半散养东北虎繁殖行为学研究
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摘要
2006年6月~2007年11月,采用全事件取样法、目标动物取样法和行为取样法对泰山东北虎园7只(3雌,4雄)半散养东北虎(Panthera tigris altaica)繁殖期各主要繁殖行为及昼间活动节律进行了研究,同时利用放射免疫分析法(RIA)对东北虎粪便中性激素水平的季节性变化进行了研究,并分析了性激素水平与繁殖行为的相关性,结果显示:
     1在时间分配方面,非繁殖期雌雄虎都将最多的时间用于休息,其次是运动,取食和其他行为时间很少;两者摄食时间差异显著(P<0.05),其他不明显(P>0.05)。雄虎发情交配期休息和运动时间最多,其次是发情交配和其他;与非繁殖期比休息明显减少(P<0.05),其他行为增加非常显著(P<0.01)。雌虎发情交配期运动和休息时间最多,其次是发情交配和其他;与非繁殖期比各种行为变化不明显(P>0.05)。雌雄虎发情交配期的各种行为时间分配差异不显著(P>0.05)。哺乳期雌虎仍然是休息所占比重最大,其次是哺乳,运动时间明显减少。雌虎在非繁殖期、发情交配期和哺乳期,休息、运动和取食时间差异极显著(P<0.01),其他行为差异不显著(P>0.05)。
     在活动规律方面,雄虎在非繁殖期卧息主要发生在11:00~13:00,睡眠多发生在13:00~15:00,运动在9:00~11:00和15:00后,其他行为分布比较均匀;雌虎非繁殖期卧息主要发生在9:00~11:00和15:00后,睡眠集中在11:00~15:00,运动也主要发生在9:00~11:00和15:00后,其他行为分布比较均匀。发情交配期雌雄虎活动规律相似,休息主要发生在11:00~14:00,睡眠多发生在12:00~13:00,运动在9:00~10:00和15:00左右为高峰期,同时发情交配行为大量增加,嗅闻、打滚、露阴和频尿时有发生。哺乳期变化较大,在室内期母虎的休息时间在一天中几乎不间断,主要集中在9:00~10:00,11:00~17:00,哺乳与休息具有同步性,运动时间主要集中在11:00~12:00和16:00~16:30;室外期休息主要集中在9:00~14:00和15:00~16:00,哺乳主要集中在9:00~13:00,运动时间比较分散,没有明显的高峰期。由于采取定时定点饲喂模式,所以雌雄虎的摄食一般只发生在17:00~18:00。
     2交配过程中,雌雄虎都存在邀配行为,但雌虎是交配行为的积极发动者,具有明显的邀配行为和特定的邀配模式。有时雌虎虽然主动触闻雄虎,也抬尾露阴但并非邀配,观察发现此时如果雄虎靠近并试图爬跨常被拒绝。
     3整个交配期,每只雌虎平均日邀配28±3次,平均邀配持续时间为20.0±1.2s。每只雄虎平均日爬跨14±1次。平均抽动持续时间为20.9±0.5s,平均交配持续时间为45.2±1.3s。
     4东北虎在交配过程中无锁结现象,雄虎通常在一次爬跨多次抽动后即出现射精。在1h内出现2次射精次数的比例占总射精次数的74.0%。东北虎的交配模式属于Dewsbury分类系统中的第11种,即无锁结、有抽动、单次插入、多次射精。
     5在分娩过程中,雌虎的梳理、卧息和舔仔行为时间最长,分别为26.90%、26.23%和23.83%;梳理、努责和舔仔行为发生频率较高,分别是37.95%、18.75%和12.50%。
     6在育幼过程中,雌虎室外期与室内期哺乳和亲仔行为的时间差异极显著(P<0.01),离开行为的时间差异显著(P<0.05)。室内期和室外期哺乳频次分别为5.75±0.36%和3.75±0.19%;室内期每次哺乳平均时间为1919.00±93.60秒,室外期885.26±38.04秒。两个时期的哺乳频次(t=4.270,P<0.01)和每次哺乳的平均时间(t=8.683,P<0.01)差异也极显著。
     7雌虎粪样中E_2的质量分数在发情交配期一直维持高水平,基本都在400 pg/g以上,峰值达1500 pg/g;育幼期水平较低,基本都在400 pg/g以下;非繁殖期含量较低,与发情前相当,一般在50~200pg/g。非繁殖期、发情交配期和育幼期雌二醇浓度差异极显著(F_(62.2)=32.513,P<0.01)。
     8雌虎孕酮的质量分数在非繁殖期、发情交配期和育幼期基本都在100 ng/g以内,育幼期稍高,峰值达400ng/g,三个时期孕酮的质量分数差异显著(P<0.05)。
     9雄虎粪样中T的质量分数在发情交配期维持了较高的水平,一般不低于200ng/g,峰值达550ng/g以上。非繁殖期T的质量分数很低,在200ng/g以下。T浓度在非繁殖期和发情交配期的差异达到极显著水平(P<0.01)。
From June 2006 to November 2007 at Taishan Amur Tiger Park,with all-occurrence recording,focal-animal sampling and behavior sampling methods,we chose 7 semi-free-ranging Arnur tigers(4 males,3 females)to investigate activity rhythms and reproductive behaviors during breeding stages and anestrous seasons,and mensurated fecal gonadal hormone concentrations in Amur tiger during different periods with radioimmunoassay method. At last,we analyse the relations between reproductive behaviors and gonadal hormone concentrations in Amur tiger.The main results are as follows:
     1 As to the time budget,male and female Amur tiger spent most time in resting,more time in moving,and less time in eating and other behaviors;and eating time(P<0.05)was significant different and time in other behaviors was similar between male and female Amur tiger,during anestrous season.During rutting period,male Amur tiger spent most time in resting and moving,more time in rutting,copulating and other behaviors;and spent less resting time(P<0.05)and more other behaviors time(P<0.01)than those in anestrous season. During rutting period,female Amur tiger spent most time in moving and resting,more time in rutting,copulating and other behaviors;and the time budget of all behaviors during rutting period and anestrous season are similar(P>0.05).The time budget of male and female Amur tiger were similar during rutting periods(P>0.05).During lactation female Amur tiger spent most time in resting,more time in nursing,and less time in moving.Resting time,moving time and eating time were all significant different(P<0.01)and time in other behaviors was similar of female Amur tiger among anestrous season,rutting period and lactation.
     For the activity rhythm,the peak of resting was recorded at 11:00-13:00;sleeping at 13:00-15:00,moving at 11:00-13:00 and behind 15:00;and other behaviors had no peaks of male Amur tiger during anestrous season.The peak of resting was recorded at 9:00-11:00 and behind 15:00;sleeping at 11:00-15:00,moving at 9:00-11:00 and behind 15:00;and other behaviors had no peaks of female Amur tiger during anestrous season. During rutting periods,activity rhythms of male and female Amur tiger were similar,and the peak of resting was recorded at 11:00-14:00;sleeping at 12:00-13:00,moving at 9:00-10:00 and behind 15:00;and many rutting and copulating behaviors which included sniffing,wallow,chinesisch and frequent urinating were recorded.Lactation changed obviously. Female tiger almost always kept rest in day before it went out.The peak of resting was recordedat9:00-10:00 and 11:00-17:00;moving at 11:00-12:00 and 16:00-16:30; and activity rhythms of resting and lactation were similar.After female tiger went out,it rested mainly 9:00-14:00 and 15:00-16:00;lactation at 9:00-13:00.Female tiger had no obvious moving peak.Eating behaviors of Amur tiger only occurred from 17:00 to 18:00 because the fixed breeding pattern.
     2 Male and female Amur tiger had sexual solicitation,and mating activities were often initiated by female tiger which had obvious sexual solicitation behaviors and given patterns. Sometimes,although female tiger sniffed the male forwardly and had tail erection,it didn't want to mate.And male was generally refused when it tried to mount at this time.
     3 During the whole mating period,every female had 28±3 sexual solicitations per day averagely with the average lasting time of 20.0±1.2 seconds,and every male mounted 14±1 times each day averagely.The average thrusting duration was 20.9±0.5 seconds,and the average mating duration was 45.2±1.3 seconds.
     4 There was no copulatory lock during mating,and the male usually ejaculated following a single intromission after several thrusts.The percentage of ejaculations happening again within an hour was about 74.0%.Amur tigers had a copulatory pattem with no lock,thrusting, multiple intromission,and single ejaculation,which belonged to copulatory pattern No.11 in Dewsbury's copulatory-patterned system.
     5 The behaviors that accounted for most of the total time budget during the parturition period were grooming,lying and licking kittens,26.90%,26.23%and 23.83%,respectively. The behaviors that occurred most frequently were grooming(37.95%),abdominal contraction(18.75%),and licking kittens(12.50%).
     6 During nursiing period female Amur tiger lactationing time and nursing time were all significant different(P<0.01)from indoor and outdoor period,went out time was significant (P<0.05).Lactating frequence were 5.75±0.36%and 3.75±0.19%in indoor and outdoor period.Lactating time were 1919.00±93.60s and 885.26±38.04s in two defferent period. Lactating frequence and lactating time were all significant different(P<0.01).
     7 Fecal E_2 concentration on the tigers kept the higher lever during rutting period.Fecal E_2 concentration was usually more than 400pg/g and the peak of it reached over more than 1200pg/g.Fecal E_2 concentration was usually less than 400pg/g during nurseing period.Fecal E_2 concentration kept usually low levers during anestrous season,and was similar with the fecal E_2 concentrations of the time before rutting period.There were very obvious difference of E_2 concentration on female tiger among rutting period,nurseing period and anestrous season.
     8 Fecal P concentrations was usually less than 100ng/g during anestrous season,rutting period and nurseing period,and the concentration during nurseing period kept higher level than those during anestrous season and rutting period.The peak of P concentration during nurseing period was more than 400ng/g.There were obvious difference of P concentration among anestrous season,rutting period and nurseing period.
     9 Fecal T concentrations of male Amur tiger kept the higher lever and were usually more than 200ng/g during rutting period.The peak of T concentrations reached over more than 550ng/g.T concentrations on males were very low and not more than 200ng/g during anestrous season.There were significant differences of T concentrations on males between rutting period and anestrous season.
引文
[1]Abramov,V K.Strategy for Conservation of the Amur Tiger in Russia.Tigers of the World,1999
    [2]Alexander G M.Sherwin B B.Sex Steroid.Sexual Behavior and SelectiveAttention for Erotic Stimuli in Woman Using Oral Contraceptive.Psychoneuroendocrinology,1993,8-(2):91
    [3]Altmann,J.Observational Study of Behavior:Sampling Methods.Behavior,1974,(49):227~267
    [4]A M Amirkhanov(Chief editor).Biodiversity Conservation in Russian.National Report,20 February,1998
    [5]Anderson J L.There establishmentnadmanagementofalion(Pnahteraelo)Populationin Zululnad,SouthArfiea.Biol.Conseyr,1981,19:107~117
    [6]Anon.Tiger Action Plan for the Kingdom of NePal.Unpubl.report to Year of theTiger Conference,Dallas,US.1997
    [7]Bardi M,Shimizu K,Fujita S,et al.Hormonal correlates of maternal style in captive macaques(Macacafuscata and M.mulatto).Am J Primatol,2001,22:647~662
    [8]Beach,F A.Sexual Attractivity,Proceptivity and Rreceptivity in Ffemale Mmammais.Horm.Behav,1976,7:105~138
    [9]Bercovitz A P.Czecala NM and Lasley BL.A new Method of Sex Determination in Monomorphicbirds.J zoo Anim Med,1978,9:114~124
    [10]Berger,J.Facilitation of Reproductive Synchrony by Gestation Adjustment in Gregarious Mammals:a New Hypothesis.Ecology,1992,73:323~329
    [11]Billing H L,Frusta and A J W.Hush Estrogens in Habit and Androgens Enhancevariant Granulosa Cell Apoptosis Endoerillology,1993,133(5):204~212
    [12]Boinski,S.Birth Synchrony in Squirrel Monkeys Saimiri Oerstedi.Behav.Ecol.Sociobiol,l 987,21:393~400
    [13]Bronson,F H.Mammalian Reproduction.Chicago:University of Chicago Press,1989
    [14]Brown,R E.The Rodents I:Effects of Odours on Reproductive Physiology.In:Social Odoursin Mammals,Vol.1(Ed.by R.E.Brown and D.W.MacDonald),pp.Oxford:Clarendon Press,1985,245~344.
    [15]Carlos.Spatial Viability Analysis of Amur Tiger in the Russian Far East.Journal of Applied Ecology;2006,43(6),1056~1068
    [16]Carosi M,Heistermann M,Visalberghi E.Display of Proceptive Behaviors in Relation to Urinary and Fecal Progestin Levels over the Ovarian Cycle in Female Tufted Capuchin Monkeys.Horm Behav,1999,36:252~265
    [17]Clutton-Brock,T H.Mammalian Mating Systems.Proc.R.Soc.Lond.B.1989,23(60):339~372
    [18]Cockerm J E,Rounce J R.Faecal Measurements of Oestradiol and Testosterone Allow the Non-invasive Estimation of Plasma Steroid Concentrations in the Domestic Fowl.Britpoultry Science,1994,35:433~443
    [19]Dewsbury,D A.Patterns Copulatory Behavior in Male Mammals.Quarterly Review of Biology,1972,7(1):1~33
    [20]Dixson A F.Primate Sexuality.Oxford:Oxford University Press,1998
    [21]Dong-Sun WON,et al.A Case of Nutritional Secondary Hyperparathyroidism in a Siberian Tiger Cub.J.Vet.Med.Sci.2004,66(5):551~553
    [22]Doty R L(ed):“Mammalian Olfaction,Reproductive Processes and Behavior.”New York:Academic Press
    [23]Evans K L,Hughes J P and Couto M et al.Pregnancy diagnosis in the Domestic Horse through direct Urinary Estrone Conjugate Analysis.Theri-ogenology,1984,22:615-620
    [24]Ganswindt A,Palme R,Heistermann M,et al.Non-invasive Assessment of Adrenocortical Function in the Male African Elephant(Loxodonta africana)and its Relation to Musth.General and Comparative Endocrinology,2003,134,156~166
    [25]Gesquiere.et al.Mechanisms of Sexual Selection:Sexual Swellings and Estrogen Concentrations as Fertility Indicators and Cues for Male Consort Decisions in Wild Baboons.Hormones and Behavior,2007(51):114~ 125
    [26]Good T,Khan M Z,Lynch,J W.Biochemical and Physiological Validation of a Corticosteroid Radioimmunoassay for Plasma and Fecal Samples in Oeld Mice(Peromyscus polionotus).Physiologyand Behavior,2003,80,405~411
    [27]Gore-Langton R E and D T.Arm Strong Folliclar Steroidogensis and its Control In:Knobil E.and TD.Neill ed.The Pysiology of Reproduction Newyork:Raven Press.1994
    [28]Goymann W,Mostl E,Gwinner E.Corticosterone Metabolites can be Measured Noninvasively in Excreta of European Stonechats(Saxicola torquata rubicola).Auk,2002,119:1167~ 1173
    [29]Goymann W,Mostl E,Vant Hof,et al.Noninvasive Fecal Monitoring of Glucocorticoids in Spotted Hyenas(Cwcuta crocuta).General and Comparative Endocrinology,1999,114:340~348
    [30]Hafez E S E.TheBehavior of Domestic Animals.Bailliere,Tindalland Cox.1962
    [31]Hall M J.Mother-offspringRelationships in Red Reer(Cervus elaphus L.)and theSocial Organization of an Enclosed Group.PhD.Dissertation,University of Sussex.1978
    [32]Haufe C.Sexual Selection and Mate Choice in Evolutionary Psychology,Biol Philos,2007
    [33]Hayward G D,Miquelle D G,Smirnov E N,et al.Monitoring Amur tiger Populations:Characteristics of Track Surveys in Snow.Wildlife Society Bulletin?2002,30(4):1150~1159
    [34]Heistermann M,Tari S and Hodges J K.Measurement of Faecal Steroids for Monitoring Ovarian Function in New world Primates,Callitrichidae.Journals of Reproduction and Fertility,l993,99:243-251
    [35]Hindle J E,Mostl E and Hodges J K.Mea-surement of urinary oestrogens and 20 α-Dihydroprogesterone during Ovarian Cycles of Black(Diceros bicornis)and White(cerratotherium simum)Rhinoceroses.J Reprod Fert.l992,94:237-249
    [36]Hunt K E,Trites A W,Wasser S K.Validation of a Fecal Glucocorticoid Assay for Stellar Sea Lions(Eumetopias jubatus).Physiology and Behavior,2004,80:595-601
    [37]Ims R A.The Ecology and Evolution of Reproductive Synchrony.Trends Ecol,1990,5,135~140
    [38]John M,Good Rich and Dale G,Miquelle D G.Short Communication Translocation of Problem Amur Tigers to Alleviate Tiger-human Conflicts.FFI,Oryx,2005,39(4):l ~4
    [39]Kerley L L,Goodrich J M,Miquelle D G,et al.Effects of Roads and Human Disturbance on Amur Tigers.Conservation Biology,2002,16(1):97~ 108
    [40]Kerley L L,Goodrich J M,Miquelle D G,et al.Reproductive Parameters of Wild Females Amur(siberian)Tigers(P anther a tigris altaica).Journal of Mammalogy,2003,84(1):288~298
    [41]Kindahl M,Knudsen O and Madej A,et al.Progesterone,prostaglandin F2 a,PMSG and Oestrone Sulphate during Early Pregnancy in the Mare.J Reprod Fert.1982,32:353~ 359
    [42]Kirkpatrick J F,Shideler S E and Lasley B L,et al.Pregnancy Determination in Uncaptured Feralhorses by Means of Fecal Steroid Conjugates.Theriogenology.1991,35(4):753~760
    [43]Klopfer,P H.Origins of Parental Care.In:Gubernick D J,Klopfer P H,editors.Parental Care in Mammals New York:Plenum Press.1981,1 ~12
    [44]Lasley B L,Kirkpatrick J F.Monitoring Ovarian Function in Captive and Free-ranging wildlife by Means of Urinary and Fecal Steroids.Zoology Wildlife,1991,22:23 ~31
    [45]Leigh Elizabeth Pitsko.Wild Tigers in Captivity:A Study of the Effects of the Captive Environment on Tiger Behavior.Master of Science in Geography,2003
    [46]Levy F R,Cervais V,Kindermann M,et al.Effects of Early Post-partum Separation on Maintenance of Maternal Responsiveness and Selectivity in Parturient Ewes.Applied Animal Behaviour Science,1999,63:3 ~10
    [47]Li C W,Jiang Z G,Jiang G H and Fang J M.Seasonal Changes of Reproductive Behavior and Fecal Steroid Concentrations in Pere David's Deer.Horm.Behav.2001,40:518-525
    [48]Linda L,John M,Dale G,et al.Reproductive Parameters of Wild Female Amur(Siberian)Tigers(Panthera tigris altaica).Journal of Mammalogy,2003,84:288~298
    [49]Liu Y X and A J W.HushSynergism between granulose and theca inierstitial cell-s in estrogen biosynthesis by n-treated rat ovaries:Studies on the Two Ceel:Tow Genadotrophin Hypothesis Using Steroid Antlsera Biology of Reproduction,1986,35:27~36
    [50]Lonnie H K,John P H and George H S,et al.Estrone Sulfate Concentrations as an Indicator of Fetal Demise in Horses.Am J Vet Res,1988,49(2):184~187
    [51]Lovari S and Locati M.Temporal Relationships,Transitions and Structure of the Behavioural Repertoire in Male Apennine Chamois during the Rut.Behaviour.1991,119:77~103
    [52]Marcot B G.Tiger Habitat Corridors in Far East Russia,Northeast China,and Northern North Kerea:Need for a Conservation Strategy.1995,1,20
    [53]Mccomb K E.Female Choice for High Roaring Rate in Red Deer,Cervus Elaphus.Anim.Behav.1991,41:99~88
    [54]Millspaugh J J,Washburn B E,Milanick M A,et al.The Relation between Urinary Cortisol Levels and Social Behavior in Captive Timber Wolves.Can.J.Zool.,1996,74:209~216
    [55]Miquelle Dale.The Use of Radio Telemetry in the Study of the Amur Tiger.Bull.Mosk.Obschestra Lspytateley Prirody.Otd.Biol.,1993,98(3):63~79
    [56]Monfort S L,Dahl K D and Czekala N M,et al.Monitoring Ovarian Function and Pregnancy in the Giant Panda(Ailuropoda melanoleuca)by Evaluating Urinary Bioactive FSH and Steroid Metabolites.J Reprod Fert.1989,85:203~212
    [57]Monfort S L,Hess D L,Shideler S E,et al.Comparison of Serum Estradiol to Urinary Estrone Conjugates in the Rhesus Macacure.Biol.Reprod.1987,37:832~837
    [58]Moore C L,Jordan L and Wong L.Early Factory Experience,Novlety,and Choice of Sexual Parter by Male Rats.Physiology and Behavior.1996,60(5):1361 ~1367
    [59]Murray M G.The Rut of the Impala:Aspects of Seasonal Mating under Tropical Conditions.Z.Tierpsychol.1982,59:319~337
    [60]Nowell K,Jackson P.Wild Cats:Status Survey and Conservation Action Plan.Gland,Switzerland,1996,54~65
    [61]Orihuela A.Some Factors Affecting the Behavioral Manifestation of Oestrus in Cattles.a Review.Appl.Anim.Behav.Sci.2000,70:1 ~ 16
    [62]Pfeifer S.Flehmen and Dominance among Captive Adult Female Scimitar-horned Oryx (Oryxdammah).J.Mammal.1985,66,160~163
    [63]Poter T A and Wilkinson G S.Birth Synchrony in Greater Spear Nosed Bat(Phyllostomushastatus).J.Zool.Lond.,2001,253:283~390
    [64]Reby D,Hewison M,Izquierdo M and Pepin D.Red Deer(Cervus elaphus)Hinds Discriminate between the Roars of Their Current Harem-holder Stag and Those of Neighbouring Stags.Ethology,2001,107:951~959
    [65]Richmond M and Stehn R.Olfaction and Reproductive Behavior in Microtine rodents.In Robert B.Asian Tigers,Lund University,Sweden,2003
    [66]Ronald L,Tilson.Tiger.American Zoo and Aquarium Association,2000
    [67]Ronald L,Tilson and Ulysses S.Tigers of the World,Noyes Publications,1987
    [68]Rosner W,Kelly W G,Deakins S M,et al.The Binding of Estrogens and Testosterone in Human Plasmas.IN:Heyes R L.Goswitz FA and Murphy BEP.U.S.Atomic Energy Commission.Division of Technical Information.1968
    [69]Russello M A,Gladyshev E,Miquelle D,et al.Potential Genetic Consequences of a Recent Bottleneck in the Amur Tger of the Russian Far East.Conservation Genetics,2004,(5):707~713
    [70]Rutberg A T.Adaptive Hypotheses of Birth Synchrony in Ruminants:an Interspecific Test.Am.Nat.1987,130:692~710
    [71]Sadleir R M.The Ecology of Reproduction in Wild and Domesticated Mammals.LondomMethuen.1969
    [72]Schwarzenberger F,Francke R and Gltenboth R.Concentrations of Faecal Immunoreactive Progestogen Metabolites during the Oestrous Cycle and Pregnancy in the Black Rhinoceros(Diceros bi-cornis michaeli).J Reprod Fert.l993,98:285~291
    [73]Snycler N R F.Limitations of Captive Breeding in Endangered Species Recovery.Conv.Biol,1996,10(4):338~348
    [74]Stocco DM.A StAR Search:Implications in Controlling Steroidogensis Bio.Reprod.1997,56:328~336
    [75]Sugimoto T.Species and Sex Identification from Faecal Samples of Sympatric Carnivores,Amur Leopard and Siberian Tiger,in the Russian Far East,Conservation Genetics,2006,10.1007/s 10592-005-9071-z
    [76]Svare B B.Maternal Aggression in Mammals.In:Gubernick D J,Klopfer P H,editors.Parental Care in Mammals.New York:Plenum Press,1981,179-210
    [77]Terio K A,Citino S B,Brown J L.Fecal Cortisol Metabolite Analysis for Noninvasive Monitoring of Adrenocorticcal Function in the Cheetah(Acxnonyx jubatus).J.Zoo.Wildl.Med.1999,30,484~491
    [78]Thirgood S,Langbein J and Putman R J.Intraspecific Variation in Ungulate Mating Strategies:the Case of the Flexible Fallow Deer.Adv.Study Behav.1999,28:333~359
    [79]Thompson K V.Flehmen and Birth Synchrony among Female Sable Antelope,Hippotragusniger.Anim.Behav.1995,50:475~484
    [80]Tilson R,Traylor-Holzer,et al.The Decline and Impending Extinction of the South China Tiger.:Prospects for the Future.in Tigers of the World.1987,35~38
    [81]Touma C,Palme R,Sachser N.Analyzing Corticosterone Metabolites in Fecal Samples of Mice:a Noninvasive Technique to Monitor Stress Hormones.Horm.Behav.2004,45,10~22
    [82]Touma C,Sachser N,Mostl E,et al.Effects ofSex and Time of Day on Metabolism and Excretion of Corticosterone in Urine and Feces of Mice.Gen.Comp.Endocrinol.2003,130,267~278
    [83]Turner J W,Nemeth R,Rogers C.Measurement of Fecal Glucocorticoids in Parrotifishes to Assess Stress.Gen.Comp.Endocrinol.2003,133,341-352
    [84]Wasser S K,Bevis K,King G,et al.Noninvasive Physiological Measures of Disturbance in the Northern Spotted Owl.Conservation Biology,1997,11:1019-1022
    [85]Wasser S K,Hunt K E,Brown J L,et al.A Generalized Fecal Glucocorticoid Assay for Use in a Diverse Array of Nondomestic Mammalian and Avian Species.General and Comparative Endocrinology,2000,120:260-275
    [86]Wasser S K,Monfort S L and Widt D E.Rapid Extraction of Faecal Steroids for Measuring Reproductive Cyclicity and Early Pregnancy in Freeranging Yellow Baboons(papio cynocephalus cyno-cephalus).J Reprod Fert.1991,92:415-423
    [87]Wasser S K,Monfort S L,Southers J,Wildt D E.Excretion Rates and Metabolites of Estradiol and Progeste-rone in baboon(Papio cynocephalus cynocephalus)faec-es.Journal of Reproduction and Fertility,1994,101:213-220
    [88]Wasser S K,Risler L and Steiner R A.Excreted Steroids in Primate Feces over the Menstrual Cycle and Pregnancy.Biol Reprod.1988,39:862-872
    [89]White P J,Fischer R B and Meunier G F.Female Discrimination of Male Dominance Byurine Odor Cures in Hamsters.Physiology and Behavior,1986,37(2):273-277
    [90]Whitten P L,Brochman D K,Stavisky R C.Recent advances in Noninvasive Techniques to Monitor Horm-one-behavior Interaction.Am J Phys Anthropol,1998,27(Suppl.):1-23
    [91]Wilson E O.Sociobiology:the New Synthesis.Cambridge:Harvard University Press.1975
    [92]Wu Changde,Pang Wanyong,Zhao Deming.Amino Acid Sequence of the Amur Tiger Prion Protein.Virus Research,2006,121(1):93-96
    [93]XiLian HU,et al.Cloning,Characterization and Tissue Specific Expression of Amur Ti-ger (Panthera tigris altaica)IGF-Ⅰ,Biosci.Biotechnol.Biochem,2006,70(8):1846-1854
    [94]Young W C.The Hormones and Mating Behavior.In:Young WC.Sex and Internal Secretions,Vol.Ⅰ.Baltimore:Williams&Wilkins.1961,1173-1239
    [95]Zou Hongfei.Recovery and Development of Siberian Tiger Populations in China.Journal of forestry Research,1997,8(1):36
    [96]Ю·A·达尔曼等.俄罗斯远东地区东北虎现状及其保护.野生动物,2006(2):19-23
    [97]白秀娟.圈养东北虎 ISSR 指纹分析初报.兽类学报,2004,24(1):90-93
    [98]蔡桂全,谢家华.糜鹿发情期主要活动时间分配及行为研究,兽类学报,1988.8(3):166-171
    [99]程会昌等.初生东北虎胃和肠的动脉分布.中国兽医科技,2004,34(6):78-79
    [100]崔卫国.动物园内动物的异常行为.生物学通报,2004,39(2):21-22
    [101]杜启艳,常重杰,牛红星,等.东北虎 Sox3和 Sox4基因的保守区序列分析.河南师范大学学报(自然科学版),2003,31(1):78-81
    [102]杜启艳,常重杰,牛红星,等.东北虎 SOX 基因的检测.动物学报,2001,47(专刊):168-171
    [103]杜卫国,盛和林.林麝哺乳期的时间分配和行为研究.兽类学报,1998,18(1):21-26
    [104]]付振和,王杰,等.东北虎先天性白内障摘除术一例.中国兽医杂志,1981,7(12):29-31
    [105]傅文源,华南虎哺乳期母幼行为的初步研究.福建林业科技,2002,29(3):62-64
    [106]高云芳,高更更,白绪祥,等.雌性川金丝猴尿液中雌二醇与孕酮水平的季节性变化.西北大学学报,2005,35(5):592-596
    [107]高云芳等,川金丝猴尿液中睾酮水平的季节性变化.动物学报,2003,49(3):393-398
    [108]高中信,马建章.世界虎的现状及保护.野生动物,1996,17(1):3-7
    [109]高中信.粪便分析法测定植食动物食性的评价.兽类学报,1991,11(3):186-193
    [110]哈尔滨动物园.东北虎的饲养与繁殖.中国动物园年刊,1978,11-18
    [111]霍军等,初生东北虎主要消化器官的观察.河南畜牧兽医,2003,24(9):5-6
    [112]贾志云,蒋志刚,王祖旺.果子狸繁殖期行为的观察.兽类学报,2000,20(2):108-115
    [113]蒋志刚.糜鹿行为谱及 PAE 编码系统.兽类学报,2000,20(1):1-12
    [114]李宝国,赵大鹏.雌性秦岭金丝猴的多次交配行为.科学通报,2005,50(10):1052-1054
    [115]李春,魏辅文,胡锦矗.雌性小熊猫粪样中雌二醇与孕酮水平的变化与繁殖启动的关系.动物学研究,2005,26(2):147-151
    [116]李春,魏辅文,李明,等.雄性小熊猫粪便中睾酮水平的变化与繁殖周期的关系.兽类学报,2003,23(2):115-119
    [117]李春旺,蒋志刚,房继明,等.麋鹿繁殖行为和粪样激素水平变化的关系.兽类学报,2000,20(2):88-100
    [118]李春旺,蒋志刚,曾岩.雄性糜鹿的吼叫行为,序位等级与成功繁殖.动物学研究,2001,22(6):449-453
    [119]李复东,彭世嫒,叶志勇,等.雌性大熊猫发情期尿中雌二醇与孕酮水平的变化及其与配种的关系.兽类学报,1993,13(3):166-171
    [120]李彤,蒋劲松,吴志刚,等.吉林省东北虎的调查.兽类学报,2001,21(1):1-6
    [121]梁冬莹等.半散放和圈养东北虎群狮弓蛔虫感染调查及综合防治.现代生物医学进展,2007,7(1):28-31
    [122]廖鸣娟,朱睦元,叶丹,等.东北虎促卵泡激素和促黄体激素基因的克隆及其序列分析.动物学报,2002,48(6):783-789
    [123]刘冰许,杨光成.幼东北虎小叶肺炎的病理学观察.云南畜牧兽医,1997,(2):20
    [124]刘丙万,蒋志刚.粪样在野生动物研究中的作用.动物学研究,2002,23(1)::71-76
    [125]刘进辉,苏泽红.果子狸繁殖期的行为观察.经济动物学报,2000,4(1):29-33
    [126]刘进辉,谭理琦,王水莲,等.东北虎胃与小肠组织形态结构的研究.经济动物学报,2004,8(1):22-25
    [127]刘丽.东北虎相残行为初步分析.野生动物,1995,16(1):23-25
    [128]刘树光,肖井贵,杨守庄,等.野生动物园东北虎繁殖行为初步观察.野生动物,2000,21(3):35-37
    [129]刘文良,王玮,刘丹,等.东北虎盆腔巨大囊肿摘除纪实.野生动物,1999,,20(6):28-29
    [130]刘昕晨,赵广英,宋惠东,等.八只东北虎肾中氨基酸的分析.野生动物,1994,15(1):29-32
    [131]刘雪卿,张泽钧,魏辅文,等.圈养小熊猫育幼行为的初步观察.兽类学报,2003,23(4):366-369
    [132]刘玉堂,应璐,徐艳春,等.东北虎雌性生殖系统的组织学.动物学杂志,2002,37(2):38-42
    [133]刘玉堂,应璐,徐艳春,等.东北虎雌性生殖系统解剖.经济动物学报,2001,5(3):40-43
    [134]刘振生,李枫,滕丽微,等.半散放东北虎行为活动时间的分配.动物学研究,2002,23(5):389-393
    [135]刘振生,李枫,滕丽微,等.饲养条件下东北虎交配行为的观察.动物学杂 志,2003,38(1):49-51
    [136]刘振生,马建章,滕丽微,等.笼养东北虎雌雄行为差异的比较.动物学研究,2002,23(4):284-287
    [137]刘振生,滕丽微,李枫,等.笼养和半散放东北虎行为时间分配及活动规律差异的比较.生态学杂志,2004,23(1):20-23
    [138]刘振生等.人工饲养东北虎幼虎的行为时间分配.生态学报,2003,23(8):1548-1553
    [139]陆佩洪,张锡然,刘淼,等.华南虎和东北虎血清同功酶的研究.南京师大学报(自然科学版),1995,18(1):81-84
    [140]马逸清.东北虎分布区的历史变迁.自然资源研究,1983(4):44-48
    [141]马逸清.老虎保护的进展.野生动物,1998,19(1)3-7
    [142]马莺.高效液相色谱/荧光检测法测定东北虎样品中的微量元素硒.分析化学,199826(4):496
    [143]梅全林,刘昕晨,赵广英,等.东北虎寄生虫病综合防治的研究.野生动物,1992,13(1):33-36
    [144]孟惠平.激素对动物行为的影响.吉林师范学院院报,1998,19(5):75-77
    [145]孟秀祥.圈养马麝行为生态研究.中科院动物所博士论文.2002
    [146]牛红星,卜艳珍,余燕,等.东北虎幼体消化系统蛋白水解酶的初步研究.兽类学报,2006,26(1):98-100
    [147]彭世媛,李复东.雌性大熊猫发情期与妊娠期尿液性激素的变化.动物学杂志,1993,28(3):25-28,59
    [148]千云松,金光凌,周爱霞.东北虎一年繁殖两胎的人工哺乳.中国动物园年刊,1982,36-38
    [149]任宝平,川金丝猴繁殖行为学研究.东北师范大学博士论文.2002
    [150]任宝平,夏述忠,李庆芬.圈养雄性川金丝猴交配模式.动物学报,2002,48(5):577-584
    [151]任宝平等.雄性川金丝猴睾酮分泌与其社群环境变化的关系.动物学报,2003,49(3):325-331
    [152]尚玉昌.行为生态学.北京:北京大学出版社,2001
    [153]沈永庆.人工饲养下虎的摄食行为障碍.大自然,1989,(1):35-36
    [154]盛和林,大泰司纪之,陆厚基.中国野生哺乳动物.北京:中国林业出版社,1999:132-133
    [155]盛和林.林麝哺乳期的时间分配和行为研究.兽类学报,1998,18(1):21-26
    [156]宋振洲,邱铁岭.东北虎一年繁殖两胎五仔.野生动物,1981,2(3):37-41
    [157]孙海义等.黑龙江省东北虎野外种群监测研究.林业科技,2005,30(6):33-35
    [158]谭超,刘进辉,王水莲,等.东北虎肾组织形态结构的研究.经济动物学报,2003,7(1):35-37
    [159]滕丽微,李枫,刘振生,等.非笼养雌雄东北虎的行为时间分配和活动节律的比较.兽类学报,2003,23(2):93-97
    [160]滕丽微,李枫,刘振生,等.笼养东北虎春季行为及活动规律观察.东北林业大学学报,2002,30(6):78-80
    [161]滕丽微,刘振生,李枫,等.半圈养条件下东北虎繁殖期的行为时间分配.生态学杂志,2003,22(6):53-56
    [162]滕丽微,宋延龄,刘振生,等.笼养东北虎行为的时间分配.兽类学报,2003,23(1):1-5
    [163]滕丽微.笼养条件下东北虎行为模式的研究.东北林业大学硕士学位论文.2001
    [164]图雅,朱伟云,陆承平,等.东北虎粪细菌区系的16SrRNA 基因序列分析.微生物学报,2005,45(5):671-674
    [165]汪松.中国濒危动物红皮书(兽类).北京:科学出版社,1998
    [166]王建辰,章孝荣.动物生殖调控.合肥:安徽科学技术出版社.1998
    [167]王松等.笼养雌性黑叶猴尿中性腺激素水平变化与性行为等级序位的关系.兽类学报,2006,26(2):136-143
    [168]王威,减著斌,夏树福,等.应用闭路电视监视系统对东北虎产仔行为的初步观察.中国动物园论文集,1990,137-141
    [169]王学高.高原鼠兔交配期及交配行为模式的研究.兽类学报,1990,10(1):60-65
    [170]王振龙.根田鼠的交配行为模式.动物学报,2001,47(专刊):187-189
    [171]王振堂,申亨哲.图门江流域人口压力对东北虎数量的影响.动物学杂志,1996,31(5):31-34
    [172]魏玉春,陈治安,何英.东北虎乳腺癌.动物学杂志,1982,17(6):35-37
    [173]巫芳来等.黔金丝猴血清雌二醇(E2)测定分析.贵州科学,1997,15(2):118-120
    [174]吴平,周开亚,王亚明,等.用 RFLP 和 PCR2RFLP 技术研究东北虎和华南虎线粒体 DNA 多态性.生物多样性,1997,5(3):173-178
    [175]吴宪忠,张明海.黑龙江省境内东北虎数量分布现状.野生动物,1994,17,(1)3-7
    [176]夏成柱.通过病毒分离鉴定和基因检测首次发现虎流感.中国兽医学报,2003.23(2):12-15
    [177]向建洲,刘进辉,刘自逵,等.东北虎肺组织结构的解剖观察.湖南农业大学学报(自然科学版),2002,28(3):227-229
    [178]修云芳,邵良平,王德春,等.小熊猫繁殖周期血清雌二醇和孕酮含量变化.兽类学报,2007,27(2):153-157
    [179]徐殿举,李宁,张忠义,等.闭路电视监视系统在东北虎繁殖上的应用.中国动物园论文集,1990,142-143
    [180]徐艳春.虎微卫星位点多态性及其在圈养种群管理中的应用.东北林业大学博士论文.2001:92-93
    [181]许宪祝,刘昕晨,赵广英.东北虎肝中氨基酸的测定.黑龙江省畜牧兽医,1997,(2):9-10
    [182]阎彩娥,蒋志刚,李春旺,等.雌性川金丝猴的邀配行为与尿液雌二醇水平的关系.动物学报,2003,49(6):736-741
    [183]杨帆,王丽红,贺倩,等.圈养亚洲象尿液中雌二醇和孕酮含量的周期性变化.兽类学报,2007,27(1):74-7 9
    [184]杨光诚等.幼东北虎多发性四肢骨折的诊治与病因探讨.郑州牧专学报,1997,17(3):63-64
    [185]杨继光,于永哲,张一.东北虎再生障碍性贫血一例报道.动物学杂志,1983,18(1)61-62
    [186]杨晓军.笼养川金丝猴哺乳行为的观察.甘肃农业大学学报,1996,季刊(4):334-338
    [187]杨晓军.笼养川金丝猴雌性性行为的研究.甘肃农业大学学报,1998,33(3):228-233
    [188]杨雪冬,赵广英,刘昕晨,等.八只东北虎肾、肝中蛋白质的分析.野生动物,1995,16(2):33-34,43
    [189]杨雪冬,赵广英,杨希彬,等.东北虎脾中蛋白质的测定.黑龙江省畜牧兽医,1994, (12):35-36
    [190]游章强,蒋志刚.普氏原羚的求偶交配行为.动物学报,2005,51(2):187-194
    [191]于洪伟.圈养东北虎幼仔行为研究.东北林业大学硕士学位论文.2006:1-34
    [192]于孝臣,孙宝刚,孙海义,等.黑龙江省东北虎的分布和数量调查.野生动物,2000,21(2):14-16
    [193]于永久、李焕敏、刘大军.东北虎对氛胺酮的耐量观察.动物学杂志,1983,18(3):35-36
    [194]曾国庆,蒋广泰,杨克勤,等.大熊猫生殖生物学研究:Ⅰ.大熊猫发情期血清和尿液中促黄体素、孕酮和17β~雌二醇含量的变化.动物学报,1990,36(1):63-69
    [195]曾国庆,张星,蒋广泰,等.大熊猫生殖生物学研究:Ⅱ.大熊猫妊娠期尿中孕酮和绒毛膜促性腺激素样物质含量的变化.动物学报,1992,38(4):429-433
    [196]张德成,陈思义,吴润培,等.东北虎、云豹、金猫红细胞上 C36受体的测定.兽类学报,1993,(2):56-259
    [197]张冠相等.野生东北虎的解剖.野生动物,1991,,12(3):40-41,15
    [198]张继忠.饲养东北虎的生物学探讨.中国动物园年刊.1983:14-16
    [199]张明海,张常智,刘丹,等.半散放东北虎产仔行为的初步观察.兽类学报,2006,26(3):255-261
    [200]张淑云,王东风.东北虎一些器官指数测定.野生动物,1993,14(6):42-43
    [201]张文平,张志和,沈富军,等.华南虎、东北虎、孟加拉虎的 D-loop 和 ND5序列及其在系统进化分析中的应用.四川动物,2005,24(1):22-26
    [202]张钖然,陈宜峰,朱红阳,等.东北虎和华南虎血象及血液生化指标的测定.兽类学报,1993,13(4):302-303
    [203]张钖然,陈宜峰,朱红阳.东北虎和华南虎染色体比较研究.动物学报,1993,39(3):334-337
    [204]张于光,李迪强,饶力群,等.东北虎微卫星 DNA 遗传标记的筛选及在亲子鉴定中的应用.动物学报,2003,49(1):118-123
    [205]张于光,李迪强,肖启明,等.饲养东北虎的微卫星变异研究.遗传,2004,26(5):620-624
    [206]张振昆.人工哺育东北虎一点滴.中国动物园年刊.1978:21-24
    [207]张志华,林安仲,吴丙新,等.母拂拂生殖周期之卵巢、生殖道与生殖内分泌素变化.台北动物园学报,1996,(8):17-38
    [208]赵广英,刘昕晨,赵淑琴,等.东北虎血液成分的测定与分析.动物学杂志,1995,30(5):38-41
    [209]赵英杰,李文发.完达山东北虎的调查报告.黑龙江八一农垦大学学报,1987(1):101-104
    [210]赵云华,刘永利,曾德生.人工饲养东北虎繁殖行为的观察.野生动物,1992,12(3):54-56
    [211]赵云华,于凯军,曾德生,等.东北虎的人工哺育.野生动物,1991,12(4):27-30
    [212]赵志刚.东北虎应激性疾病的诊疗.中国兽医杂志,2003,39(6)25-26
    [213]郑喜邦.野生动物尿和粪中类固醇激素的研究进展.青海畜牧兽医杂志,1998,28(3):31-34
    [214]仲阳康,裴恩乐,熊成培,等.圈养华南虎交配行为的初步观察.兽类学报,2006,26(3):307-310
    [215]周小平等.圈养大熊猫行为研究及其方法.四川科学出版社,2005
    [216]朱靖.大熊猫发情期叫声及其行为意义.动物学报,1987,33(3):285-292