The habitat and associated bird assemblages of the Grey-headed Parrot Poicephalus fuscicollis suahelicus in Limpopo Province, South Africa

Abstract

The Grey-headed Parrot Poicephalus fuscicollis suahelicus has a widespread distribution in sub-Saharan Africa yet is uncommon in its natural range. In some areas seasonal movements are recorded. This study, conducted in north-eastern South Africa, showed that habitat use varied seasonally between two sites, Levubu and Makuya. Differences in habitat were reflected by tree species within the breeding and non-breeding regions of the Grey-headed Parrot. Vegetation structure (tree size; mean height and DBH) was similar between sites, yet tree density differed, being greater at Levubu than Makuya. Bird communities, based on species abundance, differed between localities at each site and between sites. The temporal occurrence of Grey-headed Parrots in particular areas may be a response to the seasonal availability of food sources and nest sites.

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OSTRICH
ISSN 0030–6525
doi: 10.2989/OSTRICH.2008.79.1.2.375
Ostrich 2008, 79(1): x–x
Printed in South Africa — All rights reserved
The habitat and associated bird assemblages of the Grey-headed Parrot
Poicephalus fuscicollis suahelicus in Limpopo Province, South Africa
Craig T Symes1,2 and Michael R Perrin1*
1 School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville 3209, South Africa
2Current address: Department of Zoology and Entomology, University of Pretoria, Pretoria 0002, South Africa
* Corresponding author, e-mail: perrin@ukzn.ac.za
Introduction
The Grey-headed Parrot Poicephalus fuscicollis suahelicus
has a wide distribution from northern Limpopo Province in
South Africa, the Caprivi of Namibia and northern Botswana,
northwards through Zimbabwe, western Mozambique,
Angola, Zambia and Malawi, to north-western Tanzania,
Rwanda, south-western Uganda and south-eastern
Democratic Republic of Congo (Snow 1978, Fry et al.
1988, Forshaw 1989). Distribution covers a broad range of
general woodland habitat types and in some regions
seasonal movements are recorded (Snow 1978,
Wirminghaus 1997, Symes and Perrin 2003a). In the
tropics its range overlaps with the other member of the
P. robustus superspecies group, Jardine’s Parrot
P. gulielmi (Snow 1978). In north-eastern South Africa
there is confusion concerning the distribution of the Grey-
headed Parrot and the congeneric Cape Parrot
Poicephalus robustus, where the two species are thought
to co-occur (Wirminghaus 1997, Downs 2000). The Cape
Parrot is a habitat specialist largely confined to mistbelt
mixed Podocarpus forests in south-eastern South Africa
yet has occasionally been observed feeding away from
forest in human-altered habitats (Wirminghaus 1997,
Wirminghaus et al. 1999, Symes and Downs 2002). A relic
population occurs in the afromontane forests of
Mpumalanga Province (Wirminghaus 1997), with a popu-
lation thought to occur in the Soutpansberg (Barnes and
Tarboton 1998). In this study the habitat of the Grey-
headed Parrot was investigated at two study sites at the
southern limit of its distribution range in north-eastern
Limpopo Province, South Africa.
Grey-headed Parrots occur seasonally in the Levubu
region from August–December where post-breeding flocks
arrive to feed predominantly on mabola plum Paranari
curatellifolia (Symes and Perrin 2003a, 2003b). It was
hypothesised that while in the region they would make use
of higher-altitude afromontane habitats in the Soutpansberg,
habitat specific to the Cape Parrot. Habitats in the region
were also predicted to be distinct from those used during
the breeding season (April–August) in the Makuya region
(Symes and Perrin 2004). Bird communities often share
close affinities with specific habitat types. It was also
hypothesised that the species composition of bird commu-
nities at each site would not differ.
Methods
Study sites
The study occurred at two sites in north-eastern South
Africa (Figure 1). Levubu and Makuya were visited during
August–December 1999 and April–December 2000, when
Grey-headed Parrots were present in each region. In the
Levubu region (23°00’–23°15’ S, 30°05’–30°30’ E) vegeta-
tion is dominated by mixed broad-leaved and fine-leaved
savanna with intensive agriculture (e.g. subtropical crops)
practised in particular areas (Scholes 1997). Riparian
fringes occur along river courses (Scholes 1997). Rainfall
in the region is seasonal, falling mostly in summer
(October–March). Temperatures are hot in summer, but
cooler with increasing altitude in the Soutpansberg moun-
tains. Grey-headed Parrots occur seasonally here from
August to December, when large flocks (up to 50 birds)
with juveniles are conspicuous (Symes and Perrin 2003c).
In the Makuya and northern Kruger National Park area
(22°25–35’ S, 30°50’–31°05’ E) Grey-headed Parrots
breed from April to August (Figure 1; Symes and Perrin
2004). Altitude ranges from c.150–350 m above sea level
(asl) and vegetation is variable from broadleaved wood-
land and riverine vegetation to Mopane bushveld and
The Grey-headed Parrot Poicephalus fuscicollis suahelicus has a widespread distribution in sub-Saharan Africa yet is uncom-
mon in its natural range. In some areas seasonal movements are recorded. This study conducted in north-eastern South
Africa showed that habitat use varied seasonally between two sites, Levubu and Makuya. Differences in habitat were reflected
by tree species within the breeding and non-breeding regions of the Grey-headed Parrot. Vegetation structure (tree size; mean
height and DBH) was similar between sites, yet tree density differed, being greater at Levubu than Makuya. Bird communities,
based on species abundance, differed between localities at each site and between sites. The temporal occurrence of Grey-
headed Parrots in particular areas may be a response to the seasonal availability of food sources and nest sites.

2 Symes and Perrin
mixed bushveld. Rainfall is seasonal and temperatures in
summer may exceed 40° C.
Vegetation sampling
Vegetation was analysed at each study site as a component
of habitat (Figure 1). This was done to investigate the signifi-
cance of certain plant species (important for feeding, breed-
ing and socialising) for the Grey-headed Parrot and to
compare habitats between two separate regions that Grey-
headed Parrots use seasonally. At each site tree species
were identified, tree height estimated and diameter at breast
height (DBH) measured, 5 m each side of randomly walked
50 m straight-line transects. Tree multi-stemming was also
noted. Recent habitat use by humans (i.e. agriculture and
land clearing) can often be determined from tree architecture
(Torquebiau 1986), with trees and regenerating plant growth
indicated by those multi-stemming.
At Levubu two vegetation transects were conducted in an
area where Grey-headed Parrots were observed roosting or
feeding in the past (Prinsloo’s, 23°07.6’ S, 30°20.4’ E, alti-
tude c.620 m asl; Figure 1), and in thick secondary-growth
bush approximately 12 km away, where feeding and social-
ising were observed (Joubert’s, 23°04.3’ S, 30°14.0’ E, alti-
tude c.720 m asl).
At Makuya, three vegetation transects were conducted
near the Luvhuvhu river (Horseshoe, 22°30.9’ S, 31°01.2’ E,
altitude c.300 m asl; Figure 1) where Grey-headed Parrots
had bred previously or were observed feeding, and approxi-
mately 6.5 km away near the Mutale river (Tshikuyu,
22°25.6’ S, 30°59.7’ E, altitude c.350 m asl; Figure 1) where
Grey-headed Parrots were observed feeding and socialising.
An additional locality (Bileni, 22°27.0’ S, 30°56.2’ E, altitude
c.350 m asl; Figure 1) where a failed nesting attempt had
been recorded was also sampled (Symes and Perrin 2004).
At Prinsloo’s, Joubert’s, Horseshoe, Tshikuyu and Bileni
localities, 10, 3, 12, 12 and 5 transects, respectively, were
walked during the respective study periods (Table 1). Analyses
were completed using Statistica (StatSoft 1995).
Bird community sampling
Bird communities were investigated at two localities at each
study site. A single 300 m transect (30 min) was walked 16
times each month from October–December at Levubu and
from May–July at Makuya. Habitat characteristics negated
the use of fixed-width transects. Each day was divided into
four time periods (before 09:00, 09:00–12:00, 12:01–15:00,
and after 15:00). Four transects, each lasting 30 min, were
walked at random times during each time period to account
for temporal variation in different activity patterns of various
bird species and to attempt to detect all bird species pres-
ent in the habitat. A transect was walked each time in both
directions. Presence/absence of all bird species was
recorded visually and audibly, including birds flying over.
Species misidentification may have occurred but the data
Louis
Trichardt
LIMPOPO PROVINCE
Pafuri
ZIMBABWE
Makuya
Park
Punda Maria
1
2
Levubu
4
5
KRUGER
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PARK
3
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22°30’S
23°00’S
31°00’E
30°00’E
AFRICA
South
Africa
SOUTH AFRICA
Limpopo
Figure 1: Study sites in north-eastern South Africa showing localities at each site. Levubu: locality 1= Prinsloo’s, locality 2 = Joubert’s;
Makuya: locality 3 = Horseshoe, locality 4 = Tshikuyu, locality 5 = Bileni

Ostrich 2008, 79(1): x–x 3
are supported by bird ringing at each site (175.8 h at
Makuya; 109.2 h at Levubu; Symes 2004). This helped to
positively identify the presence of particular species and
record species not detected audibly. Relative abundance of
each bird species was calculated by determining the
number of transects each species was recorded each
month (i.e. 16 transects per month). Species recorded out
of transect sessions are indicated in Appendix 1 to indicate
rarer and more cryptic species occurring in the area. Bray-
Curtis Complete Link analysis was used to compare
monthly bird species communities at each locality using
BioDiversityPro (McAleece et al. 1997).
Results
Vegetation
At Makuya, vegetation was dominated by mopane
Colophospermum mopane, whereas at Levubu this species
did not occur (Table 2). At Levubu, although mabola plum
Parinari curatellifolia was not dominant overall, it was domi-
nant at Joubert’s. It was found to occur in moister areas north
of the Luvhuvhu River and was a common tree in Levubu
town. At higher altitudes in the Soutpansberg it was not fruit-
ing. Parinari curatellifolia was the predominant food source in
the area and observations indicate a patchy distribution due
to anthropogenic land use practices (CTS pers. obs.).
Baobabs Adansonia digitata were not common in Makuya yet
were important in providing nesting sites for Grey-headed
Parrots in the area (Symes and Perrin 2004). Baobabs were
not observed in Levubu except for three garden trees (both
DBH <100 cm) (Symes and Perrin 2004).
Similarity of localities, using Euclidean Distances
Complete Linkage Analysis (StatSoft 1995), based on tree
species abundance (Table 2) are represented in Figure 2.
Two clear vegetation groupings were identified with the
breeding area vegetation separated from the non-breeding
area vegetation based on abundance of tree species identi-
fied at each locality (Figure 2). Numerous species were
included as ‘other species’ since they were of little signifi-
cance for feeding and breeding of Grey-headed Parrots
(Symes and Perrin 2003b, 2004). Therefore, less similarity
between Prinsloo’s and Joubert’s was obvious with two
distinct vegetation types at each site. At both sites ‘other
species’ formed a major component of the tree species
identified. At Joubert’s ‘other species’ identified included
quinine tree Rauvolfia caffra, Natal forest cabbage tree
Cussonia sphaerocephala, raisin/cross-berry species
Grewia spp., Entada spicata, syringa Melia azaderach
(exotic) and guava Psidium guajava (exotic). At Prinsloo’s
‘other species’ identified included Grewia spp., common
bush-cherry Maerua cafra, cork bush Mundelea sericea,
forest num-num Carissa bispinosa, magic guarri Euclea
divinorum, common wild pear Dombeya rotundifolia, apple-
leaf Lonchocarpus capassa and Cordia spp. These were
similar to ‘other species’ identified at Makuya. At Tshikuyu
‘other species’ included common star-chestnut Sterculia
rogersi, Grewia spp., Rhigozum spp., false marula Lannea
schweinfurthii and sourplum species Ximenia spp., and at
Horseshoe ‘other species’ identified include Grewia spp.,
Boscia spp. (B. albitrunca, B. foetida and B. angustifolia),
Rhigozum spp., gardenia species Rothmania spp. and
bitterberry species Strychnos spp. At Bileni ‘other species’
identified include Grewia spp., Rhigozum spp., sausage
tree Kigelia africana and Lannea schweinfurthii.
At Levubu and Makuya, at least seven and five food
species sampled accounted for 23.8% and 22.4% of food
trees, respectively (Symes and Perrin 2003b). Of the approxi-
mately nine food tree species identified, five were shared
between sites. However, of these shared species not all were
important foods for Grey-headed Parrots during each season.
Vegetation height differed between localities (Table 1;
ANOVA, df = 4, F= 90.963, p< 0.05) and between localities
at each site (Table 1; ANOVA, Makuya: df = 2, F= 60.778,
p< 0.05; Levubu df = 1, F= 238.42, p< 0.05). Mean tree
height was greatest at Joubert’s, followed by Bileni, Tshikuyu,
Horseshoe and Prinsloo’s. However, overall vegetation
height between Levubu and Makuya did not differ signifi-
cantly (Table 1; ANOVA, df = 1, F= 0.0012, p> 0.05).
DBH of trees >2 m differed between localities (Table 1;
ANOVA, df = 4, F= 40.840, p< 0.05) as well as between
localities at each site (Table 1; ANOVA, Makuya: df = 2,
F= 18.359, p< 0.05; Levubu: df = 1, F= 139.77, p< 0.05).
Mean DBH was greatest at Joubert’s followed by Bileni,
Horseshoe, Tshikuyu and Prinsloo’s. However, overall DBH
of trees between sites differed significantly (Table 1; ANOVA,
df = 1, F= 16.617, p< 0.05).
The number of stems on multi-stemming trees differed
significantly between localities (Table 1; ANOVA, df = 4,
F= 17.396, p < 0.05) and between localities at Makuya
(Table 1; ANOVA, df = 2, F= 12.055, p< 0.05). The number
of multi-stemmimg trees was highest at Horseshoe followed
by Prinsloo’s, Tshikuyu, Bileni and Joubert’s. However, there
was no difference in the number of stems per multi-stemming
tree between localities at Levubu (Table 1; ANOVA, df = 1,
Site Trees Area Density Mean height Mean DBH No. of multi-
sampled sampled (ha) (trees ha–1) (m ± SE) (cm ± SE) stemming trees
Makuya Tshikuyu 528 0.60 880 4.3 ± 0.1 8.2 ± 0.3 213 (40.3)
Horseshoe 565 0.60 942 4.0 ± 0.1 9.0 ± 0.3 250 (44.2)
Bileni 203 0.25 812 5.9 ± 0.2 13.5 ± 1.5 42 (20.7)
1 296 1.45 894 4.4 ± 0.1 9.4 ± 0.3 505 (39.0)
Levubu Prinsloo 768 0.50 1 536 3.9 ± 0.1 6.2 ± 0.2 214 (27.9)
Joubert 156 0.15 1 040 6.9 ± 0.3 14.7 ± 1.1 6 (3.8)
924 0.65 1 422 4.4 ± 0.1 7.7 ± 0.3 220 (23.8)
Table 1: Summary of vegetation recorded at different localities at Levubu and Makuya. Figures given as mean ± SE. Percentage trees with
multi-stemming present given in parentheses

Symes and Perrin4
Scientific name Common name Tshikuyu Horseshoe Bileni Makuya Prinsloo’s Joubert’s Levubu
Colophospermum mopane mopane 48.9 57.3 46.3 52.2 – – –
Androstachys johnsonii Lebombo ironwood 1.5 – 10.8 2.3 – – –
Combretum spp. * Combretum species 17.8 14.0 12.8 15.4 12.5 – 10.4
Commiphora spp. *corkwood species 3.8 1.4 6.9 3.2 5.1 – 4.2
– other species 18.0 19.5 15.8 18.3 13.4 18.6 14.3
Acacia spp. Acacia species 2.7 1.8 – 1.9 4.9 3.2 4.7
– creeper 0.2 – – 0.1 – 18.6 3.1
– snag 0.6 1.1 1.5 0.7 0.3 2.6 0.6
Sclerocarya birrea * marula 1.9 – 1.5 1.0 0.4 – 0.3
Dichrostachys cinerea sickle bush 4.0 – 2.0 1.9 20.3 – 16.9
Terminalia prunioides * lowveld cluster–leaf 0.8 4.6 1.5 2.5 – – –
Adansonia digitata * baobab – 0.4 1.0 0.3 – – –
Terminalia sericea silver cluster–leaf – – – – 26.4 – 22.0
Crossopteryx febrifuga sand crown–berry – – – – 6.1 – 5.1
Pseudostachnostylis maprouneifolia * kudu berry – – – – 2.6 – 2.2
Burkea africana wild seringa – – – – 4.7 – 3.9
Pterocarpus spp. teak species – – – – 2.6 – 2.2
Diospyros mespilliformis * jackal berry – – – – 0.5 – 0.4
Ficus spp. * wild fig species – – – – 0.1 3.8 0.8
Celtis africana white stinkwood – – – – – 11.5 1.9
Bridelia micrantha mitseeri – – – – – 9.0 1.5
Paranari curatellifolia * mabola plum – – – – – 32.7 5.5
*species recorded in the diet of Grey-headed Parrots (see Figure 3 for associated dendrogram)
Table 2: Abundance of tree species at different localities at Levubu and Makuya study sites.

Ostrich 2008, 79(1): x–x 5
F= 0.161, p> 0.05). Between sites the number of stems
per multi-stemming tree differed significantly (Table 1;
ANOVA, df = 1, F= 39.453, p< 0.05).
Bird communities
In total, 201 bird species were identified at the two sites with
63 species (31.5%) common to both sites (Appendix 1).
Including bird species identified out of transects, 97, 103,
113 (20 migrants) and 90 (14 migrants) were recorded at
Horseshoe, Tshikuyu, Prinsloo’s and Joubert’s, respectively.
Many more species were recorded at Makuya despite
sampling occurring when migrants were absent. Twenty-
four migrant species were observed at Levubu during the
study period. Species lists are given in Appendix 1 and
include species recorded at each site when not walking
transects in a particular month. These included species
caught while bird ringing at each site and observed while
monitoring parrots. Relative abundance of each species is
represented by the number of transects each species was
recorded in each month at each locality (Appendix 1). At
Horseshoe, Tshikuyu, Prinsloo’s and Joubert’s, 11, 10, 16
(1 migrant species) and 17 species, respectively, were
recorded in more than 50% of transects at each locality.
Twenty-three species were recorded at all localities.
Seventy-eight, 70 and 31 species were identified at one,
two and three localities, respectively (Appendix 1).
At Makuya and Levubu, 27 (23 species at each locality)
and 12 (10 and 5 species at each locality) secondary
cavity-nesting species were recorded, respectively
(Appendix 1). Not all species were identified as potential
nest competitors of Grey-headed Parrots.
Inter-site similarity based on the relative abundances for
each site per month for the period of sampling is repre-
sented in Figure 3 using Bray-Curtis complete linkage
cluster analysis (McAleece et al. 1997).
Discussion
Vegetation
The Grey-headed Parrot is commonly described as a wood-
land species (Snow 1978, Symes 2001). The difference in
vegetation types sampled in the study reflect the different
types of habitat used by the Grey-headed Parrot. Seasonal
occurrence of Grey-headed Parrots in different habitats
reflect sseasonal food availability (Symes and Perrin
2003b). In this study, Grey-headed Parrots were seen feed-
ing in different habitats from mopane woodland and mixed
broadleaved woodland to disturbed secondary-growth
forest and gardens. The predominant species identified in
vegetation transects was mopane, yet Grey-headed Parrots
were seldom seen in mopane trees. In Levubu the Grey-
headed Parrot is reliant on P. curatellifolia as a food source
(Symes and Perrin 2003b). However, the Grey-headed
Parrot still occurs seasonally in the area despite major
anthropomorphic alteration to habitat through agricultural
development (Symes and Perrin 2003a). During land clear-
ing, large P. curatellifolia trees are often left standing. In
Zimbabwe, the Grey-headed Parrot has been described as
using woodland and occurs in woodland following major
river courses (Benson and Irwin 1966, Irwin 1981). In
Zambia it is a bird of Brachystegia woodland, although it is
not confined to it (Benson and Irwin 1966). In South Africa,
Bird Atlas records report the different habitat types in which
sightings were made (Wirminghaus 1997). Highest vegeta-
tion types reported include Mopane (6.9%), Miombo (5.6%),
Valley Bushveld (2.4%) and Moist Woodland (Wirminghaus
1997). In Mozambique it was not encountered in mopane
woodland and association with this habitat may have arisen
from its occurrence in woodland types overlapping with
mopane woodland (Parker 1999). Also, it may have been
recorded in vegetation types that it was seen flying over on
long distance, early morning feeding forays and late after-
noon returns to roosts (Symes and Perrin 2003c). Grey-
headed Parrots therefore make use of a variety of defined
Figure 2: Complete linkage hierarchical dendrogram (based on
Euclidean distances) reflecting similarity of the respective study
sites based on abundance (%) of most common plant species and
those important to Grey-headed Parrots Figure 3: Bray-Curtis complete linkage cluster analysis reflecting
similarity of respective study sites based on monthly relative abun-
dance of all bird species. Abundance indices were calculated as a
proportion of transects in which each species was recorded each
month. Species recorded out of transect periods were allocated an
abundance index of 0.01 (1.0%) for analysis requirements.
Levubu: J = Joubert’s, P = Prinsloo’s; Makuya: T = Tshikuyu, H =
Horseshoe
0
SIMILARITY (%)
100
50
J-Dec
J-Oct
J-Nov
P-Oct
P-Nov
P-Dec
T-Jul
T-Jun
T-May
HS-Jul
HS-Jun
HS-May
LINKAGE DISTANCE
Joubert's
Prinsloo's
Bileni
Horseshoe
Tshikuyu
020 40 60 80

Symes and Perrin6
habitat types within the woodland-savanna biome region of
southern, south Central and East Africa (Snow 1978,
Forshaw 1989).
The range of the Grey-headed Parrot overlaps with
several general habitat types of open woodland with
mopane and baobab, Brachystegia woodland on Kalahari
sand, open woodland with Brachystegia, Acacia and
Isoberlinia, forest savanna mosaic, Baikiaea dry forest and
moist woodland (Cooke 1962). Its absence from moist ever-
green forest of tropical Africa, when observing distribution
maps, is noted and it is in this habitat that Jardine’s Parrot
occurs (Snow 1978, Forshaw 1989). Therefore, the use of
habitats in two study areas in north-eastern South Africa
that appear superficially distinct is not surprising. If suffi-
cient food trees from the narrow feeding niche of the Grey-
headed Parrot are present, it will be able to persist in a
broad habitat categorisation of seasonal broadleaved
woodland. This is not withstanding additional pressures
such as illegal trade and persecution.
Multi-stemming is likely a response to disturbance confer-
ring advantages to trees exposed to damage, e.g. coppic-
ing as a result of tree felling (Johnston and Lacey 1983). At
Horseshoe (Makuya) this response is likely the result of
large herbivore damage, i.e. African Elephant Loxodonta
africana, and at Tshikuyu (Makuya) a response to tree
felling by local communities for firewood. At Levubu multi-
stemming was significantly lower, a reflection of lower
levels of recent disturbance. Tree felling may in turn affect
tree height and overall community structure and composi-
tion. These effects may have consequences on the distribu-
tion of Grey-headed Parrots in the region, because Grey-
headed Parrots prefer large trees for socialising and
feeding (Symes and Perrin 2003b, 2003c). Regions with
increased anthropogenic disturbance may therefore be
detrimental to Grey-headed Parrot populations.
Boababs, the tree species most important for Grey-
headed Parrot nest sites in north–eastern South Africa,
were only present at Makuya (Symes and Perrin 2003b).
Mabola plum, the most important food tree for post-breed-
ing Grey-headed Parrots occurred at Levubu. Therefore,
regardless of minor habitat differences between sites, so
long as certain criteria regarding breeding and feeding were
met, Grey-headed Parrots utilised the habitat.
Bird community
Bird species are good indicators of habitat type and condi-
tion (Capen 1981). Bird species communities occurring with
Grey-headed Parrots differed at each study site. Species
recorded at all localities are generally common species with
widespread distributional ranges, with those confined to
fewer localities indicating habitat preference and/or greater
rarity. Both study sites are regions of sympatry for a number
of related species. For example Green-backed Warbler
Camaroptera brachyura was recorded at Levubu, where it
was more common in thicker forest vegetation at Joubert’s
than woodland at Prinsloo’s, and Grey-backed Warbler
Camaroptera brevicaudata at Makuya. Barred Wren-
Warbler Camaroptera fasciolata and Stierling’s Wren-
Warbler Camaroptera stierlingi were syntopic at Levubu ,
whereas only Barred Wren-Warbler was recorded at
Makuya. Similarly, Southern Boubou Laniarius ferrigineus
was recorded at Levubu while Tropical Boubou Laniarius
aethiopicus was recorded at Makuya. However, this sympa-
try at Levubu results from the presence of different habitats
used by the different species in the region.
The study region appears as a transition zone for species
falling out in a southerly direction, with the Limpopo River
described as a barrier for the distribution of a number of
woodland species extending their ranges from the north
(Clancey 1994). The Limpopo has been described as an
area where the south-eastern range of Meyer’s Parrot
P. meyeri terminates (Clancey 1994). The Brown-headed
Parrot P. cryptoxanthus occurs on the eastern littoral with a
transition zone between P. meyeri and P. cryptoxanthus
where hybrids are known to occur (Clancey 1977). At
Makuya these species’ were recorded in sympatry. Hybrids
between P. robustus and P. fuscicollis are known to occur in
captivity (W Horsfield pers. comm.) but are unlikely in the
wild because of different habitat requirements. Also, no
specimens housed in museums are suspected to be
hybrids (Wirminghaus et al. 2002).
Similar species, with densities >10 birds 100 ha–1, as
recorded by Parker (1999) for Acacia, miombo, mopane and
other broadleaved woodland, were recorded in this study.
These species include Blue Waxbill, Rattling Cisticola, Dark-
capped Bulbul, White Helmet-shrike, Cape Turtle Dove, Fork-
tailed Drongo, Black-backed Puffback, Black-headed Oriole,
White-browed Robin-Chat and Emerald-spotted Wood-Dove
(see Parker 1999, p. xix for a detailed list of density esti-
mates). A decline in density estimates was recorded in late
winter (July–August) with a peak coinciding with breeding
(September–October) (Parker 1999). In this study, relative
abundances similarly reflected an increase in activity during
the spring and summer, and breeding months for most
species at Levubu (Benson 1963, Dean 1971).
The 23 species identified common to both sites are
indicative of habitat type and not necessarily habitat condi-
tion (Parker 1999). However, Levubu was depauperate of
large cavity nesting species (i.e. hornbill species), indicating
poor breeding habitat for Grey-headed Parrots (Symes and
Perrin 2004). Therefore, in establishing the likelihood of
Grey-headed Parrots breeding in an area, censusing bird
species can be used to assess habitat quality. At Levubu
there were fewer secondary nesting species recorded indi-
cating nest sites in this area are limiting. At Makuya though,
where nest sites may be more abundant, Grey-headed
Parrots are specific, selecting secure cavities in baobabs.
They may also have few competitors since their breeding
season alternates with that of most hornbill species
(Benson 1963, Kemp 1976). Other smaller species may
also be less likely to evict a parrot from a cavity. Mopane
woodland was abundant at Makuya and most likely
provided nest sites for many smaller bird species (Appendix
1; Warburton and Perrin 2005).
To conclude, the Grey-headed Parrot appears unique in its
local movement patterns in north-eastern South Africa, where
their seasonal occurrence within different habitats is strongly
linked with breeding and feeding (Symes and Perrin 2003a,
2003b, 2004). These differences in habitat are apparent in a
number of vegetation characteristics and associated avian

Ostrich 2008, 79(1): x–x 7
communities. However, bird species common to all sites
sampled indicate that these sites may not be heterogeneous
as appears. Most importantly to Grey-headed Parrots is the
presence of certain seasonally abundant fruiting trees (e.g.
P. curatellifolia at Levubu from August–October) and secure
nest sites (e.g. A. digitata at Makuya) during the breeding
season (Symes and Perrin 2003b, 2004).
Acknowledgements — The Loro Parque Fundación (Canary
Islands) is thanked for providing major sponsorship for this project.
The National Research Foundation provided study bursaries for
the duration of the study and Parrot Society (UK) and African Bird
Club provided additional sponsorship. Sarah and Casper Venter of
Louis Trichardt are thanked for their hospitality during the study.
“Boesman” and Marissa Müller, Hendrik and Lynnette Barkhuysen,
and Thinus Prinsloo and Daan Prinsloo of Levubu are thanked for
their hospitality. Wim Botha is thanked for providing a camping site
for the caravan, and ablution facilities at the SAFCOL Offices (Klein
Australie) while working in the Levubu region. Stals and Ronel
König of Tshikondeni are thanked for their hospitality, and to Stals
for a campsite at Iscor, Exploration, Tshikondeni. Iscor and
Limpopo Province Nature Conservation are thanked for granting
access to Makuya Park and National Parks are thanked for allow-
ing work to be conducted in the Kruger National Park.
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suahelicus in southern and south-central Africa. African Journal
of Ecology 41: 299–305
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in Northern Province, South Africa. Emu 103: 49–58
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behaviour patterns of the Grey-headed Parrot Poicephalus fusci-
collis suahelicus Reichenow 1898 in Northern Province, South
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Harrison, DG Allan, LL Underhill, M Herremans, AJ Tree, V
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Received February 2007, accepted August 2007
Editor: A Monadjem

Symes and Perrin8
pecies Status Prinsloo’s Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
PHASIANIDAE
Crested Francolin Dendroperdix sephaena ––––––8114223
Natal Spurfowl Pternistis natalensis ––––101057656
Swainson's Spurfowl Pternistis swainsonii –––0–1––––1–
NUMIDIDAE
Crested Guineafowl Guttera edouardi –––5*81–1–3––
Helmeted Guineafowl Numida meleagris 4120–1–––00–
TURNICIDAE
Kurrichane Buttonquail Turnix sylvaticus ––––––––––2–
INDICATORIDAE
Scaly-throated Honeyguide Indicator variegatus 10*––020*––––––
Greater Honeyguide Indicator indicator 1––––––110210
Lesser Honeyguide Indicator minor 1–––4*53––0–0–
Brown-backed Honeybird Prodotiscus regulus 11–––1––––––
PICIDAE
Golden-tailed Woodpecker Campethera abingoni 355234514–11
Cardinal Woodpecker Dendropicos fuscescens 5* – 1 2 0 1 – – – 1 – –
Bearded Woodpecker Dendropicos namaquus 21––––10–25–
Olive Woodpecker Dendropicos griseocephalus –––0*31––––––
LYBIIDAE
Yellow-fronted Tinkerbird Pogoniulus chrysoconus 1––153––––––
Acacia Pied Barbet Tricholaema leucomelas –––––––1––––
Black-collared Barbet Lybius torquatus 15 14 14 9 11 9 2 7 4 4 8 3
Crested Barbet Trachyphonus vaillantii 3–––––866*265
BUCEROTIDAE
Red-billed Hornbill Tockus erythrorhynchus 1––––––6351178
Southern Yellow-billed Hornbill Tockus leucomelas 11–––––1311108*711
Crowned Hornbill Tockus alboterminatus 1––––––1–––––
African Grey Hornbill Tockus nasutus 1––––––8946138
Trumpeter Hornbill Bycanistes bucinator 1–––––––––244
BUCORVIDAE
Southern Ground-Hornbill Bucorvus leadbeateri 1––––––111––0
UPUPIDAE
African Hoopoe Upupa africana 111––––––1–––
PHOENICULIDAE
Green Wood-Hoopoe Phoeniculus purpureus 1––––––63848–
RHINOPOMASTIDAE
Common Scimitarbill Rhinopomastus cyanomelas 1––––––––––1–
TROGONIDAE
Narina Trogon Apaloderma narina 10––260––––––
Appendix 1: Relative abundance of bird species at two localities at Levubu (Prinsloo’s and Joubert’s) covering months of the non-breeding season (October–December) of the Grey-
headed Parrot, and two localities at Makuya (Horseshoe and Tshikuyu) covering months of the breeding season (May–July) of the Grey-headed Parrot. Relative abundance figure repre-
sents the number of transects each bird species was recorded in either visually or audibly. Sixteen transects were walked per month. Zero represents species recorded out of transect
times for each month. Status: 1 = secondary cavity-nesting species; 2 = intra-African or Palaearctic migrants in the region (total = 24). An aslterisk indicates months in which respective
species were captured and ringed. See Figure 3 for the associated dendrogram. Taxonomy follows Hockey et al. (2005).

Ostrich 2008, 79(1): x–x 9
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
CORACIIDAE
Lilac-breasted Roller Coracias caudatus 1 – –––––832 210
Purple Roller Coracias naevius 1 – –––––362 –––
ALCEDINIDAE
African Pygmy-Kingfisher Ispidina picta 2 – 2* 2* – 0* – – – – – – –
DACELONIDAE
Brown-hooded Kingfisher Halcyon albiventris 910 4 8 3 5 22 6 236
Striped Kingfisher Halcyon chelicuti 1 – 03–1–––3 ––2
CERYLIDAE
Giant Kingfisher Megaceryle maximus –––––––1–––1*
Pied Kingfisher Ceryle rudis ––––––––––0–
MEROPIDAE
White-fronted Bee-eater Merops bullockoides ––––––1351––
Little Bee-eater Merops pusillus ––––––1––11–
European Bee-eater Merops apiaster 2 7 74330––– –––
COLIIDAE
Speckled Mousebird Colius striatus 3* 3 1 14 8 13* – 1 – – – –
Red-faced Mousebird Urocolius indicus 812 2 0 – – 63 – 61–
CUCULIDAE
Jacobin Cuckoo Clamator jacobinus 2 0 –––––––– –––
Red-chested Cuckoo Cuculus solitarius 2991135*7––––––
Black Cuckoo Cuculus clamosus 2 2 25––1––– –––
African Cuckoo Cuculus gularis 2 2 33–––––– –––
Klaas's Cuckoo Chrysococcyx klaas 0220––––0–––
Diderick Cuckoo Chrysococcyx caprius 2 – 34–32––– –––
CENTROPODIDAE
Burchell's Coucal Centropus burchellii –01353225023
PSITTACIDAE
Grey-headed Parrot Poicephalus fuscicollis 1 1 2–95–744 322
Meyer's Parrot Poicephalus meyeri 1 – –––––01– –––
Brown-headed Parrot Poicephalus cryptoxanthus 1 – –––––––– –1–
APODIDAE
African Palm-Swift Cypsiurus parvus 222–2–––––––
Alpine Swift Tachymarptis melba 2 1 ––21–––– –––
Little Swift Apus affinis 2 2 ––––– –– –––
White-rumped Swift Apus caffer 2 ––0–––––––––
Appendix 1: (cont.)

Symes and Perrin10
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
MUSOPHAGIDAE
Purple-crested Turaco Gallirex porphyreolophus 13 16 15 15 16 15 - 0 15 – – 0
Grey Go-away-bird Corythaixoides concolor ––––– –5611103
STRIGIDAE
African Scops-Owl Otus senegalensis 1–––––––00––0
Spotted Eagle-Owl Bubo africanus 0* – – – – – – – – – – –
Pearl-spotted Owlet Glaucidium perlatum 1–––––––00–10
CAPRIMULGIDAE
Fiery-necked Nightjar Caprimulgus pectoralis 0–––– –––––––
COLUMBIDAE
African Olive-Pigeon Columba arquatrix ––––1 –––––––
Laughing Dove Streptopelia senegalensis –0* 1 –– –– 1 –322
Cape Turtle-Dove Streptopelia capicola 10 9 12 – – – 16 14 14 16 15* 15*
Red-eyed Dove Streptopelia semitorquata 7811765202233
Emerald-spotted Wood-Dove Turtur chalcospilos 11* 11* 15 – – 2 2 2* 4* 11 12 11*
Tambourine Dove Turtur tympanistria – – 1 10 11 12– – –1* ––
African Green-Pigeon Treron calvus 247–1 114221–
OTIDIDAE
Black-bellied Bustard Eupodotis melanogaster 0–––– –––––––
RALLIDAE
Buff-spotted Flufftail Sarothrura elegans –0–17 2––––––
PTEROCLIDAE
Double-banded Sandgrouse Pterocles bicinctus ––––– –121010
CHARADRIIDAE
Crowned Lapwing Vanellus coronatus ––––– ––––111
GLAREOLIDAE
Double-banded Courser Rhinoptilus africanus ––––– –––––0–
ACCIPITRIDAE
African Cuckoo Hawk Aviceda cuculoides 10––– –––––––
Black-shouldered Kite Elanus caeruleus 11––– –––––––
Yellow-billed Kite Milvus [migrans] parasitus 2101–1–––––––
African Fish-Eagle Haliaeetus vocifer –0––– –1–––––
White-backed Vulture Gyps africanus –1––– ––1––1–
Lappet-faced Vulture Torgos tracheliotus –––– ––0––––
Black-chested Snake-Eagle Circaetus pectoralis 2–2–– ––––1––
Appendix 1: (cont.)

Ostrich 2008, 79(1): x–x 11
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
Bateleur Terathopius ecaudatus –– ––––3321––
African Harrier-Hawk Polyboroides typus –– ––––1––0––
Lizard Buzzard Kaupifalco monogrammicus 00 0–––––––1–
Dark Chanting Goshawk Melierax metabates –– ––––––––1–
Gabar Goshawk Melierax gabar –– –––––––1––
African Goshawk Accipiter tachiro –2 –021–00–0–
Shikra Accipiter badius –– ––––––––1–
Little Sparrowhawk Accipiter minullus –– –––––––––1
Steppe Buzzard Buteo vulpinus 2–1––1–––––––
Jackal Buzzard Buteo rufofuscus –– –––1––––––
Verreaux's Eagle Aquila verreauxii –– ––––1–––0–
African Hawk-Eagle Aquila spilogaster –– –––––1–––1
Wahlberg's Eagle Aquila wahlbergi –– 1–––––––––
Martial Eagle Polemaetus bellicosus –– ––––0–3–––
African Crowned Eagle Stephanoaetus coronatus –– –001––––––
FALCONIDAE
Dickinson's Kestrel Falco dickinsoni 1––––––0–––––
ARDEIDAE
Grey Heron Ardea cinerea 1– ––––––––––
Black-headed Heron Ardea melanocephala –– 1–––––––––
Cattle Egret Bubulcus ibis 02 20––––––––
SCOPIDAE
Hamerkop Scopus umbretta –– 0–0––1–135
THRESKIORNITHIDAE
Hadeda Ibis Bostrychia hagedash 02 –010––––––
CICONIIDAE
Black Stork Ciconia nigra 2–––––––1–––0
EURYLAIMIDAE
African Broadbill Smithornis capensis –– –1*–0*––––––
ORIOLIDAE
Black-headed Oriole Oriolus larvatus 6 – 3 1 – – 13 10 11 13 12 15*
DICRURIDAE
Fork-tailed Drongo Dicrurus adsimilis 43 3–––8791310*16
MONARCHIDAE
Blue-mantled Crested Flycatcher Trochocercus cyanomelas –– –265––––––
Appendix 1: (cont.)

Symes and Perrin12
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
African Paradise-Flycatcher Terpsiphone viridis 2 0 12* 12* 1 – 2* – – – – - -
MALACONOTIDAE
Brubru Nilaus afer 632–– –––––--
Black-backed Puffback Dryoscopus cubla 15 15* 11 16 16 16 8 13* 10 7 10 9
Black-crowned Tchagra Tchagra senegalus ––526 4––––--
Brown-crowned Tchagra Tchagra australis 4*76–– –614*623-
Tropical Boubou Laniarius aethiopicus ––––– –378126
Southern Boubou Laniarius ferrugineus 3 2 3 14 16 16* – – – – - -
Orange-breasted Bush-Shrike Telophorus sulfureopectus 2–––– 13321--
Olive Bush-Shrike Telophorus olivaceus –– –1111 8––––--
Gorgeous Bush-Shrike Telophorus quadricolor 67 7111212––––––
Grey-headed Bush-Shrike Malaconotus blanchoti –1––– –148*333
White-crested Helmet-Shrike Prionops plumatus 1*–––– –52*46911
Retz's Helmet-Shrike Prionops retzii ––––– ––3*2–43*
Cape Batis Batis capensis –– –12*11 7––––––
Chinspot Batis Batis molitor 576–– –165*574
Black-throated Wattle-eye Platysteira peltata ––––– ––00*–––
CORVIDAE
Pied Crow Corvus albus 010–– 1––––––
LANIIDAE
Red-backed Shrike Lanius collurio 2 ––––– 2––––––
Common Fiscal Lanius collaris –––0– –––––––
Southern White-crowned Shrike Eurocephalus anguitimens ––––– –––––00*
CAMPEPHAGIDAE
Black Cuckooshrike Campephaga flava 257–– –––––0–
PARIDAE
Southern Black Tit Parus niger 1 322–– –323435
HIRUDINIDAE
Barn Swallow Hirundo rustica 2551212 2––––––
Wire-tailed Swallow Hirundo smithii 2 ––1–– –––––––
Greater Striped Swallow Hirundo cucullata 2 –––20 0––––––
Lesser Striped Swallow Hirundo abyssinica 2–1104 4––––––
Mosque Swallow Hirundo senegalensis ––––– ––23–1–
PYCNONOTIDAE
Dark-capped Bulbul Pycnonotus tricolor 16* 16* 16 16 16 15* 16 16* 16* 14 12 15*
Appendix 1: (cont.)

Ostrich 2008, 79(1): x–x 13
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
Sombre Greenbul Andropadus importunus – – 1 16* 16 16* – – – – – –
Yellow-bellied Greenbul Chlorocichla flaviventris 4* 3 12 12* 16 13* 1 11* 11* 6 5 4*
Terrestrial Brownbul Phyllastrephus terrestris ––– 9*410––––––
SYLVIIDAE
Cape Grassbird Sphenoeacus afer –––––1––––––
Long-billed Crombec Sylvietta rufescens ––––––15*2*–22
Yellow-bellied Eremomela Eremomela icteropygialis 231–––––––––
Willow Warbler Phylloscopus trochilus 2 3*69*–––––––––
Arrow-marked Babbler Turdoides jardineii ––––––235–––
Chestnut-vented Tit-Babbler Parisoma subcaeruleum 0–– –––––––––
ZOSTEROPIDAE
Cape White-eye Zosterops capensis 10* 12 10 15* 16 14 – – – – – –
CISTICOLIDAE
Rattling Cisticola Cisticola chiniana 81111–––8*70*–33
Croaking Cisticola Cisticola natalensis – – – – – – 1* – – – – –
Neddicky Cisticola fulvicapilla 1–– –––––––––
Tawny-flanked Prinia Prinia subflava 6* 10 4 14 10 14 – – 3* – – –
Black-chested Prinia Prinia flavicans ––– ––––2––––
Bar-throated Apalis Apalis thoracica – – – 9 14* 16 – – – – – –
Yellow-breasted Apalis Apalis flavida 10* 11 9 5 8 6 – – 4 – – 1
Green-backed Warbler Camaroptera brachyura – 0* – 13* 12 11* – – – – – –
Grey-backed Warbler Camaroptera brevicaudata – – – – – – 10 11 8* 3 3* 4*
Barred Wren-Warbler Calamonastes fasciolatus –16 –––––––––
Stierling's Wren-Warbler Calamonastes stierlingi 0*97–––––1––2
MUSCICAPIDAE
Kurrichane Thrush Turdus libonyanus 10*11*9–1–463*0*50*
Pale Flycatcher Bradornis pallidus –34–––1–––––
Southern Black Flycatcher Melaenornis pammelaina 631––––––––0*
Spotted Flycatcher Muscicapa striata 2 – 1* 9* – – – – – – – – –
Ashy Flycatcher Muscicapa caerulescens ––– 1–––0*––––
Grey Tit-Flycatcher Myioparus plumbeus ––– 0––––––––
White–throated Robin-Chat Cossypha humeralis 2–– –1–––––––
White–browed Robin-Chat Cossypha heuglini ––– 342––2––1
Red–capped Robin-Chat Cossypha natalensis – – – 1 9* 10 – – – – – –
Chorister Robin-Chat Cossypha dichroa –––3*5–––––––
Appendix 1: (cont.)

Symes and Perrin14
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
Bearded Scrub-Robin Cercotrichas quadrivirgata –––1*––––––––
White-browed Scrub-Robin Cercotrichas leucophrys 10* 13* 13 – – 3 12* 12 12* 7 7 5
STURNIDAE
Red-winged Starling Onychognathus morio 1 –1––– – ––––––
Cape Glossy Starling Lamprotornis nitens 1 0–2–– – 641–1–
Meves's Starling Lamprotornis mevesii 1 ––––– –1012151––
Violet-backed Starling Cinnyricinclus leucogaster 1, 2 254–– – –0–1––
Red-billed Oxpecker Buphagus erythrorhynchus 1 ––––– – –001––
NECTARINIIDAE
Scarlet-chested Sunbird Chalcomitra senegalensis 14* 14 13* 5 4 4 – – – – – –
Collared Sunbird Hedydipna collaris – – – 7 11 10* – – – – – –
Southern Double-collared Sunbird Cinnyris chalybeus –––0* 1 – ––––––
White-bellied Sunbird Cinnyris talatala ––0–1 – 2651*45
PLOCEIDAE
Red-billed Buffalo-Weaver Bubalornis niger ––––– – –1––––
Spectacled Weaver Ploceus ocularis – – – 0* 2 7 0* 1 1 – – 1
Southern Masked-Weaver Ploceus velatus 1*11– – – ––––––
Red-headed Weaver Anaplectes rubriceps 1–––– – 210421*
Red-billed Quelea Quelea quelea 2 ––––– – –12046
Yellow Bishop Euplectes capensis 361422 6 ––––––
White-winged Widowbird Euplectes albonotatus ––––– – –––––1
Red-collared Widowbird Euplectes ardens ––––1 1 –––––1
Thick-billed Weaver Amblyospiza albifrons ––27 7 5 ––––––
ESTRILDIDAE
Common Waxbill Estrilda astrild –4484 5 ––––––
Blue Waxbill Uraeginthus angolensis – 1 9 – – – 6* 5* 12* 9 6 7
Green-winged Pytilia Pytilia melba – – – – – – 4 5* 2* 4 5* 2*
Red-billed Firefinch Lagonosticta senegala ––––– – 343*331
African Firefinch Lagonosticta rubricata 23121 1 ––––––
Jameson's Firefinch Lagonosticta rhodopareia – – – – – – 4 3 7* 4 1* 2*
Bronze Mannikin Spermestes cucullatus 1–––– – ––––––
VIDUIDAE
Shaft-tailed Whydah Vidua regia –4––– – ––––––
Pin-tailed Whydah Vidua macroura ––1– – – ––––––
Long-tailed Paradise-Whydah Vidua paradisaea ––––– – –––0––
Appendix 1: (cont.)

Ostrich 2008, 79(1): x–x 15
Species Status Prinsloo's Joubert's Horseshoe Tshikuyu
Oct Nov Dec Oct Nov Dec May Jun Jul May Jun Jul
PASSERIDAE
House Sparrow Passer domesticus 1––––––––––1–
Southern Grey-headed Sparrow Passer diffusus 1 0 0 – – – – 10 13* 14 3 8 12
FRINGILLIDAE
Yellow-fronted Canary Serinus mozambicus 10 7 11 6 7 6 3 2 3 – 1 1
Streaky-headed Seedeater Serinus gularis 0* 1 4 – – 1 – – – – –
Cinnamon-breasted Bunting Emberiza tahapisi –– –– – ––––323
Golden-breasted Bunting Emberiza flaviventris 53* 6 – – – 14* 1 448*
Total 201 83 80 78 66 68 68 66 80 72 64 75 66
Appendix 1: (cont.)