1 - Texas A&M University



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

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is entirely my own and not of any other person, unless explicitly acknowledged (including citation of published and unpublished sources).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

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Date

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Contents

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| |Contents |ii |

| |Contents |iii |

| |List of figures |iv |

| |List of tables |v |

| |Abstract |vi |

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|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 |

|1.4 |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 |

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|2.0 |AIMS |9 |

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|3.0 |STUDY AREA |10 |

|3.1 |Peru |10 |

|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 |

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|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 |

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|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 |

|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 |

|6.6 |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 |

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|7.0 |CONCLUSION |33 |

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|8.0 |Acknowledgements |34 |

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|9.0 |References |34 |

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|10.0 |Appendices |45 |

| |Appendix 1 - Example of data sheet used to record parrot abundances and flushes |45 |

| |Appendix 2 - General information about the parrot species recorded at the La Torre colpa |46 |

List of figures

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|Figure 1 |Map of Peru showing location of study site |10 |

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|Figure 2 |Satellite image of the Tambopata, La Torre colpa, Inotawa Lodge and Posada Amazonas. |11 |

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|Figure 3 |View of the La Torre colpa from the Inatowa Blind. |12 |

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|Figure 4 |Mean disturbance levels compared to mean parrot visitation numbers per day. |20 |

List of tables

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|Table 1 |Total number of individual birds observed feeding by species/day. |17 |

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|Table 2 |Mean number of individual birds observed feeding per day. |18 |

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|Table 3 |Species associations using Spearman’s rank correlation. |19 |

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|Table 4 |Species/disturbances associations calculated using Spearman’s rank correlation. |21 |

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|Table 5 |Mean and standard deviation values comparing general disturbance factors with tree and colpa flushes. |21 |

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|Table 6 |Mean and standard deviation values comparing tourist disturbance factors with tree and colpa flushes. |22 |

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|Table 7 |Mean and standard deviation values comparing boat disturbance with tree and colpa flushes. |23 |

:

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

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 1995-2000, 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).

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 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 (Iñigo-Elias 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 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.

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), red-masked 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 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 clay-consuming parrot species by enabling birds to consume previously unexploitable resources and/or increased quantities of seeds and fruits that would otherwise cause 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, scarlet-shoulded 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 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 land-use 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.

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.

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 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

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 five-minute 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 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

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 non-parametric correlation analysis for examining relationships between two variables. Each variable is ranked separately and comparison then takes place (Wheater et al 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 one-way 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, blue-headed parrot Pionus menstruus, orange-cheeked parrot Pionopsitta barrabandi, dusky-headed 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 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.

|Day No. |Red and |Chestnut |Mealy Parrot|Yellow |Blue Headed |Orange |Dusky Headed|White Eyed |Total: |

| |Green Macaw|Fronted | |Crowned |Parrot |Cheeked |Parakeet |Parakeet | |

| | |Macaw | |Parrot | |Parrot | | | |

|Boat |+0.26 p|+0.24 |-0.12 |+0.52** |-0.06 p|-0.30 p|+0.20 p|-0.45* |+0.15 |

|Disturbance |= 0.23 |p = 0.26 |p = 0.57 |p = 0.01 |= 0.79 |= 0.15 |= 0.35 |p = 0.03 |p = 0.47 |

|People |+0.35 |+0.21 p|-0.17 p|+0.44* |-0.15 p|-0.30 p|-0.07 p|-0.48* |-0.78 p|

|Disturbance |p = 0.10 |= 0.33 |= 0.43 |p = 0.03 |= 0.48 |= 0.16 |= 0.73 |p = 0.02 |= 0.71 |

|Animal |-0.16 |+0.11 p|+0.01 p|-0.07 p|+0.02 p|+0.11 p|-0.01 p|-0.17 p|+0.03 p|

|Disturbance |p = 0.47 |= 0.60 |= 0.10 |= 0.76 |= 0.94 |= 0.63 |= 0.97 |= 0.44 |= 0.90 |

Table 4: Species/disturbance associations calculated using the Spearman’s rank correlation (** = strongly associated * = associated).

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 = ................
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