Ornithological Observations
An electronic journal published by the Animal Demography Unit at the University of Cape Town
Ornithological Observations accepts papers containing faunistic information about birds. This includes descriptions of distribution, behaviour, breeding, foraging, food, movement, measurements, habitat and plumage. It will also consider for publication a variety of oth
relevant ornithological material: reports of projects and conferences, annotated checklists for a site or region, specialist other interesting or relevant material.
Editor: Arnold van der Westhuizen
INTERSPECIFIC COMPETITION
Recommended citation format:
Jubber WR 2014. Interspecific competition between Oxpecker Buphagus URL:
Ornithological Observations
the Animal Demography Unit at the University of Cape Town and BirdLife South Africa
accepts papers containing faunistic information about birds. This includes descriptions of distribution, behaviour, breeding, foraging, food, movement, measurements, habitat and plumage. It will also consider for publication a variety of oth
relevant ornithological material: reports of projects and conferences, annotated checklists for a site or region, specialist bibliographies, and any
Editor: Arnold van der Westhuizen
ITION BETWEEN OXPECKER BUPHAGUS SPECIES?
WALTER RALPH JUBBER
Buphagus species? Ornithological Observations, Vol 5: 325-335.
URL: http://oo.adu.org.za/content.php?id=142 Published online: 04 July 2014
BirdLife South Africa
accepts papers containing faunistic information about birds. This includes descriptions of distribution, behaviour, breeding, foraging, food, movement, measurements, habitat and plumage. It will also consider for publication a variety of other interesting or bibliographies, and any
INTERSPECIFIC COMPETITION BETWEEN OXPECKER BUPHAGUS SPECIES?
Walter Ralph Jubber*
Assistant Manager, Beho Beho Camp, Tanzania
* Corresponding author: [email protected] Introduction
Oxpeckers are well known for their tick foraging capabilities. They have also been seen as possible parasites on their selected host ungulates, therefore sometimes referred to as vampires (Weeks, 2000). Most of the wound opening activity, which cou detrimental to the hosts, has been seen more captive and domesticated species, rather than in natural environments
2009).
Two species of oxpecker occur in the world, both of which occur in Africa, these include: Yellow-billed Oxpecker Buphagus africanus and Red-billed Oxpecker B. erythrorhynchus. They are related to Starling family, Sturnidae; however some may argue that there is enough evidence to place them within their own family (Fry 2000; Hockey et al. 2006).
The main difference between these two species, is that the Red billed Oxpecker has a complete red beak, whereas as Yellow Oxpecker has a yellow base with a red tip. The Yellow Oxpeckers also has a pale rump, where Red-billed Oxpeckers the rump the same colour as the body (Hockey et al. 2006).
However in areas where both oxpecker species occur, what govern their specific interest in particular host ungulates, and what prevent them from competing against one another?
ITION BETWEEN SPECIES?
Assistant Manager, Beho Beho Camp, Tanzania [email protected]
heir tick foraging capabilities. They their selected host ed to as vampires (Weeks, of the wound opening activity, which could be more captive and domesticated species, rather than in natural environments (Plantan,
both of which occur in Buphagus africanus . They are related to the Starling family, Sturnidae; however some may argue that there is enough evidence to place them within their own family (Fry et al.
s that the Red- reas as Yellow-billed
. The Yellow-billed billed Oxpeckers have
2006).
oxpecker species occur, what govern host ungulates, and what prevent
Fig 1 – A Yellow-billed Oxpecker (left) and a Red-billed Oxpecker (right) show the different characteristics between the species.
© Trevor Hardaker
Ecologically no two species within the same area
same ecological or realised niche for a long period, otherwise one will out compete the other (Hardin, 1960). Some reference is
body mass and surface area ratio of the host specie higher percentage of ectoparasites, and theref
tick yield. For the Yellow-billed Oxpecker their slightly larger body mass may indicate slightly greater needs than
Oxpecker, therefore a higher tick yield of engorged adult ticks and immatures may be more beneficial (Mooring & Mundy, 1996; Nunn al. 2011). In captivity Yellow-billed Oxpeckers
billed Oxpecker (right) to different characteristics between the species.
wo species within the same area can hold the exact ong period, otherwise one will out compete the other (Hardin, 1960). Some reference is made to body mass and surface area ratio of the host species, thus having a higher percentage of ectoparasites, and therefore having a greater their slightly larger body than that of the Red-billed a higher tick yield of engorged adult ticks and (Mooring & Mundy, 1996; Nunn et xpeckers was recorded as
eating 13 000 nymph ticks or 100 engorged adult ticks
billed Oxpeckers 12 500 tick larvae or 100 engorged adult ticks day (Stutterheim et al., 1988). Both oxpeckers have been observed on a large variety of ungulates, and in some studies it is shown that the presence of Yellow-billed Oxpeckers on Impala
melampus has assisted with reducing their grooming needs by 36% (Mooring & Mundy, 1996). It is well documented though that, larger ungulates are definitely more preferred by Yellow Oxpecker.
In cases where both oxpecker species have been seen on th host species together, principally in captivity, and in particular Greater Kudu Tragelaphus strepsiceros the Yellow-billed Oxpecker was always dominant. However, few interactions like th
been observed in the wild (Stutterheim & Panagis, 1988).
This study was done to:
· establish which species within the study area are host(s) to one or both oxpecker species;
· investigate which host species they share;
· explore any interspecific competition between the two oxpecker species; and
· verify feeding requirements and needs as observed by Mundy (1996) and Nunn et al. (2011). These researchers
at body mass as well as parasite yield of the host species and to what extent these factors influence preference by the oxpeckers Study Area
The study area is the Makuleke Contract Park in nor
National Park. The area is situated between the Limpopo River, which forms the border between Zimbabwe and South Africa, and the Luvuvhu River, which separates the Makuleke Contract Park ticks and Red- 500 tick larvae or 100 engorged adult ticks per
oth oxpeckers have been observed large variety of ungulates, and in some studies it is shown that on Impala Aepyceros has assisted with reducing their grooming needs by up to 36% (Mooring & Mundy, 1996). It is well documented though that, rred by Yellow-billed
both oxpecker species have been seen on the same in particular on billed Oxpecker interactions like these have been observed in the wild (Stutterheim & Panagis, 1988).
establish which species within the study area are host(s) to one
any interspecific competition between the two oxpecker observed by Mooring &
These researchers looked species and to by the oxpeckers.
orthern Kruger een the Limpopo River, which forms the border between Zimbabwe and South Africa, and the Luvuvhu River, which separates the Makuleke Contract Park
from the Greater Kruger National Park. This contract area
mately 24 000 ha in extent and incorporates five main landscapes (Gertenbach, 1983). The Makuleke Contract Park
Pafuri management area of the Kruger National Park. The total area of Pafuri is said to cover only 1% of the Greater
surface area, yet 75% of the biodiversity can be found in this region This can be contributed to the various landscapes and geology of the area (Gertenbach, 1983; Venter, 1990).
The Makuleke Contract Park became a contractual park in 1998, when the Makuleke community had won their land claim submitted in 1994/95. It was then decided to manage the area under the commercial rights of the Makuleke community, with the environ mental management still under the South African National Parks (SANParks) (Harries, 1987; De Villiers, 1999; Child, 2004).
Within this study area there is a great variety of ungulates, some of which were reintroduced back into the area. White or Square
Rhinoceros Cerototherium simum simum was reintroduced in 2005 (Pedersen, 2009). Efforts were made to re
Wildebeest Connochaetus taurinus taurinus and Southern Giraffe Giraffa camelopardalis giraffa in the area. Neither of the two
did well, and Southern Giraffe left the area soon after were made.
The Makuleke Contract Park has a large diversity of species – it includes both oxpecker species. Yellow was first re-discovered in South Africa in 1979 in the n National Park (Hall-Martin, 1987; Whytte et al.
contract park was selected as the study site.
influence in the area made access to the site relatively easy.
ruger National Park. This contract area is approxi- tes five main landscapes (Gertenbach, 1983). The Makuleke Contract Park forms part of the management area of the Kruger National Park. The total area to cover only 1% of the Greater Kruger National Park can be found in this region.
This can be contributed to the various landscapes and geology of the
The Makuleke Contract Park became a contractual park in 1998, land claim submitted in 1994/95. It was then decided to manage the area under the commercial rights of the Makuleke community, with the environ- mental management still under the South African National Parks
Child, 2004).
a great variety of ungulates, some of White or Square-lipped was reintroduced in 2005 reintroduce both Blue and Southern Giraffe in the area. Neither of the two species fe left the area soon after these attempts
rk has a large diversity of over 420 bird Yellow-billed Oxpecker in South Africa in 1979 in the northern Kruger 1987). Therefore the contract park was selected as the study site. Wilderness Safari’s
made access to the site relatively easy.
Fig 2 - Vegetation and area map of the Pafuri area, containing the Makuleke Contractual Park (Gertenbach, 1983).Vegetation and area map of the Pafuri area, containing the Makuleke Contractual Park (Gertenbach, 1983).Vegetation and area map of the Pafuri area, containing the Makuleke Contractual Park (Gertenbach, 1983).
Methodology
The study was done over a period of 9 months - April to December 2008.
All observations were recorded on field data sheets.
10x42 binoculars were used to aid with viewing. Surveys were on walks as well as drives through the study area. Records include all instances where oxpeckers were seen on a host ungulate, as well as those birds seen perched or flying overhead.
The following data were recorded:
· Observer
· Time and date
· Oxpecker species observed
· Number of individuals seen, adult and juveniles were recorded in separate columns
· Location in the study area (GPS coordinates)
· Host species
· Any interspecific competition observed, and which species was exerting it over which.
· Any additional comments and observations.
All manually recorded data were captured in MS Excel
analysis. Guides working at Pafuri Camp assisted with da recording in the field.
Distribution and presence of both oxpecker species Makuleke Contractual Park
Both species of oxpecker occur within the study area. For more than 60 years (1915-1979) Yellow-billed Oxpecker was absent from South April to December
data sheets. Swarovski viewing. Surveys were done through the study area. Records include een on a host ungulate, as well
were recorded in
Any interspecific competition observed, and which species was
xcel® for data assisted with data
r species within the For more than was absent from South
Africa (Stutterheim & Brooke, 1981) – the area south of the Limpopo River. It was first observed again in these northern parts of the Kruger National Park, which included the Makuleke Contractual Park.
The study area was proclaimed part of the Kruger National Park in 1969 (Harries, 1987). It was assumed the oxpeckers had re
from Zimbabwe just north of the Limpopo River. The
records after re-establishment were documented in 1985 (Hall Martin, 1987; Whyte et al. 1987).
The present range of the species is further south into the Kru National Park and they had been re-introduced into the KwaZulu Natal Parks as well (Stutterheim & Panagis, 1985).
Red-billed Oxpecker were always present in these areas, as long as there were enough ungulate hosts in the region.
Red-billed Oxpecker has suffered in the past due to cattle dipping practices. Birds disappeared from certain areas and
declined in other regions due to this farming practices (Bezuidenhout
& Stutterheim, 1980). As farming practices changed
friendly the populations have recovered over the years. In some areas where they were affected most re-introduction programs were implimented (Lockwood, 1988).
Observed host preference displayed by both Oxpecker species
Yellow-billed Oxpecker
Yellow-billed Oxpeckers were more often observed on African (Cape) Buffalo Syncerus caffer (56%), followed by Greater Kudu (22%), Plains Zebra Equus quagga (16%), and
sightings on Impala. The findings corresponded
the area south of the Limpopo northern parts of the d the Makuleke Contractual Park.
of the Kruger National Park in ries, 1987). It was assumed the oxpeckers had re-entered from Zimbabwe just north of the Limpopo River. The first breeding establishment were documented in 1985 (Hall-
urther south into the Kruger introduced into the KwaZulu- (Stutterheim & Panagis, 1985).
in these areas, as long as n.
the past due to cattle dipping certain areas and numbers declined in other regions due to this farming practices (Bezuidenhout As farming practices changed to be more eco-
have recovered over the years. In some introduction programs were
reference displayed by both Oxpecker species
billed Oxpeckers were more often observed on African (56%), followed by Greater Kudu ), and two separate . The findings corresponded with that of Grobler
and Charsley (1978), and Campbell (1989). Althoug
Square-lipped Rhino was present in the area, it was only observed after the study period that Yellow-billed Oxpecker was present on them. The numbers of Rhino are low in the study area, sightings were not regular and all were made when observers were on foot.
Table 1: Results of sightings during field surveys for Yellow-billed Oxpecker Host species Number of
sightings
Number of individual birds seen Impala
Aepyceros melampus 2 4
Plains Zebra
Equus quagga 5 8
Warthog
Phacochoerus africanus 0 0
African (Cape) Buffalo
Syncerus caffer 18 115
Nyala
Tragelaphus angasi 0 0
Eland
Tragelaphus oryx 0 0
Bushbuck
Tragelaphus scriptus 0 0
Greater Kudu
Tragelaphus strepsiceros 7 26
TOTAL 32 153
From the results (Table 1 and Fig 3) it is clear that African
was by far the more frequented host species. In the case of Yellow billed Oxpeckers it is not only tick load which is important, but also host species mass, which matches with the high percentage of sightings of Yellow-billed Oxpecker on African Buffalo. These results correspond with Mooring & Mundy (1996) and Nunn et al.
rsley (1978), and Campbell (1989). Although White or in the area, it was only observed billed Oxpecker was present on low in the study area, sightings t regular and all were made when observers were on foot.
billed Oxpecker Average number of birds seen per host
2.00 1.60 0.00 6.39 0.00 0.00 0.00 3.71
it is clear that African Buffalo ase of Yellow- tick load which is important, but also host species mass, which matches with the high percentage of . These results et al. (2011).
Fig 3 – Yellow-billed Oxpecker host preference as a percentage.
Red-billed Oxpecker
The results of the fieldwork for Red-billed Oxpecker are summerised in Table 2 and Fig 4.
Red-billed Oxpeckers have a wider range of h
Yellow-billed Oxpecker - with Impala (31%) being the most host, Greater Kudu (21%) and African Buffalo (16%
Red-billed Oxpeckers were observed on the follo
(Yellow-billed Oxpeckers were not seen on these hosts) Phacochoerus africanus, Nyala Tragelaphus angasi Tragelaphus oryx, and Bushbuck Tragelaphus scriptus
preference is similar to what was recorded and seen in Hwange National Park, Zimbabwe (Hustler, 1987), and the Moremi Game Reserve, Botswana (Stutterheim & Panagis, 1985
Both Bushbuck and Warthog tried to prevent Oxpeckers from landing and feeding on them.
6%
22%
16%
56%
Oxpecker host preference as a percentage.
billed Oxpecker are summerised
a wider range of host species than being the most preferred 16%) of the sightings.
owing species as well seen on these hosts): Warthog Tragelaphus angasi, Eland Tragelaphus scriptus. This species preference is similar to what was recorded and seen in Hwange and the Moremi Game 1985a).
tried to prevent the Red-billed Impala
Kudu Zebra Buffalo
Table 2: Results of sightings during field surveys for Red-billed Oxpecker Host species Number of
sightings
Number of individual birds seen Impala
Aepyceros melampus 31 79
Plains Zebra
Equus quagga 10 39
Common Warthog
Phacochoerus africanus 3 3
African Buffalo
Syncerus caffer 16 84
Nyala
Tragelaphus angasi 13 26
Eland
Tragelaphus oryx 5 78
Bushbuck
Tragelaphus scriptus 1 1
Greater Kudu
Tragelaphus strepsiceros 21 47
TOTAL 100 357
Even though there were only 5 different sightings of
Oxpeckers on Eland, in all cases they were in large flock densities.
This resulted in the highest average number of 15.6 birds per host species.
Observations showing interspecific competition between the species of Oxpecker?
In Fig 5 the number of sightings per oxpecker species observed on the different host species is compared. Fig 6 compares the numbers of oxpeckers per oxpecker species observed on the different species. From these graphs it is apparent that Red-billed have a wider range of host species. It is clear that Yellow
billed Oxpecker Average number of birds seen per host
2.55 3.90 3.00 5.25 2.00 15.60
1.00 2.24
ent sightings of Red-billed y were in large flock densities.
birds per host
Observations showing interspecific competition between the oxpecker species observed on 6 compares the numbers observed on the different host billed Oxpeckers It is clear that Yellow-billed
Fig 4 - Red-billed Oxpecker host preference as a percentage Oxpeckers prefer the larger angulates – African Buffalo Kudu and Plains Zebra, with African Buffalo the favourite host.
Where both species were observed together on the same host at the same time, they were detected on African Buffalo
In the first instance Yellow-billed Oxpeckers occurred in hi numbers and in the second instance the number of birds species was the same on the host.(Fig 7).
In all the observations, it was never detected
oxpecker would attempt to chase away the other. Interspecific competition therefore is limited to one species (Yellow
present in higher numbers than the other when the host (African Buffalo) was shared.
31%
10% 21%
13%
5%
1%
16%
3%
preference as a percentage
African Buffalo, Greater Plains Zebra, with African Buffalo the favourite host.
Where both species were observed together on the same host at the , they were detected on African Buffalo and Greater Kudu.
billed Oxpeckers occurred in higher umbers and in the second instance the number of birds of both
er detected that one species of attempt to chase away the other. Interspecific is limited to one species (Yellow-billed) being the other when the host (African
Impala Kudu Zebra Nyala Eland Bushbuck Buffalo Warthog
Fig 5 - Comparison of number of sightings per oxpecker species observed on different host species
Fig 7 - Comparison of numbers of both 0
5 10 15 20 25 30 35
Comparison of number of sightings per Buphagus sp. for each host species. Red billed Oxpecker
Comparison of number of sightings per Buphagus sp. for each host species. Yellow billed Oxpecker
Buffalo Buffalo 19/04/2008 16/05/2008 Red Billed Oxpecker numbers 3 3 Yellow billed Oxpecker numbers 5 11
0 2 4 6 8 10 12 14 16
son of number of sightings per oxpecker species observed Fig 6 - Comparison of individual number of oxpecker species observed host species
Comparison of numbers of both oxpecker species occurring on same host species Comparison of number
of sightings per Buphagus sp. for each host species. Red billed Oxpecker
Comparison of number of sightings per Buphagus sp. for each host species. Yellow billed Oxpecker
0 20 40 60 80 100 120 140
Buffalo Buffalo Buffalo Buffalo Buffalo Kudu Buffalo 16/05/2008 04/06/2008 21/06/2008 09/07/2008 10/08/2008 13/08/2008 04/09/2008
6 3 7 2 1 8
15 11 1 13 1 15
rison of individual number of oxpecker species observed per Comparison of
individual numbers of Oxpecker spp. per host species. Red billed Oxpecker
Comparison of individual numbers of Oxpecker spp. per host species. Yellow billed Oxpecker
Buffalo Kudu 04/09/2008 27/10/2008
2 2
Feeding biology of Oxpecker species
There are no clear indications of positive dominance or aggressive behaviour displayed between the two Oxpecker species
share the same host species. What else could be governing the host preference and selection?
A number of papers on the feeding behaviour of oxpeckers pose the suggestion that oxpeckers are "vampires" and that they
target host species to feed on blood by deliberately opening wounds to obtain fresh blood. Ticks drink blood, therefore they are targeted as a "bycatch" (Weeks, 2000). This may be true of
domesticated host species as in the Weeks study (1999), and observed on captive Black Rhino (McElligott et al., 2004).
papers conclude that oxpeckers are in fact more detrimental host species than beneficial.
On the other hand there are accounts and other studies
this view. In studies on wild species parasitism was found to be of a lesser extent. Oxpecker species were more focussed on feeding on host species with a larger tick yield, African Buffalo being a very good example (Nunn et al., 2011). I did not observe any
wounds on purpose during this current study.
However other observers have mentioned blood feeding by both Yellow-billed Oxpecker (McClean, 1993) and Red-billed Oxpecker (Van Someren, 1951; Bezuidenhout & Stutterheim, 1980
birds fed on (and around) wounds and opened them allowing the blood to flow, the wounds were also cleared of fly larvae
was beneficial to the host species. In the study carried out by Plantan (2009), it was confirmed that Red-billed Oxpeckers
species in Kruger National Park exploited wounds, rather than creating wounds. This behaviour was observed on donkeys
positive dominance or aggressive Oxpecker species when they hat else could be governing the host
on the feeding behaviour of oxpeckers pose the that they actually by deliberately opening wounds they are targeted of captive or (1999), and as , 2004). These more detrimental to their
studies to contradict found to be of a species were more focussed on feeding on host species with a larger tick yield, African Buffalo being a very observe any opening of
have mentioned blood feeding by both billed Oxpecker ezuidenhout & Stutterheim, 1980). Where the and opened them allowing the of fly larvae. This in itself the study carried out by Plantan
on wild host ited wounds, rather than onkeys as well.
Red-billed Oxpeckers have been recorded to feed on four main genera of ticks: Amblyomma, Hyalomma, Boophilus and Rhipicephalus (Plantan, 2009). Some species are preferred with herbreaum, B. decoloratus, H. truncatum and
(Brown-eared Tick) being well documented as
hout & Stutterheim, 1980; Stutterheim, 1982; Stutterheim & Panagis, 1985; Stutterheim et al. 1988; Plantan, 2009). Red
are further described as having 4 feeding mechanism plucking, pecking and insect-catching (Bezuide
1980). They obtain most food requirements from the host and this may include ear wax as well as flies and lice
(Bezuidenhout & Stutterheim, 1980; Stutterheim &
1988; Plantan, 2009). This species may also feed on termite alates, as a substitute food source. I observed this behaviour on 28 December 2008, when I came across Red-billed
termite alates.
Both Oxpecker species have the same diet (Stutterheim however the intake by Yellow-billed Oxpecker was larger. Th be related to its larger body size to that Red
Yellow-billed Oxpecker weighs 60 g on average
Oxpecker approximately 50 g (Hockey et al., 2006) and thus hav greater nutritional needs.
Conclusion
During this study, no observations were made of
oxpecker species actually chasing one another away particular host. Nor were evidence of any other form of physical competition on a specific host.
Yellow-billed Oxpeckers were present in higher billed Oxpeckers when they shared their host
have been recorded to feed on four main genera of ticks: Amblyomma, Hyalomma, Boophilus and ome species are preferred with A.
and R. appendiculatus nted as favourites (Bezuiden- Stutterheim, 1982; Stutterheim & Panagis,
). Red-billed Oxpeckers 4 feeding mechanisms: scissoring, catching (Bezuidenhout & Stutterheim, equirements from the host and this lies and lice Damalinia bovis Stutterheim & Bezuidenhout, also feed on termite alates, . I observed this behaviour on 28 d Oxpeckers hawking
(Stutterheim et al., 1988) billed Oxpecker was larger. This could Red-billed Oxpeckers.
on average and Red-billed 2006) and thus having
, no observations were made of any of the two tually chasing one another away from a any other form of direct
higher numbers than Red- (Fig 7) – in particular
African Buffalo. At 45% more, it is almost double the nu average number of birds per host species sighting, Yellow Oxpecker had an average of 6.39 individual birds per sighting, compared to 5.25 individual Red-billed Oxpeckers birds p
This larger number densities where they shared the
species and then only on African Buffalo (Fig 7) may be the only form of "interspecific competition" – and this was in no instance related to physical interaction. The higher numbers can be related to larger nutritional needs as Yellow-billed Oxpeckers are sligthly larger than their cousins.
In all studies Yellow-billed Oxpeckers preferred larger ungulates larger body mass and higher tick yield (Grobler & Charsley, 1978 Stutterheim & Panagis, 1985; Hall-Martin, 1987; Hustler, 1987 Campbell, 1989; Mooring & Mundy, 1996; Nunn et al.,
billed Oxpeckers on the other hand has a wider selection of species.
- oo0oo -
Acknowledgements
Fellow guides at Wilderness Adventures Pafuri Camp helped with the recording of data in the field. Chris Roche (Wilderness Safaris) commented on the draft of this paper and supported with advice during the study. Trevor Hardaker gave permission to use photos to illustrate the different species.
almost double the number. In the sighting, Yellow-billed idual birds per sighting, birds per sighting.
the same host may be the only form and this was in no instance related to The higher numbers can be related to larger billed Oxpeckers are sligthly larger than
preferred larger ungulates –
& Charsley, 1978;
Martin, 1987; Hustler, 1987;
, 2011). Red- selection of host
helped with the s Roche (Wilderness Safaris) e draft of this paper and supported with advice the study. Trevor Hardaker gave permission to use photos to
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