69EG3191 Third Year Project

Factors affecting flight ecology of macaws, parrots and parakeets on the La Torre
colpa, Rio Tambopata, Peru.

Mr D.E. Crabtree

A project submitted in partial fulfilment of the requirements for the degree of Bachelor of
Science (Honours) in Ecology and Conservation. The
Manchester Metropolitan University

Department of Environmental and Geographical Sciences

The Manchester Metropolitan University

April 2007

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 Ymroddedig i’r Cymry

Acknowledgements

My project coordinator Dr. Stuart Marsden for the help and advice, Alan Lee and for
aiding in data collection, Rolando and the family for the accommodation and a great time
in the jungle, and finally Richard Amable for teaching me about Peruvian culture and
customs.

Declaration of originality

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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explicitly acknowledged (including citation of published and unpublished sources).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 of for any other purpose.

Signed ………………………………….....

Date ……………………………………..

Abstract

This study explored patterns of early morning flight activity and flocking to and from a clay
lick in a diverse community of parrots in a Peruvian Amazonian lowland rainforest. Parrots
were most active just after sunrise with a second peak of flight activity half way through
morning feeding. Most parrot species flew in groups of one to four individuals; suggesting
that mated pairs are stable and that family groups remain together post-fledging. The flight
patterns of each species except for spp. Artinga were non-random, suggesting a particular
nesting location for the species. The birds were observed daily as they were disturbed from
the ‘clay lick’ by either the local boats with peke-peke motors, or by the boats with outboard
motors transporting tourists to and from Puerto Maldonado. The former (i.e. local boats)
were found to cause the greater disturbance suggesting that the birds' behaviour had adapted
to recognise the locals as a potential threat. This may be because the macaws and parrots
were hunted or caught for the pet trade in the recent past, or may indeed be due to continued
exploitation.
Table of Contents

1. Introduction (Page 6) 1.1 Introduction

1.2 The Psittacine Order

1.3 Psittacine behaviour

1.4 Psittacine conservation status

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 1.5 Geophagy in south-eastern Peru

1.6 Conservation and economic value of ‘clay licks’

1.7 Aims

1.8 Objectives

2. Methodology (Page 14) 2.1 Study Site

2.2 Experimental Design and Implication

2.3 Analytical and Statistical Methods

3. Results (Page 20) 3.1 Avian Abundance

3.2 Incoming Flight Directions

3.3 Landing Locations

3.4 Species and Weather Associations

3.5 Disturbances

3.6 Direct Flight Results

4. Discussion (Page 28) 4.1 Abundance, Incoming flight directions and Landing
Locations

4.2 Large Species of Macaw

4.3 Associations

4.4 Weather

4.5 Disturbance

5. Conclusion (Page 35) 5.1 Conclusion

5.2 Improvements to the design of the study

5.3 Management implications

6. Reference List (Page 38)

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Tables

Table 1. Scientific and common names of the species studied, including geographical range and IUCN conservation status.
Detailed information in Appendix 1.
Table 2. Historical investigations regarding geophagy within avifauna in the Tambopata area.
Table 3. The number of recordings of each of the eight species plus the number of groups recorded over the 14 day period.
Table 4. The number of groups recorded on the incoming flight from each zone, including the percentage for that species:
Table 5. The number of groups recorded landing in each section surrounding the colpa, including the percentage for that
species:
Table 6. Tests of association between species with the chi-squared (X²) and p – values.
Table 7. The total mean percentages of the birds leaving the colpa site when a flush was initiated, and the percentage that
returned from the original number that left.
Table 8. The recorded causes behind the flushes
Table 9. The abundance counts for the three large species of macaw. Included are group sizes and individual numbers:
Table 10. This table shows the flight direction these large species of macaw flew.

Figures

Figure 1. The Tambopata river, colpa and research site.

Figure 2. How the sky surrounding the colpa was fragmented in to sections
Figure 3. The colpa and the surrounding 6 zones of vegetation
Figure 4. Activity graph representing the mean number of individuals from each data set plotted against the morning time

1.1 Introduction

Peru is located on the Pacific coast of South America and is the third largest country on the
continent. Two thirds of Peruvian territory is located within the Amazon basin. The Amazon
rainforest is well known for being the zenith of faunal and floral biodiversity (De Castro
1985) including bird species that surpass a fifth of all avifauna that exists (Beiregaard et al
2001). It is said to be one of the most biotically rich terrestrial biomes in the world
(Heywood 1995), but at the same time, it has been impinged on by heavy anthropogenic
disturbances and remains a high priority for conservationists (Hubbell & Foster 1992). This
area is also home to a number of landmark animals listed in the IUCN's Red Data Book.
Amongst them the giant armadillo (Priodontes maximus), ocelot (Leopardus pardalis),

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jaguar (Panthera onca), giant river otter (Pteronura brasiliensis), harpy eagle (Harpia
harpyja) and black caiman (Melanosuchus niger).

1.2 The Psittacines Order

The study concentrated on several species from the order Psittacines (which contain macaws,
parrots and parakeets). This order is one of the most diverse orders of birds containing a total
of around 353 species (Forshaw 1989). They are found in most neo-tropical areas of the
world and have the characteristic curved beak (Juniper & Parr 1998). Their origin is still a
mystery but there are suggestions that they originated from the precedent Gondwanan
supercontinent, due to the present diversity of Psittaciformes in South America and
Australasia (Juniper & Parr 1998).

1.3 Psittacine behaviour

Social behaviour varies between different psittacine species. Solitary behaviour is the
exception (Kakapo, Strigops habroputilus) with most species pairing for life and showing
complex social organization (Luescher 1997). Flock formation is also important, helping
with predator detection and avoidance, access to mates, defence of territories and foraging
efficiency (Wilson 1975). Previous studies on psittaciform species have shown their habitat
preferences are primary and secondary forest with a relatively high number of palm species
and emergents for nesting sites (Enkerlin-Hoeflich et al 1999). Eucalyptus plantation
habitats were the only totally anthropogenic habitat used by the macaws in this study
(Marsden et al 2005).

Griffin et al (2001) reviewed the importance and meaning of dominance interactions between
psittacine social groups. A dominance relationship exists when predictable dominance-sub
ordinance responses occur between members of a stable social group, based on the prior

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interactions between the individuals. Once relationships are established, there is consistency
in social interactions, resulting in fewer, or less intense, aggressive assertions of dominance
(Luescher 1997). Another important behaviour of the Psittaciformes order is allopreening
which occurs when an individual uses its beak to groom another bird. It is cited as the most
important mechanism for maintenance of the pair bond (Gill 1995)

1.4 Psittacine conservation status

Eight of the sixteen species of macaws are classified as Endangered and most populations of
macaws are declining (www.birdlife.org). Parrots have the highest percentage of threatened
species of all the world's bird families with the 17 macaw species are among the most
endangered (Collar 1997). There are currently fewer than 3,000 hyacinth macaws in the wild,
red-fronted macaws and blue-throated macaws are both thought to number fewer than 1,000,
and, most distressing of all, only one Spix's macaw remains in the wild (www.birdlife.org).
The reason is due to habitat destruction and the popularity of the wildlife trade (González
2003). One over-looked threat is demand for Mauritia or Aguaje palm fruit, which is one of
the macaw's favourite nesting sites and important food sources. The quickest way to collect
the fruit from these trees is to cut them down, which the locals undertake, thus eliminating
suitable nesting sites (Bonadie & Bacon 2000). The majority of macaws are dependent upon
forests both for food and for nesting sites making them a danger since the world's forests are
being destroyed at rapid speeds. The main causes are commercial logging and clearance for
agriculture (Burslem & Whitmore 1999).

The survival rate for young birds captured in the wild is very low, sometimes as low as 1%,
prompting trappers to often focus their attention on adults, which is extremely damaging to
populations (Nilsson and Mak 1980). To combat illegal hunting and exploitation an
organisation called CITES was set up in the early 1960s. The ‘Convention on International
Trade in Endangered Species’ is an international agreement between governments to ensure
that international trade in specimens of wild animals and plants does not threaten their
survival. They manage this by subjecting international trade in specimens of selected species
to certain controls. (www.birdlife.org)

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Their diet is relatively diverse, with most Psittacines feeding on fruits, berries, seeds and nuts
as well as unripe seeds and fruits, those with tough shells and spines, or distasteful ones
containing poisons (Renton 2001). This is one of the reasons why they carry out geophagy,
which is explained in the next section.

1.5 Geophagy in south-eastern Peru

Many of species from the Psittacidae genera congregate in large numbers around clay licks
(Colpas) (Brightsmith 2004a). It is here where they undertake geophagy, which is defined as
the intake of clay or soil into a species diet. It happens to mostly vertebrates, and has been
recorded in animals such as birds, sheep, zebra and even bears (Diamond et al. 1999; Cooper,
2000: Low, 2003). Geophagy is widespread and well documented in mammals, but avian
geophagy has only recently become the subject of serious scientific investigation
(Brightsmith 2004). In the Tambopata region of Peru alone there are many species of birds
and mammals that congregate around colpas regularly, including peccary and tapir (Rolando
pers. Comm). There are many a theory behind why certain species do this, with journals
demonstrating 1) how the clay contains minerals that the species necessitate (Jones &
Hanson 1985) 2) others say it aids digestion (Best & Gionfriddo 1991), 3) neutralises dietary
toxins (Diamond et al. 1999) 4) or protection of the gut lining (Gilardi 1996). Jones &
Hanson (1985) demonstrate how clay can contain trace amounts of minerals such as Sodium,
Iron and Calcium and are used as supplements to the diets of these species, as these are
usually hard to come by through customary food selection. Best & Gionfriddo 1991
comment on how clay can be used as a mechanical aid to assist digestion. It is common in
other species of bird in which the small stones are held in the muscular gizzard and used to
grind seeds before they are passed to the stomach (Moss 1989). Brightsmith 2002 explains
that plants can be protected by toxins such as tannins and caffeine and in large quantities can
be poisonous to the organism. When clay is eaten the very fine negatively charged particles

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bind with the positively charged dietary toxins. When united they prohibit entry into the
bloodstream and are passed out in the faeces. He also explains that clay can protect the gut
lining by sticking to the lining of the stomach providing a physical barrier between the toxic
food and the stomach. Geophagy has been found to be important in domestic animals and
therefore seeming logical to assume that this mechanism of lining the stomach is also helping
some parrots and macaws in neo-tropical areas (Klaus & Schmid 1998).

It is probable that any of the above theories are correct under the right circumstances, but
geophagy in most species is poorly understood and may be a complex combination of all the
above theories (Gilardi 1996).
Brightsmith (2004) explains that colpa use in southeastern Peru is highly seasonal. Most
species show simultaneous annual lows in lick use during the end of the wet season and
beginning of the dry season (April – June). Annual highs in lick use occur between July and
February and differ among species. The peaks in colpa use for most species studied coincide
with breeding. In particular it seems that adults are feeding clay to their young chicks during
the period of maximum growth and perhaps lowest resistance to natural toxins found in their
diet.

Previous studies in the Tambopata area have shown that Macaws have a very low intake of
sodium in their diet (Brightsmith & Aramburú 2004) however the soil located at the clay
licks are rich in sodium, thus possibly explaining the geophagy in the Tambopata area. It has
also been suggested that social interactions are grounds for meeting at these colpas. Birds
may arrive at the colpas only to interact with members of the same species i.e. finding a mate
or learning of new food sources (Hammer 2001).

1.6 Conservation and economic value of colpas

This event gives the area huge ecotourism potential with visitors attracted by the spectacular
bird gatherings. Ecotourism is economically important and is now the third leading foreign
currency gainer for Peru (Stronza 2000). New eco-tourist lodges along the river help to
protect the rainforest of Tambopata while meeting the economic needs of the people in the

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 local community (Piana 1999). They try to provide training to community members in all
aspects of ecology management. A short distance from the research site there was and eco-
lodge called Posada. After 3 year of ownership the operators have made sure that all profits
and operations are handed over to the local community. (Posada Lodge is illustrated in Plate
1).
One area of concern is that as part of the holiday, Posada Amazonas organizes daily trips to
the colpas. There are no restrictions in place (pers. obs.) as to the number of visitors at the
site and there is obviously a conflict of interest in wishing to maintain the colpa in their
undisturbed state against possible disturbance by visitors. Scientists at the Tambopata
Research Centre have concerns that birds are being frightened away from the colpas before
having a chance to feed (Alan Lee Pers Comms)..
The names, geographical ranges and IUCN listings of the species studied are illustrated in
table 1. Illustrations and detailed information of these species are displayed in Appendix
One.
There is a large Macaw Project in the area, conducted by ecologists from the Tambopata
Research Centre at many different colpa locations. Manchester Metropolitan University have
a strong link with the project as one of the Ph.D. students is carrying out a research study in
the area. Present in table 2 are some of the key studies on geophagy in avian fauna.

Table 1: Scientific and common names of the species studied, including geographical
range and IUCN conservation status.

Scientific Name Common Name Weight (g) Range Population IUCN Redlist
Ara severa Chestnut-Fronted Macaw 430 5,800,000 km² Stable Least Concern
Aratinga weddellii Dusky-Headed Parakeet 110 2,300,000 km² Increase Least Concern

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Amazona farinosa Mealy Parrot 800 7,200,000 km² Decline Least Concern
Pionus Menstrus Blue-Headed Parrot 293 8,300,000 km² Increase Least Concern
Ara chloroptera Red and Green Macaw 1250 8,100,000 km² Decline Least Concern
Ara macao Scarlet Macaw 1015 6,700,000 km² Decline Least Concern
Ara ararauna Blue and Gold Macaw 1125 7,800,000 km² Stable Least Concern
Brotogeris cyanoptera Cobalt-Winged Parakeet 67 2,600,000 km² Stable Least Concern

Table 2: Historical investigations regarding geophagy within avifauna in the

Tambopata area.

Authors Title Summary of findings

Effects of weather on parrot Discovered there was greater colpa use on
D. Brightsmith 2004 geophagy in Peru sunny mornings than on rainy ones
Parrot colpa and geophagy The most likely reason for clay eating is
M. Hammer 2001 behaviour from the El Gato detoxification at this site.
Biochemical Functions of
JD Gilardi et al 1999 Geophagy in Parrots Role of geophagy is dependent on location
Avian geophagy and soil Preferred soils were deficient in particles large
characteristics on a colpa in enough to aid in the mechanical breakdown of
D. Brightsmith 2004 South-eastern Peru food and help digestion

1.7 Aims of this project

This paper presents a study of flight ecology on species from the Psittacidae genera around a
colpa in south-eastern Peru. The aim is to illustrate if there are any factors present at the site
that disrupt or disturb the feeding of flight ecology of the species.

1.8 Objectives

Through the use of various methods of non-parametric analysis, the objectives are (1 to
quantify the factors affecting the flight and feeding of various species of Psittacidae landing
at the colpa 2) to propose a scenario to explain these factors, for the benefit of future
sustainable colpa use for tourism.

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2. Methodology

2.1 Study Site

The research site was located at the edge of the buffer zone of the Tambopata National
Reserve (26K south-west of Puerto Maldonado, SE Peru, 12º 48’ S, 69º 18’ W). The reserve
originated in the late 1970s as the small Tambopata Reserved Zone comprising of about
5,000 hectares, but was enlarged to 1.5 million hectares in 1990. The altitude of the park

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ranges from 240 to 3,500 metres, with habitats ranging from sub-tropical moist forest, cloud
forest, to tropical savannah. (Treves et al 2003). The study site was located in lowland
tropical forest and situated on the banks of an upper tributary to the Amazon, the Tambopata
River, near the La Torre Community in the Department of Madre de Dios. Annual
precipitation is around 2810 mm per year (Pearson & Derr 1986). There is a well-defined dry
season that starts in April and ends in October during which time rapid changes in
temperature are common, with a minimum of 8 ºC a maximum of 34 ºC and a mean of 24 ºC
(Kirkby et al. 2000).

The dimensions of the colpa were approximately 7meters in length by 3meter in height ,
situated on the bank of the Tambopata River. The river was about 30m wide at this point. It
meandered strongly with high mud cliff walls on the outside bends, with flat terrain on the
inside bend. This pattern was present at the site and the research area was set up on the
riverbank opposite the colpa. The distance between the research site and the colpa was about
40 metres. The colpa faced west; therefore sunrise was behind the colpa as seen from the
research site. Due to the sunrise being in the west in the southern hemisphere the colpa was
therefore in the shade in the morning and only getting some weak sunlight in the afternoon.
The location of the colpa relative to the Tambopata River is illustrated in Fig 1.
The habitat around the colpa consisted of mixed primary and secondary rain forest with
relatively little human impact. Human activity was concentrated along the Tambopata River
on the western bank, with the relatively unspoilt reserve on the eastern bank. The understory
plant species on the site consisted mostly of Giant Heliconias (Heliconia imbricate). Before
the creation of the National Park, large-scale timber extraction was common, but this has
since disappeared apart from some small-scale subsistence extraction. Some localized small-
scale gold mining is also practiced illegally (Crabtree pers obs).

Figure 1: The Tambopata river, colpa and research site. The east side of the river is
the Tambopata-Candamo Reserve which is mostly untouched by human development. The
south border of the reserve is at the La Torre River, which is visible at the bottom of the
figure (Google Earth 2007):

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2.2Experimental Design and Implication

The aim of this project was to quantify the macaws found at the Inotawa colpa, and where
possible to assess the biotic and abiotic factors that affect their abundance and flight ecology.

The survey began by selecting a site directly opposite the colpa that entailed a high quality
view of the surrounding sky. The sky was sectioned into 6 different zones, each covering
approximately 36°. This categorization is illustrated in Fig 2. The site was chosen and then
monitored collectively every morning for a total of 14 days between the dates of 02/07/06
and 16/07/06.

The morning study protocol was as follows: Arrival was shortly before sunrise and
observations were made until the early morning lick activity ended (usually between 05h30
and 07h30 EST). The weather conditions were noted as either rain (rain falling on the

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observer), sun (sun hitting the ground anywhere in site), or cloud (if neither of the others
applied). The presence or absence of fog was also noted. Flock activity was then monitored
over the study area and the following would be recorded for further study:

 Species of Psittaform
• Size of flock
• In-flight calls
 Flight direction and angle of incoming birds
• Flight direction and angle of outgoing birds
• Location around the colpa on which the species landed
• Whether boat traffic and noise from the hide had any impact on the majority of birds?

Due to the site being next to the river, other factors which affected the bird while they
initiated in geophagy would also be recorded. This included boat traffic and tourists at the
colpa hide as there was a growing concern for the welfare of Psittaciformes in the area due to
anthropogenic disturbance and their vulnerability at the colpas (Masello & Quillfeldt 2003).

To record the location where the species landed the adjacent trees surrounding the colpa was
divided into 6 zones. Zone 1 consisted of tall reed type vegetation at around 2.5 meters, Zone
2, 3, 4 and 6 consisted of mixed primary and secondary forest which enclosed several tall
emergent trees. Zone 5 contained a single tree that was approx 6 meters of height that
overshadowed the river.
Every group that landed was then labelled as a number between 1 and 6 corresponding to the
respective zone. The zones at the colpa are illustrated in Fig 3.

It was hoped also that incidental observations could be made from the e.g. the average time
taken for the birds to land on the colpa after arriving at the site. Previous studies by have
shown that larger species of macaw tend to be more nervous at colpas due to predator
evasion.

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The equipment used was writing apparatus, binoculars and a watch to record the time. For
the first 6 days a local bird guide was present to teach me how to distinguish the species
solely by the in-flight calls. This was necessary as low light levels at dawn cast the birds in
silhouette, making it essential not to rely solely on the visual data. Nineteen days of data was
collected and was analyzed using software such as SPSS with hoping to create a
comprehensive picture on factors affecting flight ecology of the psittaciform family in South-
eastern Peru.

Figure 2: How the sky surrounding the colpa was fragmented in to sections(Google Earth
2007)

Figure 3: The colpa and the surrounding 6 zones of vegetation (Authors photograph)

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2.3 Analytical and Statistical Method

Following the collection of the data, it was input in to SPSS Statview, which is a computer
statistical programme. The abundance and incoming/outgoing flight data for each species
was analysed using simple cross tabulations and percentages. This made it easy to compare
the different species as from which direction they originated from.

Because the number and nature of the parameters in the data were flexible and not fixed in
advance, and not normally distributed, they are known as non-parametric statistics. Because
of this, the Kruskal-Wallis Test and Spearman’s Rank Correlation Coefficient were used.
Wheater & Cook (2000) describe the Kruskal-Wallis Test as a one-way analysis of variance
and a non-parametric method for testing equality of population medians among groups by
ranks. It is an extension of the Mann-Whitney U test to 3 or more groups. Since it is a non-
parametric method, the Kruskal-Wallis test does not assume a normal population, unlike the
analogous one-way analysis of variance. However, it still assumes that population
variability’s among groups are equal.

Spearman's rank correlation coefficient is also a non-parametric measure but determines

correlation. It assesses how well an arbitrary monotonic function could describe the

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relationship between two variables, without making any assumptions about the frequency
distribution of the variables (Wheater & Cook 2000).

3. Results

3.1 Avian Abundance

3.1.1 General Description:

A total of 8 species, 1548 individuals and 332 groups were observed in the study time
frame. 4 out of the 8 species were not analysed in detail because the recorded quantity was
very low. Having plotted an activity graph denoting the mean number of species groups
against the time, it suggests that the busiest peaks in morning activity were at 6:05 AM and
6:45 AM. This is illustrated in Fig.1

3.1.2 Overall Species Abundance

Of every species recorded, Ara severa was the most abundant, closely followed by
Aratinga weddellii. The least abundant recorded were Brotogeris cyanoptera with 11
individuals. The numbers of the remaining species studied varied between these figures,
illustrated in Table 3.

3.1.3 Daily Species Abundance

All 8 study species combined to give an average of 111 individuals and 24 groups per
day. The mean group size was 4.71 individuals. The species of large macaws; Ara ararauna,
Ara macao and Ara chloroptera did not land in the area but flew directly over the site.

3.1.4. Group Sizes

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 All species except for A. weddellii flew in groups of 1 to 4 individuals. Artinga
weddellii had an average group size of 8 individuals. This are illustrated in table 4.

Table 3: The number of recordings of each of the eight species plus the number of
groups recorded over the 14 day period.

Scientific Name Common Name # of Groups # Of Individuals

Ara severa Chestnut-Fronted Macaw 156 642
Aratinga weddellii Dusky-Headed Parakeet 68 545
Amazona farinosa Mealy Parrot 63 213
Pionus menstruus Blue-Headed Parakeet 26 93
Ara chloroptera Red and Green Macaw 9 18
Ara macao Scarlet Macaw 6 14
Ara ararauna Blue and Gold Macaw 4 12
Brotogeris cyanoptera Colbolt-Winged Parakeet 1 11
Total 333 1548

Figure 4: Activity graph representing the mean number of individuals from each data
set plotted against the morning time

140 species
BHP
CFM
DHP
120
MP

100
M ean # O f G rou ps

80

60

40

20

5.55 6.05 6.15 6.25 6.35 6.45 6.55 7.05 7.15 7.25
Time

Table 4: Mean group sizes of the species studied. Brotogeris cyanoptera is not shown
as only one group was recorded.
Ara Aratinga Amazona Pionus Ara Ara Ara
severa weddellii farinosa Menstrus chloroptera macao ararauna
4.16 8.01 3.38 3.15 2 2.33 3

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3.2 Incoming Flight directions
Because the large macaw species did not land on or around the colpa they were excluded
from the flight analysis.

3.2.1 General description

Zone 3 recorded the most incoming flights with a total of 112 groups over the 14 day period.
The zone with the least incoming flights was Zone 5, with a total of 39 recorded groups. The
numbers of the remaining species studied in these zones varied between these figures,
illustrated in Table 4.

3.2.2 Species incoming flight directions

Ara severa
The incoming flight results showed that 76% of Ara severa groups passed via Zone 3 and
Zone 4 on route to the colpa area. Figures illustrated in Table 4.

Aratinga weddellii
The species Aratinga weddellii were more balanced in their directions. Each zone had
roughly the same amount of birds passing through. Figures illustrated in Table 4.

Pionus menstruus
Pionus menstruus was found to fly in from a 90° section of sky, ranging from North to South
West. This ranged from Zone 1 to Zone 3 with a total of 96% of the groups for this species
recorded coming from this direction. Figures illustrated in Table 4.

Amazona farinose

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89% of this species flew in from Zone 1 to Zone 3. This might suggest that the nesting sites
of Pionus menstruus and Amazona farinose might be located in this direction. Figures
illustrated in Table 4.
Table 4: The number of groups recorded on the incoming flight from each zone, including
the percentage for that species:

  Zone 1 Zone2 Zone 3 Zone 4 Zone 5

Ara severa 7 10 73 45 20
% 4 7 47 29 13
Aratinga weddellii 15 8 15 14 15
% 23 12 22 21 22
Amazona farinosa 9 30 15 3 4
% 15 49 25 5 6
Pionus menstruus 7 9 9 1 0
% 26 35 35 4 0

3.3. Landing locations

3.3.1 General Description.

A total of 313 groups were recorded landing in the vicinity of the colpa. The most popular
section was number 3, in which a total of 143 groups were recorded landing. The least
popular area for the birds was section 6, in which 7 groups were recorded. The numbers of
the remaining species are illustrated in Table 5.

3.3.2 Landing preferences

Ara severa
121 groups of Ara severa were recorded landing in section 3 and 4. This was 77.6% of all the
species recorded at the site. Section 1 and 5 were least used by this species, with 9 groups
landing in that area.

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Aratinga weddellii
This species preferred to land in section 3 with 53% landing in this area. Section 6 was not to
contain this species at any time. A total of 68 groups of Aratinga weddellii were recorded
on/near the colpa.

Pionus menstruus
This was the least abundant of all the regular colpa species. A total of 24 groups out of 26
landed in Section 2 – 4. No Pionus menstruus were seen landing in Section 5 or 6.

Amazona farinoa
The majority of Amazona farinose landed in section 2, with the remaining groups evenly
distributed in the other 5 sections. The percentage landing in section 2 was nearly 60%.

Table 5: The number of groups recorded landing in each section surrounding the
colpa, including the percentage for that species.
Section Section Section Section Section Section
  1 2 3 4 5 6
Ara severa 4 8 89 32 18 5
% 2.6 5.1 57.1 20.5 11.5 3.2
Aratinga weddellii 1 13 36 11 7 0
% 1.5 19.1 52.9 16.2 10.2 0
Amazona farinosa 9 37 8 3 4 2
% 14.2 58.7 12.7 4.8 6.3 3.2
Pionus menstruus 2 8 10 6 0 0
% 7.7 30.8 38.5 23.1 0 0

3.4. Species Associations and influence of weather

Note: Four species, Ara chloroptera, Ara macao, Ara ararauna and Brotogeris cyanoptera
were too rare to be analyzed; and therefore discussion on species associations was not
possible.

3.4.1 Species with associations

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Every species were positively associated with each other, however to different degrees.
Amazona farinose had the highest positive association with the other species with a p value
of 0.01 or less. Association between the remaining species were also positively correlated.
The figures illustrated in table 6:

Table 6: Tests of association between species with the chi-squared (X²) and p –
values.
Species Name Association against X value p-value Association
Ara severa Aratinga weddellii 0.655** < 0.001 Yes
Amazona farinosa 0.788** < 0.001 Yes
Pionus menstruus 0.431* 0.016 Yes
 
Aratinga weddellii Amazona farinosa 0.613** < 0.001 Yes
Ara severa 0.655** < 0.001 Yes
Pionus menstruus 0.414* 0.21 No
 
Amazona farinosa Ara severa 0.788** < 0.001 Yes
Aratinga weddellii 0.613** < 0.001 Yes
Pionus menstruus 0.561** 0.001 Yes
 
Pionus menstruus Ara severa 0.431* 0.016 Yes
Aratinga weddellii 0.414* 0.21 No
  Amazona farinosa 0.561** 0.001 Yes
*- Correlation is significant at the 0.05 level (2-tailed)
**- Correlation is significant at the 0.01 level (2-tailed)

3.4.2. Weather associations.

The numbers of groups seen each day were not positively associated with a certain type of
weather. 13 Days were recorded as sum and very little cloud, 3 days with heavy cloud cover
and 1 day with heavy mist.

3.5. Disturbances

23
3.5.1 General Description
A flush is described as a large flock of birds leaving the colpa due to a predator avoidance
technique. A total of 28 flushes were recorded over the 14 day period. This resulted in a large
number of birds leaving the colpa with only a small percentage returning. If the flush was
initiated before 7 am, there were a higher percentage of birds returning than if it had
happened later in the morning. Percentages illustrated in Table 7. The causes of the flushes
were either noted down as 1) boat traffic, 2) predators 3) noise from the hide and 4)
unknown. These are illustrated in Table 8.

Table 7: The total mean percentages of the birds leaving the colpa site when a
flush was initiated, and the percentage that returned from the original number that
left.

Time Total Flushes Mean % Flush Mean % Returned

After 6am 14 81.4 38.6
After 7am 14 91.4 31.4
Total 28 86.4 35

Table 8: The recorded causes behind the flushes

Boats Hide Predators Unknown

9 7 4 8

3.6. Species which made direct flights over the colpa

3.6.1 These are species that were not analyzed above and were recorded making a direct
flight over the colpa site. These species were Ara chloroptera, Ara macao and Ara ararauna,
which are all large species of macaws.

3.6.2. Species abundance

24
These species were relatively rare with not many individuals recorded over the study period.
The most abundant was Ara chloroptera with 18 individuals, and the least abundant was the
Ara ararauna. The abundance measures are illustrated in Table 11.

3.6.3. Species flight route.

A high percentage of these species flew in from zone 3, directly over the colpa to the forest
beyond. They did not land on the colpa and possibly went to the larger colpa at Posada
tourist lodge. Figures are illustrated in table 12.

Table 9. The abundance counts for the three large species of macaw. Included are
group sizes and individual numbers:

  Individuals Groups
Ara ararauna 10 3
Ara chloroptera 18 9
Ara macao 14 6

Table 10. This table shows the flight direction these large species of macaw flew.

Incoming Zone Groups %

1 1 8.3
2 0 0.0
3 12 66.7
4 4 33.3
5 1 8.3
Total 18 100

Outgoing Zone Groups %

1-5 4 22.2
6 14 77.8
Total 18 100

4. Discussion

4.1. Abundance, Incoming flight directions and Landing Locations

25
Exact population counts in avifauna are next to impossible due to the geographical range
of some species and the ease to which they move from one area to the next (MacArthur et
al 1962). The census methods can also produce unreliable results as mistakes in counts
are common and may be revealed by simultaneous observations by pairs of observers
(e.g. Spearpoint et al. 1988). By being a lone observer this source of error was
eliminated. A further important source of error for population estimates concerns the unit
of study. It is usually assumed that the group of species studied constitutes a single
population. In reality, populations are distributed heterogeneously in space and show
differences in behaviour that effect their ability to be detected by census workers (Bell et
al. 2006).

The birds started arriving at the site at roughly 05h55 in groups which consisted mostly
of Ara severa and Aratinga weddellii species. On an average morning 4 species and 111
individual birds were observed landing in the area with flights to the colpa tending to be
most frequent between 05h55 and 7h00. Flocks were exclusively monospecific except
when the birds were foraging in trees or eating soil at colpas. Markedly increases in flight
activity happened at 06h05 and 06h45 in every species except for Pionus menstruus. This
is illustrated in Figure 1. Most parrot species flew in groups of one to four individuals;
suggesting that mated pairs are stable and that family groups remain together post-
fledging. Most of the species (33%) flew in directly overhead from Zone 3. Presumably
the reason for seeing exceptionally small numbers of Brotogeris cyanoptera is because
they prefer to feed on colpas in early afternoon instead of early mornings (Gilardi &
Munn 1998).

Feeding at the site did not commence at once; but after a period of loud calls and possible
social interactions the first species started to land. The smaller species seemed more
accustomed to disturbance as these were always the first to feed, whereas it is known that
the larger species tend to be wary and feed less often (Hammer 2001).
Ara severa was the most abundant species recorded and the second largest species
seen at the site. Most of the species flew in from zone 3 and zone 4 (77%) which led to
the assumption that the species used the Tambopata River as a geographical feature to

26
find the colpa. This is also documented for other species of avifauna (Alerstam 1996;
Griffin 1969) and it is not unreasonable to suppose members of the Psittacidae family
undertake the same technique. Ara severa usually flew in groups of even numbers (X=
4.16) and were one of the first species groups to arrive. Most of the species landed
Aratinga weddellii is thought to have a large population as the species is
described as 'common' in at least parts of its range (Stotz et al. 1996). They were by far
the most vocal species recorded and were one of the first to land in the area each
morning. Their incoming flight directions were entirely random and usually arrived in
relatively large groups (X= 8.01) in which over half landed in the tall emergent tree in
zone 3. They seemed to wait in the trees until what appeared to be a critical number of
the same species were gathered, before moving onto the colpa. They were always the first
species to feed and suggesting that larger species possibly used Aratinga weddellii as a
type of indicator as whether it was safe to land on the colpa.
Pionus menstrus was the least abundant found at the site, with nearly all species
flying in from a westerly direction (96%). The species nests and roosts in un-flooded
primary forest (Sosa 2000), at approximate canopy height of 21m – 30m (Bjork & Powell
1995). This does not explain why the species come from this direction as the west suffers
from relatively high levels of disturbance due to the Trans-Amazonian highway not being
very far away to the north: Over half of the P. menstrus landed in zone 2.
Amazona farinose also had a higher number flying in from the west than would be
expected at random (89%). Again there is no plausible explanation for this as it prefers
nesting habitats such as gallery forest (Bjork 2000). 213 individuals were recorded and
the mean group size for this species were 3.38 individuals per flock.

However the numbers recorded in the surrounding area and vegetation were not the
actual number seen feeding. Data from a study conducted by another student on the
Inotawa colpa at the same time concentrated on the actual individual numbers feeding on
the clay. The mean number of individual birds in that study was 32 with a maximum and
minimum count of 72 and 0 respectively (Lovesey pers comm). Comparing it with the
data in this study, showed that only around 32% of all birds arriving at the colpa actually
fed. This suggests that social interactions at these congregations might be of more

27
importance to the birds than actual feeding. If this is the case, then disturbance by tourism
might not be as damaging to the species, as feeding at the sites may not be as important
as originally thought.

Studies into parrot behaviour at colpas have shown that large species of macaw behave
aggressively towards the smaller species (Maloney & Mclean 1995). This may explain
why there are large numbers of smaller avifauna species at this colpa as they are not
subjected to the aggressive behaviour of the larger species. The small surface area of the
lick probably displaced the larger macaw species to a larger site due to the risk of
predation.

It is relatively hard to compare macaw abundance measurements at this particular colpa

with other studies at the site as so many different biotic factors affect the feeding
numbers. The greatest factor affecting feeding numbers and species at the colpa is the
overall surface area of the clay accessible and the season (Brightsmith 2004). Previous
studies in the area (Brightsmith 2002) have shown that the numbers and species of birds
at the colpas change with the time of year. At the Tambopata Research Centre (TRC)
colpa use is busiest in the months of August and September in which over 350 birds can
be seen per day. In the month of May, this can drop to an average of 100 birds each
morning. December has the highest number of species seen each day (>9) while May still
has the least with about 5. Since this study took place in August it is reasonable to
assume that the data collected illustrates the colpa at probably the busiest time of year.

There have been many authors who have studied a range of colpa variables such as
topography, clay composition, weather and colpa surface area, and found significant
relationships in the Tambopata area between the composition of the clay and species
found at the site (Brightsmith 2004; Gilardi et al 1999; Burger & Gochfeld 2003),
whereas other studies in other areas of the world have found little evidence for the
mineral composition being the main reason for geophagy. (Hammer 2001; Nycander et
al 1995)

28
4.2 The large species of macaw.

Renton (2002) explains that larger species of macaws do not use small colpas due to the
risk of predation and hence this might be a factor in explaining the absence of the large
macaw species seen landing at the Inotawa colpa. It was possible that the birds were
flying to another colpa next to the Posada Amazonas lodge, where they have been
recorded before, and the data supported this. The larger species of macaw are very
nervous at colpas compared with the smaller species e.g. Aratinga weddellii. As
mentioned in the previous section, the large number of smaller species seen at this small
colpa might be explained by a lack of competition from the larger species.
4.3 Associations

In this study there were only two species that did not show significant associations:
Aratinga weddellii and Pionus Menstrus. Ara severa and Amanoza farinosa displayed
significant associations with all species suggesting a positive relationship between them
e.g. to aid predator response. Burger & Gochfeld. (2003) described the larger species of
macaw using the smaller species as a sort of predator indicators due to their quicker
responses to disturbance or noise. This could be called an association because the larger
species will not start to feed till there is a critical number of smaller species on the colpa.
However, a possible explanation for Pionus menstrus not having an association with
Aratinga Weddellii is that P. menstrus arrived slightly later in the morning and Aratinga
Weddellii being one of the first to arrive.

4.4 Weather

In line with previous studies on days with weather other than sun, or cloud resulted in
fewer recordings of parrots and macaws feeding (Hammer & Tatum-Hume 2005,
Brightsmith 2004). However, contrary to expectation, on one morning following bright
sunshine, no feeding was observed. This was attributed to other unidentified factors such
as unknown disturbances. Thus, although proving that weather is a factor in influencing
overall patterns, it appears that birds will not feed when they feel threatened, especially

29
the larger species. Mist had a far greater influence on the number of birds eventually
feeding. On day 7 heavy mist was encountered at sunrise, and the birds were observed
arriving at 06h49, a full 20 minutes after the sun had appeared through the clearing mist.
This might suggest that had the mist persisted then no species would have arrived at all
throughout the morning. This might be useful information for the surrounding eco-lodges
who might wish to cancel the colpa trip if mist was predicted.
The Kruskal-Wallis test was used with weather as the grouping variable to calculate
association between weather and the species (X² = 2.3, d.f = 4, p = 0.67) but the results
indicate that this was not significant. This might have been because the weather changed
from being very misty to sun in a space of 30 minutes after which many birds arrived.
This trend was not significant for the total number of species recorded.

4.5 Disturbance

As with many areas in the Amazon basin, increasing economic development is putting a
strain on the natural resources of Peru’s rainforest. Logging, farming and tourism are a
growing threat to its wildlife, particularly in the area west of the Tambopata River (pers
obs). Responses to intruders and predators varied by species of parrot/macaw and type of
intruder, but more importantly, disturbance by humans would have a negative effect on
feeding patterns and could result in the abandonment of the site if disturbance levels are
high and persistent (Mee 2005). Further, the increasing presence of humans and the
known whereabouts of the site might result in increasing effort to trap birds for the pet
trade. Munn (1992) has shown that large macaw species avoid areas of permanent human
settlement and his study revealed that fewer individuals were seen the closer one surveys
to the town of Puerto Maldonado. This suggests that mass tourism on small colpas would
almost certainly destroy natural behaviour patterns and may deprive parrots of a much-
needed resource.

At the site a variety of occurrences were described as disturbances since they had a
profound effect on the feeding on the colpa. Some were natural and some anthropogenic.
These include boat traffic, noise at the hide and natural predators, e.g. falcons. When a

30
flush was initiated, responses could be partial (some individuals flew away, circled, and
returned), temporary (all individuals flew away but returned within a few minutes), or
total (all flew away and abandoned feeding for at least a 15 minutes). The results show
that flushes were initiated by passing boats or noise from the hide. Flushes were twice
initiated when a falcon was present and the rest were due to unknown causes. The
macaws in particular used nearby trees before flying down to feed at the Inotawa colpa
and as a retreat when disturbed. The most frequent disturbances were caused by the boats
carrying tourists from the lodges arriving each morning. The birds were not disturbed by
the many commerce and trade boats due to the fact that they depart Puerto Maldonado in
the early morning, and therefore would only pass the colpa in the late afternoon or
evening, when no birds where present.

The first daily boat to arrive each morning was the Inotawa lodge boat, arriving at the
colpa early enough as not to create a disturbance (05h55 – 06h10 EST). However the boat
from Explorers Inn arrived considerably later (06h10 – 06h25 EST) to which the birds
responded with loud alarm calls. This delayed them from landing for another 20 minutes
or so, probably longer for the larger species of macaw and parrots on site. The noise from
the petrol motors attached to these boats and the movement of the tourists from the rivers
edge up to the hide in full view disturbed the birds greatly. This might explain why a
peak was shown on the activity graph at roughly 6h00, and then a drop until 06h45 as the
Inotawa boat had arrived but not the Explorers Inn boat. From personal observation, the
birds also seemed to be disturbed more by a boat with a peke-peke motor (a long-shafted
boat motor of either 10 or 16 horsepower) compared with a modern 25hp – 50hp
outboard motors. The peke-peke motors are used by the local La Torre tribe and the
increased disturbance caused by this particular motor might suggest they associate this
with predation i.e. still exploited in the area for food or feathers, or long term memory in
the birds.

When a flush was initiated, typically 80% - 90% left the area with an approximate 30%
returning to the area surrounding the colpa immediately. The species that returned in their

31
greatest numbers were Aratinga weddellii, and most commenced to feed again following
the disturbance. This was rarely recorded for the other species.

Over 60% of boats passing when birds were present caused birds to leave the vicinity of
the colpa. Birds that remained were often in the trees far back from the river’s edge.
Although not quantified in this study, there may be a correlation between size and the
probability of a flush, where smaller parakeets and parrots are less likely to flush
compared to larger parrots and macaws. In a few instances feeding activity has continued
with Aratinga weddellii and Pionus menstrus in the presence of boats, while this was
never recorded for the Amazona farinose, which is many times larger.

Conclusion

1 Concluding remarks

This study has shown that most of the species fly in each morning from a particular area
of rainforest. However, this direction might only be representative of the species at this
time of year.
In a recent study, McLoughlin (1970) explains that the diets of the birds may shift
with the seasons, meaning that it is possible that most of the members of a species may
temporarily leave the Tambopata Region and go elsewhere. Their incoming direction
might also change if some resources are scarce following land use change, which is not
improbable in the rainforest. This requires individuals of some species to either alter their
diets or change their movement patterns to utilize resources elsewhere (e.g. Galetti.1993;
Renton 2002). The study also shows that the birds were disturbed on a daily basis and
sometimes up to 3 times daily. This disturbance restricted the birds from feeding on the
clay for long periods of time. However, as only a certain percentage of the species were
actually recorded feeding at any one time, it may be that at this time of year, social
interactions at the site was also of great importance.

5.2 Improvements to the design of the study.

32
If this study were to be repeated, then improvements to the amount and to the accuracy of the
data would be obtained by conducting it over a longer time period. Also better preparation
prior to the study by thoroughly revising identification techniques for the avifauna of the
region. More emphasis may also be placed on the effect that human disturbance had on the
birds.

5.3 Management implications and recommendations

A number of papers present compelling evidence that certain species of birds, mammals
and amphibians are strongly influenced by the presence of tourists and hide/trail
infrastructure in their habitat (Kirkby et al 2000). There are a many recommendations that
could ease the anthropogenic pressure on these sites including 1) small scale tourism
conducted sensibly on colpas. This includes making sure that the observation points are
sufficiently far enough away, making it less likely to have a detrimental effect on the
avifauna. Another idea is to introduce a hide ‘carrying capacity’, where tourist numbers
at the hides are regulated. This ultimately reduces noise by reducing group sizes and
improving group management, through improved guide training. Another proposal is that
the visits to the colpa are timed better e.g. before 06h00; 2) continued wildlife monitoring
at these and other lodges, at least for the more easily identifiable species and more
especially those identified as impact indicators. This would indicate the status and health
of local avifauna populations and eventually improve encounter predictability in time and
space thus improving the number of times tourists can successfully observe wildlife; 3)
increased protection for threatened species and communities. Some of the vegetation
surrounding the lodges in the Tambopata area can harbour rare and endangered, and in an
area that has one of the highest diversities of avifauna in the world. Lodge administration
should set aside preferred habitats and colpas for vulnerable species that are off limits to
tourism, although one or two well managed sites with appropriate viewing facilities could
be implemented for those specialist tourists interested in observing rare bird species;
finally 4) Markedly improved guide training. Tour guides provided by the lodges should
have good knowledge of the general ecology at the vicinity of the lodge and increased

33
information concerning the home ranges of specific animals or groups nearest the lodges.
Personal experience at the Inotawa Lodge has suggested that the tour guides have very
limited knowledge and may not be appropriately trained being mostly from the local
town of Puerto Maldonado and not from the jungle. Efforts should be made to conserve
those sites most likely to have become damaged / disturbed by the presence of tourists.
Better prediction of diurnal and seasonal behaviour in order to increase encounter
frequency with minimal disturbance. Workshops should also be set up for the guides as it
is important that sufficient training is provided.

It may be suggested that studies of this nature will continue to be ever more important in
evaluating the impact of man on the natural environment. Particularly in the study area, to
find ways of managing tourism, which is important to the local economy in an
environmentally sensitive way. This can only be done by conducting the right research to
provide the information needed.

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