the mountain tapir, endangered flagship species of the ... · the mountain tapir, endangered...

ORYX VOL 30 NO 1 JANUARY 1996

The mountain tapir, endangered 'flagship'species of the high Andes

Craig C. Downer

The mountain tapir has already disappeared from parts of its range in the highAndes of South America and remaining populations are severely threatened byhunting and habitat destruction. With an estimated population of fewer than 2500individuals, urgent measures are necessary to secure a future for the species. Thispaper presents an overview of the species throughout its range as well as some ofthe main results of the author's studies on tapir ecology. Finally, a plea is made forconservation action in Sangay National Park, which is one of the species's mainstrongholds.

The mountain tapir: an overview

The mountain tapir Tapirus pinchaque, was dis-covered by the French naturalist Roulin nearSumapaz in the eastern Andes of Colombia(Cuvier, 1829). The species is poorly knownand most information has come from oc-casional observations or captures in the wild,reports from indigenous people in thespecies's range, and observations and studiesin zoological parks (Cuvier, 1829; Cabrera andYepes, 1940; Hershkovitz, 1954; Schauenberg,1969; Bonney and Crotty, 1979).

Like lowland tapirs, mountain tapirs areprimarily solitary, herbivorous browsers.They prefer moist habitats and bathe fre-quently. In spite of their thick woolly fur, theydepend on cloud forests for shelter from thefierce storms that assail the Andes.

The most distinctive feature of the tapirfamily is the prehensile proboscis, which isemployed in browsing, for orientation,bathing, and even squirting pursuers withwater. Their sense of smell is keen (Walker,1964) and allows the tapir to orientate at nightor in dark forest. It usually moves with itshead and trunk close to the ground (D'Aulaireand D'Aulaire, 1979; Janis, 1984).

The species has been recorded between 1400and 4700 m in the Andes of Colombia andEcuador and parts of northern Peru, but oc-curs most frequently between 2000 and 4300m. However, it is rapidly disappearing

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throughout its range. There is limited evi-dence to indicate that it may have occurred inwestern Venezuela about 20 years ago.However, Venezuelan authorities indicate atotal absence of mountain tapir remains fromVenezuelan territory, either from the recent ordistant past (J. Paucar, F. Bisbal, O. Linares,pers. comms). Northern Peru contains only asmall population (Grimwood, 1969).

In common with many montane forestsworld-wide, those of the Andes are beingrapidly destroyed, causing the decline of thethe tapirs (Poore, 1992). The mountain tapir isone of the world's most critically endangeredlarge mammals; the IUCN/SSC TapirSpecialist Group estimates its population asfewer than 2500 individuals and that it has agreater than 20 per cent probability of disap-pearing within the next 20 years (Mace andLande, 1991).

A study of the mountain tapir

The author first observed the mountain tapirin 1978 in Colombia's Las Hermosas NaturalNational Park (Downer, 1981) and since thenhas surveyed the species in northern Peru,Ecuador, Colombia and western Venezuela,listing about 20 remnant populations for theIUCN Species Survival Commission andWildlife Conservation Society. Field obser-vations formed a major part of the surveys

45

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C. DOWNER

85° 75° 65°

10°

Sta. Marta ^ < fGuajira

Atlantic OceanCaracas

11-,©,

BuenaventuraUCaJi^t * & COLOMBIA

San Cristobal_ VENEZUELA

2 Llanos Orientales

I

Guayaquil'^ ( C o r d i l .

desertl^^F del CondorPERU

I Alitiplano

BRAZIL

Amazon Basin

Occurrence reported in past 10 years

Possible former occurrence but no firmevidence

Firm evidence for former occurrence butdoubtful occurrence now

Reported occurrence but needs verification

Figure 1. Mountain tapir distribution.1. Perija PN, Venezuela, and Catatumbo Bari PNN, Colombia.2. El Tama PN, Venezuela, and Tama PNN, El Cocuy PNN and Pisba PNN, Colombia.3. Chingaza PNN, Colombia.4. Sumapaz PNN, Colombia.5. Cordillera Los Picachos PNN, Colombia.6. Los Nevados PNN and Ucumari RP, Colombia.7. Las Hermosas PNN, Colombia.8. Nevado del Huila PNN, Colombia.9. Purace PNN, Colombia.

10. Cueva de Los Guacharos PNN, Colombia.11. Macizo de Tatama PNN, Colombia.12. Farallones PNN and Munchique PNN, Colombia.13. Galeras Flora and Fauna Sanctuary, Colombia.14. El Angel RE, Ecuador.15. Cotacachi Cayapas RE, Ecuador.16. Cayambe-Coca RE, Sumaco-Napo PN and Antisana RE, Ecuador.17. Cotopaxi PN, Ecuador.18. Llanganatis Bosque Protector and Sangay PN, Ecuador.19. Chimborazo Reserva de Production de Fauna, Ecuador.20. Cajas Area Nacional de Recreacion, Ecuador.21. Podocarpus PN, Cordillera del Condor, and Zamora province, Ecuador; Piura state (national border),

Cajamarca state (national border), Tabaconas-Namballe RE and Cordillera del Condor, Peru.

PN, Parque Nacional; PNN, Parque Nacional Natural; RP, Regional Park; RE, Reserve Ecologica.

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THE MOUNTAIN TAPIR IN THE HIGH NORTHERN ANDES

and involved expeditions into parks and un-protected areas throughout the species's geo-graphical range (Figure 1). This papersummarizes some of the findings from thesesurveys and from an intensive study inSangay National Park in Ecuador.

Sangay National Park

Sangay National Park in the eastern Andeswas designated a World Heritage Site largelybecause of its diversity of ecosystem types(Fundacion Natura, 1992; UNESCO, 1993).Recently expanded to nearly double its formersize, the park covers more than 5180 sq km(Fundacion Natura, 1992; Figure 2). Themountain tapir occurs at higher elevationsthroughout the park, but is more frequent inthe east. Sangay may be the species's most im-portant stronghold.

Apart from the mountain tapir, mammals ofnote include the endangered spectacled bearTremarctos ornatus, white-tailed deerOdocoileus virginianus, red brocket deerMazama rufina, pudu Pudu mephistopheles, rab-bit Sylvilagus braziliensis and a rare endemicsubspecies of guinea-pig Cavia apertea patzelti.Pumas Felis concolor are common, particularlyin areas where their numbers have increasedin response to cattle invasion of the park.

Bird life includes the Andean condor Vulturgryphus, carunculated caracara Phalcobaenuscarunculatus, red-backed hawk Buteo polysoma,white-collared swift Streptoprocne zonaris,sword-billed hummingbird Ensifera ensiferaand the torrent duck Merganetta armata (Maceyet al., 1976; Patzelt, 1989).

Study site in Sangay National Park

The main study site lies between 3300 and4200 m in the Rio Culebrillas sector of the park3 km west of the 5230-m-high Sangay Volcanoand covers approximately 6 x 9 km (Figure 2).To the west, the land is occupied almost ex-clusively by pastoral Puruhaes Indians and afamily hacienda. The Amazon rain forests - ortheir remnants - extend east to the horizon.

© 1996 FFI, Oryx, 30 (1), 45-58

Sangay Volcano, 5230 m, erupting. The Culebrillasstudy area frequently receives ashy fallout, killingits plants and animals, but enriching its soils forlater ecological succession (Craig C. Downer).

The study area encompasses a broad, gentlysloping river plain which extends along theCulebrillas River and its tributaries, includingthe Ramos-Tambo River. The plain is coveredwith grass and scrub while the adjacent slopesrise abruptly and are covered with densecloud forest, giving way to bushy chaparraland treeless paramo at higher altitudes.Periodic flooding allows limited meadow de-velopment in the riverine areas but overgraz-ing by cattle is causing stream bank erosion.

A combination of heavy rains, fragile ashysoils and precipitous slopes make the areahighly prone to earth slippage and erosion.The Sangay Volcano is a major influence onthe successional level of the biotic communityand over the centuries has enriched the soilsof this region with minerals for hundreds of sqkm around the volcano. The soils generallytend to be 'black Andean' with some rockylithosols in the paramos (Macey et al., 1976).The ecological sere is young because whenSangay erupts the lava and ash kill all her-baceous species, many shrubs and trees andmost animal life (Archivos Nuevo Mundo,Madrid; Lewis, 1950). The last major eruptionswere in 1947 and 1988, but the latter did notdeposit ash to the west and the study site wasunaffected.

The vegetation comprises cloud forest andwet paramo (Ramsay, 1992), although for thepurposes of this study five habitat types wereidentified: Andean forest, riverine meadow,chaparral, paramo and pampas. Montane

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C. DOWNER

Atillo Lakes

I New Soutfiern'I Extension ofI Sangay Nat. Park.

I

Park Control^guamote-Macas Road under construction Scale 1:378 000

Amazon Basin

N

Northern Sangay NationalPark (UNESCO World HeritageSite)

: K;"o Macas

1 cm - ~9 km= Approximate

mountain tapir study area.= Pa r k l i m i t

^ = Hut or cabinBoxed area = Main study site

Figure 2. Sangay National Park,Ecuador, showing study site.

cloud forest receives 2000^1000 mm precipi-tation annually and has a temperature rangeof 6-12"C, while subalpine rain paramo re-ceives 1000-2000 mm precipitation a year andhas a temperature range of 3-6°C (Holdridgeet al., 1971; Macey et al, 1976). Snow some-times descends to 3500 m and Sangay peak iscovered to its base during certain months. Iftotal condensation on plants is included, thecloud forests probably receive at least doublethe above figures of moisture (Cavelier, 1991).

Culebrillas receives rain year-round, withstorms arising frequently from the east, al-though relatively less between October toJanuary (Macey et al., 1976; M. Mejia, SangayNational Park superintendent, pers. comm.).

Methods

In 1989 the author started a long-term studyon the mountain tapir, which involved radio-

48

tracking individuals, behavioural observationsand a survey of food plants. The first successat radio-collaring the mountain tapir wasachieved in the Culebrillas sector of SangayNational Park. Three adult mountain tapirswere captured, examined, radio-collared andreleased in December 1989. In July 1990, a fe-male subadult was radio-collared in the samearea. Another three mountain tapirs were cap-tured and tracked, two in September andNovember 1992, and one in December 1993.Two of these were in an area known as LaPlaya near the base of Sangay Volcano,slightly higher (c. 3700 m) than Culebrillas.The other was in a lower (2800 m) montaneforest area to the north of the volcano, knownas El Placer. Signals from the radio collars pro-vided activity readings and allowed a numberof topographical and ecological variables to bemeasured. Time-sequence data were also kept.

Eight vegetation transects were surveyed inthe study area to produce a plant inventory

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and 37 faecal samples collected for dietaryanalysis. Germination experiments were con-ducted on tapir faeces at the InstitutoNacional de Investigacion Agropecuario(INIAP) experimental station south of Quito,where elevation and daily watering simulatedcloud forest conditions. Throughout the studytapirs were observed whenever possible andtheir behaviour recorded, including territorialmarking: rubbing tree trunks, urination, anddung piles.

Results

Vegetation surveys

A total of 264 species in 53 vascular plant fam-ilies were recorded (Table 1). Several of theseare believed to be new species including twospecies of Fuchsia and one new genus relatedto the filmy fern, Hymenophyllum The familyAsteraceae predominated, with 27 genera and59 species, and the Gramineae were well-rep-resented, with 13 genera and 23 species.

The mountain tapir as consumer, germinator andseed disperser

Table 1 shows species that were observed tobe eaten by the mountain tapir, or to germi-nate from its faeces. Figure 3 shows their rank-ing in order of dietary preference ratios(Crawley, 1983). This shows which foodgroups are most sought relative to their pres-ence in the habitat, and is not to be confusedwith absolute quantity of consumption (Figure4). The mountain tapir consumed 205 (77.7 percent) of 264 identified vascular plant speciesfrom the Culebrillas study area. One-hundred-and-ten (84 per cent) of 131 genera from 53vascular plant families were represented inthe diet. The mountain tapir clearly favouredcertain taxa: 35 of 59 species (59 per cent) ofAsteraceae (the most common family) wereeaten and 13 of 14 recorded species ofRosaceae were taken. In absolute quantity,leaves of the silvery-leaved aster tree Gynoxysaccounted for 24 per cent of the diet, whilefern fronds accounted for 19 per cent (Figure4). As indicated by the faecal analysis, these

© 1996 FFI, Oryx, 30 (1), 45-58

A 4-month-old, subadult mountain tapir eatingleaves of Micronea crocea (Melastomataceae) in cloudforests of Ecuador, c. 3000 m (Craig C. Downer).

two categories comprised the greatest bulk ofdietary consumption. Regarding preferenceratios, however, mountain tapirs expend con-siderable effort to select certain plants, for ex-ample the legume Lupinus, which is rare in thehabitat, while other food items are taken withmuch less frequency than would be expectedfrom their abundance in the tapir's habitat(Figure 4).

The other three extant tapirs, Tapirus indicus,T. terrestris and T. bairdii, have also been re-ported to act as seed dispersers for many plantspecies (Williams and Petrides, 1980; Janzen,1982; Fragosa, 1983; Williams, 1984; Bodmer1989, 1991; Rodrigues et al, 1993). The resultsof the present study show that 42 per cent ofthe species of vascular plants eaten in thestudy area (32.6 per cent of those recorded)germinate in the mountain tapir's faeces(Table 1). This is comparable to informationreported for Baird's tapir (Williams, 1984).

Observations of plants germinating fromtapir dung and their proximity to mineralseeps, suggests that the mineral-rich faeces ofthe tapir, particularly those having recentlyvisited mineral licks, provide a nutritive sub-strate for germinating seeds. There is someevidence that the mountain tapir is an obligatesymbiont for certain species of high Andeanplants, such as the Colombian wax palmCeroxylon quindiuense (H. Acosta, pers. comm.).

Like other members of the Perissodactylathe mountain tapir possesses a post-gastric di-gestive system, which allows it to ingest large

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C. DOWNER

Table 1. Identified vascular and non-vascular plants in or near Culebrillas study area, Sangay National Park,that were seen to be eaten by mountain tapirs or were observed to be germinated from their faeces

Class

All fungiMuscopsidaHepaticopsidaFilcopsida

LycopsidaSphenopsida

Liliopsida

Magnoliopsida

Family

AdiantaceaeBlechnaceaeDennstaediaceaeHymenophyllaceaeLomariopsidaceaePolypodaceaeThelypteridaceaeLycopodiaceaeSelaginellaceaeEquisetaceaeBromeliaceaeCyperaceaeGramineaeJuncaceaeIridaceaeAsteraceaeCampanulaceaeValerianaceaeGentianaceaeLoganiaceaeLamiaceaePlantaginaceaeConvolvulaceaeSolanaceaeRubiaceaeScrophulariaceaePolygonaceaeAmaranthaceaeCaryophyllaceaeCruciferaeEricaceaeElaeocarpaceaePrimulaceaeActinidaceaePassifloraceaePiperaceaeCoriaraceaeRanunculaceaeGeraniaceaeOxalidaceaeTropaeolaceaeHalogoraceaeMelastomataceaeOnagraceaePolygalaceaeVitaceaeCunoniaceaeFabaceae

Found

Genera

662211113111125

1322

2711312112252121411111111111221114

Species

89223141521112

122344

5928722316293145811113144312572128

Eaten

Genera

262211113111125

1322

2011212111131111411111111111211114

Species

28223131521112

1023

22

3526422314273122511112143312552128

Germinated from faeces

Genera

01 -2000000000122700611012101121110400010101101210102

Species

012000000000122

1200723012301163110500010101301320103

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Table 1. (continued).

Class Family

RosaceaeSaxifragaceaeLoranthaceaeApiaceaeAraliaceae

TotalsMinus non-vascular plantsTotal vascular plants

Found

Genera

62251

145-14131

Species

144272

283-19264

Eaten

Genera

51121

120-10110

Species

133132

217-12205

Germinated from faeces

Genera

51010

56-353

Species

123020

89-386

quantities of coarse vegetation (Janis, 1976;Bodmer, 1990). Observations in this study ofmountain tapirs eating tough, fibrous leavesof shrubs is consistent with this. The ingestionof a wide variety of plant species may serve toreduce toxicity by avoiding the accumulationof any particular toxin (Crawley, 1983).Mineral seeps and licks may also serve to neu-tralize plant toxins, although this remains tobe proven (Acosta et al., in press).

As Eisenberg (1989) stated for tapirs in gen-eral and as Terwilleger (1978) found for T.bairdii, the current studies found mountaintapirs to be selective, although their choice isbroad. The perambulating foraging style ofthe mountain tapir, as revealed by both radio-tracking and field observation, may relate asmuch to predator avoidance as to optimal for-aging, especially in regions where the puma

population has increased due to the preva-lence of livestock prey or where there ishuman disturbance.

By snapping off branches and topplingtrunks, the mountain tapir makes additionalforage available to smaller herbivores such asthe diminutive Pudu mephistopheles (compareWilliams, 1978, for T. indicus, and Williams,1984 for T. bairdii). Along with its role as aseed disperser, this tends to substantiate themountain tapir's keystone species role for thehigh northern Andes.

The mountain tapir as prey

The study showed that pumas both stalk andkill mountain tapir. Tapir fur was found intwo of 15 puma scats, and puma prints follow-ing the exact course of some mountain tapir

Figure 3. Dietary item preferenceratios for mountain tapirs.Preference ratio: a 1 = preferred;s 1 = avoided, i.e. eaten lessfrequently than encountered.

• 3

1

GamochaetaBuddleja

CortaderiaLuzula

NeurolepsisEricaceae

CarexRubus

Acaena-AlchemJungia

CalceolariaLasiocephalus

GunneraFerns

BromusOreopanax

BrachyotumStachys

EquisetumGynoxysLupinus

]

ai3n~~!~ 1' 1

;

1_J

!:

1

1

j

10 12Preference ratio

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Muhlenbergia — iSatureja —Trifolium —Agrostia —Buddleia —

Rubus —Luzula —

Gaidendron —Juncus —

Hypochoeris —Moss —

Eleocharis —Jungia —

Ericaceae —Cortaderia —Neurolepra —Munnozia —

Bromus —Stachys —

Oreopanax —Acaena-Alchemilla —

Lasiocephalus —Festuca —

Calceolaria —Carex —

Gamochaeta —Lupinus —

Gunnera —Brachyotum —

Equlsetum —Ferns —

Gyrtoxys

10 20

Peroentaae of total for each food item

Figure 4. Relative proportion offood items in the diet ofmountain tapir as determined byfaecal analysis, Culebrillas sectorand adjacent areas, SangayNational Park. No. faecal samples= 37.

prints were discovered on two separate oc-casions in widely separated areas. In the ElPlacer region of Sangay National Park adulttapirs were encountered bearing fresh scars,resembling the arched, thin claw-marks of thepuma. Pumas stalking and killing mountaintapirs have also been reported fromPodocarpus National Park (P. Moran pers.comm.) and in the Ucumari (H. Acosta, pers.comm.) and Las Hermosas (pers. obs.) parksof Colombia. Peyton (1980) documented thekilling and devouring of mountain tapir byspectacled bear Tremarctos ornatus at 2200 m inCajamarca, Peru. The presence of the spec-tacled bear was noted in nearly all areas in-habited by mountain tapirs that the authorvisited between 1978 and 1993.

In the Valladolid region of Loja state, south-ern Ecuador, unconfirmed reports of jaguarPanthera onca ascending to elevations occupied

by the mountain tapir and killing them havebeen received from local hunters (G. Capa,pers. comm.). One major cause may be the de-struction of forest habitats occurring inEcuador's portion of the Amazon Basin(Simpson, 1990), which may also be drivingTapirus terrestris higher, thus causing possiblecompetition with the mountain tapir. Localpeople and zoologists have suggested that hy-bridization occurs between the latter two con-geners (G. Capa, P. Mena, L. Albuja, per.comms).

Behaviour and habitat association

Activity data (c. 3500 telemetric readings)were produced during the study. Tapirs aged3-10 years slept most between midnight anddawn and had a peak of activity between15.00 and 21.00 h. Another consistent pattern

Table 2. Home ranges ofmountain tapirs radio-collaredand tracked in the Culebrillassector, Sangay National Park

Name

SambitaRobertoGabriel

Sex

FMM

Average core range

Age(years)

3-68-103-6

880

Corerange (ha)

1,020846775

Datestracked

12/89-12/9212/89-12/9112/89-12/92

No.locations

205152206

Tracking of these three adults and a subadult female was terminateddue to their being poached. Culebrillas is heavily invaded by cattle.Three additional adult female mountain tapirs are being radio-tracked(June 1995) in two other areas of the park that are still cattle-free.

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was increased activity between dawn and09.00 h. During the rest of the daylight hours,activity decreased slightly, with a restingperiod between 12.00 and 15.00 h. The twosubadults tracked were predominantly inac-tive during all periods of the day, rather thanshowing the above adult pattern. Thesubadult female collared in Culebrillas,showed a more nocturnal pattern.

Activity during daylight and nocturnalhours was approximately equivalent. This isin distinct contrast to a predominantly noctur-nal activity pattern among the other tapirspecies (Fradrich and Thenius, 1968;Terwilliger, 1978; Williams, 1978) and may bedue to the higher, colder altitudes occupied bythe mountain tapir, although T. bairdii hasbeen recorded as high as 1500 m (D. Brooks,pers. comm.) and T. terrestris may ascend tosimilar heights (P. Mena, L. Albuja, O. Linares,pers. comms).

The adult female is often found in partial as-sociation with a male territory, but seems tohave a generally larger home range overall(see Table 2). On several occasions a male andfemale were observed grazing or interactingtogether, but the adults are typically solitary,as is generally true of the Tapiridae (Walker,1964; D'Aulaire and D'Aulaire, 1979).

The mountain tapir has a gestation periodof 13 months (normally single birth) (Bonneyand Crotty, 1979). Field observations inSangay National Park showed that the femalegenerally secludes herself when giving birthand during the first 6 months of nursing thecalf, which is camouflaged with stripes andspots. Tracking indicated that the young staysclose to the mother during the first year of life,particularly during the first 6 months.

Evidence of territorial marking was particu-larly strong in the Purshi sector of the parkand may relate to a higher observed density ofmountain tapirs in this previously virgin habi-tat within the park (now violated by a road)This relates to the longer history of ecosystemstability in this area, where volcanic disturb-ances are not as frequent as in Culebrillas(Macey et al, 1976).

During the study Sangay Volcano fre-quently showed signs of activity and its

© 1996 FFI, Oryx, 30 (1), 45-58

surges affected not only my radio-receptor,but also the tapirs themselves, causing in-creased activity and general restlessness.

Observations, and tracking records fromhundreds of locations, revealed that averagehabitat use in five identified types of habitatwas approximately 28.7 per cent in Andeanforest, 22.9 per cent in riverine meadow, 22.3per cent in chaparral, 19.7 per cent in paramoand 6.4 per cent in pampas. (For the purposesof this study pampas was defined as grasslandcreated by burning and cattle occupation, andchaparral as the scrub ecotone between forestand paramo; riverine meadow occurred in allhabitat types.) Subadults under 2 years spentmost of their time concealed in forest and sel-dom ventured beyond the cover of the bushychaparral. In the wet season greater use wasmade of Andean forest and less of paramo,while during the dry season greater use wasmade of the higher open paramo. The tapir'savoidance of cold by sheltering in forests inthe wet season provides the primary expla-nation for this pattern. The fruiting of shrubbymembers of the Ericaceae and Bromeliaceaeduring the dry season may also help to ex-plain the greater occupation of paramo then(C. Downer, pers. obs; cf. Klein, 1965).

The results of this study suggest that theAndean forest is the most important habitattype for the mountain tapir because of its rolein providing food, shelter and concealment. Itis precisely this ecotype that is being de-stroyed most rapidly (Poore, 1992; Cavelierand Etter, 1995; Associated Press, 1995).

Threats facing Sangay National Park andmountain tapirs

There are three main problems facing SangayNational Park and the mountain tapirs thatlive there: deforestation, encroachment bylivestock and humans, and lack of awarenessof the plight of the tapir.

Deforestation

Ecuador currently has the second highest rateof deforestation in Latin America and lost 48.7

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C. DOWNER

per cent of its forest in the period 1980-89(Simpson, 1990). Sangay National Park isbeing invaded from all sides. Apart fromEcuador, the only other country still contain-ing at least a minimum viable population(Soule, 1986) of 1000 or more (author's esti-mate) tapirs is Colombia, which lost 65.4 percent of its forest during the same period(Simpson, 1990). Deforestation in tropicalmountains has proceeded at faster rates thanin lowland tropical forests according to theWorldwatch Institute (Associated Press, 1995).Currently, an estimated 600-1000 sq km ofmontane forest in the three Andean Cordillerasof Colombia are destroyed each year. If de-struction continues at this rate, the mountaintapir and other high Andean-dependentspecies will soon disappear (Cavelier andEtter, 1995).

Encroachment by livestock and hunters

During the study period the number of cattleestimated by the author to be present in thestudy area and adjacent park areas towardsthe Sangay Volcano rose from 300 to over1000, and this trend shows no signs of slowingdown (V. Alvarez, Sangay National Parksuperintendent, pers. comm., 1995). Because ofthe fragile soils and steep slopes, this is notonly causing habitat conversion but severeerosion in places. Hunting is concomitant withlivestock invasion, as is burning of forest andparamo to clear vegetation. All these activitiesare having devastating effects upon the moun-tain tapirs and their habitats.

Concomitant with lack of park protection,the mountain tapir suffers range constriction,exposure to disease (Hernandez-Camacho,pers. comm.) and increased risk from pre-dation by both humans and pumas, althoughprimarily the former. The tapir is a trophy-species for many collectors, who pay largesums to local people to act as guides (A.Castellanos; A. Zegarra, pers. comms). Thesethreats prevail in most areas where the moun-tain tapir survives and certainly in SangayNational Park.

54

Lack of awareness of the tapir's plight

National legislation is seldom enforced to pro-tect the park, while the Puruhaes people ap-pear to be dominated by a small group whoshow a marked reluctance to consider alterna-tives to increasing dependence on livestockagriculture. This is unfortunate, because themain value of the high Andes lies in its serviceas a watershed and wildlife reserve, similar tothe very strictly protected reserves of the Incas(Vega, 1966; Morrison et ah, 1976).

Conservation requirements

Mountain tapir numbers are falling belowminimum population levels. The isolation ofthe remaining populations is leading to gen-etic bottlenecking and affords the species littlefuture (Downer, 1995a, b). Population andconcomitant habitat fragmentation is clearlyleading rapidly to extinction and remedial ac-tion must be taken soon (Wilcox and Murphy,1985; Janzen, 1986).

Range protection

Mountain tapirs are extremely vulnerable toinvasion of their habitat (IUCN, 1972, 1982).The present author's observations indicatethat after 10-20 per cent of their home rangehas been occupied, tapirs either die (mainlydue to direct human detection and killing) orabandon their territory for safer ground, ifpossible (Figure 5). Extensive, persistent dis-turbance by humans and their livestock invad-ing mountain tapir habitat in the high Andesis the primary cause of the decline of moun-tain tapirs (UN, 1978; Castano-Uribe, 1991;Downer, 1995a, b). This highlights the neces-sity of protecting large areas of natural habitatwhere appropriate buffer zones isolate thecore populations.

It is suggested that zones of limited agrariandevelopment, where ecotourism supplementsfamily income, be developed around moun-tain tapir areas, starting with Sangay NationalPark. These buffer zones would contain natu-ral forests and paramos inhabited by the

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Figure 5. Effect of cattle invasionon home-range size and viabilityof three adult mountain tapirs,radio-tracked in 1990-93,Culebrillas sector, SangayNational Park, d, confirmeddeath of mountain tapir, (cf.Table 2).

2000

f 1800

jjj 1600to

• D 1400

§ 1200

| 1000

m 800

« 600

•5 400c3 200

2000

1800 So

1600

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mountain tapir and associated species. Somesustainable harvest of certain plants and ani-mals could be conducted in the buffer zones.

A programme to encourage organic farmingand use of native plant species to revegetateeroded mountain slopes is in its early stages inthe community of Alao bordering the park(Guinan, 1992; C. Rundo, S. McCarthy, V.Alvarez, pers. comms.).

Habitat corridors linking refuges shouldalso be established whenever possible toensure against inbreeding (Soule, 1986) andefforts undertaken to prevent further fragmen-tation of the mountain tapir's finest reserves.For example, in the Purshi section of SangayNational Park mentioned above, road build-ing threatens to divide and damage one of themost extensive and pristine areas of high-Andean forest and paramo remaining(Downer, 1995b). This area is of criticalimportance both to the mountain tapir and tomany other disappearing species (Downer,1991, 1995b). The Ecuadorean Governmenthas undertaken an Environmental ImpactStudy on the effects of the road and SangayNational Park has been designated as WorldHeritage Site in Danger (UNESCO, 1993).

Education of local people

Amerindian hunters have played a major rolein many large animal extinctions since thePaleolithic, both in North America (Martinand Wright, 1967) and South America. Boththe local haciendas and the local Puruhaes

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'Sambita • Roberto A Gabriel A Cattle numbers

tribal communities are implicated in the de-struction of the high-Andean ecosystem.International commerce in cattle, particularlybulls, raised in the region is also directly in-volved in the destruction of the ecosystem.

Highland Indians use the whole body of themountain tapir: meat, hide and internal or-gans (including intestines made into a soup).They use the hooves and snout as medicinalremedies for epilepsy and heart problems. Thelatter are also sold in urban centres. This com-merce is proving particularly devastating tosurviving populations, such as those in north-ern Peru (A. Zegarra, pers. comm.).

Despite legal protection, there is littleawareness or concern for the endangeredstatus of this animal. Any form of harassment,killing or commerce involving the mountaintapir is illegal in all three countries of its range(IUCN, 1972,1982). However, due to the long-established tradition of mountain tapir hunt-ing and given increasing human pressures,some conservationists consider the implemen-tation of sustainable harvest models in bufferzones outside parks to be a more realistic wayof saving this endangered species (R. Bodmer,L. Salas, D. Brooks, pers. comms).

Certainly no realistic attempt to conservethe mountain tapirs of Sangay National Parkcan or should be made without the involve-ment of the local people. Conservation of thepark should be part of a co-ordinated pro-gramme, including the restoration of erodedsoils by agroforestry programmes using nativetrees and encouragement of ecotourism by

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which local people would benefit directlyfrom their proximity to one of the world'smost spectacular wildlife areas (Guinan, 1992).

Sangay National Park and its tapirs couldbe saved by means of a contractual agreementwith local people, who would receive rev-enues generated by the park in exchange forwithdrawing their cattle, ceasing to huntwithin the park, and acting as park guides,hosts and protectors. It is critical that such asolution be implemented immediately.

Maintenance of population strongholds

From telemetric data collected betweenDecember 1989 and December 1992, theauthor estimated an average horizontal homerange of 880 ha for the adult mountain tapir(Table 2). The highly accentuated topographyof the habitat actually gives the mountaintapir a much greater surface area and veg-etation resource (S. Matola, pers. comm.).Taking into account a conservative, inter-malemean range overlap of one-third, 587 ha rep-resents the exclusive area required for eachpair of breeding adults and their subadult off-spring at any given time. Multiplying this fig-ure by 500 (given sharing of territory by amale-female pair) gives a total of 293,500 ha,which would be the total area of suitable habi-tat needed for one minimum viable popu-lation of approximately 1000 breeding adultsof equal sex ratio (based on interpretation ofSoule, 1986). An emphasis on protecting areasof suitable, contiguous habitat of at least thissize and as nearly circular in shape as is con-sistent with natural wilderness boundaries isneeded in order to make the most efficient useof conservation action (Boyce, 1992).

Much of Sangay National Park's 5180 sq kmha lies above 2000 m and the volcanic peakstower over 5000 m. This park could probablysupport a viable population of mountaintapirs; although 1000 is considered by somepopulation biologists to be insufficient for thelong-term survival of such a slowly repro-ducing mammal (Newmark, 1986). Given thelevel of cattle and hunter invasion with the as-sociated habitat destruction in Sangay, the

56

critical situation of the species can be well ap-preciated. The park could serve as a modelexample to other nations of effective conser-vation if a plan is successfully proposed andexecuted. Moreover, it could serve as a sourceof animals for restocking regions formerly in-habited by tapirs (Griffith et ah, 1989). It ap-pears that Venezuelan park authorities wouldlike to accomplish reintroductions of moun-tain tapirs (E. Yerena, pers comm.).

Conclusion

Experiments and field observations haveshown that the mountain tapir plays a vitalecological role in the high-Andean ecosystem.Its survival depends upon the preservationand restoration of the forest and paramos ofthe northern high Andes, which serve as vitalwatersheds for all the ecosystems lying belowthem, including those upon which humansdepend. The mountain tapir is a large, easilyrecognizable animal and its image appears onthe logo of Sangay National Park. It would bean ideal 'flagship' species for the conservationof both the park and the high northern Andesas a whole. I recommend that, as part of an in-ternational effort, a survey of the remaininghigh-Andean forest and paramo habitats beundertaken, including by satellite, in order toplan essential conservation efforts for themountain tapir. These would benefit the manyother species, including humans.

Acknowledgements

I wish to thank the following individuals and organ-izations that have helped over the past few years.The generous aid of NYZS-The WildlifeConservation Society is much appreciated and I amgrateful for the considerable help of the EcuadoreanNational Parks and Wildlife Department and thevarious officials and park rangers and willing andable Puruhaes guides and assistants from the villageof Alao. I am indebted for the very special support,faith and generosity of the Georgia Earth Alliance ofAtlanta and its founder Gloria Wilkins. I greatly ap-preciate the support of the World NatureAssociates, Silver Spring, Maryland, ChicagoZoological Society for their contributions and con-tinued interest, the Los Angeles Zoological

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Foundation and the San Diego Zoological Society. Ithank the High Desert Rainforest Group of Reno,Nevada, for their benefit reggae concert and contri-butions, the Nevada Humane Society, Ed Chipp,Ruska Bogdanovich, Chuck and Donna Steadman,and Ria Katz for their faith, concern for the tapirs,and generous contributions. Bernard Peyton pro-vided valuable advice and equipment and the USPeace Corps provided me with my first encounterwith the mountain tapirs. I thank the MissouriBotanical Garden Herbarium for help in identifi-cation, as well as the Catholic University Herbariumand INIAP of Quito. Also in Ecuador, thanks aredue to the Escuela Politecnica Nacional and to theecological groups Ecociencias and Cordavi of Quito,Banos 2000 of Tungurahua, and Arcoiris of Loja. Iowe a special debt of gratitude to the School ofBiological Sciences at the University College ofNorth Wales, Bangor, particularly Ralph Oxley,Terry Williams and Rob McKay, for their help indata analysis. I thank my assistants, biologistsAlfredo Lopez and Armando Castellanos-Penafiel,for their courage and faith. I express my gratitude tomy parents, Robert C. Downer and Alice G.Downer, who have greatly aided me, both morallyand financially.

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Craig C. Downer, School of Biological Sciences,University College of North Wales, Bangor,Gwynedd LL57 2UW, UK. Current address: PO Box456, Minden, NV 89423, USA.

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