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Tuatara aims to stimulate and widen interest in the natural sciences in New Zealand, by publishing articles which (a), review recent advances of broad interest; or (b), give clear, illustrated, and readily understood keys to the identification of New Zealand plants and animals; or (c), relate New Zealand biological problems to a broader Pacific or Southern Hemisphere context. Authors are asked to explain any special terminology required by their topic. Address for contributions: Editor of Tuatara, c/o. Victoria University of Wellington, Box 196, Wellington, New Zealand. Enquiries about subscriptions or advertising should be sent to: Business Manager of Tuatara, c/o. Victoria University of Wellington, Box 196, Wellington, New Zealand.
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is the journal of the Biological Society, Victoria University of Wellington, New Zealand, and is published three times a year. Joint Editors:
Because of the Relative Ease with which pond organisms can be reared under laboratory conditions, the rearing of stream-dwelling organisms tends to be neglected. This article provides brief descriptions and a literature list of techniques used to rear stream organisms in the laboratory.
One difficulty in setting up an artificial stream is the transfer of stream fauna from their natural habitat to the laboratory. The death of fauna during transfer is caused usually by an inadequate oxygen supply and/or rapid increases in temperature.
The oxygen supply can be maintained at an adequate level if a small volume of water with a large surface area is continually agitated. Rapid agitation, however, may cause damage to delicate insect larvae. If large volumes of water are necessary for transport of fauna, an adequate supply of oxygen can be maintained by using an air bubbler. It is possible to construct a simple air bubbler from a large plastic funnel, by attaching a bubbler tube to the neck and directing the open end of the funnel into the windstream created by the vehicular transport (Fig. A.). This method of aeration, however, rapidly raises the water to air temperature.
Temperature may be controlled by using large mouthed “Thermos” flasks during transport. Large volumes of material can be transported with good temperature control by using expanded polystyrene “Chillybins.”
Growths of algae for browsing insect larvae can be obtained by seeding the artificial stream with natural stream water. However, some species of Ephemeroptera larvae eat considerable amounts of algae and can easily eat all available food if large numbers of larvae are placed in the stream. In artificial streams where filtered tap water is used, food must be added for filter feeders. Yeast suspension has been used successfully by D. M. Woods (pers. comm.) to feed Simuliidae larvae.
Some town water supplies are highly toxic to many stream animals because of such additions and impurities as hypochlorite and heavy metal ions. Copper retaining screens were found to be highly toxic to Simuliidae larvae by Hartley (1955), who also mentions that other workers have experienced difficulties with contamination when the water supply was in contact with copper. D. M. Woods (pers. comm.) uses activated charcoal filters successfully to remove both hypochlorite and heavy metal ions. Because of possible contamination no metal should be in contact with the water contained in artificial streams.
Techniques for rearing stream organisms in the laboratory vary considerably in the way the currents are produced, but can be placed into two broad categories:—(1) Open Systems and (2) Closed Systems.
(1) Open Systems: These are the simplest types of artificial streams, but depend on the abundance and purity of local water supply. The simplest type of Open System stream splashes tap water over rocks and stones which have been brought to the laboratory with the organisms still attached. B. Mannheims (pers. comm.) has used this arrangement to rear Blepharoceridae larvae to adults. A similar arrangement uses a stream of water directed into jars or Petri dishes and allows the water to overflow to waste (Fig. B). Hartley (1955) and Carlsson (1962) used jars and
Water flowing rapidly down plastic troughs has been used by D. M. Woods to rear Simuliidae larvae, similarly concrete channels with glass observation ports have been used by Dorier and Vaillant (1954) to study the effect of current on numerous aquatic invertebrates. Large (approximately lin. internal bore) plastic tubing with the water flowing through has been used by Carlsson to rear Simuliidae. and by the writer to rear Chironomidae, Blepharoceridae and Simuliidae.
A glass aquarium with a jet of water directed against one side, below the water line, provides a good basis for a general stream aquarium (Fig. D.). The depth of water, current and substrate can be varied with ease. Both Tonnoir and Carlsson used similar methods to rear Blepharoceridae and Simuliidae and the writer has reared Chironomidae, Blepharoceridae, Ephemeroptera and Trichoptera using this arrangement.
(2) Closed Systems: These types of artificial streams allow greater control of temperature, mineral nutrients, food and hydrogen ion concentration than Open System streams. Unfortunately, nearly all of the numerous methods of producing currents in these streams are completely dependent on electrical power.
By using a stirrer to produce a circular current in a jar (Fig. E.), Philipson (1953) was able to study the feeding habits of Trichoptera larvae. However, to investigate the effects of water flow and oxygen concentrations on 6 species of Trichoptera larvae, Philipson (1954) used jars with rotating glass stoppers which were lubricated with paraffin oil, the stirrers being attached to the inside centre of the stoppers. Whitford and Schumacher (1961) used a modified magnetic stirrer to study the effect of current on algal metabolism. In a similar method immobile plates were suspended within a rotating jar of water (Fig. F.). By oscillating jars in an horizontal plane a gentle circular current can be created. Freeden (1959) used these last two methods to rear certain species of Simuliidae larvae.
Air bubbled through water will produce sufficient turbulence or current to keep most stream organisms alive. However, Freeden reports that apparently the upper velocity limit of the water current
Recirculating apparatus usually consists of pump, a straight trough and a reservoir. The water is pumped from the reservoir either directly into the trough or firstly into an upper head-tank, and thence into the trough. Troughs in artificial streams of this type have been constructed from wood, earthenware, asbestos-fibre and perspex; self-priming centrifugal pumps appear to have been used rather than other types of pumps. Refrigeration units for temperature control are usually placed in the reservoir. Using apparatus of this type, Zahar (1951), Wright (1957) and Hall and Harrard (1963) have reared Simuliidae; Moore (1964), freshwater snails; Whitford, Dillard and Schumacher (1964), lotic organisms, and Lauff and Cummins (1964), have studied lotic ecology.
Apparatus based on the recirculation principle has been used to study the metabolism of running water organisms. In these cases the stream was completely enclosed and airtight. Odum and Hoskins (1957) have studied algal metabolism, Gaufin and Gaufin (1961) the effect of oxygen concentration on Trichoptera larvae, and Brett (1965) the metabolism of Canadian salmon. Sudia (1951) used an oval galvanised iron tank to study the effects of water flow on mosquito larvae. The current was produced by nozzles directing the water around the trough.
The writer has constructed an artificial stream of the circulation type incorporating many of the ideas from other workers (Fig. J.). The stream has been designed to provide a flexible piece of apparatus for use in as many fields of research as possible.
A fibre-glass centrifugal pump capable of delivering 700 gallons of water an hour, though not running to full capacity, is powered by a ⅓ h.p. intermittent-running electric motor. The water is pumped into a head-tank and is maintained at a set level by an off/on float switch which controls the motor and pump. A ball-cock valve which is connected to the tap-water supply becomes activated if the water in the head tank falls below a minimum level. This ensures a water flow in case of power failures or breakdown of the pump or motor. From the head-tank the water flows down large-bore plastic tubes into the stream portion of the apparatus,
This stream has performed without major trouble for approximately twelve months. Stream organisms reared successfully so far are algae, Hepaticae, Turbellaria, Gastropodia, Ephemeroptera, Plecoptera. Neuroptera, Trichoptera, Chironomidae, Blepharoceridae and Simuliidae. The stream has also been used to study the effects of current on various species of N.Z. Galaxidae fish.
Maritime ‘waifs and strays’ of the animal world not infrequently strand on New Zealand's shores; among them we find four species of marine turtles and two sea snakes. Whether these animals should be regarded as accidental visitors, i.e. fallouts from shoals bypassing New Zealand, or part of the maritime fauna of the country is a moot point. I favour the latter proposition, for if we accept the former then some of the pinnipeds and cetaceans would have to be excluded from the New Zealand fauna! Our records of marine turtles and snakes are based mainly on the dead and dying which strand on our shores and come to the notice of some interested person, and not on the possible, regular off shore visitations of the animals.
As both marine turtles and snakes are normally inhabitants of the warmer tropical and subtropical waters of the globe, it is natural that their visits to New Zealand should vary annually in response to the flow of the currents, and possibly also to the movement of their food supply. Their strandings may be due to sudden changes of temperature which can spell disaster to most repitilian life. Strandings are commonest on the northern shores of New Zealand, but there are also some on the southern shores; the latter animals may have reached there by way of the East Australian Current.
The order Testudines is divided into two main suborders, Athecae comprising a single family, genus and species, the Luth, Leathery Turtle or Leatherback; and the Thecophora embracing all other living species. In addition to the Luth, only the members of the family Cheloniidae, the marine turtles, concern us.
Four species of marine turtles have been recorded from New Zealand: Dermochelys coriacea (Linn.), the Luth; Chelonia mydas (Linn.), the Green Turtle; Eretmochelys imbricata (Linn.), the Hawksbill Turtle; and Caretta caretta (Linn.), the Loggerhead Turtle.
The carapace of the Luth is composed of hundreds of polygonal, irregular bony plates, the largest forming seven regular longitudinal keels along the dorsal surface; these keels converge posteriorly into the free projection of the carapace. The plastron is devoid of dermal bones except along five longitudinal ridges in which the bony series is incomplete. These ventral ridges are not always present in the adults.
Colour: The adult is dark-brown or bluish-black above, spotted or blotched with pale yellowish or pale bluish-white about the throat, flippers and on part of the carapace.
Size: The Luth is the largest of the living chelonians attaining a length of over two metres and an estimated weight of 1800 Ibs.
Distribution: The Luth is an inhabitant of tropical waters throughout the world. In New Zealand it appears as an occasional straggler, most frequently seen in northern waters, but also recorded as far south as Otago and Foveaux Strait.
Carapace with four pairs of costal shields, an intergular shield and a series of inframarginal plastral shields. Head covered with symmetrical shields; one pair of prefrontals. Dorsal shields juxtaposed. Jaws not hooked. Tail short.
Colour: Adult olive or brown, the shields with more or less distinct brown rays; yellow beneath.
Size: The carapace attains a length of 1,100 mm.
Distribution: The Green Turtle is worldwide in tropical and subtropical seas, and is an occasional visitor to New Zealand.
Note: The Green Turtle derives its name from the colour of its fat. It is the animal which provides the famous Turtle Soup of gourmets — the fat imparts its green colouring to the soup.
In the young the carapace is three-ridged and the dorsal shields are strongly imbricate; in the adult the ridges disappear and the scutes or shields may remain imbricate or become juxtaposed. There are twenty-five marginals, including the nuchal, strongly serrated posteriorly. Snout compressed in adults, jaws hooked (hence the popular name!. edges not or but feebly denticulated. Head with two pairs of prefrontal shields.
Colour: Young brown above, blackish below. Carapace of adult marbled yellowish and brown, plastron yellow; shields or scales of the head and limbs dark brown, margined with yellow.
Size: An adult carapace may attain a length of 900mm.
Distribution: Widely distributed in tropical and sub-tropical seas. In New Zealand the species is almost restricted to the north. Young occasionally strand along Ninety-Mile-Beach.
Note: The Hawksbill Turtle is the species that provides the ‘Tortoise shell’ of commerce — ‘Turtle shell’ of commerce is derived from the other species. In the past this species was decimated for the sake of its shell, but in modern times synthetic materials have given it a respite. The beautiful translucent shields are used for ornamental purposes and for the manufacture of jewellery. The flesh is not palatable.
Carapace of young three-keeled, the keels disappearing with age or only the median one surviving as an elevated ridge on the first two anterior neurals; nuchal hexagonal; costals five or more pairs, symetrically or asymmetrically arranged. Marginals 26, occasionally asymmetrically arranged, 25 or 27. Inframarginals four, occasionally three or five, the fifth partly covering the bony structure beneath. Head large with strongly hooked jaws, symphysis of lower jaw very long; two pairs of prefrentals, occasionally subdivided. Tail short, not exceeding the carapace in females, extending beyond in males.
Colour: In the adult the carapace is a drab olive or olive grey or reddish brown; the under surface is yellowish or buff suffused in places with pink; beak yellowish horn.
Size: The carapace of the adult attains a length of 1,050 mm.
Distribution: The tropical waters of the Atlantic, Pacific and Indian oceans, and the Mediterranean Sea. It occurs along the Australian and Tasmanian coasts and is an occasional visitor to the New Zealand area.
Only two species of marine snakes have been recorded from New Zealand waters: The Black-banded Sea Snake, Laticauda colubrina (Schneider) and the Yellow-bellied Sea Snake, Pelamis platurus (Linn.) Very occasionally a terrestrial snake may appear in cargoes of timber, fruit and other goods.
All sea snakes are poisonous and some of them are particularly deadly. However, the venom of Laticauda does not appear to be strongly toxic to human life. Sea snakes generally require a considerable amount of provocation before they can be induced to bite.
Description: Body subcylindrical. only slightly compressed, rostral scale higher than broad; an azygous scale separating the prefontals, sometimes absent. Labials 7 or 8, third (sometimes third and fourth) below the eye. 21-23 (rarely 25) rows of imbricate scales around body. Caudal scales along tail, 37 to 47 in males, 22 to 35 in females; anal scale divided.
Colour: Light or dark bluish grey above, yellowish below, with black bands of more or less uniform width throughout, or narrowing across the belly.
Size: Males smaller than females, 875mm long, tail 130mm; females 1.420 mm, tail 145 mm.
Distribution: Ranges from the Bay of Bengal to the seas of southern Japan, the coasts of Australia and islands of Oceania. An occasional visitor to New Zealand.
Head narrow, snout elongate; body much compressed, the greatest height posteriorly being more than twice that of the neck. Supralabial scales 7 to 8, fourth and fifth below the eye, usually separated from it by subocular scales. 49 to 67 scales around the thickest part of the body. Scales more or less hexagonal or quadrangular, the lowermost each with two or three small tubercles, strongest in adult males.
Colour: there are numerous colour varieties of this species and all possible intergrading takes place. The colour form occurring in the New Zealand area is described as ‘black above, yellowish or brown yellow below, with a lateral series of black spots which may be confluent into stripes, head black above, upper lip usually yellow.”
I have seen specimens from our area which were truly bicoloured, blackish above and yellow beneath, the two colours well demarcated.
Size: Total length of male 720 mm, tail 80 mm; female 880 mm, tail 90 mm.
Distribution: Pelamis is the most widely distributed of all the sea snakes. It is truly pelagic and has been found hundreds of miles out at sea. It is confined to salt water and does not frequent the mouths of rivers.
Note: (A more comprehensive paper on the Marine Turtles and Snakes of New Zealand is awaiting publication.)
New Zealand Forest Service, Wellington.
Zoology Department, Victoria University of Wellington.
FallowDeer are one of the most attractive of the deer family and were among the first of the eight species of deer that have been successfully introduced to New Zealand. Although several liberations were made, fallow deer have not become widespread but remain restricted to areas near where they were liberated. Introduced for sporting purposes, the bucks were eagerly hunted when fallow deer became established. Despite the hunting pressures, in most cases the herds built up in numbers and all protection of them ceased in 1925.
Fallow deer belong to the genus Dama Frisch, 1775, which is placed in the Order Artiodactyla, Family Cervidae, Sub-Family Cervinae. The peculiar shape of antlers is the basis for placing fellow deer in a genus district from Cervus. There are 2 species in the genus Dama; D. dama L., 1758 (Fallow deer) and D. mesopotamica Brooke, 1875 (Persian fallow deer). Dama mesopotamica is given specific rank because of its greater size, difference in shape of antlers, and more particularly in the shape of the nasal bones which are broader across the proximal ends (Ellerman and Morrison-Scott, 1951). Dama mesopotamica is probably extinct now although there may be a few in captivity (Walker, 1964). The range of D. mesopotamica formerly extended to Palestine where it could have come into contact with D. dama.
The adult male (called a buck, Vesey-Fitzgerald, 1946) stands between 35-37 inches at the shoulder, with the female (doe) averaging a few inches less. The weight of an average buck is between 170 and 180 lb, with large animals up to 240 lb (Southern, 1964). Body length is between 65 and 75 inches (Fig. 1.)
There are many colour varieties, but they may be divided into 3 main types: (1) Dark grey, often appearing to be true black,
(2) Mottled with prominent white spots on a fawn background, and (3) Pure white, de Nahlik (1959) states that the white fallow deer are all of park blood; the spotted form comes from South European imported stock, and the dark grey represents the wild variety. The most common summer coat for fallow deer in New Zealand is a deep fawn with conspicous white spots on the flanks, and a black or dark grey brown ridge of hair along the centre of the back and tail. The winter coat is a greyish-fawn with the spots far less prominent. Fawns are very heavily spotted.
As with most other members of the Cervidae, antlers are carried only by the males. Antlers appear in the second year and are fully developed by the sixth; they are flattened but have a cylindrical stem, usually with no bez tines and palmated above the trez. The length of full-grown antlers averages between 25 and 31 inches (exceptionally up to 36 inches), with a circumference of 4 to 6 inches. Distance between the tips is 15 to 20 inches (exceptionally 28 inches). The palmation of a trophy head should be as solid as possible with a number of tines along the posterior edge. The palm should be concave and spoonlike, bulging outwards (de Nahlik, 1959). Antlers are cast in November-December (in New Zealand), regrown by February, and are clear of velvet by the end of February. The age of a fallow deer is usually judged on the size and shape of the antlers.
Vesey-Fitzgerald (1946) states that fallow deer have as good a power of scent and hearing as red deer but far better eyesight. This will be confirmed by anyone who has stalked both fallow and red deer. Fallow deer are usually silent except for a bark-like call used when alarmed; a rutting call which is a mixture of a grunt and a bark and does not carry very far; and a whining bleat used occasionally by does and fawns.
The gait of the fallow deer is characteristic, appearing stilted or jerky. They run by jumping on all fours. Even when undisturbed they are able to jump 6-foot obstructions and when pressed can clear 7-foot obstacles with ease.
The original home of fallow deer is around the Mediterranean region of Southern Europe and Asia Minor (Ellerman and Morrison-Scott, 1951), but they have been widely introduced to other European countries and are now found wild throughout Western Europe and the Baltic States. Ellerman and Morrison-Scott (1951) also state that introductions were made in North Africa but it is doubtful if they survived. The present distribution in Asia Minor is not clear. Southern (1964) concludes that the evidence for fallow deer being indigenous in the British Isles is doubtful, but they have been present there for some time, as it is recorded in the Doomsday Book that they were well established. Ingersoll (1906) states that fallow deer were taken to the British Isles probably by the Roman colonists. Some herds still live wild in parts of England, and herds resulting from park escapes are common in most of the woodland areas; they appear to be less common in Wales, Scotland and Ireland. Fallow deer were liberated in Tasmania, from England, about 1850 (Bently, 1957) and are now well established. Small herds are also present in other Australian States. Distribution in New Zealand is shown in the accompanying map (Fig 2) and is discussed further below.
The first fallow deer introduced to New Zealand came from Richmond Park. Surrey, and were liberated in 1864, in the Aniseed Valley a few miles from Nelson (Donne. 1924). Twenty-five known
The following notes indicate briefly the present status of the herds.
Wairoa River (North Auckland) — This herd is probably on the borderline of extinction with only a few old footprints at present evident.
Great Barrier (Selwyn Island) — The status of this herd is not known but only a few animals, if any, now exist there.
Puriri (Coromandel Peninsula) — The numbers of fallow deer are very low.
Kawau Island — Only a very light population present.
Kaipara — Stable population, present in moderate numbers.
Manakau Heads — Probably not more than a dozen fallow deer exist in this herd at present.
Rangitoto and Motutapu Islands — Only very few deer now present.
Matamata — Light to moderate population present.
Wanganui — This herd is thriving and is probably extending its range.
Mt. Arthur (Nelson) — Moderate, stable population.
Aniseed Valley (Nelson) — Light, stable population.
Paparoa Range (Westland) — Light, stable population.
Opihi River (South Canterbury) — Light, stable population.
Blue Mountains (Otago) — one of the most extensive herds in New Zealand and has been subject to control measures when some 6,000 fallow deer were killed by Government operations in 1961-62.
Wakatipu — Moderate population, and increasing its range.
It is noticeable that fallow deer have colonised only the country close to the point of original liberation and thus appear as discrete populations — Caughley (1963) gives a mean rate of dispersal for fallow deer as 0.5 miles per year.
Fallow deer kept in deer parks are gregarious, but in the wild are less so than red deer. The herds are rarely large, except locally in late winter or spring (Vesey-Fitzgerald, 1946). They prefer extensive deciduous or mixed woodlands, preferably with a thick herb and shrub layer (Southern, 1964). During daylight fallow deer remain in the cover of the forest and their feeding is usually confined to dusk and dawn, although where undisturbed they may become more diurnal, especially in winter.
Sexes keep together from the beginning of the rut (April) until early spring; the remainder of the period they usually separate. In New Zealand the rut begins in late April and generally is of short duration (i.e. when compared to that of red deer). As with red deer, frosty nights and clear days result in earlier rutting, while warm weather delays the rut. Whitehead (1950) points out that a definite order of precedence exists among the bucks. Dispositional behaviour changes with age, with young fallow bucks alert and timid while old bucks become more sullen and moody. At times fallow bucks are aggressively pugnacious and are able to cause extensive injury to one another.
Fawns are born in December after a gestation period of 230 days (Ashdell, 1946). They can run within an hour or so of birth and are able to jump well when only a few hours old. Usually only one fawn is born, rarely two (triplets have been known, [Donne 1924]).
Disease is unknown in New Zealand fallow deer, but in a recent study by one of the present authors (Andrews, MS) three nematode parasites were recorded from this host. These worms were: Apteragia quadrispiculata and Spiculopteragia asymmetrica from the abomasum; and Oesophagostomum venulosum from the caecum. Sweatman and Williams (1962) found cysts of the dog tapeworm, Taenia hydatigena, in New Zealand fallow deer.
When first liberated fallow deer were protected, but from 1890 onwards could be shot by licensed stalkers. This control was not always effective and at times Acclimatization Societies found it necessary to destroy considerable numbers. From 1923 to 1930 fallow deer were declared vermin by various Acclimatization Societies, and in 1930 the Government completely removed protection from all deer. At present fallow deer are declared ‘noxious animals’ by the Noxious Animals Act, 1956 which is administered by the New Zealand Forest Service. This Act makes provision for the control and eradication of harmful species, including fallow deer.
Fallow deer are able to build up to very high numbers in some areas, e.g. Blue Mountains (Otago) where 6,000 were killed in 2 years on a 20,000 acre block (unpub. report N.Z. Forest Service). Fortunately, although fallow do cause severe damage to the vegetation, they do not occupy high altitude forest and alpine grassland where the erosion risk is most severe. The total number of fallow deer killed each year, by either government employed hunters or private hunters, is not known.
Fallow deer have no commercial value in New Zealand. Bucks are hunted for their heads. The meat is palatable though not especially prized by hunters.
The Authors wish to thank the members of the Division of Protection Forestry, New Zealand Forest Service, for their continued help.
New Zealand has a small fauna of native fresh-water fishes, consisting of about 34 species grouped in 8 families. The fauna includes none of the primary fresh-water fishes — groups which are restricted entirely to fresh-water; in fact, all the families are described as peripheral fresh-water fishes, characterised by the ease with which they tolerate brackish or fully marine waters. Increasing numbers of New Zealand fresh-water species are being found to have marine dwelling stages.
The fauna is a mixture of Indo-Pacific and southern temperature derivatives which have dispersed to New Zealand, probably from Australia (the southern forms) or South-east Asia via Australia (Indo-Pacific forms). The southern temperate forms are the most numerous. One family (Cheimarrichthyidae) is endemic to New Zealand and its affinities are uncertain. Fresh-water fishes have been able to reach New Zealand, in its isolation from other land masses, because of high tolerance of sea water, and it seems unnecessary to postulate land connections between New Zealand and other land areas to explain the presence of fresh-water fishes in New Zealand (see McDowall, 1964a).
Numerous attempts have been made to introduce fresh-water fishes to New Zealand; Thompson (1922) listed 30 species, mostly from the Northern Hemisphere. Many of these were Salmonidae, and of the 13 species which have succeeded in New Zealand, most are Salmonidae. As a result of these introductions, New Zealand has a famed brown and rainbow trout fishery; also a likely result of introducing these fishes is the almost certain extinction of Prototroctes oxyrhynchus. This species, once abundant, has not been recorded alive since 1923. Other species (e.g. Galaxias argenteus) also appear unable to compete successfully with trout and are rapidly becoming rarer. Such happenings sound a warning which should be heeded when the introduction of further predatory game fishes into New Zealand is considered.
The native fauna appears to be peculiarly restricted to smaller streams, although this may be only a reflection on the problems of collecting small fishes from large swift rivers. Most species
Cheimarrichthys fosteri, which has adapted to the unstable environment of riffles in shingle rivers. Very few of the native fishes are commonly known, even to fishermen; some like the eels (f. Anguillidae), the whitebait (f. Galaxiidae) and smelts (f. Retropinnidae) are often seen and well known but others are secretive and dart for cover when disturbed. Further species are seldom seen because they are nocturnal hunters. The smelts and Galaxias attenuatus are the only truly shoaling species, and Prototroctes oxyrhynchus, the so-called ‘grayling’, is probably the only species to have had trout-like, ranging, pool dwelling habits.
The aim of this guide is to enable the reader to determine the families of the native and introduced fresh-water fishes. Two common estuarine families are also included. For the introduced and estuarine families, the species occurring in fresh-water are listed and brief notes appended. For identification below family level, the keys have been confined to the native fresh-water families. Having determined the family to which the native fishes belong, the reader is referred to secondary keys for each family which will usually enable identification to species level. In some cases present descriptions do not permit identification to species by means of a key; hence species groups are indicated in the keys. This problem is particularly severe in the f. Retropinnidae, in which distributional data have been heavily relied upon for specific identification. By necessity, some of the key characters are not easy to determine in specimens, and for this reason notes on distribution, habitat, life history and structure are given in brackets below the key characters for each species. Difficulties in the construction of keys also occur when juveniles are taken into account. Except in the case of Geotria australis, no allowance has been made for the separation of juveniles by means of the keys. Geotria australis has three distinct life-history stages in fresh-water, and notes have been included for the recognition of these stages. Sexual dimorphism occurs in all New Zealand Eleotridae, and it may therefore appear confusing that two fish with quite different colouration and rather different proportions key out to the same species (compare Figs. 11 and 12). Literature on New Zealand fresh-water fishes is listed in a recent bibliography (McDowall, 1964b).
Abdominal pelvic fins — pelvic fins inserted on abdomen, usually well behind pectoral fins.
Adipose dorsal fin — a small fleshv fin without supporting rays, dorsally behind rayed dorsal fin.
Anadromous — species migrates upstream, usually from the sea or a lake.
Anal fin — fin mid-ventrally, behind vent.
Biserial teeth — teeth in two, usually parallel rows.
Canines — enlarged jaw teeth.
Catadromous — species migrates downstream usually into the sea or a lake.
Caudal fin — tail fin.
Caudal peduncle — posterior part of trunk between anal and caudal fins; length measured from base of last ray of anal fin to hypural crease; depth measured as least vertical dimension.
Colouration — refers to colouration of fresh specimens.
Compressed — laterally flattened.
Depressed — dorso-ventrally flattened.
Dorsal fin — median, unpaired fin on dorsum of trunk; one or more may occur.
Entopterygoidal teeth — present on paired bones forming roof of mouth behind vomer.
Flange of caudal peduncle — compressed, fleshy, dorsal and ventral expansions of the caudal peduncle.
Genital papilla — small protuberance behind vent, carrying the openings of the reproductive organs.
Gill rakers — bony filaments along the anterior edges of the gill arches.
Head length — distance from tip of snout to posterior margin of opercular membrane (H.L.).
Hypurals — the modified last few vertebrae which support the caudal rays.
Hypural crease — crease formed at the end of the caudal peduncle by bending the caudal fin to one side; the crease forms at the junction of the hypurals and the caudal fin rays.
Jugular pelvic fin — fin occurs anterior to pectoral fin base.
Keel — a semi-bony ridge along the mid-ventral abdomen in front of the vent.
Lacustrine — lake dwelling.
Maxilla — paired bone in upper jaw, behind and adjacent to premaxilla.
Median — in dorso-ventral midline.
Origin of fin — anterior point of attachment of fin to trunk.
Pre-maxilla — paired bone forming front of upper jaw: toothed.
Pyloric caeca — tubular sacs arising from posterior end of stomach.
Ray — soft, bony rod supporting fin; may be branched or simple, but always in adjacent pairs and segmented; such a fin is soft rayed.
Relative positions of fins — refers to the relative positions of vertical lines through fin origins.
Snout profile — outline of head from tip of upper jaw to above eyes.
Spine — firm, bony rod supporting fins, unbranched, unpaired and unsegmented; such a fin is spinous.
Standard length — distance from tip of snout to hypural crease (S.L.).
Thoracic pelvic fin — fin positioned below pectoral fin bases.
Uniserial teeth — teeth in a single row.
Vomerine teeth — teeth on vomer, a bone situated medially in the front of the roof of the mouth behind the pre-maxillae.
Vent — posterior opening of the alimentary canal.
Vertebral count — number of vertebrae from head to caudal fin, not counting hypurals.
Some of the criteria used in the keys are relative, e.g. jaw length, fin positions, and others are a little diffcult to use, e.g. entopterygoidal teeth, gill rakers, making the keys more difficult to follow. To minimise this, notes have been added below the key characters for each species. These notes include information which helps to confirm identifications, e.g. general shape and form, structural characteristics, colouration, distribution, habitat, marine phases, migratory habits and maximum size attained. While these do not produce absolute separation of species by themselves, when they are used in conjunction with the key characters, they should make identification more simple.
F. Ameiuridae Ameiurus nebulosus — catfish (N.Z.) or brown bullhead (U.S.A.). Introduced from the Great Lakes region, U.S.A. Stoutly built, scaleless, with the first dorsal fin having a stout erectile spine anteriorly; 4 pairs of barbels occur around the mouth. Brown to olive green in colour, growing to about 40 cm. It is uncommon in New Zealand except in parts of the Waikato River system.
F. Cyprinidae. Tinca tinca — European tench. Stockily built with a pair of barbels at the corners of the mouth, small, deeply embedded scales and truncated caudal fin; olive-green in colour. It grows to about 75 cm., but is usually smaller; rare in New Zealand.
Carassius auratus — Prussian carp or goldfish.
Carassius carassius — Crucian carp. Both species introduced from Europe or Asia. They are stout bodied with prominent scales and a long, high dorsal fin; olive to silvery-grey in colour, variants orange-red or pearly; known to grow to 75 cm., but usually to about 20 cm. Common in some localities.
F. Percidae. Perca fluviatilis — European perch. A deep bodied fish, well covered with moderately large scales; a distinct hump often occurs behind the head, carrying the large, flag-like dorsal fin which has sharp, rigid spines. Beautifully coloured; silvery green with a darker back and bright orange-red on the pelvic, anal and lower caudal fins. Grows to 25 cm., or more. It is successful in New Zealand in lagoons, coastal lakes and sluggish, weedy rivers.
F. Poecilidae. Gambusia affinis — mosquito fish. A small, live bearing species introduced from North America. It has a flattened head and upturned terminal mouth; the male has the anal fin modified to form an intromittent gonopodium. Silvery in colour. The male is smaller than the female, lengths 35 mm., and 60mm., respectively. Not widespread in New Zealand, but locally very abundant in northern areas.
F. Salmonidae. —
Salmo trutta —brown trout.Salmo gairdneri — rainbow trout.Salmo salar — Atlantic salmon Salvelinus fontinalis — brook char.Oncorhynchus tschawytscha — quinnat salmon.Oncorhynchus namaycush — Great Lakes char or mackinaw.
The trouts, salmons and chars were introduced from North America and Europe with varying success. Salmo trutta and S. gairdneri are by far the most successful and widespread in New Zealand. They are large, rather slender, graceful fish with a good covering of small, firmly embedded scales. River dwelling fish are usually dark coloured with a variety of coloured spots on the dorso-lateral trunk; lake-dwelling and sea-run fish are usually silvery coloured with less prominent markings. In contrast with many native fishes, they inhabit open pools and tend to be more mid-water swimming fishes. They grow to a large size, up to 100 cm., but are usually smaller, commonly 30-75 cm.
F. Blenniidae. Tripterygion sp. — blennies or cockabullies. Small, stout, heavily scaled active fishes, bottom dwelling in tidal estuaries
F. Mugilidae. Aldrichetta forsteri — yellow eyed mullet. Erroneously called herring, it is a slender, fast swimming, shoaling species which moves into river estuaries with the rising tide and will move beyond estuarine waters into lowland rivers. Its scales are easily dislodged. It is bright silver in colour with a deep bluegreen back and a bright yellow eye; grows to about 45 cm., and is New Zealand-wide.
Mugil cephalus — grey mullet, in some localities known as fresh-water or jumping mullet. Similar to A. forsteri, but is more heavy bodied with a broader head and larger scales. The eye of M. cephalus is covered by a thick, fleshy, adipose eyelid which is divided into two halves by a vertical slit. M. cephalus may penetrate many miles up rivers and may be seen jumping from the water as it moves. It grows to about 70 cm., and is mostly a northern species in New Zealand.
F. Geotridae — lamprey.
Partially fresh-water dwelling and semi-parasitic. There is one species in New Zealand; genus Geotria. G. australis Gray.
(Three life-history stages:—
Form slender, trunk rounded with swollen branchial region, a hood surrounding a small oral sucker, no obvious eyes, one continuous fin-fold; grey-brown in colour; muddy backwaters and pools; 85 mm. Ammocoete — larva.
Form slender, slightly compressed, no hood around suctorial mouth, branchial swelling lost, prominent eyes, two separate dorsal fins; trunk silver laterally, iridescent blue dorsally; 120 mm.; catadromous to sea. Macrophthalmia — juvenile. Form eel-like, with expanded suctorial mouth, eye present, male with prominent pouch ventrally on head; colour when leaving the sea similar to macrophthalmia, becoming dull grey; anadromous from sea to breed; N.Z.-wide; 55 cm., adult). (Fig. 1.)
F. Retropinnidae — smelts, cucumbers or silveries.
Small, slender, silvery fishes, usually with a purplish lateral sheen and a cucumber-like smell; pelvic fins abdominal. The males have larger fins than the females and in some species have small nuptial tubercles on the latero-ventral trunk and the lower fins. Seven species occur in New Zealand, some of these being migratory estuarine forms, the others lake dwelling. There are two genera, Retropinna and Stokellia.
F. Aplochitonidae — ‘Grayling’.
(trout-like with small scales and under-shot jaw; grey to red-brown on back, silver to golden on belly; formerly
F. Galaxiidae — includes whitebait, kokopu and mudfishes.
Mostly small, scaleless fishes with thick fleshy fins, rounded trunks and sometimes quite stout build. The sexes are similar and the colouration generally sombre. They are usually solitary, secretive, fast-swimming fishes. Five species have marine juveniles (whitebait) which migrate upstream from the sea in the late winter and spring in mixed-species shoals.
(Fig. 5.). (slender, pectoral fin lateral; pale with darker mottling, silver belly; shoaling; lowland streams; N.Z.-wide; juvenile, marine whitebait; 160 mm.)
Canines strongly developed; gill rakers long, caudal fin slightly concave, fins thick and fleshy G. fasciatus Gray
(Fig. 7.) (stout, pectoral fins ventro-lateral: colour dark
(similar to N. apoda but much darker in colour; north of North Island; small holes in peat swamps, sometimes swamp creeks; 120 mm.)
F. Pleuronectidae — flounders.
One species commonly found above tidal estuaries; genus Rhombosolea. R. retiaria Hutton
(Fig. 4.). (rounded outline with blunt snout; grey-brown to black on right side with many brick red blotches, each surrounded by a dark halo, left side grey to yellowish-brown; estuaries and upstream; quiet muddy pools to rapid gravelly rivers, also lowland lakes; N.Z.-wide; juvenile probably marine; 75 cm.)
F. Anguillidae — fresh-water eels.
Two species occur in New Zealand, one growing to large size; apparently scaleless but possessing small scales embedded in skin. The pelvic fins are absent; genus Anguilla.
Dorsal fin extends much further forward along the body than anal fin, vomerine teeth in a narrow band, cleft of jaw extends past eye A. dieffenbachi Gray
(features gross, thick lips, broad bulbous head, especially in large examples; dark dorsally, yellowish-brown ventrally, estuaries to upland lakes and streams; N.Z.-wide; juvenile marine; 180 cm.)
Dorsal fin extends only a little in advance of anal fin, vomerine teeth in a club-shaped mass, cleft of jaw just reaches eye A. australis schmidti Phillipps
(Fig. 3.). (features more slender; greenish-brown to grey dorsally, grey-white ventrally; estuaries, lowland streams and lakes; N.Z.-wide; juvenile marine; 90 cm.)
F. Eleotridae — bullies.
These are mostly small fishes with large scales, thoracic pelvic fins and two dorsal fins. The sexes are dimorphic, the male being more brightly coloured than the female, having larger fins, stouter build and growing larger than the female; two genera are recognised — Philypnodon and Gobiomorphus.
(Fig. 14.). (fairly stout, depressed and ventrally flattened, under-shot jaw, fins large and fleshy, pelvic fins jugular; light grey with several oblique irregular dark bands across trunk, fins and belly reddish; mostly in swift often unstable gravel rivers; N.Z.-wide; juvenile probably marine; 175 mm.).
Financial support of a New Zealand National Research Fellowship during the completion of this paper is gratefully acknowledged. The author wishes to thank the following for critically reading the