Order Chiroptera.

94 | Order Insectivora Stream Mainland Island overlapped by 70— 80% t2081. Competition from hedgehogs could limit kiwi numbers in the long term, and, because kiwi and hedgehogs nest in similar sites, hedgehogs may compete for nests, disturb incubating kiwi, or even damage kiwi chicks. Competition for nests and food between hedgehogs and kiwi, and other native insectivores, needs to be formally investigated. Control Hedgehogs have been added to the list of target species in many trapping programmes designed to protect indigenous fauna.'1650' However, the level of hedgehog control needed to gain any benefit has not been established. There are no data describing the relationships between population densities of hedgehogs and the extent of consequent damage to vulnerable fauna, and no estimates of how well, if at all, trapping regimes can reduce local hedgehog densities. Hedgehogs can tolerate high levels of brodifacoum.'-At Boundary Stream where brodifacoum was always available in bait stations, 37 radiotagged or marked hedgehogs were observed over 4 months but none died.'208' No formal research on toxins effective against hedgehogs has been done. Hedgehogs are also tolerant of a variety of other toxins including the venom of snakes, bees, wasps and scorpions; toxins of bacterial origin; and arsenic and strychnine, although they are susceptible to molluscicides.'2261' Literature resources and reviews Reeve;'226" Brockie.'303' C.J & M.D.S. ORDER CHIROPTERA Bats are the only mammals capable of sustained flight. They are easily distinguished from all other mammals by the extension of the finger bones to support the wing membranes (the name Chiroptera means 'hand-wing'). Bats have evolved many structural and physiological modifications, in addition to wings, to adapt them to their unique way of life. Some of their most unusual features are the specialised ears and elaborate facial skin growths of the many bat species that have evolved echolocation. The variation in form, size, and structure of bats is enormous; from the tiny bumble-bee bat (Craseonycteris thonglongyai), which weighs about 2 g, to the giant flying fox (Pteropus giganteus), which weighs over 1.2 kg and has a wingspan of 1.7 m. At present, 1001 species of bats are recognised in the world (and new, cryptic species are added as they are discovered), and many are classed as threatened. They form two quite different groups, the Suborders Megachiroptera (167 species) and Microchiroptera (834 species).'1316' The earliest known fossils, from the early Eocene (55 million years ago), are in some characters as advanced as many living species of Microchiroptera. Therefore the earliest bats must have arisen long before this, possibly in the mid to late Cretaceous (70-100 million years ago). Their immediate ancestors might have been small arboreal insectivores possessing gliding membranes and an early form of echolocation."248' The New Zealand bat fauna is very small and unlike that of any other country. Since there are over 90 species of bats in Australia,'821' it is surprising that not more of them have been blown to New Zealand in the past. At least three species have reached Lord Howe I., 500 km from Australia, and two have reached Norfolk I., 1300 km from Australia;'703' but, aside from Chalinolobus tuberculatus, there is only one record of a single vagrant flying fox reaching New Zealand, 2000 km from Australia.'693' Advances in field technology have made study of these cryptic nocturnal mammals both feasible and accessible, and enabled a huge increase in research on New Zealand bats during the 1990s.'2032' Small portable bat detectors, which record the ultrasonic sounds emitted by bats, are now used widely throughout the country to study distribution and habitat use.'2042; 2119' Radio transmitters small enough (0.4-0.7 g) to attach to <15 g bats are providing a wealth of information on home range, movements, use of roosts, and social structure for periods of up to a month at a time.'2033; 4991 Specialised harp traps for catching bats allow systematic capture programmes, including long-term banding studies of long-tailed bats.'2425' Infrared, time lapse video technology and low-cost, miniature video cameras and portable recorders allow monitoring of behaviour, population sizes, and activity of bats at roosts in remote locations.'2421' Maintenance of New Zealand bats in captivity is becoming routine.'24,7; 24l9; 1598' DOC has an active bat recovery programme,'18331 which aims to ensure the survival of all extant bat species and subspecies throughout their present ranges and, where feasible, to establish new 95 96 | Order Chiroptera populations within their historical range. The objectives of the current Recovery Plan are to (1) undertake or promote research on bats, which will assist in their management; (2) evaluate the status of both short-tailed and long-tailed bats; (3) establish populations of short-tailed bats on suitable islands; (4) select, protect, and monitor populations of short-tailed and long-tailed bats throughout their geographic range; and (5) raise public awareness of bats and involve the public in bat conservation. Since the publication of the Recovery Plan, researchers, conservation managers, and members of the public have been active in following up reports, identifying populations, recording distribution, and locating and monitoring roost sites. There is a growing interest in incorporating knowledge of bats into learning programmes."33'1389' The burgeoning of bat research in New Zealand also raises more questions and opens new avenues of research, and will provide a solid base against which to test more general theories of bat behaviour. Live bats are protected by the Wildlife Act 1953 and must not be handled or disturbed. Any dead bats found must be sent for study, either to one of the major museums or to the nearest office of DOC, together with information on date and locality. SUBORDER MICROCHIROPTERA The Microchiroptera comprise 834 taxa from 17 families and 137 genera, distributed almost worldwide. Most feed on insects, and the rest on a range of foods from fruit, nectar, and pollen, to blood, birds, small mammals (including other bats), amphibians, and fish. All navigate by echolocation, emitting ultrasonic sounds through the mouth or nose. The distinguishing characters of the three endemic species of New Zealand Microchirop teran bats are summarised in Table 22. All three are classed as threatened"833' 126" and one is probably extinct. Two belong to the Family Mystacinidae (short-tailed bats, genus Mystacina), which have evolved in isolation for so long that they possess characters not found in bats anywhere else in the world. The ancestors of the third species, the longtailed bat Chalinolobus tuberculatus (Family Vespertilionidae) are believed to have reached New Zealand from Australia by chance dispersal across the Tasman during the Pleistocene (p. 113), as have those of many of our birds.[2907] Figure 12 Echdccation coll sl^res lor New Zealand batsPlots of audio frequency against time, with amplitude represented by the intensity of shading, o) Twenlyfive millisecond sequence, showing individual pulses, b) fair second sequences showing pulse trains 100 kHz 0 kHz 100 kHz 0 kHz Mystacina Chalinolobus 10 15 Millisecs b) 100 kHz Mystacina i I • 11 i 111. 11 i i i II | h i n | HE li,,; ii iii in in n11n1111 1 '•' • i i1 iii i i-' 111 111 • i - - - ^ •-1 ,— , .. .. ... 100 kHz Chalinolobus 20 —* — , - 100 200 300 400 500 Millisecs Suborder Microchiroptera | 97 -S £ '1 s -Q p o _c to u_ o ,c J> o c 5 a> 2. 6 5 I J 3 I II If 5 a J I 3 1 J I .§ = J 3 <5 o CD | 3 XI X < "O £= o < Z 0) <= p [O X CD O E CJ _c CD £ _D 3 X o 3 p o "o CD E o X own 3 0 c X CD c o X "o E E 0 CD ci 3 CD P K 2r- <D 0 > o cJ > CD 0 b c CL u .5L X c: O O o Qo >. oo JD > CD c p -Q 0 >- X o X E CD £ > O X c ~p V o o a> X o CD E JD 0 X o CD 5 u to c o O X X CD ID c Q_ X _o O _c c to O O > CD to > o CD O E ro Z < _b < j= 3 o o to C= X X C o < O X CD X CD cz . 0 CD cz 3 c 0 X S_CD c 3 o o CJ c o CJ u X c CD r— o X to O O _0 3 O £ 3 5 x c= o X _o CL £ 5 I 9 E o - I 3 8.-5 -§ | Q—sz 9 E "o <D 5 o u o Jl> 3 X _o Jx ==" at >~ _g 8 x m >- X X 0 o CJ 0 tr c 0 p >- X ~3 E 0 o E cz p O) _0 o E . 0 "to 3 0 X c o '— 0~ pu _o x CD 0 <D •s-: = o O 0 _Q £ a) o E M -o -8 | 0 O E oo CO 5 .E O X X E "X c o I S CI 0 0 N N I 3 E CJ o 0 o c O X ~6 0 X o CL I 0 CT) X _g x c: o >- Q) X !i Ji ^2 0 T2 - o x — 0 c 1 O ? r-i > 5 to n 5 U LLI ~o o o X x c o o _ >o X c o E £ O I o o o X 16 mm "o E oo Smal N f\ oo 98 | Order Chiroptera Four exotic species of bats from two Microchiropteran families have arrived dead in New Zealand. Three were vespertilionids: a Japanese pipistrelle (Pipistrellus javanicus abramus) arrived in a cargo of car parts,17041 an Australian lesser long-eared bat (Nyctophilus geoffroyi) in a cargo of timber,1 031 and an Australian little forest bat (Vespadelus vulturnus) in a crate of aircraft parts.' ~51 A small unidentified bat belonging to the Molossidae arrived among a shipment of bananas from Ecuador in 2002.120371 FAMILY VESPERTILIONIDAE The Vespertilionidae (evening bats) is the largest of the Microchiroptera families, and includes about 357 species in 37 genera.113161 Vespertilionid bats are among the most widely dispersed of mammals, found almost worldwide (except for the polar regions), including remote oceanic islands. Genus Chalinolobus Five of the seven species of Australasian wattled or lobe-lipped bats live in Australia (including Tasmania and Norfolk I.) and New Guinea; one newly recognised species in New Caledonia; and one, Chalinolobus tuberculatus, in New Zealand.'825' 1316 They are small to medium-sized bats which have a fleshy lobe on the lower ear margin, and often another at the corner of the mouth. The ears are short and broad, the tragus curves inwards, the forehead is high, and the muzzle has a glandular appearance.1"581 New Zealand long-tailed bat Chalinolobus tuberculatus (Forster, 1844) Synonyms Vespertilio tuberculatus Gray, 1843; Vespertilio tuberculatus Forster, 1844; Scotophilus tuberculatus Tomes, 1857; Chalinolobus tuberculatus Peters, 1866; Chalinolobus morio Thomas, 1889. Also called New Zealand short-eared bat or New Zealand wattled bat (English); pekapeka (Maori). The name Chalinolobus tuberculatus should be attributed to Forster (1844) rather than Gray (1843) as previously.16971 Vespertilio (Chalinolobus) tuberculatus was described by Forster in his journal, and illustrated by his son G. Forster, during Cook's second voyage to New Zealand in 1773.12522:13421 There are significant differences in size (Table 23), calls,121201 and genetic diversity129141 among populations in North and South Is, but it is not yet clear whether these warrant subspecific status. Description Distinguishing marks, Table 22 and Plate 3. A small, delicate bat, in contrast to the more robust Mystacina. The head is broad and the forehead is high. The face is moderately hair)', with the nostrils set on prominences. The eyes are small with well-formed fleshy lids. The small ears and short tragus are rounded distally, and the outer margin of the ear continues along the face, beneath the eye, as an antitragus, which terminates just behind the liplobule. The more pronounced tragus extends from within the ear above the antitragus. It is narrow at the base, but widens and is rounded distally.120321 Reproductive females are larger than males (Tables 23, 24). Pelage colour is variable in living wild bats, and changes with age. Adult females usually have rich chestnut upperparts, sometimes with white tips to the fur. Males, and 1-3 year olds of both sexes, are darker, with dark brown upperparts and blackish fur around the head.'20321 Underparts are pale brown in both sexes, paler about the pubic region. The fine, soft dorsal fur is up to 7 mm long, with no differentiation into overhair and underhair. The fur of preserved museum specimens is often bleached to a light reddish-brown. The limbs, wing, and tail membranes are almost naked and blackishbrown in colour. New Zealand long-tailed bat | 99 ° ~E ai ,-s .2* 05 TJ 5 IO 8. 1? £ i •£ o> c _a> E c £ "E o E u_ — CO K CM — — — CO O CO CM CO — o O o o o o o o O o o -H -H -H -H -H -H -H -H -H -ft -H -H o CM O o CM CO o -q ocq cq a" d o o o ,—" CO ,— CM CM CO ot of CO CO CO CO CO CO CO CO co co o o Mf CM K K — — -H -H -H •H Mf K K co <i CM cb K K CO K K CM CM CM CM CO CO CM CO O Mf •o CM o o o o o o o o o o o o o o o o -H +i -H -H -H -H -H -H -H -H M -H -H -H CO Of CO CO CO o CO CO CO K o o CO d o-' d d d o d o d d CO d d o CO co CO CO CO Mf CO Mf co co CO CO CO "Cf CM co K CO K — •— — — , — o CM o o o o o o o o o +1 o -H O -H o -H -H -H -H +i -H -H -H o -H co -H CM -H CO CO O CO CO O CO •o K o K o co o d d CO d d d d — CO CO d O O o o o o v v o o o o o o v v o v K o K O — P ca K O O co -OK CO "cf — cr> •— o a O Mf CM CO '— K — ^r I J£ 0) 0£ 0) ~o 5 o co K CM MjK CO •st O i a U o (J ~o c g O d 13 _Q c O U 100 | Order Chiroptera J £ C O ~a> -8 -Q O .C ~6 <5 o ~D O a CD U o E J. "5> •£ io a •£ Q. B •o cn c I E J. •£ o> c _S> E "8 o o o o — — — U) +i -H -H -H o o o o ft CO wo •o -H -H -H o 1 CN o o o CO — o o 1 1 • o' a CO V _o V 5 m O D) < o -H W) K K O +1 W) o +t wa O -H nt CN o -H CO CN — — V) W") i/") W") CO NT CN IS +1 K K K CN CN -H CN CO •o CN -o •o a O — — o o o o o -H -H -H -H -H W-) wo o — CO d o o-' o o' NT co NT CO wo NT NT CN oco CN CO CN -H +t -H -H -H wo CO -O WO K O co CN WO o wo wo wo NT wo "O NT CN CN CN io •— CO .— ~s K CN "8 o z K CO O o o Nf "a < o o v T5 o The bones of the legs and forelimbs (except the thumb), are long and slender. The small thumb projects free from the wrist and carries a long curved claw. The small hind foot is turned outwards. The calcar extends from the heel as a strong process and supports almost half of the posterior border of the large interfemoral membrane. A small, rounded post-calcareal lobe is present near the base of the foot. The relatively long, pendant penis of the male makes for easy distinction of the sexes. Females without visible nipples, or with nipples covered in fur, are non-reproductive J20431 Nipples remain conspicuous after females have given birth once. Nonvolant young are recognisable by their small size, patches of bare skin, and short, grey-black fur. Young-of-the-year are recognisable until phalangeal epiphyses are fully fused at c. 3 months old. Search phase echolocation calls of long-tailed bats (Figure 12) begin with a steep, downward frequency-modulated sweep, followed by a short, less modulated component at lower frequencies. Up to three harmonics are present, although some calls have none. Peak amplitude is in the shallow tail of the fundamental call at about 36-40 kHz, depending on population. Search phase calls averaging 6.3 ms long sweep through c. 30 kHz from 65-34 kHz.'2l20; 2,23; 2I221 Dental formula I2/3 C1/, Pm2/2 M3/, = 34. The complete mitochondrial genome of the long-tailed bat has been mapped (GenBank accession number AF321051).'1566' Field sign Long-tailed bats leave little sign of their presence because they are nocturnal and rare. They are visible flying in twilight for about 45 minutes after sunset. Most roosting cavities are high in trees.1242 1 More accessible roost sites in caves may be marked by a pile of guano under a crack or fissure occupied by sleeping bats, or by audible squeaking if bats are active.[702: 20361 Individual New Zealand long-tailed bat | 101 droppings are small and mouse-like (usually 2—4 mm long and 1 mm wide). Flying long-tailed bats can be detected with hand-held heterodyne Batbox III bat detectors set at c. 40 kHz. There is some overlap between the fundamentals and harmonics of the calls of Mystacina tuberculata and C. tuberculatusp-x201 but they can be distinguished because the peak amplitude of M. tuberculata calls is c. 27-28 kHz, and the call rate of M. tuberculata is twice as fast as C. tuberculatus.[2120; 20421 Measurements Adult head and body length 42—63 mm; forearm 36.7-46.0 mm; wingspan 240—298 mm; wing depth at fifth digit 48-62 mm; tail 30-46 mm; ear 7.2—11.0 mm; condylobasal length 13.6-14.3 mm; adult mass (pre-feeding and nonbreeding) 7.1—12.5 g; juvenile mass (nonvolant) 3—6 g.I825; 826; 1089; 2032] Long-tailed bats have medium wing loading and aspect ratios, typical of bats that have moderate to fast flight speeds, relatively low energetic costs of flight, and that forage along forest edges and in gaps (Table 25). Their manoeuvrability within dense vegetation is limited. Significant sexual dimorphism and morphometric differences among populations may have consequences for wing characteristics, flight behaviour, and manoeuvrability. Juveniles develop isometrically once volant, although relationships among forearm, body length, and wingspan growth are highly variable.[1089] Variation Within populations, individual body mass varies with season11089; 2035' and among bats of different age and sex classes (Table 24). Males accumulate body fat reserves in autumn (late March). Weights of reproductive females vary according to the stage of the breeding season; they increase in mass during pregnancy (to 16.0 g, Puketitiri; 17.4 g, Eglinton Valley). In the Eglinton Valley, lactating females weigh 10.0-12.5 g (pre-feeding weights), non-reproductive females 8.7—10.0 g. Bats of either sex can weigh up to 3 g more after successful foraging, but return to basal weights by dawn. Comparisons among populations (Table 23) show that long-tailed bats from Eglinton Valley (45°S) were significantly larger in key morphological characters than those from three other sites (Ruakuri, 37°S; Grand Canyon Cave, 38°S; South Table 25 Wing morphology of adult male (M) and female (F) long-tailed bats from the Eglinton Valley, Fiordland. Analysis excludes visibly pregnant females, and females caught in November-December (parturition period). Character Sex Mean ± 1 SD Range n P Wing Area (m2) M 0.011 0.0006 0.010-0.013 19 F 0.012 0.0006 0.011-0.013 58 <0.0001 Tip Shape Index M 1.55 0.34 1.02-2.05 19 F 1.29 0.35 0.71-2.94 58 0.015 Wing Loading (N rrf2) M 8.42 0.45 7.60-9.07 19 F 8.91 0.62 7.60-10.81 58 0.003 Aspect Ratio M 6.63 0.26 6.09-7.33 19 F 6.52 0.25 5.96-7.19 58 0.12 Wing area, wing loading, tip-shape index, and aspect ratio calculations follow Norberg and Rayner.'1954P = differences from 2-sample Wests.'2026! significance of inter-sexual 102 | Order Chiroptero Canterbury, 44°S). Ruakuri and Grand Canyon bats were indistinguishable. South Canterbury bats have the smallest forearms but intermediate mass and wing depths. Diversity of mitochondrial DNA varies significantly among five populations,'2914' to the extent that divergence between North and South I. longtailed bats is comparable with that between the accepted subspecies of short-tailed bats. Variation in echolocation calls between North and South I. populations is also significant,'2120' and associated with significant size differences (Table 23). Because body size and echolocation call design in bats are interrelated within the same adaptive complex,17' variability may demonstrate adaptation to different habitats. History of colonisation The long-tailed bat is probably derived from an ancestral form of Chalinolobus windblown across the Tasman Sea some time during the Pleistocene. It has evolved in isolation in New Zealand for at least a million years. Figure 13 Distribution of reports of long-toiled bats in 10(300 yard grid squares on the New Zealand national grid. Records pre-1930 and 1930-1960 are from DwyerJ826! Records 1980-95 are from the National Bat Database administered by DOC. New Zealand long-tailed bat | 103 Distribution The long-tailed bat is found only in New Zealand, where it is widely distributed from the north of the North I. (35°S), through the western South I., to Halfmoon Bay on Stewart I. (47°S). It is also present on Great Barrier I., Little Barrier I., and Kapiti I. (Figure 13).[825; 826: 690; 7015 703: 2028] The surprisingly few known subfossil or recent skeletons (one, from Metro Nile River cave, completely enclosed in calcite inside a stalagmite) are all from localities within the present range (Grand Canyon Cave, Waipuna Cave, Gardners Gut Cave, and Porthole Cave, total n = 9).'577; 702;697' Historical anecdotes and monitoring since 1990 indicate that long-tailed bats are now rare or absent at many sites they formerly occupied, e.g. Banks Peninsula.'2028' There are few records now from the eastern South I. A review of reports lodged in the National Bat Database (administered by DOC) revealed many errors, now corrected.'2028' Habitat Long-tailed bats are associated with indigenous forest of various types,'2028' from sea level to the treeline (c. 1000 m asl), including northern coastal pohutukawa,"3' kauri-dominant remnants,'8' mixed beech,'242 ' podocarp1'089' and podocarphardwood forests,'203 ' and indigenous shrublands dominated by regenerating manuka/kanuka.',3,6: 2030; 8] preqUenCy Qf records of bats is inversely correlated with distance from forest.'2509' Long-tailed bats survived in some cities up to the 1920s and 1930s, but there are no records since then.'l53:2028' In rural landscapes, they forage mainly in remnants of indigenous forests and riparian vegetation.'1138: 268; 2030' In Kinleith Forest, central North I., they foraged within exotic plantations of various species including pine and eucalypts, most actively along roads through mature P. radiata forest.'1840' Straight-line canopy gaps created by roads were also favoured for hunting in kauri forest.'8' In Nothofagus rainforest in the Eglinton Valley, Fiordland, foraging was concentrated along linear landscape features such as forest edges (43% of bat passes) and roads (46%) in all seasons. Relatively little use was made of forest interior (3% of passes) in any season,'21'3" but above-canopy activity was observed both in the Eglinton'2026' and in Peel Forest."138' Long-tailed bats roost primarily in cavities in trunks and large limbs of trees within indigenous forest,'1089: 2426; 8; 2045' and less often in caves, buildings, under bridges, and on cliffs,' 01-703' presumably because in cold climates there are significant thermal benefits to roosting in a small enclosed space,'20/°' even though caves are common in parts of New Zealand. For example, of >300 recent excavations of caves, none has recorded long-tailed bat bones except in fossilised owl pellets, whereas the bones of other small animals of similar size were common;'29""1 of 221 large caves (>250 m long) listed in King Country, long-tailed bats were reported from only six.'2044' Long-tailed bats have used Grand Canyon Cave for at least the last 40 years, but not in large numbers except in spring and early summer when ambient temperatures within the cave were an optimum 10-13°C.'2036' The 371 bat roosts found in the Eglinton Valley in summer were not simply a random subset of all available cavities;'2427,2426' all were located in knot hole cavities, were relatively high from the ground, had smallto medium-sized entrances, and were internally dry and well insulated (warmer at night). These characteristics provided a specialised, more stable microclimate suitable for breeding,'2420' conservation of energy in cold climates, and likely increased breeding success. Most roost trees were large (>80 cm dbh), old (100—>600 y), standing in mature, open-structured lowland forest on the relatively flat valley floor within 500 m of the forest edge. Similar characters define roost trees in forests elsewhere in New Zealand.'108* 8; 20321 104 | Order Chiroptera In South Canterbury, 97% of the forest cover has been removed,120301 so 23% of breeding roosts were in small cracks in limestone bluffs. The oldest cavity-bearing trees in this agricultural landscape were strongly preferred, but offered poor quality roosts because most were < 80 cm dbh and poorly insulated. Exotic trees rarely reach sufficient age in New Zealand to provide high quality cavities.124281 Food The long-tailed bat is a moderately fast-flying aerial insectivore. Most identified fragments in faeces sampled from Grand Canyon Cave were Diptera (29%), Coleoptera (25%), and Lepidoptera (17.5%).110891 Various other terrestrial and aquatic flying insects were found less frequently, including Mycetophilidae, Culicidae, Psychodidae, Calliphoridae, Hemiptera, Trichoptera, and Ephemeroptera, mostly small (<10 mm). In the Eglinton Valley, food remains dropped at nightroosts (Table 26) included a variety of large (10^0 mm) invertebrates; insects >20 mm were common, particularly Tipulidae (20-25 mm). Large body parts are often discarded, so would not be easily detected using faecal analysis.120321 Social organisation and behaviour Activity. Emergence from roosts varies geographically, averaging 17 min after sunset in Hawke's Bay,1108''] 24 mjn after sunset at Qran(l Canyon Cave,1203 ], 30 min after sunset in South Canterbury,1"381 and only 2 min after (occasionally up to 54 min before) sunset in the Eglinton Valley.120321 Solitary bats, and all bats in winter, emerge significantly later.120321 Dusk temperature, invertebrate availability, cloud cover, wind strength, and illumination from the moon influence emergence times on any given night, and the amount of activity through the night. Minimum temperature determined whether or not bats flew at night, whereas invertebrate activity determined how long bats remained active.120311 Long-tailed bats may remain active all night throughout New Zealand, but with local and seaTable 26 Large invertebrates consumed by Chalinolobus tuberculatus that were identified from remains dropped in roosting cavities (n = 71 cavities examined)120371 ORDER, Family Common Name PLECOPTERA, Eustheniidae MEGALOPTERA, Corydalidae COLEOPTERA Lucanidae Scarabidae Tenebrionidae Elateridae Oedemeridae DIPTERA, Tipulidae LEPIDOPTERA Psychidae Noctuidae Stoneflies Dobsonflies Stag beetles Chafer beetles Darkling beetles Click beetles Lax beetles Craneflies Bag moth Owlet moth Species Approx. Body Length (mm) Stenoperla prasina 32 Archichauliodes diversus 35-40 Dorcus sp. 15 Odontrio sp. 12-14 Unidentified sp. 15 Unidentified sp. 20 Thelyphassa sp. 20 Unidentified sp. 20-25 Unidentified sp. 25 Unidentified sp. 25 Number of individuals 1 2 23 New Zealand long-tailed bat | 105 sonal variations in timing. During spring and summer in the Eglinton Valley, much activity was recorded in the first two hours after sunset, but in winter very little during the first three hours after sunset or in the five hours before dawn.120311 Similar patterns were recorded in Kinleith pine plantations,118401 and kauri forest,181 but in South Canterbury and at Puketitiri, activity had a bimodal pattern with dusk and dawn peaks in activity.1108* 11381 Radio-tagged long-tailed bats made on average four foraging flights interspersed with three roosting periods per night in the Eglinton Valley.120341 Bats were active for an average of 71% of the night (354 minutes) and stationary for 29% of the night (146 minutes) regardless of gender, reproductive condition, and weather. Reproductive females balance their high energy requirements throughout their reproductive cycle by increasing foraging efficiency, using torpor, and selecting thermally beneficial roosts.12841:24211 Hence there were few differences in activity patterns between reproductive classes, and energy demands limit all classes in a cold temperate climate when food is in short supply.120341 Roosts. The night-roosts used between foraging flights are usually different from day-roosts.I2036: 20341 At Grand Canyon Cave, up to c. 360 bats may enter the cave at night. Bats come and go from the night-roost throughout the night, although most arrive between 2300 and 0200 hours and depart for their day-roosts in trees before dawn. The ecological function of night-roosting in long-tailed bats is unknown, but in other bats is thought to be important for physiological (digestion, resting, grooming, shelter, energy conservation) and behavioural (social interaction, information transfer, mating) reasons.1149*233* 26031 Hibernation. Activity is reduced significantly in winter, but usually does not cease completely; feeding has been recorded in winter and in temperatures of -1.5°C.[1138; 20311 Wild long-tailed bats commonly use daily torpor during the summer to save energy.12838' 28411 Adult males and females roosting alone entered torpor on 80% of days, for virtually the whole day (c. 12 hours), but only on 35% of days, and not all day (c. 9 hours), when roosting in groups. Duration of torpor was inversely correlated with ambient temperature. The body temperatures of captive torpid adults fell to within a few degrees of external ambient; energy expenditure at 10°C was c. 2% of that when euthermic. Periods of apnoea (a transient cessation of respiration) were observed within minutes of bats entering torpor.128381 Social structure. Long-tailed bats form complex social groups. Radio-tagged bats in the Eglinton Valley in 1993-96 occupied colonial roosts on 63% of nights but also often (on 37% of nights) roosted alone.120431 All sex and age classes switched frequently between solitary and colonial roosts. During the breeding season colonial roosts held on average 34.7 ± 23.4 (SD) (max = 123) bats. Adults in colonial roosts comprised 62.8% reproductive females, 22.1% non-reproductive females, and 15.1% adult males. Males roosted more often alone.120431 At Puketitiri, bats were more often solitary (70% of nights) than colonial.110891 Long-term non-random associations among individuals defined three distinct social groups in the Eglinton Valley, comprising on average 72, 99, and 132 individuals/year.12029' 20451 The collective foraging ranges of the three groups overlapped, but they roosted in three geographically distinct adjacent areas. Bats switched occasionally between groups, potentially linking the local population assemblages into demes. Non-reproductive females and males switched between groups more often than reproductive females, but individuals switched only once or twice during the study and then just for one night (total 1.6% of 3102 captures at colonial roosts). Juveniles of both sexes 106 | Order Ch iroptera were associated with their natal group as 1-yearolds, and then later when breeding. Social groups were cryptic because foraging ranges of the groups overlapped, bats belonging to each group were spread over many roosts each day, and these roost sites changed from day to day.'20291 In South Canterbury in 1998-2002, two distinct social groups averaging 55 and 29 bats lived in the vicinity of Hanging Rock. Their foraging ranges overlapped, and their distinct group roosting areas occupied adjacent sections of the Opihi River. Only three individuals have so far been detected roosting outside their natal groups.120371 Roosting behaviour. Long-tailed bats move frequently between roost-sites, in highly structured groups of small average size.120291 Residency times of colonial summer roosts averaged 2.0 ± 0.4 SE days (range 1-4) in the Waitakere Ranges, 1.2 ± 0.1 days (1-2) in the Western King Country, 1.7 ± 0.3 days (1-5) in Puketitiri, 1.6 ± 0.2 days (1-8) in South Canterbury, and 1.7 ± 0.2 (1-5) days in the Eglinton Valley. Occupancy was longer (3.6 days) in winter J1138' an(J longer in males, juveniles, and solitary bats than in reproductive females (up to 16 days/roost, compared with 1-2 days).'2043' Bats shifted to new roosting sites virtually every day carrying non-volant young, moving simultaneously as a group.'2425; 20431 They seldom reused a given roost in the same season, but usually returned to it at about the same time each year.'20 0' Suitable roost trees were abundant in the Eglinton Valley, so long-tailed bats could move frequently in response to social and thermoregulatory requirements.'20'131 Avoidance of predators or parasites are possible reasons for shifting roosts in other bats,'1560' but neither explains why longtailed bats would move every day.'2045' Dispersion. In the Eglinton Valley, 50 radio-tagged bats ranged collectively over 11 700 ha, and their home range sizes were among the largest published for Microchiroptera.120331 Median range sizes for adult males were 1589 ha (max = 5629 ha), 1361 ha for post-lactating females, and 657 ha for nonreproductive females. Ranges of lactating females (median = 330 ha) were significantly smaller. Juveniles that had been volant for <2 weeks had the smallest ranges (median = 237 ha), but their movements increased significantly after c. 2 weeks to a median of2006 ha. Movements were frequent and rapid within the range (mean = 790 m/15 min) and average range lengths were 3.3-10.9 km (max =19 km). Maximum flight speeds of 60 km/h were recorded. Despite their large home ranges, Eglinton Valley bats concentrated their activity (85% of fixes) in small core areas that represented a mean of 5.7% of their ranges. Roosting sites for each individual were spread over relatively small areas (20-129 ha) averaging 9.4% of the area of their foraging ranges,'2033' and bats were faithful to the same areas of forest each year.'2045' Individual bats followed similar movement patterns each night, and they always returned to their day-roosting area rather than remaining in their foraging areas. Home ranges and their core areas did not overlap extensively, suggesting a degree of spatial segregation among foraging bats. In South Canterbury, home ranges were much smaller and overlapped considerably, ranging from 322-642 ha with maximum range widths of 2.8-4.4 km.'1138' Measured range sizes were probably underestimates of seasonal, annual, and life-time range requirements, because the transmitters can remain attached for <1 month.'2033' Reproduction and development Breeding has been studied only in the Eglinton Valley.'2035' Males were classed as sexually active when their epididymides were grey and distended, and as inactive when they were black and regressed.'2"43 Males had distended Cauda epididymidis at 1 year old (mean =1.6 years). Sperm New Zealand long-tailed bat | 107 can be found in the epididymides in late summer—early autumn in South Canterbury, the Eglinton Valley, and the King Country.'2031 Copulation has not been observed. As in Chalinolobus morio and C. gouldii, females probably store sperm over winter, but the mode of storage, dates of ovulation and fertilisation in spring, and length of gestation, are all unknown. Females produce a single young once a year, presumably in the colonial nursery roosts dominated by reproductive females, whereas some other Chalinolobus spp. breed only every second year or produce twins. In the Eglinton Valley, all nonreproductive bats were nulliparous and usually 1-2 years old, implying that all adult parous females reproduce annually after 2—3 yrs of age. Visibly pregnant females may be found from early to late November. Parturition dates range from mid November to mid December depending on the population.'10891 2030; 20351 In Hawke's Bay (39°S), median parturition date was about the last week of November, and young made their first flights 5 weeks later in early January (extrapolated from Gillingham'1089'); in South Canterbury, despite its southern latitude (44°S), births began in the first week of November, c. one month earlier than other populations, and first flights of young were recorded in the second week of December. In the Eglinton Valley (45°S), 70% of births fell during a 10-day period in mid December, and most of the rest throughout December; two bats visibly pregnant in the first week of January were first-time breeders. Sex ratio at birth is equal.'203"'' Palpation of abdomens of 52 near-term females indicated one embryo. Counts using infrared video, and captures of banded bats at roosts, confirmed a 1:1 ratio between parous females and young. Lactation coincided with highest annual air temperatures and peak abundance of flying invertebrates. Reproductive females can carry non-volant young until the young weigh up to 80% of their own body mass.'2023' Most lactating females return to roosts to feed and collect their young 1-3 hours after first emergence.'2043' Development is rapid, and young begin flying at 5-6 weeks old, when their forearm length reaches 35-38 mm, and mass 6.5—7.5 g. 203"'1 The average number of different newly volant juveniles captured was similar to the number of reproductive females (0.91 juveniles/female). Nipples of reproductive females were regressing two weeks later, but it is not known whether young continue to suckle once they begin flying. Date of first birth in the Eglinton Valley varied by 4 days annually, but date of first flight varied by 17 days. The short (<3 months), highly synchronous breeding season, birth of single young, early mating, and late age of sexual maturity compared with similar-sized vespertilionids may be related to low food availability and unpredictable cold-temperate weather conditions.'2035' Population dynamics Populations appear to be structured as a series of local demes,120291 of unknown size. Banding studies of some of the larger ones suggest a minimum of 800 bats at Grand Canyon Cave'2036' and 150-200 at Hanging Rock in South Canterbury.'1557' In the Eglinton Valley, three social groups averaged 303 banded individuals alive/year (pooled total), but there were other groups in the valley that were not banded.'2029' Average counts at colonial roosts are poor estimators of total population size, because groups moved to new sites virtually every day, and there is a daily turnover of individuals using each site, representing different combinations of subsets of each group. Thus, average counts of 86 bats emerging from colonial roosts in Hawke's Bay (range = 5-208)'1089' and 14 bats in the Waitakere Ranges (range = 2—24)'8' are certain to be underestimates of total local population size. Survival. Of 107 reproductive females banded as adults in 1993 in the Eglinton Valley, a minimum 108 | Order Chiroptera of 58 (54%) were still alive after 6 years and 34 (32%) after 7 years. Given that most females begin reproducing at 2 years old, some bats still alive are a minimum of 9 years old. The oldest known-age banded bat in the Eglinton Valley was 11 years old in January 2004.120371 Annual productivity did not differ between the three social groups in the Eglinton Valley, and the probability of survival of juveniles in their first year was generally high, varying from 0.26—0.88 (mean = 0.53), inversely proportional to size of natal group.120351 Variability in breeding parameters was unrelated to climate, but the social group with highest survival rates lived in the best habitat and contained reproductive females and juveniles with significantly higher body condition than other groups. In 1996-97 a sudden collapse in numbers, and poor productivity and survival, coincided with an irruption in the numbers of stoats, and a second one in 2000-01 coincided with an irruption in numbers of ship rats.'2035' 22021 In South Canterbury, productivity was low (0.22-0.24 young per parous female reaching volancy: n = 163 bats observed over 3 summers). Probability of young surviving their first year was only 0.23.1204,1 These low figures probably reflect the poor insulation of their roosts, and predation by introduced possums and feral cats. Predators, competitors, parasites, and diseases Owls (Ninox novaeseelandiae and the extinct Sceloglaux albifacies) are the natural predators of long-tailed bats,'826'29451 and even introduced owls (Athene noctua) have been observed attempting (unsuccessfully) to catch long-tailed bats near Geraldine."1381 Ten of 36 freshly dead long-tailed bats (28%) had been killed and partly eaten by feral or domestic cats.1 031 One particular domestic cat near Geraldine killed three long-tailed bats (two adult females and one juvenile) under an outside house light, where the bats were apparently feeding on moths attracted to the light.17011 In South Canterbury, possums were observed attempting to reach into the cavities containing young bats on 50% of nights when roosts were monitored using video cameras.'2030 Significant mortality of bats followed irruptions of ship rats and stoats in the Eglinton Valley.'22021 In the limestone areas in South Canterbury, starlings (Sturnus vulgaris), house sparrows (Passer domesticus), feral pigeons (Columba livid), and introduced wasps (Vespula spp.) all occupied cavities that appeared to be suitable as bat roosts but were no longer available to bats.'11381 In the Eglinton Valley, starlings and ship rats made nests in roost cavities,'24261 although it is not known if these species actually displaced bats from the roosts or found them empty. Disturbance and competition may displace or kill individual bats, but the consequences of these losses on the longterm viability of entire populations are unknown. The flea Ponnibiuspacificus}xi901 is common on and apparently specific to the long-tailed bat.'8261 Several species of mites have been observed, including Ornithonyssus spinosa,1,6331 a large mite probably belonging to the family Trombiculidae,18261 and an as yet undescribed new species of spinturnicid mite (Spinturnix sp.) found on a long-tailed bat from Fiordland.1'2131 The only endoparasite known is the cestode Hymenolepis chalinolobi.',o1 Significance to the New Zealand environment The North I. form of the long-tailed bat is classed by DOC as 'Nationally Vulnerable', while the rarer South I. form is 'Nationally Endangered'.'12611 Both are fully protected by the Wildlife Act 1953, but threatened with extinction in the medium term if nothing is done to reverse their decline.'20281 Causes of decline include loss of foraging and roosting habitats through clearance and logging of lowland forests, predation by introduced animals, competition for roost sites by introduced mammals, Family Mystacinidae | 109 birds and wasps, and human interference and disturbance at roost sites.[2028; 2030; 22021 Several roost trees, reported to contain 'hundreds' or 'thousands' of bats, were cut down.'3601 Some of these records specifically concerned long-tailed bats.'1 0lCases of trees containing roosts being felled for firewood and timber production are still being reported,'20301 but declines have also been documented in areas with little forest modification.'2028; 22021 Conservation status. Long-tailed bats were common throughout New Zealand in the 1800s, but by 1900-1930 bats were becoming scarce in many districts, and they are now rare or absent at many sites where, even as late as 1990, they were still common and their conservation status regarded as secure'.'703'6971 Formal surveys in the South I. since then have failed to find long-tailed bats, or recorded them in low numbers, at 13 of 15 sites.'20281 Conservation programs are being implemented by DOC'18331 and some regional authorities.'20301 These include protection of roosting sites from disturbance or habitat loss, habitat enhancement, predator control programs in important habitats, and education packages.'153' 13891 Monitoring to measure the response of some populations to management has started.'2044'20401 Captive husbandry is possible over the short term,'241 1 though captive breeding has not been attempted. C.F.J.O'D. SUPERFAMILY NOCTILIONOIDEA The Noctilionoidea (previously called Phyllostomoidea) is a large superfamily, more diverse than any other superfamily of bats, divided into four families with 162 extant species: Mystacinidae (1 species), Noctilionidae or fishing bats (2 species), Mormoopidae or moustached bats (8 species), and Phyllostomidae or New World leaf-nosed bats (151 species)."3161 The group includes insectivorous, carnivorous, sanguinivorous, nectarivorous, frugivorous, and omnivorous species, and all except Mystacina are now restricted to central and South America, most in the tropics or subtopics. Although attempts to determine Mystacinas phylogenetic relationships using morphological features have produced conflicting results,'1468' molecular analyses consistently assign Mystacina to the superfamily Noctilionoidea. Mystacina is regarded as close to Noctilio,[2X7X] probably representing the most basal group in the superfamjjy [1468] phylogenetic analyses of DNA sequences from mitochondrial"412' 27621 and nuclear'26,4: genes confirm an ancient link between Mystacina and the Noctilionoidea. These phylogenies supersede classifications based on morphological features, which place Mystacina in the superfamily Vespertilionidae.'24621 FAMILY MYSTACINIDAE The family Mystacinidae is a distinctive southern lineage now represented only by the genus Mystacina. Three fossil species attributed to Mystacinidae have been identified from early to middle Miocene deposits in Australia."1661 The fossil species, at present placed in the genus Icarops, have a combination of characters shared only with M. tuberculata and M. robusta (in particular, the loss of two lower incisors, a large single rooted P,, and a moderately reduced M3). Genus Mystocino This endemic New Zealand genus contains a single surviving species, the lesser short-tailed bat Mystacina tuberculata. A second, larger species, the 110 | Order Ch iroptera greater short-tailed bat, M. robusta, is probably extinct (p. 129). Mystacina has a mosaic of morphological adaptations, including derived and convergent features, which arose as a result of its diverse lifestyle.'14681 Mystacina is a genus of medium-sized microchiropterans with a distinctive stocky body shape. Their fur is grey-brown, short (c. 7 mm), dense, and velvety, sometimes appearing frosted. The bare skin of the ears, wings, nose, legs, and tail is grey-brown. The forehead slopes steeply and the muzzle (c. 11 mm long) has a conspicuous array of whiskers. The incisors are sturdy and chisel-like. The nostrils are prominent and tuberculate (wart-like). The ears are large (c. 18 mm long X c. 9 mm at base), oval, and simple, with a long (c. 10 mm) tragus. The propatagium (leading edge of the wing) and proximal regions of both the plagiopatagium (hand-wing) along the arm and body, and uropatagium (tail-wing) close to the legs, are thickened and striated. When not in flight the wing is folded under these thicker regions to protect the delicate flight membranes. The second digit of the wing is reduced to a metacarpal and a single minute phalanx. The proximal section of the tail lies within the uropatagium, but the distal end projects c. 7 mm from its dorsal surface. The legs are unusually robust for a microchiropteran species, and the feet are stout and broad (c. 6 mm). A very fine talon at the base of the inside curve of each claw, only just visible to the unaided eye, is a unique characteristic of the genus. Lesser short-tailed bat Mystacina tuberculata Gray, 1843 Synonyms Mystacina velutina Hutton, 1871; Mystacina tuberculata Dobson, 1873; Mystacops tuberculatus Lydekker, 1891; Mystacops velutinus Thomas, 1905; Mystacops tuberculatus Miller, 1907; Mystacina tuberculata Simpson, 1945. Also called northern short-tailed bat, New Zealand long-eared bat (English); pekapeka (Maori). A recent ruling by the International Commission for Zoological Nomenclature'13421 conserving usage of the names Mystacina and M. tuberculata, clarifies a longstanding confusion. Gray,'111 i believing he was dealing with only one species of bat from New Zealand, mistook an illustration by G. Forster of New Zealand's longtailed bat Chalinolobus tuberculatus (then called Vespertilio tuberculatus J.R. Forster, 1844) with two specimens of short-tailed bat, which he described as M. tuberculata. The difference between them was soon recognised,'2'0"' and Hutton'131 ] proposed the name M. velutina for M. tuberculata, so that the two species of New Zealand bats would have different specific epithets. Kirsch et al."40' supported by Mayer et al." °4' revived Hutton's idea, but the Commission accepted"342' Spencer and Lee's[2522' argument for retention of the name M. tuberculata. Description Distinguishing marks, Table 22 and Plate 3. Specific characters, in addition to those given in the genus description, are as follows. M. tuberculata is smaller than M. robusta, with longer, narrower nostrils and proportionately longer ears, forearms, and wing elements."24/1 The ears extend to, or beyond, the muzzle, when laid forward. The rostrum is lighter and less massive, and the braincase rises more abruptly from the rostrum than in M. robusta. Chromosomes 2n = 36'230' Dental formula I'/j C1/, Pm2/, M3/3 = 28. Field sign Daytime. Occupied or recently occupied large colonial roost trees are scarce (i.e., two or three occupied roosts at any one time in >100 km2 of central North I. forest"589'). Some occupied colonial roosts can be easily recognised by the Lesser short-tailed bat | 111 accumulation of droppings at their base, the noise of bats within the roost, and a distinctive musky smell, but this is not true for all colonial roosts. Often there is little or no visible build-up of droppings, because they remain within the roost tree. If the group is small, the roost is high above the ground, or the bats are several metres above the roost entrance, there will be little detectable noise or smell. Roosting bats are usually quiet in the mornings, but during afternoons and early evenings they can be heard moving around within the roost and making 'chirupping' social calls, which have audible and ultrasound components, allowing bats' presence to be confirmed using a bat detector. There is a wide variety of small colonial and solitary roosts, but they are not distinctive. Mating roosts can usually be recognised by the stained area around the small roost entrance (c. 25 mm diam.) polished with a brown musky secretion from the male bats. When fresh, individual droppings are dark brown pellets, 4—5 x 3 mm. Typically, they contain finely comminuted insect fragments, recognisable only microscopically. During summer and autumn they may contain fruit pulp and small seeds. In early summer the droppings are sometimes almost exclusively chewed wood from roost excavation, and in late summer, when the bats moult, there may be large amounts of fur mixed in with the droppings. The presence of the short-tailed bat's distinctive and flightless commensal batfly Mystacinobia zelandica is convincing evidence that short-tailed bats have used a cavity as a roost. Individual batflies, or their exoskeletons, may persist either around roost entrances, or in the guano below the roost, for several weeks after roosts have been abandoned by the bats. Swarms of a small black fanniid fly (Fannia sp.) specific to roosts of the short-tailed bat"290' often hover round the entrance of active roosts. For echolocation calls, see Figure 12 and below. Night-time. Short-tailed bats do not usually fly until after it becomes dark in the forest, and they are not easily seen or identified in flight, though occasionally they will circle an observer as if inspecting her. The most effective way to detect short-tailed bats is using bat detectors at night. The echolocation calls of long-tailed and short-tailed bats are different (Figure 12), so it is usually possible to distinguish the two species reliably using narrow band (heterodyne) bat detectors. Short-tailed bat echolocation calls can be detected over a wide range of frequencies extending from 20 to 100 kHz. At the optimum frequency, 28 kHz,"833,2120' short-tailed bat echolocation calls are a rapid series of short staccato clicks (20-50/second), whereas long-tailed bat echolocation calls are a slower series of soft 'thwacks' (c. 8/second). Changes in the nature of long-tailed bat calls during prey capture (feeding buzzes) can confound species identification, but feeding buzzes are rarely heard in the absence of other, more characteristic, calls. Feeding buzzes are not often heard from shorttailed bats."389' Monitoring flowering plants fed on by shorttailed bats (e.g., Dactylanthus taylorii and Metrosideros spp.) can be used as a supplementary method of locating short-tailed bats,'845' but is not as effective as wide-scale surveys of large stands of old growth forest with automatic bat detecting systems. From November to April, male bats may be heard singing a high-pitched repetitive audible call resembling that of the rifleman (Acanthisitta chloris). A strong broadband ultrasound component to the song can be heard using a bat detector. Both audible and ultrasound components can be detected over a range of 50 m. The males generally sing in small groups, either from small cavities in trees (mating or singing roosts) or while moving 124 | Order Chiroptera bat would receive the median lethal dose (LD50) of 1080 from as little as 0.04 g of arthropods. The risk of secondary poisoning with brodifacoum may be higher, as it has a cumulative effect and persists in the environment. However, aerialbroadcast 1080 had no measurable effect on a bat population at Rangataua,1159S1 and bat populations persist at Pureora and north Waitaanga despite extensive possum control operations with a variety of toxins and baits. The bat population on Codfish I. survived the aerial-broadcast brodifacoum used to eradicate kiore.'2424' Predators In pre-European New Zealand, at least three native avian predators hunted lesser short-tailed bats: the extinct laughing owl Sceloglaux albifacies, New Zealand falcon Falco novaeseelandiae, and morepork Ninox novaeseelandiae. Until kiore arrived in New Zealand, the laughing owl took large numbers of bats and left piles of Mystacina bones in their middens;129441 falcons also took them occasionally.129431 More recently, morepork have been observed hunting outside colonial bat roosts, apparently targeting juvenile lesser shorttailed bats learning to fly.115891 The effects on lesser short-tailed bats of mammalian predators introduced since human settlement are uncertain. Kiore are probably not now a serious threat, since bats have coexisted with them at high densities on Little Barrier I. and (formerly) on Codfish I. for many decades. The cats, mustelids, and rodents brought by Europeans have probably reduced lesser short-tailed bat populations, either by direct predation or by competition for invertebrates, but the evidence is largely circumstantial. Introduced predators have not had any catastrophic effects on lesser short-tailed bats in the central North I., where large populations of the bats have persisted in their presence for over 150 years. Nevertheless, introduced predators could have contributed to their disappearance from other mainland forests. Six of 23 dead lesser short-tailed bats handed in by the general public were caught by domestic cats,169 1 but short-tailed bat remains have not been recorded in diet studies on feral cats. During video surveillance of short-tailed bat roosts at Rangataua Forest, stoats and ship rats were recorded visiting roosts, though only stoats actually entered them.1'5891 No kills by either species were recorded. There is circumstantial evidence that ship rats prey on short-tailed bats. Daniel and Williams1703' reported that both species of Mystacina disappeared from Big South Cape and Solomon Is following an irruption of ship rats in 1965. In Northland, ship rats were seen inside two fallen hollow roost trees occupied by several hundred lesser short-tailed bats. Twenty-nine (72%) of 40 bat skeletons collected from these roosts had a hole chewed in the back of the skull and the brains removed, presumably by ship rats,1696:7031 but it is not known whether rats killed the bats or scavenged them. Colonial summer roosts are particularly vulnerable to predators, because they are often noisy and smelly, and have large entrances low to the ground. Yet, long-term video surveillance shows that predation at these roosts is rare. Lesser shorttailed bats are extremely aggressive and could deter a predator entering the roost by attacking in large numbers. Large group sizes in colonial roosts, and frequent movements between roosts several kilometres apart, reduce individual predation probability by predator confusion and satiety. Bats leave a roost in small groups at random intervals, and individuals fly at high speed in different directions ('burst-emergence'). During winter hibernation, roosting bats immobilised by torpor for long periods are vulnerable to predators, but the entrances to winter roosts are usually too small for rats or stoats to enter. Terrestrial and arboreal foraging might be expected to make short-tailed bats vulnerable to Lesser short-toiled bat | 125 mammalian predators,1703; 18331 but they would not be easy prey because they are cryptic, fast moving, with acute hearing and sense of smell, and can take to flight easily. Parasites and associated insects A single record of the long-tailed bat flea {Porrihius pacificus), recorded on a museum specimen of M. tuberculata,18261 is probably an accidental contamination; otherwise, M. tuberculata has no host-specific fleas, and does not share any fleas with other native or introduced species.16961 It does host an undescribed species of tick belonging to the genus Argas (Carios), which includes four other species found on several Australian bat species (G.W. Ramsay16961). Several undescribed species of mites have been found in the fur of M. tuberculata.1826: 6961 A new species and genus of mesostigmatid mite Chirolaelaps mystacinae was implicated in the death of six M. tuberculata in Wellington Zoo.11214; 12151 A second species of Chirolaelaps remains to be described.112131 A new species of sarcoptic mange mite Chirophagoides mystacops, belonging to a new genus and subfamily, was described from the wing of a single M. tuberculata in the British Museum (Natural History) collected on Solomon I.1903:9041 An unidentified sarcoptic mange mite, found on bats from the Rangataua Forest, caused high levels of mortality in young captive-born bats as a result of massive subcutaneous secondary infection.119 1 A sarcocystis parasite has been identified in muscle tissue of specimens collected from Codfish I.1819' The New Zealand batfly. M. tuberculata has a unique, and possibly symbiotic, association with the New Zealand batfly, Mystacinobia zelandicay2891 a wingless dipteran, with a cream and brown, spider-like body, 4—9 mm long. Although superficially similar to other batflies (superfamily Hippoboscoidea), New Zealand batflies evolved separately, and are placed in their own family, Mystacinobia, within the superfamily Oestroidea, where they are most closely associated with the Calliphoridae, or blowflies.[2076; 2323; 10951 Unlike hippoboscoid batflies, New Zealand batflies are not parasitic, but feed on bat guano throughout their life cycle. Their gut contents include all the components present in guano (pollen, fruit pulp, insect parts, and fungal hyphae) except seeds. 12S91 Batfly development depends on the high temperatures (27-30°C) and humidity found in occupied or recently occupied bat roosts. When bats move to another roost, they carry some of the flightless adult flies, particularly gravid females, clasped to their fur with specially adapted claws. For a few days after a bat roost has been deserted, large numbers of batflies congregate around roost entrances after dark to alight on any bats that land there. Batflies may be found in the fur of flying bats at any time of the year; commonly there is only one fly per bat, but up to 60 were observed on one bat on Little Barrier Id691 The relationship between batflies and bats in occupied roosts is symbiotic. The flies accelerate the breakdown of bat guano, and adult batflies appear to groom the bats. The flies feed or drink from the mouths of torpid bats, and in maternity roosts they also drink milk, spilled by suckling young. Noticeable vibrations, emitted from the flies' thorax, may deter bats from eating them. Batflies are associated with short-tailed bat populations throughout the North I.1'5891 and in the Eglinton Valley,124181 but not on Codfish I. although fragments of Mystacinobia sp. were reported from there.169 1 Adaptations to the New Zealand environment The wide range of foraging modes found in M. tuberculata, and their associated morphological adaptations, are presumed to be unique adaptations to New Zealand's pre-human environ- 126 | Order Ch iroptero ment.'694; I589i Terrestrial and arboreal foraging is made easier by furling of the wing and tail membranes, increased freedom of movement for the forelimb permitted by a reduced propatagium, robust hind limbs, and a wide range of movements of the femur. The unspecialised wing morphology permits a variety of flight strategies. The extensile tongue and the brush of fine papillae on its tip are suited to feeding on nectar, while the rough transverse ridges on the tongue may help in extracting juice from fruit. The dentition is essentially insectivorous, but one upper premolar and a lower premolar and incisor have been lost, and the remaining lower incisor is small. The resulting gap in the teeth at the front of the lower jaw facilitates use of the extensile tongue to feed on nectar. These adaptations could have evolved in response to the absence of significant ground predators and nocturnal competitors in pre-human New Zealand.'694' Alternatively, they could be a response to the extreme climate oscillations of the late Pleistocene, when a wide range of foraging behaviour would have been advantageous. Significance to the New Zealand environment As the only extant species of an ancient endemic family, lesser short-tailed bats provide a globally significant contribution to New Zealand's biodiversity. They also pollinate several indigenous forest species, including kiekie Freycinetia bauriana, wood rose Dactylanthus taylorii, pohutukawa and rata Metrosideros spp., rewarewa Knightia excelsa, and the perching lilies Collospermum hastatum and C. microspermum. None of these species is completely dependent on lesser short-tailed bats for pollination, but bats are often more effective pollinators than birds, and some plant species (e.g., F. bauriana and D. taylorii) are adapted for pollination by short-tailed bats."60 ; 8461 Lesser shorttailed bats disperse seeds of C. microspermum, C. hastatum, and F. bauriana in their droppings and adhering to fur and wing membranes. Conservation status. Previously abundant throughout New Zealand, lesser short-tailed bats are now endangered. The <50 000 surviving individuals are scattered between 13 known populations, of which only seven include >1000 bats. The other six populations each include only a few hundred bats, and are especially vulnerable to extinction. Various conservation ranking systems list M. tuberculata as: threatened,"881 vulnerable,'1139' a species of highest conservation priority,'183"'' and facing a high risk of extinction in the medium-term future."1'1' The most recent New Zealand Threat Classification System"261' lists the three subspecies separately. M. t. rhyacobia is classified as 'range restricted' due to 'human induced loss of range'. M. t. aupourica and M. t. tuberculata are both classified as 'nationally endangered', having 'small population subject to moderate to high recent or predicted decline' as a consequence of'human induced loss of range'. For M. t. aupourica and M. t. tuberculata, it is noted that: 'Confidence in the listing is poor due to the poor data available for assessment'. The core ranges of all known populations of short-tailed bats are within forest areas now afforded full legal protection, which should halt declines caused by deforestation. The prospects for long-term survival of the species will be improved by increasing the number and geographic range of populations with translocations, both to oldgrowth forests managed to reduce alien predators and competitors, and to restored forests on fertile lowland sites. For translocations and other conservation purposes, existing phylogeographic patterns"59" l590' should be preserved and, because all populations are demographically distinct,""'90' each population should be considered as a separate management unit."866' The four outlying mainland populations (Omahuta, Tararua, north-west Nelson and Eglinton Valley) deserve the highest priority for conservation effort. B.L. Greater short-tailed bat | 127 Greater short-tailed bat Mystacina robusta Dwyer, 1962 Synonym Mystacina tuberculata robusta Dwyer, 1962 Also called southern short-tailed bat, Stewart Island short-tailed bat (English); pekapeka (Maori). Originally described by Dwyer'826' as a large subspecies of short-tailed bat, M. t. robusta, restricted to the Muttonbird Is, and elevated to species status as the greater short-tailed bat M. robusta by Hill and Daniel."247' Description Distinguishing marks, Table 22 and Plate 3. Overall M. robusta was larger and stouter than M. tuberculata, especially in the skull.'2946' However, it had proportionately shorter ears, forearm, and wing elements, so the two species overlap in these measurements. Skulls of M. robusta have a slightly heavier and more massive rostrum with wider, shorter nostrils, and the braincase generally rises less abruptly from the rostrum. The ears do not reach the muzzle when laid forward.'1247' Dental formula I1/, C1/, Pm2/2 M3/3 = 28. Measurements Measurements are of recent specimens collected from the Muttonbird Is (DM 1083, 1553, 1554, 1555.1,2 and 1629.2-4; MNZ, Wellington): total length 70-85 mm (up to 90 mm);'826' snout to vent length 65.4—72.4 mm (n = 8; mean 68.6 ± 2.86 mm);"592' wingspan 290-310 mm; tail 15 mm;[826:6971 forearm 45.3-47.5 mm (n = 8; mean 46.4 mm); tibia 18.2-19.1 (n = 8; mean 18.7 mm); ear 17.7-18.6 mm (n = 8; mean 18.3 mm); greatest length of skull 22.2-23.5 (n = 5; mean 23.2); condylobasal length 21.0-22.5 (n = 5; mean 22.0)."247' No live specimens have been weighed. Estimates of the body mass of M. robusta, 25—35 g'696' or 15.1-15.8 g,"589' calculated from the regression of body mass against forearm length of M. tuberculata are invalid, because the forearm is proportionally shorter in M. robusta. The overall linear dimensions of M. robusta are c. 20% greater than for M. tuberculata, so the normal cubic relationship between linear dimension and mass suggests that M. robusta was 70% heavier (c. 24 g). Variation After comparing size variation among recent specimens of M. robusta from the Muttonbird Is and fossil remains from throughout New Zealand, Worthy'2939' concluded that the greater short-tailed bat was a single species with a clinal variation in body size of 10-20% decreasing from the most northerly specimens down to the most southerly ones. History of colonisation See p. 113. Distribution M. robusta was endemic to New Zealand, often sympatric with M. tuberculata,'2938' Recent fossil remains (<20 000 years old) have been found in caves, on rock ledges, and in swamp sites in Waitomo, Hawke's Bay, and Wairarapa in the North I., and north-west Nelson, Westland, Canterbury, and central Otago in the South Island.'294" 522: 29441 No live or freshly dead specimens have been reported from the three main islands since European settlement. From 1840 until the early 1960s, living M. robusta were found only on the rat-free Muttonbird Is of Big South Cape I. (Taukihepa, 930 ha) and Solomon I. (Rerewhakaupoko, 32 ha), 2—10 km off the south-west coast of Stewart I.'703; 69 ' There have been no positive sightings of it since 1967. Direct evidence that M. robusta and M. tuberculata were formerly sympatric on the Muttonbird Is'1247; 697] j-,een controversial. All nine specimens of Mystacina collected on Big South Cape I. and held in MNZ are M. robusta,"5921 including a juvenile, 128 | Order Chiroptera DM 1629.1, originally identified as M. t. tuberculatum ' But other collections reportedly include M. tuberculata from the Muttonbird Is, e.g. two females from Solomon I. (BMNH 89.10.27.2—3).129461 Habitat There are differences in the relative proportions of greater and lesser short-tailed bats in fossil deposits around New Zealand, e.g., more M. robusta in Canterbury, fewer in north-west Nelson. 2939' The Muttonbird Is are granite, overlaid by a deep mantle of peat, covered mainly by muttonbird scrub (Olearia lyallii and O. angustifolia) and southern rata (Metrosideros umbellata), with some podocarp-broadleaf forest. There are numerous sea caves along their rugged shorelines. Abundant soil arthropods nourished by guano from breeding seabirds, especially sooty shearwaters (Puffinus griseus), may have been important to sustaining the bat population in these southerly latitudes. Food M. robusta probably ate the same wide range of foods as M. tuberculata-. arthropods, fruit, nectar, and pollen.1696' Pollen analysis of stomach contents of two M. robusta found both rata pollen and fern spores.1694' Reports that Mystacina are carnivorous and scavenge vertebrate carcasses1694' 6971 stem principally from observations of greater short-tailed bats on Solomon I.1211 ' Seven bats were held in captivity for a few days. On the first night the only food provided was the skinned carcass of a diving petrel: some flesh was consumed on the first night, but none subsequently. Reports that short-tailed bats on the Muttonbird Is had been observed chewing fat and meat off plucked muttonbirds (fledgling sooty shearwaters) hung out overnight182,i were unsubstantiated. Social organisation and behaviour Flight. Estimates of wing loading and aspect ratio for M. robusta[]C>c"'] based on a body mass estimate of 24.5 g, were higher (9.2 and 20.2 Nm"2) than for M. tuberculata, indicating that M. robusta would have been faster in flight and more agile, (i.e. with more rapid turns), but less manoeuvrable (i.e., requiring larger turning radii). Echolocation calls. There is a linear relationship between the frequency of maximum intensity of the echolocation calls of insectivorous bats and their forearm length.12"' ' It predicts that the frequency of maximum intensity for greater shorttailed bats should be 1-2 kHz lower than that of lesser short-tailed bats, i.e., 26-27 kHz. Roosting. Abundant remains of greater short-tailed bats in limestone caves around Waitomo show that they roosted in caves on the mainland.12942' Presumably they also roosted in tree cavities, although evidence for this would not persist. On Big South Cape and Solomon Is, bats were observed roosting in large colonies in sea caves along the granite shoreline, in cavities in the trunks of southern rata125"7'7031 and in seabird burrows.1697' Torpor and hibernation. Stead1255 1 describes removing cold and sluggish (i.e., torpid) bats from a day-roost on Solomon I. during early summer. Like many other temperate-zone microchiropteran species, they almost certainly also used seasonal hibernation, with occasional flights during the winter months, observed on Solomon I. between March and August.11641' Reproduction and development There are three pregnant greater short-tailed bats in museum collections, dated December 1931, May 1951, and August 1964, but these could be the dates of accession, not of collection.169 1 Field observations are scanty. J.A. Mackintosh collected a juvenile specimen (1-2 weeks old) on Solomon I. in late May, and also observed several nursery colonies with young, in hollow southern rata trees and in the burrow of a sooty shearwater, from late April to mid May of Suborder Megachiroptera | 129 1963—65.1697' These observations suggest that M. robusta gave birth in autumn on the Muttonbird Is, presumably to coincide with peak soil arthropod abundance during autumnal fledging of muttonbirds, rather than midsummer as observed in M. tuberculata throughout New Zealand. Population dynamics Decline. In 1961, several hundred bats were observed in Puai Cave on Big South Cape I., and bats were also regularly seen flying on both Big South Cape and Solomon Is.169/1 Ship rats arrived on the two islands in the early 1960s, and reached plague numbers between 1964 and 1967.11861 Bat numbers on the two islands declined rapidly: by 1965 very few flying bats were seen, and in 1966 no bats were reported from either island. The last confirmed sighting was in November 1967, when, although no bats were seen flying, four unidentified mystacinid bats were observed roosting in Puai Cave. The last confirmed reports of M. robusta are: one collected from Big South Cape I. in August 1964, and another mist-netted on Solomon I. in April 1965.1'641' Predators Laughing owl Sceloglaux albifacies, morepork Ninox novaeseelandiae, and falcon Falco novaeseelandiae all hunted M. robusta.129421 Abundant robusta remains were found in the middens of laughing owls in north-west Nelson and Canterbury,12941' 2943"2945' and smaller numbers in falcon deposits in north Canterbury.12943' Stead12557' reported a short-tailed bat (presumably robusta) killed by a morepork on Solomon I. The simultaneous disappearance of M. robusta from the main islands and the appearance of kiore in laughing owl middens suggests that kiore wiped out M. robusta shortly after they arrived.1 03; 69" 2945] -phg jarger yif robusta would have been more vulnerable to predation in their hibernacula than M. tuberculata, as crevices accessible to them would also be accessible to kiore. An alternative hypothesis, that M. tuberculata and M. robusta are not separate species,182 ' but the result of selective predation by kiore against larger bats now recognised as M. robusta,[xm] is not supported by analyses of new Mystacina skeletal material.129'161 Ship rats caused the final extinction of the greater short-tailed bat on its last two island refuges.1'861 Significance to the New Zealand environment Conservation status. M. robusta is assumed to be extinct, as there have been no confirmed records since 1965-[703:697; 1,41 Searches for it continue, and two possible sequences of mystacinid-like calls were recently recorded on Putauhina I., close to the last island refuge of greater short-tailed bats.12027' The most recent New Zealand Threat Classification System11261' lists the species as 'data deficient'. B.L. SUBORDER MEGACH I ROPTE RA FAMILY PTEROPODIDAE This family, the only one in the Megachiroptera, includes about 173 species in 44 genera, all native to the Old World tropics from Africa to the Pacific including Australia. All members of this family feed on fruit, pollen and/or nectar, have large eyes and, except for one genus (Rousettus), navigate by vision rather than echolocation. Genus Pteropus Of about 65 species of Old World giant fruit bats or flying foxes, four are found in Australia, and the rest in Africa, Madagascar, India, South-East Asia, and the Pacific. These are large to very large bats with heads and faces resembling those of foxes, prominent eyes, large simple ears with no tragus, and no noseleaf. All have a claw on the second finger as well as a claw on the thumb. 130 | Order Ch iroptera Little red flying fox Pteropus scapulatus Peters, 1862 Also called little red fruit bat, collared flying fox. See Plate 3. This is the smallest of the four species of Australian flying foxes, with reddishbrown fur, a light brown-yellow mantle and pale fur round the eyes. When in flight, the wings appear partly transparent. Dental formula I 2/2 C '/, Pm3/3 M2/3 = 34. Measurements Head and body length about 220 mm, tail very short, and forearm length 118-132 mm.'2'78' Distribution Widespread in eastern and northern Australia, from dry inland areas to the coast. This nomadic species can migrate long distances within Australia, following the irregular flowering of eucalyptus blossom. There is only one vagrant record from New Zealand. A specimen was found electrocuted under powerlines after a storm in Hamilton East in about 1927-29.'693' Unusually large migrations of this species into New South Wales and Victoria were observed a few years after a severe drought in Queensland in 1926-27;'2234' this specimen may have been the only survivor of a group blown across the Tasman Sea at that time. A photograph of it, taken by J.E.C. Flux after it had been hanging on a barn wall for many years, was included in Daniel's description of this incident.'697' This is the only documented record of an Australian bat reaching New Zealand alive since European settlement, although undoubtedly other Australian fruit bats have been blown to New Zealand in the past but died out unrecorded. In the 1870s sightings of large unidentified bats were reported near Wellington, Wanganui, and the Clarence River.'2574' C.M.K ORDER LAGOMORPHA Lagomorphs are smallto medium-sized herbivores that, like rodents, have a pair of large, chisel-shaped incisors separated by a long diastema from the grinding molars. In other respects they are distinct from rodents: the molars (as well as the incisors) grow continuously throughout life; there is a second pair of small, peg-like incisors directly behind the first pair; the nostrils are covered by a retractable flap of skin; the gut has a large caecum with a spiral septum inside; and the tail is very short. They maximise the value of their food by reingestion (also called refection, or coprophagy). There are two families: the Ochotonidae (c. 25 species in one genus), the short-eared pikas of montane Asia and western North America, and the Leporidae (56 species, 11 genera), which includes all rabbits and hares, distributed globally. FAMILY LEPORIDAE The Leporidae contains one large genus, Lepus, mainly open-country hares, and ten smaller genera of mainly burrowing rabbits, including Oryctolagus. All have long ears, hind legs, and feet. Two species of leporids are present in New Zealand, distinguished as in Table 29. Genus Oryctolagus The genus Oryctolagus has only one species, the European rabbit. It is distinct from the hares, but superficially resembles some other genera of rabbits such as the American cottontails (Sylvilagus). European rabbit Oryctolagus cuniculus cuniculus (Linnaeus, 1758) Table 29 Distinguishing marks of lagomorphs in New Zealand Rabbit Oryctolagus cuniculus Brown hare Lepus europaeus Weight (adult male) Length of ears Length of hind feet Body colour Tips of ears Colour of eyes Gait 1.3-2.1 kg 60-70 mm 75-95 mm Grey-brown Narrow black rim Brown Bobbing, tail up, showing white underside when unhurried; scuttling rush, tail down, when alarmed 2.4-4.8 kg 90-105 mm 1 30-155 mm Tawny Black patch at tip Yellow Loping, tail down showing black upper surface 131