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Road kills of animals by public traffic in Mikumi N atio ~'-.
Park, Tanzania, with notes 011 baboon mortality
Department of Zoology, UniversitJ' of Cambridge. Cambridge CE23EJ, U.K

The Tanzania-Zambia highway crosses Mikumi National Park over a stretch of
50 km. The road was rehabilitated between May 1990-November 1991 resulting
in higher average driving speeds. Data were collected on the incidence of road
kills of animals within the park boundaries during this period. The list of 183
mammals, birds and reptiles killed by vehicles included at least 52 species, two of
which are endangered: the African elephant (Loxodonta africana Blumenbach)
and the African hunting dog (L,vcaon pictus Temminck). The road-kill rate
increased and was estimated at a minimum of three road kills per day (21'8 road
kills/km/yr) by the end of the study. Road kills were more frequent near water
holes and traditionally used animal tracks. The demographic effect of road kills
on troops of yellow baboons (Papio cynocephalus Linnaeus), which include the
highway in their ranges, is in the order of 10% of the total yearly losses. High
speed and the danger of collisions with wildlife are causing regular accidents in
Mikumi National Park. Adequate signposts along the road and speed bumps are
recommended in order to reduce the average driving speeds for the sake of
humans and wildlife.

Key words: conservation, demography, endangered species, wildlife management

La grand route qui va de Tanzanie en Zambie traverse Ie Parc National de
Mikumi sur une distance de 50 km. La route a ete refaite entre mai 1990 et
novembre 1991, ce qui a entraine une augmentation de la vitesse moyenne des
vehicules. On a recolte des donnees sur Ie nombre d'animaux tues sur la route
dans les limites du parc pendant cette periode. La liste des 183 mammiferes,
oiseaux et reptiles tues par des vehicules comprend au moins 53 especes dont 2
sont menacees: l'elephant africain (Loxodonta africana Blumenbach) et Ie lycaon
(Lycaon pictus Temminck). Le taux d'animaux tues sur la route a augmente, et
on l'estimait a environ trois morts par jour (21,8 animaux tues/kmlan) a la fin de
l'etude. Les accidents mortels etaient plus frequents pres des points d'eau et des
pistes traditionnellement utilisees par les animaux. L'effet demographique des
accidents sur les groupes de babouins olive (Papio cynocephalus Linnaeus) dont
l'aire de distribution couvrait la grand route est de l'ordre de 10% des pertes
annuelles totales. La grande vitesse et Ie risque de collision avec des animaux
*Present address: Programa de Vida Silvestre. Universidad Nacional. Apdo. 1350-3000. Heredia.
Costa Rica.
90 C. Drews Road kills in a Tanzanian nalional park 91

provoquent regulierement des accidents dans Ie Pare National de Mikumi. On in the park. Between 16 May 1990 and 16 August 1991 the species and location
recommande de placer des panneaux routiers ct des casse-vitesse pour reduire la of each road kill encountered were recorded while driving at 40 km h - \ on the
vitesse moyenne, pour la securite des gens et des animaux. highway to and from the baboon study area (5-15 km north of the park
headquarters). as well as during trips to Mikumi town along park territory. On
Introduction average, a 20-km stretch (40'Yo of the highway within the park) was sampled
'v1ikumi National Park in Tanzania is one of a number of African conservation regularly. The northern sector of the road was sampled much less often. Sampling
areas which are traversed by a public road. The TANZAM (Tanzania-Zambia) occurred on 65% of the total number of days covered by the study period. In
highway traverses the reserve over a stretch of 50 km. Anecdotal reports order to estimate the absolute rate of road kills along the 50-km road stretch
of vehicles killing elephants (Loxodonta ajricana Blumenbach), giraffes within the park, the number of kills recorded was corrected for the proportion of
(GiraOu camelopardalis Linnaeus) and hippopotamus (Hippopotamus amphibius highway covered during regular sampling as well as the proportion of days
Linnaeus), among others, indicated that the traffic was having an impact on the during which it was surveyed. This yields a correction factor of 3·75. Road kills
protected wildlife. Although public highways and railways pass through nature generally disappeared from the road within one day as a result of the actions of
reserves in Africa (e.g. Kenya: Tsavo National Park, Marsabit National Park; scavengers. Occasional road-kill reports by other drivers are also included in this
Tanzania: Serengeti National Park, Selous Game Reserve; Uganda: Queen study. The recording effort remained more or less constant throughout the study.
Elizabeth National Park; Zaire: Virunga National Park; Zambia: Kafue The records were limited to the more conspicuous animal classes: mammals, birds
National Park), the range of species affected by vehicle traffic and the demo­ and reptiles. However. frogs and invertebrates, particularly ants, grasshoppers,
graphic implications of road kills has rarely been quantified in this continent and millipedes were also commonly found killed by vehicles. The speed of
(1. Lopez, pers. comm.; see review of road-kill literature by Lopez & Roviralta, vehicles was measured by driving behind them and reading the speedometer.
1993). This study complements the scarce information available about road kills Unfortunately, the road toll office in Mikumi town was closed down during the
in Africa (e.g. Broekhuysen, 1965; Lewis, 1989; Drews, 1991). study, and highly desirable information on traffic rates became unavailable.
The highway was in operation prior to the establishment of Mikumi National Quantitative information on road proximity and group size of baboons was
Park in 1964, 300 km south-west of Dar es Salaam. The park was expanded in used to assess the possible demographic effects of road kills on a gregarious
1975 to a present area of 3300 km 2 , which includes grassland, open woodland, mammal. A visit to the highway by a baboon troop was defined as any instance
miombo and montane rain forest (Norton et al., 1987; Lovett & Norton, 1989). in which at least 10% of troop members was within 10 m of the road. Baboon
The highway is of great economic importance in that it provides the main access troops were followed each day for seven hours, on average 0745-1545 h). When
to southern Tanzania and its bordering countries of Zambia and Malawi, which a troop visited the highway more than once during an observation day (9% of all
utilize the road for the transportation of goods to and from the port of Dar es cases, N= [44). the single durations of these visits were added to calculate the
Salaam. Drivers in transit through the park can contemplate African wildlife at daily stay on the road (Table 3). The highway stretch considered for this analysis
no cost, an unusual privilege given the dwindling number of large mammals in was 20 km long: from 15 km north of the park headquarters, to 5 km south of
the rural areas of Tanzania. the headquarters. Non-parametric tests were used in all statistical analyses.
The public road stretch inside the park was repaired between May 1990 and
November 1991 as part of the rehabilitation scheme for the TANZAM highway.
In early 1990 the poor condition of the road forced vehicles to cross the park at Results
average speeds of 30 km h - I. By the end of 1991, however, buses, lorries and Location of road kills
private vehicles passing at speeds above 90 km h ~~ 1 on the immaculate tarmac A total of 183 road kills were recorded within the national park over 15 months.
were common. During the same period, information on the occurrence of road The bulk (98°/,,) of the road kills occurred on the public highway. Kills were
kills within the national park boundaries was collected in order to assess (i) particularly common along road stretches near waterholes and where traditional,
which taxa are affected by traffic in Mikumi; (ii) the road kill rate; (iii) the effect small-scale migration tracks cross the highway. Three birds (2%) were found
of an improvement in road condition on road kill rates; and (iv) to quantify the dead on the unpaved tourist road system.
local effect of road-kill mortality on a subpopulation of yellow baboons (Papio
cynocephalus Linnaeus). This report provides a basis for the design of manage­
ment measures targeted at improving road safety, the conservation of threatened
species, and animal welfare in conservation areas intersected by public roads. Taxa represented among road kill records are listed in Table I. At least 52 species
of mammals (41°/<)), birds (42%) and reptiles (17%) were among the sample.
Mammals accounted for 61 % of the total number of road kills, birds for 30% and
reptiles for 9%. The most commonly killed mammal was the African civet
Road-kill data were collected during a study on the behaviour of yellow baboons (Viverra cil'effa Schreber), followed by spotted hyaena (Crocuta crocufa
in Mikumi National Park. The two-lane public highway is the only paved road Linnaeus), yellow baboon and African buffalo (Syncerus caffer Sparrman). Other
92 C. Drews Road kills in a Tanzanian national park 93

Table I. Number of road kills Table I. Conlinued

Number of Number of
for each species recorded in
Species kills Species kills
~likumi National Park.
Tanzania (16 May 1990-16
August 1991) Birds Mammals continued
Birds of prey Other con I inued
Gyps aFicanus Salvadori 2 Lepus sp. Linnaeus 6
Gypoheirax angolensis Gmelin 1 Canis sp. Linnaeus 5
Circus ranivorus Daudin 2 Giraffa camelopardalis Linnaeus
Aquila wahlbergi Sundevall 1 Helogale parvula Sundevall 2
Mil1'lIs migrons Boddaert i Loxodonta aFicana Blumenbach 2
Falco dickinsoni Sclater 1 Connochaeles taurinus Burchell o
Aquila rapax Temminck 1 Panlhera pardus Linnaeus 2
Glaucidium capense Smith I Procavia capensis Pallas
unidentified vultures 3 Cephalophus sp. Hamilton Smith
unidentified raptors 2 Mel!ivora capensis Schreber
Panthera leo Linnaeus
8 Total mammal road kills 111
Coracias cal/data Linnaeus
Number of species=22
SlI'cplOpelia sp. Bonaparte 4
Merops apiasler Linnaeus 3
Melaenomis pallidus Muller 2
Cenlropus superciliosus Hemprich & Ehrenberg 2
Varanus exanthematicus Schmidt 2
Hirumlo sp. Linnaeus
Varanus sp. Merrem
,Numida meleagris Linnaeus
GUllera pucherani Hartlaub
Francolinus afer Statius Muller
Cicon;a ciconia Linnaeus
Macrodipleryx vexillaria Gould
1 Snakes
Macronv.\' sp. Swains
Python sebae Gmelin 2
Prinia subf/ava Gmelin 1
Rhamphiophis oxyrhynchus Reinhardt 2
Lanius sp. Linnaeus 1
Bilis Gl'ielans Merrem
Lamprolomis chalibaeus Hemprich & Ehrenberg I
Naja nigricol!is Linnaeus
unidentified birds 10
Psammophis phillipsii Hallowell
Total bird road kills
55 Psammophis sp. Boie
Number of species=23
unidentified snakes

Total reptile road kills 17
Number of species=7
Viverra civella Schreber 20 Total of road kills recorded
Crocuta croc1I1a Linnaeus 17 (III! taxa) 183
Genella sp. Oken 2 Total number of species
Hystrix crislata Linnaeus identi{!ed= 52
Ietonyx striata Perry
Triaenaps pers!c'a Dobson
mammals recorded more than twice include impala (Aepyceros melampus
Diurnal Lichtenstein), hare (Lepus sp. Linnaeus), jackals (Canis sp. Linnaeus), giraffe
Papio cynocephalus Linnaeus 15
and African hunting dog (Lycaon pictus Temminck). The lilac-breasted roller
Lycaon pictus Temminck 3
Plwcoc!lOerus aelhiopicus Pallas 1 (Coracias caudata Linnaeus) was the most common bird species among the road
kills (Table I). The bird list includes two species of palaearctic migrants, the
Other white stork (Ciconia deania Linnaeus) and the European bee-eater (Merops
Syncerus caffer Sparrman 11 apiaster Linnaeus). Birds of prey accounted for 27% of all bird records. Most
Aepyceros melampus Lichtenstein 7 reptiles killed were snakes, among a list which included rock monitor lizard
(Varanus exanthematicus Schmidt) and terrapin (Pelomedusidae).
94 C. Drews Road kills in a Tanzanian national park 95
Table 2. Baboon troops which include in their range part of a 20 km stretch of public highway in Mikumi
National Park. Juvs. =juveniles and infants. The categorization of troops with respect to frequency of
occurrence on the road is subjective (F. frequently; R, rarely). (-j no data
Troop On road Size Males Females Juvs. Dates of census
0 30
Mgoda F 28 4 8 16 26 March 1991
~ Viramba 2 F 32 3 II 18 16 August 1991
'" Viramba I F 21 I 8 12 20 August 1991

20 Yuma Hill F 60 6 17 37 27 April 1991
0:: I ./ Tented camp F 16 I 8 7 22 February 1991
Kikoboga F 20 3 8 9 26 October 1991
10 Highway F
Shy R 21 3 19 March 1991
Mwanambogo R 66 II 29 26 2 March 1991
Bwawadogo R 7 I 4 2 27 March 1991
1m I 1~ Kambini R
Troopx R
Fig. I. Monthly number of road kills recorded in Mikumi National Park, Tanzania, showing a significant
increase with time (Spearman's 1'=0'878. n= 15. P<O·OOI). Troop size: 0'=29'1 (SD=19'lj, Median=21. N=IO

traffic was diverted during road work. Temporary dirt tracks made up 45% of the
total stretch considered. Thus, good road condition (i.e. higher driving speeds) is
Road kill rate associated with significantly more road kills (X 2 = 5,69, df= I, P<0·05).
Data from the last 90 days of the study (N= 73 road kills) yield a rate of 91 road It is noteworthy that doves (Streptopelia sp. Bonaparte) and impalas started
kills per month, i.e. about three road kills per day within the park boundaries to appear repeatedly in the records during the last months of the study when
during that period. This figure is equivalent to 21·8 road kills/km/yr, consisting most of the highway had been newly paved. These animals are extremely agile
of 13·3 mammals/km/yr, 6·5 birds/km/yr and 2·0 reptiles/km/yr. These rates were and fast, which suggests that they are killed only by vehicles going at high
calculated by taking into account the percentage of road distance sampled speeds. On 3 November 1991 the road work inside the park was completed.
regularly as well as the proportion of days when it was sampled (see Methods). Subsequently, vehicles going at more than 90 km h - I were regularly seen on the
The rate calculated above is probably an underestimate given that an unknown 50 km of excellent tarmac that traverses the park.
proportion of animals were hit by vehicles and died in the nearby vegetation,
without being recorded. In particular, this negative bias affects the bird counts Demographic effect of road kills in yellow baboons
and perhaps large mammals which were not killed outright and died later from The sample for this analysis is composed of, firstly, 15 baboons found killed
injuries obtained on the road. within a 20 km road stretch over a period of 15 months, and secondly, of 13
Over the course of the study, the number of road kills increased significantly troops that were seen more or less regularly within that 20 km road stretch
(Spearman's r=0'878, N= 15, P<O'OOI; Fig. I). This positive correlation was still (Table 2). The size of troops which were seen frequently on the road did not
significant when the extremely high value for July 1991 was omitted from the differ significantly from those troops which were only seen rarely on the highway
sample (Spearman's r=0'849, N= 14, P<O·OOI). The road-kill rate was five times (Mann-Whitney U-test, two-tailed, P=0·67). The estimated total of baboons
higher at the end of the study than at the beginning (first three months N= 15, killed along the 20 km stretch is 23, when the fact that the road was not sampled
last three months N=73). The obvious correlate of this increment is the on 35% of the time is taken into account (i.e. 15 x 1'5).
improving conditions of the road during the study period and the associated Three adult males and 10 juveniles, no infants and no adult females, were
increase in average driving speed. The road work in the park started early 1990 found among the dead baboons. The pooled troop compositions of nine troops
at the northern boundary. During the second half of the study the new paving (Table 2) indicate that the proportion of juveniles to adults in the subpopulation
started to include the main sampling road stretch and in November 1991 it is 139 (51%) to 131. The proportion of juveniles among the road kills (77%) was
reached the southern boundary of the park. higher than in the overall population. This difference just missed statistical
During March, April and May 1991, the condition of the road where a kill significance (X 2 =3'309, df= I, P=0·0653). A larger sample size is required for a
was found was recorded. Out of 30 records, 23 (77%) were found on good more powerful test of the susceptibility of certain age and sex classes to be killed
tarmac, whereas only seven occurred on the temporary dirt tracks to which the by vehicles.
96 C. Drews Road kills in a Tanzanian national park 97

Given a mean of 29 baboons per troop and 291 baboons among 10 troops of Table 3. Frequency and duration of highway visits by three baboon troops in Mikumi National Park
known size (Table 2), the estimated total of baboons among the 13 troops of the
sample is approximately 378 (i.e. 291 +(29 x 3)). Thus, the estimated total of Percent Min. per day
Troop Days observed on road on road SD Median Maximum per visit
23 baboons killed over the IS months of the study represented 6% of the
baboon subpopulation that frequents the road. On average, 1·53 baboons were
killed each month on the road or about 18·4 baboons per year (4'9% of Viramba 1 20 35,0"';, 36 16 30 60
subpopulation). Consequently, each troop lost, on average, 1·4 baboons per year Viramba 2 90 50·0";', 58 52 46 220
Mgoda 34 76'5'~';' 17 23 9 94
to the traffic on the highway, i.e. 4·8% of the average troop size, when
recruitment from births and immigrations is not accounted for.
Days observed=average duration of daily observation was seven hours (0745-1545 h); Percent on
The estimated rate of baboons killed per troop and knowledge about the road=proportion of days during which the road was visited by the troop at least once; Min. per day on
demography of baboon troops in Mikumi enables a crude assessment of which road=mean daily time spent on the road in minutes on days when the road was visited; SD=standard
proportion of overall losses in a troop could be attributed to road kills. deviation; Maximum=maximum duration of a single visit in minutes
Demographic data over the same period of the road kill sample are available
from two medium-sized troops eViramba I' and 'Viramba 2', Table 2). A 'loss'
Road safety
is defined here as any baboon which died or disappeared from the troop, except
subadult males which left their natal troop upon attaining sexual maturity (four Any collision with an animal entails the risk of causing material damage
cases, all resighted later in other troops, pers. obs.). Adult males that transferred and human casualties. Eighteen collisions between vehicles and large animals
out of the troop are treated as 'losses'. 'Viramba I' lost four adult males, four (elephant, buffalo, giraffe and wildebeest (Connochaetes taurinus Burchell» were
adult females and eight juveniles (total of losses= 16, i.e. 12·8 per year). 'Viramba recorded during this study. Two human fatalities were associated with these
2' lost eight adult males, three adult females and seven juveniles (total of collisions. Several additional accidents occurred when the driver successfully
losses= 18, i.e. 1404 per year). Thus, the estimated troop average of I A baboons avoided hitting an animal but left the road after losing control of the vehicle. On
killed by vehicles each year could represent 10·9% and 9'7% of the total yearly average, at least one large animal collision occurred each month on the highway
losses, respectively. stretch inside Mikumi National Park.
None of the losses in the Viramba troops was confirmed to be attributed to Expensive freight losses and pollution due to wildlife-related accidents are
road kills. Carcasses of baboons killed by vehicles in the study area could not be further reasons for concern. In October 1991 a petrol tanker rolled over when the
assigned to any particular troop unambiguously because (i) the killing was not driver avoided a buffalo crossing the highway. Several tons of crude oil were
witnessed; and (ii) damage to the baboon in question did not allow identification. spilled onto the grassland as a consequence. In another accident, a large load of
Thus, the demographic data of Viramba troops were used to perform an indirect DDT was spilled in Mikumi National Park in 1982 (G. Norton, pers. comm.).
estimate of the potential extent of road kill mortality as a proportion of overall
troop losses. Deliberate killing of animals by drivers
Two factors should be considered for the interpretation of the average values On two occasions drivers were seen attempting to hit yellow baboons with their
calculated above. Firstly, the overall road-kill rate increased throughout the vehicles. It is conceivable that species considered as vermin (e.g. baboon), dirty
study (Fig. 1); towards the end of the study the road-kill risk was five times (e.g. spotted hyaenas), or good to eat are killed deliberately by some drivers (c.
higher than at the beginning (Section 3). Thus, the mortality risk that baboons Kidung'ho, pers. comm.). Impala was the only road kill observed to be, illegally,
were experiencing on the highway at the end of the study is higher than the mean carried away by drivers for meat. In five of the seven impala kills recorded, only
value calculated. Eight (53%) of all baboon kills recorded occurred during the a few pieces of blood-stained fur enabled identification after the carcass had been
last three months of the study. Only 33% of the IS kills occurred during the first taken by drivers. A similar situation applies to buffalo, which was occasionally
half of the study. Second, the troops differed considerably in the frequency and found to have been butchered prior to the carcass being discovered. These
duration of highway visits, i.e. some troops were at much higher risk than others observations suggest that there is a realistic potential for meat poaching on the
(Table 2, Table 3). highway by using vehicles as weapons.
Troops which were seen regularly on the highway differed significantly in
their mean daily stay on the road (Kruskal-Wallis one-way ANOVA, X2 =22'9,
df=2, N=76, P<O·OOl). Frequencies and daily stay on the road are underesti­ Discussion
mates since daily observations took place during 7 out of 12 daylight hours. The different species observed killed on the highway are a representative sample
Although Mgoda troop had the highest frequency of visits to the highway, it of the mammalian diversity of Mikumi National Park. Interestingly, the bat
spent the shortest time on the road during each visit. Most of the visits recorded Triaenops persica Dobson could be the first record of the species in the region
for Mgoda troop were an uninterrupted crossing of the road as the troop moved (Kingdon, 1974; K. Howell, pers. comm.). Most mammals were probably killed
from a sleeping site to the foraging grounds. at night when visibility is poor and the animals are blinded by the headlights of
98 C. Drews Road kills in a Tanzanian national park 99

the vehicles. The most commonly killed mammals were nocturnal species, vehicle losses in wildlife-related accidents should be reduced. Secondly, the list of
namely African civet and spotted hyaena. Although not strictly nocturnal, road kills included at least two endangered species, namely African hunting dog
buffalos too were generally killed at night. Interestingly there were no small (IUCN, 1990), and African elephant (Appendix I of CITES, 1991). Five
rodents among the road kills in Mikumi National Park (cf. Lewis, 1989). This elephants are known to have been killed on the highway inside the park between
taxonomic group, together with domestic carnivores, represents the majority of 1989-1992 (Mikumi National Park staff, pers. comm.). The killing of endangered
mammal records in some road-kill studies in Europe (e.g. Frean et al., 1993; animals is of immediate concern to conservation-oriented management plans.
Lopez, 1993). Thirdly and finally, from an ethical point of view, every individual animal
The relatively high number of killed yellow baboons, a strictly diurnal deserves protection from unnecessary, human-induced suffering or death.
species, may be partly explained by the acquired habit in some troops of
expecting food from passing vehicles. The feeding of baboons, prohibited in all Recommendations
Tanzanian national parks, is still a common practice along the highway despite
warning signs. In addition, drivers are often unaware of their accidental loss of Undoubtedly, the best way of reducing the rate of collisions between vehicles and
food. On at least four occassions baboon troops stayed on or next to the road for animals is to reduce the driving speed. The existing speed limit of 80 km h - I on
up to 3·5 h when maize fell from passing lorries. Baboons utilized the highway in the TANZAM highway is too high within the park. A speed limit of 50 km h - I,
additional ways, including walking on the tarmac to cover long distances and which is the limit in all Tanzanian national parks, should be implemented on the
foraging on the lush roadside vegetation. Reptiles seek the warm tarmac to public highway within the park boundaries. Adequate signposts at regular
absorb heat. This basking behaviour on the road makes them particularly intervals should remind the drivers to reduce speed and of the presence of wild
vulnerable to passing vehicles. animals on the road. The most effective way of reducing average driving speed,
Overall. a large proportion (27%) of the killed birds were birds of prey. These however, is speed bumps. These should be placed close to areas identified as 'hot
exemplify the fact that road kills facilitate more road kills: dead animals on the spots' for road kills, such as waterholes and traditional, frequently used animal
road attract scavengers, which during the day are mainly raptors and vultures. tracks. A further step towards reducing additional road kills is to remove from
These are, in turn, killed next to the carcass by passing vehicles. Some of the dead the road any carcass found, so that its scavengers are not exposed to the traffic
jackals (Canis sp.) and hyaenas were also in the proximity of fresh carcasses. (also Barragan & Lopez, 1993).
The estimated rate of three road kills per day and its increasing trend indicates
that the effect of public transport on the wildlife of Mikumi National Park is not Acknowledgments
negligible. The increase of the road-kill rate was associated with the rehabilitation I am grateful to the Tanzanian Commission for Science and Technology,
of the highway which, in turn, led to higher average driving speeds. Also, the Tanzania National Parks, and the Serengeti Wildlife Research Institute for
additional traffic resulting from the vehicles of the road workers could have permission to work in Mikumi National Park. The national park authorities and
contributed to the increase in road-kill rate. By the end of the study, the rate of rangers provided invaluable logistical support and background information
large mammals killed on the road probably exceeded that of poached wildlife in the about the road-kill problem. The Long Term Mikumi Baboon Project kindly
park (D. Njao, warden in charge of anti-poaching in Mikumi N.P., pers. comm.). provided unpublished demographic information on two of the baboon troops
Highway mortality is evidently one more demographic parameter in some of the cited (Viramba 1 and 2) as well as accommodation in the park. Special thanks to
populations which include the road in their range. For some of the baboon troops Astrid Schmidl-Drews, Charles Kidung'ho, William Marwa and Ayoub Njalale
which frequently visit or cross the highway, road kills may have a measurable who helped regularly in spotting, identifying and removing carcasses from the
influence on survivorship, potentially accounting for about 10% of annual losses. road. I thankfully acknowledge Michael O'Meara, who collected road-kill
The relatively high incidence of bone fractures among the adult male baboons of information during by temporary absence in October-November 1990. My
the Highway troop is probably associated with their high frequency of road visits sincere thanks to all those who reported road kills to me throughout the study,
(pers. obs). Although not fatal, impacts by vehicles are impairing these males in and to Keith Eltringham, Dawn Hawkins, Guy Norton, Sam Wasser, Javier
respect of their competitive ability and reproductive performance. Lopez and two anonymous referees for their valuable comments on a previous
The problems associated with road kills in African protected areas are similar draft. The Leakey Foundation, SigrnaXi, Durham Fund, Bedford Fund and
to those identified in a recent, nationwide survey of road kills in Spain Kings College, Cambridge made this study possible by funding my stay in
(Coordinadora de Organizaciones de Defensa Ambiental CODA, 1992; Mikumi National Park.
Asociacion Tecnica de Carreteras AIPCR, 1993) At least three reasons call for
attempts to reduce the rate of road kills in Mikumi National Park. First, the risk References
of collisions with large animals within the park boundaries is increasing as traffic BARRAGAN. B.I. & LOPEZ. J. (1993) Soluciones a los atropellos de vertebrados en carreteras. In: II
Simposio Naciollal sobre Carreteras y Medio Ambiel1le. Asociacion Tecnica de Carreteras (AIPCR),
on the TANZAM highway increases in parallel with the development of
southern Tanzania and its neighbouring countries. In addition to human BROEKHUYSEN, G. (1965) An analysis of bird casualties on the road in the southwestern Cape Province,
casualties, the risk of pollution in the park and of considerable freight and South Africa. L·Oiseall. Rev. ji". Omit/wI. 35,35-51.
100 C. Drews Afr. 1. £Col. 1995, Volume 33, pages 101-113

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estudio v prevencion de la mortalidad de rerlebrados en carreteras, Vols. 1-3. CODA, Madrid.
ASOCIACION TECNICA DE CARRETERAS AIPCR (1993) II Simposio Nacional sobre Carreteras y Media significance of plant height in Aloe candelabrum
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Department of Botany and UN-FRD Research Unit for Plant Growth and Development.
en las riberas del Millo en Orense. In: If Simposio Nacional sabre Carreteras y Media Ambiel11e.
Asociaci6n Tecnica de Carreteras (AIPCR). Madrid. University of Natal, Pietermaritzburg 3200, South Africa
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IUCN-World Conservation Union. Gland. Switzerland. Summary
KINGDON. I. (1974) East African Mammals. An Atlas oJ Evolution in Africa. II (A). Insecrivores and Bals. Pollinator exclusion experiments showed that insect pollination of Aloe
Academic Press. London.
LEWIS. A.D. (1989) Road kills and other records of mainly smaller mammills from Kenya: data for a
candelabrum Berger is negligible and that birds are the only effective pollinator
Kenyan mammal atlas. Eas/ Afi'. Nat. Hisr Soc. Bull. 19. 20-22. guild. Bird activity is greatest where flowering aloes are most dense. Also fruit set
LOPEZ. I. (1993) Metodologia y resultados del proyecto de seguimiento de la mortalidad de vertebrados en is lower in those aloes that bloom later in the season after most of the population
carreteras (P.M.V.C.le.O.D.A.). In: II Simposio Nacional sohre Carreteras y Medio Ambiente. has already flowered. Thus the reproductive fitness of aloes is dependent on the
Asociaci6n Tecnica de Carreteras (AIPCR). Madrid. density of flowering plants and there is selection for limited seed dispersaL As a
LOPEZ. J. & ROVIRALTA. F. (1993) Banco de datos y centro de documentaci6n acerca de la mortalidad de
vertebrados en carreteras. In: II Sirnposio Nacional sobre Carre/eras F Media Amhien/e. Asociaci6n
result, spatial distributions show clumping in young populations. These clumps
Tecnica de Carreteras (AIPCR). Madrid. act as nuclei from which plants spread slowly over time with mature individuals
LOVETT. I.e. & NORTON. G.W. (1989) Afromontane rainforest on Malundwe Hill in Mikumi National forming the centres of the densest stands.
Park. Tanzania. BioI. Conser". 48. 13-19. As a further consequence of limited seed dispersal, intraspecific competition
NORTON. G.W .• RHINE. R.I., WYNN. G.W. & WYNN. R.D. (1987) Baboon diet: A five-year study of is intense, resulting in self-thinning of the population. The height of A.
stability and variability in the plant feeding and habitat of yellow baboons Papio cynocephalus
of Mikumi National Park. Tanzania. Folia primarol. 48, 78-120.
candelabrum does not enhance pollinator attraction. In closed environments,
aloe height serves only to allow the persistence of those plants that establish
(Manuscript accepted 3 March 1994) before the development of a tree canopy. Under the constraint of limited seed
dispersal capabilities, selected for by pollinator activity, the height of A.
candelabrum serves as an adaptation to spread seeds away from the parent plant
and from each other. Variability in seed morphology enhances the latter effect.

Key words: Aloe, plant height, seed dispersal, competition

Des experiences sur l'exclusion des pollinisateurs ont montre que la pollinisation
par les insectes est negligeable chez Aloe candelabrum Berger et que les oiseaux
sont les seuls pollinisateurs reellement efficaces. L'activite des oiseaux est
maximale quand la floraison des aloes est la plus intense, et la fructification est
plus faible chez les aloes qui fleurissent plus tard dans la saison, lorsque la plus
grande partie de la population a deja fleuri. Done, l'aptitude reproductrice des
aloes depend de la densite des plantes en fleurs et il y a une selection dans la
dispersion limitee des semences. Par consequent, la distribution spatiale presente
des agglomerats de jeunes populations. Ces groupes torment un noyau a partir
duquelles plants se repandent lentement au cours du temps, avec des individus
adultes qui forment Ie centre des peuplements les plus denses.
Consequence ulrerieure d'une dispersion limitee des semences, la competition
intraspecifique est intense et aboutit a un affaiblissement de la popuation. La
hauteur de A. candelabrum n'augmente pas l'attraction sur les pollinisateurs.
Dans des environnements fermes, la hauteur des aloes sert seulement a permettre