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Hibernation and Daily Torpor in Two Pygmy Possums (Cercartetus Spp., Marsupialia) Author(S): Fritz Geiser Reviewed Work(S): Source: Physiological Zoology, Vol

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Hibernation and Daily Torpor in Two Pygmy Possums (Cercartetus Spp., Marsupialia) Author(S): Fritz Geiser Reviewed Work(S): Source: Physiological Zoology, Vol Hibernation and Daily Torpor in Two Pygmy Possums (Cercartetus Spp., Marsupialia) Author(s): Fritz Geiser Reviewed work(s): Source: Physiological Zoology, Vol. 60, No. 1 (Jan. - Feb., 1987), pp. 93-102 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/30158631 . Accessed: 16/04/2012 20:11 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to Physiological Zoology. http://www.jstor.org HIBERNATION AND DAILY TORPOR IN TWO PYGMY POSSUMS (CERCARTETUS SPP., MARSUPIALIA)' FRITZ GEISER2 Schoolof BiologicalSciences, Flinders University, Adelaide, South Australia 5042, Australia (Accepted6/24/86) The physiologyof torporwas investigatedin the pygmypossums Cercartetus con- cinnus(18 g) and C. lepidus(12 g). Body temperatures(Tb) during torpor remained within 1 C of air temperature(Ta) and showed a minimum of about 5 C in both species.Oxygen consumption (Vo2) duringtorpor was reducedto about 0.05 liters 02/kg h, which amountsto only 1%of the rate of normothermicanimals. Below Ta of 5 C, the metabolismduring torpor increased with a furtherdecrease in Ta,while Tbwas regulatedat about 5 C. Ratesof arousalwere fasterthan predictedfor endo- thermicvertebrates of comparablesize. Durationof torporbouts rangedfrom less thana day to a week,with the longestbouts occurring at Ta< 10 C. The physiological characteristicsof torporin these two marsupialspecies are qualitativelyand quanti- tativelysimilar to those of placentalhibernators. INTRODUCTION nounced reduction of body temperature Pygmy possums, small (<50-g) marsu- and mass-specific oxygen consumption pials of the family Burramyidae, are found during torpor (Bartholomew and Hudson in Australiaand New Guinea. Seven species 1962), a shorter duration of torpor (Hick- are recognized, with Distoechuruspennatus man and Hickman 1960; Strahan 1983), occurring only in New Guinea. The taxo- and a slower rewarming from torpor (Bar- nomic membership of the two feathertail tholomew and Hudson 1962; Fleming possums (Acrobates pygmaeus and Disto- 1985a, 1985b) than in placental mammals. echurus pennatus) to burramyids is cur- Furthermore, there are no experimental rently under review (Archer 1984), and the data on the thermoregulatory ability of remaining species belong to the genera pygmy possums during torpor and thus no Cercartetus and Burramys. indicationthat the reductionin Tbis a con- Torpor has been observed in all Austra- trolled process. Some of these quantitative lian burramyids and is qualitatively similar differencesled to the conclusion that torpor to hibernation in placental mammals (Bar- in burramyid marsupials may be less "re- tholomew and Hudson 1962). It has been fined" than in the "perfect" model of hi- suggested, however, that quantitative dif- bernating rodents (Lyman 1982). ferences do exist between the two groups. Physiological data are available for three Such differences appear to be a less pro- of the six Australian burramyids (Bartho- lomew and Hudson 1962; Fleming 1985a, 'I wishto thankMeredith Smith for herloan of 1985b). Because large interspecific differ- severalpygmy possums and John Coventry for allowing ences in the pattern of thermoregulation me to use hispitfall traps in the BigDesert and partake have been observed in rodent families that of his dinkumbush chooks. Bronwyn McAllan helped contain hibernators and me to set up and checktrap lines. RussellBaudinette may "deep" spe- providedmany helpfulsuggestions during the exper- cies that enter shallow, daily torpor (Lyman imentalperiod, and SaraHiebert, Jim Kenagy,and 1982), it was of interest to investigate two anonymousreferees constructively criticized the whether such specific differencesin the manuscript.The study was supportedby a Flinders thermalphysiology also occur in the bur- UniversityResearch Scholarship and the paperwas compiled while the author held an Alexandervon ramyids.I also examinedin greaterdetail Humboldt-LynenResearch Fellowship at the Uni- the suggesteddifferences in the physiology versityof Washington. between placental hibernators and the 2 Present address: Department of Zoology, Univer- marsupialfamily Burramyidae by studying sity of Washington, Seattle, Washington 98195. temporaland thermalaspects of torporin Cercartetus the western Physiol. Zool. 60(1):93-102. 1987. concinnus, pygmy - 1987 by The University of Chicago. All possum, and C. lepidus (syn. Eudromicia rightsreserved. 0031-935X/87/6001-8636$02.00 lepida), the little pygmy possum. 93 94 F. GEISER MATERIALAND METHODS certain nights. Food was exchanged daily in the afternoon. THE ANIMALS Cercartetusconcinnus is found in South EXPERIMENTALDETAILS Australiaand WesternAustralia and occurs Diurnal fluctuations in oxygen con- in mallee heath and dry sclerophyll forest. sumption (V02) were determined at Ta 6- It is nocturnal, uses tree hollows as daytime 31 C for 30 periods of 21.1 - 2.1 h (range shelters,eats insects, pollen, and nectar, and 15.5-29.25 h, C. concinnus) and for 20 pe- storesfat in its prehensiletail (Smith 1983). riods of 22.0 - 2.3 h (range 17.25-25.75 h, Three males and two females were live C. lepidus). These experiments began in the trapped at various locations on the south- late afternoon. Food and water were not eastern coast of South Australia and the available during measurements of Vo2. To Eyre Peninsula and kept in the laboratory determine whether burramyids regulate at Ta 22 C under natural photoperiod. In their Tbduring periods of torpor at a specific autumn (March) they were transferredto a "set point," torpid animals were cooled constant Ta of 19 C with a photoperiod of (<0.05 C/min), and the Vo2 was continu- 12L:12D, a condition similar to the natural ously monitored. Additional measurements photoperiod at that time of the year, and during shorter periods were used to deter- held for 3 wk before the experiments com- mine the V02 Of postabsorptive,normo- menced. All experiments were performed thermic, inactive animals at T, around and in autumn between March and May 1984 above thermoneutrality. The Vo2 of one C. and 1985. The mean body mass during the lepidus was measured during prolonged experimental period was 18.6 - 1.8 g (SD) torpor over a 7-day period. Flow rates of (range 16-22 g). air in the open flow system were between Cercartetus lepidus occurs in Tasmania 0.15 and 0.30 liters/min and were adjusted and two areas of southeastern Australia. It and measured with calibrated rotameters. is the smallestof all possums.It is mainly Measurements of V02 were used to deter- nocturnal and has been found sheltering in mine the metabolic rate of normothermic, hollow logs and birds' nests (Green 1983). inactive animals measuredwhen a variation It appearsto feed primarilyon invertebrates of less than 5%over at least 15 min occurred and stores fat in its tail. One male and one within an inactivity period of at least 30 female were live trapped in the Big Desert, min; the metabolic rate of torpid animals Victoria, in September and transferred to was determined at times of constant Vo2 the laboratory, where they were kept at Ta over at least 30 min. Furthermore, the du- 19 C and a photoperiod of 12L:12D (similar ration of torpor and the time that was re- to natural photoperiod at the time of cap- quired to arouse from torpor to normo- ture) for 5 wk before measurements com- thermia were determined from these mea- menced. Experiments with these two in- surements (the Vo2 peak during arousalwas dividuals were conducted in spring and assumed to be the end point of the arousal early summer (October to beginning of De- period). In some instances the displacement cember) 1984. Another male was caught at of sawdust from the animal's back was used Messent Conservation Park, South Aus- to determine the duration of torpor. All tralia, and was L.eptin the laboratory at Ta measurements were conducted in a quiet 22 C and natural photoperiod. In autumn controlled-temperature room (Ta + 0.5 C) (March) it was transferred to Ta 19 C and that was acoustically insulated from the re- a photoperiod of 12L:12D and was kept cording equipment. A video camera was under these conditions for 3 wk before the used to observe the animals during the ex- experiments began. The mean body mass periments. was 12.6 - 1.9 g (range 10-15 g). A Servomex Model OA 184 paramag- The animals were fed with Heinz canned netic oxygen analyzer was used for the Vo2 baby food (fruits), a mixture of Heinz high- measurements together with a Rikadenki protein cereal and honey, and apples. Be- potentiometric recorder. The Vo2 was de- cause pygmy possums held in captivity termined from the difference between the become extremely fat, their weight was oxygen content in two parallel open-flow controlled by feeding them only apples on circuits, one being a room air reference and TORPOR IN PYGMY POSSUMS 95 the other containing the animal. All gas tendency to enter torpor spontaneously volumes were correctedto dry standard than the largerC. concinnus. temperature and pressure (STPD).The Ta Most entriesinto torporin both species was measuredcontinuously in the respi- occurredbetween 2400 and 0600 hours, ratorychamber (volume 3 liters)with cal- beforethe onset of light. The time of day ibratedthermocouples. The Tbwas mea- when torpor commenced was not related sured with calibrated 0.5-mm diameter to Ta. The nocturnalpeaks of Vo2 during thermocouplesinserted 20 mm into the activityof C. concinnusoccurred between rectum. Esophageal temperatureduring 1845 and 0130 hours,with a mean of 21.1 torpor was also determinedoccasionally. + 2.1 h, anddid not appearto be dependent During induced arousal the rate of re- on Ta.
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