Mystacina tuberculata

MystacinaGray, 1843:296. Type species Mystacina tuberculataGray, 1843, by original designation (International Commission on Zoological Nomenclature [ICZN] 2002; Mayer et al. 1999). MystacopsLydekker, 1891:671. Unavailable replacement name for MystacinaGray, 1843:296, proposed under the incorrect premise that Mystacina Gray is preoccupied by MystacinusBoie, 1822 (Lydekker 1891; Mayer et al. 1999). Mystacops is on the Official Index of Rejected and Invalid Generic Names in Zoology (ICZN 2002). Order Chiroptera, suborder Yangochiroptera (Teeling et al. 2005), superfamily Noctilionoidea (Kennedy et al. 1999; Kirsch et al. 1998; Pierson et al. 1986; Teeling et al. 2003; Van Den Bussche and Hoofer 2000), family Mystacinidae (Miller 1907). Genus Mystacina includes 1 extant species, M. tuberculata, and 1 recently extinct species, M. robusta, which has not been sighted since 1967 (Molloy 1995). M. robusta was originally described as a subspecies, M. t. robusta (Hill and Daniel 1985). Mystacina tuberculataGray, 1843:296. Type locality “New Zealand;” and 1st use of current name combination. Mystacina velutinaHutton, 1872:186. Type locality “[T]he Hutt Valley, near Wellington,” North Island, New Zealand, and “Milford Sound, on the southwest coast of the South Island,” New Zealand. This name is on the Official Index of Rejected and Invalid Specific Names in Zoology (ICZN 2002). Mystacops tuberculatus: Lydekker, 1891:671. Name combination. Mystacops velutinus: Thomas, 1905:423. Name combination. Content as above. Three morphologically distinguished subspecies (Hill and Daniel 1985) are supported by some molecular data (Winnington 1999). Mitochondrial gene sequences distinguish 6 lineages (NW, NL, NE, S1, S2, and SN), with divergence dates of 0.93–0.68 million years ago (Lloyd 2003a). These lineages do not align with morphological subspecies listed below. M. t. aupouricaHill and Daniel, 1985:294. Type locality “Omahuta Kauri Sanctuary, Northland, North Island, New Zealand, 35°10′S, 175°37′E.” M. t. rhyacobiaHill and Daniel, 1985:295. Type locality “Te Rimu area, upper Waimarino River, Kaimanawa Forest Park, SE of Lake Taupo, central North Island, New Zealand, 39°02′S, 175°56′E, ca. 700 ft.” M. t. tuberculataGray, 1843:296; see above. Mystacina tuberculata (Fig. 1) can be distinguished from the extinct M. robusta using the following measurements (range, in mm, M. tuberculata and M. robusta, respectively—Hill and Daniel 1985): length of forearm, 40.0–45 and 45.3– 47.5; condylobasal length, 17.3–19.1 and 21.0–22.5; length of mandibular toothrow (c–m3), 7.3–8.0 and 8.9–9.4. Ears of M. tuberculata extend to, or beyond, tip of muzzle when laid forward; ears of M. robusta do not reach tip of muzzle when laid forward. Skeletal measurements distinguish the 2 species better than external features (Worthy and Scofield 2004). Adult female Mystacina tuberculata tuberculata from the Eglinton Valley of Fiordland, New Zealand. Photograph by Daniel K. Riskin. Adult female Mystacina tuberculata tuberculata from the Eglinton Valley of Fiordland, New Zealand. Photograph by Daniel K. Riskin. Mystacina tuberculata is readily distinguished from Chalinolobus tuberculatus, the only other extant species of bat in New Zealand; M. tuberculata has pointed ears that extend past nose when laid forward, prominent tubular nostrils, and tail that extends 6–8 mm from dorsal surface of uropatagium, whereas C. tuberculatus has a tail fully enclosed in the uropatagium (Daniel 1990; Lloyd 2001). Mystacina tuberculata and C. tuberculatus also can be distinguished by echolocation calls using a heterodyning bat detector. At 28 kHz, M. tuberculata echolocation flight calls are a series of short staccato clicks (20–50/s), whereas C. tuberculatus calls are a series of soft “thwacks” (8/s—Lloyd 2001). Pelage is very thick and velvety. Dorsal fur is grayish to dark brown and ventral fur is paler. Bare skin of ears, wings, nose, legs, and tail is gray-brown (Lloyd 2001). Unique external features of M. tuberculata include well-defined, thick, and striated proximal region of wing membranes and small basal talons at inside curve of toe and thumb claws. Length of body is 60–70 mm plus ca. 20 mm with extended uropatagium; wingspan is 280–300 mm (Lloyd 2001). Tail reaches a length of up to 12 mm (Lloyd 2001) and projects 6–8 mm from uropatagium (Daniel 1990). Tibia is 14.5–17.0 mm in length and feet are stout, broad, and ca. 6 mm long (Lloyd 2001). Loose wrinkled skin covers soles of feet. Ears are 17.5–19.1 mm in length and 9 mm wide at base (Daniel 1990; Lloyd 2001). Tragus is simple, pointed, and ca. 10 mm in length (Lloyd 2001). Nostrils are large and vertically oblong. Muzzle is conical, obliquely truncated, relatively long, and has an array of whiskers encircling mouth and nostrils (Dwyer 1962a). Skull is twice as long as broad (Fig. 2), and width is ca. 1.33 times height of cranial cavity (Dwyer 1960a). Condylobasal length is 17.3–19.1 mm (sample size unknown—Hill and Daniel 1985). Dorsal, ventral, and lateral views of cranium and lateral view of mandible of an adult male Mystacina tuberculata aupourica collected at Omahuta Kauri Forest, New Zealand (Royal Ontario Museum specimen ROM80308). Greatest length of skull is 20.3 mm. Used with permission of the photographer J. L. Eger. Dorsal, ventral, and lateral views of cranium and lateral view of mandible of an adult male Mystacina tuberculata aupourica collected at Omahuta Kauri Forest, New Zealand (Royal Ontario Museum specimen ROM80308). Greatest length of skull is 20.3 mm. Used with permission of the photographer J. L. Eger. Adults weigh 10–22 g before foraging (Lloyd 2001; O'Donnell et al. 1999). Body mass can increase by 20–30% while foraging, and preforaging body mass can increase by 20–30% during preparation for periods of torpor (Lloyd 2001). Female body mass can increase up to 35% during pregnancy (Lloyd 2001). Morphometrics of M. tuberculata vary across subspecies and populations. Forearm measurements (mean, SD, and parenthetical range, in mm, sexes combined) are as follows: M. t. aupourica (n = 49): 40.90, 0.89 (36.95–42.00); M. t. rhyacobia (n = 999): 43.64, 1.24 (39.88–46.90); M. t. tuberculata (n = 304): 42.23, 1.01 (39.44–45.10—Lloyd 2001). Specimens of M. tuberculata from the Eglinton Valley of Fiordland had significantly greater weights, larger forearms, and smaller ears, than those from Codfish Island and Little Barrier Island (O'Donnell et al. 1999). M. tuberculata in the Eglinton Valley is sexually dimorphic with respect to mass and forearm length, with females being significantly heavier and having longer forearms (O'Donnell et al. 1999). M. tuberculata on Little Barrier Island showed no difference in forearm length between males and females, although females were significantly heavier than males (Arkins 1996; Winnington 1999). M. tuberculata in Eglinton Valley also showed greater variation in size between age classes than did M. tuberculata on Codfish and Little Barrier Islands (O'Donnell et al. 1999). Mystacina tuberculata is endemic to New Zealand (Fig. 3), where its distribution is highly correlated with presence of indigenous forest (Daniel and Williams 1984; Greaves 2005). On North Island, confirmed populations of M. tuberculata include a population at Omahuta-Puketi Forest in Northland, 7 large populations in central North Island, and a small genetically distinct population in the Tararua Ranges at the southern end of North Island (Lloyd 2001, 2003a). On South Island, isolated populations have been found in the Oparara Basin of Northwest Nelson, Eglinton Valley of Fiordland (O'Donnell et al. 1999), and Dart Valley of Aspiring National Park (C. F. O'Donnell, pers. comm.). Calls also have been recorded at Punakaiki on the West Coast (Lloyd 2001), and populations may exist in large unsurveyed forests of South Island (Greaves 2005). No calls have been recorded from surveyed forests around Mt. Taranaki, Wanganui River Trench, Matemateonga, Tongariro Forest, and the Ruahines (Lloyd 2001). Populations also exist on Little Barrier Island and Codfish Island, and a population is being established by the New Zealand Department of Conservation on Kapiti Island (B. D. Lloyd, in litt.). M. tuberculata is found from sea level to ca. 1,100 m, the highest altitudinal limits of New Zealand forest (Daniel and Williams 1984). Geographic distribution of populations of Mystacina tuberculata in New Zealand. Dots represent locations of known populations. Ranges of morphologically distinguished subspecies (1, M. t. aupourica; 2, M. t. rhyacobia; 3, M. t. tuberculata) are separated by dashed lines. Modified from the work of Lloyd (2003a;2003b), with inclusion of a recently discovered colony at Dart Valley (C. F. O'Donnell, pers. comm.). Geographic distribution of populations of Mystacina tuberculata in New Zealand. Dots represent locations of known populations. Ranges of morphologically distinguished subspecies (1, M. t. aupourica; 2, M. t. rhyacobia; 3, M. t. tuberculata) are separated by dashed lines. Modified from the work of Lloyd (2003a;2003b), with inclusion of a recently discovered colony at Dart Valley (C. F. O'Donnell, pers. comm.). Six genetically distinguishable subspecies (Lloyd 2003a) are sympatric within most populations (Fig. 3) and correlate only roughly with distribution of 3 original morphologically based subspecies. Within central North Island, phylogeographical structure and demographic history of these 6 lineages of M. tuberculata is with from following (Lloyd On South Island, phylogeographical structure a from in South Island of South Island years ago (Lloyd for an for and from 3 species in the extinct have been in et al. were found in at and and et al. of Mystacina from the have been collected from a series of New Zealand (Daniel 1990; and Mystacina tuberculata is of and and on the to the of M. tuberculata only 1 distinguishable a that did not include an the and of the of mass did not in a of mass were than and did not from to with et in external and (Dwyer have to and M. tuberculata can its within a of wing of and of wing (Dwyer are within a proximal region of and of wing when and are with greater of and a can be by only of and a wide of by and basal talons on inside curve of and in and (Dwyer 1962a). the is by the pers. comm.). Mystacina tuberculata has thick fur and of structure exist (Dwyer 1962a). are short and of is with 2 of near base and is and is to to (Dwyer 1962a). Mystacina tuberculata in and and also from to foraging O'Donnell et al. wing of flight for these et al. 2003; et al. and wing and from the flight during et al. 2003; and features of include a from and 3 in (Dwyer 1962a). Mystacina tuberculata has for with echolocation and small ventral and large upper (Dwyer 1960a). M. tuberculata on and in to has an (Daniel is 28 (Daniel 1990). are between and in and can be (Daniel is 7 6 (Dwyer 1960a). and 1st are (Dwyer 1960a). are with and 1st (Dwyer 1960a). 9 are 2 are and 2 are (Dwyer 1960a). is and of an surface on dorsal and to and an is to is ca. 12 mm in length, and has a tip in (Daniel New Zealand bat had a and an ca. mm in length Mystacina tuberculata has a long with external of extend most of length of is in of where in only and are (Lloyd 2001). Mystacina tuberculata is with in and by a in or during (Daniel 1990; Lloyd 2001). in and from to and during and from to (Lloyd 2001). are (Lloyd 2001). or in and a is in (Daniel 1990; at ca. from to (Daniel 1990; Lloyd 2001). Within a during a (Lloyd 2001). No New Zealand (n = during the on Codfish Island were or and and were in J. and M. in litt.). the were females, were were females, were of the and were adult males (n = in J. and M. in litt.). At Forest in central North Island, ca. of females whereas are (Lloyd 2001). On Little Barrier Island, of female were or in (Winnington 1999). At weigh ca. g and have a forearm ca. mm are 2 (Daniel Lloyd 2001). of are on dorsal and grayish on ventral surface (Daniel Lloyd 2001). of a in that and for 3 legs, and are at ears to of for and are and 1 (Daniel 3 are by and are and (Lloyd 2001). At ca. 6 the and by 12 are fully in size (Daniel 1990; Lloyd 2001). is with adult structure than adult mass (Arkins 1996; Daniel 1990). and foraging in forest populations of M. tuberculata are to of New forest that include large for and (Lloyd 2001). New Zealand in the Eglinton Valley in forest with a relatively structure Mystacina tuberculata up to 2 wide to reach foraging (O'Donnell et al. 1999), and small have been recorded in and near forests (Lloyd 2001). On Codfish Island and Little Barrier Island, have been foraging by and in (Arkins 1996; Daniel 1990; Winnington 1999). Mystacina tuberculata has a and foraging to and other small M. tuberculata and (Arkins et al. 1999; Daniel M. tuberculata is an with (Arkins 1996; et al. 1999). a of on of 2001). and on fur of M. tuberculata in Northland, of and and an that M. tuberculata also on (Lloyd 2001). that M. tuberculata on and of of and do not (Arkins 1996; et al. 1999; Daniel Lloyd 2001; and New Zealand are can in a an of to of its body mass (Daniel was ca. 20 (Daniel and an adult female bat 20 g in to g of (Daniel M. tuberculata g of or of body mass (Lloyd and No in were found age and classes of New Zealand on Little Barrier Island (Arkins of in at times or of of the following and and and et al. 1999). were from the and and et al. 1999). by M. tuberculata and and et al. 1999). by M. tuberculata include of of and C. and and of are also to be (Daniel Lloyd 2001). in have to or and from a wide of in and fur of M. tuberculata from central North Island (Lloyd 2001), Little Barrier Island (Arkins 1996; et al. 1999), and (Daniel M. tuberculata of (C. and and during periods (Daniel Lloyd 2001). central North Island, the only of (C. 2001). from and the and endemic is found only (Lloyd 2001). M. tuberculata a wide of and (Arkins et al. 1999; of and in that M. tuberculata and and other and the from its body fur during (Arkins et al. 1999; Daniel Lloyd 2001). Mystacina tuberculata a of indigenous forest (C. and C. and and (Lloyd 2001). M. tuberculata is an of D. New only fully and the only known to be by a bat 1995). M. tuberculata also of New Zealand and (C. and C. Lloyd 2001). of M. tuberculata had no with other for and of by (Daniel 1990; in M. tuberculata populations have been to of the and the smaller or species of Mystacina from South and Islands of (Daniel 1990). of M. tuberculata have with on Little Barrier Island and Codfish Island (Lloyd 2001), in New Zealand bat populations of from Codfish Island (C. F. O'Donnell, pers. comm.). On central North Island, and other have not had on populations (Lloyd 2001). of M. tuberculata include the New Zealand and extinct and 1996; Lloyd 2001). 6 were by and 1 by a (Daniel and Williams 1984). and have been at M. tuberculata has not been confirmed (Lloyd 2001). M. tuberculata species of described from specimen of M. tuberculata, a of (Daniel 1990; colony of 6 New Zealand from by the et al. species of on M. tuberculata (Daniel 1990). include found in the fur found in of and D. found in near is a large bat that on No or were found in 2 of M. tuberculata and Daniel and no were found in from from a colony in Omahuta (Daniel of in M. tuberculata from Codfish Island and found no of bat and no did a of of the 1st of in et al. of population size at Forest a between and (Lloyd 2001). at other locations have population from to (Lloyd 2001; O'Donnell et al. 1999). with are to forearm wing and forearm of New Zealand (Lloyd 2001). sexually males in to or J. Daniel and D. in Lloyd 2001; O'Donnell et al. 1999). Eglinton Valley, were in and and were smaller than those for (O'Donnell et al. 1999). in are with a brown from of males (Lloyd 2001; C. F. O'Donnell, pers. comm.). This has a and may females (Lloyd 2001). At males from for periods of at a (O'Donnell et al. 1999). calls are to the and have a (Lloyd 2001). in and (Daniel in litt.). to of males (Daniel 1990). are from other males and are in years J. Daniel, in litt.). with locations of female near are only (Lloyd 2001). on are to be (Winnington 1999). Mystacina tuberculata has echolocation calls with of et al. 2003; 2001). New Zealand also have calls with a of and a et al. 2003; Lloyd 2001; O'Donnell et al. 1999; Calls a and 2 other with an (Lloyd 2001; 2001). with the most is in the may be found in Calls by 1st from to is from to kHz, and when is from to with the most ca. and for and 2001). Calls have an of and an of 2001). Calls and of New Zealand in forests are than those in from et al. and of significantly between from calls of New Zealand that were in a flight or from (Arkins 1996; O'Donnell and 2001) are to M. has a bat and a that the and is to M. an and using of calls from M. tuberculata, can of calls to species (n = 2001). Mystacina tuberculata when foraging for and the and et al. 2003; Lloyd 2001; New Zealand are to foraging as follows: and and (Daniel M. tuberculata ca. of foraging on the or and as an of of foraging on the M. tuberculata is the most of M. tuberculata and and are to under or in et al. M. tuberculata near and from the or under and et al. is the (Lloyd 2001). and M. tuberculata in the relatively between and also can in Lloyd 2001). New Zealand in central North Island and in southern South Island from to (Lloyd 2001; O'Donnell et al. 1999). flight is ca. long at of up to in of M. tuberculata near the southern of the in the Eglinton Valley in et al. are M. tuberculata a length of the with in a smaller (O'Donnell et al. 1999). 12 the of a male = range, et al. 1999). New Zealand (n = an of and most an of 2003a). from to with a of 2003a). Mystacina tuberculata on as using as and (Lloyd 2001; O'Donnell et al. 1999; in are larger than with respect to size of cavity and in and may up to (Lloyd 2001). New Zealand a of than those in have in larger at cavity greater cavity and cavity than other in are in the of the with larger than a greater of and in species as vary and across forest and include in Northland, and in forests of Little Barrier Island, in and and in forests (Lloyd 2001). On Codfish Island, was as a than be New Zealand based on in species as include southern 2001), and F. O'Donnell, pers. comm.). and are as (Daniel are for (Lloyd 2001). are (Lloyd 2001). the 2 females with and to times females may in other and only the to (Lloyd 2001). At females and males may to use the as a (Lloyd 2001). in the the do not in and in is and and at the southern of the on Codfish Island are known 2001). At New Zealand relatively and of the with a of at 2001). in central North Island, M. tuberculata are smaller than in and are for up to in a (Lloyd 2001). the M. tuberculata in forest of Fiordland were by females and with size = = New Zealand the for an of = = and female M. tuberculata in Fiordland do not in with during the in M. tuberculata in small in have in large and under in small in inside the of at the base of and in in the (Lloyd 2001). on Codfish Island, M. tuberculata on of = in 2001). in southern South Island, New Zealand on of (O'Donnell et al. 1999). of on Codfish Island are by New Zealand in Mystacina tuberculata in and that (Daniel and the on may be to (Dwyer At a colony of ca. M. tuberculata in a and at a and had been (Daniel of a was described as to a of 1 with and by the (Daniel of with bat and in ca. 10 by wide and up to were found with smaller a 6 by New Zealand in a to the adult females and had the of had been than twice in a in 1 was when were and recorded when were was was to be (Daniel in M. in litt.). of Mystacina in and small in of (Daniel Daniel and Williams 1984). No have been found in in large of in (Daniel 1990; Lloyd 2001; and Mystacina tuberculata is highly between and at for up to a at a (Lloyd 2001; O'Donnell et al. 1999; in large of New Zealand in and of most of the 2001). New Zealand (n = on = and between 2001). by New Zealand may be the or is a known for (Lloyd 2001). can be in for a 1st found in on Codfish Island was in New Zealand that from on Codfish Island, up to a 2001). of and on = of the with of correlated with 2001). of M. tuberculata is highly of from to (Arkins 1996; in Daniel 1990; O'Donnell et al. 1999). the southern of range, was by an = range, a of ca. = range, and a of ca. = range, O'Donnell et al. 1999). in the found that New Zealand the with only short and periods of = = in On Little Barrier Island, near the of range, around was with between and 6 (Winnington 1999). are on and although on are with 1 greater than the other (Arkins M. tuberculata is in J. Daniel, in torpor is by New Zealand in not at M. tuberculata extended torpor to during periods (Lloyd 2001) and has a and long of and foraging, inside and of 2001). M. tuberculata between of and of and to of may to be and for longer 2001). includes foraging, and 2001). central North Island, periods of torpor up to 10 are with periods of from 1 to (Lloyd 2001). At the southern of its on Codfish Island, M. tuberculata had at a of = of (n = on when was = On the of New Zealand 2001). M. tuberculata in the Eglinton Valley was than on Codfish Island using bat 2001). of M. tuberculata, based on 1 adult has of and of et al. with 6 of large 6 of 2 of and of small were not morphologically et al. Mystacina tuberculata is listed by the Conservation (International for the Conservation of and as et al. 2001). M. t. rhyacobia is listed as a subspecies, whereas M. t. aupourica and M. t. tuberculata are listed as 2002). population of M. tuberculata is at ca. (Lloyd 2001, of populations in the central and southern of North Island is the most for those populations before is million (Lloyd of a of for divergence and the of sympatric M. tuberculata no (Lloyd and isolated populations at Northwest Nelson, and Eglinton Valley were for highest by Lloyd to M. tuberculata include species, and in et al. 2001). M. tuberculata large of forest Lloyd 2001; O'Donnell et al. 1999), and forest and have (Molloy 1995). M. tuberculata to of and in large (Lloyd 2001, 2003a). that on is of New Zealand (n = an of did not (Lloyd and 2002). Mystacina tuberculata is under the of was in and a bat has been established et al. 2001). New Zealand Department of Conservation is to a population on Kapiti Island by 20 (B. D. Lloyd, in litt.). history of M. tuberculata from the original of the species by Gray in et al. 1999). Gray only 1 bat species on New Zealand, is listed under the name tuberculatus, a species to which to 2 proposed the Gray the 2 and did not had discovered a species, the of Mystacina tuberculataGray, 1843, has been of the Commission on Zoological Nomenclature Mystacina tuberculataGray, 1843, was as the and name for name Mystacina from the to the name tuberculata to small on the upper of C. tuberculatus and was to M. tuberculata the 2 species were originally Mystacina has been in 7 and (Daniel 1990). (Kennedy et al. 1999; Kirsch et al. 1998; Pierson et al. 1986; Teeling et al. Den Bussche and Hoofer using a of molecular for M. tuberculata as a of most of divergence from other is million years ago (Teeling et al. 2005). for is on the name include New Zealand New Zealand New Zealand New Zealand and New Zealand bat (Daniel 1990). of subspecies are forest New Zealand bat t. New Zealand bat t. and southern New Zealand bat t. name of M. t. aupourica is from the name or for the of New Zealand (Hill and Daniel 1985). name M. t. rhyacobia is from the a of and a of a to the in which the specimen was found (Hill and Daniel 1985). Lloyd, J. J. and C. O'Donnell that had in North J. C. O'Donnell, and 2 on of J. L. of the Ontario Museum the skull and

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