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Afr. J. Eeo!. 1995, Volume 33, pages 89-100 "'::-.~
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Road kills of animals by public traffic in Mikumi N atio ~'-.
Park, Tanzania, with notes 011 baboon mortality
CARLOS DREWS*
Department of Zoology, UniversitJ' of Cambridge. Cambridge CE23EJ, U.K

Summary
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

Resume
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
provoquent regulierement des accidents dans Ie Pare National de Mikumi. On
recommande de placer des panneaux routiers ct des casse-vitesse pour reduire la
vitesse moyenne, pour la securite des gens et des animaux.
Introduction
'v1ikumi National Park in Tanzania is one of a number of African conservation
areas which are traversed by a public road. The TANZAM (Tanzania-Zambia)
highway traverses the reserve over a stretch of 50 km. Anecdotal reports
of vehicles killing elephants (Loxodonta ajricana Blumenbach), giraffes
(GiraOu camelopardalis Linnaeus) and hippopotamus (Hippopotamus amphibius
Linnaeus), among others, indicated that the traffic was having an impact on the
protected wildlife. Although public highways and railways pass through nature
reserves in Africa (e.g. Kenya: Tsavo National Park, Marsabit National Park;
Tanzania: Serengeti National Park, Selous Game Reserve; Uganda: Queen
Elizabeth National Park; Zaire: Virunga National Park; Zambia: Kafue
National Park), the range of species affected by vehicle traffic and the demo­
graphic implications of road kills has rarely been quantified in this continent
(1. Lopez, pers. comm.; see review of road-kill literature by Lopez & Roviralta,
1993). This study complements the scarce information available about road kills
in Africa (e.g. Broekhuysen, 1965; Lewis, 1989; Drews, 1991).
The highway was in operation prior to the establishment of Mikumi National
Park in 1964, 300 km south-west of Dar es Salaam. The park was expanded in
1975 to a present area of 3300 km 2 , which includes grassland, open woodland,
miombo and montane rain forest (Norton et al., 1987; Lovett & Norton, 1989).
The highway is of great economic importance in that it provides the main access
to southern Tanzania and its bordering countries of Zambia and Malawi, which
utilize the road for the transportation of goods to and from the port of Dar es
Salaam. Drivers in transit through the park can contemplate African wildlife at
no cost, an unusual privilege given the dwindling number of large mammals in
the rural areas of Tanzania.
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
average speeds of 30 km h - I. By the end of 1991, however, buses, lorries and
private vehicles passing at speeds above 90 km h ~~ 1 on the immaculate tarmac
were common. During the same period, information on the occurrence of road
kills within the national park boundaries was collected in order to assess (i)
which taxa are affected by traffic in Mikumi; (ii) the road kill rate; (iii) the effect
of an improvement in road condition on road kill rates; and (iv) to quantify the
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.
Methods
Road-kill data were collected during a study on the behaviour of yellow baboons
in Mikumi National Park. The two-lane public highway is the only paved road
Road kills in a Tanzanian nalional park 91
in the park. Between 16 May 1990 and 16 August 1991 the species and location
of each road kill encountered were recorded while driving at 40 km h - \ on the
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
average, a 20-km stretch (40'Yo of the highway within the park) was sampled
regularly. The northern sector of the road was sampled much less often. Sampling
occurred on 65% of the total number of days covered by the study period. In
order to estimate the absolute rate of road kills along the 50-km road stretch
within the park, the number of kills recorded was corrected for the proportion of
highway covered during regular sampling as well as the proportion of days
during which it was surveyed. This yields a correction factor of 3·75. Road kills
generally disappeared from the road within one day as a result of the actions of
scavengers. Occasional road-kill reports by other drivers are also included in this
study. The recording effort remained more or less constant throughout the study.
The records were limited to the more conspicuous animal classes: mammals, birds
and reptiles. However. frogs and invertebrates, particularly ants, grasshoppers,
and millipedes were also commonly found killed by vehicles. The speed of
vehicles was measured by driving behind them and reading the speedometer.
Unfortunately, the road toll office in Mikumi town was closed down during the
study, and highly desirable information on traffic rates became unavailable.
Quantitative information on road proximity and group size of baboons was
used to assess the possible demographic effects of road kills on a gregarious
mammal. A visit to the highway by a baboon troop was defined as any instance
in which at least 10% of troop members was within 10 m of the road. Baboon
troops were followed each day for seven hours, on average 0745-1545 h). When
a troop visited the highway more than once during an observation day (9% of all
cases, N= [44). the single durations of these visits were added to calculate the
daily stay on the road (Table 3). The highway stretch considered for this analysis
was 20 km long: from 15 km north of the park headquarters, to 5 km south of
the headquarters. Non-parametric tests were used in all statistical analyses.
Results
Location of road kills
A total of 183 road kills were recorded within the national park over 15 months.
The bulk (98°/,,) of the road kills occurred on the public highway. Kills were
particularly common along road stretches near waterholes and where traditional,
small-scale migration tracks cross the highway. Three birds (2%) were found
dead on the unpaved tourist road system.
Taxa
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
(Viverra cil'effa Schreber), followed by spotted hyaena (Crocuta crocufa
Linnaeus), yellow baboon and African buffalo (Syncerus caffer Sparrman). Other
92 C. Drews
Table I. Number of road kills
for each species recorded in
~likumi National Park.
Tanzania (16 May 1990-16
August 1991)
Species
Birds
Birds of prey
Gyps aFicanus Salvadori
Gypoheirax angolensis Gmelin
Circus ranivorus Daudin
Aquila wahlbergi Sundevall
Mil1'lIs migrons Boddaert
Falco dickinsoni Sclater
Aquila rapax Temminck
Glaucidium capense Smith
unidentified vultures
unidentified raptors
Others
Coracias cal/data Linnaeus
SlI'cplOpelia sp. Bonaparte
Merops apiasler Linnaeus
Melaenomis pallidus Muller
Cenlropus superciliosus Hemprich & Ehrenberg
Hirumlo sp. Linnaeus
,Numida meleagris Linnaeus
GUllera pucherani Hartlaub
Francolinus afer Statius Muller
Cicon;a ciconia Linnaeus
Macrodipleryx vexillaria Gould
Macronv.\' sp. Swains
Prinia subf/ava Gmelin
Lanius sp. Linnaeus
Lamprolomis chalibaeus Hemprich & Ehrenberg
unidentified birds
Total bird road kills
55
Number of species=23
Mammals
Nocturnal
Viverra civella Schreber
Crocuta croc1I1a Linnaeus
Genella sp. Oken
Hystrix crislata Linnaeus
Ietonyx striata Perry
Triaenaps pers!c'a Dobson
Diurnal
Papio cynocephalus Linnaeus
Lycaon pictus Temminck
Plwcoc!lOerus aelhiopicus Pallas
Other
Syncerus caffer Sparrman
Aepyceros melampus Lichtenstein
Road kills in a Tanzanian national park 93
Table I. Conlinued
Number of Number of
kills Species kills
Mammals continued
Other con I inued
2 Lepus sp. Linnaeus 6
1 Canis sp. Linnaeus 5
2 Giraffa camelopardalis Linnaeus
1 Helogale parvula Sundevall 2
i Loxodonta aFicana Blumenbach 2
1 Connochaeles taurinus Burchell o
1 Panlhera pardus Linnaeus 2
I Procavia capensis Pallas
3 Cephalophus sp. Hamilton Smith
2 Mel!ivora capensis Schreber
Panthera leo Linnaeus
8 Total mammal road kills 111
Number of species=22
4
3
Reptiles
2
Lizards
2
Varanus exanthematicus Schmidt 2
Varanus sp. Merrem
Turtles
Pelomedusidae
1 Snakes
Python sebae Gmelin 2
1
Rhamphiophis oxyrhynchus Reinhardt 2
1
Bilis Gl'ielans Merrem
I
Naja nigricol!is Linnaeus
10
Psammophis phillipsii Hallowell
Psammophis sp. Boie
unidentified snakes
Total reptile road kills 17
Number of species=7
20 Total of road kills recorded
17 (III! taxa) 183
2 Total number of species
identi{!ed= 52
mammals recorded more than twice include impala (Aepyceros melampus
Lichtenstein), hare (Lepus sp. Linnaeus), jackals (Canis sp. Linnaeus), giraffe
15
and African hunting dog (Lycaon pictus Temminck). The lilac-breasted roller
3
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
white stork (Ciconia deania Linnaeus) and the European bee-eater (Merops
11 apiaster Linnaeus). Birds of prey accounted for 27% of all bird records. Most
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
50
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
40
Troop On road Size Males Females Juvs. Dates of census
if)
>Q
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

.
i<
-0
20 Yuma Hill F 60 6 17 37 27 April 1991
0
0
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
MAY JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR APR MAY JUN JUL Lodge R 20 2 6 12 23 July 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

Road kill rate
Data from the last 90 days of the study (N= 73 road kills) yield a rate of 91 road
kills per month, i.e. about three road kills per day within the park boundaries
during that period. This figure is equivalent to 21·8 road kills/km/yr, consisting
of 13·3 mammals/km/yr, 6·5 birds/km/yr and 2·0 reptiles/km/yr. These rates were
calculated by taking into account the percentage of road distance sampled
regularly as well as the proportion of days when it was sampled (see Methods).
The rate calculated above is probably an underestimate given that an unknown
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
and perhaps large mammals which were not killed outright and died later from
injuries obtained on the road.
Over the course of the study, the number of road kills increased significantly
(Spearman's r=0'878, N= 15, P<O'OOI; Fig. I). This positive correlation was still
significant when the extremely high value for July 1991 was omitted from the
sample (Spearman's r=0'849, N= 14, P<O·OOI). The road-kill rate was five times
higher at the end of the study than at the beginning (first three months N= 15,
last three months N=73). The obvious correlate of this increment is the
improving conditions of the road during the study period and the associated
increase in average driving speed. The road work in the park started early 1990
at the northern boundary. During the second half of the study the new paving
started to include the main sampling road stretch and in November 1991 it
reached the southern boundary of the park.
During March, April and May 1991, the condition of the road where a kill
was found was recorded. Out of 30 records, 23 (77%) were found on good
tarmac, whereas only seven occurred on the temporary dirt tracks to which the
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
associated with significantly more road kills (X 2 = 5,69, df= I, P<0·05).
It is noteworthy that doves (Streptopelia sp. Bonaparte) and impalas started
to appear repeatedly in the records during the last months of the study when
most of the highway had been newly paved. These animals are extremely agile
and fast, which suggests that they are killed only by vehicles going at high
speeds. On 3 November 1991 the road work inside the park was completed.
Subsequently, vehicles going at more than 90 km h - I were regularly seen on the
50 km of excellent tarmac that traverses the park.
Demographic effect of road kills in yellow baboons
The sample for this analysis is composed of, firstly, 15 baboons found killed
within a 20 km road stretch over a period of 15 months, and secondly, of 13
troops that were seen more or less regularly within that 20 km road stretch
(Table 2). The size of troops which were seen frequently on the road did not
differ significantly from those troops which were only seen rarely on the highway
(Mann-Whitney U-test, two-tailed, P=0·67). The estimated total of baboons
killed along the 20 km stretch is 23, when the fact that the road was not sampled
on 35% of the time is taken into account (i.e. 15 x 1'5).
Three adult males and 10 juveniles, no infants and no adult females, were
found among the dead baboons. The pooled troop compositions of nine troops
(Table 2) indicate that the proportion of juveniles to adults in the subpopulation
is 139 (51%) to 131. The proportion of juveniles among the road kills (77%) was
higher than in the overall population. This difference just missed statistical
significance (X 2 =3'309, df= I, P=0·0653). A larger sample size is required for a
more powerful test of the susceptibility of certain age and sex classes to be killed
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
the vehicles. The most commonly killed mammals were nocturnal species,
namely African civet and spotted hyaena. Although not strictly nocturnal,
buffalos too were generally killed at night. Interestingly there were no small
rodents among the road kills in Mikumi National Park (cf. Lewis, 1989). This
taxonomic group, together with domestic carnivores, represents the majority of
mammal records in some road-kill studies in Europe (e.g. Frean et al., 1993;
Lopez, 1993).
The relatively high number of killed yellow baboons, a strictly diurnal
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
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
food. On at least four occassions baboon troops stayed on or next to the road for
up to 3·5 h when maize fell from passing lorries. Baboons utilized the highway in
additional ways, including walking on the tarmac to cover long distances and
foraging on the lush roadside vegetation. Reptiles seek the warm tarmac to
absorb heat. This basking behaviour on the road makes them particularly
vulnerable to passing vehicles.
Overall. a large proportion (27%) of the killed birds were birds of prey. These
exemplify the fact that road kills facilitate more road kills: dead animals on the
road attract scavengers, which during the day are mainly raptors and vultures.
These are, in turn, killed next to the carcass by passing vehicles. Some of the dead
jackals (Canis sp.) and hyaenas were also in the proximity of fresh carcasses.
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
negligible. The increase of the road-kill rate was associated with the rehabilitation
of the highway which, in turn, led to higher average driving speeds. Also, the
additional traffic resulting from the vehicles of the road workers could have
contributed to the increase in road-kill rate. By the end of the study, the rate of
large mammals killed on the road probably exceeded that of poached wildlife in the
park (D. Njao, warden in charge of anti-poaching in Mikumi N.P., pers. comm.).
Highway mortality is evidently one more demographic parameter in some of the
populations which include the road in their range. For some of the baboon troops
which frequently visit or cross the highway, road kills may have a measurable
influence on survivorship, potentially accounting for about 10% of annual losses.
The relatively high incidence of bone fractures among the adult male baboons of
the Highway troop is probably associated with their high frequency of road visits
(pers. obs). Although not fatal, impacts by vehicles are impairing these males in
respect of their competitive ability and reproductive performance.
The problems associated with road kills in African protected areas are similar
to those identified in a recent, nationwide survey of road kills in Spain
(Coordinadora de Organizaciones de Defensa Ambiental CODA, 1992;
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
of collisions with large animals within the park boundaries is increasing as traffic
on the TANZAM highway increases in parallel with the development of
southern Tanzania and its neighbouring countries. In addition to human
casualties, the risk of pollution in the park and of considerable freight and
Road kills in a Tanzanian national park 99
vehicle losses in wildlife-related accidents should be reduced. Secondly, the list of
road kills included at least two endangered species, namely African hunting dog
(IUCN, 1990), and African elephant (Appendix I of CITES, 1991). Five
elephants are known to have been killed on the highway inside the park between
1989-1992 (Mikumi National Park staff, pers. comm.). The killing of endangered
animals is of immediate concern to conservation-oriented management plans.
Thirdly and finally, from an ethical point of view, every individual animal
deserves protection from unnecessary, human-induced suffering or death.
Recommendations
Undoubtedly, the best way of reducing the rate of collisions between vehicles and
animals is to reduce the driving speed. The existing speed limit of 80 km h - I on
the TANZAM highway is too high within the park. A speed limit of 50 km h - I,
which is the limit in all Tanzanian national parks, should be implemented on the
public highway within the park boundaries. Adequate signposts at regular
intervals should remind the drivers to reduce speed and of the presence of wild
animals on the road. The most effective way of reducing average driving speed,
however, is speed bumps. These should be placed close to areas identified as 'hot
spots' for road kills, such as waterholes and traditional, frequently used animal
tracks. A further step towards reducing additional road kills is to remove from
the road any carcass found, so that its scavengers are not exposed to the traffic
(also Barragan & Lopez, 1993).
Acknowledgments
I am grateful to the Tanzanian Commission for Science and Technology,
Tanzania National Parks, and the Serengeti Wildlife Research Institute for
permission to work in Mikumi National Park. The national park authorities and
rangers provided invaluable logistical support and background information
about the road-kill problem. The Long Term Mikumi Baboon Project kindly
provided unpublished demographic information on two of the baboon troops
cited (Viramba 1 and 2) as well as accommodation in the park. Special thanks to
Astrid Schmidl-Drews, Charles Kidung'ho, William Marwa and Ayoub Njalale
who helped regularly in spotting, identifying and removing carcasses from the
road. I thankfully acknowledge Michael O'Meara, who collected road-kill
information during by temporary absence in October-November 1990. My
sincere thanks to all those who reported road kills to me throughout the study,
and to Keith Eltringham, Dawn Hawkins, Guy Norton, Sam Wasser, Javier
Lopez and two anonymous referees for their valuable comments on a previous
draft. The Leakey Foundation, SigrnaXi, Durham Fund, Bedford Fund and
Kings College, Cambridge made this study possible by funding my stay in
Mikumi National Park.
References
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(Manuscript accepted 3 March 1994)
Afr. 1. £Col. 1995, Volume 33, pages 101-113
Population dynamics, pollination ecology and the
significance of plant height in Aloe candelabrum
C. J. STOKES and R. 1. YEATON
Department of Botany and UN-FRD Research Unit for Plant Growth and Development.
University of Natal, Pietermaritzburg 3200, South Africa
Summary
Pollinator exclusion experiments showed that insect pollination of Aloe
candelabrum Berger is negligible and that birds are the only effective pollinator
guild. Bird activity is greatest where flowering aloes are most dense. Also fruit set
is lower in those aloes that bloom later in the season after most of the population
has already flowered. Thus the reproductive fitness of aloes is dependent on the
density of flowering plants and there is selection for limited seed dispersaL As a
result, spatial distributions show clumping in young populations. These clumps
act as nuclei from which plants spread slowly over time with mature individuals
forming the centres of the densest stands.
As a further consequence of limited seed dispersal, intraspecific competition
is intense, resulting in self-thinning of the population. The height of A.
candelabrum does not enhance pollinator attraction. In closed environments,
aloe height serves only to allow the persistence of those plants that establish
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
Resume
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