curved sharp-edged tooth on the dorsal margin of the land. The fingers are heavy and black.
F. Grapsidae. Dr. Isabella Gordon of the British Museum has written me that there can be no certainty of the correct application of the name Plagusia chabrus and this name is lapsing. Our Plagusia (p.32) is better recognised now as P. capensis. Dr. Gordon has kindly re-examined for me material of the genera Brachynotus and Hemigrapsus, supports Rathbun in maintaining the distinctness of the genera, and has demonstrated to me what our Hemigrapsus sexdentatus (p.34) cannot by priority retain its specific name and must now be known as H. edwardsii as proposed by Hilgendorf.
F. Hymenosomidae. I am indebted further to Dr. Gordon for comparing specimens of Halicarcinus with material in the British Museum. It is now clear that the key must be corrected since the H. ovatus (p.68) in the key is actually H. planatus, and as I had suspected, the species listed as H. planatus in the key is a new species for which I propose the name of H. innominata and which I will describe in full elsewhere. Mr. Forster of the Canterbury Museum informs me that the types of Chilton's H. marmoratus are no longer available. The species referred to as 15 (Fig. 45) in the key (p.67) agrees well with Tesch's description of H. edwardsi and in my experience is characterised by the heavily furred hand of the male. Tesch has synonymised E. quoyi (p.67) with H. pubescens but without reference to actual material.
Mosses together with the liverworts form a very natural and usually distinct group, the Bryophytes. The mosses, especially, can usually be placed as such in the field and are easily collected, requiring only drying and packeting, as they moisten out well for later examination. Their main drawback is their small size, some being only a millimetre or two high, even when in fruit, so that a microscope is required for their examination, as also for the minute structure of leaves and peristomes of the larger species, some of which reach the unusual size for mosses of 1 ½ feet or more.
Mosses are widely spread in New Zealand, ranging from mountain rocks to coastal sands. We have our share, too, of very interesting species, and a high endemism, given as nearly 40 per cent, by Martin, but, of course, there is the usual cosmopolitan element.
Some of our mosses were included in Dr. W.
Various collections were made after this, notably by T.W.N. Beckett; by
H. N. Dixon, however, laid the foundation for an up to date Manual of our mosses with his “Studies in the Bryology of New Zealand” which was published in six parts, 1913-1929, by the N.Z. Institute as Bulletin No. 3. In this Dixon collected and critically examined all records of our mosses and brought the nomenclature up to date. He credited us with under 500 mosses, grouped into about 170 genera. This work revived interest in the mosses and a small keen band of collectors has since carried out extensive field work which it is hoped will make the compilation of a manual possible.
At present no such recent work exists but students will find the following works useful:
Dixon's work mentioned above, which has six plates and mentions all the mosses then known to grow in New Zealand, but, while discussing critical species in detail, it often gives no description of the well known species. Dixon's “Student's Handbook of British Mosses,” 1924, is an excellent book so far as it deals with our genera. W. Martin published a check list of our mosses with much other information in Volume 76 (1946) of the Transactions of the Royal Society and also a good popular account of our mosses with several plates in his “Flora of New Zealand” (3rd edition).
Mosses are divided into three sub-classes, based on the structure and development of the capsule: (i) Sphagnales (ii) Andreaeales and (iii) Bryales, the first two containing only one genus each. The Bryales are divided up into two clans, based on peristome characteristics: the Nematodonteae have solid teeth, not tranversely barred, derived from several concentric layers of the cells of the Sporogonium and include seven families: the Arthrodonteae have the teeth (sometimes absent) thin and membranous, derived from a single layer of cells of the sporogonium, and are transversely barred and articulate and include the rest of the mosses—some 67 families. These latter are again divided up into 3 Sub-clans on peristome characteristics, so that it can be seen what an important part the capsule plays in the classification of mosses and it should be collected whenever possible. The Sub-clans are divided into orders, families and genera on characters taken from the gametophyte as well as from the sporophyte.
The main divisions of mosses can be told as follows:
Sphagnales: White or whitish green mosses with fascicled or whorled branches, growing in bogs or wet ground. Leaves of very narrow long cells which anastomose to form a network of large wide thin-walled cells which are often strengthened with evident spiral thickenings: operculum falling: no peristome (Figs. 17, 18) Sphagnum.
Andreaeales: Small reddish to dark purple or black mosses usually growing on alpine rocks, very occasionally larger and growing in mountain streams. Leaves inserted all round the stem, cells dot like. Capsule black, erect, dehiscing by 4 lateral slits, the operculum never being shed. Andreaea.
Bryales: The operculum is usually shed, though in exceptional cases this is not so and then the capsule wall rots or breaks irregularily (not by regular slits) to release the spores. Includes all the other mosses.
Miss Adams' contribution in supplying the drawings is gratefully acknowledged.
The Key below gives characteristics for 123 of the Commonest Genera of New Zealand Mosses out of the total of about 170.
Transverse section of leaf of Polytrichum commune showing lamellae on the upper surface.
Leaf of Dicnemoloma showing papillose lower surface.
Leaf of Leptobryum pyriforme.
Leaf of Pterigophyllum dentatum with forked nerve and denticulate margin.
Leaf of Tortula princeps showing the nerve excurrent in a long dentate, flexuose, piliferous point.
Leaf of Hypnum cupressiforme, strongly curved and nerveless.
Leaf of Acrocladium auriculatum with short nerve and conspicuous alar areas.
Leaf of Fissidens leptocladus showing the nerve reaching the apex, vaginant leaf base (stippled) and thickened margins.
Leaf of Polytrichum commune showing dentate margins and upper surface opaque owing to the numerous longitudinal lamellae.
Leaf of Calyptopogon mniodes showing the apical mass of gemmae on the upper surface.
Leaf of Catharomnion ciliatum showing the remarkable ciliation.
Leaf of Hymenodon piliferus showing piliferous apex and nerve not reaching the apex.
Leaf of Rhizogonium mnioides with thickened, dentate margins.
Leaf of Cladomnion ericoides, nerveless, plicate and with recurved apiculus.
Leaf of Ctenidium pubescens showing the margins dentate almost to the base.
Leaf of Dicranoloma dicarpum with strongly spinulose upper margins.
Branch leaf of Sphagnum cuspidatum.
Sphagnum: part of a branch leaf surface, strongly magnified to show the long chlorophyllose cells enclosing the larger hyaline ones which are strengthened with “spirals” or cell wall thickenings.
Capsule of Polytrichum juniperinum and front view of its mouth showing the (ruptured) circular membrane held in place by the 64 short teeth.
Capsule of Tortula princeps, erect, with twisted peristome rising from the entire basal part.
Capsule of Ceratodon purpureus, inclined and ribbed.
Capsule of Bryum truncorum, pendulous with conical operculum.
Capsule of Weisia viridula, erect, with short peristome teeth.
Capsule of Conostomum pusillum, striate and horizontal.
Capsule of Polytrichadelphus magellanicus with operculum.
Capsule of Trematodon suberectus, inclined with long neck.
Capsule of Hypnum cupressiforme, inclined and with conicomamillate operculum.
Capsule of Ptychomnion aciculare, inclined and striate with the strikinly long operculum.
Capsule of Leptobryum pyriforme.
Capsule of Orthotrichum hortense, erect and ribbed with short, recurved outer peristome. The inner peristome consists of 8 filiform processes which remain more or less horizontal across the mouth, but may be soon lost.
(a) Archephemeropsis trentepohlioides, part of tuft with young capsules and male flowers. (b) Enlarged piece showing male flowers.
Fissidens leptocladus showing terminal seta.
Cyathophorum bulbosum, ventral side, showing the three rows of leaves and capsules.
Hypopterygium setigerum showing umbrella-like growth habit.
Pterigophyllum dentatum showing leaves inserted in several rows but spreading laterally to form a flattened “frond”.
Hedwigia albicans with immersed capsules.
Leptobryum pyriforme.
Macromitrium longipes.
Physcomitrium conicum with gymnostomous capsule.
Thuidium furfurosum with bi-pinnate branching.
Weymouthia mollis showing irregular or sub-pinnate branching.
Dicranoloma fasciatum showing the perichaetial leaves sheathing the whole seta.
Tetraphidopsis pusillus showing terminal heads of gemmae.
Hypnum cupressiforme with the densely placed curved leaves giving the stems a turgid appearance.
Fringed cucullate calyptra of Campylopus torquatus.
Entire mitriform calyptra of Cyathophorum bulbosum.