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Transcript
69EG3291 Third Year Project
An Investigation into the Effects of Tourist Related
Disturbances on Parrot Abundance and Behaviour at a
Peruvian Geophagy Site
S. Lovesey
A project submitted in partial fulfilment of the requirements for the degree of
Bachelor of Science (Honours) in Physical Geography, The Manchester
Metropolitan University.
Department of Environmental and Geographical Sciences
The Manchester Metropolitan University
April 2007
1
Declaration of originality
This is to certify that the work is entirely my own and not of any other person,
unless explicitly acknowledged (including citation of published and unpublished
sources). The work has not previously been submitted in any form to the
Manchester Metropolitan University or to any other institution for assessment or
for any other purpose.
Signed
------------------------------------------
Date
------------------------------------------------
2
Contents
Page
Contents
ii
Contents
iii
List of figures
iv
List of tables
v
Abstract
vi
1.0
INTRODUCTION
1
1.1
Diversity in the rainforest
1
1.2
Threats to tropical wildlife
1
1.3
Parrots
2
1.3a.
Parrot diversity
2
1.3b.
Parrot ecology
2
1.3c.
Threats to parrots
3
1.3d.
Conservation of parrots
4
1.3e.
Parrots of Peru
5
Geophagy
5
1.4a.
What is geophagy?
5
1.4b.
Geopagy in parrots
5
1.4c.
The absorption of dietary toxins and gastrointestinal protection
6
1.5
Parrot abundance and geophagy in southeastern Peru
7
1.6
Ecotourism and parrots in Peru
8
1.7
Effects of tourist visitation on parrot geophagy sites
9
2.0
AIMS
9
3.0
STUDY AREA
10
3.1
Peru
10
1.4
3
3.2
Study site
11
3.3
The La Torre colpa
12
3.4
Tourist visitation
13
4.0
METHODOLOGY
13
4.1
Parrot observations
13
4.2
General disturbances
14
4.3
Tourist disturbances
14
4.4
Boat disturbances
15
4.5
Animal disturbances
15
4.6
Unknown disturbances
15
4.7
Statistical analysis
15
4.7a.
Associations
15
4.7b.
Variances between disturbance factors
16
5.0
RESULTS
16
5.1
Numbers recorded
16
5.2
Species associations
18
5.3
Parrot abundance and days of disturbance
19
5.4
Species associations with different disturbance factors
20
5.5
General disturbance factors
21
5.6
Tourist disturbance factors
22
5.7
Boat disturbance factors
22
6.0
DISCUSSION
23
6.1
Abundance
23
6.1a.
Most abundant species
23
6.1b.
Least abundant species
24
6.1c.
General abundance of species
24
6.2
Species associations
25
6.3
Species interdependence
25
6.4
General disturbance and parrot abundance
25
6.4a.
Associations with general disturbance factors
26
6.4b.
Differences observed between general disturbance factors
26
4
6.5
Tourist disturbance factors
27
6.5a.
Cough/sneeze and dropped objects
27
6.5b.
Loud talking
27
6.5c.
Quiet talking
28
6.5d.
Arrival and departure
28
Boat disturbance
29
6.6a.
Loud boats not stopping
29
6.6b.
Quiet boats not stopping
29
6.6c.
Tourist boats stopping
30
6.7
Limitations of the study
30
6.8
Management implications
31
6.8a.
Abundance of species
31
6.8b
Tourist disturbances
31
6.8c
Boat disturbances
32
7.0
CONCLUSION
33
8.0
Acknowledgements
34
9.0
References
34
10.0
Appendices
45
Appendix 1 - Example of data sheet used to record parrot abundances
and flushes
Appendix 2 - General information about the parrot species
recorded at the La Torre colpa
45
6.6
5
46
List of figures
Page
Figure 1
Map of Peru showing location of study site
10
Figure 2
Satellite image of the Tambopata, La Torre colpa, Inotawa Lodge and
Posada Amazonas.
11
Figure 3
View of the La Torre colpa from the Inatowa Blind.
12
Figure 4
Mean disturbance levels compared to mean parrot visitation numbers
per day.
20
6
List of tables
Page
Table 1
Total number of individual birds observed feeding by species/day.
17
Table 2
Mean number of individual birds observed feeding per day.
18
Table 3
Species associations using Spearman’s rank correlation.
19
Table 4
Species/disturbances associations calculated using Spearman’s rank
correlation.
21
Table 5
Mean and standard deviation values comparing general disturbance
factors with tree and colpa flushes.
21
Table 6
Mean and standard deviation values comparing tourist disturbance
factors with tree and colpa flushes.
22
Table 7
Mean and standard deviation values comparing boat disturbance with
tree and colpa flushes.
23
:
7
Abstract
Geophagy, the intentional consumption of soil or clay, plays a vital role in maintaining parrot
health. This research investigates the effects that tourist related disturbances and boat traffic are
having on parrots at a geophagy site, situated on the river Tambopata, southeastern Peru. Five
minute counts were used to establish parrot abundance with flushes being recorded as signs of
disturbance. Impulse noises such as the onset of loud talking, coughing/sneezing and dropped
objects resulted in the greatest tourist disturbance to parrots overall. Boats using peke-peke motors
attributed the most disturbances by boats to parrots at the site. Simple guidelines on talking levels
inside the blind should be put into place. An alternative floor covering should also be used to
reduce the impact of dropped objects. Speed limits and passing distances for boats could also be
used to reduce disturbance further. Overall the study was successful but further research needs to be
undertaken on the actual quantities of clay needed for parrots to remain healthy. Studies similar to
this at other parrot geophagy sites would also greatly contribute to the limited knowledge that has
already being gained.
Word count: 9,627
8
1. INTRODUCTION
1.1 Diversity in the rainforest
Tropical rainforests are the most diverse habitat on the planet (Bierregaard et al 1992).
Even though they only cover 7% of the planet’s landmass, they are home to half to two
thirds of the plant and animal species on Earth (Wilson 1988, Raven 1988). Due to this
great diversity of life they have become known as biodiversity hotspots, which are vital
when considering the conservation of many tropical species (Mittermeir et al 1998).
Pressure on the world’s rainforests is ever increasing due to expanding human
populations who require land to live on and provide income (Laurance 2001).
1.2 Threats to tropical wildlife
Globally tropical rainforests are coming under continuing threat from human activities
(Turner 1996). Deforestation is the main threat faced by tropical rainforests and
wildlife (Geist et al 2002, Putz et al 2000). Logging for the international timber trade,
forest conversion for crop cultivation, clear felling for grazing arable land and
traditional practises such as agroforestry and swidden farming techniques, all combine
to fragment and disturb a rainforest’s natural dynamics and habitats (Sivakumar 2000,
Blockhus et al 1992, Bowles et al 1998). It is estimated that over the period 19952000, 1.9 million ha per year of Amazonian rainforest were lost to these activities
(Laurance et al 2001).
Habitat loss and conversion are the direct effects of logging on tropical rainforest
wildlife, however there are secondary factors to consider. Associated with logging and
deforestation is an increase in hunting pressure for food and the bush-meat trade
(Sandercock et al 2000). This is due to once inaccessible areas of forest being opened
up through road building in order to access and remove timber or crops (Bennet et al
2000, Uhl et al 1989). People will expand along new infrastructure clearing patches of
forest for personal home-gardens and subsistence agriculture (Kellman et al 1997,
Johns et al 1996). Many will rely on bush meat and trapping to provide extra income,
increasing the pressure on wildlife resources in that area (Fa et al 2002, Carpaneto et
al 2004).
9
1.3 Parrots
1.3a Parrot diversity
Parrots (of the order Psittaciformes) are one of the largest and uniformly distinctive
groups of birds in the world (Juniper & Parr 1998). There are around 353 different
species that can be divided into two families (Snyder et al 1996): Cacatuidae
(cockatoo) and Psittacidae (true- parrots). Members of the parrot family include
macaws, parakeets, and parrotlets (Snyder et al 2000). They can be be found in most
warm regions of the world, including India, southeast Asia, the Neotopics and west
Africa (Juniper et al 1992). Parrots become increasingly diverse in tropical and
subtropical lowland forested areas, with the most speciation occuring in the New
World and Australia (Karr 1976).
1.3b. Parrot ecology
Most parrots dwell in forest habitats and are threfore largly or exclusivly arboreal,
however there are exceptions, such as the kakapo Strigops habroptilus
of New
Zealand (Clout et al 1995), and the ground parrot Pezoporus wallicus Kerr of
Australia (Meredith et al 1984). Most parrot diets are comprised of plant parts in the
form of seeds, fruits, blossoms, nectar, pollen, buds, leaves, berries, nuts and
sometimes bark (Juniper & Parr 1998, Galletti 1993). Many are generalist feeders with
a wide dietary flexiblity that allows them to spread over large and ecologically diverse
ranges, whilst others are specialists associated with a small habitat range and a less
varied diet, such as the endangered Lear’s macaw Anodorhynchus leari (Yamashita
1987). The majoritory of species are gregarious for at least part of the year and are
usually encountered in small flocks or pairs, attributed to foraging effectivness and
anti-predator defence (Monterrubio-Rico et al 2006, Gilardi et al 1988, Burger et al
2003). Social roosting is common in parrots (Chapman et al 1989). Some species like
the African grey Psittacus erithacus, spend the night in tree tops (Juniper & Parr
1998), other species in small groups in tree hollows (Pyrrhura parakeets, Best et al
1996), some on cliffs (Brightsmith 2005), or others in communial nests (monk
parakeet Myiopsitta monochus, Sol et al 1997). Parrots are mainly monogamous and
in the case of many larger species will pair for life (Loffredo et al 1986). The vast
10
majoritory of species are cavity-breeders, with nests located in tree hollows (Bessinger
et al 1992, Brightsmith 2005). The availability of suitable nest-sites is a limiting factor
in the breeding density of many parrot species due to very few activly constructing
nests (Renton et al 1999).
1.3c Threats to parrots
The World Conservation Union (IUCN), has identified two main threats to parrot
species, trapping for the live bird trade, and habitat loss and fragmentation (Snyder et
al 2000). Due to their attractiveness and intelligence parrots have always been highly
desired as pets (Cooney et al 2005). This has led to vast quantities of new world
parrots being exported from Third World to First World countries in order supply
private buyers and aviculturlists (Guix et al 2004). More than 1.8 million parrots
legally entered the international trade from 1982-1988, most imported into the United
States (80%), European Union countries (15%) and Japan (3%) (Bessinger et al 1992).
Estimates of mortality rates and illegal smuggling indicate that the actual number of
birds taken from the wild may be two to three times greater than this figure (IñigoElias et al 1991).
As the international trade in parrots depletes numbers in the wild they also face the
effects of habitat destruction and conversion that threatens all wildlife in the tropics
(Geist et al 2002, Putz et al 2000, Turner 1996). Research into bird extinctions in
tropical rainforests suggests that a 1000 ha area of fragmented rainforest will support
only 50 percent of the original bird species recorded before fragmentation occurred
(Brooks et al 1999). Every parrot species has its own reaction to forest disturbance
according to habitat selection, foraging behaviour, dietary adaptability and sensitivity
to microclimatic conditions (Thiollay 1997). Habitat loss and conversion combined
with legal and illegal trade has left at least 30% of the 140 parrot species found in the
Western Hemisphere now being threatened with extinction (Collar et al 1992).
Research suggests that 40% of these species are threatened primarilly by habitat
destruction, 17% by trade, 36% by a combination of the two causes and 7% by other
factors (Collar et al 1992), making neotropical Psittacidae one of the most threatened
groups of birds in the world (Bennet et al 1997). This illustrates the need for
continuing research into the many effects that human acivities are having on the
11
world’s parrot species. This can then be used in creating effective management plans
to aid in the global conservation in parrots.
1.3d. Conservation of parrots
Conservation efforts to ensure the protection of parrots and other species have been
undertaken since the 1960s (Myers et al 2000). The Convention on International Trade
in Endangered Species of Wild Fauna and Flora (CITES), is an international
agreement between governments (CITES 2007). Its aim is to ensure that international
trade in specimens of wild animals and plants does not threaten their survival. Along
with the establishment of conventions like CITES, organisations such as the IUCN and
BirdLife International campaign and fund for the preservation of parrot species
(Jenkins 1996, BirdLife International 2006).
Along with environmental conventions and the activities of other organisations, more
general practises are evolving that aid in parrot conservation, due to the growing
international concern over the destruction of the world’s tropical rainforests (Smyth et
al 2004). Measures are being taken to reduce the levels of deforestation that are
currently occurring, and the habitat loss that is associated with it (Bawa et al 1998,
Lambert 1992). Practises such as selective logging and supervised logging regimes are
increasingly being used to reduce damage to forest habitats that is usually associated
with traditional logging regimes (Lewis 2001, Whitman et al 1998, Asner 2004). This
involves directional felling to reduce damage done to the remaining stand, liana
cutting to avoid destroying the forest canopy, more care in planning roads and skid
trails, leaving refuge stands to initiate forest regeneration and not logging on steep
slopes to prevent soil erosion (Packer 1967, Burke 1973, Holmesay et al 2002). This
reduces the habitat loss and disturbance that occurs through traditional logging
practises, and promotes faster forest regeneration times (Torquebiau 1992). This
benefits the whole wildlife community that depend on tropical rainforest habitats for
survival (Hamer et al 2003, Dah 2004). This however, will only go part of the way to
aiding in parrot conservation, as it is far to general. Many species will need
independent research into specific management plans to help individual species
depending on habitat preference, ecology and abundance.
12
1.3e. Parrots of Peru
Peru has an incredible diversity of bird species, approximatly 1878 species inhabit
Peru with 139 of those being endemics (Clements 2001). 52 parrot species have been
recorded with one being endemic to the country (yellow-faced parrotlet Forpus
xanthops). There are 10 globally threatened species that have been identified on the
IUCN’s Red List of Threatened Species (IUCN 2007). These are military macaw Ara
militaris (vulnerable), blue-headed macaw Primolius couloni (endangered), redmasked parakeet Aratinga erythrogenys (near threatened), golden-plumed parakeet
Leptosittaca branickii (vunerable), white-necked parakeet Pyrrhura albipectus
(vulnerable), yellow-faced parrotlet Forpus xanthops (vulnerable), gray-cheeked
parakeet Brotogeris pyrrhopterus (endangered), amazonian parrotlet Nannopsittaca
dachilleae (near threatened), spot-winged parrotlet Touit stictopera (vulnerable) and
red-faced parrot Hapalopsittaca pyrrhops (vulnerable).
1.4 Geophagy
1.4a. What is geophagy?
Geophagy is the intentional consumption of soil or earth (Brightsmith et al 2004). This
has been recorded primarily in the tropics amongst many different species. These
include mammals (Jones et al 1985, Klause et al 1998), birds (Symes et al 2006,
Montenegro 2004), reptiles (Brown 1981) and invertebrates (Wolters 2004). The
principle theories as to why this behaviour occurs include mechanical enhancement of
digestion, mineral supplementation, acid buffering, the absorption of dietary toxins and
gastrointestinal cytoprotection (Gilardi et al 1999, De Souza et al 2002). However,
different animals consume clay or soil for different reasons so one hypothesis cannot be
true for all species.
1.4b. Geophagy in parrots
Geophagy amongst birds has been described from observations of many species, but is
particularly well known to occur in the parrot family. Reports of parrot geophagy in the
13
Neotropics have come from Mexico, Bolivia, Brazil and Peru; however parrot geophagy
is not limited to this region. Observations of soil consumption have been recorded in the
palm cockatoo Probosciger aterrimus, of Papua New Guinea (Symes et al 2005), and
the grey lourie Corythaixoides concolor, in Botswana (Pyrce 2004). As this behaviour is
often overlooked, it is expected that new observations in varying species will be made
that support the theory that avian geophagy is widespread, and has evolved several times
independently (Gilardi et al 1999).
1.4c. The absorption of dietary toxins and gastrointestinal protection
The most extensive data on parrot geophagy comes from Peruvian clay-lick sites
(Brightsmith et al 2004, Gilardi et al 1999, Hammer 2002). Studies indicate that there
are two main reasons for geophagy in parrot species, the absorption of dietary toxins and
gastrointestinal protection (Brightsmith et al 2004, Gilardi et al 1999). Soil samples
collected and analysed from several Peruvian clay-lick sites demonstrate that soils
consumed have the ability to absorb large quantities of alkaloid quinine (Brightsmith et
al 2004). This toxin occurs naturally at low levels in most parrot diets. Soil analysis
showed that the clays could absorb ~100 mg of alkaloid quinine per gram of clay
consumed. This indicates that the consumption of several grams of clay per day could
absorb biologically significant quantities of this toxin, enabling an increase in dietary
capability.
The consumption of clay to absorb dietary toxins can also be linked to gastrointestinal
protection (Gilardi et al 1999). The presence of clay in the gut increases mucus secretion
and enhances the mucus barriers ability to protect the gut lining from chemical attack
(Diamond et al 1999). Research into the passage rate of clays in captive parrots
indicates that large amounts of clay were still present in the gastrointestinal tract for at
least 12 hours after consumption (Gilardi et al 1999). Most parrot species that exhibit
geophagy behaviour consume clay daily, with most consumption occurring in the early
morning. This can be used as evidence for clay being used for the absorption of dietary
toxins and gastrointestinal protection throughout the day as birds feed (Brightsmith et al
2004). The absorption of dietary toxins and gastrointestinal protection benefits clayconsuming parrot species by enabling birds to consume previously unexploitable
resources and/or increased quantities of seeds and fruits that would otherwise cause
14
illness or death (Gilardi et al 1999). It also allows birds to consume nutritionally rich but
highly toxic resources during the dry season when food is a limiting factor to other
frugivores (Terborgh 1986). Therefore, in the case of parrots, geophagy extends their
dietary capacity and may increase distributions and abundances of certain of species.
This highlights how important geophagy is too many parrot species, which have evolved
this behaviour to increase their health and survival rates. Due to this it is vital to
understand any effects that human disturbances at parrot geophagy sites are having.
1.5 Parrot abundance and geophagy in southeastern Peru
Roughly twenty members of the parrot family inhabit the dense lowland tropical
rainforest of southeastern Peru (Rainforest Expeditions 2001). These are (largest to
smallest), blue and yellow macaw Ara ararauna, red and green macaw Ara
chloroptera, scarlet macaw Ara macao, blue-headed macaw Ara couloni, red-bellied
macaw Ara manilata, chestnut-fronted macaw Ara severa, mealy parrot Amazona
farinosa, yellow-crowned parrot Amazona ochrocephala, blue-headed parrot Pionus
menstruus, white-bellied parrot Pionites leucogaster, orange-cheeked parrot
Pionopsitta barrabandi, white-eyed parakeet Aratinga leucophthalmus, dusky-headed
parakeet Aratinga weddellii, black-headed parakeet Pyrrhura rupicola, painted
parakeet Pyrrhura picta, cobalt-winged parakeet Brotogeris cyanoptera, scarletshoulded parrotlet Touit huetti, tui parakeet Brotogeris sanctithomae, Amazonian
parrotlet Nannopsittaca dachilleae and dusky billed parrotlet Forpus sclateri
(Rainforest Expeditions 2001). Only two species are on the IUCN Red List of globally
threatened species, the blue-headed macaw Ara couloni (endangered) and Amazonian
parrotlet Nannopsittaca dachilleae (near-threatened) (IUCN 2007). The reason for this
diversity in parrot species is due to the varied habitats and food sources that are to be
found in lowland tropical rainforests (Terborgh et al 1990). This allows for vast
amounts of evolutionary diversity to occur over other habitats that have fewer and less
varied food sources available.
As seen in afore mentioned literature on parrot geophagy, it is vital for some parrots to
consume soil or clay as a regular part of their diet (Brightsmith et al 2004, Gilardi et al
1999). This is true for many species that inhabit southeastern Peru. As such there are
15
many macaw, parrot and parakeet clay-lick sites, locally called colpas (referred to as
in remaining literature), to be found in the area. These are usually located on exposed
sections of river bank kept free from vegetation by erosion from the river, but can also
be found inland in some cases. Locally the colpas are used on a daily basis by feeding
birds and this regularity has led to extensive research been undertaken at certain sites
(Tambopata Research Centre, the largest macaw colpa in the world, studied for over
15 years). It has also led to the development of ecotourism in the area with many
lodges been built to cater for tourists who want to observe rainforest flora and fauna.
Excursions to observe the many species of parrot that utilize colpas in the region are
an integral part of this. Due to these factors it is an ideal location to perform a study on
the extent of human disturbances at a parrot geophagy site.
1.6 Ecotourism and parrots in Peru
Ecotourism is hard to define, but can losely be described as “nature-based tourism,
which is protective of nature as well as enjoying it” (Valentine 1992). The use of the
term “ecotourism” can only be traced back as far as the late 1980s as a reaction to the
negative impacts of mass tourism to natural areas (Richardson 1993). An ecotourist
should be seen to practise a non-comsumptive use of wildlife and natural resources, and
contribute to the visited area through labour or financial means, aimed at directly
benifiting the conservation of the site (Ziffer 1989).
Since the 1970s ecotourism has been expanding in Peru. The country has varying
habitat types from high-altitude mountain ranges and altiplano, to dense, lowland
tropical rainforest. This combined with a deep anthropological history ensures that Peru
is a magnet for many travellers. Ecotourism in the Peruvian Amazon has been used as a
way to enhance the value of intact wildlands, promote conservation and stabalise landuse patterns (Yu et al 1997). Many ecotourist lodges in the Peruvian Amazon rely on
parrots and their associated geophagy behaviour as a selling point to visitors. This is due
to the unique abundance and variety of parrot species that will reliably consume clay
from known geophagy sites on a daily basis. As such many colpas in southeastern Peru
will have associations with one or several lodges, who take tourists to observe parrots
feeding. It is the disturbances caused to feeding birds during these visits that this study
will be focusing on.
16
1.7 Effects of tourist visitation on parrot geophagy sites
There is very little known about the effects of tourist visitation on parrot colpa feeding
behaviour. The quantity of clay and the time needed on geophagy sites for parrots to
remain healthy is unknown. Due to this factor it is vital to try to minimise any
disturbances that are attributed to tourist visitation. Studies on parrot geophagy sites
have indicated that increased disturbances will result in a decrease in parrot abundance
(Tatum-Hume et al 2004, Hammer 2002). It is not known how much this decrease
affects parrot health, due to the possible effects of deficencies in clay usually sort from
colpas. Different factors such as tourist arrival time and behaviour will all cause
different levels of disturbance to any parrots present at the site. At this moment there are
no studies quantifing tourist disturbances or highlighting the factors responsible for the
most disturbance. Due to the importance of geophagy for many parrot species to survive
and remain healthy, it is vital to understand any disturbances caused to them and try to
reduce these factors to a minimum.
2.0 AIMS
The aim of this investigation is to evaluate tourism related disturbances on a parrot
geophagy site in southeastern Peru. Boat, tourist and natural disturbances will all be
analysed in order to identify the factors that attribute most disturbances to parrots on
and around the colpa. The abundance and number of species will be recorded to assess
which use the colpa and whether any are more susceptible to the certain disturbances.
Management recommendations will then be made that will help to remove or at least
reduce avoidable human disturbance factors. This will reduce the impacts that current
visitation is having and benefit the whole parrot community that utilize the study site.
17
3.0 STUDY AREA
3.1 Peru
Peru, officially the Republic of Peru, is the world’s 20th largest country, with a
landmass of 1,285,220 km2 (BBC 2007). It borders Ecuador and Columbia to the
north, Brazil and Bolivia to the east and Chile to the south (Figure 1). To the west lies
the Pacific Ocean. It has a population of over 28 million people (United Nations
2004), who speak Spanish, with others bilingual in Quechua, Aymara or other native
languages. Eastern Peru consists mostly of the moist tropical rainforest of the Amazon
Basin whilst western areas are dominated by the Andes mountain range and other high
altitude habitats.
Figure 1: Map of Peru showing location of study site (BBC 2007).
3.2 Study site
The study was carried out at the La Torre colpa (S 12° 49’38, 09’, W 69° 17’23, 49’)
on the river Tambopata, southeastern Peru (Figure 2). The site is roughly 2 hours by
boat upstream of the town of Puerto Maldonado (Figure 1), which is located on the
confluence of the Amazonian rivers of Madre de Dios and Tambopata (S 12° 36’12,
55’, W 69° 11’31, 79’). It is in the tropical zone 205 metres above sea level, in the
Department of Madre de Dios, near to the borders of Brazil and Bolivia. The colpa is
situated on the edge of the Tambopata-Candamo Reserve Zone (TCRZ) that originated
in the 1970s. Initially it comprised about 5,000 hectares but this area was enlarged to
1.5 million hectares in 1990. The TCRZ is spread over two Departments, that of
Madre de Dios (Province of Tambopata), and Puno (Provinces of Carabaya y Sandia),
and covers approximately 1.5 million hectares. Habitats range from sub-tropical moist
18
forest, to cloud forest and tropical savannah (Hammer 2002). Rainfall averages 2,000
mm per year and humidity is roughly 75%.
Figure 2: Satellite image of the Tambopata , La Torre colpa, Inotawa Lodge and Posada Amazonas
(GoogleEarth 2007).
3.3 The La Torre colpa
The La Torre colpa is a small, exposed clay cliff set back from the river Tambopata’s
eastward bank by 20-25 metres (Figure 2). At this point the river is roughly 50 m
wide. The cliff is approximately 5-8 m high and 15m wide (Figure 3). A broad sandy
beach made up of fluvial deposits runs for a width of 6-10 m for approximately 30 m
along the river’s edge. This is predominantly exposed but after heavy rainfall becomes
flooded. The beach is backed by dense secondary rainforest consisting mainly of a
variety of palms (Palmaceae), Cercropia and Balsa (Ochroma). The colpa is only
visable from about 25 m on either side. However it is exposed and visable from the
river and opposite dank due to a depression that is directly in front of the cliff face
(approximatly 30 m2). This is mainly vegetated by understory species such as
heliconia (Heliconiacae) and legume (Leguminosae). The colpa is used by two lodges
in the area and as such two blinds have been constructed to allow for tourist visitation.
These are situated 20 m to the left of the colpa, both having capacity for 8-12 people.
Figure 3: Veiw of the La Torre colpa from the Inotawa Blind (Authors photograph).
3.4 Tourist visitation
19
There are two lodges in the area that use the La Torre colpa. Inotawa Lodge is situated
roughly 500 m downstream of the colpa (Figure 2), on the westward side of the river
Tambopata (S 12° 48’36,87’, W 69° 18’11,32’). The lodge has capacity for 30 people
and provides ecotourists with the opportunity to observe different varieties of
rainforest flora and fauna. This includes guided walks through the forest, observing
different forms of native cultervation, and visiting the colpa to watch macaws, parrots
and parakeets. In peak tourist season the blind is used nearly everyday with variations
in tourist numbers depending on group size. In the off season the blind is used
approxinatly every 2-3 days depending on numbers of tourists staying at Inotawa.
Posada Amazonas (Figure 2), is much larger than Inotawa Lodge with 30 rooms and a
capacity of 100 people (S 12° 48’08,22’, W 69° 17’59,37’). It is situated approximatly
20 minutes walk from the colpa on the eastward side of the river Tambopata. It
Opened in 1998 and is run by Rainforest Expeditions (Est. 1989). It is part of a
community partnership ecotourism project that is jointly owned by the local Ese-Eja
community of Infierno, and is situated inside the communities private reserve on the
eastward side of the river (Rainforest Expeditions 2006). It offers similar trips to
Inotawa and also has a blind at the La Torre colpa. The Posada blind is situated next to
the Inotawa’s 20 m left of the colpa. It is used most days during the peak season, with
visitation numbers to the colpa varying according to how many people are staying at
the lodge.
4.0 METHODOLOGY
4.1 Parrot observations
The study was conducted in early July 2006 over a 28-day period, with a total of 24
observation days. Recordings of parrot species and abundance were made from the
Inotawa blind using a data sheet designed for the Tambopata Research Project
(Appendix 1). The species and number of birds feeding were recorded using fiveminute counts (Brightsmith 2004). The moment the first bird of the day landed on the
colpa a one-minute count of the species feeding and number of individuals involved
20
was started. A gap of 4 minutes was then left until another one-minute count was
started. This was repeated until all birds had left the colpa and surrounding trees. This
method allowed for identification of variation in abundance between days of high and
low disturbance. Observations were made using binoculars and Tambopata area parrot
field guide to ensure accurate species and number recognition (Rainforest
Expeditions). Arrival at the study site was by canoe, approximately 6 AM (EST) just
before sunrise. This was to ensure arrival before any tourists and to cause as little
disturbance to any birds that may have already been present. Departure time from the
site was roughly 9-10 AM (EST), half an hour after the last bird to leave the study
area. This was to ensure that feeding activities at the colpa had totally finished.
4.2 General disturbances
Disturbances to parrots were divided into four broad categories: tourist disturbances,
boat disturbances, animal disturbances and unknown disturbances. Parrot flushes were
used as an indicator of disturbance. A flush is when a congregation of birds suddenly
takes flight from a settled position (Bessinger & Casagrande 1997), in this case on the
colpa or surrounding trees. These were recorded in order to assess the effects of
disturbances on feeding and non-feeding birds.
4.3 Tourist disturbances
Tourist disturbances were divided into arrival/departure, quiet talking, loud talking,
dropped object and cough/sneeze. Every time one of these factors occurred any
associated flushes from the colpa or surrounding trees were recorded. Any species and
the number of individuals involved in the flush were also recorded. This was to ensure
accurate identification of the tourist factor that caused the most disturbances, and
whether any species is affected more than another.
4.4 Boat disturbances
21
Boat disturbance was divided into three different factors; loud boat not stopping, quiet
boat not stopping and tourist boat stopping. Loud boats were classed as those that used
traditional peke-peke motors (8-16 BHP), quiet boats were ones that used more
modern outboard motors, and tourist boats were recorded as any boat that was used at
the pick-up point on the beach, for tourist arrival and departure. Any boat disturbance
factor that caused a flush to birds on the colpa or in the surrounding trees was
recorded. The species and number of individuals involved were also recorded. This
was used when establishing whether one boat disturbance factor was affecting the
birds more than another, and whether any species were particularly affected.
4.5 Animal disturbances
Animal disturbance was recorded as any flushes caused to birds on the colpa or in the
surrounding trees by wildlife that occurs naturally in the surrounding area, such as
monkeys, birds of prey and snakes. When a flush was caused in this way any species
and the numbers of individuals involved were recorded. This was used to establish the
extent of natural animal disturbance caused to the parrots that use this site.
4.6 Unknown disturbances
Unknown disturbances were classed as flushes by birds on the colpa or in the
surrounding trees without any obvious association with any of the disturbance factors
already listed in this study. When this was observed, any species involved and the
numbers of individuals were recorded to establish the amount of apparently unknown
disturbances.
4.7 Statistical analysis
4.7a. Associations
Associations of species and species associations with disturbances were calculated
using Spearman’s rank correlation coefficient (Wheater & Cook 2000). This is a nonparametric correlation analysis for examining relationships between two variables.
Each variable is ranked separately and comparison then takes place (Wheater et al
22
2000). A P value of less than 0.01 shows a highly significant difference between
variables, less than 0.05 is a significant difference and a P value greater than 0.05
shows no significant difference between variables.
4.7b. Variances between disturbance factors
Variances between the different disturbance factors and amount of disturbance caused
to birds on the colpa and surrounding trees was calculated using Kruskal-Wallis oneway ANOVA analysis. This is a non-parametric statistical test to examine differences
between more than two samples comprising unmatched, independent data (Wheater et
al 2000). Any significant or non-significant differences between data sets will be
highlighted using P value results.
5.0 RESULTS
5.1 Numbers recorded
Parrot numbers feeding on the colpa were recorded over a 28-day period with a total of
24 observation days. Overall 8 species of parrot were recorded feeding on the colpa,
these were red and green macaw Ara chloroptera, chestnut-fronted macaw Ara severa,
mealy parrot Amazona farinosa, yellow-crowned parrot Amazona ochrocephala, blueheaded parrot Pionus menstruus, orange-cheeked parrot Pionopsitta barrabandi, duskyheaded parakeet Aratinga weddellii and white-eyed Parakeet Aratinga leucophthalmus
(for general information see Appendix 2). In total, 775 individuals were recorded
feeding on the colpa (Table 1). Dusky-headed parakeet Aratinga weddellii were the
most commonly observed with 337 individuals recorded over the study period.
Combined with 209 blue-headed parrot Pionus menstruus individuals they make up over
50% of total parrots recorded feeding. Red and green macaw was the least common with
only 2 individuals being observed on day 27 (Table 1).
Dusky-headed parakeets Aratinga weddellii had a mean value of 1.48 individuals
recorded feeding per day (Table 2). This makes them the most commonly observed
species feeding on the colpa. However, they show a high standard deviation of 1.33
23
suggesting high variability between visitation numbers per day. Blue-headed parrots
Pionus menstruus record the second highest value with a mean of 0.74 (Table 2). Red
and green macaw Ara chloroptera was the species with the lowest mean value recorded
of 0.004 individuals per day, making it the least abundant species (Table 2). A standard
deviation of 0.18 shows little variance in number of individuals observed between days.
Table 1: Total number of individual birds observed feeding by species/day.
Red and
Green
Macaw
Chestnut
Fronted
Macaw
Mealy
Parrot
Yellow
Crowned
Parrot
Blue
Headed
Parrot
Orange
Cheeked
Parrot
Dusky
Headed
Parakeet
White
Eyed
Parakeet
1
0
0
0
0
0
2
0
0
0
0
7
0
0
0
0
0
21
0
28
3
0
0
4
2
13
0
14
5
38
4
0
2
1
5
0
2
6
2
2
0
0
0
7
1
13
3
19
7
51
6
0
3
3
3
21
2
23
3
58
7
0
8
0
0
0
0
2
0
14
0
16
0
0
0
7
0
16
0
9
23
0
0
0
1
4
0
6
0
11
10
0
0
0
0
12
0
4
16
32
11
0
0
0
0
2
0
11
0
13
12
0
0
1
2
9
0
7
0
19
13
0
2
0
1
10
0
5
2
20
14
0
0
13
6
10
0
27
6
62
15
0
18
5
0
17
2
32
0
74
16
0
0
0
0
0
0
2
0
2
17
0
1
0
0
0
0
0
0
1
18
0
4
5
7
12
0
14
0
42
23
0
25
0
2
22
0
0
0
49
24
0
13
2
7
8
0
29
0
59
25
0
11
2
3
21
0
13
1
51
26
0
0
0
2
0
0
11
0
13
27
2
8
1
5
8
1
35
0
60
28
0
1
0
2
9
0
34
0
46
Total:
2
90
43
46
209
8
337
40
775
Day
No.
Table 2: Mean number of individual birds observed feeding per day (± Standard Deviation).
24
Total:
Species:
Mean:
Red and
Green
Macaw
Chestnut
Fronted
Macaw
Mealy
Parrot
Yellow
Crowned
Parrot
Blue
Headed
Parrot
Orange
Cheeked
Parrot
Dusky
Headed
Parakeet
White
Eyed
Parakeet
0.0038
± 0.18
0.3730
± 0.68
0.1447
± 0.26
0.1441
± 0.19
0.7374
± 0.74
0.0345
± 0.08
1.4799
± 1.33
0.025
± 0.17
5.2 Species associations
Species associations were calculated using Spearman’s rank correlation analysis.
Mealy parrot Amazona farinosa were the species most associated with other species
feeding on the colpa (Table 3). They showed a correlation at 0.01 levels with duskyheaded parakeet Aratinga weddellii, blue-headed parrot Pionus menstruus and yellowcrowned parrot Amazona ochrocephala. At the 0.05 level correlations were seen with
chestnut-fronted macaws Ara severa, orange-cheeked parrot Pionopsitta barrabandi
and white-eyed parakeet Aratinga leucophthalmus (Table 3). Chestnut-fronted
macaws were also strongly associated with other species being present. Associations
were seen with blue-headed parrots Pionus menstruus and yellow-crowned parrots
Amazona ochrocephala at the 0.01 level and at the 0.05 level correlations were seen
with mealy-parrots Amazona farinosa and orange-cheeked parrots Pionopsitta
barrabandi. White-eyed parakeets Aratinga leucophthalmus showed little species
associations; only at 0.01 levels with mealy parrot Amazona farinosa and blue-headed
parrot Pionus menstruus. Red and green macaw Ara chloroptera was the only species
to have no associations (Table 3).
Table 3: Species associations using Spearman’s rank correlation
25
(** = strongly associated * = associated).
Red and
Green
Macaw
Chestnut
Fronted
Macaw
Mealy
Parrot
Yellow
Crowned
Parrot
Blue
Headed
Parrot
Orange
Cheeked
Parrot
Dusky
Headed
Parakeet
-0.13
p = 0.54
-0.01
p = 0.99
0.41*
p = 0.04
0.11
p = 0.63
0.51*
p = 0.01
0.22
p = 0.30
0.20
p = 0.35
0.35
p = 0.10
0.36
p = 0.08
0.59**
p = 0.01
0.46*
p = 0.02
0.51*
p = 0.01
0.51*
p = 0.01
0.39
p = 0.60
0.49*
p = 0.02
0.50*
p = 0.02
0.10
p = 0.64
0.42*
p = 0.04
0.20
p = 0.36
0.67**
p = 0.00
0.61**
p = 0.01
0.51*
p = 0.01
0.33
p = 0.12
0.52**
p = 0.01
0.53**
p = 0.01
Mealy
Parrot
0.05
p = 0.82
0.44*
p = 0.03
Chestnut
Fronted
Macaw
0.24
p = 0.26
White
Eyed
Parakeet
Dusky
Headed
Parakeet
Orange
Cheeked
Parrot
Blue
Headed
Parrot
Yellow
Crowned
Parrot
5.3 Parrot abundances and days of disturbance
Day 5 had the highest mean total parrot numbers with a value of 6.8 (Figure 4). On the
same day an extremely low disturbance value of 0.2 was recorded. Day 1 and 17 have
the lowest mean value for total parrots, however recorded large disturbance figures of
0.8 and 1.0 respectively (Figure 4). These results seem to show links between low
disturbance levels and high parrot visitation numbers. However the results are
extremely varied over the whole study period. Day 27 recorded the second highest
mean total parrot value of 6.2 but it also recorded the highest disturbance levels over
the whole observation time with a figure of 1.5 (Figure 4).
Figure 4: Mean disturbance levels compared to mean parrot visitation numbers per day.
26
7.00
Mean Disturbance Levels/Mean Total Parrots
Mean Total Parrots
Mean Disturbance
6.00
5.00
4.00
3.00
2.00
1.00
0.00
1
2
3
4
5
6
7
8
9
10 11 12 13 14 15 16 17 18 23 24 25 26 27 28
Day Number
5.4 Species associations with different disturbance factors
Species associations with disturbances were calculated using Spearman’s rank
correlation analysis. Yellow-crowned parrot Amazona ochrocephala and white-eyed
parakeet Aratinga leucophthalmus were the only species to show any correlation with
any disturbance factors (Table 4). Amazona ochrocephala were correlated at the 0.05
level with boat disturbances and at the 0.01 level with tourist disturbances. Aratinga
leucophthalmus were also correlated with tourist and boat disturbance but only at 0.01
levels (Table 4). No species showed any association with animal disturbances. The
results of unknown disturbances were minimal and no data analysis has taken place.
27
Table 4: Species/disturbance associations calculated using the Spearman’s rank correlation (** = strongly
associated * = associated).
Red and
Green
Macaw
Chestnut
Fronted
Macaw
Mealy
Parrot
Yellow
Crowned
Parrot
Blue
Headed
Parrot
Orange
Cheeked
Parrot
Dusky
Headed
Parakeet
White
Eyed
Parakeet
All
Parrots
Boat
Disturbance
+0.26
p = 0.23
+0.24
p = 0.26
-0.12
p = 0.57
+0.52**
p = 0.01
-0.06
p = 0.79
-0.30
p = 0.15
+0.20
p = 0.35
-0.45*
p = 0.03
+0.15
p = 0.47
People
Disturbance
+0.35
p = 0.10
+0.21
p = 0.33
-0.17
p = 0.43
+0.44*
p = 0.03
-0.15
p = 0.48
-0.30
p = 0.16
-0.07
p = 0.73
-0.48*
p = 0.02
-0.78
p = 0.71
Animal
Disturbance
-0.16
p = 0.47
+0.11
p = 0.60
+0.01
p = 0.10
-0.07
p = 0.76
+0.02
p = 0.94
+0.11
p = 0.63
-0.01
p = 0.97
-0.17
p = 0.44
+0.03
p = 0.90
5.5 General disturbance factors
Non-parametric tests using Kruskal-Wallis analysis shows a significant difference
between the general disturbance types and bird disturbance (flushes from trees X2 = 52.9
df = 3 P = <0.001, flushes from colpa X2 = 63.3 df = 3 P = <0.001). Boats attributed the
least disturbances to both birds in trees and on the colpa, with low means of 1.25 and
0.18 respectively (Table 5). Tourist disturbances were responsible for the greatest
amount of overall bird disturbance, with large mean values of 3.04 flushes from trees
and 2.85 flushes from the colpa (Table 5). Animal disturbances had a high mean value
of 3.33 flushes from trees, but a low value of 1.67 from the colpa. Both means for
animal disturbance have high standard deviations indicating that animal disturbances
resulted in variation between flushes in the trees and colpa (Table 5).
Table 5: Mean and standard deviation values comparing general disturbance factors with tree and colpa
flushes.
Boats
(N = 103)
Tourist
(N = 46)
Animal
(N = 3)
Unknown
(N = 15)
Mean
Std. Dev
Mean
Std. Dev
Mean
Std. Dev
Mean
Std. Dev
Flush From
Trees
1.25
1.22
3.04
1.56
3.33
2.08
3.40
1.30
Flush From
Colpa
0.18
0.76
2.85
2.27
1.67
2.90
2.60
2.29
28
5.6 Tourist disturbance factors
Results of the tourist disturbance factors examined in this study, using Kruskal-Wallis
non-parametric analysis, shows there is no significant difference between different
tourist disturbance types and flushes from trees or the colpa (flushes from trees X2 =
0.486 df = 4 P = 0.975, flushes from colpa X2 = 9.908 df = 4 P = 0.042). Mean analysis
of tourist attributed flushes show varied disturbance types affect birds in the trees and on
the colpa differently. Quiet talking had a mean value of 3 flushes from the trees
compared to a 0 value of flushes from the colpa (Table 6). Coughing and sneezing were
the largest disturbance factor overall with mean figures of 3.75 from the colpa and 3.5
from the surrounding trees (Table 6). Loud talking affected birds on the colpa the most
recording a mean of 3.95. The least disturbance caused was by the arrival and departure
of tourists, however a high standard deviation of 2.52 for colpa flushes shows variances
in disturbance between days.
Table 6: Mean and standard deviation values comparing tourist disturbance factors with tree and colpa
flushes.
Human
disturbance factor:
Mean
Quiet talking (N = 3)
Std. Dev
Mean
Loud talking (N = 18)
Std. Dev
Mean
Cough/sneeze (N = 4)
Std. Dev
Mean
Dropped object (N = 9)
Std. Dev
Mean
Arrival/departure (N = 11)
Std. Dev
Flush from trees
3.00
2
3.06
1.35
3.5
1.74
3.00
1.21
2.82
1.10
Flush from colpa
0.00
0
3.95
1.59
3.75
2.5
2.33
2.30
1.82
2.52
5.7 Boat disturbance factors
Kruskal-Wallace non-parametric analysis shows there is no significant difference
between boat disturbances and flushes from trees or the colpa (flushes from trees X2 =
2.391 df = 2 P = 0.302, flushes from colpa X2 = 5.487 df = 2 P = 0.064). Mean analysis
shows that loud boats not stopping was the boat disturbance factor that affected birds on
29
the colpa and surrounding trees the most (Table 7). Tourist boats not stopping and quiet
boats stopping both have low mean disturbance values for colpa flushes. Quiet boats not
stopping caused the least disturbance to birds in the trees (Table 7). Tourist boats
stopping have a higher mean disturbance value for tree flushes but a lower standard
deviation, indicating variability in bird reactions to different boat disturbance.
Table 7: Mean and standard deviation values comparing boat disturbance with tree and colpa flushes.
Human
disturbance Type:
Quiet
boat
not Mean
stopping (N = 56)
Std. Dev
Loud
boat
not Mean
stopping (N = 8)
Std. Dev
Tourist boat stopping Mean
(N = 103)
Std. Dev
Flush from trees
1.11
1.22
1.63
1.77
1.38
1.10
Flush from colpa
0.18
0.72
0.88
1.81
0.06
0.32
6.0 DISCUSSION
6.1 Abundance
6.1a. Most abundant species
In total eight parrot species were recorded consuming clay from the La Torre colpa.
Dusky-headed parakeet Aratinga weddellii and blue-headed parrot Pionus menstruus
were the two most abundant species to use the study site, with 337 and 209 individuals
respectively recorded over the study period. Numbers feeding per day varied from 034 Aratinga weddellii and 0-22 Pionus menstruus. This is similar to group sizes
recorded from other similar sites in the area (Hammer 2002, Brightsmith 2004). On
days when both species were absent from the colpa very low numbers of other species
were recorded. This is because larger species rely on Aratinga weddellii and Pionus
menstruus as an indicator of danger on or around the colpa, due to their relatively
smaller size and therefore increased risk of predation (Burger et al 2003, Blumstein et
30
al 2005). Hammer (2000) also recorded this use of “lead” species when studying a
similar colpa in the El Gato region of the TCRZ.
6.1b. Least abundant species
Red and green macaw Ara chloroptera was the least abundant species observed using
the colpa, with only 2 individuals being recorded on one day. This is due to its large
size and the relatively small size of the colpa. Larger colpas in the area attract vast
gatherings of Ara chloroptera, and other large macaw species (Brightsmith et al 2004,
Tatum-Hume et al 2003). Larger macaw species have slower reaction times than the
smaller parrots and parakeets, so favour colpas with a large surface area, allowing for
easier predator vigilance and escape (Renton 2002).
6.1c. General abundance of species
Mealy parrot Amazona farinosa, yellow-crowned parrot Amazona ochrocephala,
orange-cheeked parrot Pionopsitta barrabandi and white-eyed parakeet Aratinga
leucophthalmus all were similar in abundance to each other. Each species had some
days when no individuals were present and other days when 5-20 individuals would
use the colpa. This may be due to the fact that there are several colpas in the
immediate area that the birds can visit with no preference over which one (Duffie
2003, Brightsmith et al 2004). All these species are similar in body size (Appendix 2),
and are smaller than the least abundant species Ara chloroptera, and larger than the
smallest and most abundant species of Aratinga weddellii. This shows how this small
colpa is dominated by parrot and parakeet species over larger macaws. As said in the
above literature large macaws do not use this colpa due to fear of predation (Renton
2002). Studies into avian behaviour at limited resources show that larger macaw and
parrot species may act aggressively towards other smaller, less dominate species
(Burger et al 2003, Mac Nally et al 2005). The lack of these larger species and
associated aggression can explain the large abundance of smaller species using this
colpa (Burger et al 2003). This is similar to other studies, where sites with a smaller
surface area are dominated by smaller species, whilst others with a larger and more
exposed area have the most abundance occurring in larger species (Tatum-Hume et al
2003, Duffie 2003, Gilardi et al 1998).
31
6.2 Species associations
Species association was calculated using Spearman’s rank correlation analysis. All
species recorded apart from red and green macaw Ara chloroptera showed association
with blue-headed parrot Pionus menstruus and dusky-headed parakeet Aratinga
weddellii. These were present most observation days and were usually one of the first
and most confident species to land on the colpa (pers. obs), as is the same from studies
into similar sites (Duffie 2003, Hammer 2002, Brightsmith 2004). This links with the
abundance data, as when these species were absent from the colpa, very low
abundances were recorded for the other species. This seems to prove that the larger
species are relying on these smaller, more predatory at risk species as indicators of
safety (Lima et al 1999, Karubian et al 2005). This has also been recorded in research
into other avian species feeding behaviour (Morse 1977, Brown 1969). With relevance
to this study it is obviously vital that disturbances cannot reach the level where they
completely deter Pionus menstruus and Aratinga weddellii from feeding. If this
happens the other species that use this colpa will not feed.
6.3 Species interdependence
In general associations were seen amongst all species, except for red and green macaw
Ara chloroptera, owing to it only being observed on one day. The associations shown
between the other species in this study illustrates how interdependent on one another
they are as indicators of safety (Lima et al 1999, Hilton et al 1999). This is explained
by the decreased risk of predation when flocking behaviour is adopted, due to
collective vigilance, dilution of risk and predator confusion (Hilton et al 1999, Barbosa
1997, Szekely et al 1989). This is important when considering disturbance factors at
this site, as a disturbance that deters one species, will consequently help to deter
another through its lack of presence.
6.4 General disturbance and parrot abundance
Mean disturbance levels show very little correlation with parrot abundance. Some
days with high mean disturbance levels still recorded high parrot visitation numbers.
This indicates that it is not the general level of disturbance but certain disturbance
32
factors that influence parrot abundance on the colpa. Previous studies on the effects of
human disturbance on avian behaviour have highlighted this (Webb et al 2005,
Lafferty 2004), as well as the need to identify the key factors that cause the most
disturbances (Guillemain et al 2007, Heil et al 2006, Stillman et al 2007).
6.4a. Associations with general disturbance factors
Only two out of the eight parrot species in this study showed any associations with the
three main disturbance factors tested (boat, tourist or animal). Flushes by yellowcrowned parrot Amazona ochrocephala and white-eyed parakeet Aratinga
leucophthalmus were correlated with boat traffic and tourists. It has been well
documented that boat traffic and human presence do negativly affect bird abundance
and behaviour (Stalmaster et al 1997, Bratton 1990). However, it has also been shown
that there is variation in response to disturbance between different species (Gill et al
1999). This depends on their natural reaction to predation and species life history in
the area (Hansen et al 1992). In this case Amazona ochrocephala and Aratinga
leucophthalmus species may associate boats and people with increased danger due to
previous exploitation by humans (Burger et al 2003). This may explain their more
extreme reactions to tourist and boat disturbance over the other parrot species studied.
6.4b. Differences observed between general disturbance factors
A significant difference was found between disturbance type (boat, tourist, animal)
and overall bird disturbance. Boats caused the least disturbance whilst tourists caused
the most. This stretch of the River Tambopata is used regularly by boat traffic with an
average of about 8 boats going past the colpa during each day of observation (pers.
obs). The lack of overall reaction suggests that some habituation to boat disturbance
has taken place in the species recorded at this site (Carney et al 1999). This has been
illustrated in other studies of repeated boat disturbance on avian species (Burger et al
2003, Guillemain et al 2007). Animal disturbance only recorded three incidences over
the study period, accounting for a very small percentage of overall disturbances. This
makes this irrelevant to this study, as they are not a major disturbance issue at this site.
When considering management implications for this site this needs to be taken into
account, as tourists are disturbing birds the most.
33
6.5 Tourist disturbance factors
6.5a. Cough/sneeze and dropped objects
Tourist disturbance factors were split into five categories; Arrival/departure, quiet
talking, loud talking, cough/sneeze and dropped object. Overall coughing and sneezing
were the factors that caused the most disturbances to birds on or around the colpa.
These types of disturbances can be classified as “impulse” noises, due being noncontinuous and originating suddenly (Larkin 1996). Extensive research undertaken on
the subject has been focused mainly on the effects of military impulse noises such as
artillery and gunfire (Tazik et al 1992, Schomer 1994). These have shown that impulse
noises cause greater disturbance to wildlife than continuous levels of sound (Tazik et
al 1992, Schomer 1994, Pater et al 1999, Larkin 1996). Dropped objects can also be
classed as impulse noises, and in this study resulted in relatively large amounts of
disturbance to birds on the colpa and in the trees. This highlights the need for the
reduction in impulse noises when observing parrots at the colpa to minimise
disturbance.
6.5b. Loud talking
Loud talking was the most commonly recorded tourist factor, and disturbed birds on
the colpa and surrounding trees to similar levels. Loud talking can be described as a
continuous noise but also has impulse noise properties due to its often-sudden onset
(Surman 2006). Flushes were caused at the start of loud conversations, and commonly
resettlement into at least the surrounding trees took place whilst the talking continued
(pers. obs.). Other studies into noise disturbance on birds have found similar behaviour
to occur in the European starling Sturnus vulgaris (Rich et al 2005) and marbled
murrelet Brachyramphus marmoratus (Singer et al 1995). This indicates that some
habituation to the levels of noise has taken place but the natural response to the sudden
onset of loud talking is still present. This needs to be considered in this study as loud
talking forced birds of the colpa and therefore reduced their feeding time.
34
6.5c. Quiet talking
Quiet talking was the least disturbing of the tourist factors analysed in this study. Birds
feeding on the colpa saw no reaction, and only minor reactions were seen to birds in
the surrounding trees. This can be attributed to parrot habituation to tourist visitation
at this site (Yorio et al 2001). This has been well documented in other avian species
where blinds are used for tourist observation (Walker et al 2006, Bouton et al 2005,
Hidinger 1996). Quiet talking can be described as a continuous noise factor with no
defining start and finish (Larkin 1996). As seen in the above literature continuous
noise factors cause fewer disturbances to birds than sudden impulse noises (Tazik et al
1992, Schomer 1994, Pater et al 1999). This has been illustrated in this study when
comparing parrot reactions to coughing and sneezing with reactions to quiet talking.
6.5d. Arrival/Departure
Tourist arrival and departure resulted in minimal disturbances to the various species
studied at this site. Disturbance was minimised to feeding birds by tourist arrival
usually being before feeding had started and departure once feeding had finished (pers.
obs). This meant that there were few, if any birds in the area when these activities
were taking place. It also meant that most birds arrived after tourist settlement into the
blinds had occurred. This has been found to significantly reduce the impact of humans
at other wildlife study sites (Tershey et al 2002, Wasser et al 1997). Birds in the trees
showed some responses but reactions were not as strong as with the other disturbance
factors of loud talking, dropped object or coughing/sneezing. These responses (or lack
of) can be attributed to the habituation of human activity at the landing site on the
beach (Pfeiffer 2004). Reduced reactions to disturbances caused by the arrival and
departure of tourists from the same frequently used point have been recorded from
similar studies into the effects of human disturbance (Yosef 2000, Pitts 2001,
Grossberg et al 2003). Intolerance of tourists has been observed from other wildlife
studies and is usually marked with a decline in abundance of the species concerned
(Enzenbacher 1994, Tershy et al 1997). However the continuous use of this colpa by
various parrot species indicates that the disturbances attributed to tourist arrival and
departure does not unduly deter birds from feeding.
35
6.6 Boat disturbance
6.6a. Loud boats not stopping
Disturbances by boats were assessed by noise, and whether or not they stopped at the
beach in front of the colpa. Quiet boats were classed as ones that used outboard motors
and loud boats were ones that used more traditional peke-peke motors. Loud boats
going past the colpa resulted in the most flushes to feeding birds as well those perched
waiting to feed. This was the least common form of boat disturbance and may account
for increased bird reactions due to habituation to the other forms (Belanger et al 1989).
Peke-peke motors are significantly louder and deeper in pitch than outboard motors
(pers. obs). Research shows that it is a combination of both the sound and sight of
boats that causes disturbance, with increased reactions to fast moving power boats
over quieter, slower moving vessels, such as sail boats or canoes (Rogers et al 2002,
Mosisch et al 1998). The length of disturbance time is increased with peke-peke
motors as they are slower and take longer to pass the colpa (pers. obs). This increases
exposure time to human sight and sound disturbance and therefore gives more time for
a flush reaction to take place (Draulans et al 1985, Gill et al 1999). Species life history
may also be taken into account when considering boat disturbance. The species in this
study may associate the traditionally used peke-peke motor with previous hunting and
exploitation (Martin 1995, Hansen et al 1992). Increased reactions to this type of
disturbance may have been passed from generation to generation contributing to the
lesser reaction to newer outboard motors over traditional peke-peke (Dobson 1989).
6.6b. Quiet boats not stopping
This factor resulted in the least disturbance to birds in the trees, and birds on the colpa
showed only minor reactions. Disturbance levels varied in scale with some days
having only a few boats going past whilst others recorded more than ten incidences.
The high level of use of the river for transport and tourist lodges can account for this
(pers. obs.), with decreased bird reactions been attributed to the habituation of
outboard powered boats (Bright et al 2003). Literature about boat disturbance on
parrots is limited, but habituation of this type has been recorded amongst many other
36
aviforme species (Vos et al 1985, Watson et al 1999, Newbrey et al 2005, Peters et al
2007). As written above, the decreased volume of outboard motors compared to pekepeke’s will also have affected parrot behaviour, due to increased flush reactions
caused by louder, longer lasting peke-peke disturbance events (Draulans et al 1985,
Gill et al 1999).
6.6c. Tourist boats stopping
This factor caused the least disturbance to birds on the colpa and resulted in minimal
disturbance to birds in the surrounding trees. As seen when considering tourist
disturbances in the above sections, the lack of reaction by birds on the colpa was due
to the times of tourist arrival and departure (Tershey et al 2002, Wasser et al 1997).
Arrival would usually be before any birds had started feeding and departure would
usually be after all the birds had stopped feeding (pers. obs.). This resulted in the
minimum of disturbance to feeding birds. Disturbance to birds in the surrounding trees
was greater than that of quiet boats not stopping but less than that of loud boats not
stopping. The close proximity (within 30 m of the colpa) of the landing site and clear
view of tourist activities by perched birds would have attributed to this (FernadezJuricic et al 2001, Hill et al 1997). However, the lack of overall negative reactions to
tourist boats landing indicates tolerance and habituation to these activities (Burger et
al 2003). This behaviour has been found to occur commonly in many bird species
exposed to high rates of tourist visitation (Carney et al 1999, Bright et al 2003,
Watson et al 1999)
6.7 Limitations of the study
This study was relatively successful in establishing the main effects that tourist
visitation is having on at this site, however there are improvements that could be made
in future research. More analysis of tourist disturbances should take place, such as
amount of activity in the blind and colour of clothing worn. These factors may
influence parrot behaviour due to their being some outside visibility into the blind. If a
future study was to take place more information on boat disturbances should be
gathered. This should include clear identification of who is responsible for the boat,
whether it be tourist lodges or locals. This can then be used to clearly establish the
37
effects of the many tour-operated boats that rely on the river to transport guests.
Identification is needed on the actual quantity of clay certain parrot species need to
consume to remain healthy. This is currently unknown and it is therefore hard to
assess whether parrots are actually consuming enough clay. A better understanding of
this would greatly improve the current knowledge of the effects of tourism on parrot
behaviour at geophagy sites.
6.8 Management implications
6.8a. Abundance of species
Research into geophagy in parrot species has illustrated the importance and necessity
of this activity in order to survive and remain healthy (Brightsmith et al 2004, Gilardi
et al 1999). If disturbances were too great birds would be driven away to other quieter
colpas in the area, or at worst possibly not feed at all. In this study, general boat and
tourist disturbances didn’t seem to be deterring feeding parrots at this site. The study
illustrates that on days with high disturbances recorded, high parrot visitation and
feeding numbers could also be observed. The species that utilize this site are fairly
common and none are threatened in the near future (IUCN 2007). Key species such as
Aratinga weddelli and Pionus menstuus need to have their presence maintained at this
site in order to attract other species. This site can continue to be used for tourist
visitation, as disturbances caused are not unduly deterring the various parrot species
from feeding.
6.8b. Tourist disturbances
There are few papers available on the impacts of tourist visitation to parrot geophagy
sites (Tatum-Hume et al 2004, Duffie 2003, Hammer 2002). The results of this study
have highlighted factors that cause the most disturbances to feeding and perched birds
at the La Torre colpa. Impulse noises such as coughing/sneezing, dropping objects and
the onset of loud talking are all clearly identified as major sources of tourist
disturbance. Although little can be done about coughing/sneezing simple measures
could be taken to reduce the impact of other tourist created impulse noises. A tougher
38
policy and enforcement of talking volumes when observing parrots from the blinds
should be employed. During this study, some tour guides from the lodges were good at
controlling sound levels coming from the blind, but others were not. This study has
demonstrated that quiet talking does not cause significant amounts of disturbance.
Therefore tighter control of noise volume at a quiet level would significantly reduce
the disturbance that tourist visitation is currently having. Inotawa Lodge and Posada
Amazonas could easily do this, by informing tourist guides of appropriate talking
levels and making sure they enforce them. As well as loud talking, dropped objects
also represented a significant proportion of tourist created disturbance. There is little
that can be done about this, as it usually resulted from clumsiness and accidents.
However, padding of some sort could be used as a floor cover in the blind, on top of
the wooden planks that are currently in place. This would provide some insulation
from dropped objects and hopefully reduce the impact that this type of disturbance is
having.
6.8c. Boat disturbances
There has been much research undertaken on the impacts of boat disturbance on avian
species such as waterfowl and shorebirds (Belanger & Bedard 1989, Lafferty 2004,
Stillman et la 2007). However there have been few studies on the impacts of boat
disturbance to feeding parrots at geophagy sites (Tatum-Hume et al 2004). In terms of
the management implications of this study there is little that can be done about boat
disturbance. Arrival and departure of tourists was usually before and after any birds
had started feeding, so the impacts of this activity had already been reduced as much
as possible. Boats that used quiet, outboard motors (usually operated by a tourist
lodge) caused very little disturbance to feeding birds. Studies have however shown
that increases in boat traffic in the future could be enough to deter birds from feeding
permanently (Tatum-Hume et al 2003). Monitoring the levels of boat traffic on the
river should be undertaken to ensure it remains under control. Boats using peke-peke
motors caused the most disruption to parrots at this site. As these are mostly used by
locals there are limited options when considering management implications for this
factor. Local people in the area cannot afford newer outboard motors, so there is no
option for the discontinuation of peke-peke use. Education and information about this
site and other sites in the area could be provided to locals with recommendations for
39
speed limits and passing distances (Fernadez-Juricic 2001). This has been shown to
work in other situations where boat disturbance is affecting wildlife (Bratton 1990).
Inotawa and Posada, as a way of helping to insure that parrots are not driven away
from the colpa, could provide this.
7.0 CONCLUSION
This study has highlighted the clear necessity of geophagy in the parrot species that
use the La Torre colpa. It has also identified the tourism related factors that are
responsible for the most disturbances to birds using the site. Analysis of these has
shown that management strategies can be put into place to reduce the amount of
disturbances that are currently occurring. Simple guidelines on sound levels when
observing the colpa and floor padding to reduce the impact of dropped objects, would
dramatically reduce the amount of disturbance that tourist visitation is contributing.
Boat disturbance affected birds to a lesser degree than tourist visitation. Habituation to
quiet boats and the times of tourist arrival and departure meant that few disturbances
could be associated with these factors. Locally run boats with peke-peke motors were
the boat factor most associated with disturbance. It is not economically viable to
discontinue the use of these so information needs to be provided about the importance
and location of colpa sites, appropriate speed limits and passing distances. This should
jointly be undertaken by the lodges along the river that rely on the tourism provided by
the many parrot geophagy sites in the area.
Future research into other tourist
disturbance factors and tourism related boat traffic should be undertaken to help
further the current knowledge on tourism disturbances at geophagy sites. This should
be accompanied by studies into the quantity of clay and the time needed to consume
enough to maintain parrot health. If these suggested management strategies are
implicated disturbances to the many parrot species that consume clay on the La Torre
colpa can be better understood. This will help to reduce the effects that tourism at this
site and others like it is having.
40
8.0 Acknowledgments
Thanks to my project supervisor, Dr. Stuart Marsden, for all the expert help and advice
kindly given to me throughout this project, even at times when he didn’t have to. Alan
Lee for putting up with me trying to learn parrot calls and identification in the field, as
well as providing an excellent data sheet to use during my study. Lastly I would like
thank Rolando, Maria and Frieda, the family I stayed with when conducting my field
research, they made me very welcome.
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10.0 Appendices
Appendix 1: Example of data sheet used to record parrot abundances and flushes
(provided by Alan Lee)
52
Appendix 2: General information about the parrot species recorded at the La Torre
colpa
53