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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 University of Wellington, Box 196, Wellington, New Zealand, should be sent to: Business Manager of Tuatara, c/o Victoria.
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New Zealand plants loomed large in the life of Thomas Kirk. His earliest collectings were made within a month of his arrival in Auckland in 1863, and when his field days were over, until the last week of his life, he requested from correspondents certain specimens that ‘would be highly prized.’ Amongst New Zealand botanists he will always occupy a key position.
The Transactions of the New Zealand Institute, its first volume published in 1869, was an instrument ready to Kirk's hand, and he made good use of it, contributing some 138 papers during nearly 30 years of membership. It is in these papers, and in the herbarium specimens supporting the records he made there, that his labours for New Zealand botany are well represented. His two books, the large Forest Flora and the incomplete Students' Flora, were the first of their kind written in this country for this country, and their wealth of local knowledge and references endeared them to users
The broad pattern of Kirk's botanical interests is clear from the Transactions alone, though he did publish a few papers elsewhere. Following chronologically through, one sees his interest and experience develop along a number of parallel lines, all of them fairly well defined from the beginning, and the discussion of his work falls naturally under equivalent headings. No attempt is made here to assess his contributions to more strictly economic botany, for example on grasses, flax, sand-reclamation and forest products.
Kirk obviously had a tidy mind and it came naturally to him to classify. He liked to be able to name any plant that he met. If in a group that was generally satisfactory he found an obvious novelty he did not hesitate to describe it as new, but he also made a practice (as Veronica 21 (7); Celmisia 17 (6); Gentiana 17 (7); Carmichaelia 17 (11); Olearia 16 (4); Pittosporum 15 (5); Lepidium 10 (4); Ligusticum 9 (5 in several genera); Panax and Pseudopanax 8 (5); Cotula 8 (3); Gunnera 7 (4); Colobanthus 6 (4). In all he described more than 380 new taxa, including two genera (Huttonella in Leguminosae and Simplicia in Gramineae), many species and varieties and some formae. More than 50 species and 70 subspecific entities were published in the Students' Flora, his unfinished last work. Some 200 of his names survive, many of them in familiar present use, for example: Astelia trinervia, Carmichaelia williamsii, Celmisia walkeri, Coprosma arborea, Dacrydium intermedium, Drimys traversii (Pseudowintera), Gahnia pauciflora, Gentiana corymbifera, Gunnera hamiltonii, Hymenophyllum rufescens, Isoetes alpinus, Pseudopanax ferox, Pilularia novae-zelandiae, Pittosporum ralphii, Podocarpus acutifolius, Podocarpus hallii, Ranunculus paucifolius, Veronica canescens (Parahebe), Vitex lucens.
It is mostly Kirk's varieties that have been dropped, some because they are now considered to be hybrids, a possibility that he does not seem to have contemplated. For him the criteria for distinguishing
Astelia (see Skottsberg, 1934). His keys are real keys, setting out parallels of firmly contrasting characters, and they are much easier to use than Cheeseman's abbreviated diagnoses arranged in key form.
The herbarium was a result but not the principal objective of his botanical investigations. His main collection, now at the Dominion Museum, Wellington, was referred to by Cheeseman (1906:v) in these words: ‘I am indebted to the Education Department for the loan of that portion of the herbarium of the late Mr Kirk which after his death was purchased by the New Zealand Government. Although comprising only a small part of the collections formed by this active and enterprising botanist, it nevertheless includes excellent and well-selected specimens of most of the species of the flora, including the types of the new species described by him.’ Other portions are to be found in many parts of the world; in the genus Miss Pittosporum, for instance, Cooper (1956) lists 76 Kirk specimens in nine different herbaria outside New Zealand. Specimens that he gave away are in fact often better than those in Wellington where unfortunately there was some confusion of labels and some damage amongst unmounted specimens (Hamlin 1965:93).
Kirk had no doubt about the importance of plant geography. He felt under an obligation to explore as many parts of his adopted country as possible and to record his observations on the general appearance of the vegetation, to list the species noted, and often to indicate the type of habitat favoured by each. Lists were made under such headings as ericital, sylvestral, uliginal, littoral. Districts for which he provided surveys more or less of this kind included the northern part of the Province of Auckland, Great Barrier Island,
Kirk tried to define the northern or southern limits of species of restricted range and drew attention to extensions of range that came to his notice, but he placed less emphasis on altitude. In general he did not offer hypotheses to explain the limits of distribution but he did correlate the presence of maritime littoral plants far up the Waikato Valley with the presumed earlier intrusion of the sea.
These surveys cover naturalised as well as indigenous plants as a matter of course, and a fair share of attention was given to the frequently overlooked water-plants.
Early expeditions were in the Auckland Province and Kirk's appreciation of the whole plant life of the country became balanced only as his explorations took him further south. In 1872, for instance, in a comparison between the indigenous floras of New Zealand and the British Islands, tussock grassland is not even mentioned among the major vegetation types of New Zealand. Though later he climbed many mountains he nowhere gives any detailed account of alpine plants in their natural habitats. Wherever he went he found plenty that was new and noteworthy, and certainly he did not ignore mountain species of the genera that he monographed.
Kirk obviously arrived in New Zealand with a good knowledge of European plants and immediately he began to record those he found playing an important part in New Zealand vegetation. One of his first papers was on naturalised plants, especially those in the environs of Auckland, and later accounts deal with the naturalised plants of Chatham Islands and of Port Nicholson. In 1896 he enumerated 104 species of plants collected on a heap of earth ballast deposited by a ship from South America on the Wellington waterfront, where he followed their progress through two or three years. His Presidential Address to the Wellington Philosophical Society in July, 1895 was on ‘The Displacement of Species in New Zealand,’ a
The Students' Flora of New Zealand and the Outlying Islands is still unique amongst New Zealand Floras in that it sets out to describe native and alien plants in one sequence. Had Kirk succeeded in completing this project the pattern would undoubtedly have been accepted for later volumes to the great advantage of the study of plants in this country. Such a flora not only teaches about a great many more of the plants to be met with but also introduces additional families and gives a broader view of the structure and content of the families to which New Zealand indigenous species belong. But, as Kirk found, the preparation makes great demands.
The botanical writings of Thomas Kirk are on the whole very impersonal, revealing little of the man except his intense interest in all plants. He refers warmly to certain friends — Bishop Williams, Mr Justice Gillies, Mr Enys — but rarely mentions Cheeseman or Petrie, except for a minor tiff with the latter over Gunnera ‘obovata.’ He had an obvious affection for Buchanan whose botanical errors and omissions he corrected gently though always firmly. His correspondence with botanists abroad would undoubtedly provide much of interest, but that lies beyond the scope of the present note. One learns something of his methods of handling plants from his article on the preparation of botanical specimens (1895) and very occasionally he allows himself to recall some circumstances of collecting. For example, writing of Gleichenia dichotoma at Rotomahana: ‘In several instances steam jets had burst through spots occupied by this fern, and destroyed patches of it two or three feet in diameter. The temperature endured by the roots must have been over 100°F. In one spot the ground gave way under my feet, when a steam jet immediately broke through and destroyed the fern all round. My natives did not approve of fern-collecting in such situations.’
Of Mount Anglem, Stewart Island, he records: ‘Before reaching the crest we encountered a dense snowstorm, which later on was varied by fierce blasts of sleet and hail, so that hands and faces were stung almost past endurance, and the use of note-book and pencil became impossible for the rest of the day… The rapid approach
In 1897, in an essay on the history of botany in Otago, Kirk wrote ‘It is well that the memory of the pioneers in any branch of research should be treasured by those who reap the benefit of their labours.’ Thomas Kirk will not be forgotten.
Porrhothele antipodiana (Mygalomorphae, Dipluridae), is the large, orange-backed tunnel web spider commonly found under logs and rocks in the lower North Island and also in the South Island. Along with Aparua and Hexathele, the other two New Zealand genera in the Dipluridae, Porrhothele spins a tubular web which often leads into a burrow in soil or wood. All three genera contain large spiders, and P. antipodiana is no exception on this characteristic, some individuals of this species attaining a body length of 30mm. Their size, coupled with an aggressive manner and a covering of long black hairs on the legs and abdomen makes these spiders an impressive and readily identifiable sight.
Research on the New Zealand mygalomorphs has been mainly along taxonomic lines, although some observations have been made on their biology. Todd (1945) observed some of the habits of P. antipodiana but on the whole her paper is more extensive in its consideration of the biology of the Ctenizidae. Forster (1967), and Forster and Wilton (1968) included some interesting observations on the natural history of the New Zealand mygalomorphs; but ecological and behavioural data were still sparse at this date. Accordingly, the present investigation was undertaken to determine what prey P. antipodiana took and what particular adaptations this spider has for prey capture. The paper comprises three sections: prey types taken, the use of the web in prey capture, and the behaviour of the spider.
P. antipodiana carries its prey into the tunnel of the web so this is the portion that must be examined to find out what the spiders have been capturing. Identification of prey is often difficult because small portions or segments are all that are found in some cases. Hard parts such as beetle elytra and the thoracic parts of most insects do survive. Slater and millipede segments usually become detached and identification in these groups is particularly difficult.
The following is a list of prey species recovered from 50 webs in the Wellington area:
Coleoptera: Ochosternus zealandicus, Oemona hirta, Odontria sylvatica, Cecyropa lineifera, Cecyropa sp., Philoneis aucklandicum, Cilibe sp., Holcaspis sp., Xyloteles humeratus, Xyloteles griseus, Artystona rugiceps, Navomorpha sp., Coptomma variegatum.
Homoptera: Scolypopa australis.
Hymenoptera: Vespula germanica, Amblyopone australis, Bombus sp., Salius monachus, Salius fugax.
Isopoda: Porcellio scaber.
Mollusca: Helix aspersa.
Araneae: Miturga sp., Dolomedas sp. Ixeuticus sp.
Various Unidentified: F. Hepalidae at least two species, Diplopoda at least two species, Isopoda at least one species, Diptera at least one species, Araneae: some crab spider fragments.
This list is by no means exhaustive. P. antipodiana seems to eat almost anything that comes near enough to its web and so it probably preys on the majority of beetles, millipedes and slaters that happen to be in the same locality as the spider. Of the hymenoptera, the Salius species found were probably the result of unsuccessful forages into the tunnel after the spider. V. germanica investigates most objects, so it is likely that it often gets caught in P. antipodiana webs; and from my own observations, the larger spiders have little trouble in dealing with this wasp once it has entangled itself in their web. Bumble bees are picked out of the web and carried back to the tunnel without difficulty too. In the case of V. germanica and the bumble bee, the spider invariably grasps them from behind the thorax thus avoiding the sting. The presence of land snails in the webs may be a surprise on the grounds that the spider would find difficulty in gaining access to the soft parts of the snail.
The New Zealand Ctenizidae seem to take similar prey to P. antipodiana. Todd (1945), in an examination of burrows of South Island trapdoor spiders reported finding beetles, a weevil, large numbers of Odontria remains, caterpillar remains and a small fly.
Of the fifty webs examined in the present investigation, the different prey groups comprised the percentages shown in table one.
The proportions of the different prey taken does vary from area to area. For example, at Paremata, there are large numbers
The web of P. antipodiana, as in other spiders is an essential part of prey capture and has particular adaptations for this specific purpose. The basic pattern consists of a woven silk tunnel in which the spider shelters. In addition there may be a sheet of silk extending beyond the opening of the tunnel. The extent of the sheet is very variable and seems to be related to the local environment of the spiders. For example, webs found under logs or rocks have little if any development of the sheet; whereas webs situated on banks or in trees usually have an extensive sheet development. Thirty webs on a cliff face at Paremata showed a range of from 10cm to 40cm in the width of the sheet, measured at the widest point. Measurements taken of thirty webs from under rocks and logs on the hills around Ngaio (Wellington) showed that the sheet could be absent or up to 10cm in width. Fig. 1. gives some web shapes found in a variety
Of a hundred webs surveyed at Paremata, the following distribution of sites was found: in crevices of the bank 60; up trees 14; under logs or stones 26. An investigation of the hills around Wellington shows quite a different distribution of sites. Here the only sites available seem to be those under rocks and logs so almost 100% of the population is found in these sites.
Studies on mygalomorphs such as the Ctenizidae and the Atipidae have indicated that these spiders remain in the one burrow for the duration of their life (Forster and Wilton 1968, Savory 1926). This is not the case with P. antipodiana, for movement from one site to another seems to be common. In an investigation at Paremata lasting from March to August of 1971, 30 webs on an area of bank were pegged for identification and weekly inspections were made with a torch at night to determine if spiders were still in residence. This disturbance of the spider was minimal, causing them to retreat into their tunnels for 10-15 minutes at the most. At the end of the investigation, fifteen of the sites were no longer occupied, while ten new sites had been taken over by spiders. While this study did not identify which spiders moved where, it at least points out the mobility of the population.
Mobility is also evident in those populations living under logs and stones. Invariably there will be a number of web extensions to be seen when a log is lifted. Many are old and disused and the movements of the spider under the log can be traced by following the older tunnels to the newest, occupied part.
P. antipodiana, in its mode of dealing with prey, shows some adaptations not so apparent in spiders such as the orb web spiders (Epeiridae) and the common house spiders (Dictynidae). While it is common to see these spiders in their webs during daylight, it has not been my experience to see a single P. antipodiana either mending its web or waiting for prey while the light is still strong. P. antipodiana waits until dark and then comes out to the entrance of the tunnel. Occasionally, individuals may be found further out on the sheet web, presumably being engaged in mending the web. This very obvious difference between day and night behaviour of P. antipodiana is shown in table 2. which was based on the dropping of a slater into each of twelve webs in daylight and then repeating the procedure at night.
In the daylight part of the experiment, four spiders failed to appear at all so the mean figures given in the table are based on eight responses — not twelve as in the night responses.
Interpreting the table, the most obvious feature is the time taken to reach the prey. The night spiders reached the slater in minimum time, running at high speed the moment it landed in the web. The daytime spiders — if they appeared at all, were very cautious and waited at the entrance of the tunnel for some time before making a rapid dash to the slater. During the day the spiders spent far less time dealing with the slater, whereas at night the spiders handled the slater in a leisurely manner, taking much longer to deal with it.
The speed of the spider is probably significant in prey capture so timings were carried out to determine the spider's maximum speed. Webs with long sheets were selected, and a slater was dropped in at a point approximately 30cm from the tunnel entrance, and the time taken for the spider to reach the slater was estimated using a stop watch. Some spiders reached the slater in 1.5 seconds which meant they were running at a speed of at least 0.2metres-sec. For comparison, slaters and millipedes were timed over flat level ground to get an idea of the maximum speed they might be capable of running at. The fastest slaters ran at a speed of 0.1 metre-sec.; while 0.02metre sec. was the fastest speed recorded for the millipedes. These speeds are a generous estimate of the speed that slaters and milipedes could move at in a P. antipodiana web, but are the best that can be offered because of the difficulties experienced in getting prey species to move in a straight line in a web.
At night the spider has to contend with the stimuli provided by leaves, small pieces of wood, etc., dropping into the web. Initially, these are treated as prey — the spider approaching and assuming a position with the palps touching the object. This affords at least a tactile investigation of the object and possibly also a gustatory or an olfactory investigation. Blumenthal (1935) considered that the palps had an olfactory function, but this has been disputed. Inanimate objects are left alone after a few seconds investigation. Sometimes a prey animal will stop moving as the spider approaches and if this happens, the spider ‘tests’ it with the palps and within a short time it usually stabs the animal with its chelicerae. The stabbing of the prey is done more than once; this probably ensures deep penetration of the venom and would help in dealing with hard-bodied forms such as the beetles where the spider may have to manoeuvre the fangs until they slide into a joint or a soft spot. The chelicerae, along with the palps serve to hold the prey for carrying it back to the tunnel where it will be eaten.
At night the spider does not always find the prey on its first attempt. It is common for the spider to overshoot the prey or to misjudge the direction of the prey. Fig. 2. contains some prey-seeking movements of P. antipodiana that were observed during this study.
The ability of P. antipodiana to accept a wide variety of prey types must be of significance in its success as a predator. The presence of millipedes as prey may surprise some biologists, on account of the unpalatable secretions of these animls, which apparently deter some predators (Forster, 1971). Petrunkevitch (1952) has recorded that the tarantula Cyrtopholis portoricae feeds on insects and millipedes, so it may be that many of the mygalomorphs can exploit the millipedes as a source of food. Land snails in the tunnels of P. antipodiana present a problem of interpretation; it may be that the spider carries them there or it may be a similar case to that of worms in the burrow of the purse web spider Atypus. Savory (1926) recorded that partly devoured worms were found in the tunnel of Atypus; however Bristowe (1958) discounts records of Atypus feeding on worms. He observed that worms taken into the
P. antipodiana tunnels, like worms in Atypus tunnels are intruders that may or may not be killed by the spider.
The web characteristics must rate highly as a factor in P. antipodiana's success too. The extensive sheet webs of those living on exposed sites on banks or trees contrasts strongly with the often complete lack of development of the sheet in webs found under logs or stones. The reason for this is probably to be found in the distribution of prey species; for instance, under logs the spider is right amongst its prey — the beetles, slaters, etc., that shelter and feed there too. Whereas on the exposed sites the spider must rely on those animals that are moving over the surface of the bank for its food. There is no concentration of prey here on the bank as there is under the log. Consequently the spider on the exposed site requires a more extensive web to cover a wider area and so increase its access to prey. A similar situation has been described by Main (1957) in a study on the trapdoor spiders of South Western Australia. She found that those forms inhabiting damper, richer litter having an adequate prey supply, seized prey that were within reach of the burrow entrance. Those forms living in dry environments lay out lines of silk to act as indicators of when prey animals are passing by. These spiders would rush out several inches from their burrows to grab prey — an obvious advantage where prey are scarce.
The speed at which P. antipodiana is capable of running must be important too, for in cases of an insect wandering across the edge of the web, it is probably the speed at which the spider chases the insect that will determine whether or not a capture is made. The majority of prey species do seem to be able to cross a P. antipodiana web without becoming entangled; and it is here that comparisons can be made between P. antipodiana and the aran-eomorphs such as the orb-web spider, Aranea. The sticky webs of the latter, by holding prey, make prey capture so much easier; while P. antipodiana has to have the extra speed and hunting ability to ensure that captures are made.
Migratory behaviour probably gives a more efficient utilisation of prey resources, in that the spider goes to the prey and not vice-versa as is the case with the trapdoor spiders. Like the building of extensive sheet webs, mobility functions to cover a wider area of possible prey movement.
The behaviour of the spider in daylight must have considerable survival value. That few individuals give an immediate response to the presence of prey in the web probably reduces the susceptibility of P. antipodiana to predation by the Pompilid wasps Salius monachus and S. fugax (as well as other possible predators).
In conclusion it is recognised that a paper on prey capture could well include reference to other features such as the toxicity
P. antipodiana's venom; the postures assumed by the spider and its sensitivity to stimuli from the web. It is hoped that these features will form the substance of another paper at a later date.
P. antipodiana, commonly known as the tunnel web spider, feeds on a wide variety of prey types. The list includes the Isopoda, Hymenoptera, Coleoptera, Lepidoptera, Homoptera, Diplopoda, and other spiders as well. The web of P. antipodiana figures strongly in prey capture and shows considerable variation in differing environments. Mobility from site to site is a feature of this spider's behaviour. A strong photo-negativism is evident in its behaviour; which when coupled with the speed of running results in minimum exposure to predators. The manner in which P. antipodiana subdues its prey is described.
My thanks are due to Dr P. antipodiana; and to Miss Logan Hudson. Dominion Museum, who identified the prey remains, read the manuscript and offered valuable criticisms of it.
The larva of the endemic species Ixodes jacksoni Hoogstraal is described. I. pterodromae Arthur is recorded from sea birds in the New Zealand Subregion. The Neotropical species I. auritulus Neumann s.s. is recorded from a land bird in the Auckland Islands. The taxonomic status of I. auritulus zealandicus Dumbleton is discussed. Haemaphysalis bispinosa Neumann is replaced by H. longicornis Neumann.
The tick fauna of the New Zealand Subregion has been treated in previous papers (Dumbleton, 1944, 1953, 1958, 1961, 1963; Roberts 1970). The present article can be regarded as supplementary to the author's review of the New Zealand tick fauna in Tuatara 11: 72-78. A new endemic sea bird tick, Ixodes jacksoni Hoogstraal (1967), has recently been described from the subregion and the previously undescribed larva of this species is described below. The auritulus-percavatus species group of Ixodes is a difficult one taxonomically and an attempt is made to clarify the identity, occurrence and distribution of those species of this group which occur in the New Zealand Subregion.
J. Med. Ent. 4: 37-41. (male, female).
The larva, unknown until now, is described below.
Body (Partly fed) length 1.1 — 1.4mm, width 0.9 — 1.0mm. A smaller species than I. uriae White on the dimensions given for the latter by Arthur (1963).
Capitulum (Fig. 1): ventral length 0.2mm, width 0.13mm, with a posteroventral cordiform swelling. First palpal segment short, transverse, without setae; segments 2 and 3 not separated, without apicomesal gibbosity, length 0.08mm, with 5 setae about mid-length and 5 subapical; segment 4 with 2 long and 4 shorter setae apically and 1 or 2 subapical. Hypostome length 0.1mm, dentition 2-2 with 6 teeth in each file. Only one pair of posthypostomal setae present, on anterior margin of swelling on venter of basis capituli.
Scutum (Fig. 2): length 0.3mm, width 0.29mm, widest anteriorly, with 9 structures (5 setae, 4 sensillae) on each side. Eyes absent.
Body setae: (Figs. 3 and 4): 5 pairs centrodorsal, 7 pairs margino-dorsal; 3 pairs sternal, 2 pairs preanal, 1 pair anal, 4 pairs premarginal, 3 pairs marginoventral.
Legs: coxae without internal or external spurs, each with 2 setae. Fore tarsus (Fig. 5) length 0.22mm; dorsal setae; 4 prehalleral, 4 posthalleral, 2 basal; ventral setae; 4 apical, 4 median, 4 basal; Haller's organ with 5 internal processes.
Described from 2 larvae collected from a dead Pied Shag (Phalacrocorax punctatus punctatus Sparrman) on New Brighton Beach by Mr J. Ixodes eudyptidis Maskell were also present on this bird.
The larva resembles the nymph and female of I. jacksoni, and those of the same stages of I. uriae, in having a scutum which is widest anteriorly. Like that of uriae it is distinguished from most other Ixodes species except that of I. kopsteini Oudemans (Roberts, 1969), by the presence of only one pair of posthypostomal setae. Only 3 marginoventral setae are present and the presence of 5 centrodorsal setae is unusual in the genus, but both these conditions exist in I. brunneus Koch (Clifford et al, 1961).
The chaetotaxy of the jacksoni larva coincides with that of uriae as figured by Wilson (1967) but Filippova (1958) figures fewer structures on the scutum of uriae (4 setae only on one side and 4 setae plus 2 sensillae on the other) and additional dorsal setae (1 supplementary seta and 8 marginodorsal setae). The two species are undoubltedly con-subgeneric on characters of all stages. The apparent subdivision of the ventral plates of the males of uriae appears to be due to differences in pigmentation colour rather than in structure. I. jacksoni, endemic on the New Zealand mainland, is the only species which is closely related to the distinctive seabird species I. uriae, which in spite of its bipolar distribution gives no evidence of subspeciation. The only host of jacksoni known so far is a rather sedentary cormorant species nesting on rocky coasts, and the only locality records are from Banks Peninsula, or its vicinity, in the South Island, though colonies of the host species occur in the northern part of the North Island. Ixodes uriae on the other hand occurs commonly on various seabirds, including cormorants, in the New Zealand Subantarctic but appears rarely to breed on the New Zealand mainland. There are three records (Dumbleton 1953, 1961) of the occurrence of uriae in mainland localities but one of these was based on a misidentification of jacksoni, another
Eudyptes pachyrhynchus Gray) which does breed on the mainland.
Additional records of jacksoni are:—
Lake Forsyth: female ex Pied Shag, J. Ornithodorus capensis Neumann); Birdlings Flat; nymphs ex Pied Shag nests, I. eudyptidis and O. capensis).
Parasitology 50: 217-23 (female). Roberts 1960, Aust. J. Zool. 8: 414-7; 1964 J. ent. Soc. Qd. 3: 75-6 (male).
Tick specimens collected from sea birds in the New Zealand Subregion and belonging to the auritulus-percavatus group of Ixodes were originally considered (Dumbleton, 1953) to be I. auritulus Neumann 1904, a species nearly related to I. percavatus Neumann 1906 from sea birds on Nightingale Is. (Tristan da Cunha) by their common possession of an anterior prolongation of the inner side of the first palpal segment (Fig. 6). Zumpt (1952) and Arthur
percavatus to be a synoym of auritulus but Arthur (1960) re-established percavatus as a valid species distinguished by the possession of a basodorsal spur on the first palpal segment (Fig. 7) and described the females of three new species which possess a similar structure. The identity and host relationships of I. cornuae are in some doubt since no holotype was designated. The syntypes include 3 females ex a species of quail Equateur, Dr Rivet 1901, and 1 female and nymphs, Baie Orange, Mission du Cap Horn 1882-8, Dr Hyades and Sauvinet. The Equateur specimens are not only from a landbird but are apparently from Equateur since Dr G. Rivet collected there in this period with a French Army Mission measuring an equatorial are of meridian (Becker, 1919). I. zumpti Arthur was based on three females from sea birds (Puffinus, Diomedea, Phebetus spp.), Nightingale Is., Tristan da Cunha. I. pterodromae Arthur was based on two females from sea birds (Pterodroma spp.) from Marion Is, Indian Ocean. Roberts (1960, 1964) recorded pterodromae and described the male from Macquarie Is. and Wilson (1964) recorded the species from Campbell Is.
Three records of auritulus s.s. from the New Zealand Subantarctic (Dumbleton, 1953) are now considered to be based on probable misidentifications of pterodromae. The Antipodes Is. record was based on a nymph, a stage which is difficult to identify with certainty in this species group.
The specimens on which were based two early records of auritulus from Macquarie Is. have not been located but later collections from there have been exclusively pterodromae (Roberts, 1960, 1964a). It is possible that pterodromae itself may eventually be found to be a synonym of the older species percavatus.
Specimens determined as pterodromae have been collected as follows:—
Ocean Is. Auckland Is.): male among plants, P. M. Johns, 28.xii.62, det. Pelecanoides exsul Salvin).
Birdlings Flat: females ex Sooty Shearwater (Puffinus griseus Gmelin) M. Fitzgerald, 16.i.63, det.
Arch. de. Parasitol., 8: 450 (female); Arthur 1960, Parasit. 50: 201-12; Kohls and Clifford 1966, J. Parasit. 52: 815-7 (male).
Female: Body length (excluding capitulum) 6.4mm, width 5.0mm in engorged female; 2.6 × 1.8 to 3.8 × 2.6 in others. Colour yellowish-white in unengorged females, greyish-white in engorged female.
Capitulum ventral length 0.79mm. Basis widest 0.47mm across bases of cornuae; cornuae equilateral-triangular with slightly convex
Hypostome long, only slightly spatulate, rounded apically, dentition 4-4, about 12 teeth in outer file. Scutum suboval, longer than wide, length - width ratio 1.2 — 1.5, little anterior emargination, scapulae small, widest at midlength, anterolateral side slightly concave, lateral angle rounded, posterolateral side with slight, concavity at end of cervical groove, posterior margin broadly rounded. Colour yellowish brown except narrow cervical groove, a transverse anterior band and the outer half of the sub-marginal area between the cervical groove and the margin on which the punctation is most evident. Legs: Coxae with 4 prominent stout setae on posterior submargin, 3 between the internal and external spurs and one on the outer base of the external spur. The external spur on fore coxae not markedly larger than those on other coxae. Coxa I with 1 anterointernal seta and several large and subacute, the posterior margin between them deeply concave. Coxa II with shorter blunter internal spur and large external spur, the posterior margin between them more widely but less deeply concave. Coxa III with short blunt internal spur but large external spur, the margin between them straighter and shallower. Coxa IV with a broad trenchant edge or sometimes a very small spur internally, the external spur large and the posterior margin between them nearly straight. All trochanters with ventral spurs and first two with small dorsal spurs.
Genital aperture between coxae III. Genital grooves straight.
Anal grooves subparallel posteriorly. Spiracular plate subcircular, the macula slightly anterior of centre.
Described from 5 females ex Auckland Is. Snipe (Coenocorypha aucklandica Gray), collected by P. M. Johns on Ewing Is. (Auckland Is.) on -.i.63.
The characters of the first palpal segment in these specimens conform with those of the predominantly land bird infesting American species I. auritulus, and the coxal characters are very similar to those of Guatemalan specimens of this species figured by Arthur (1960). Males are not known and there is insufficient evidence to determine whether the Auckland Is. population should be regarded as a distinct subspecies. The probable validity of the record is strengthened by Robert's (1964b) record of auritulus s.s. from two land birds, Strepera fuliginosa Gould (Black Currawong) and Sericornis
humilis Gould (Brown Scrub Wren), in Tasmania. Dr Roberts who examined an Auckland Is. specimen agrees on its identity with his Tasmanian specimens.
It is not possible to decide between the possible explanations of the disjunct distribution of auritulus s.o. It seems improbable that the Auckland Is. and Tasmanian occurrences could be attributed to chance dispersals of auritulus from America on migrating birds, since it occurs characteristically on land birds and the migratory land bird species coming to New Zealand are largely or exclusively from Asia rather than America. A second possibility is that the Auckland Is. population segregated from I. pterodromae in an environment where the habitats of land and sea birds were contiguous, and that its morphological similarity to auritulus resulted from convergent evolution. Such a segregation has been suggested by Smit (1965) to explain the origin of the New Zealand parrot flea Parasyllus nestoris Smit from one of the other species of the endemic P. cardinis group, all of which occur on sea birds. It seems unlikely however that a species which segregated on land birds in the Auckland Is. could have dispersed to Tasmania (or vice versa), or that there were independent segregations of morphologically identical species on both islands. The occurrence of auritulus in southern Chile, Auckland Is., and Tasmania suggests the austral type of distribution which characterises many other southern hemisphere taxa. The Tasmanian and Auckland Is. populations would then be regarded as relict. The fact that auritulus occurs in both North and South America, while inconsistent with the restricted southern hemisphere temperate zone distributions of typically austral taxa, does not necessarily negative this explanation.
N.Z. Jl. Sci. 4: 765-6.
A male Ixodes from the nest of a sea bird (Pelecanoides chathamensis) on Snares Is. 90 miles south of New Zealand was described and figured (Dumbleton, 1953) as the previously unknown male of I. auritulus Neumann sensu stricto, a species described from an unknown host in southern Chile but occurring commonly (possibly exclusively) on land birds in both the Neotropical and Nearctic Regions. Associated females from the Snares Is. and from a burrow-nesting sea bird (Pachyptila turtur (Kuhl)) on Stephen Is. in Cook Strait (in addition to others from islands in the New Zealand Subantarctic) were also considered to be auritulus s.s., Arthur (1960) who redescribed and figured the Snares Is. specimens and accepted their close affinity with auritulus s.s. but pointed out morphological features in which the New Zealand specimens differed. Dumbleton (1961) gave the Snares Is. (male holotype and female paratype) and
zealandicus of auritulus. Zumpt (1952) mentioned two females in the British Museum from a ‘dove petrel’ on Stephen Is. which he identified as auritulus s.s. These are probably zealandicus since they appear to be from the same host and locality as paratypes of this subspecies.
The anterior prolongation of the inner side of the first palpal segment of the female places the subspecies as belonging to the auritulus-percavatus group. The absence or weak development of the baso-dorsal spur on this segment in the Snares Is. paratype female (Arthur, 1960) suggests affinity with auritulus. This structure is however more strongly developed in two Stephen Is. paratype females, and its range of variation in pterodromae and other members of the percavatus complex is imperfectly known. The female of zealandicus differs from that of auritulus s.s. in the greater length and subrectangular shape of the 2nd and 3rd coxae and resembles percavatus rather than pterodromae in the absence of acute internal spurs on both of these.
The presumed male of auritulus s.s., subsequently described by Kohls and Clifford (1966) from an unknown host in southern Chile, was stated to differ from that of zealandicus in the more numerous hypostomal teeth (3-3 in anterior third, increasing to 6-6 or 7-7 at midlength, decreasing to base), the presence of a distinct median scutal elevation and a pit-like dorsolateral depression on each side of the scutum just before midlength, and the longer and more densely setose fore tarsi. As figured it differs also in that the anterior mesal margin of the epimeral plate does not terminate on the anterior mesal margin of the spiracular plate but continues to the 4th coxae. The median plate is longer in relation to its width (L-W 1.4) and is as wide at midlength as posteriorly. The anal grooves are strongly converging posteriorly and all coxae have small external spurs. These differences are greater than would be expected if zealandicus were a subspecies of auritulus.
The male of zealandicus moreover closely resembles the male of pterodromae described by Roberts (1964a) from nests of Pachypila desolata Gmelin on Macquarie Is., though the latter is stated to have a 5-5 — 6-6 hypostomal dentition.
The taxon is retained as a subspecies of auritulus only because the evidence is inconclusive. Specimens from the New Zealand Subregion which are morphologically identical with those of the nominate subspecies of auritulus from the Americas are also consistent in their host association in that they are from a land bird, but the occurrence of auritulus zealandicus on sea birds is anomalous and considerations of distribution also suggest that its relationships, whether as a distinct species or as a subspecies, may be with pterodromae.
Haemaphysalis longicornis Neumann, 1901, p.261, fig. 2 (Neumann 1901) Hoogstraal et al (1968) consider that the species from Australia, New Zealand, Fiji and Noumea, determined as bispinosa is not bispinosa of Neumann (1897), and have resurrected for it longicornis Neumann 1901.
I am indebted to Messrs M. Fitzgerald, J. R. Jackson, and P. M. Johns of Christchurch for the gift of specimens, and to Dr.
A plea is made for more adequate descriptions of extant leaves as an aid for overseas and New Zealand palaeo-botanists who have no or limited access to New Zealand material. As a guide, a list of preservable dicotyledonous characters is given. An experiment on the identification of extant leaves carried out in 1968 is briefly discussed in an appendix.
It is confirmed that the ratio of entire to non-entire leaf margins forms an important ecological indicator that is not invalidated by a lack to taxonomic determinations.
Generally speaking the taxonomic descriptions of plant taxa deal inadequately with the morphology of their leaves. Venation characteristics, which include some of the main features used in the identification of plants from leaves, are often omitted. For example, the description of leaves of
I do not wish to press this point too hard because arguments can be put forward for better descriptions of other parts of plants, e.g. seeds, fruit, wood anatomy and morphology. There are, however, two main reasons why, from a palaeobotanical viewpoint, I consider it necessary to improve leaf descriptions of extant plants: —
The abundant fossil angiosperm leaves, in Quaternary and Tertiary sequences, are very difficult to study without detailed knowledge of the morphology of extant leaves. Reference material and literature are not always available.
Study of fossil leaves may lead to a better understanding of the present ecological distribution of species when used as a supplementary technique to palynology in investigations of past distribution and palaeoclimatology.
One of the most accurate ways of describing leaf characters is by illustration. Clearly it would be impractical to include photographs and/or drawings of all species in the various books on the
This section is essentially a review of an excellent paper by J. A. Mouton (1966/1967) which is not apparently well-known to New Zealand botanists, supplemented by work done by myself in 1968. Madler and Straus (1971) have done similar work and have prepared a list of 109 characters for use in the description of leaves.
The study of leaf systematics poses many complicated problems that are not discussed here in any great detail; this has already been done elsewhere by Mouton (1966/1967) and Wolfe (1971).
Firstly, there are problems inherent in both fossil and extant leaf material:
1. Variability in size and shape:
This may be caused by growth deficiencies or excesses, periods of arrested growth, polymorphism and heterophylly. The New Zealand flora has a high percentage of juvenile leaf forms, some forms lasting well into the adult stage of tree or shrub growth. Some 10 per cent or about 200 species have this character (Millener, 1960). Also, species of many New Zealand genera are unusually diverse, both morphologically and ecologically, e.g. Coprosma, Hebe, Senecio, Plagianthus, etc.
2. Convergence of forms from different families:
Parallel development may be inherited or environmentally induced. Many of the strange features of the New Zealand flora involving parallelism have been listed by Millener (1960). They include cushion plants, climbing plants, epiphytes, divaricating plants and others, often occurring in genera which do not exhibit such habits elsewhere. The conifer-like appearance of the whip-cord hebes is a further example of convergence.
Secondly, there are problems peculiar to fossil leaves:
3. Loss of characters by fossilisation:
This problem could be rather serious in New Zealand since tectonic activity, even in Quaternary rocks, has caused small scale
4. Distinguishing compound leaves from simple leaves.
5. Present method of identification:
Fossil leaves are simply compared with extant herbarium material without detailed study of either.
Many characters used in the description of fossil leaves can also be used to identify extant leaves. These are as follows:
Well-preserved fundamental characters:
Primary and secondary venation
type
number of pairs of secondary veins
distance between veins
mode of attachment of secondary veins to midrib
angle of emergence of secondary veins (where parallel)
density (see Manze, 1968).
Tertiary venation
type
angle of emergence from base of secondary veins.
density (see Manze, 1968).
General leaf form and position of midrib.
Primary and secondary domatia.
Leaf margin.
Well-preserved characters of secondary importance:
Leaf biometry
surface area
length to breadth ratio.
Morphology of base and apex.
Secondary intercalary veins.
Geometry of the areole and the intersecondary field.
Poorly-preserved characters:
Biometry
length of leaf blade
apical ratio (length of drip-tip to length of leaf blade)
Basal ratio (length of base to widest part of leaf blade to the length of leaf blade).
Petiole length and presence or absence of a pulvinus.
Internal glands and cystoliths.
Hairs and stomata.
Phyllotaxy and stipules.
These characters can all be recorded in a punch card system. A preliminary attempt has been made to do this for all extant New Zealand dicotyledonous species. At this stage the cards only contain characters used in published descriptions so that little is yet known about venation details. Only a few species are based on herbarium specimens as the N.Z. Geological Survey does not
There are about 1,300 extant dicotyledonous species in New Zealand, including the off-shore and Sub-Antarctic islands. Disregarding altitude and overlapping ranges (both geographically and in the size range of the leaves) the distribution of species and their margins and leaf size classes can be set out as follows:
Further, and probably more meaningful, results can be obtained by dividing the lowland trees into their botanical districts and working out the distribution of various leaf classes from north to south.
The percentage of plants with particular characteristics or combinations of characteristics can be readily worked out. Of particular interest to palaeobotanists and palaeoecologists are the percentages of plants with leaf morphology associated with particular climatological and ecological niches. For example, plants in tropical areas generally have large (mesophyllous) leaves with long drip-tips, irrespective of their position within the various forest strata. In a normal tropical rain forest about 70-80% of the species fall into this category (Richards, 1952; Wolfe, 1971). Taking the overall flora in New Zealand and off-shore islands there is a decrease in the percentage of species exhibiting acute apices from the Kermadecs to the Sub-Antarctic Islands. The following table shows that the percentage does not reach more than 60% suggesting that the New Zealand forests could not be regarded as tropical in this respect. The percentages could reflect tropical affinities as they are characteristic of sub-tropical floras, especially those listed by Wolfe (1971).
Table listing percentages of species with acute apices. The percentage of species that exhibit acuminate and/or acicular apices are shown in brackets (based on information in Allan, 1961).
If the same and similar studies could be applied to fossil leaf material then comparisons could be made with the present day climate. In this case it would not be necessary to identify the leaves but just to distinguish between different leaf forms. A preliminary attempt has been made on a fossil flora from Whakarongo Stream, which drains into the Waikato River (N51/f894). No serious attempt was made to identify the leaves but twenty-five different leaf forms were recognised.
The marginal conditions and leaf size ranges are given in the following table and are compared with present day data given by Dawson and Sneddon (1969) for a forest covering the Maungataniwha Ranges, near Kaitaia, some 280 km to the north.
The percentages of the marginal characteristics of the Kaitaia Forest include compound leaf types which were not distinguished in the fossil flora. The fossil flora has more mesophyllous and entire leaves than the Kaitaia flora, suggesting warmer temperatures, and the temperature difference would presumably be even greater at the locality of the fossil flora.
For further assessment of the palaeoclimate, the pollen flora was studied to see if it supported the climatic interpretation based on the leaves. The pollen also assisted in determining the age of the rocks. The pollen spectrum was dominated by extinct beeches (Nothofagus) of the 'brassi' group (some of the fossil leaves were of extince ‘brassi’ group beeches with typically broad leaves).
In fact 47% of the total pollen flora was of this pollen form, suggesting warmer temperatures than at present in the same locality, since the Nothofagus ‘brassi’ group is now found in New Guinea and New Caledonia in warm moist conditions (although in climatic zones no different from North Auckland today). The presence of this group, as well as other extinct species — Haloragacidites harrisii (Couper) Harris, H. trioratus Couper, Proteacidites minimus Couper (cf. Knightia excelsa R. Br., which was found as a leaf fossil),
Slide number L5736. Locality N51 / F895
Adopted floral age: Opoitian to Waitotaran
Investigations have shown that other leaf floras from the South Island, ranging in age from Miocene to Eocene, exhibit a high percentage of leaf forms with entire margins but most were microphyllous in size. These floras may represent slightly cooler conditions than the one described above but more detailed investigations are needed, particularly of the associated pollen flora. An Upper Eocene sample from the Temuka Pottery Clay Pit contained a microphyllous flora with serrate/dentate leaves forming a large percentage.
I hope that these results will persuade taxonomists to give fuller descriptions of extant leaves enabling a more precise identification to be made of fossil leaves and thus allowing a more meaningful interpretation of palaeoclimate. It is possible that leaves of present day species were much larger in the warm phases of the upper part of the Tertiary and possibly Quaternary. Since this method has been used with great success overseas, particularly in America (Wolfe and Hopkins, 1967; Wolfe 1971), it is certain to be of value in New Zealand, where plants either had to migrate or change their habit during unfavourable periods in order to survive.
In order to discover if botanists can identify leaves without being able to see the growth form of the parent tree or shrub, or the colour, taste, feel and smell of their leaves, plaster-cast moulds of one hundred native New Zealand leaves were made and given to five botanists to identify. The botanists all had some experience in New Zealand taxonomy and ecology and were given unlimited
The models were made by pressing a leaf into soft plaster-of-Paris, placing a strip of thin plastic on top and weighing the leaf down with additional plaster-of-Paris. Few leaves failed to form good impressions. After setting, the plaster-of-Paris easily separated along the ‘plane’ formed by the plastic.
The one hundred examples included some taxonomic replicas, i.e. different leaves from the same species. The results of this study are set out in the following table, arranged in taxonomic order after Allan (1961) to whom the reader is referred for author citations of the species.
o identified to genus
- identified to species
x incorrectly identified
A blank signifies that no attempt was made in identification
The botanists were reluctant to fully commit themselves into identifying many of the moulds, probably due to the amount of time that it would have involved. The plaster-cast models show far more and better accentuated characters than leaves fossilised and impressed under natural conditions, so that it would be easier to identify plaster-of-Paris models than fossilised leaves of the same species. Thus it is surprising that the five botanists could only identify 60% to genus and 55% to species, while individually no one scored more than 40% correct to genus. The result was even more surprising as the specimens were, with few exceptions, well-known, common, native species. Poor casts may account for the failure of all botanists to identify numbers 44 (Hoheria angustifolia) and 74 ( Dysoxylum spectabile).
We can look at the results from the point of view of operator error and the difference in their approaches to the problem, and from the view of leaf variability and difficulty of identification. For example, operator 2 attempted to identify 38 specimens and made only six mistakes while the other operators attempted 62-70 specimens and mis-identified 23-30 of them. Quite obviously operator 2 is the more accurate worker even though not proficent
Nothofagus species (Fagaceae) but no operator identified any of the Escalloniaceae (Ixerba, Quintinia and Carpodetus) even to genus.
Disregarding the operator variability, the problem caused by taxonomic replicas and the fact that 70% of the first fifty moulds were correctly identified to genus and only 50% of the last fifty, possibly suggesting a growing disinterest, there are some points which may be noted. These are as follows:
The following common and distinctive species were not identified at all — Agathis australis, Brachyglottis repanda, Carpodetus serratus, Corynocarpus laevigatus, Elaeocarpus dentatus, Fuchsia excorticata, Hoheria populnea, Olearia arborescens, Phymatodes diversifolium, Pseudopanax lessonii and
Only a few of the moulds, including Lophomyrtus bullata, Nothofagus solandri, Phyllocladus trichomanoides and
Vitex lucens was identified as
Rhabdothamnus solandri was identified as
Weinmannia silvicola was identified as
The three genera Carpodetus, Ixerba and Quintinia of the family Escalloniaceae were not identified at all.
Podocarpus, Dacrydium and Phyllocladus species were relatively easy to distinguish.
Metrosideros, a common leaf fossil, was readily identified in most cases to species and is readily distinguished from species of Pittosporum which were poorly identified.
Nothofagus species were identified readily.
The following general conclusions were made on the identification of fossil leaves.
To be sure of an accurate identification a palaeobotanist needs not only vein patterns, leaf margins, shapes and sizes (preservable characters), but also, in some cases, non-preservable characters as well as the use of extensive herbaria and personal experience.
Leaves of different genera may seem identical when fossilised.
Conspecific leaves of slightly varying shapes and sizes when found as fossils can readily be attributed to different species or genera.
A large herbarium of native and overseas plants must be available locally for comparisons of fossil leaves. For New Zealand the following overseas collections are essential for accurate comparisons:
Fagus, Ficus); an extensive collection of Malayan, Polynesian and Australian genera (especially such genera as Acacia, Casuarina, Protea, Banksia, Dryandra, Nothofagus ‘brassi’ group, etc.); and a collection of ‘primitive’ genera still surviving today (including Cycadaceae, various pteridophytes, Magnoliaceae, Winteraceae, etc.).
The inevitable possibility of mis-identification means that the palaeobotanist is giving names not only to true genera and species but also to varieties of leaf form within species. This could account for the large numbers of species within a genus in many overseas Cretaceous beds, particularly those described at the end of last, and the beginning of this, century. It would be realistic to admit that macropalaeobotany deals solely with form taxa throughout the geological column as is customary in palynology. Obviously the variation between congeneric specimens is far greater than conspecific leaves and this increases the magnitude of the identification problem many times.
It may be profitable to repeat this experiment using new plaster-cast models and more controlled conditions to test both operator experience and recognition of variations in leaf form. If any senior botanists would like to participate in this experiment I would be keen to hear from them. If five or more are willing to be tested I will prepare new models and send them to the interested parties.
Thanks are extended to the five botanists who gave generously of their time of participate in the experiment described above. The help of Dr
Edited by Howard M. Lenhoff, Leonard Muscatine, and Lary V. Davis. Published by the University of Hawaii Press, Honolulu, October 1971, 281 pp.
This book is the result of research on coelenterates done by the editors, together with a select group of graduate students, at the Hawaii Institude of Marine Biology during the summer of 1967. The book is presented as a series of papers, and is divided into four parts dealing with, respectively, growth and development of coelenterates; feeding behavious, food transport, and metabolism; endosymbiosis with algae; and calcification. Many studies, particularly those comprising parts two to four, provide useful examples of the application of quantitative biochemical methods, and autoradiography, to the study of some of the ‘classical’ problems of coelenterate biology. The presentation of the papers is concise, and illustrations are of very good quality. Graphs and Tables are well laid out and are easily read. Full references are given after each paper, and a subject index is provided. One criticism of the book, which probably reflects the time limit of the whole research programme, is that some of the studies are very short and tend to leave interesting questions unanswered. Nevertheless such studies at least give some direction to subsequent research in this field.
The specialised nature of this book limits its use primarily to those interested in coelenterate research. It would also be a useful reference work for anybody teaching invrtebrate biology.
Zoogeography has been a fruitful preoccupation of New Zealand biologists throughout the history of New Zealand biology, and this is probably no accident. New Zealand's geographical isolation, and some striking peculiarities in its biota, both have made zoogeography a fascinating and rewarding pursuit. Furthermore, taxonomy has a very strong heritage in New Zealand, and taxonomy and zoogeography are closely allied and interdependent fields of study.
Zoogeography comes at two levels. In modern times, P. J. Darlington Jr., exemplifies the synthetic zoogeographer and there seems little doubt that his training and experience as a taxonomist (even though he is an entomologist and much of his zoogeography has dealt with vertebrates), has been fundamental to the success of his zoogeography. Most of us must confine our zoogeographical dabblings to dealing with the zoogeography of particular taxonomic groups in which we have a special knowledge and prime interest. And this is where the groundwork of zoogeography is largely done.
Zoogeography is one of the great synthetic sub-disciplines of biology and it requires an appreciation and knowledge of many other fields, both within and beyond the realms of biology — taxonomy, phylogeny, genetics, evolution, ecology, meteorology, geology, oceanography, and so on. The zoogeographer must have at his fingertips some knowledge of all these fields to facilitate interpretation of the distribution patterns he observes. Above all he must have an appreciation of the principles of logic and must examine the whole of the available evidence if he is to avoid falling into difficulties. It is so easy to argue circularly, or to accidentally ignore a part of the evidence. For instance, Ball and Fernando (1969) looked at the distribution of Dugesia, a freshwater triclad, and found that it occurs on all the southern continents, and New Zealand. Customarily, triclads are intolerant of sea water and are very poor at crossing even small ocean gaps, and the occurrence of a species on both sides of a body of water is taken to indicate that the two land areas where the triclad species is present were formerly connected by land. So Ball and Fernando concluded that the Southern Hemisphere distribution of Dugesia was a product of the former existence of Gondwanaland. However, they also pointed out that Dugesia occurs on the Crozet Islands, a group of very isolated islands that seem to be oceanic and relatively young. This suggests that Dugesia must sometimes cross ocean gaps. This being the case, we must consider the likelihood that the distribution of Dugesia
could be due to transoceanic dispersal. I'm not saying that continental drift and Gondwanaland did not have anything to do with the dispersal of Dugesia, but the evidence from the presence of Dugesia on the Crozet Islands forces us to consider that continental drift may not have. There is a basic need to consider all the implications of any zoogeographical hypothesis, and in the case of Dugesia, we need to examine how Dugesia got to the Crozet Islands, and consider the impact of this on the primary hypothesis, that the range of Dugesia is a result of the existence of Gondwanaland and subsequent fragmentation and dispersal of the continent.
We must be aware of the dangers of zoogeography as a science, but not be put off by these dangers. In fact, I believe that it is one of the responsibilities of good taxonomists to develop theories of evolution, relationship and zoogeography for the groups with which they work. No other worker is as capable of deciding where evolutionary relationships lie, and most zoogeography is an interpretation of phylogenetic relationships, dispersal, and geology.
Mayr (1969) subdivided taxonomy into ‘alpha,’ ‘beta,’ and ‘gamma’ taxonomy. Alpha taxonomy is the description of taxa, especially new ones, beta taxonomy is the study of the relationships of taxa, and gamma taxonomy is the study of intraspecific variation, evolutionary studies and zoogeography. Some taxonomists argue that a taxonomist should restrict his attention to alpha taxonomy because so many new species remain to be studied and described. They argue this way because their specialisation in some taxon allows them to make their greatest contribution to biology by describing and classifying unknown species. Mayr (1971) questions this attitude, noting that no one is better equipped to derive generalisations from taxonomic data than the specialist — ‘Indeed in 99 cases out of 100, if he does not do it, no one else will do it either… Let us remember that almost all of the major questions of evolutionary biology discussed during the past 50 years were either raised by taxonomists or were a product of taxonomic findings, (Mayr, 1971). Taxonomists do the groundwork necessary for all zoogeography, and it is their loss, as well as biology’s as a whole, if they do not take the extra trouble to determine what their taxonomic findings mean in a zoogeographic sense. It seems to me that zoogeography is a taxonomist's responsibility, and it is unfortunate that many a fine taxonomic treatment of some group or other, complete with elaborate distributional data has no discussion of the zoogeography of the group.
Apart from the fact that taxonomists have much to say zoogeographically, it seems to me that the taxonomist can better understand and deal with his taxonomic problems if he looks at the zoogeography of the animals he studies. This is a dangerous area, as it is very easy to argue circularly, and very easy to be badly
Let's take a look at New Zealand moas. On the basis of morphological evidence, 28 species have been described, almost all recent or subfossil, so essentially contemporaneous in New Zealand (Oliver, 1955). And remember that it seems that New Zealand was a very small land area in the Oligocene (Fleming, 1962). So, probably since then, as the land area expanded again, the moa fauna evolved to reach a total of 28 species. From an evolutionary-zoogeographic viewpoint, it is very hard to understand how this number of very large and rather specialised birds could have evolved and lived contemporaneously in the small area of New Zealand. What is apparently clear to the ornithologists on morphological evidence is rather incomprehensible from an evolutionary-zoogeographic viewpoint. The ornithologist must have the last word, but zoogeographers can, I think, have useful things to say about the taxonomies and may ask questions that can assist the taxonomist. I would very much like to see a comprehensive series of measurements taken on available moa bones, and have them submitted to detailed computer analysis to see if any of the material can be tied together as growth, sexual or geographic variations. It seems most unsatisfying that New Zealand has 28 moa species but less than 100 other land birds of diverse character and habits, and that New Zealand has more than threequarters of the entire ratite bird group.
Recently, I have had a look at the scope of the galaxiid fish genus Brachygalaxias. Several species have, at times, been included in this genus. The type species in South America has one unique character, it shares another peculiarity with two Australian species that are very similar to each other, and shares a further peculiarity with both Australian species and a further species in South Africa, (unpublished data). The question the taxonomist must answer is where to limit the genus. But the answer to the question has far-reaching zoogeographic implications. If the Australian and/or South African species are included in Brachygalaxias with the South American species, we are indicating strong belief in close phylogenetic relationships, and posing a zoogeographic question for which we
Brachygalaxias creates zoogeographic problems that the morphological evidence on relationship probably does not warrant. The alternative is to retain the South American type species in Brachygalaxias, and to regard the other species as fringe species of Galaxias, which, on the basis of certain morphological evidence, may be regarded as related to Brachygalaxias. By choosing this alternative, the taxonomist is not forcing the zoogeographer to seek explanations of relationships and distribution patterns that may be a taxonomic artefact. And yet, the question of relationship, and the intriguing zoogeographic problems remain open for consideration. The interplay of taxonomy and zoogeography in this way should produce taxonomies that have the greatest utility and retrieval value for users of the taxonomic treatment, for zoogeographic or other purposes.
While concordance between taxonomy, phylogeny, and zoogeography is nice — it's very satisfying to draw a cohesive and consistent picture — I think that we must beware of organising taxonomies to produce nice, perhaps preconceived zoogeographic pictures. This is a very real danger. Taxonomies must be based on taxonomic evidence and taxonomic principles. If the taxonomy results in a satisfying zoogeography, that is nice. If it doesn't, may be we can and should take another look at the taxonomy, but we should not and must not reorganise the taxonomy to tidy up the zoogeography.
Zoogeography in our day is undergoing a profound revolution. The edifice of modern historical zoogeography is very clearly built upon the foundation of Darlington (1957) ‘Zoogeography — the geographical distribution of animals.’ This book deals only with land and freshwater vertebrates, and it follows as a basic premise, the view that the continents have always been roughly where they are now. And it is no discredit to Darlington that, largely since this book was written, geologists have become increasingly convinced that the continents have, in fact, not always been where they now are. Darlington, told by geologists, that the continental positions had not changed, had to rationalise the distribution of animals on this basis. Now, we are told that the continents have dispersed away from Gondwanaland, a large southern continent, and we will now have to establish a whole new set of rationalisations. Stable continent zoogeography, confined largely to vertebrates, left plenty of perplexing problems — characid fishes and ratite birds are two of the more horrendous zoogeographic problems for which neither Darlington nor anyone else has had any really satisfying answers. It is interesting that invertebrates, especially insects, had little wide-ranging attention from stable continent zoogeographers, and I suspect that the problems of characid fishes and ratite birds may have
The increasing confidence of geologists and geophysicists that continental drift is a fact, means that drift warrants, or rather demands serious attention from zoogeographers. The impact of this on zoogeography is going to be substantial. I believe that zoogeography has very little indeed to say about the former positions of the land masses, and at this point, we are very much at the mercy of the geologists. And if they say that Gondwanaland split up some time during the Cretaceous, and that various fragments moved in certain directions, at given rates, at various times in the past, then we have little option but to accept this, and investigate the impact of these theories on zoogeography. Perhaps zoogeographers will help to refine the dating of continental fragmentation using paleontological evidence, but I believe that we can have little more to say than that.
Following his 1957 synthesis of animal distributions on the basis of stable continents Darlington (1965) has begun to restructure his zoogeography. This 1965 book is a reflection of Darlington's efforts at an approach to continental drift zoogeography, built upon the foundations of his earlier work, and the tensions between these two mutually exclusive approaches are evident.
The acceptance of continental drift has, as I see it, three impacts on existing zoogeographical knowledge:—
It may help to explain the previously inexplicable.
It may lead to the replacement of old explanations for patterns of distribution by new explanations.
It may produce a whole new array of inexplicables.
Early attention is bound to be paid to the first of these, but I have little doubt that all three of the above will apply.
What is the impact of continental drift on New Zealand zoogeography? New Zealand is believed to have moved away from Antarctica about the Cretaceous, about 50-60 million years ago. It can be regarded as having been an isolated land area ever since, although the area of land is believed to have fluctuated broadly (Fleming, 1962). Thus the effect of continental drift, and the departure of the New Zealand land mass from Antarctica is related to: 1. The initial departure from Antarctica; and 2. The increasing distance between New Zealand and Antarctica.
There should be evident in the New Zealand fauna signs that New Zealand was suddenly cut off from other land areas. A whole series of taxa should show signs of having been contemporaneously isolated in New Zealand. However, because it was so long ago, the
The frog is another problem, as are the crayfish, mussels and earthworms. These seem unlikely to have been able to cross oceans in the way the tuatara could have. Their very presence in New Zealand, has been enigmatic, but probably can be explained by New Zealand's involvement in Gondwanaland.
Much of the New Zealand freshwater fish fauna has relationships with southern areas (McDowall, 1964). The Southern Hemisphere has a characteristic southern temperate freshwater fish fauna that is very small. This comprises the galaxioid fishes (Sub-order Galaxioidei, Families Galaxiidae, Retropinnidae, Aplochitonidae, Proto-troctidae) and the southern lampreys (Family Geotridae). The galaxioid fishes are a southern temperature radiation comparable with the northen temperate salmonoid fishes (Sub-order Salmonoidei, Families Salmonidae, Osmeridae, Plecoglossidae, Salangidae); but the lampreys are merely a family in the south comparable with a family in the north (Family Petromyzonidae). These southern freshwater fishes appear as if they could have attained their present distribution in relation to the fragmentation of Gondwanaland, and there has been much interest in the past especially in the distribution of the Galaxiidae (Australia, New Caledonia, New Zealand, South America and South Africa). This distribution has been taken as evidence that there have been connections between these
Galaxias maculatus) common to Australia, New Zealand and South America, without any evidence of even separate subspecies; or races. If the range of this species is related to Gondwanaland, how can we explain the conspecificity of these highly disjunct populations after about 50 million years of isolation? If we take another look, we also find that this species has a marine stage in its life cycle, so is easily capable of transoceanic dispersal. Boulenger pointed this out as long ago as 1902. but no one seemed to take any notice of him. Another galaxiid (Galaxias brevipinnis) is present in Tasmania, New Zealand and the Sub-Antarctic Islands, and it too has a marine stage in its life cycle. Species common to western and eastern Australia and/or Tasmania also have marine stages. If we look at the distribution of galaxiids in New Zealand, we find two types of distribution — widespread species and localised species — and we find that the widespread species have marine stages and the localised ones do not. All these facts indicate very clearly that there has been dispersal of galaxiid fishes through the sea (McDowall, 1970). The southern lamprey Geotria australis is found in western and eastern Australia, Tasmania, New Zealand, Chile and Argentina, and like the galaxiids, the lamprey has a marine life history stage; the conspecificity of populations in all these areas shows that it also has dispersed through the sea.
Caughley (1964) wrote a paper under the stimulating title ‘Does the New Zealand vertebrate fauna conform to zoogeographic principles?’ With the admitted tremendous advantage of hind-sight, Caughley asked us to go along with him as he constructed a theoretical New Zealand vertebrate fauna, on the basis of disperal from the faunas of likely contributing land areas. Caughley assumed that New Zealand is an oceanic island, ruling out land connections and continental drift, for his exercise. For what they are worth, his predictions were good, and his failures were heavily weighted towards the archaic elements in the New Zealand vertebrate fauna, i.e.: the frog, tuatara, moas and kiwis. These are the sections of the vertebrate fauna that are most likely to have had origins in Gondwanaland. As Caughley excluded land connections of any type, we would have expected him to have had difficulty predicting the presence in New Zealand of any flightless animals that don't disperse across the sea, and which must have reached New Zealand by land routes. Note that the group of animals that: Caughley had difficulty predicting for the New Zealand fauna is almost identical with the archaic element in the fauna Fleming, 1962).
Thus, to answer my earlier question, the impact of continental drift on the zoogeography of New Zealand, from the point of view of vertebrates, is relatively slight in terms of the entire vertebrate
One of the interesting impacts on zoogeography of continental drift is on the origins of bipolar groups. It has long been recognised that there are pairs of groups of organisms with one of the pair in the north-temperate — sub-Arctic and the other in the south-temperate — sub-Antarctic. Gill (1893), and more recently Hubbs (1953) have examined this subject, the problem being to explain how groups apparently intolerant of the high temperatures of the tropics, could have dispersed across the tropics. Hubbs suggested that some of the fishes could have crossed the tropics by descending into deeper, cooler water, and others may have done so when the tropics were cooler, as during the Pleistocene glaciations. But for many groups, neither suggestion is helpful.
For some old groups, e.g. the galaxioid (southern) — salmonoid (northern) pair, or amongst the plants the Nothofagus (southern) — Fagus (northern) pair, the former existence of a formerly united continental mass may assist the explanation of how and why there are northern temperate groups with obvious, closely related radiations in the southern temperate.
The field of New Zealand zoogeography is wide open, and as we see the taxonomy of New Zealand invertebrates develop, I think we will see a great flowering of New Zealand zoogeography. If this flowering had taken place 20-25 years ago when the continents were regarded as having been stable, and when we knew little about the taxonomy of our invertebrate fauna, I think that we would have had zoogeographical chaos. Now, the time seems ripe for this zoogeography really to flourish.
The arrival of a live male specimen of tuatara (Sphenodon punctatus punctatus (Gray) at our Department early in August, 1963 provided an opportunity for the study of the following aspects of that animal. Firstly, there was the problem of the animal's reaction to the sudden reversal of its daily and seasonal rhythmicity following a flight from New Zealand to Poland of some 40-50 hours. Secondly, as the existing information on the tuatara's biology is still relatively scarce, an attempt was made to collect more data. Finally, we were interested in the susceptibility of the tuatara's senses and particularly to find out whether the animal's parietal eye would react to thermal rays.
Our specimen was an adult male of the typical form of S.p. punctatus (Gray) conforming to the descriptions by Wermuth and Mertens (1961) and Dawbin (1962). It was kept in a large metal vivarium of 110 × 70 × 70cm, specially constructed, provided with glass panes on two sides, netting on the remaining sides and a 25cm high ridgeroof. During the first months, the vivarium's bottom was covered with a 15cm deep layer of soil to allow the animal to burrow underground gangways and holes. At a later stage when the tuatara became infected with a kind of scurvy, the soil was removed, the vivarium was disinfected, a wooden shelter was added and the animal remained in this accommodation until the end of its life.
1. Food and Water
Initially our tuatara was fed with earthworms, which it took readily like Merten's (1958) specimen, but once a start was made feeding it with larvae of meal worms (Tenebrio molitor), it much preferred these to earthworms. At a later date, our tuatara, like that of Werner (1926) also took young, still naked mice. An attempt was made to feed the tuatara with snails but it refused to eat them and even drew away their mucous bodies with obvious dislike each time a snail managed to creep on to it.
Our tuatara maintained a reasonably good appetite while it remained in good health. For instance, in August, 1963 it consumed a record daily amount of food consisting of 17 earthworms and 29
The tuatara regularly took some water but the amount consumed could not be recorded. In natural conditions the tuatara has been reported to consume water but mainly after eating marine foods. Our specimen liked bathing in a tank provided for this purpose in the vivarium. It also swam well though it disliked diving.
2. Hollow Dens
In natural conditions the tuatara dwells in burrows which it digs out in soil, sand or clay. In the vivarium the tuatara dug out initially one gangway some 60-70cm long running horizontally though somewhat obliquely in which it could hide itself. Its usual position was to have only its head sticking out of the burrow's entrance. At a later stage, our specimen dug out another gangway which it used in the same way as the first. When digging the animal used each foreleg alternately, throwing soil to the back. Upon leaving the burrow it was covered with dirt, but this was soon washed off by bathing in the tank. One of the gangways had a second opening and could be used by the animal as a tunnel.
3. Defence
The tuatara defends the entry to its burrow against any intruder. A hand levelled at the burrow's entrance was immediately attacked by the tuatara's strong teeth. Mertens (1958) recorded in his male tuatara such aggressive behaviour that he could not get close to it. Our specimen, though also a male, behaved differently: from its arrival it was quite tame and did not attack anything and sometimes would even accept food presented on forceps by its keeper. Exceptional behaviour was noted when the animal was being caught: it assumed a warning position by lifting itself on its forelegs and opening its mouth widely to show its teeth and also its strong, thick tongue. This threatening behaviour was followed by an attack with a sudden, sidewise twist of the head allowing the hand to be caught from one side. In addition to this type of defence, the tuatara can hit its adversary strongly with its snout to the extent of causing bruises. An excited tuatara, particularly when being caught utters sounds that can be described as short, strong croaks.
4. Curiosity
Despite an association with marine birds living in the same burrow, the tuatara has an essentially solitary way of life, and except during the mating season one does not find two animals together. Males are known to be unfriendly towards other males
5. Sounds
The tuatara utters two different sounds: one emitted in excitement is not unlike a loud short croak; and the second sound is much weaker and reminds us of a growl or groan. A bioacoustic analysis of the first sound has been given elsewhere (Wojtusiak and Majlert in press) but there was no opportunity for recording the second sound, nor for its analysis. It is possible that the second kind of sound may be of some importance during mating displays or during feeding but no proof has been found so far for this.
6. Senses
A. Sight: Our observations show that the sight of the tuatara is well developed. The presence of food could be recognised from a distance of 50-60cm and this would confirm previous observations (Werner, 1912 and 1925). However, in catching a prey it seemed to be more directed by movement than the perception of shape. It has been reported to have better sight at night and this prompted us to experiment with the sensitivity of this reptile to infra-red radiation.
B. Sensitivity to Infra-red Radiation. The tuatara has a well developed pineal gland or parietal eve, the structure of which reminds us of a third eye (Dendy 1910 and 1911, Stebbins and Eakin 1958, Stebbins 1944, Cans and Parsons 1970). It is well developed in young specimens and not covered by scales (Anonymous 1954) which grow upon it at the age of six months. The presence of a retina and lens (Gabe and Saint Girons 1961) in the parietal eye located beneath the skin but not covered by one, make it conceivable that it could function as an eye sensitive to infra-red radiation and be of assistance to the animal in orientating itself during the night. According to Anthony (1970) it was Ruckard (1886) and Francotte
C. Other Senses. It would appear that the tuatara's taste is also well developed. The preference for mealworms over earthworms, which the animal refused to eat, would support this statement. This is in accordance with observations of other authors (Werner 1913 and 1923 and Wettstein 1931) who reported cherries taken being distastefully spitted out, as were snails in the case of our animal. This is interesting because in its home, the tuatara has been reported to consume snails of the species Janella schaumslandi Plate, but European snails were obviously disliked.
7. Daily (Twenty-four hours) Activity
The tuatara is a nocturnal animal and this is indicated by its pupils which narrow during the day into a vertical slit. In subdued light the pupil is round which, together with the relatively large size of its eyes gives the tuatara a more “human” expression than other reptiles. Merten's (1958) male specimen was timid, hiding either in a burrow or in a shelter made out of boards which it left only at dusk. It took that specimen several months before it would emerge in the daytime and it never came out in sunlight or even during a bright day. Our tuatara did not avoid people, did not hide nor avoid diffused light or sunlight. It was active during the day taking food, climbing up the branch set in the vivarium, bathing in the tank and resting on the ground of the vivarium.
It should be borne in mind that our tuatara had been shifted from the full winter in the Southern Hemisphere to mid-summer in the Northern one. As there is between New Zealand and Poland a time difference of about 11-12 hours, our tuatara, which in its home country was active at night, had fairly suddenly found itself at the same time of the day in conditions of full daylight.
The behaviour of our tuatara described above seems to indicate that the animal during the first weeks or even months of its stay in Poland adhered to its former activity period which involved a change from nighttime activity in New Zealand to daytime activity in Poland. The fact that the tuatara did not avoid sunshine or light would indicate that its nocturnal activity is not entirely dependent on the day and night rhythm. Several months later our tuatara would usually leave its burrow in the morning, about noon and in the evening. A particularly high nocturnal activity occurred during the heatwave during the summer. This was shown by the soiling of the vivarium's walls: foot marks could even be seen on the hip of the vivarium's roof which would indicate a climb of 60-70cm above the terrarium's surface.
Observations carried out by Madame J. Gieszczykiewicz during the first four months of the tuatara's stay in our laboratory provide some information on the gradual change in its activities.
During this time the animal had been kept at a constant temperature of about 20 C and meteorological observations were recorded on a thermo-hydrograph and a barograph. Lack of space prevents us from including diagrams from which we can conclude that changes in atmospheric pressure and humidity had little or no effect on the daily activities of the animal.
The activity of the tuatara has been divided into the following five categories:
1 Rest or inactivity 2 Walking or bathing 3 Eating and drinking 4 Burrowing 5 Climbing and attempts to escape.
As a daily formulation of the observations taken would present a somewhat confused picture, the results have been compiled in ten-day periods, which are set out in Table 1 and Figs. 1-3.
Fig. 1 shows that initially our tuatara's greatest activity occurred during the early hours of the morning when it was enjoying all kinds of activity from walking to climbing and attempts to escape. Gradually we observed a shift of activity into the afternoon and
Fig. 2 shows the totals of the number of hours of activity and
It would have been of considerable interest to find out how the tuatara's activity gradually shifted into the night, unfortunately no appliance for an automatic registration of its activity during the night was available.
8. Annual Activity
In New Zealand the tuatara was reported to hibernate for a four-month period, from the middle of April to mid-August. While hibernating, the tuatara remains in its burrow and does not take food. However, during warm days it is known to leave the burrow and even to prey for quarry. However, to achieve full activity it is known to require a temperature of at least 10°C and it does not leave its burrow at temperatures below. Upon the tuatara's arrival from New Zealand, despite the change of season, it immediately exhibited quite considerable activity: it walked around the
In concluding, it would appear that the rapid adaption of our tuatara to its new environment was facilitated by the summer conditions and particularly by the high temperatures that it encountered on its arrival in Poland.
9. Adaptation to Temperature
The tuatara differs from other reptiles in its adaptation to relatively low temperatures (Batham 1960). Its optimum ambient temperature is between 12 and 13°C which according to Bogert (1949) and Mertens 1958) conditions its longevity. Our specimen came from Stephens Island, where April temperatures are reported to be between 9.4 and 14.0°C with a mean of 12.2°C, but in November, when tuataras are most active, temperatures from 8.8 to 13.7°C, with a mean of 11.4°C occur. The tuatara's body temperatures recorded in November (Dawbin 1949 and Bogert 1953 a and b) were 7.6° to 13.3°C with a mean of 10.6°C.
Our own observations seem in indicate that at temperatures below 16°C a decline in the tuatara's activity was noticeable. Temperatures
10. Illness and Death
The tuatara lived in our laboratory for nearly three and a half years and died on February 9, 1967, as the result of disease. Upon the animal's arrival the presence of several, large grey-green, 5mm wide ticks, well adapted in colour to the skin of the host, were noted and removed. (Apedonoma sphenodoni). The scurvy of the mouth proved to be a serious ailment. From the pus the following bacteria were identified: the pathogenic Staphylococcus pyogenes, var. aureus and two saprophytic bacteria Micrococcus sp. and Sarcina lutea. An antigramme of S. pyogenes has shown an average sensitivity to standard antibiotics.
Our tuatara was attended by veterinary surgeons, trained in pathological aspects of zoological gardens. Fig. 4 shows the regeneration after the operation on the upper jaw. During its convalescence, the tuatara resumed feeding and soon recovered a healthy appearance. However, later it became ill with scurvy again and this time the scurvy attacked the inner organs and caused its death. It would be of some interest to find out whether similar cases of scurvy occur not only in captivity but on Stephens Island and other islands with tuataras in New Zealand.
The specimen and its skeleton have been presented to and are deposited in the Zoological Museum, the Jagellonian University in Cracow.
Our specimen of tuatara was presented by the Government of New Zealand to the Jagellonian University in Cracow on the occasion of the 600th anniversary of its foundation. I would also like to thank Professor Zygmunt Grodzinski, Director, K. Hoyer's Institute of Comparative Anatomy, Jagellonian University for kindly handing the specimen to our Laboratory and for reading early drafts of this text.
Our thanks are due to Mr D. M. Luke, New Zealand Department of Internal Affairs for all the arrangements connected with the catching of the specimen and its transport from Stephens Island
Last but not least I would like to thank my friend and colleague Professor K. Wodzicki, F.R.S.N.Z., for his initiative in the procurement of the specimen and for the translation of the text into English.
According to local fishermen, sponges containing small fishes are sometimes found in Tasman Bay, New Zealand. This was confirmed on February 9, 1970, by a trawl catch from that area, taken at a depth of 40 m and containing, among other things, many small tarakihi, Cheilodactylus macropterus (Bloch and Schneider), and a quantity of the sponge Suberites australiensis Bergquist. I found small fishes in the exhalant cavity of two sponges: one contained three tightly packed tarakihi, the other a scaly gurnard Lepidotrigla brachyoptera Hutton. All these fishes were between 10 and 12cm long. They sat in the cavity with the head inwards, and were almost completely enveloped by the sponge. On the outside only the tails could be seen, protruding through a narrow opening (Figs. 1-3). The inner end of the cavity occupied by the scaly gurnard had the same size and characteristic angular shape as the head of the fish itself, so that the cavity was almost a
It is unlikely that these fishes used the sponges for shelter, because then they would have been more likely to occupy the cavity with the head outwards. The cavities were so narrow that, once inside, the fishes could not have turned round. A more likely explanation follows from the fact, mentioned to me by Professor P. R. Bergquist, that the exhalant canals of Suberites australiensis are sometimes inhabited by small animals such as worms. Tarakihi feed on this kind of prey and may thus be attracted to enter the exhalant cavity of the sponge.
Suberites australiensis has much contractile tissue around the exhalant cavity. If irritated by the presence of a fish inside the cavity, the sponge may contract in an effort to rid itself of the source of irritation, with the result that the fish becomes trapped and will remain so unless the sponge relaxes again. Both tarakihi and scaly gurnard have spines which may make it difficult for them to move backwards through a narrow opening in spongy material, and wriggling movements intended to free the fish may have the effect of pushing it further inwards and of irritating the
The tendency to explore narrow openings in search of food has been observed in other fishes. For instance, the yellow-eyed mullet, Aldrichetta forsteri (Curier and Valenciennes), can be caught in baited milk bottles, and a ‘bait-catcher’ (a small cylinder with two funnel entrances) sold commercially in New Zealand operates on the same principle. The presence of one fish in such a trap seems to attract others, because many can be caught at a time. This may account for the presence of three fishes in one of the sponges. I have also found a tarakihi of 12cm trapped inside a somewhat larger salp (Salpa sp.) which shows that this phenomenon is not restricted to sponges but can also occur in other animals.