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The Thermal Physiology of the Mountain Pygmy-Possum Burramys Parvus (Marsupialia: Burramyidae)

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The Thermal Physiology of the Mountain Pygmy-Possum Burramys Parvus (Marsupialia: Burramyidae) THE THERMAL PHYSIOLOGY OF THE MOUNTAIN PYGMY-POSSUM BURRAMYS PARVUS (MARSUPIALIA: BURRAMYIDAE) MICHAEL R. FLEMING Fleming, M. R. 1985. The thermal physiology of the Mountain Pygmy-possum Burramys parvus (Marsupialia: Burramyidae). A ust. Mammal. 8: 79-90. The Mountain Pygmy-possum Burramys parvus is restricted to the Australian alpine region. Laboratory studies of the thermo-physiology of this species found that body temperature (T") was tightly regulated at 36.1 oc, but animals quickly become hyperthermic at ambient temperatures Cta) above 30 °C, causing the thermal neutral zone to be truncated. Basal metabolic rate was 2.15 W kg -"· ' 5 (mean body mass 44.3 g) and weight-specific thermal conductance was 0.112 rnl 1 O, g-' h-' oC- . These values are 9% and 44% lower, respectively, than the mass predicted value for a marsupial, showing that the overall rate of energy expenditure is considerably reduced in this species. Huddling also reduces individual rates of energy expenditure. Burramys parvus enters prolonged bouts of deep torpor lasting up to one week, during which T" is very low and close to T. and the rate of oxygen consumption greatly reduced. Spontaneous arousal occurred from a T" as low as 6oc and the overall rate of rewarming was 0.17 oc;rnin. Attainment of a critical body mass appeared to be necessary before an animal would enter torpor. Burramys parvus shows physiological adjustments similar to that described in many placental mammals from cold climates and this species represents a true marsupial hibernator. M. R. Fleming, Department of Zoology, Monash University, Clayton, Victoria, Australia 3168. Present address: Wildlife Research Section, Conservation Corn­ mission of the Northern Territory, P.O. Box 1046, Alice Springs, Northern Territory, Australia 5750. THE comparative thermophysiology of Localised, low-density populations have many northern hemisphere placental mam­ been recorded from Mt Hotham and the mals has been studied in detail with parti­ Bogong High Plains in Victoria and in the cular attention being given to arotic mam­ Kosciusko National Park in New South mals (reviews: Hart 1964, 1971 and Miller Wales (Mansergh 1984). Fossi-l remains 1978). The physiological adjustments found close to the modern form of B. parvus in placentals f·rom cold environments have have been found in owl deposits at much been used to infer general corrdations be­ lower altitudes (Broom 1896 and Wakefield tween climate and single physiological para­ 1972). These deposits are Pleistocene and meters (Scholander et al. 1950, Irving et Recent in age indicating that this species al. 1955, Hart 1964, McNab 1966 and had a much wider distribution in the recent Withers et al. 1979). These generalized past. placental 'adaptat•ions to cold' include in­ Extant populations are strongly associat­ creased insul•ation of the surface layers, ed wit1h peri-glacial boulder streams which peripheral cooling, increased body size, a are usually shrub covered (Gullen and basal metabolic rate higher than predicted Norris 1981) . This habitat would provide by mass, increased summit metabolism and an extensive sub-nivean environment during the ability to become torpid. the five months of t•he year that the ground Burramys parvus Broom is the only is usually snow-covered. The ecology of marsupial restricted to the alpine and sub­ B. parvus is reviewed by Kerle ( 1984) and alpine habitats of southeastern Australia. Mansergh ( 1984). This study describes 80 AUSTRALIAN MAMMALOGY thermoregulation and torpor in B. parvus The chamber was placed in a lighted as an example of a small marsupial adapt­ constant-temperature cabinet regulated ed to a cold environment. within ± 0.5°C of the desired setting. Temperature within the chamber was METHODS recorded on a Leeds and Northrup Speedo­ max W multipoint recorder. Dried room METABOLIC MEASUREMENTS OF air passed through a copper heat exchanger NORMOTHERMIC ANIMALS before entering the metabolic chamber. The animals used in this study were Oxygen consumption rates were deter­ obtained from two sources. Five captive­ mined on resting post-absorptive animals bred adults were obtained from a colony in held in the metabolic chamber for more Canberra. J1he original stock for this colony than I hour at a selected Ta. The minimum came from a number of sub-alpine local­ metabolic rate for a Ta was calculated ities within the Kosciusko National Park, from the lowest value of the fractional N.S.W. A further three adult female B. difference lasting for at least 5 min. All parvus were live-trapped at Mt Hotham, measurements were made between 0800 Victoria. One female had four pouch young and 1900 h. vo2 was calculated from the when captured and these were successfully equations of Withers (1977) with all gas reared. The animals were housed as pairs volumes corrected to STP. or individuals in steel mesh cages (I m) in a constant temperature room. The female METABOLIC MEASUREMENTS OF with pouch young was housed in a large TORPID ANIMALS cage (1.5 x 2 x 1 m) in the same constant temperature room. All cages contained The initial study on torpor was made in nestbox-metabolic chambers fitted with the Department of Zoology, Australian thermocouples for the continuous measure­ National University, Canberra in 1976. ment of fur surface temperature (Godfrey Three captive-bred adults which had been 1968) . The B. parvus were maintained on maintained in a large outdoor cage were a diet of raw peanuts, sunflower seed, and individually housed in a 2 x I x I m cabinet commercial baby food with added vitamin made from 50 mm thick polystyrene foam. powder. A calcium supplement was added The cabinet was placed in an isoiated room to the drinking water which was available and a constant temperature of 5-8 °C was ad libitum. maintained within the cabinet by the daily renewal of tubs of crushed ice placed above Oxygen consumption rate ('V02 , irrcli­ the animal cages. A little light penetrated cated hereafter by symbol V02) was from an external light source set at 14 measured continuously using an open-circuit hours dark, 10 light. system connected to a paramagnetic oxygen analyser (see Fleming 1980 for details). Animals were supplied with a nestbox­ T,he oxygen consumption of indivi­ metabolic chamber made from a hemis­ duals and groups was determined by the pherical lidded plastic bowl (diameter = following procedure. Animals were weighed 100 mm) with a perspex entrance tube to the nearest 0. I g and placed in an air­ attached (30 x 50 mm). Torpor was detect­ tight perspex metabolic chamber equipped ed by continuously monitoring temperature with inlet and outlet ports and a wire mesh within the nest (Godfrey 1968). Oxygen platform. Single animals were measured in use during torpor was measured by draw­ a 0.5 I metabolic chamber with a flowrate ing air through a small hole in the Ed, over of about 300 ml air min-1 and groups of H1e animal and out through a rubber bung, four animals were measured in a 1.2 1 which was used to seal the entrance tube. chamber with a flowrate of about 900 ml Jlhe air was dried and sampled for oxygen air min·1 . content every two minutes with a Beckman FLEMING: THERMAL PHYSIOLOGY OF BURRAMYS PARVUS 81 (.) 0 41... • :I • ~ 41 c. • • E • ~ .... • • • .. .... t•• ,> .. •.. .. • .. • .... •t 0 .... • 0:1 .. .. • .. • • 'I J: ~ 'j' • Ol rS E d.j ·> .. 1 .. .. ..~·.... .... 0 15 20 25 30 35 Ambient Temperature ( °C) Fig. 1. The relationship of body temperature and oxygen consumption rate to ambient temperature for individual Burramys parvus. The solid line in the lower part is the regression equation VO, = 4.24-0.123 T •. 82 AUSTRALIAN MAMMALOGY E2 paramagnetic oxygen analyser. To chamber. Respiration rate was measu reel measure the very low rates of oxygen use by a pressure transducer connected to the during torpor, the nestbox-metabolic cham­ metabolic chamber and the output was dis­ ber was sealed for up to l hour, then un­ played on a Grass polygraph. sealed and oxygen use recorded until the value prior to sealing was reached. The STATISTICAL AN A LYSIS area under the curve was integrated over All averaged data is presented as Mean time to measure total oxygen use. ± Standard Deviation (n = number of The induction of torpor in B. parvus was measu rements, N = number of indivi­ attempted at the Department of Zoology, duals). Areas under oxygen consumption Monash University, under three different curves were determined by finding the area conditions. For the first winter ( 1978), five of an irregular polygon. Linear regressions animals were housed in a constant tempera­ were fitted by the least squares method and ture room set at 7 ± 2 oc with the light inflexion points were determined by a pro­ regime identical to natural photo-period. gramme called TURNPOINT (Fleming Torpor was monitored from outside the 1985) . constant temperature room by continuous­ ly recording nestbox temperature. The constant temperature room also housed RESULTS other species being used for other experi­ mental work. For the second winter, four N::>RMOTHERM IC: INDIVIDUALS animals were kept in the same constant Burramys parvus strictly regulate their temperature room in 0.30 x 0.45 x 0.15 m body temperatures at ambient temperatures tubs with wire mesh tops and nestbox­ (T,,) between 2°C and 28 °C (Fig. l). With­ metabolic chambers. The tubs were placed in this range T" is 36.1 ± 0.53 oc (n = within a cabinet (2 x l x I m) made from 28, N = 6). A pronounced inflexion in 5 em thick polystyrene foam to reduce the body temperature curve occurs at Ta disturbance by noise. For the third winter, = 28.0 oc with T" increasing at a rate of the polystyrene cabinet was placed in an 0.68 oc per degree Ta.
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