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A Species Survival Plan _SSP_ for Snow Leopard' Panthera uncia

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A Species Survival Plan _SSP_ for Snow Leopard' Panthera uncia Powered By Docstoc
					Int. Ped. Book of Snow leopards3: BI - 102' 1982




     A SpeciesSurvivalPlan (SSP)for SnowLeopard'
        Pantherauncia: Geneticand Demographic
               Analysisand Management
ThomasJ. Foose
Ph.D., AAZPA ConservationCoordinator, ISIS Office, MinnesotaZoo, Apple Valley, MN
55124,USA




                                                      been initiated primarily as a North American
TheSpecies SurvivalPlan (SSP)                         program, there is no desireor need to be pro-
of the AAZPA                                          vincial. It has merely seemedmore feasibleto
 As affirmed in the World Conservation Stra-          organize programs of the scope proposed by
 tegyof IUCN, captive populations and propa-          the SSPon a continentalbasisbefore there is
 gationcan and must be an integral part of the        the more ambitiousattempt to developa fully
 global programs to protect and preserve en-          international effort. Hopefully, the SSP can
 dangered    and rare species wildlife. Zoos and
                              of                      serveasa foundation,a modeland/ora compo-
 aquariums    are "an ark" that can provide sanc-     nent for more internationalprogramsby zoos
 tuary for speciesuntil or unless their natural       a n da o u a r i u m s .
 habitats  can be securedor restored. Four ways           A collective     strategyby zoosand aquariums
 in which zoos can directly contribute to the         is absolutelyvital if captive institutions are
 WCS Objective of "Preservationof Genetic                                                      of
                                                      really to contributeto conservation wildlife'
  Diversity"are:                                      Preservation viablelevelsof geneticdiversi-
                                                                         of
  1. to serve as refugia for speciesthat are des-      ty and demographic        security           main-
                                                                                         necessitates
      tinedfor extincionin the wild;                   tenance of large and stable populations.No
  2. to provide sourcesof propagulesfor repop-         matter how dedicatedor resourceful,indivi-
      ulation of natural habitats;                     dual institutions annot,for many reasons'
                                                                               c
  3. to reinforce natural populations which may        maintainlargeenoughpopulations manyor  {or
      be so small and fragmentedthat they are not      most species preservetheselevelsof diversi-
                                                                         to
  , viablegeneticallyor demographically;               ty over long periodsof time. Nor shoulda spe-
  4. to maintain repositories of germplasm in          cies be dependenton a singleor few institu-
      addition to or as an alternative to popula-      tions for its survival.Suchlimited distribution
      tionsof animals.                                 is very vulnerableto variouskinds of disaster'
     Further, zoos can perform a significant role         There are two general functions of the SSP'
   in two other parts of the WCS by conducting         One is to assistwith development of scientific
   researchthat will improve wildlife manage-           and cooperative programs to manage specles
   ment and by educatingthe public to support           as biologicalpopulationsin captivity. Just as
   conservation.                                        importantly, the SSPwill also attempt to pro-
      To better contribute to all of theseconserva-     vid-e some coordination for strategic assign-
   tion objectives, the American Associationof          ment of priorities and allocationof resources   to
   Zoological Parks and Aquariums has com-              species by the caPtivePrograms.
   mittedto a SpeciesSurvival Plan (SSP).The               Unfortunately, evenwith scientificand coor-
   SSPis an attempt to provide a strategic and          dinated management,the capacityof zoos and
   supportive  framework for programs to propa-         aquariumsfor populations large enough to be
   galeand   preserverare and endangeredspecies         viible is very limited in relation to the great
   in nnZpe institutions.Although the SSPhas            and growing number of species       requiring sanc-


                                                                                                      8l
                                     year'.lhe                                 the captive habitat optimally' The SSP will
tuary in captivity. During the last                                            uit"^pt to provide guidance suchstrategic
                                                                                                                 for
AAZPA      has sponsorea i "Wtrite ?aper". to
                                                                               -
                                                                               selection.
assess what resourcesare availablefor capttve
                                   inslitutions'                                   ittir processcan be iilustrated with an anal-
l.lounu,ion in its constituent                                                 vsisof ih. lutg.t felids' Table I assembles       in-
il".;?;th"r    information.this report' whichis
                                      indicates                                iormation on ihe       numberof taxa generallyre-
t"i"n F".ttt"r refined and extended'
                                   currentlv1al1ta1n
                        institutions                                           cosnizedin relation to the apparentcapacltyoI
ir,'"fTeZia                                                                     zois. Capacity, in this case, is being crudely
;;r;;i;;.ly             47.00-0^    mammals' 33'000bi rd^s'
                                                               159'000          ."urrr."d by tire populations that are currently
iblOOO eptiles.4.500 amphibians'
             r
                                                                                being mainiained' Specieshave been further
                          i
f i s h ,a n d 7 q . 0 0 0 n v e r t e b r a t e s '                                         into groupswith approximatelysim-
                                                                                orga'nized
     Thus selectionof speciesbecomesa process
      -assigning
                                                                                ilai requirementsfor facilities and resources'
 of                    priorities for allocation of the
                                                                                    For eachspecies     and category,there is then a
 ,ou.a u-ndresources               availablein zoos' For ge-
                                                                                 calculationof the number of taxa that could be
 ,r'.ii. ."uton t,      captive populationsshould be.as
                                                                                 accommodated the habitat estimatedfor the
                                                                                                    in
 large as possible. But there are many specles                                          institutions the populationof eachform
                                                                                                     if
                                                                                 trSIS
 competingfor this space'So it seemsnecessary
                                                                   every         has to be maintainedat one of three mtntmal
 to establish a carrying capacity for
                                                                                 sizesfor genetic (and demographic) viability'
 species.      T h i s c a r r y i n g a p a c i t y u s lb e a c o m
                                      c            m
                                                               popu.ta-          Consideringthe proportion of the total captive
  oromise between maintaining large
                                diversity and demographic                        facilities ttrit ttre ISIS institutions constitute'
  iio"t to. genetic                                                                        level of minimal population is.probably
                                                                    man-y        itt" ZSO
  stability aid providing sanctuaryfor a1                                                                   No computatlonsnave
                                                                                 the most reasonable.
  ;;t;          a s p o s s i b l e . S l r a t e g i c s e l e c t i o no f
                                                                                 U.Ltt p"ttot-ed to include the non-ISIS insti-
  sbecies     would, therefore,seemto requlre: as-
                                                                 that 1s         tutions sincegenerallydata on capacityare not
  slssment of the space and resources
                                                              of what             available.
   available in zoos; ascertainment
   ao""l".           in need of sanctuary,allocation of
               "."




                                                                        TABLE I
                                                                FOR LARGER FEL]DS
                            CAPACITY OF ISIS CAPTIVE FACILITIES

                                                                                         SUBSPECIES
                                                                                   NUI'IBER
                                                          IN     ISIS       ISIS     IF POPULATION
                                          EXTANT SUBSPECIES                                  5OO
                                                     DATA
                                                   RED   BOOK INSTITUTIONSPOPULATIONlOO U.
               I ES
            SPEC                         SUBSPECIES
                                                                  97        381      ll   11
           leo                              11         1
   Ponthero
                                                                 110        450      421
           tisrls                            8         8
   Ponthero
                                                                 r20        8J1      832
                                            19         I
   L i o n sa T i g e r s
                                                                    8                   65            178           2r0
           onco
    Ponthero                                  8
                                                                                        72            2U6           270
           oordus
    Ponthero                                 15                    I)


                                                                    I                                 r28           100
           unclo
    Ponthero                                   1

    Fells concolor                           ?9                     2                   69             173
                                                                                                       :-)
                                                                                                                    ?!9
                                             -
                                             ))                    26                  N,C,            tt           731
    " l n t e r m e d l o t eC o t s
                             "
                                                                                                        A?
                                               4                        ll              20
    N e o f el ls n e b u l o s o
                                               6                        b               32             IDO          1_q9
    Aclnonyx ubotus
                  i
                                                                    10                 N .C ,                        2r0
    " 0 t h e r " L o r s eF e l i d s        10                                                       "r"
                                                                                                      ,*            1873
                                              82                    45
    TOTALS


    82
   Admittedly, there are defects in this type of              Speciesare being selectedfor the SSP pro-
analysis.One problem is that there is incom-               grams by a comprehensive and coordinated
plete information of various taxa being main-              systemaccordingto criteria that reflect the gui-
tained. Only for thoseinstitutionsparticipating            delines of the World Conservation Strategy
in ISIS and/or for those speciessurveyed by                and objectives of the IUCN,ryYWF.Thus, the
I.Z.Y. or Studbooks there evenan estimate.
                      is                                   selectionprocessis: responsiveto statusin the
Moreover, this kind of analysisconsidersonly               wild; representativc of taxonomic, zoogeo-
quantity, not quality of habitat. It is fallacious         graphic,and other kindsofbiologicaldiversity;
to considerthat the speciesas categorizedare               6uLrealisticabout the feasibility of and capaci-
really interchangeablein their captive ecolo-              ty for propagatingparticular species  captivity.
gies.A further refinement that should normal-              Evaluations of potential candidates are de-
ly tre included in the table is the number of              pending greatly upon information provided by
subspecies each speciesthat are endange-
            of                                             the IUCN,^ilWF through the Red Data Books,
red. However, with the exception of Panthera               the SSC, the CBSG, the ICBP, and other re-
leo and Felis concolor. mosl taxa are consi-               lated agencies.  Indeed, the SSPis being deve-
dered in some degree of endangerment.The                   loped in very close consultation with the SSC
future doesnot appearoptimistic for any of the             CaptiveBreedingSpecialist    Group.
Lrigcatsin the wild into the next century.                     SSPspecies   are           by
                                                                                designated the AAZPA
   Despite all its defects,this analysisdoes, in            Wildlife Conservationand Management Com-
my opinion, provide an informative, if rough,               mittee. Currently,designation  may be initiated
picture of the situation for large felids in capti-         in two ways:
vity. Patently,not all taxa, even thosepresent-             1. The WCMC selectsspecies,more or less
ly endangered,   could be maintained in popula-                 systematically,according to criteria deve-
tions large enough to be viable in the facilities               loped.
 that appear available. Even for a specieslike              2. The WCMC considers candidates nomi-
 the snow leopard where there will be no con-                   nated by other interested persons and
 troversy or competition about subspecies,                      groups.
 there is going to have to be allocation of con-               Initial selectionshave concentratedon spe-
 siderably more habitat than is currently oc-               ciesthat can be employed asmodels for the en-
 cupied (Table2).                                           tire program and hencerepresentboth a diver-


                                                  TABLE 2
                                                   OF    LEOPARDS REGION
                                          POPULATION SNOII      BY




                                 Reql0n           Instltutl0ns         I ot
                                                                    Popu lon

                              l{orth Anerico                        69/ 59 = 118

                              Europe                  r4            25/ 32 = 57
                              USSR                    12            22/ \U = 36
                              Aslo                     2             2/ 2=      4

                                                      65           II8/II7   = 235
                              TOTAL



sity of organismsand a variety of problems' To              particularly high priority for the SSPprogram.
date, about 30 species have been selected,inc-                Each SSP program is organized around a
luding: Siberiantiger; Asian lion; black rhino;             SpeciesCoordinator who will be assisted a  by
Asian two-horned (Sumatran) rhino; Asian                    managementcomittee known as a Propagation
wild (Przewalski) horse; Grevy zebra; bata-                 Group. Further, to facilitate development and
singha; okapi; gaur; Scimitar-horned oryx;                  operation of theseprograms, the AAZPA has
Arabian oryx; Asian smallclawedotter; ruffed                created a position of ConservationCoordina-
ldmur; black lemur; golden lion tamarin; lion-              tor, who is located in the ISIS Offices. Species
tailed rnacaque; gorilla; orangutan; Roth-                  Coordinators must be approved by the
schild's (Bali) mynah; white-naped crane;                    AAZPA WCMC. Again, the ProcssmaY be
Humbolt's penguin; Chinese alligator. The                   initiated by the WCMC or by interestedindivi-
 snowleopard has been designateda speciesof                 duals and/or institutions. The SpeciesCoordi-


                                                                                                        83
                                                                                FIGURE 3
nator will be primarily responsiblefor develop-
ment and application of a Populational Mas-
terplan.                                              z
   The Propagation Group should include a             9r.
                                                      F

Studbook Keeper. In many cases, the Stud-             F
                                                      z
book Keeper and SpeciesCoordinator will be            u
                                                      o20
                                                      U
the sameperson. Where there is an Internatio-
                                                      u
nal Studbook Keeper outside Ihe AAZPA,hel
she is being invited and encouragedto serveon         u-
                                                      o
the Group. Indeed, where possible,it would be         F
                                                      z
optimal and appreciatedif a Studbook Keeper           U,N
                                                      o-
outside the AAZP Awould serve at least as Co-
                                                      CE
                                                      U
                                                      c
Coordinator for the SSP programs. A repre-
sentative from the SSPSubcommitteeand the
 AAZPA Conservation Coordinator also serve
in an ex officio capacity on the Propagation
Group for each species.
    Other members on the Propagation Group
are elected by and from the participating insti-     application               ,".t;;""0"t,"t            from the
tutions. The size of the Propagation Group is                          ", "
                                                     SpeciesCoordinator and Propagation Group.
limited to a workable number of persons.             Transactionsderiving from recommendations
 1. For SSPprograms involving 10 or fewer ins-       from the Propagation Group to relocate ani-
     titutions, there is at least 1 representative   mals will be between the institutions involved
     from each on the Propagation Group, with        and may entail sales,           exchanges,       donations,or
     a maximum of 10 members.                        loans    at their discretion. An example of a
 2. For SSP programs involving more than 10          Memorandum of Participation prepared for
     institutions, eachnominatesa candidatefor       possible application to the snow leopard
     the Propagation Group. The list of nomi-        a p p e a r s l t h e e n d o f t h i sP a P e r .
                                                                 a
     nees is circulated to participating institu-        The real substance an SSPprogram is the
                                                                                     of
     tions, each of which vote for 10. The ten re-   Populational Masterplan. Collections of spe-
     ceiving the most votes serveon the Group.        cies in captivity can be preserved viably for
    The terms of servicefor the SpeciesCoordi-        long periods of time only if they are managed
 nator on the Propagation Group are limited           as biological populations. Thus, the SSP pro-
 but renewable.All thesepersonsare subjectto          grams are being predicated on multigenera-
 regular review by the WCMC. The SSP Sub-             tional masterplans for genetic and demo-
 committee participates in this review by eva-        graphic managementformulated with data and
 luating the Populational Maserplan. Length of        analysisprovided by the International Species
 the terms of serviceare a matter for further de-     Inventory System(ISIS) and Studbooks.
 termination.                                             An SSPprogram must insure that the basic
    A "Memorandum of Participation" docu-             data (individual identification, sex, parentage,
 ments the commitment of an institution to par-       birthdate, and deathdate)requisitefor popula-
 ticipate in an SSPprogram. The Memorandum            tional analysisand managementare compiled-
 of Participation commits each participant to         This objective will normally require that data
 manage their animals in accordancewith the           be entered and improved in the International
 guidelines of the Populational Masterplan and         Species Inventory System (ISIS) and that a
 ihe recommendationsof the Species      Coordina-      Studbook is being well maintained. With the
 tor and the Propagation Group. This docu-             principal support of AAZPA, ISIS has been
  ment emphasizesthat the commitment is to             developedto compile and analyze,in a centra-
  cooperatein the program, not to transfer of          lized and computerized manqer, the informa-
  ownership or relinquishment of control of ani-       tion necessary populational managementof
                                                                           for
  mals. There is a definite but renewableterm of       captive species.      The SSPand ISIS are thus very
  commitment and ample provision for the zoos          interrelated and both will continue to develop
  to terminate participation. Further' a petition      together. For snow leopard, close cooperation
  ratified by a simple majority of participating       has alreadybeen developedbetween the Inter-
  institutions (1 vote each) can nullify general       national Studbook and ISIS.


 84
                                                                       3.         provide for the population's rapid expan-
                                                                                  sion to and stabilization at the carrying
                                                                                  capacity with an age and sex structure
                          TABLE 4
                                                                                  that will optimize genetically effective
         OF
 I{UI'IBER LIVING ANIt.lALS
                          I.IIIH REPRESENTATION
                                            OF VARIOUS
                                                     FOUNDERS                     size and demographic stability; this will
                                                                                  normally entail both removal of animals
                                                                                  from certain age and sex classes well as
                                                                                                                   as
         Foundel                USSR                    Asio                      regulationof reproduction;
     Kounos     I          2/1,= 3  (87)                               4.         analyzeeach speciesgeneticallythrough
                2          2/I=    3(82)
                q          2/1,='t
                                                                                 geneological as well as electrophoretic
                                    187,)
                    d
     L e n i n g r o9      I/3= 4   l11I)                                        and karyotypic studiesto assess diver-
                                                                                                                  the
                   I0               (I1 L)
                           0/ tr = t!                                            sity and distribution of the genepool;
     lvloscow  Ll          5/7= 6   tI/7,)
               5           \/2=    61177")                             5.        maximize preservation of genetic diver-
     Novosibirsk      l    2/I=     (87,)                                        sity in the species
                                   3
                                   (87")
                                                                                                    by:
                      4    J/0= 3
                                                                           a.    insuring that there are an adequatenum-
     Nul ioorous
           I
     1 4l id b o r n s    II/6=17       ltl7Z)   2/2=    \   (t]OI)              ber of founders, where available,for the
                                                                                 captivepopulation.
                                                                           b.    attempting to perpetuateequal represen-
                                                                                 tation of thesefoundersin the population
   The masterplans themselves will presentde-                                    through time.
mographicand geneticanalyses   and will provi-                             c.    retarding genetic drift by optimizing ef-
de recommendationsfor both general strate-                                      fective population size through regula-
giesand specifictacticsfor management,i.e.:                                     tion of family sizes, sex ratios, and age
I.   what the sizeand structure of the popula-                                  structure.
     tion is presently, potentially, and opti-                             d.   minimizing or at least managingconsan-
     mally, in terms of numbers, ages, and                                      guineous reproduction by rearrange-
      SCXCS;                                                                    ments of animalsto separaterelated spe-
2.    how many institutions should be accom-                                    cimens.
      modatingthe species:                                                 e.   avoidingmost artificial selection.
3. which animals should reproduce, how                                     f.   optimizing the number of demes (sub-
      often and with whom;                                                      populations or groups) into which the
4. which animalsshould be maintainedin or                                       population is divided.
     removedfrom the population;                                      6.        in addition to maintenance of popula-
5. what basic standards of husbandry and                                        tions of animals, direct collection and
     considerations sociobiologyshould be
                     of                                                         preservationas much germplasmas pos-
     emphasized.                                                                sible.
   More techncally, the populational master-
plansmust:                                                            An SSPProgram for Snow Leopard
1. determine an optimal carrying capacity                              In the caseofthe snow leopard, the processof
     for the captive population so as to maxi-                         developing a cooperativeprogram for popula-
     mize its genetically effective size under                         tional management is already well advanced
     the constraint that many other species                            under the guidanceof the International Stud-
     must be provided sanctuaryby the limited                         book Keeper. Institutions have been consult-
     space and resources of zoos and aqua-                            ing Leif Blomqvist for advice and/or approval
     riums. (Normally, there would have to be                         on management and transactions with their
    very cogent arguments and exceptional                             animals. This activity is an excellent example
    circumstances for this carrying capacity                          of the implementation of the recommendati,on
    size to be more than 500 animals.) Also                           that emanatedfrom the International Union of
    recommendedshould be the number of                                the Directors of Zoological Gardensthat Stud-
    habitats or institutions over which the                           book Keepers should increasinglyoperate less
    species  shouldbe distributed;                                    as data compilersand more as species     mana-
2.   analyzeeach speciesdemographicallyto                             gers.
    determine patterns and potentials of sur-                            Nevertheless,there would be advantaqes   to
    vivorships and fertilities, and hence, of                         organizing the snow leopard p.ogrurnhor.
    change;                                                           formally under the auspices of, or at least


                                                                                                                     85
according to the model of, the SSP.Whily in-                          POPULATIONAL ANALYSIS
breeding has certainly been avoided or alle-                          AND MANAGEMENT OF
viated by the endeavors of the International                          THE SNOW LEOPARD
Studbook, a number of genetic and demo-
graphic problems remain or impend. More-                              As a basis for an SSPprogram, some prelimi-
over, a more formal structure should facilitate                       nary analyseshave been performed on the ge-
communication and improve to cooperation.                             netic and demographicstatusand prospectsof
An SSPprogram would also constitute an en-                            the snow leopard in captivity. Hopefully, these
terprise for this speciesthat could be endorsed                       analyses can be extendedand applied to impro-
by the IUCN SSC Captive Breeding Specialist                           ve the population managementof this very en-
Group which appearsto be emergingas the in-                           dangered species.These analyseshave been
ternational authority for the captive compo-                          conducted with the assistance   and facilities of
n e n t so f w i l d l i f ec o n s e r v a t i o n .                 the International Species Inventory System
   The International Studbook Keeper should                           which will be available for further support of
serve as at least the Co-Cordinator of any SSP                        the Snow Leopard Program.
Program organized. It would, however, also
appear constructive to appoint a Co-Cordina-
tor from North America to be able to coordi-                          Genetic Analysis and Management
nate, more closely, management of the large                           The major objective of genetic management
part of the captive population located on this                        for endangeredspeciesin captivity is to pre-
continent.                                                            serveas much aspossibleof the heritable diver-
                                                                      sity that has evolved and exists in the wild
                  FIGURE 4                                            populations. There are three more or less
                                                                      interrelated components of a strategy to
                                                                      achievethis objective:
                   U.S.S.R.                                           L. Maximizing the geneticallyeffective size of
                                                                          the population.
                            &                                         2. Equalizing the representation of the wild
                       ASIA                                               born founders.
                                                                      3. Minimizing inbreeding coefficients of pro-
                                                                          genyproducedby captivereproduction.
                                                                         Loss of genetic diversity depends upon the
z                                                                     sizeof a population. In general,the smaller the
o
tr                                                                    population, the faster genetic diversity is lost.
F                                                                     However, the size of significanceis not merely
z
uJ                                                                    the total number of animals in the population
U)
lJ.t                                                                  (Seal and Flesness 1979; Soul6 and Wilcox
E                                                                     1980). Rather, the genetically effective size,
(L
UJ                                                                    Np, is also a function of the structure and the
(r
                                                                      dynamics of the population. The same total
u,                                                                    number of animalscan produce very disparate
o
                                                                      Np's. Populationscan be managedso their ge-
F
z                                                                     netically effective sizecan very from a tenth to
1!
o                                                                     twice the actual number of animals.Significant
cc                                                                    factors determining genetically effective size
LU
o-                                                                    are the number, sex ratio, and family sizls (i.e.
    PARITY
                                                                      total number of offspring produced by an indi-
                                                                      vidual in its lifetime) of the animalsthat actual-
                                                                      ly reproduce. The smaller the fraction of ani-
                                                                      mals that actually reproduce and the rnore dis-
                                                                      parate their sex ratio and family sizes,the lo-
                                                                      wer the genetically effective size will be.
                    I104            5 3 411u11]o"rou"                    Maximizing effective population number for
                   L g r a d M o s c o w N o v o sW i l d B o r n s
                                                                      the carrying capacity size of the population will
                   FOUNDERS                                           therefore require:
1. As many animalsas possiblebe recruited to                             (Because of both geographic and gene.tic
    the reproducingpoPulation;                                       s e p a r a t i o ni.t s e c m si n f o r m a t i v e o r e c o g n i z e
                                                                                                                             t
    -fhe                                                             regional populationsfor some of these ana-
2.         sex ratio, and especially the family
    sizes, these
            of           individuals equalized far
                                        be                as         lyses).Only wild born animalsthat have actu-
    asis feasible.                                                   allv reproducedhave been included in these
   For snowleopards,           ofthe 323registered         ani-      computations.           There are another 2lll4 : 35
malsthat were born before 1976,only Il7, or                          nulliparouswild borns in captivity that might
36 "/", have reproduced. The sex ratio of the                        potentially be recruited as founders' If well
breeders not been too uneven(54/63)and
             has                                                     managed,this number of foundersshould be,
would havedepressed only slightly.Much
                                Ng                                   as Leif Blomqvist (1981)has observed,more
more serious       hasbeen and continues be the        to            than adequatefor a viable populationin cap-
very disparate       family sizesof individuals(Fig-                 tivitv.
ure 1). It will be importantto attcmptrectifica-                         More important than the actual number of
tion of this pattern in the future . This desidera-                   f o u n d e r si s t h e i r p r o p o r t i o n a lr e p r e s e n t a t i o n
tum should be facilitated by the further im-                          t h r o u g ht i m e . F u n d a m e n t a l l y .r e s e r v a t i o o I
                                                                                                                        p                     n
provement of husbandry that will promote                             genetic diversity will be maximized by equa-
betterreproduction more individuals.
                             by                                       lizing the representation founders. In this
                                                                                                                of
   Logically, no more genetic diversity can be                        respect, the situation for the captive popula-
preserved      than hasbeenimportedinto the cap-                      tion of snowleopards lesssanguine. is                          Propor-
tive population.But, comparedto someother                             tional representation foundersis very dispa-
                                                                                                        of
endangered        species     that are evidentlyviable,               rate in all three major regions(Figures2 4).
the captive population of snow leopards de-                           S o m e b l o o d l i n e sa r e e x t r e m e l y o v e r r e p r e -
rives from a relatively ample number of                               sentedwhile severalothers are on the verge of
f o u n d e r sF i g u r e s - 4 . T a b l e s - 4 ) :
              (            2                  3                       disappearance.This pattern is evident from
         North America 15117                :32                       anotherperspective, number of living ani-
                                                                                                        the
          Europe                            :
                                    9 t r 12 0                        mals with representation the various foun-of
          USSR                      5t4: 9                            ders (Tables3-4). In North America, the Co-
          World                    21t24:      45                     penhagen,         and to a lesser       extent,the SanAnto-


                                                                       FIGUBE I


                                                       PROGENY




                o

                =15
                2
                l!
                o
                E10
                ul
                o
                =
                z




                                                                  FAMILY      SIZE

                                                                                                                                                81
nlo and San Francisco,bloodlines are particu-
                                                                                                          TABLE 3
larly predominant. Management of founder
stock needsto be improved for better preserva-                        I U S E i O F l I V I i l G A N I M L S H I T H R I P R S € f l T A T I OOl F V A R I O I S O U N D E R S
                                                                                                                                               i                 F



tion of the genetic material that has been ob-             Founder                          !0rtlr,!]fts                           t@e                                 I9t!l

                                                       Bronx 15                            9/5=rq            (10:)                                          9/5-14

tained from the wild.                                           t8
                                                       C e n t e rN l l l I
                                                                                           7/5=12
                                                                                           t/I-       2
                                                                                                             197)
                                                                                                             lLTl          1/0=         1        (21)
                                                                                                                                                            // 5 = 12
                                                                                                                                                            2/l=    J
     However, equalizingfounder representation                   2
                                                                                           7/8=15
                                                                                           u/2= 6
                                                                                                             (lIt)
                                                                                                             141)
                                                                                                                                                            7/ 8 . 15
                                                                                                                                                            4/2-    6
may not be as simple as it may appear in prin-                   t                        to/ / - 17
                                                                                           0/]=       I
                                                                                                             lr2 7t
                                                                                                             (I1)
                                                                                                                                                           ro/7-\1
                                                                                                                                                            0/i-    I
                                                       BrookfieldI
ciple. Not all of the founders enumerated in                       2                        0/t=      I      {11)
                                                                                                                          II/ / - 18 (12 l)
                                                                                                                                                            0/t=    I
                                                                                                                                                           2A/24 - 4\
                                                       tPl t                                9/\/ - 26        lI9 7)
Table 3 and 4 and Figures 2 to 4 are indepen-               5                               5/6=11           (8r)
                                                                                                             (8t)
                                                                                                                           1/0= I (2I)                      6/6.12
                                                            8                               2/9-11                                                          2/ 9' \I
dently or completely representedin the extant          clnclnnot' I                       I 5 l 1 1= 2 6     (19;)          2/Q=         2       t\Xt      )7/I\ - 28
                                                                                          l5lll = 26         (19 l)                                        \//)I - 28
population. Some, such as Bronx 15 and 18,
                                                                   2                                                        2/A-         2       l\2,
                                                       Derroit I                            0/l=      I      (iz)                                           a/\.
                                                                                                             (11)
have passed through bottlenecks of a few,
                                                                2                           0/l=      I
                                                        tomonton I                          t/J=      \      ta7)
                                                                                            \/ t = \
sometimesa single animal. Hence, only a frac-                    2                                           | 37)                                           I/3-
                                                                                           2/t=    5         l\7,                                            2/ t -

tion of the original geonomesof these "foun-           f l 1l * o l k e e 2
                                                                              l            2/3= 5
                                                                                           8//.15
                                                                                                             t47)
                                                                                                             (1Il)                                          8//=1   5

ders" could possibly have been transmitted to          o k ) o h o m o2
                                                       hoho I
                                                                                          t5llq = 29
                                                                                           2/t=    5
                                                                                                             1 2 1r )
                                                                                                             tu7\
                                                                                                                            2/2-         \       \7Xl      \//16 .
                                                                                                                                                            2/ t =
                                                                                                                                                                    t1
                                                                                                                                                                     5

the present. In suchcases, doesnot seempru-
                              it                                              2
                                                                                           9/r7 - 26
                                                                                           5/t6 = 2l
                                                                                                             t\9 7l
                                                                                                             (15;)
                                                                                                                            0/5.5
                                                                                                                            0/u-    4
                                                                                                                                                 (9!)
                                                                                                                                                 t77t
                                                                                                                                                            9/22 =
                                                                                                                                                            ,/2A =
                                                                                                                                                                    3\
                                                                                                                                                                    2'

dent or even possibleto accordequality of rep-                                    5
                                                                                          22/t\ - tJ
                                                                                          22/\r - t3
                                                                                                             t?4 7)
                                                                                                             t2\ ll
                                                                                                                            ?/2-
                                                                                                                            2/2-
                                                                                                                                    \
                                                                                                                                    4
                                                                                                                                                 t/7\
                                                                                                                                                 \/7)
                                                                                                                                                           ?q/rt -
                                                                                                                                                           zq/I) =
                                                                                                                                                                    37
                                                                                                                                                                    37
resentation. Thus, while equalizing founder             seottle I
                                                                2
                                                                                            6/6=t2
                                                                                            6/E-r?
                                                                                                             19t)
                                                                                                             191)
                                                                                                                            2/O= 7
                                                                                                                            2/Q= ?
                                                                                                                                                 t4Z)
                                                                                                                                                 t\71
                                                                                                                                                             A/6=lq
                                                                                                                                                             8/6=Iq
representation is perhaps the major objective                                 2
                                                                                           t//J/ - /\
                                                                                           tl/1/ = /q
                                                                                                             t5\ t)
                                                                                                             15\ 7i
                                                                                                                           Ii/ 9 = 22
                                                                                                                           It/ 9 . 2?
                                                                                                                                                 tt9 7)
                                                                                                                                                 (t9 7)
                                                                                                                                                            50/q6 = 96
                                                                                                                                                            50/q6 = 96
for geneticmanagement,there are thesecomp-              Helslnkl I
                                                                 2
                                                                                            q//.11
                                                                                            q//=I\
                                                                                                             (8i)
                                                                                                             \87'
                                                                                                                           ]!/20 = J4
                                                                                                                           tql20 - lq
                                                                                                                                                 (60 X)
                                                                                                                                                 (60 t)
                                                                                                                                                            !3/27 . 45
                                                                                                                                                            Ia/27 = \5
lications. Severalgroups, including the staff at        Zurich I
                                                               2
                                                                                                                            o/\=
                                                                                                                            A/2.2
                                                                                                                                    \            tTtl
                                                                                                                                                 \\l)
                                                                                                                                                             o/q= \
                                                                                                                                                             A/2= 2
 ISIS, are at work on this problem and hopeful-                                                                              0/I=        |       l27l        0/l=         I
                                                        Krereld I                                                            0/\.        t       t27)                     I
 ly will be able to provide refined advice in the
                                                                                                                                                             0/1=
                                                                                            q/ I =       5    | \7t          q/5=            9   u5t,         8/6=I\177'
 near future. In the meantime, there is much
 that can and should be accomplished redress
                                          to
 the imbalanceamongbloodlines.
     Very critically, founder or bloodline analysis    animals merely for this purpose, there should
 identifies certain specimensas extremely im-          at leastbe provisionsto collect sperm opportu-
 portant in terms of the genetic material they         nistically should animals be tranquilized for
 possess.                                   in
             The founder representation every          other reasonsor should animals die. Research
 animal known aliveis presented      inTables5-7.      should be conductedon artificial insemination
 (Simple 1 to N numbershavebeenassigned          by    to permit utilization of sperm that can be col-
 ISIS to specimens facilitateanalysis.)
                       to                      Vari-   lected. Superfertilization and embryo trans-
 ous sorts of immediate action should be consi-        plants should be explored as a meansof rapidly
 dered to securethe genetic material these ani-        expanding the representation of rare blood-
 mals contain. Where feasible, every effort            lines, especiallythose survivingonly in females
 should be exerted to activate these animals           whose productivity is, of course, more limited
  reproductively. In some cases, relocation of         than in the caseof males.
 animals would be advisableto produce better              Equality of founder representationis impor-
 pairs genetically,behaviorally, and physiologi-       tant for the geneticdiversity of the population.
 cally.                                                Inbreeding, in its strictestsenseofconsanguin-
      Better interactions between the major re-        eous reproduction, is important for the genetic
 gions would be especiallyconstructive. North          diversity or heterozygosityof individuals. For
  America and Europe have actually been en-            the optimal vitality of individuals, inbreeding
  gaging in considerableexchangeof bloodlines          should be minimized or at least closely ma-
  recently. However, the collections in the            nagedin captivepopulations.
  U.S.S.R. and Asia remain isolatedfrom the               Inbreeding in captive snow leopards is still
  rest of the captive population. Gene flow be-        modest. Only 24 (- tO %) of 235 animals
  tween these regionswould be mutually benefi-         known to be alive, as of this analysis,are defi-
  cial.                                                nitely inbred, i.e. possessinbreeding coeffi-
      But, the situation is so critical for many of    cients )0 (Table 8). Indeed, of the 470 alive
   these specimensthat various technologicalas-        and dead animalsknown to be registeredin the
   sistanceshould perhapsbe invoked. Certainly,        International Studbook, only 38 are definitely
   sperm should be collected from critical males.       inbred. Inbred animals still believed alive are
   If risks are consideredtoo sreat to immobilize       identified in Table 9. The situation may be
                                                                 FIGURE 2


                                                            NORTH AMERICA
z
9
F                  I


F
z                  I

llJ
o
IJJ
TT                 o
(L
UI
E                  5



lrJ
(t
      P A R I TY
F
z
l!
o
tr
UJ
o.
                       t5t8 t 123l2               35         |2121213                       2 2'1  1'2'3' 5't'2      1'2   l 2Nrriiparors
                       Bronr Ctr Choyenne Brkfld.  L.P.Z.   Clncln, Oetrolt Edmntn. L.A.           S.Ant.
                                                                                           Mlw,OKOma          Seaitl€Cpnhcn.
                                                                                                        S.Fran,                 Wild8o.ns
                                                                                                                           Hlenk.
                             Hlll
                                                                    FOUNDERS




somewhat    worse than computed sincethis ana-                                                               TABLE 8
lysis presumesthat all wild born founders of
the captive population have been unrelated                                                                 OF     ANII'IALS
                                                                                                     NUI1BER INBRED
and noninbred. However, the acquisitiondates
for severalpairs of the founders (Brookfield 1                                                N o r t hA m e r i c o :   10/9=19
& 2; Seattle 1 & 2; Edmonton 1 & 2) provoke                                                       Buffolo                0/r
suspicion that they were littermates. Since                                                       Houston                v0
these pairs, particularly the Seattle animals,                                                    Bronx                  7/ 3
have reproduced together, their progeny may                                                       11i
                                                                                                    lrioukee             UO
be the offsprng of full sibs and hencebe inbred                                                   0klohomo               r/3
at the .25 level.                                                                                 S o nA n t o n i o     0/2
   Nevertheless,  consanguinitydoesnot yet ap-
pear to have developed into the serious prob-                                                 Europe:                    4/ I=5
lem afflicting many other endangeredspecies                                                       Bekesbou   rne         YI
in captivity. In part, this sanguinesituation is                                                  L e i p zs
                                                                                                          i              u0
due to the fact that snow leopardshave not yet                                                    lvluni ch              v0
propagated   through many generationsin zoos.                                                     Zu ich
                                                                                                     r                   v0
But, no small credit is due to the efforts of the
International Studbook Keeper and the snow                                                    U.S.S.R.:                    0
leopard institutions in managing their collec-
tions to avoid consanguineous      reproduction.                                              Asio                         0
However, 28 of the 65 current collections do
                                                                                              TOTAL                      Iq/10 = 2\
contain related pairs (Table 10). As propaga-
tion continues more care will be necessary     to
minimizeinbreedins.                                                                        Ronse,008 Io ,25t - lOUpopulotionlnbred


                                                                                                                                      89
                                   TABLE 9                                                                                      TABLE TO

                                        ANIIVIAIS                                                                                 PAIRS
                                                                                                               INSTITUTIONS RELATED
                                                                                                                          l,lITH
                                   INBRED
                                                                                                                         ( 6 5 I n s t i t u t l o n s T 0 t 0 l)
                            I
                     I nbreedns
                     coefflcient                    Anlmol
 Anlmol                                                                      I nbreedng
                                                                                    I
                                                                             Coefflclent           BotonRouge                                           Konsos ltv  C
                        ,25                Bekesbourne 1
Bronx 22                                                        2                                  Bekesbou      rne                                    K0un0s
          t7            ,25                                                                                                                             KrqfeI d
                        ,25                l l e l si n k l     8                                  c e n t e rH i l l
          2U                                                                    ,25                               l'lountoln                            Leipzi9
                        ,25                                    22                                  Cheyenne
          25                                                                     .E
                                                                                                                                                           Iw0ukee
                                                                                                                                                        l'11
                        aq                                     23                                  L l n c o l n P o r kZ o o
          zo                                                                    ,25                Cinclnnot      I                                     NettI no
                        .008                                   28
          27                                                                     )q
                                                                                                   c I eve lond                                         Bronx
          28            ,008               0klohomo
                                                               1l                                  coI umbl    o                                        0klohomo
          29            .008                                                    ,r25                                                                    Progue
                                                               T2                                      I
                                                                                                   Co umbus
          31            ,008                                                    , \t)                                                                   S o nA n t o n l o
                                                               73                                      I
                                                                                                   Do los
          )z            ,008                                                    ,r25                                                                    s e o t tI e
                                                               Ill                                 Detr0it
          33            .008                                                                                                                            St, Louis
                                                               15                                  D u biln
          34            .008                                                     1r(
                                                                                                                                                        l.llnnlDeg
                                                                                                   Helslnkl
                                                                                                   Houston                                                 r
                                                                                                                                                        Zu l ch


                                                                                                           (28 tnstltutlonslllth Rel0ted olrs)
                                                                                                                                        P




                                                              FIGURE 5
                                                                                                                                         Total
                                                                                                                                         Populatlon




                                                                                                                         ',              Captlve
                                                                                                                                         Born

                                                                                                                                t

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                                                              YEAR

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100
                                                       AGE STRUCTURE
                                                       SNOW LEOPARDS
                                                        1 J A N U A R Y 19 8 1

                                                                                          Females

       urrrowtl aoE
               i?-la
               It-17
               It-lt
               1a-15
   o           It-ltl
   o           12-14
               t l-12
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   o             e-to
                  t-e
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   o               G-7
                   6-6
                  a-5
                   3-a
                   2-3
                   1'2
                   O-l
                      20
                                                                               o

                                                                            NUMBER
                                                                      FIGURE 6



                                                                     STABILIZATION
                                 SOME OPTIONSTO ATTEMPT DEMOGRAPHIC
                                                           CARRYINGCAPACITYOF 25O
                                 OF AMURTIGERPOPULATION AT

                   oPTroN I       oPTloN 2            3
                                                 OPTION       oPTtoN 4-OPTtoN 5.o
                                   DJUS'EO IO                                                                     rHBEE IITTE'S     LITIEi OF
                                                               ONE LI?IEi    ONE II'TEi
     IF                                          ofl€ lfrEh
                                                                                                                   2.43 CIAS        2.43 CUBS
REPRODUCTION                                                                              2.41 C0A3   2.a! coBs
                     1056-A0
        ts                                                      a cuas        3 CUBS
                                                                                                        EACX
                                                                                                                     €ACH
                    LtrTEB SEE
                                   oFlfilaur€D




THE REMOVAL
                                                                                             30%        35%             53%           72%
O.I YEAR OLDS            46%            o%                         o%
REQUIBEO FOR
STABILITY     ts
              IS
       -oR-

FEMOVAL FROM
EACH AGE                                                                                   4.85%      7.75%           9.5%            | 5.5%
                         7%            o%
REQUIRED FOR                                                       o%
STABILITY     ts


GENERATION

                                                                                                                                  TABLE IT



                                                                                                                                                l0l
Demographic Analysis                                   conspicuously,the age distribution appearsto
and Management                                         be developingthe potential for stability, with a
                                                       healthy baseof younger animals.
For genetic reasons, captive populations
                                                          Until the patternsof survival and fertility are
should be as large as possible. But size is not
                                                       better established,it is probably too soon to
quite enough. The population must also be
                                                       compute various schedules of reproduction
maintained stably at this size. Genetically, if
                                                       and removal that would stabilize the popula-
populations fluctuate significantly, Np will be
                                                       tion. However, as a preview of what will be in-
closer to the minimum than to the maximum.
                                                       volved, Table 11 arrays various options that
Stability is also vital to avoid population explo-
                                                       have been projected for stabilization of the
sions and collapsesthat can causeextinction of
                                                       North American population of Siberian tiger
speciesin captivity. Regulation of population
                                                       (Panthera tigris altaica). A similar program
size and stability is the function of demo-
                                                       ;iI .*"t*lly      b"            for the snow"leo-
graphic management.                                                         ""*tsary
                                                       pard. Almost certainly, there will be a need to
    Naturally, the first demographic objective
                                                       remove animalsfrom the population eachyear
for a captive population must be the develop-
                                               rep-    for demographiccontrol. Naturally, this stock
ment ofhusbandry conduciveto successful
                                                       could be available for reintroduction of the
roduction by the species. To a considerable
                                                       species  into its natural habitat.
extent, this objective hasbeen achievedfor the
snow leopard although appreciable problems
                                                                                        XTMiAMUX Of PAiIICIPATIOi
must still be resolved.                                                                               f$5
    The captive population of snow leopards                                                                                                        !
may be developingthe pattern and potential of
                                                             r i e ( :

vigorous growth that is requisite for demo-                  n
                                                                "]tli6Drmif$-t66ffi
                                                                                                                                               .l

graphic security. Figure 5 depictsthe history of                                                          rn a proerah or Doplrilloi'r         I


the captive population, and various compo-                   cooai..b..            desien.ted ty      rh€ rirdriie                              g



nents thereof, from 1960to 1980. Clearly, the                ::ii.:i'":"Hl;:1i;,:'i#i";."1,::r.T::t?,il.i"".1'::";:Jl.1i,::
                                                                                                                     iJ
proportion and size of the population that is                                                              ttr   sor.ie!   Coordli.!or   .nd
                                                                                                                                               :
captire born is on the increase,significantlyso
                                                             Proprqrrioi          Grou9 riil
                                                                                                                                               .9
                                                             obiecr,!r       of    roiq-teri   i.Jn
                                                                                                                                               'a
in recent years.                                                                                                                                   9
                                                                                                                                                   a
    With theseprospects,it is not too early to be
consideringmanagementto stabilizethe popu-                                                                                                          E
lation at a determined carrying capacity. Such
                                                             iffiiffir




stabilizationwill eventuallyrequire that repro-
duction be restricted and/or that stock be re-                                           ./o Nrrnesotr
                                                                                         rppt. v,|.r,
                                                                                                         roor09ic.r
                                                                                                         ril 55124
                                                                                                                      G.rden



moved from the population.                             References
    Stabilizationalso requiresthat agestructure,       BLOMQVIST, L. 1978a: First report on the snow leopard,
as well as total numbers, be regulated. Cons-          Panthera uncia, studbook and the 1976 world. register. Int.
tancy of numbers is impossiblewithout stabili-         Zoo Yearb. 18:227-231, 1978
ty of the agedistribution. The nature of a stable      BLOMQVIST, L. 1978b: The snow leopard, Panthera
                                                       uncia. in capitivity and the 1977 world register. Int. Ped.
 age distribution has been discussedin detail          Book of Snow leopards1: 22-24, 1978
 eliewhere(Foose1981a & b; Goodman1980).               BLOMQVIST, L.: The snow leopard register. Int. Ped.
 It will suffice here to statethat a stabledistribu-   Book of Snow leopards2: 218-238, 1980
 tion will have a broad base of young animals          BLOMQVIST, L.: The 1980 annual report of the captive
                                                       snow leopard (Pantherauncia.)population and a review at the
 and a gradual decline in numbers through the          breeding resultsduring th? 1970's. Helsinki Zoo Annual Re'
              age
 successive classes.                                   port 1980:32-50, 1981
    Figure 6 presents the age distribution pub-        FOOSE, T. 1981a: Demographic management of endan-
 lished by the International Studbook for the          geredspeciesin captivity. I nt. Zo o Yearb. 20: I 54 - I 65, I 98I
                                                       FOOSE, T. 1981b: Demographic Problems and Manage'
 captive population alive on 1 January 1981
                                                       ment in Captive Populationsl Proceed. of the 1980AAZPA
 (Blomqvist 1981).Lack of data on European             Annual Conference.AAZPA. Wheeling, WV: 46-68, l98l
 and Soviet collectionsduring 1981and t982ha-          GOODMAN, D.: Demographic interventionfor closelyma-
 ve prevented an update. However, even this            nagedpopulations. In: Conservation Biology. M. Soul6 and
                                                       B. Wilcox (ed.). Sinauer Assoc., Sunderland, Mass.: l7l-
 age distribution is most encouraging. More
                                                        195.1980
 sophisticatedanalyseswill be required to de-          SEAL, U. and N. FLESNESS:Noah'sArk* Sex               and Surui-
 fine exactly what the stable configurationsfor        val. Proceed. of the 1978 AAZPA Annual Conf. AAZPA.
 this speciesreally are (Foose 1981a & b). But         Wheeling, WV.



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