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118 ELEPHANTS AND RHINOCEROS

Int. Zoo Yb. (2006) 40: 118–129 © The Zoological Society of London

An integrated management strategy for theconservation of Eastern black rhinocerosDiceros bicornis michaeliin KenyaR. AMIN1, B. OKITA-OUMA2, K. ADCOCK3, R. H. EMSLIE3, M. MULAMA2 &P. PEARCE-KELLY1

1Zoological Society of London, Regent’s Park, London NW1 4RY, United Kingdom,2Kenya Wildlife Service, PO Box 40241-00100, Nairobi, Kenya, and 3IUCN/SSCAfrican Rhino Specialist Group, Box 1212, Hilton 3245, KwaZulu-Natal, South AfricaE-mail: [email protected]

At time of writing Kenya holds c. 84% of theremaining in situ population of Critically Endan-gered Eastern black rhinoceros Diceros bicornismichaeli. The Kenyan 5 year Black Rhino Conser-vation and Management Strategy has given highestpriority to the biological management of the subspe-cies in order to help achieve and maintain rapidmetapopulation growth to increase rhinoceros num-bers. Specific training and capacity building in mon-itoring of rhinoceros, from collection of field data tostatus reporting, have been implemented. Proceduresto assess ecological carrying capacity for Easternblack rhinoceros habitat have been developed tohelp establish new viable populations and to opti-mize the management of existing sanctuaries, someof which face loss of suitable habitat owing to com-petition for resources between rhinoceros and otherbrowsers. The management objective is to maintainrhinoceros populations below the ecological carryingcapacity of a reserve in order to increase reproduc-tion in ‘donor’ populations as well as the trans-location of animals to create new populations withthe potential for rapid growth. The conservationvalue of establishing site-level support relationshipsis also discussed. Kenya, for the first time since the1970s, achieved a growth rate of �5% over theperiod 2004 and 2005 (the national goal is to increasenumbers by at least 5%).

Key-words: biological management, capacitybuilding, eastern black rhinoceros, ecologicalcarrying capacity (ECC), population estimation,rhinoceros monitoring, status reporting

In 1970 the Eastern black rhinocerosDiceros bicornis michaeli population inKenya was c. 20 000 but by the late 1980sthis population numbered �400 animals(Foose et al., 1992; Anon., 1993; Emslie &

Brooks, 1999; Brooks, 2002; Mulama &Okita-Ouma, 2004) (Fig. 1). At the end of2003 the c. 450 Eastern black rhinocerosin Kenya (c. 84% of the total in situ popu-lation) are spread across 14 sanctuaries,reserves and national parks, with habitatsranging from dense forest to semi-aridscrubland (Fig. 2; Plate 1). The Easternblack rhinoceros is considered CriticallyEndangered by IUCN (2004). This drasticdecline was initially the result of culling inorder to clear land for agriculture and wasexacerbated by intense poaching, whichoccurred both within and outside thenational parks and reserves. In 1984, inresponse to this rapid decline, Kenyaimplemented a strategy aimed at restoringthe rhinoceros population. The strategyincluded the establishment of sanctuaries,collaboration between the Kenya WildlifeService (KWS) and the private and com-munity sectors, heightened protectionthrough increased law enforcement,improved staff densities and capacities,revision of staff remuneration and drasticchanges to the management of nationalparks.

This strategy halted the decline of therhinoceros population in Kenya and since1987 numbers have increased, albeitslowly, at an average annual growth rateof �5%. However, if it had been possibleto achieve even the AfRSG minimum

REVIEW: EASTERN BLACK RHINOCEROS INTEGRATED MANAGEMENT STRATEGY 119

Fig. 1. Eastern black rhinoceros Diceros bicornis michaeli trend across Africa and in Kenya 1970–2003, in alogarithmic scale, showing the sharp decline and slow recovery. Data from Anon. (1993), Foose et al. (1992),Emslie & Brooks (1999), Brooks (2002) and Mulama & Okita-Ouma (2004).

acceptable (Emslie & Brooks, 1999) meta-population growth rate of 5% per annumthe population could have still grownsignificantly (Fig. 3). The value of 5% waschosen as the minimum acceptable growthrate as it is only just over half of r-max(9%) and should be attainable. In practice,one would hope to achieve higher rates ofincrease (6·5–9%). Given an expandingpopulation with a young age structure ingood habitat, one can temporarily achieveeven higher rates of growth (10%+). As itis, even at the 5% rate it would take30 years to achieve the target populationlevel of 2000 animals; a level that could beachieved in 15 years if annual metapopu-lation growth rate could increase to 9%(i.e. near the biological maximum growthrate for non-sex-biased black rhinocerospopulations). Large, rapidly breedinghealthy populations not only provide thebest possible insurance against futurepoaching losses but also preserve geneticdiversity, or at least minimize loss of

heterozygosity, by ensuring maximumrate of gene transfer to future generations(Gilpin & Soule, 1986). In recognition ofthe importance of striving to maintainrapid growth to retain genetic diversity,Kenya’s current Black Rhino Conserva-tion and Management Strategy placesgreater emphasis on improving metapo-pulation performance through effectivebiological management (Anon., 2003)(Fig. 4). The strategy advocates managingfor metapopulation growth of at least 5%per annum and preferably higher. Toachieve this, specific capacity and proce-dural mechanisms were urgently requiredin: (1) field monitoring of rhinoceros,data collection, analysis and reporting,(2) production of standardized annualstatus reports that assess the numbers,performance and population dynamics ofEastern black rhinoceros populations inKenya in order to aid biological-manage-ment decision making and (3) assessmentof habitat conditions and ecological


Fig. 2a. As at end of 2003 there were c. 540 and c. 170 Eastern black rhinoceros in situ and ex situ, respectively(AfRSG, 2004); b. Kenya is the stronghold of D. b. michaeli, with 458 rhinoceros in the population, as at end of2003, mostly within sanctuaries, both in protected areas and on private land, and in a free-ranging population oncounty-council land. Tanzania has an estimated 42 Eastern black rhinoceros, mostly in free-ranging populationsin unfenced protected areas and a few in one sanctuary. Rwanda and Ethiopia hold relict populations of one (ina protected area) and two to four (on community land) animals, respectively. As at 2003 South Africa had anestimated 36 D. b. michaeli, of predominantly Kenyan origin, out-of-range on private land.

carrying capacities in fenced sanctuaries,allowing the management of habitats byadjusting rhinoceros and other browserpopulation densities, and fire regimewhere relevant, and assisting in develop-ment of new viable populations. SeveralKenyan sanctuaries have experienced orare experiencing high densities of rhino-ceros and competing browsers, or veryhigh/low fire frequencies, and there is arisk of the long-term productivity of thehabitat being negatively affected.

MONITORING FOR MANAGEMENTIn order to monitor rhinoceros popu-lations, Kenya has implemented astandardized programme of patrols toobtain information on rhinoceros sight-ings. Rhinoceros are identified individu-ally and registers of the features ofindividual animals are maintained. Whereneeded, recordings are also made of sight-ings of ‘clean’ rhinoceros (i.e. those thatare not individually recognizable). These

monitoring data are used to provide esti-mates of population size, age and sexstructures, calving rates (i.e. � breedingperformance), mortality rates (by age andsex), and the distribution and movementof rhinoceros. This information is used togauge the performance of each populationand guide biological-management deci-sion-making processes, such as introduc-tions and removals, to realize the nationalconservation strategy of increasing thetotal rhinoceros population as rapidly aspossible.

The individual identification of rhino-ceros requires properly skilled and moti-vated observers, a system of strict controlon data quality at observer and data-recording levels, and the support ofthe wider conservation-managementstructure.

DEVELOPING MONITORING CAPACITYTraining A formalized training pro-gramme for personnel can greatly


Plate 1. An ear-notched and, thus, recognizable Eastern black rhinoceros Diceros bicornis michaeli on the plainin Lake Nakuru National Park, Kenya. ZSL, London, UK.

accelerate the process of acquiring highstandards of observational and data-col-lection skills. The project, funded throughthe UK government’s Darwin Initiative,has implemented a sustainable field-stafftraining programme and, at the time ofwriting, 24 officers from national parksand sanctuaries have been trained asaccredited rhinoceros-monitoring instruc-tors in the continental-wide modularAfrican Rhino Specialist Group (AfRSG)course. These field instructors are, in turn,training the c. 200 rangers in the field onan on-going basis. The process of testingand accrediting both trainers and traineeshas helped institutionalize the process andprovide formal recognition to those whogain accreditation.

A significant advantage of thisapproach is that staff can be trainedwhere they are located, thus saving timeand money and minimizing daily opera-tional impact on the field teams. In addi-tion, with continuous teaching of themodules it is possible to maintain consis-tency as well as deal with the inevitable

staff turnover. Standardized data collec-tion, including the use of the continentalrhinoceros ageing and body-conditionassessment systems, also means results canbe compared between parks, includingthose in other range countries (Adcock &Emslie, 2004).

Tools and procedural mechanisms TheKenyan Rhino Programme has imple-mented data quality-control procedures inall its reserves to ensure that the data arecollected on an on-going basis and are ofthe best possible quality. Field rangershave been trained to approach andobserve rhinoceros, and accurately com-plete the standardized sighting forms(Fig. 5). This information is then checkedby experienced accredited observers andthe sightings are classified in accordancewith the ‘ID’ master files, which are con-tinually updated by the field officers.

The extensive sets of field data arestored and analysed using a compre-hensive geographic information databasemanagement system (KIFARU). Standard-


Fig. 3. Estimates of Eastern black rhinoceros populations after 1%, 5% and 9% growth rates. Since 1987 popu-lation numbers of the Eastern black rhinoceros have increased at an annual rate of 1% and, without biologicalmanagement to improve population growth, it will take 170 years to reach the target population of 2000 rhino-ceros. However, if population growth rates in Kenya increased to 5% per annum, this target population will beachieved in 30 years and at a rate of 9% the target population could be achieved 15 years earlier than this.

ized monthly reports provide key informa-tion, such as sighting frequencies ofindividual rhinoceros, patrol movementsand the availability of manpowerresources, which are used to optimizedeployment of patrols and analyse popu-lation performance (Amin et al., 2001).

Monitoring practicalities Tsavo EastNational Park, Kenya, is vast(c. 11 000 km2) and the relatively lownumbers of monitoring staff, coupled withinternational boundary-related problems,such as infiltration of firearms from neigh-bouring countries, have made the effectivemonitoring and security of the rhinocerospopulation in this area difficult.

Rhinoceros are rarely sighted duringthe day in the difficult terrain of theChyulu-Hills National Park, or in thedense vegetation of Aberdare National

Park and Ngulia Rhino Sanctuary inTsavo West National Park, and thismakes it difficult to estimate populationsize and structure accurately. However,these areas are important to the long-termconservation of rhinoceros in Kenya.Tsavo East and West National Parks areover 20 000 km2 in total and have a veryhigh capacity to assimilate rhinocerosfrom smaller, overstocked sanctuaries andparks. This area used to have over 6000black rhinoceros before the poachingonslaught in the 1970s and 1980s.

The practical issues of achieving effec-tive monitoring in these areas need to beaddressed before such moves can takeplace. A number of initiatives are beingdeveloped and implemented. For example,the dry season full-moon night census isbeing improved using better planning andequipment to obtain more comprehensive


Fig. 4. The Conservation and Management Strategy for Eastern black rhinoceros sets six strategic objectives toachieve the national goal of increasing numbers of the subspecies to 1000 by 2020 and eventually achieving thetarget population of 2000 in natural habitat in Kenya (Anon., 2003).

Fig. 5. One side of the standardized form used to rec-ord details of rhinoceros sightings.

population information in the Tsavo WestNational Park Ngulia Rhino Sanctuary.Systems to achieve more effective night-monitoring programmes are also beingdeveloped in Aberdare and Chyulu-HillsNational Parks. A combination of directand indirect methods for deriving indicesof rhinoceros abundance, and relatingthese to indices of patrol and poachingactivities, are also being considered totrack population distributions and densi-

ties, and allocate law-enforcement effortsappropriately in order to protect thesepopulations.

Estimating population size A problemexperienced in many Kenyan reserves isthat an increasing number of animals thatare not individually recognizable (i.e.clean). Young clean rhinoceros becomeunrecognizable once they leave theirmothers. In the past, population estimateshave only been based on identifiableanimals, which provided a minimumindex rather than an estimate of the truepopulation size.

Reliable population-size estimates(within 90% of the true total) are requiredevery year (or at least every 3 years) inorder to assess how well a population isperforming and to manage populations atthe metapopulation level. This informa-tion, along with the ecological carryingcapacity (ECC) estimate of each park andsanctuary, can assist programme man-


Fig. 6. K-selected species like rhinoceros are likely to have a plateau of nearly constant growth rate (density-independent phase), followed by a ramp of density-dependent decline once the maximum sustained yield level(c. 75% of carrying capacity) has been exceeded.

agers in making decisions that are neces-sary for achieving and sustaining highpopulation growth rates.

The individual-identification-basedmonitoring method undertaken in allKenyan reserves enables the use ofmark–recapture methods for estimatingpopulation size. Field staff are beingtrained on a continual basis to ensureaccurate recording of sightings of bothidentifiable and clean rhinoceros, withequal emphasis on each sighting. RhinoProgramme scientists have also beentrained to use the continental level RHINO

Bayesian Mark–Recapture software tool(Emslie & Amin, 2003), which calculatespopulation size using ongoing sightingdata. To improve population estimates allrhinoceros immobilized for translocationor veterinary treatment are ear-notchedfor identification purposes as standardpractice. When resources are available,specific ear-notching exercises are under-taken in areas with a high proportion ofclean animals.

Annual status reporting National statusreporting is a key component of imple-menting the conservation strategy. In thepast metapopulation performance hasbeen ‘averaged out’ in Kenya, thusperformance issues within individualpopulations may have been overlooked.To promote optimal metapopulationperformance it is necessary to look at theage and sex composition, calving rates of��, and causes and rates of mortalitywithin each population. Reasons for sub-optimal performance can then be deter-mined and solutions put in place.

A formal national-status reporting pro-gramme has been implemented. KenyaWildlife Service scientists have beentrained to analyse population perform-ance data and synthesis of the national-status report. At time of writing, 26officers from 13 reserves have also beentrained in population data analysis andpreparation of annual park-status reports.The training introduces the principles ofstatus reporting, the concepts aroundmetapopulation management and, in


Fig. 7. Amounts of available browse in each of the nine main Kenyan black rhinoceros areas and the compositionalbreakdown of this browse by palatability class (i.e. the suitability of the browse component for black rhinoceros).Each class comprises several woody or forb species. Black rhinoceros browse is almost totally within a 0–2 mheight range and the browse availability (BA) measure indicates the percentage to which this 2 m layer is filledby browsable plant canopies. BA was sampled by vegetation type and a weighted average BA was obtainedaccounting for the proportional area of each vegetation type within each rhinoceros sanctuary.

particular, the critical need to maintainrapid population growth rates (for demo-graphic, strategic and genetic conserva-tion reasons). Trainees are shown how torecognize the main causes of poor growthand how to interpret various performanceindicators derived from their own ongoingground-based monitoring of rhinoceros.The practical application of these indica-tors is also highlighted, as trainees areshown how monitoring assists with thedecision-making process on sanctuarymanagement and translocations.

The park-level status reports supplyinformation on population size, age andsex structure, translocations and mortali-ties (including causes), as well as a numberof standardized biological performanceindicators (age at first calving, percentage�� calving, proportion of adult �� withcalves, intervals between calving, mor-

tality rates and net population growthrates) (Okita-Ouma, 2004).

The individual park reports are synthe-sized and analysed at a national level. Theresultant national report interprets andcontrasts the status, performance andpopulation dynamics of all Eastern blackrhinoceros populations in Kenya. Thefeedback from the national-statussummary report is vital to programmemanagers and staff because it places theresults of individual-reserve reports into ametapopulation context.

Without regular park-level statusreporting and the production of inter-preted national status report summaries, aproblem may also remain undetected farlonger. The first detailed status reportshave shown clear density dependence in anumber of populations stocked at or nearECC. The reports have indicated that a


number of Kenyan rhinoceros popu-lations could become donor populations.In addition, the c. 5% underlying popu-lation growth in Nairobi National Parkfollowing a period with an average 5%annual removal of rhinoceros has pro-vided empirical support to the SetPercentage Harvesting approach advo-cated by the IUCN/SSC’s AfRSG(Emslie, 2001).

ESTIMATING ECC OF SANCTUARIES ANDPARKSAchieving and maintaining a high meta-population growth rate can only be real-ized if the areas maintaining rhinocerosare not overstocked relative to availablehabitat resources. This depends on an esti-mate of the ECC of each area.

It is not easy to measure ECC. Past esti-mates of Eastern black rhinoceros ECChave differed widely in the reports of dif-ferent observers for the same area. Inparticular, non-experts have tended tooverestimate substantially the ECC.Nevertheless, it is easy to conceptualizethat a habitat at a given point in timemust be able to support sustainably alimited number of any given herbivorespecies. This ECC is probably mostclosely defined as ‘the maximum numberof animals of a species (sustainably)supportable by the resources of a specifiedarea’ (Caughley, 1976; McCullough,1992). Once the ECC is known, existingpopulations can be managed formaximum productivity and excess animalsremoved to enhance other populations orestablish new populations as necessary.Estimation of ECC is also a necessary pre-requisite for making decisions about thesuitability of an area for establishing anew subpopulation. For large mammals,such as rhinoceros, evidence suggests thatthe maximum sustainable yield or MSY[commonly called maximum productivitycarrying capacity (MPCC)] is c. 75% ofestimated (long-term) ECC (Owen Smith,1988; McCullough, 1992), although

optimal stocking levels may be lower thanthis during drought years (Fig. 6).

Managing populations at or below 75%of ECC should minimize the risk of den-sity-dependent effects negatively affectingpopulations (e.g. through reduced calvingsuccess and increased mortalities andsocial problems). By reducing densities ofEastern black rhinoceros and/or otherbrowsers to below habitat carryingcapacity, there is also a reduced risk ofnegatively affecting the long-term rhino-ceros carrying capacities of the areas.

Performance in a number of Kenyanblack rhinoceros populations may be poorbecause, in the past, inadequate attentionwas given to habitat conditions. Severalreserves face loss of suitable habitatthrough high rhinoceros and competingbrowser densities.

Kenyan Eastern black rhinoceros ECCmodel The Kenyan Darwin Initiativeproject has developed an ECC model forthe region. A similar model has beendeveloped for the Southern African rangestates (Adcock, 2001). The estimation ofECC is multi-faceted. For a given amountof standing ‘browse-plant biomass’ thebrowse productivity (growth rate) andquality are primary determinants of blackrhinoceros carrying capacity. Rainfall,soil quality and temperature influencethese parameters on a broad scale andmeasures of these factors are alsorequired. All nine well-established Kenyanrhinoceros sanctuaries were surveyed forbrowse availability and species composi-tion (Fig. 7). About 100–150 detailed veg-etation plots were assessed in each areaand this information was combined withLandsat-7 satellite imagery data to giveoverall browse-availability and browse-suitability index maps. Auxiliary data onvariables linked to Eastern black rhino-ceros ECC were also compiled for theareas (game count data, soil and geologydata, long-term rainfall and temperaturerecords).


Plate 2. Eastern black rhinoceros and African elephant Loxodonta africana (competing browsers) observed duringdry-season full-moon night census at Tsavo West National Park, Kenya. ZSL, London, UK.

Monitoring habitats Habitat monitoringprocedures have also been developed, andpark ecologists and monitoring staff aretrained in their use. This is important asECC will change over time (in the mediumto longer term) in response to habitatchanges, which for Eastern black rhino-ceros may be positive (increased browseavailability of suitable plant species) ornegative [favoured food plants decliningor growing out of reach, unpalatablespecies increasing at the expense of morepalatable species, increases in alien plants(e.g. Lantana camara), frequent fire,increased grass interference and/or anincrease in browsing pressure following abuild-up in numbers of competitivebrowsers (e.g. African elephants Loxo-donta africana, Giraffe Giraffa camelopar-dalis and Cape buffalo Syncerus caffer)](Plate 2). To be able to update estimatesof ECC continually, therefore, requires anunderstanding of browser/browsedynamics, the nature of vegetationchanges and their likely impact on ECC.

The estimates of ECC allow managersto take 75% of this figure as a ballparkestimate of the MSY level. Excess animalsare then removed to maintain densities

near the estimated threshold MSY level,thus maximizing rhinoceros productivity.

Monitoring the habitat resources aswell as rhinoceros-population perform-ance is useful in the light of changing veg-etation in East African ecosystems and theinfluences of competing browsers. Largelong-lived animals like rhinoceros mayalso overshoot carrying capacity before apotential problem in the population-performance indicators is identified.

SITE-LEVEL SUPPORT PARTNERSHIPSExperience has demonstrated that evenmoderate additional external institutionsite-linked assistance can make a signifi-cant impact at the park operational level.KWS’s collaboration with The ZoologicalSociety of London (ZSL) and other part-ners to provide additional support forNgulia Rhino Sanctuary, serves as anexample of the value of such site-level ini-tiatives. In addition to helping supportinfrastructural elements, such as water-system maintenance and vehicle provi-sion, site-level links can assist theimplementation and evaluation of the bio-logical management tools and associatedresearch areas. Securing out-of-region


grants (e.g. the UK government’s DarwinInitiative) is another important benefitthat can arise from developing such part-nerships which, in turn, benefit the widernational programme.

CONCLUSIONThe successful development and imple-mentation of the Kenyan Rhino Pro-gramme’s integrated biologicalmanagement and monitoring strategyacross all rhinoceros parks and sanctu-aries is key to providing decision makerswith the information necessary to managethe national herd at the metapopulationlevel. Specific staff training in the field hasmeant that there is a consistency of datacollection, which facilitates the ability tomake accurate assessments of the status ofthe whole population. There has been theadded benefit of gathering information onunrecognizable (clean) animals andincluding this in the population estimatesfor the region. Training has also beengiven in habitat assessment and data ana-lysis, population data analysis and pre-paration of reports, so any problems withrhinoceros habitats, population status,performance or dynamics can be identifiedand acted on as quickly as possible.Ongoing evaluation and refinement ofthese tools and associated approachesprovides the mechanism for optimizingKenya’s Eastern black rhinoceros annualgrowth rate as well as informing widerregional rhinoceros conservation efforts.The 2005 (end of year) national popu-lation estimate compiled at time of writingis 539 animals, which is an annualincrease of over 5% (the national strategygoal is to increase numbers by at least 5%)(Okita-Ouma & Wandera, 2006). The endof 2004 national estimates also showedsimilar increase, for the first time since the1970s.

ACKNOWLEDGEMENTS

The authors would like to highlight the support ofUK government’s Darwin Initiative, African RhinoSpecialist Group (AfRSG), The Zoological Society

of London, Kenya Wildlife Service Director JuliusKipng’etich, and staff of all Kenyan rhinocerosreserves for their continued commitment to rhino-ceros conservation.

REFERENCESAdcock, K. (2001): Black rhino carrying capacityand performance. In Proceedings of a SADC rhinomanagement group (RMG) workshop on biologicalmanagement to meet continental and national blackrhino conservation goals: 9–13. Emslie, R. (Com-piler). KwaZulu Natal: Giants Castle Game Reserve.Adcock, K. & Emslie, R. (2004): MonitoringAfrican rhino. In ‘Sandwith’ training course for fieldrangers (5th edn). Malilangwe, Zimbabwe: IUCN/SSC, African Rhino Specialist Group.AfRSG (2004): Working group report. 5. Meta-population planning for D. b. michaeli. In IUCN/SSC-AfRSG meeting. Leader-Williams, N. &Okita-Ouma, B. (Eds). Kilaguni, Tsavo West:African Rhino Specialist Group.Amin, R., Okita-Ouma, B. & Mulama, M. (2001):Kenya implementing a new black rhino informationmanagement system. Pachyderm 30: 96–97.Anon. (1993): Conservation strategy and manage-ment plan for the black rhinoceros (Diceros bicornis)in Kenya. Nairobi: Kenya Wildlife Service, RhinoProgramme.Anon. (2003): Conservation and managementstrategy for the black rhino (Diceros bicornismichaeli) in Kenya 2000–2005: ix, 39. Nairobi:Kenya Wildlife Service.Brooks, M. (2002): Proceedings of African RhinoSpecialist Group meeting. In IUCN-SSC-AfRSG.Emslie, R (Ed.). Malilangwe, Zimbabwe: IUCN/SSC, African Rhino Specialist Group.Caughley, G. (1976): Wildlife management and thedynamics of ungulate populations. In Appliedbiology: 183–246. Coaker, T. H. (Ed.). New York,NY: Academic Press.Emslie, R. (Compiler) (2001): Proceedings of aSADC rhino management group (RMG) workshop onbiological management to meet continental andnational black rhino conservation goals. KwaZuluNatal: Giants Castle Game Reserve.Emslie, R. & Amin, R. (2003): rhino rewrite: anupdate. Pachyderm 30: 100–101.Emslie, R. H. & Brooks, P. M. (1999): African rhino.Status survey and action plan—IUCN/SSC AfricanRhino Specialist Group. Gland and Cambridge:IUCN.Foose, T. J., Lacy, R. C., Brett, R. & Seal, U. S.(1992): Kenya black rhino meta-population ana-lysis—briefing book. In Kenya black rhino meta-population workshop. Nairobi: Kenya WildlifeService, IUCN/SSC/CBSG.Gilpin, M. E. & Soule¬, M. E. (1986): Minimumviable populations: processes of species extinction. InConservation biology: the science of scarcity and


diversity: 19–34. Soule, M. E. (Ed.). Sunderland,MA: Sinauer Associates.IUCN (2004): 2004 IUCN red list of threatenedspecies. Gland and Cambridge: IUCN. http://www.redlist.orgMcCullough, D. R. (1992): Concepts of large her-bivore population dynamics. In Wildlife 2001: popu-lations. Barret, R. H. (Ed.). London: ElsevierApplied Science.Mulama, M. & Okita-Ouma, B. (2004): Black rhinoannual status report summaries: 16. Nairobi: KenyaWildlife Service.Okita-Ouma, B. (2004): Population performance ofblack rhinoceros Diceros bicornis michaeli) in six

Kenyan rhino sanctuaries. MSc dissertation, DurrellInstitute of Conservation and Ecology, University ofKent, UK.Okita-Ouma, B. & Wandera, A. (2006): Status andmanagement of black rhino in Kenya 2005–2006:Kenyan black rhino status report summary. Nairobi:Kenya Wildlife Service, Darwin Report.Owen-Smith, N. (1988): Megaherbivores. The influ-ence of very large body size on ecology. Cambridge:Cambridge University Press.

Manuscript submitted 13 July 2005;accepted 13 February 2006; 9 May 2006

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