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Now at University of New England, Armidale, N.S.W. Publication of this paper has been assisted by a grant from the Victoria University Publications Fund.
Now at University of New England, Armidale, N.S.W. Publication of this paper has been assisted by a grant from the Victoria University Publications Fund.
Mystacina tuberculata Gray is represented in New Zealand by two subspecies. Differences between these in overall size and in relative proportions of extremities of the body are discussed.
Mystacina tuberculata Gray is a small bat with a close, compact layer of fur which is generally frosted in appearance. The elongate head tapers conspicuously forwards to prominent nostrils. The simple ears have a long, narrow tragus. The thin flight membranes are exceptionally tough and the reduced propatagium, medial portions of the plagiopatagium, and the basal part of the uropatagium are conspicuously thickened. With the exception of the slender metacarpals and phalanges of the chiropatagium limb elements are robust; the feet being very large and turned outwards. The short tail penetrates the uropatagium dorsally.
This species is represented in New Zealand as two distinct forms which are here described as subspecies. One ranges throughout the North Island and is present in at least northern areas of the South Island. but the other is at present only recognised from Stewart Island and a few neighbouring islets. A specimen from Okarito, Westland, was in too poor condition to enable subspecific determination.
In Vespertilio tuberculatus G. Forster of .the family Vespertilionidae, which he described as "Yellowish brown; ears small, rounded. Inhab. Dusky Bay, N.Z." In an appendix on page 296 of the same journal he stated that having received two specimens of the New Zealand bat he found it belonged to a new genus and accordingly described it as Mystacina tuberculata. He maintained this synonymy in the Catalogue of Mammalia in the British Museum (Gray,
Had Gray been able to examine Forster's Vespertilio tuberculatus he would have realised that his animal was distinct. It remained, however, for Tomes (Mystacina tuberculata Gray. Later Tomes (Mystacina to the family Phyllostomidae but Gray (Mystacina velutina be adopted in reference to the velvet-like nature of the fur of the short-tailed bat.
Dobson (Mystacina tuberculata to belong to the family Emballonuridae, but to differ from typical members of that family in possessing a third phalanx in the middle finger, a feature which he described for the American Phyllostomidae. He separated Mystacina as a group Mystacinae (subfamily Molossinae) of the above family.
Lydekker (in Flower and Lydekker, Neave (Mystacina Gray was preoccupied by Mystacina Boie, Mystacinus Boie, Mystacina Gray.Mystacops tuberculatus (Gray). His change was generally adopted. Winge (Mystacina to the Vespertilionidae. Thomas (Vespertilio tuberculatus this made Gray the author of that name and at once invalidated Gray's own name for the short-tailed bat. On these grounds he reinstated Hutton's name of Mystacops velutinus (Hutton). Miller (
Eleven specimens of Mystacina were available for examination. These are itemized in Table I, in which the abbreviations D.M., A.M., C.M., and V.U.W. are used to indicate Dominion Museum, Auckland Museum, Canterbury Museum and Victoria University of Wellington collections respectively. I would like to thank Dr R. A. Falla, Dr G. Archey and Mr E. G. Turbott for making this material available, and Professor L. R. Richardson for his supervision and guidance throughout the study.
Muzzle conical, obliquely truncated, with conspicuous array of stiffened hairs radiating from posterior margins of glandular eminences. Prominent nostrils opening sublaterally. Ears simple and extending well beyond the fur of the head; tragus long and attenuate. Fur close, grey to black. Coarse overhairs, with long distal thickening, scattered amongst short, wavy underhairs. Scale form entire to repand coronal. Robust forelimb; upper arm slightly less than 3/5 forearm length. Long metacarpals shorter than forearm. Second digit with single rudimentary phalanx Third digit with three bony phalanges. Cartilaginous tip of fifth digit extending beyond margin of chiropatagium. Wings from metacarpal of thumb and from ankles. Propatagium reduced and thickened. Plagiopatagium and uropatagium with numerous ridges on upper surface of proximal portions. Leg short and robust with large feet about 3/4 shank length. Basal talons present on the claws of toes and thumbs. Short tail perforating uropatagium. Weak calcars.
Range: North Island, N.Z., and at least northern areas of the South Island.
A small delicately proportioned bat characterised by the length of the ears, which reach to or beyond the tip of the muzzle, and by the remarkably prominent, though narrow, nostrils. Total length up to about 75 mm, wing span to about 280 mm. Gray's plate (
Range: Only known from the Stewart Island region, N.Z.
In contrast to the northern subspecies this animal is larger and extremely robust.
Type. Male from Big South Cape Island, Dominion Museum Specimen
Table I shows the relationship of ear length (from junction with head and from meatus), forearm length, digital length (third metacarpal and associated phalanges) to total length for the eleven specimens of Mystacina available during this study. Comparable date from the literature has been included and where possible the subspecies has been indicated. "Ear length" of Hutton (
With the exception of this last specimen the values for ear or forearm length compared with total length fall into two groups corresponding with the known ranges of the two subspecies. Ear length from junction with head as a percentage of total length is above 20% in M. t. tuberculata but below this value in M. t. robusta. The values for ear length from meatus are more distinct, above 25% in the northern subspecies, below 23.5% in M. t. robusta. For the forearm corresponding values are 62.5% and 60%. The relationship between digital length and total length is not so marked. Values for M. t. robusta are generally lower, but Knox's (
The distinction between the subspecies relates both to overall size and to the relative proportions of exposed parts such as nostrils, ears and limbs. Overall size is greater and the extremities relatively shorter in M. t. robusta than in the northern M. t. tuberculata. Such differences are in accordance with Bergmann's and Allen's et. al., et. al., that hibernation may reduce the effect of temperature upon structure in mammals by avoiding exposure of the animals to winter minimum temperatures. It is of interest, therefore, that there is some evidence that Mystacina does not undergo a strict period of hibernation during winter months.
Mystacina tuberculata Gray is one of the two bat species present in New Zealand, and is the sole representative of the family Mystacinidae. Some attention has been directed to a number of peculiarities in this bat which have been interpreted as adaptations suitable for terrestrialism. These conclusions were first indicated by Dobson (Mystacina hunts for its insect food, not only in the air, but also on the branches and leaves of trees, among which its peculiarities of structure most probably enable it to walk about with security and ease."
Dobson stated that in repose the first phalanx of the third digit is folded forwards upon the ventral surface of the metacarpel, the second phalanx is folded backwards on the first and the third forwards on the second. Hutton and Drummond (
The following description is based upon a single female specimen of M. t. tuberculata in which the left wing was in the position of repose. The form of the metacarpophalangeal and the phalangeal joints would not permit the wing to fold otherwise. (Figs. A, B, C.)
In folding, the membranous portions of the wings are carried beneath the forearms and against the body and the uropatagium is rolled forwards beneath the tail. Ventrally the reduced propatagium extends beneath the stout forearm as a strong band which takes origin from the metacarpal region of the thumb. On each side this is produced beyond the posterior margin of the forearm so that a distinct concavity is formed between the band and the plagiopatagium. The delicate proximal portions of the wings are concealed within these concavities. The tips of the wings are contained within small lateral pouches present at the sides of the body, just forward of the thighs, and extending along the underside of the thighs. Only the thickened leathery portions of the wings remain exposed. Behind the wrist the ventral and medial aspects of the folded palm are protected externally by a thickened region of the membrane between the second and third metacarpals and by the fifth metacarpal together with a narrow, thickened ridge present immediately lateral to this metacarpal. This ridge extends onto the upper surface of the membrane for almost half the length of the metacarpal.
The folding process commences with the proximal phalanx of the third digit being turned inwards beneath the membrane. This carries with it the proximal phalanx of the fourth digit and the second (terminal) phalanx of the fifth digit. The short first phalanx of the fifth digit remains extended along the line of the metacarpal. The second phalanges of the third and fourth digits, and the flexible
The distal portion of the uropatagium is rolled forwards beneath the tail so that it lies close against the body at the base of the tail. Only the thickened basal portion remains exposed. The calcar lies along the inner surface of the shank.
Mystacina is not, as far as can be ascertained, a crevice dweller, and the extreme protection of the delicate membranes from possible injury correlated with a maximum degree of freedom for the limbs must be interpreted in relation to terrestrial locomotion. Similar protection of wing membranes occurs in the molossid Cheiromeles torquatus with the wing tips being folded into pouches, but here the pouches are in the armpits and open backwards (Kitchener, Eumops, for instance, is unable to take flight from a level surface (Vaughan, Mystacina freedom of the limbs provided by the reduced propatagium and wing folding process, the robust limbs, the specialized claws, and the wide range of movement of the femur permitted by the form of the acetabulum are terrestrial or arboreal adaptations. The ability to take flight from level ground has been recorded for Mystacina and terrestrialism in this bat is not connected with roosting but is rather to be associated with foraging habits. The scanty information available pertaining to the ecology of this species does appear to substantiate Dobson's earlier conclusions.
Abbreviations: 3m, 4m, 5m, metacarpals of digits 3, 4 and 5; 3 ph. 1, 3 ph. 2, 3 ph. 3, phalanges of digit 3; 4 ph. 1, 4 ph. 2, phalanges of digit 4; 5 ph. 1, 5 ph. 2, phalanges of digit 5.
Hair structure of Chalinolobus tuberculatus (Forster) is distinct from that of C. gouldii Gray; in some respects it agrees with the generalized vespertilionid structure but differs in the lack of separate overhair and underhair. In Mystacina tuberculata Gray, both hair types are present; overall structure is clearly different from that of the Molossidae but corresponds closely to that of the Noctilionidae and Nycteridae.
The present paper describes the hair structure of the two New Zealand bats Chalinolobus tuberculatus (Forster), F. Vespertilionidae, and Mystacina tuberculata Gray, F. Mystacinidae, and examines the affinities of these species in the light of this. Hairs from the mid-dorsal region at scapula level were examined and sketched using a camera lucida. The terminology of Benedict (
In C. tuberculatus there is no differentiation of hair as overhair and underhair. The long, fine filaments are of fairly uniform diameter and lack a medulla. Filament length is from 4 to 7 mm with the maximum diameter, 15μ, occurring both above and below the midpoint (diameter 12μ). Distal scales are about 15μ in diameter and the hair ends as an acute projection formed from two or three incompletely developed scales.
Melanin is concentrated in the proximal half of the filament; the maximum density occurring about a third of the distance along the hair. Within each scale the melanin is arranged dis.tally except in the distal portion of the hair where the few granules are scattered irregularly. Scale arrangement is annular. Scale form in the proximal half of the hair is medium, divergent or divaricate, equal or unequal hastate coronal but distally it becomes short, slightly divergent, equal hastate coronal. The terminal scales are slightly longer than those preceding them.
The hair structure of C. tuberculatus agrees in some respects, such as the hastate coronal scale form, with the generalized vespertilionid structure described by Benedict (C. gouldii as being the only bat in which the hair scales show entire margins in the proximal third and denticulate margins thereafter. In this respect and in the coarser nature of the filaments C. gouldii is strikingly different from C. tuberculatus. In the lack of differentiation of overhair and underhair and in the arrangement of the melanin within the filament the two species agree. The two species of the African Glauconycteris described by Benedict are clearly separated from Chalinolobus in details of hair structure. In both G. humeralis and G. varigata papilio overhair and underhair are present and the melanin is dispersed generally, not in bands proximally as in Chalinolobus. The scale form of the underhair is short, appressed, hastate coronal except in the distal third of the filament of G. varigata papilio where it becomes lobate.
The two trends away from the typical vespertilionid scale form which occur in these closely related genera are represented more completely in other genera. Thus Scotophilus has lobate coronal scales and like Glauconycteris has both overhair . and underhair and Mimetellus has dentate coronal scales and corresponds with Chalinolobus in the absence of distinct hair types.
Overhair and underhair are represented in the fur of Mystacina. The coarse dark overhairs are scattered amongst the close layer of short and wavy underhairs. They are long and straight and of irregular diameter, having a long, club-like thickening in their distal two-thirds and one or two shorter thickened portions near the base. No medulla occurs in either hair type.
Filament length of overhairs is 7-8 mm. Maximum diameter of the distal club is 30μ, that of the shorter proximal swellings 27μ. Terminal scales have a diameter of 6μ, the base of 11μ and the constrictions between the swellings 13.5μ. The tip is bluntly rounded. Melanin granules are scattered in irregular longitudinal bands through the length of the filament with the greatest concentration about midway along the distal club. This club has a narrow outer zone free of granules. Scale arrangement is annular. Scale length varies considerably along the filament from 7.5μ at the base to 17.5μ in the proximal swellings and thereafter decreases to about 6μ, toward the tip. Scale form is appressed to divergent, entire to repand coronal at the base and at the constrictions between the swellings, but is appressed, entire to repand coronal in the swellings and terminally.
In all the overhairs examined a long distal club-like portion was present, but some variation in the presence or absence and in the number of proximal swellings occurred. Usually one or two of these latter were present. In some individuals a number of the overhairs possessed a swollen bulb-like portion at some interval along the length of the distal club.
Filament length of underhairs is fairly uniform, about 6 mm. Diameter is greatest, about 12μ, at the middle. The filament tapers proximally to a diameter of about 7μ and distally to about 5μ. The tip of the hair is oblique. Melanin granules are scattered in irregular longitudinal bands through the length of the hair with the greatest concentration present at the middle of the filament. Scale arrangement is annular. Scale form is long (about 18μ), divergent, entire (oblique or rounded) coronal for the greatest part of the filament but becomes shorter (about 7μ), appressed distally.
The peculiar nature of the fur of Mystacina has caused comment by several previous authors. Gray (in Dieffenbach,
My findings differ from the description given by Benedict (
Benedict (Myotis, to which Miller (Mystacina, has a hastate scale form. The Molossidae which compare with Mystacina in the robust form of the hind limb are strikingly different in the dentate coronal scale form and the lack of differentiation of overhair and under-hair. The Emballonuridae, like the Molossidae, lack distinct hair types and have divaricate, dentate coronal scales. Mystacina compares with the Desmodontidae, Natalidae, Noctilionidae and Nycteridae both in the presence of distinct overhair and underhair and also in having a scale form varying only from entire to sinuate coronal. The greater abundance of melanin granules in the central core of the filament of
Desmodus corresponds to the presence of an outer granule-free zone in the distal club of Mystacina, but the presence of a basal bulb in the former genus clearly distinguishes the two. In the Natalidae the straight overhairs are slightly coarser than the wavy underhairs but the hair types correspond in length and the underhairs have some hastate scales. The Noctilionidae and Nycteridae correspond closely to the form described for Mystacina, particularly in the possession of an unusually long, club-like portion in the distal region of the overhair. In general the hairs of noctilionids are somewhat shorter, and those of nycterids somewhat longer than is the case in Mystacina.
The present area of distribution of New Zealand bats is considerably less than it was over a century ago. The decrease is correlated with the restriction of forest during the last century and the failure of either species to survive in open country or to urbanise. The Urewera in the North Island and the Buller River drainage in the South Island appear to support the highest densities of bats in New Zealand. Field observations in relation to behaviour and ecology are reviewed, and probable species differences in hibernation, foraging times, and diet are indicated. Fur mites are reported as parasites of both species.
During a recent investigation of the two New Zealand bats, Mystacina tuberculata Gray and Chalinolobus tuberculatus (Forster), an attempt was made to survey the distribution of these species, and to review the fragmentary knowledge of their ecology and behaviour. A brief account of some of the results of this survey has already been published (Dwyer, Mystacina was obtained.
Both species of New Zealand bat are endemic. The long-tailed bat, C. tuberculatus, is one of six species of the Australasian genus Chalinolobus (F. Vesper-tilionidae) ; a genus closely related to the southern African Glauconycteris. Tate (C. tuberculatus is a comparatively recent immigrant to New Zealand. The shorttailed bat, Mystacina tuberculata, is the sole species of the family Mystacinidae. Within New Zealand Mystacina is represented as two subspecies. M. t. tuberculata is recorded from the North Island and the northern parts of the South Island, while M. t. robusta is known only from the Stewart Island region.
There are some unconfirmed records suggesting the existence of other species of bat in New Zealand. Polack (Hipposideros cervinus (Gould) extends south to New Zealand, but I have been unable to find any evidence for this remark. Tate (Hipposideros and does not include New Zealand within the range. It is possible that records such as these could be based on chance sightings of occasional wind-borne Australian species.
The information upon which the following account of distribution is based was derived from various sources. Records have been supplied by the Auckland Institute and Museum, Dominion Museum, Canterbury Museum and Southland Museum. Phillipps' file at the Dominion Museum, Internal Affairs Department file No. 46/108 and Animal Ecology Division (D.S.I.R.) file No. 6/24/2 have been valuable references. The reports obtained from the New Zealand Deer Stalkers' Association were summarized in Roaring Stag. Several records have been supplied by the New Zealand Speleological Society.
Distribution records have also been obtained from the following: Buller (
Maps 1 and 2 record the localities from which sightings of bats have been reported, and Table I summarises the details of the distribution records for each of the seventeen districts shown on these plates. Details of the actual distribution records are included as an appendix to the thesis "Studies on New Zealand Ghiroptera", by P. D. Dwyer, deposited at Victoria University of Wellington, New Zealand. Since the completion of this thesis the following additional records have been made available by the New Zealand Deer Stalkers' Association. These records are incorporated in the totals of Tables I and II, but are not shown on the distribution maps.Mystacina, Chalinolobus, and unidentified bats respectively. Large flights are taken, quite arbitrarily, as those including twelve or more animals.
Two individuals of Mystacina tuberculata on Little Barrier Island during early Chalinolobus tuberculatus) sighted in a limestone tunnel (Wai-tomo district) during February,
Note that sightings along the Wanganui River on the boundary of Taranaki and North Wellington are considered in Table I as being in North Wellington.
Maps 3 and 4 illustrate the relationship of observations to forested areas and to rivers and lakes. In Map 3 only observations since
A number of problems in interpretation are at once apparent from an examination of the distribution records available. The most obvious of these is that records must tend to reflect the distribution of observers, and for animals such as bats which become active after dusk this is particularly relevant. Both species are forest inhabiting and the greater percentage of observations of bats in flight have been made over lakes, in river valleys, or in open clearings within or adjacent. to forest. It is only in such open areas that the rapid movement of the bats may be observed against the background of the sky. There is a tendency, therefore, for observations. to fringe the forests (post
Table II groups the total numbers of observations in five yearly intervals. The discrepancies in the totals between Tables I and II and between the tables and the maps result from differences in the details supplied by individual records.
The second problem arising in interpretation of distribution records is that the total number of observations for different districts may give a false impression of the relative densities of bats in these districts. Thus a comparison of the records for the South Auckland and Gisborne districts (Table I) indicates fewer observations, 22 against 26, for Gisborne, but more colonies or large flights, 6 against 3. In addition the percentage of observations since
Sight records are not sufficient to enable species identification. Therefore most of the observations are of unidentified bats. Species records have only been accepted in the light of good evidence. Mystacina has long been popularly legarded as the rarer species, but I feel this stems from its structural peculiarities and the popular synonymy of "unusual" and "rare". There is no great difference between the numbers of records since Mystacina, 20 for Chalinolobus.
With these problems in mind, I have attempted to assess the distribution of bats in New Zealand, in relation to time and physical characters of the environment.
The present area of distribution of New Zealand bats is considerably less than it was over a century ago. Early records are available from many major urban centres such as Wellington and Dunedin and a well authenticated colony beneath a bridge over the river Avon in Christchurch persisted to the early 1900's. Smaller townships such as Dannevirke (Hawke's Bay) and Ross (Westland) also had their colonies.
Before
There is no evidence that the density of bats has decreased within unmodified forest. Rivers and lakes provide an abundance of suitable insect food and clearings near bush provide favourable hunting grounds. Bat colonies have therefore survived in these localities. In open limestone country isolated colonies sometimes occur in caves. Such small colonies are present in the Te Kuiti-Waitomo district (South Auckland).
Altitudinally the bush line limits the distribution of bats. The single record. above this limit is at 3,460 feet in the Tararua Range (South Wellington).
In the North Island bats are regularly reported from southern South Auckland, Rotorua-Taupo, Gisborne and the northern parts of North Wellington and Hawke's Bay. Within. the Rotorua-Taupo and Gisborne districts the greatest number of sightings are from the Urewera and Lakes Waikaremoana-Rotorua districts.
In the South Island, north-western Marlborough, Nelson and northern West-land seem to be well populated. The Wairau, Pelorus, Motueka, Karamea, Buller and Grey River drainage systems account for these records. The few recent records for Canterbury come from the Geraldine County. In Southland the few records are perhaps deceptive and the recent nature of these, excepting those coastal, suggests that a general absence of observers in previous years accounts for an apparent absence of bats. Stewart Island and its subsidiary islets still support small but flourishing colonies.
I would suggest that the Urewera in the North Island and the Buller River drainage in the South Island support the highest densities of bats in New Zealand. For only two localities is it possible that one species may occur in the absence of the other. In the North Island only Chalinolobus has been recorded from the Wanganui River drainage, and in the Stewart Island region only Mystacina has been taken.
Little has been recorded of the behaviour or general ecology of the New-Zealand bats. Chance encounters with colonies, often as a result of felling timber, provide some information relating to habitat and gregarity. Stead (Mystacina near Stewart Island, and a few brief accounts, such as Roach and Turbott (
Fig. 1 records the activity of bats in terms of the number of observations for each month of the year. The basic pattern, reflected in the value for unidentified bats, is of low numbers during the winter months (May to August) followed by a steady increase in numbers from September to mid-summer and autumn. Though values are low, it is noticeable that records of Mystacina are lacking for only two months, March and September, whereas Chalinolobus is not recorded during five months, three of these being the winter months, May, July and August. Fig. 2 records the yearly activity of a small colony of unidentified bats at Puke-titiri, Hawke's Bay, observed by Miss P. Lewis during the years
Of the numerous observations of unidentified bats, 22 record dusk, 22 evening and 10 summer evenings as the sighting time. A few claim that the bats were seen after dark. Confirmed sightings of Chalinolobus state evening as being the period of activity. Roach and Turbott (
There are few references to Mystacina in flight. Two records of the entry of this species into lighted huts, and Stead's (Mystacina in the Rotorua-Taupo district consistently roosted till 8 or 8.30 p.m. in the early stages of its capture.
Fig. 3 shows the relationship between the times of appearance of the Puketitiri bats and the times of sunset for Chalinolobus. Phillipps' record for the 19th February coincides with the Puke-titiri times for the corresponding period of the year. A winter sighting of an unidentified bat in the Rotorua-Taupo district during June,
The duration of the foraging period is difficult to assess. Observations of bats usually last between a few seconds and fifteen or twenty minutes. Longer observations are seldom possible because of the rapidly failing evening light. Captive animals are usually active for only short periods each day. McKay (Internal Affairs file) records that for a captive Mystacina feeding, drinking, and exercise usually occupied three hours. Stead (Chalinolobus specimen. Phillipps' Chalinolobus remained away from the roost for six hours. At Puketitiri sightings have been made at 4.45 a.m. in January, 5 a.m. in March, and early dawn in June.
Bats are generally reported on fine warm nights. Miss Lewis (personal communication) states that no bats are seen at Puketitiri on misty damp evenings and suggests that on cloudy evenings the bats appear earlier than usual. A record of bats flying high in drizzle, and a record of an injured Chalinolobus found on the floor of a shed after a rough night, are the only reports of activity in adverse conditions.
At Puketitiri the bats appear regularly from the same section of forest. Feeding areas are established and are not more than about 800 yards from the point at which the bats appear. Three areas serve as the main feeding grounds. These
A few other evening observations record the following distances from the nearest forest: 150 yards and 350 yards, ¼, ½, and ¾ of a mile. Of fifty reports which include details of the habitat, 28 refer to the presence of water either as swamps, lakes, streams, etc., and 25 refer to the presence of forest. Of these latter, all but three indicate that the bats were observed at the forest margin or in open areas adjacent to forest. The remaining three are sightings in dense forest.
The flight of both species is erratic. Colenso (Chalinolobus as quick, soft, and noiseless. It is unaffected by full daylight. When resting between flights this bat hangs upside-down with the wings furled.
Frequent changes of direction are also characteristic of Mystacina but Stead (Chalinolobus. Mystacina may pursue insects in the vicinity of lights, flying at times quite close to the ground, and a number of specimens have been captured in forest when stunned after hitting against torches or lanterns, or after flying into lighted tents or huts. In captivity this species may rest between flights with the wings partially extended. When roosting it suspends itself upside down, using the recurved claws of both feet. The head extends straight downwards. The thigh is directed outwards from the body, at right angles to it, and the shank extends posteriorly and slightly medially. The foot is turned only slightly outwards from the line of the shank.
A number of structural peculiarities in Mystacina have been interpreted as adaptations suitable for terrestrialism (Dobson, Mystacina taken on Solomon Island. "They were placed in a box with a wire-netting front, and were most active in their efforts to escape, running head first with a curious stiff action and quite fast, using their folded wings as forelegs, the wrist joints coming in contact with the ground. They climbed the wire-netting or the smooth wooden sides tail first and with remarkable agility."
Mr H. E. Grubner (Whakatane) who kept a single bat of this species for two weeks during October,
A slight "clacking" emitted during flight has been noted for Mystacina. Stead (Mystacina when handled, and Grubner (personal communication) describes ear twitching as a reaction to light scratching or high pitched sounds. He describes the reaction of this bat to radio static as "panic", and describes a behaviour pattern before flight in which the animal "opened its mouth and turned its head from side to side". This latter would facilitate echo-orientation. Phillipps (Dominion Museum file) records a high pitched squeak and a low grunt from a colony of Chalinolobus.
The numerous evening observations of bats above water suggest that adult instars of insects with aquatic larvae may form a major portion of the diet. Bats hunting the forest margin in the evening probably obtain moths. Moths, mosquitoes, and midges are all recorded as food items by observers, and Colenso (
Cheeseman (Chalinolobus, and Roach and Turbott (Chalinolobus are small, about 4 to 8 mm long and, though usually tapered at either end, they may be somewhat irregular. Droppings from the surface of a cave pool at Orakei-Korako (Rotorua-Taupo district) consisted almost entirely of lepidopterous and dipterous remnants.
Items accepted as food by captive specimens of Mystacina include earwigs, spiders, moths, grasshoppers, crickets, wood grubs, and wire worms. Of these, spiders, moths and crickets were apparently most acceptable. McKay (Internal Affairs file) states that crickets "were quickly killed and, like all other insects that were offered, they were devoured head first with a distinct crunching noise". Spiders and crickets were readily taken from the floor of the cage.
Stead (Mystacina and carcasses of mutton birds hung out overnight to dry have been damaged by these bats. The strong transverse tongue ridges (Knox,
Parham (personal communication) describes drinking habits of Mystacina. "The animal was offered half a teaspoonful of water. It would take a drink, then wipe its mouth on the floor of the cage before taking the next. This process was repeated four or five times. During this period it was sitting with the toes of each foot splayed out, and the arms resting on the floor."
The number of bats flighting together varies considerably. Table III is prepared from 132 sightings. Nearly all the larger flights have been noted above water. Pairs of bats and solitary bats are frequently seen hunting along the forest margin.
Stead (Mystacina. The seven bats, including both males and females, were packed closely together. In
Stead's small colony was at the end of a hole in a rata limb. The hole, about I8in long, with the opening about five inches in diameter, was five feet from the ground. Other colonies are recorded from hollow ratas, totaras and puriri trees, as well as from caves. A musty smell is usually associated with long standing colony sites. The colony of thirty or more bats discovered at Matahina during
Many of the bats from this last colony flew directly to a neighbouring totara when they were disturbed. Stead (Mystacina in Milford Sound in
The body of Mystacina is cold to the touch during the roosting hours and if disturbed at this time movements are extremely sluggish. Waking involves stretching and head shaking. The animals are difficult to rouse when cold but become very active and quite warm after handling.
There are several records of large colonies of Chalinolobus in trees and caves. A colony numbering several hundreds is mentioned by Buller (
Colonies of unidentified bats have been found in caves and hollow trees and in the leafy upper portions of tall trees. Totara, rimu, kahikatea, pukatea and rata are all recorded as sites; totara being reported most frequently. Specific identification of trees was not available in any instance.
Moreporks have been frequently reported as bat predators. Stead (Mystacina in a morepork's nest. The bat had a small claw puncture in its neck. A pair of moreporks nested for a number of seasons at the entrance to a cave known to support a colony of Chalinolobus. Grubner (personal communication) has observed a bat escape an attack from a morepork by flight in a fast upward spiral.
Two records, one for each species, are known of capture by cats. Three Mystacina were killed by a dog when a tree containing a colony was felled. Stead (Mystacina. Mustelids could similarly cause extensive loss to accessible colonies, but there are no confirmed records of predation by any of these mammals.
Both species of New Zealand bat carry external parasites. Fleas are known from both, but till now records of mites exist only for Mystacina. Several observers refer to this bat as being "infested with vermin" and Grubner (personal communication) , who reports "a large spidery thing, reddish brown, and about ⅛in or more in size" from a captive bat, states that the animal was able to scratch any portion of its back with the foot.
Most specimens, juvenile and adult, of Chalinolobus examined during this study carried fleas (Order Aphaniptera). These occurred in the fur on all portions of the body, being commonest on the ventral surface in the axillary and pubic regions. From four animals (3 females, 1 male) which were thoroughly combed 23 fleas (17 females, 6 males) were obtained. All were Porribius pacificus described by Jordan (Mystacina examined. It was a female P. pacificus. Jordan includes five species in the genus Porribius, all as parasites of Australasian bats. Host species include Eptesicus pumilus, Nyctinomus (= Tadarida) australis, and Chalinolobus morio (Rothschild, P. pacificus is at present in the process of transferring to Mystacina.
Grubner's description of a parasite of Mystacina as "spidery" and "about ⅛th inch or more in length" is strikingly suggestive of the family Nycteribidae, Order Diptera. Allen (
Mites (Order Acarina) were common on Mystacina, as many as 15 being obtained from a single specimen. The three species recognised belonging to the family Laelaptidae (Mesostigmata), suborder Sarcoptiformes (perhaps family Acaridae) and the group Parasitoidea (Mesostigmata). The laelaptid mite species was the most common. Only a single specimen of the last was obtained. Mites were present in the fur and none were found on the membranes.
A single specimen of a large mite was obtained from a juvenile Chalinolobus from Pelorus Valley (Marlborough). A member of the suborder Trombidiformes, the extremely dense coat of setae on the dorsal surface of this mite suggest that it belongs to the family Trombiculidae. Three genera of this family are recorded (Domrow, Trombicula thomsoni known from Chalinolobus gouldii.
I would like to thank Professor L. R. Richardson for his guidance throughout this investigation and Professor A. F. O'Farrell and Dr J. le Gay Brereton for suggestions during the preparation of the manuscript.
Numerous persons have provided the information upon which this survey is based, and in particular acknowledgment is due to Mr W. J. Phillipps, who amassed most of the early records discussed here. Other important records have been provided by the Dominion Museum, Canterbury Museum, Internal Affairs Department, Animal Ecology Division, New Zealand Deer Stalkers' Association, New Zealand Speleological Society, Miss Pam Lewis, Mr D. Arthur and Mr W. Parham.
Not available during this study.