analysis of hunting options good taboos colding 1998

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Ecological Modelling 110 (1998) 5 – 17 Analysis of hunting options by the use of general food taboos Johan Colding * Department of Systems Ecology, Stockholm Uni6ersity, S -106 91 Stockholm, Sweden Abstract A hypothetical model was built, using the STELLA II software program, to test several hunting options for a human hunting group. Different outcomes of possible hunting modes are analysed, such as a change in hunting rate, prey hunted, or species avoided or not avoided by taboos. The model consists of five sectors that reflect a short food chain in an upper Amazonian ecosystem. There is a vegetation sector, a predator sector, and two sectors consisting of browsers and grazers. The last sector represents a human group, known as the Ecuador Achuar. The critical factor analysed is how differences in hunting rate affect a target resource, and how this resource may be affected by general food taboos. The major results of the model are that general food taboos may not be an adaptive short term strategy for hunters, but that a ‘moderate’ hunting mode may be the most effective option for the human group. Since the model is a simplification of the real world, no general conclusions for management should be drawn from the results. © 1998 Elsevier Science B.V. All rights reserved. Keywords: General food taboos; Hunting options; Amazone ecosystems 1. Introduction Many traditional societies employ food taboos on animal species for a number of reasons. Some researchers suggest that there are nature manage- ment motives behind them (Rappaport, 1967, 1968; Reichel-Dolmatoff, 1976; McDonald, 1977; Johaness, 1978; Ross, 1978; Harris, 1979), while others resent any such ecological motives (Rea, 1981; Edgerton, 1992). Hunting among some tra- ditional groups may be conducted in a highly efficient way, consistent with predicators of forag- ing theory (Alvard, 1993, 1994). While species are thus pursued with short-term harvest rate maxi- mization, one may very well ask why some native hunting groups employ food taboos. McDonald (1977), in a study of 11 South American tropical groups, suggests that taboos may function as con- servation mechanisms for reducing the hunting pressure on larger mammals in environments where such species are low in abundance. General food taboos—applying to all members within a community — may play a role in biodiversity con- * Present address: The Beijer International Institute of Eco- logical Economics, The Royal Swedish Academy of Sciences, PO Box 50005, S-104 05 Stockholm, Sweden. Tel.: +46 8 6739500; fax: +46 8 152464; e-mail: [email protected]. se 0304-3800/98/$19.00 © 1998 Elsevier Science B.V. All rights reserved. PII S0304-3800(98)00038-6

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  • Ecological Modelling 110 (1998) 517

    Analysis of hunting options by the use of general food taboos

    Johan Colding *

    Department of Systems Ecology, Stockholm Uni6ersity, S-106 91 Stockholm, Sweden

    Abstract

    A hypothetical model was built, using the STELLA II software program, to test several hunting options for ahuman hunting group. Different outcomes of possible hunting modes are analysed, such as a change in hunting rate,prey hunted, or species avoided or not avoided by taboos. The model consists of five sectors that reflect a short foodchain in an upper Amazonian ecosystem. There is a vegetation sector, a predator sector, and two sectors consistingof browsers and grazers. The last sector represents a human group, known as the Ecuador Achuar. The critical factoranalysed is how differences in hunting rate affect a target resource, and how this resource may be affected by generalfood taboos. The major results of the model are that general food taboos may not be an adaptive short term strategyfor hunters, but that a moderate hunting mode may be the most effective option for the human group. Since themodel is a simplification of the real world, no general conclusions for management should be drawn from the results. 1998 Elsevier Science B.V. All rights reserved.

    Keywords: General food taboos; Hunting options; Amazone ecosystems

    1. Introduction

    Many traditional societies employ food tabooson animal species for a number of reasons. Someresearchers suggest that there are nature manage-ment motives behind them (Rappaport, 1967,1968; Reichel-Dolmatoff, 1976; McDonald, 1977;Johaness, 1978; Ross, 1978; Harris, 1979), whileothers resent any such ecological motives (Rea,

    1981; Edgerton, 1992). Hunting among some tra-ditional groups may be conducted in a highlyefficient way, consistent with predicators of forag-ing theory (Alvard, 1993, 1994). While species arethus pursued with short-term harvest rate maxi-mization, one may very well ask why some nativehunting groups employ food taboos. McDonald(1977), in a study of 11 South American tropicalgroups, suggests that taboos may function as con-servation mechanisms for reducing the huntingpressure on larger mammals in environmentswhere such species are low in abundance. Generalfood taboosapplying to all members within acommunitymay play a role in biodiversity con-

    * Present address: The Beijer International Institute of Eco-logical Economics, The Royal Swedish Academy of Sciences,PO Box 50005, S-104 05 Stockholm, Sweden. Tel.: 46 86739500; fax: 46 8 152464; e-mail: [email protected]. se

    0304-3800:98:$19.00 1998 Elsevier Science B.V. All rights reserved.

    PII S 0304 -3800 (98 )00038 -6

  • J. Colding : Ecological Modelling 110 (1998) 5176

    Fig. 1. Approximate location of the Ecuador Achuar territory (marked in grey).

    servation. For example, Colding and Folke(1996), found that a number of threatened popu-lations of species, including endemic and keystonespecies, benefit from such taboos.

    The purpose of this modelling exercise withSTELLA II, has been to analyse possible relationsbetween general food taboos and different hunt-ing modes. The STELLA II program may be asuitable tool for an investigation of this kind.While real world data on wild populations areavailable in the literature (see Emmons, 1990;Nowak, 1991; Redford and Eisenberg, 1992), it isnevertheless problematic to generalise from suchdata. For example, data on numbers of densityare measured in habitats where species thrive. Togeneralise from such data, from one habitat toanother, may be highly awkward. Among otherthings, this is due to the differences in the hetero-genity of landscapes within and between ecosys-tems. Perhaps more realistic data for thepresented model could be obtained using site-spe-

    cific data from field studies. Unfortunately, thishas not been possible. Thus, the estimates usedhere are highly artificial.

    The Ecuador Achuar represents the humangroup of this model. This group, inhabiting a vastregion at the Ecuador:Peru border (see Fig. 1),has been well studied by DeScola (1986). TheEcuador Achuar impose general food taboos onseveral species of mammals. They abstain fromthe consumption of all carnivorous mammals andregard several other such species as inedible, refer-ring to them as sickening meat (DeScola, 1986).Among such species is the rodent Capybara, Hy-drochoerus hydrochoerus, which is widely huntedby other neotropical forest groups (see Redford,1993; Bodmer 1994). However, other species ofrodents are hunted, e.g. agoutis (Agouti paca) andacouchis (Dasyprocta and Myoprocta). Ungulates,such as Brocket deer (Mazama Americana) andTapirs (Tapirus terrestris and T. pinchaque) areavoided by general food taboos, due to reincarna-tion beliefs (DeScola, 1986).

  • J. Colding : Ecological Modelling 110 (1998) 517 7

    Fig. 2. The basic structure and key actors of the model.

    narios are presented in Section 3. Individualmodel runs will also be discussed in Section 3,while an overall discussion will follow in Section4.

    2. Basic structure and data

    The main sectors and key actors of the modelare presented in Fig. 2. This figure illustrates thebasic food chain investigated. The population ofjaguars is regarded as the top-predator in thisextremely simplified ecosystem. Jaguars prey onboth the rodent and tapir populations respec-tively, that, in turn, feed from the vegetationsector. Although all carnivore species areavoided by the Achuar hunter, the jaguars inthis model are thought of representing a fractionof all the carnivores in this ecosystem. The sameholds true for both the rodent sector and the

    The results obtained in this study are highlyartificial and should not be taken as indicatorsof the Ecuador Achuar hunting mode. Themodel is far too incomplete for any such infer-ences to be made. This complies for both thestructure of the ecosystem, as well as for thevarious populations of species described in themodel. Some possible hunting modes and sce-

    Fig. 3. A model for examining different hunting options for a human group.

  • J. Colding : Ecological Modelling 110 (1998) 5178

    Table 1Sensitivity runs on population stocks of the model, using STELLA II

    3: Vegetation 2: Rodent 3: RodentMonths 1: Rodent1: Vegetation 2: Vegetation

    9 000 000 9000 18 000 27 0000 3 000 000 6 000 00011 955 12 00950 6 617 864 6 618 420 6 618 413 12 035

    11 79111 772 11 785100 6 605 2776 605 182 6 605 27814 512 14 521150 6 557 093 6 557 171 6 557 170 14 52512 272 12 279200 6 620 185 6 620 226 6 620 225 12 281

    1: Jaguar 3: Jaguar2: Jaguar3: Tapir1: Tapir 2: Tapir10 250 300 600 40900

    28282850 651300 541566 28 28 28100 294 511

    31 3131150 561335 5302727200 338 503 521 27

    1: Tapir* 2 Tapir* 3: Tapir*0 900300 600

    50 218 485 603100 48767 410

    463150 0 400200 3920 333

    An increase in the rate of tapirs per jag from RANDOM (0, 0.4, 0.4) to RANDOM (0, 0.5, 0.5) is marked as * in the table.

    tapir sector. The former represents mainly alumping of larger rodents that the EcuadorAchuar pursue, and the latter a fraction of theungulates avoided by general food taboos.

    2.1. Climatic conditions of the ecosystem

    The ecosystem is a simplified version of theEcuador Achuar territory, situated at theEcuador:Peru border. It lies north of the Pastazariver, and covers an area of about 9000 km2 (seeFig. 1). The region has a typical equatorial cli-mate, constantly humid, no dry season, and amonthly rainfall always over 60 mm. It is situ-ated 2 south of the equator, where days andnights are nearly the same length, and tempera-tures remain constant throughout the year. De-spite its proximity to the Andean barrier, thisecosystem is not directly affected by the specialmeteorological conditions of the foothills. Meanannual daytime temperatures vary between 24and 25C. The annual average low temperaturefluctuates between 19 and 20C, depending onaltitude, and the average annual high tempera-

    ture is between 29 and 31C. It is marginallywarmer from October to February.

    The average annual rainfall is no more than3000 mm for the highest latitudes, and not lessthan 2000 mm for the lowest. However, rainfallvaries from year to year. Periods of too little ortoo much precipitation have no noteworthy ef-fect on the vegetative activity of wild and culti-vated plants, since the duration is too short tohave any long-term influence. However, animalpopulations may be affected by this variation,since droughts rapidly dries up secondarybranches of rivers, normally filled with water,affecting the fish that live there, and other spe-cies of animals that normally receive water atthese places. In the opposite case, heavy continu-ous rainfall tends to accelerate the decompositionof organic bedding, rapidly destroying the fruitand seeds eaten by large terrestrial herbivoressuch as tapirs and peccaries.

    The ecosystem is in a state of dynamic equi-librium, as its system of energetic exchange oper-ates theoretically in a closed circuit (Odum,1971). The organic matter and minerals are con-

  • J. Colding : Ecological Modelling 110 (1998) 517 9

    Fig. 4. A case with a zero hunt on all the three populations of species. The ecosystem is in balance.

    tinually recycled by a complex network of micro-organisms and specialised bacteria. Given thatalmost 90% of the nutrients of tropical forestecosystems are stored in the standing plantbiomass (Begon et al., 1990), this closed circuitsystem is sensitive to larger clearings and defor-estation activities.

    2.2. Description of model sectors

    A detailed structure of the model, describingthe various in- and outflows between and withinstocks and sectors is presented in Fig. 3. Themodel has been run at a monthly time step for200 months. This length was chosen on the basisthat it would allow a human population to adjustto changes in hunting modes, since possible effectsof a taboo would thus be known in a fairly shortnotice of time.

    2.2.1. The 6egetati6e sectorBased on the climatic conditions described in

    Section 2.1 the vegetation sector is considered tobe largely unaffected by climatic variability. Nu-trients are thus tightly recycled and closely associ-ated to the standing living biomass. This has beenaccounted for in the model by the design of anutrient cycle circuit (see Fig. 3). The vegetationsector was constructed to be robust, and largelyunaffected by different levels of populationchanges within other sectors. Sensitivity runs indi-cate that population levels of the herbivores (ro-dents and tapirs) are constrained by theirrespective carrying capacities, and do not pose athreat to the vegetation, even if initial numbersgreatly overshoot carrying capacity levels (seeTable 1). This sector is thought to consist largelyof evergreen leaves, flowering plants, fruits andseeds, and constitutes the first trophic level of thisecosystem (for information on input data usedthroughout Section 2.2, see equations in the Ap-pendix).

    2.2.2. The rodent sectorThe rodent sector of the model represents ro-

  • J. Colding : Ecological Modelling 110 (1998) 51710

    Fig. 5. A case where rodents are hunted at the hunting rate of 0.17, equal to an average of pursued rodents by other neotropicalforest groups (Redford 1993). Jaguars and tapirs are not hunted at all.

    dents that may be hunted by the EcuadorAchuar (mainly those of a weight between 2and 10 kg). Species under consideration includeAgouti paca, and Dasyprocta spp. and My-oprocta spp. They represent part of the secondtrophic level. Rodents are widely hunted amongtropical Amazonian groups (Redford, 1993) andare important sources of protein in the EcuadorAchuar diet.

    Initial population of rodents were set at18 000. This estimate is based on adding thedensity numbers for the above rodent species(see estimates of Nowak (1991), Redford (1993),and Bodmer (1994)) (Density of Agouti is 5.1per km2, for Dasyprocta and Myoprocta 6.1 perkm2.) For an ecosystem 9000 km2 in size, theestimate will end up being about 110 000 ro-dents, which is about 12.2 individuals per km2.This is considered as an extremely high estimate.Densities for rodents such as hares in Sweden,

    are considered very high at 4 per km2 (ibid),which is about 1:6 of the number above. Forthis reason the estimated number on density hasbeen set at about 2 rodents per km2, whichgives a total value of about 18 000 rodents.Mean litter size of rodents was set at four(Eisenberg, 1981), with 34 litters per year(ibid). Based on these numbers the specific birthrate was set at about 6.0, which gives a monthlybirth rate of 0.5. The death rate was estimatedby using the survivorship curve based on that ofOdum (1983), and set at a monthly rate of 0.46.

    2.2.3. The tapir sectorTapirs are both grazers and browsers. They

    represent part of the second trophic level of thismodel. The Ecuador Achuar employ a general foodtaboo on the killing of tapirs. Densities for tapirshave been estimated at 0.4 per km2 (Bodmer, 1994).As was the case for rodents, 1:6 of this estimate has

  • J. Colding : Ecological Modelling 110 (1998) 517 11

    Fig. 6. A case displaying the maximum hunting rode on rodents. At a hunting rate of 0.92, both rodents and jaguars are severelyaffected. Above this rate these populations go extinct. Jaguars and tapirs are not hunted at all.

    been used as an input value for this model. There-fore, the initial population of tapirs is set at about600. For the calculation of birth and death rates,the survivorship curve of the species of Black-taildeer was used (see Odum, 1983). This species is aherbivore of approximately the same size, and wasassumed to have a survivorship curve of the samekind as tapirs. Litter size of tapirs was set at one,with an average of about one litter per year(Eisenberg, 1981). Monthly birth rate wascalculated at 0.04, and monthly death rate at0.017.

    2.2.4. The jaguar sectorJaguars represent the top-predator of this

    ecosystem, preying on both rodents and tapirs.No information on rates of predation, birth anddeath were found in the literature. Ecologicaldensity of jaguars is estimated at 0.013 per km(Redford and Eisenberg, 1992). One sixth of thisvalue sets the initial jaguar population at about 20individuals.

    2.2.5. The Achuar sectorThe Achuar of this particular region of

    Ecuador has a steady population of about 1000(DeScola, 1986). The group has many options forhunting which will be analysed below.

    3. Scenarios and individual outcomes of runs

    The results of the different runs are presented inFigs. 49. Following are three different scenarios,each representing possible hunting options for theEcuador Achuar population.

    3.1. Scenario one

    As has been stated, the Ecuador Achuar obeygeneral food taboos on the hunting of tapirs andjaguars. In this first scenario, this has been ac-counted for by setting hunting rates on thesespecies at zero (see Figs. 46). In this scenario,consisting of three runs, the Achuar only huntrodents. Different outcomes will be obtained, de-

  • J. Colding : Ecological Modelling 110 (1998) 51712

    Fig. 7. A case where jaguars are pursued at a hunting rate of 0.0036, which turned out to be the maximum rate allowed for notcreating a collapse of the jaguar population. Rodents are pursued at a hunting rate of 0.17. Tapirs are not pursued at all.

    pending on the different hunting rates of rodents.Fig. 4 represents a case where the Achuar em-

    ploy a zero value in the hunting rate of rodents.This run indicates that the model ecosystem is inbalance.

    In the second run (see Fig. 5), the hunting rateon rodents employed by the Achuar was set at170 per month, which is about the normal hunt-ing rate on rodents by other neotropical forestgroups (Redford and Robinson, 1987).

    In the third run (see Fig. 6), the numbers ofrodents hunted was set at 920 per month, whichwas the highest number of rodents that could behunted without collapsing this population. If therodent population collapses it will make thejaguar population collapse as well, since jaguarspredominantly prey on rodents in this model.These results indicate that the Ecuador Achuarsector of the model may hunt rodents sustainablyat a rate of \0.17 to 0.92.

    3.2. Scenario two

    If rodents are hunted at a rate of 170 permonth, and the taboo on tapirs is still employedbut the taboo on jaguars is not obeyed, the runpresented in Fig. 7 is obtained. In this run thehunting rate on jaguars is set at 3.6 per month,which is the maximum hunting rate allowedwithout collapsing the population.

    3.3. Scenario three

    In this last scenario two different runs weremade. In Fig. 8 the taboo on the hunting ofjaguars is observed by the hunters, but not thetaboo on tapirs. The result from this run indi-cates that if tapirs are hunted at a rate of twoper month, or greater, the number of the tapirpopulation will gradually decline and eventuallygo extinct. Additional runs were made in which

  • J. Colding : Ecological Modelling 110 (1998) 517 13

    Fig. 8. A case displaying a zero hunting rate of jaguars (by taboo). With a hunting rate on tapirs of 0.002, the population declinessteadily over time. Rodents are hunted at the rate of 0.17.

    it was possible for hunters to hunt all three popu-lations of species simultaneously at rather highhunting rates in a sustainable manner. For exam-ple rodents, jaguars and tapirs, may all be pur-sued at corresponding hunting rates of 170, 3 and2 per month, without threatening the long timesurvival of these populations.

    Fig. 9 shows that over-hunting of jaguarsmay be a rational hunting option for groupsthat hunt species, such as rodents and ungu-lates. The interspecific competition between thehuman hunters and jaguars is thus reduced. Thissituation does not pertain to the Ecuador Achuar.

    4. Discussion

    There are several limitations to this exercise.For example, input data on consumption ratesare simply estimated without any real refer-ences. The system was sensitivity tested andanalysed in this regard, which demonstrated

    that if the numbers of tapirs taken per jaguarare slightly increased from a random value of0.4 to 0.5, it could have a dramatic impact ona tapir population of about 300 individuals (seeTable 1). At this level the whole tapir popula-tion collapsed and became extinct.

    Estimates of vegetation, below and abovevalues in the model, revealed that carrying ca-pacity is set at about 6.6 million vegetativeunits. Sensitivity runs on the rodent populationat numbers below or above the estimated popu-lation size, showed that the rodent populationestablished itself close to a carrying capacity atabout 12 00014 500 in most runs (see Table 1).

    Despite the limitations of this exercise, whatcan be said about the overall results of thismodelling exercise? Firstly, the results indicatethat general taboos imposed on species ofjaguars and tapirs protect them from becomingextinct. However, the taboos of this modelseem not to have any hidden ecological func-tions by, e.g. enhancing other populations of

  • J. Colding : Ecological Modelling 110 (1998) 51714

    Fig. 9. A case showing that it may be advantageous for hunters to pursue a predator species such as the jaguar. This may reducethe interspecific competition between two populations feeding on the same resource. Hunting rate on jaguars is set at 0.0036, onrodents at 0.17, and on tapirs at 0.002.

    species pursued by the Ecuador Achuar. It seemsto be more efficient to pursue jaguars wheneverencountered, than not to. As Fig. 8 also indicates(and several other runs have demonstrated), it maybe most efficient to pursue all three species ofmammals at moderate hunting rates.

    5. Conclusion

    General food taboos may be ecologically adap-tive for hunters in ways which are not disclosed bythe nature of this simplified model. The STELLAmodel used here indicates that a moderate huntingmode may be the most effective option for theneotropical group of this model. However, noinferences should be drawn based on this result fortropical forest groups such as the Ecuador Achuar.In the model of this paper, general food taboos donot increase the yield for hunters of species whichare not surrounded by taboos. Cultural reasons,

    based on mythologies and perceptions of reality,may be the reason behind these taboos in the realworld (see also DeScola, 1986). On the other hand,for the preservation of species that may go extinctfrom over hunting, the use of a moderate huntingmode, or the use of general food taboos, seem tobe viable options. Any final conclusion on theserelationships cannot be drawn from this model.

    Acknowledgements

    I would like to thank Professor Robert Costanza.at the Institute for Ecological Economics, Univer-sity of Maryland, for giving me valuable advice andfeedback in the construction of this model. Withouthis insistence this paper would never have beencompleted. Warm thanks also to Professor CarlFolke, at the Beijer Institute of Ecological Econom-ics, Stockholm, for valuable suggestions andthoughtful considerations.

  • J. Colding : Ecological Modelling 110 (1998) 517 15

    Appendix A. Equations for the model

    Human Human

    group(t)Human

    group(tdt)INIT Human

    group=1000 Jaguar

    kill

    per

    human=0 Rodent

    kill

    per

    human=0 Tapir

    kill

    per

    human=0Jaguars Jaguar

    pop(t)=Jaguar

    pop(tdt)

    +(births

    jaguardeaths

    jaguar)*dtINIT Jaguar

    pop=20INFLOWS:

    births

    jag=Jaguarpop*(birth

    frac+jag

    of

    tap+birth

    fract

    jag

    of

    rod)

    *(1(Jaguar

    pop/car

    cap

    jag))OUTFLOWS:

    deaths

    jaguar=Jaguarpop*(death

    frac

    jag

    of

    rod+death

    fract

    jag

    of

    tapirs)+Human

    group*Jaguar

    kill

    per

    human car

    cap

    jag=40* birth

    fract

    jag

    of

    rod=GRAPH(Rodent

    pop)(0.00, 0.0265), (100, 0.0285),(200, 0.033), (300, 0.0375),(400, 0.0425), (500, 0.048),(600, 0.0515), (700, 0.0575),(800, 0.059), (900, 0.06),(1000, 0.06)* birth

    frac

    jag

    of

    tap=GRAPH(-Tapir

    pop)(0.00, 0.13), (10.0, 0.145),(20.0, 0.174), (30.0, 0.205),

    (40.0, 0.245), (50.0, 0.305),(60.0, 0.35), (70.0, 0.37),(80.0, 0.383), (90.0, 385),(100, 0.385)* death

    fract

    jag

    of

    tapirs=GRAPH(Tapir

    pop)(0.00, 0.11), (60.0, 0.105),(120, 0.09), (180, 0.075),(240, 0.06), (300, 0.05),(360, 0.04), (420, 0.032),(480, 0.0235), (540, 0.016),(600, 0.00)* death

    frac

    jag

    of

    rod=GRAPH(Rodent

    pop)(0.00, 0.91), (1800, 0.855),(3600, 0.76), (5400, 0.625),(7200, 0.45), (9000, 0.295),(10 800, 0.18), (12 600, 0.105),(14 400, 0.06), (16 200, 0.025),(18 000, 0.0005)

    Rodents Rodent

    pop(t)=Rodent

    pop(tdt)+(birthsdeaths)*dtINIT Rodent

    pop=18 000INFLOWS:

    births=Rodent

    pop*birth

    fraction*(1Rodent

    pop/car

    cap

    rod))OUTFLOWS:

    deaths=Jaguar

    pop*rod

    per

    jaguar+Rodent

    pop*death

    fraction+Human

    group*Rodent

    kill

    per

    human car

    cap

    rod=36 000 rod

    per

    jaguar=RAN-DOM(0.100, 50)* birth

    fraction=GRAPH(Vegeta-tion)(0.00, 0.12), (700 000, 0.165),(1.4e+006, 0.23), (2.1e+006, 0.285), (2.8e+006, 0.33),(3.5e+006, 0.395), (4.2e+

  • J. Colding : Ecological Modelling 110 (1998) 51716

    006, 0.43), (4.9e+006, 0.455), (5.6e+006, 0.485), (6.3e+006, 0.5), (7e+006, 0.5)* death

    fraction=GRAPH(Vegetation)(0.00, 0.815), (80 000, 0.63),(160 000, 0.49), (240 000, 0.395),(320 000, 0.32), (400 000, 0.27),(480 000, 0.24), (560 000, 0.215),(640 000, 0.213), (720 000, 0.215),(800 000, 0.213)Tapirs Tapir

    pop(t)=Tapir

    pop(tdt)+(births

    tapirdeaths

    tapir)*dtINIT Tapir

    pop=600INFLOWS:

    births

    tapir=Tapir

    pop*birth

    fract

    tapir*(1

    (Tapir

    pop/car

    cap

    tapirs))OUTFLOWS:

    deaths

    tapir=Jaguar

    pop*tapirs

    per

    jag+Tapirpop*death

    fract

    tapir+Human

    group*Tapir

    kill

    per

    human car

    cap

    tapirs=1200 tapirs

    per

    jag=RAN-DOM(0, 0.4, 0.4)* birth

    fract

    tapir=GRAPH(Vegeta-tion)(0.00, 0.0005), (80 000, 0.0185),(160 000, 0.0275), (240 000, 0.033),(320 000, 0.0358), (400 000, 0.0388),(480 000, 0.0403), (560 000, 0.0425),(640 000, 0.0433), (720 000, 0.0438),(800 000, 0.044)* death

    fract

    tapir=GRAPH(Vegeta-tion)(0.00, 0.9), (600 000, 0.8), (1.2e+006, 0.7), (1.8e+006, 0.4), (2.4e+006, 0.0498), (3e+006, 0.0305),(3.6e+006, 0.019), (4.2e+

    006, 0.016), (4.8e+006, 0.0155),(5.4e+006, 0.015), (6e+006, 0.0145)

    Vegetation Vegetation(t)=Vegetation(tdt)+(regenerationconsumption)*dtINIT Vegetation=6 000 000INFLOWS:

    regeneration

    =seeding+(Vegetation*regen

    per

    plant)

    *(1(Vegetation/car

    cap

    veg))OUTFLOWS:

    consumption=Rodent

    pop*veget

    per

    rodent+Tapir

    pop*veget

    per

    tapir car

    cap

    veg=7 000 000 seeding=1 veget

    per

    rodent=20 veget

    per

    tapir=20* nutrients=GRAPH(Vegetation)(0.00, 0.015), (600 000, 0.07),(1.2e+006, 0.135), (1.8e+006, 0.23), (2.4e+006, 0.365), (3e+006, 0.565), (3.6e+006, 0.745),(4.2e+006, 0.815), (4.8e+006, 0.855), (5.4e+006, 0.88), (6e+006, 0.9)* regen

    per

    piant=GRAPH(nutri-ents)(0.00, 0.085), (0.1, 0.125),(0.3, 0.145), (0.3, 0.185),(0.4, 0.215), (0.5, 0.275),(0.6, 0.385), (0.7, 0.505),(0.8, 0.645), (0.9, 0.705),(1, 0.735)

    References

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