Distribution
and habitat use of Crested Larks
Galerida cristata at Loiyengalani,
Kenya
Leon Bennun, Luca Borghesio, Luca Biddau, Edward Waiyaki and Luc Lens
The Crested Lark Galerida cristata
is
a widespread and polytypic species
(Cramp 1988).
In Africa, the
bird ranges widely to the north and south of the Sahara, along the Nile Valley, and
—
—
the race
Kenya, southeastern Ethiopia, and northern Somalia (Keith et al. 1992). In
Kenya, Crested Larks are common in sandy semi-desert in the northwestern quarter of the country
(Lewis & Pomeroy 1988). The species is considered mainly resident (Cramp 1988, Keith et al. 1992).
In Europe, family units persist until autumn (Dolgushin et al. 1970) and movements away from the
natal area involve wandering juveniles (Labitte 1957). Few data are available on Crested Lark densities
and distribution patterns outside the breeding season. Scattered observations mainly involve single
individuals or small flocks (Abs 1963, Kozlova 1975, Viulleumier 1979). Temporal aggregations and
short-distance movements are assumed to be related to bad weather (Porter et al. 1969), abundant food
supply (Cramp 1988) and/or water supply (Rustamov 1958).
somaliensis
in northern
In this paper,
we
(i)
introduce a
new census
technique, specially designed for standardized counts
of ground-dwelling passerines in arid areas, and
(ii)
analyse non-breeding population densities and
spatio-temporal distribution patterns of Crested Larks in relation to the characteristics of semi-desert
habitat near Loiyengalani,
Lake Turkana.
Study area and methods
Lake Turkana, 250
is
km long and reaching the Ethiopian border along its northern end,
the largest of the lakes in the
region
is
April) and
high, with
1992,
Kenyan
Rift Valley
characterized by extreme dryness (<200
often subject to long periods of drought
is
mean maxima over 32°C (Hopson
maximum
area
was
some
Hughes 1992). The
(Hopson 1982). Temperatures
are
1982); during our study in February
be considered a semi-desert
3.2
(Pratt et al. 1966).
in the southeastern part of the basin, close to the fishing village
of Loiyengalani (2°46N, 36°43E).
shoreline
&
daytime temperature ranged from 35 to 39°C. With these climatic
features, the region can
Our study
(Hughes
mm annual rainfall, peak March-
km long,
We
and inland
censused Crested Larks along a stretch of
to
some
3
km from the lake. Most of this
area
consisted of large sand plains with sparse, scattered vegetation and variable amounts of
pebble and rock. Acacia
tortilis is
the dominant tree, with
Doum palm Hyphaene
compressa and thickets of Salvadora persica growing around springs and along dry
riverbeds.
Meadow-like Sporobolus spicatus grass covers a
belt
some hundreds of
metres inland from the lake, petering out as one moves further away from the shore.
Crested Lark populations were sampled between 14 January and 18 February 1992,
using a specially-designed technique that
we termed
a circular-plot count (CPC).
To
perform a CPC, one observer stands stationary holding the end of a stretched string
56.4
m long
(i.e.,
the radius of a 1-ha circle).
The other walks around in a circle holding
the opposite end. Birds are flushed or observed as the string passes over them, and all
the birds seen inside the circle are recorded
CPCs
in
50
different plots.
For each
CPC
date, time of day, percentage of terrain
by the
central observer.
We carried out 78
the following parameters
were recorded:
covered by grass/pebbles, and distance from
-69-
Leon Bennun
70
human dwellings/lake (estimated to
(HMSO, 1977) where appropriate).
et al.
the nearest 10 per cent, with the aid of a
map
Results
Table
1
shows the
results of a Poisson regression
model, selecting the best predictors
of the number of birds observed in the different CPCs. The parameters of the model
maximum
were estimated by
goodness-of-fit of a
in the
likelihood,
model including
model were date (10
which provides a convenient index of
a particular set of parameters. Variables included
different sampling days), period of
day (morning: 07:00-
11:00; mid-day: 11:00-15:00; evening: 15:00-19:00), percentage of terrain covered
by grass, percentage of terrain covered by pebbles, distance from human dwellings and
distance from the lake.
program
GLIM (NAG
We
used a stepwise backward procedure
1986) to select a
final
in the statistical
model containing only
significant
parameters. In the final model, period of day and percentage of terrain covered by grass
significantly affected the
number of
birds observed, with period being the strongest
predictor.
Table
1.
Poisson regression model selecting parameters which best predict the number of
CPC. All parameters are tested with a stepwise backward procedure,
birds per
(approximately equivalent to scaled deviance) when a
calculating the change in
parameter
is
excluded from the model.
Parameter
*,
Adf
P < 0.05
P
Percentage pebbles
0.58
1
0.45
Distance from habitation
1.54
1
0.21
Distance from the lake
1.88
1
0.17
15.35
9
0.08
4.04
1
0.04*
13.89
2
0.001*
Date
Percentage grass cover
Period of day
= 0.35 (± 0.21) + 0.008 (± 0.003)
0.44 (± 0.26) period of day.
Selected model: G(x.)
grass cover
-
The percentage of grass cover was positively, though weakly, related to number of
CPC (« = 78, r = 0.215, P = 0.059). To examine the effect of period of day on
birds per
number of birds per CPC in more detail, we selected 14 plots for which CPS were
made in all three periods. The number of birds per CPC markedly decreased from
the
morning
to
mid-day, and increased again from mid-day to evening, leaving no
statistical difference
between morning and evening (Repeated measures
F2 2,= 5.02,/' = 0.014; Fig.
ANOVA:
1).
The presence of human dwellings did not tend to affect the distribution pattern of
Crested Larks. However, our CPC technique could not be applied within villages. We
often observed Crested Larks gathering around shops or houses were food was
available,
and densities could have been high.
Distribution
and habitat use of Crested Larks
71
1. Mean number of Crested Larks
per plot, estimated with the circular plot
count technique, in 14 plots counted in
each period of the day. The V -values
shown correspond to single paired t-tests
Figure
morning
mid-day
Time
The mean
of
evening
day
overall density of birds (individuals ha
')
during the course of the day,
estimated through CPCs, was 1.45. In the selected Poisson model, distance from the
number of
lake did not significantly affect the
densities
were recorded
in a belt of
100
birds per
although only during the morning period (Two-way
3.64,
P = 0.031;
CPC. However, higher
m from the shore compared to further 'inland',
ANOVA:
factor Period,
from Lake (<100 m versus >100 m), F^ .^^=
F^^^= 2.68, P = 0.07).
factor Distance
0.001; two-factor interaction,
/^2 72
~
UM^ P =
Discussion
Crested Larks are the commonest passerines around Lake Turkana. If both the overall
density observed in the course of our study (1.45 individuals ha
structure are similar to those
an area of 2000 km^
(i.e.,
lake), the population
found in other parts of the region,
a belt of about 4
amounts
to about
km
extended
300 000
all
')
and the habitat
we can estimate that
individuals. This very tentative
estimate could be improved by carrying out additional counts in other areas.
technique would be appropriate for
passerines in open habitats.
and accurate
CPCs
It
this,
in
around the shores of the
The
CPC
and for censusing other ground-dwelling
proved easy to perform, and appears
to give consistent
results.
carried out at different distances
movement between
from the lake suggest a circadian pattern of
the lakeshore and the surrounding areas.
Our
the birds were concentrated along the lakeshore during the early
results suggest that
morning hours. Some
of these birds apparently scatter inland during the day. During the hottest hours,
when
CPC numbers were at a minimum, the birds may concentrate around dwellings,
in the
shade of rocky
cliffs,
or at favoured drinking sites around springs or standpipes.
observed concentrations of birds
census with the
CPC
technique.
at all
One
We
these types of sites, but they are difficult to
lark,
captured and ringed
at
a drinking-site,
was
Leon Bennun
later
observed feeding
km away,
1
so
et al.
movements could be
substantial.
The afternoon
appears to see the larks feeding in the open again as they begin a generalized
movement back towards
the lakeshore.
Acknowledgements
This study was carried out during an expedition organized by the Istituto Nazionale Biologia
Museums
Selvaggina, Bologna, Italy; the National
Universita Pavia, Italy.
Lombarde
(Pavia),
We
of Kenya, and the Dipartimento Biologia Animale,
thank Ferrino (Torino), Zeiss (Milano), Cassa Risparmio delle Province
Agip (Kenya) and
Olivetti
(Kenya) for support, and James Wachira, Patrick
field. We are grateful to Michel Louette for his
Gichuki and Syingi Kathele for assistance in the
comments on
earlier drafts of this paper.
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Museums of Kenya, Box 40658, Nairobi
Received 10 September 1996