A comparison of Aguaje Palm fruit consumption by terrestrial mammals, focusing on the three main habitats in the Lago Preto Concession Area



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Hypotheses


1.H1 There is a significant difference between Aguaje palm fruit consumption between frugivorous terrestrial mammals.

1.H0 There is not a significant difference between Aguaje palm fruit consumption between frugivorous terrestrial mammals.

2.H1 There is a significant difference between Aguaje fruit consumption between habitat types.

2.H0 There is not a significant difference between Aguaje fruit consumption between habitat types.


Study Sites


The three habitat types were located in the Lago Preto Conservation Concessions located on the Upper Yavari River, in Loreto, Peru. Lago Preto is located approximately 177.61 km away from the capital Iquitos. The study site is located on the Peruvian side of the river, near the mouth of the Yavari Mirin River. The site contains a high diversity of flora and fauna species. The concession is a 10,000-hectare block of tropical forest. The Lago Preto includes some of the most biodiverse habitats in the Amazon, and is divided into 3 main habitat groups; upland non-flooded terra firma forests, seasonally flooded varzea and palm swamps Aguajal. The temperatures vary from 30-40 degrees Celsius during the day with high humidity (Expedition Briefing 2007).

The Habitat Types: Terra Firma

Upland non-flooded terra firma forests are made up of undulating hills separated by streams, valleys and ravines. The canopy can reach up to 30 metres high as each flora species competes with the others for sunlight (Expedition briefing 2007). The under storey is not as dense as the other habitats as the sunlight is sparse amongst the under storeys. The ground does not hold the moisture as well as other habitats, and the water drains away quickly. Precipitation hardly penetrates the canopy due to the thick foliage. Out of the three habitats terra firma has the most diverse ecosystem.



Varzea


These habitats are flooded for up to 6 months a year, from the Yavari river water. These forest habitats are located parallel to the river, and are easily affected by increased precipitation. The average height of the canopy is a lot lower than that of terra firma. These forests have a lower floral diversity then that of terra firma (Expedition Briefing 2007). In areas the water will remain, long into the dry season.
Palm Swamps (Aguajal)

Palm swamps occur in areas between terra firma and Varzai. These habitats consist mainly of Palm species, meaning they have the lowest floral diversity of all the habitats. The understorey keeps water all year round, making it tough terrain for some species. However these areas provide numerous species with Aguaje fruit ranging from ants to tapirs. The canopy does not reach the great heights like the terra firma, providing light to the understorey. This allows the growth of young palms making the understorey dense.


Fig.6: Highlighted is the area of Loreto, Peru


Fig.7: The map shows the Lago Preto Conservation Concessions


Methods


In order to understand which of the terrestrial mammals feed upon the Aguaje fruit, a number of fruit baited tracking stations were placed in each of the three habitats from the 27/05/07 to the 14/06/07. Firstly 125 Aguaje palm fruits were collected from the Aguajal habitat (this was due to there being a large number of ripe fruits in this area). The fruits were located by looking for female Aguaje palms, and then searching the forest floor to find the suitable fruits that were ripe. It was essential that the fruits were ripe, squeezing the fruit slightly tested this. To begin with the tracking stations were made in the Varzea habitat. The area was selected approximately 5m away from the track, this made sure that the people utilising the tracks during the sampling period did not disturb the animals. Making the tracking station involved clearing a 1m2 area with a machete, and moistening the ground. Once damp enough the area was flattened with the edge of the machete. Then 5 Aguaje palm fruits were then neatly placed into the centre of the trap. A blue ribbon was tied onto a tree along the path so that the trap could be found the next day. Another 24 tracking stations were set up in the similar way, making sure that the traps were 60m apart from one another. Each of the traps was recorded in a book so, they could be easily found the next day. After all the traps had been made, 125 more Aguaje fruits were collected from Aguajal. The fruits were stored in a thick black bag and kept out of the sun. This was important, as the sunlight would continue to ripen the fruits, making them inuseable. The following day the traps were checked at around 0700 hours. Upon arrival of the traps the number of fruits taken, number of whole fruits and the number of fruits eaten onsite were recorded on a data sheet. If the fruits had been eaten or taken then the footprints were analysed using the “Neotropical Rainforest Mammals-A Field Guide” and the expertise of the field guide Anderson Cariajamo Ijuma. If the species could not be identified then a photograph was taken with the use of a scale for further analysis. Once all the footprints had been analysised then the trap was wipe clear and flattened with the machete and rebaited with 5 fresh Aguaje. This process continued for the next 24 traps. Once all the traps had been checked and rebaited then more fresh fruit was collected from Aguajal. The whole process was repeated every day at the same time over a 5-day period (23/05/07-31/05/07).

Towards the end of the sampling stage in Varzea, new traps were made in the terra firma habitat. This worked along the same principles as in Varzea. However only 20 traps were made due to the lack of moisture in the ground. All of the traps were cleared in the same way and baited in the same way. Once the sampling session had finished in Varzea, then the sampling began in Terra firma. In exactly the same way as before the traps were checked, species identified and then rebaited. The same process continued for a 5-day sampling period (03/06/07-07/06/07).

Towards the end of the sampling session in Terra firma, new traps were made in the Aguajal habitat. Again this worked along the same principles as in Terra firma, as only 20 traps were made, due to lack of ideal areas to form traps, and to much water. All the traps were cleared in the same way and baited the same. Once the sampling session had finished in Terra firma then the sampling began in Aguajal. In the same way the traps were checked, species identified and then rebaited for the 5 day sampling period (10/06/07-14/06/07). Once all the data had been collected then it was collated in to tables so that statistical analysis could be completed.

Results


To begin with a Shannon-Weaver index was completed to determine the variation of diversity between the three habitat sites.

H=-spI ln pI

The results showed that there was a greater diversity of terrestrial mammals in the Aguajal habitat (0.88) then Terra firma (0.74) and finally Varzea (0.43).

Varzea (all data found in appendix 5)


Of the 625 Aguaje fruits placed on the tracking stations 371 (59%) were taken away from the trap. Of that the identity of the fruit taker was known for 288 (78%). Of the Aguaje fruits placed on the tracking station 126 (20%) were consumed on site. Of that the identity of the fruit eater was known for 88 (70%). Spiny rat accounted for 255 (89%) of the fruit removed and 62 (70%) of the fruit eaten onsite. Agouti accounted for 5 (2%) of the fruit removed and 2 (2%) of the fruit eaten onsite. Armadillo accounted for 19 (7%) of fruits removed and did not eat any fruits onsite. Paca accounted for 5 (2%) of fruits removed and 4 (5%) of fruits eaten onsite. Ants accounted for none of the fruits removed from the site but ate 20 (23%) onsite. Birds accounted for 4 (1%) of the fruits removed and none of the fruits eaten onsite.
Terra firma (all data found in appendix 5)

Of the 500 Aguaje fruits placed on the tracking stations 331 (66%), were taken away from the trap. Of that the identity of the fruit taker was known for 226 (68%). Of the Aguaje fruits placed on the tracking station 54 (11%) were consumed onsite. Of that the identity of the fruit eater was known for 38 (70%). Spiny rat accounted for 179 (79%) of the fruit removed and 27 (71%) of the fruit eaten on site. Agouti accounted for 24 (10%) of the fruits removed and 8 (21%) of fruits eaten onsite. Armadillo accounted for 15 (7%) of the fruits taken and 1 (3%) of the fruits eaten onsite. Paca accounted for 8 (4%) of the fruit removed from the site and 2 (5%) of the fruit consumed onsite. There were no records for the ants and birds in terra firma.


Aguajal (all data found in appendix 5)

Of the 500 Aguaje fruits placed on the tracking stations 244 (49%), were taken away from the trap. Of that the identity of the fruit taker was known for 166 (68%). Of the Aguaje fruits placed on the tracking station 43 (9%) were consumed onsite. Of that the identity of the fruit eater was known for 35 (81%). Spiny rat accounted for 94 (56%) of the fruit removed and 32 (91%) of the fruit eaten on site. Agouti accounted for 61 (37%) of the fruits removed and 3 (9%) of fruits eaten onsite. Armadillo accounted for 1 (0.7%) of the fruits taken and none of the fruits eaten onsite. Paca didn’t account for any of the fruit removed from the site but did eat 10 (6%) of the fruit onsite. There were no records for the ants and birds in Aguajal.


Other then the species mentioned above, the only other species that visited the stations was an Ocelot. However no fruits were taken or touched therefore the visit was not included in the statistical analysis.
The Spiny rat was the primary fruit taker in all 3 habitats and the primary fruit eater in all three habitats, then Agouti, Armadillo, Paca, Ants and Birds respectively.


Testing the Hypotheses.

In order to see if there was a significant difference between the fruit consumption between the 3 habitat types, a two-way ANOVA was completed (see table 1). The data for the fruits removed and the fruits eaten onsite were collated.




Source Of Variation

Sum of Squares

d.f

Variance

F-Value

Tabled F-Value

Significance

P-Value

Between Samples

22235.78

11
















Variable A (Sites)

636.13

2

318.06

2.35

19.47

Not Significant




Variable B (Species)

17878.98

3

3959.66

44.05

26.36

Significant

0.01

Interaction

3720.66

6

620.11

4.58

3.77

Significant

0.05

Within Samples

64.94

48

135.29













Fig.9 shows the results of the ANOVA statistical analysis.
For the first hypothesis the null is rejected as the f-value (F 17878.,3 = 44.05, p= 0.01) exceeds that of the p-value meaning that there is a significant difference between Aguaje fruit consumption between terrestrial mammals.

For the second hypothesis the null is accepted, as the f-value (F 636,2 = 2.35, p=0.05) does not exceed that of the p-value meaning that there is not a significant difference between Aguaje fruit consumption between the habitat types.


Spiny rat Abundance’s

Location

Visitations

Sampling Time (Days)

Lago Preto Conservation Concessions

649

15

San Pedro, northeastern Peru

5362**

6*

Pacaya-Samiria National Reserve

Tamshiyacu-Tahuayo Community Reserve



93

34

Fig. 10 Spiny rat visitations * trapping **Estimated densities per km2 not visitations.



Fig. 11Shows the Aguaje fruit consumed on site and removed by the four main terrestrial mammals in the Terra firma habitat type.





Fig. 12 Shows the Aguaje fruit consumed and removed by the four main terrestrial mammals in the Varzea habitat type.



Fig. 13 Shows the Aguaje fruit consumed and removed by the four main terrestrial mammals in the Aguajal habitat type.


Discussion

Testing Hypotheses


The results of the ANOVA reject the null of the first hypothesis, meaning that there is a significant difference between the Aguaje fruit consumption between species. Looking at the data collected in the field it is obvious that the spiny rat consumed and removed the most fruit from the tracking stations. This could be related to a number of reasons. One reason could be that the spiny rat densities are higher than that of the other main terrestrial species. The high density could be related to the morphology and biological processes of the species. Histological experiments have highlighted that the male and females of the species remain sexually active throughout the year, therefore they are not seasonally dependent. This could explain a high density in the species if the reproduce all year round. In addition the species can produce 2-5 offspring at one time (Hon-Tsen & Yao-Sung 1999). Another reason for large densities could be related with the density and availability of the fruit. Studies in Panama showed a positive correlation between Spiny rats and the density of large fruited fig trees (Adler 2000). This could be similar to the relationship between the Spiny rat and the Aguaje Palm.
The results of the ANOVA accept the null of the second hypothesis, meaning that there is not a significant difference between Aguaje fruit consumption between the three habitats. This proves that there is not a habitat preference between the frugivorous terrestrial mammals. However the similarity between visitations between the three habitats highlight the importance of the Aguaje fruit in each of them. The fruit was eaten in each habitat, even though it only grows in flooded soils (Delgado et al 2007). Therefore a prediction would be that the visitations in terra firma would be significantly lower than in the other two. However this was not evident in the results, highlighting the importance of Aguaje fruit for frugivorous terrestrial mammals inhabiting each of the three habitats.
Diversity

The diversity of the three habitats was tested using the Shannon-Weaver diversity index, which compares the species richness of multiple areas (Fowler 1999). The number of species, which visited the fruit baited tracking station, was used for the analysis. The index highlighted that Aguajal had the highest species richness with 0.88, than Terra firma with 0.74 than finally Varzea with 0.43. One possible explanation of this result is the availability of food sources. Aguajal is abundant with palm species, which provide numerous terrestrial and arboreal species with food. The Aguaje palm is the most abundant within the Aguajals (Kahn 1990). The species is considered a keystone species because of the number of animals that need it for their existence, as it produces food through times of scarcity (Galetti & Aleixo 1998). The female palms can have up to 8 inflorescences per year producing 900 fruits at a time (Delgado et al 2007). A number of species compliment their diet with the Aguaje fruit. These are the Spiny rat (Prochimys sp.), Agouti (Dsayprocta fulignosa), Paca (Agouti paca), Lowland Tapir (Tapirus terrestris) and Collared Peccary (Tayassu tajacu) (Bodmer 1990 & Brooks et al 1997). Therefore is would be assumed that the diversity and richness of these species will be highest in the Aguajal habitat.


Agouti Visitations

The number of Agouti visitations was higher in the Aguajal habitat then in the other two, which could be related to the diet of the agouti. The agoutis feed primarily of seeds and fruits. One of the main fruits of their diets is Aguaje. Therefore this could be the reason why the visitations were higher in Aguajal.


Armadillo visitations.

There was a high frequency of armadillo visits, then previously recorded in other studies. Although the armadillo species can compliment it’s diet with Aguaje fruit, the possible reason for the high number of visits could be correlated with the number of ants present on the tracking station. Each of the stations that had a high number of ants over the course of the data collection received at least one visit from the armadillo. One possible explanation of this result could be related to the ripeness of the fruits. The fruits, which were used, were ripe on collection. However over the course of the day the fruit would continue to ripen, which attracted the ants. A large amount of ants may have attracted the armadillo, therefore producing a high result.


Lack of Visitations by larger bodied mammals.

One surprising finding was the lack of visitations by the larger bodied mammals like the Tapir. No visitations were recorded and no footprints were found around the tracking stations. One reason for this finding is that the larger bodied mammal has a larger home range, meaning that the forage furthers. Therefore they may cover larger areas over the course of a day and not necessarily remain in one area. Studies of the Bairds Tapir of Costa Rica proved that the monthly home range of the species was approximately 55 hectares (Foerster 2002). Even though there wasn’t any visitations to the fruit baited tracking stations during the field study, this does not mean that the Aguaje palm is not important for the larger bodied terrestrial mammals. Stomach samples have shown percentage of Aguaje consumed by larger mammals. The lowland tapir stomach sample included 76% of Aguaje fruit, the white lipped peccary had 45% Aguaje fruit, the collared peccary had 12% and the red brocket deer had 9% (Bodmer 1989). This highlights the importance of the fruit within the mammal’s diet in particular the lowland tapir. Another reason for the lack of visitations could be related to the reduction in numbers across the main species, which could be through over hunting, this is unlikely as the field study occurred in a protected area.


Lower visitations in Aguajal.

The visits by all the species were a lot lower in Aguajal than in the other two habitats. The reason for this could stem from the food availability within the area. Aguajals primarily consist of the Aguaje palm, with a small frequency of other palms, therefore the Aguaje fruit is available throughout the habitat. Therefore the fruit placed on the fruit baited tracking stations isn’t essential to the species, as they can easily forage the fruit themselves. With the high presence of palms, the less likely the animals will target and visit the tracks. However in other habitats like terra firm the Aguaje palm isn’t present, therefore opportunist feeders will scavenge the fruit from the tracking stations. This again highlights the huge presence of Aguaje within the Aguajal habitat.


The spiny rat

The results show that the Spiny rats ate and removed the most Aguaje fruit from the fruit baited tracking stations. One possible reason for the increased visitation by the spiny rat could be related to its size and home range. Due to their small bodies the spiny rat’s home range will be significantly smaller than that of a larger bodied animal. Meaning that the species only travels short distances and remains in small areas (Russo 2005). The spiny rat species is also considered scatter-hoarders (Russo 2005), which means the seeds are removed and buried for times of food scarcity. This would explain the high percentage of fruit removal from the three habitats. In terra firma 79% of all fruits removed from the habitat was from the spiny rat, which is considerably higher than any other species. High fruit removals by the species was also recorded in the other two habitats where 89% was removed in varzea and 56% in Aguajal (data derived from appendix 4). These percentages are higher than the percentage of fruit eaten on site thus further confirming the scatter hoarding behaviour by the species. Another reason could be related to the slight fluctuation in resources available to the mammals as the reproductive rate of the spiny rat is positively correlated with that of food availability. During seasons of resource scarcity the density of the spin rat is reduced, due to unavailability of food to support the young. During seasons of resource abundance the populations increase, as the reproductive rate increases (Adler 1998). Therefore densities are at their greatest at the end of the season with resource abundance, which could be related to results of the study. The data was collected at the end of the rainy season, which could be considered the resource abundance season. Therefore the data was collected when spiny rat densities were at their greatest, which is representative of the results.


A comparison of spiny rat densities of two other studies.

The results collected highlight the high density of spiny rat in the Lago Preto conservation concession. A thesis by Michael Valqui highlighted the high density of spiny rats in San Pedro northeastern Peru. By using traps, Valqui estimated the density per km2 at 1508 in terra firma and 3854 in varzea (Valqui 2001). The large density estimation could be linked to the use of traps. Using traps instead of fruit baited tracking stations may give a better estimation to the correct densities of the spiny rat.

However research by Petra Mikkolainen showed a low density of spiny rats in the Pacaya-Samiria National Reserve and Tamshiyacu-Tahuayo Community Reserve, Peru. Mikkolainen’s combined results for the spiny rat were a lot lower, with only 93 spiny rat visits being recorded in all three habitats. The research also showed that the agouti visited the tracking stations more than the spiny rat (Mikkolainen 2004).

The co-existence of frugivores and the Aguaje palm.

Frugivorous species and fruit producing species like the Aguaje palm depend on each other for their species survival. If one were to become extinct than the other would be greatly effected. The study has highlighted the dependence of the spiny rat, agouti and paca on the Aguaje palm. If the population of the palm species were to alter, than the spiny rat would be the most greatly affected.

Frugivores disperse a number of seeds through their digestion, therefore dispersing the seeds. This allows the fruiting species to remain abundant throughout their habitat. This mutualistic relationship also benefits the frugivorous species, by providing them with food. The Aguaje palm is of great importance for the frugivorous community, as it produces fruit through times of scarcity (Galetti & Aleixo 1998).

The spiny rat has also been associated with the dispersal of vesicular-arbuscular mycorrhizal fungi. This fungus enhances the uptake of mineral nutrients for plants. VAM increases the rate of survival for the plant, as it improves the uptake of phosphorus from the soil (Janos 1995). An increase in the dispersal of the fungus leads to plants becoming stronger and taller, which increase competition within the floral community. Therefore the dispersal of the fungus by the spiny rat increases the diversity of the rainforest (Janos 1995).


The effects of an unsustainable harvest of the Aguaje palm.

From the results it is clear to see that the Spiny Rat consumed the highest amount of Aguaje fruits, then the Agouti, Armadillo, Paca, Ants and birds respectively. This highlights the fact that Aguaje is an essential part of the frugivorous terrestrial mammal’s diets. The results indicate which species will be greatly affected by an unsustainable harvest of the Aguaje Palm tree. The rodent species will be worst affected, as the species are relatively small bodied therefore only having a small home range. Therefore the smallest of harvests for example the removal of one tree could have a devastating affect on the surrounding Spiny Rat individuals. The slightly larger bodied animals for example the tapir would not be as directly affected because their home ranges cover a vast amount of forests (Foerster 2002). This does not suggest that the larger bodied terrestrial mammals will not be affected, as they eventually will become threatened if the Aguaje Palm fruit harvesting does not become sustainable. Although none of the tracking stations were visited by tapirs, it does not imply that they do not depend on the Aguaje palm fruit as it has been proven that the tapir’s diet consists of 76% of Aguaje fruit (Bodmer 1989).

The Iquitos market sells approximately 193 fruit species, and of those 120 are wild harvested. With approximately 57 species belonging to 24 different plant species. These species are important to the economy and the diets of the local people. However some of these species are beginning to be grown as crops, however the destructive methods of collecting the wild species are leading to the depletion in resources (Vasquez 1989).

The current destructive harvesting techniques will eventually lead to the destruction of the palm communities amongst the neo-tropic regions. Eventually this will lead to the reduction of the ecosystem’s carrying capacity, lowering the abundance of the some of the most vital species. The effect of an unsustainable harvest will affect numerous species within the community. The spiny rat not only disperses the seeds of the Aguaje fruit, but also the VAM fungus that is essential to plant growth (Janos 1995). Therefore if the Aguaje palm reduced in numbers then so would the frugivorous mammals and the plant species that have their seeds dispersed. However more studies need to be completed in order to determine a better understanding of the relationship between palms and frugivores. The mutualistic relationship between frugivore and plant has been identified in previous studies. One research project looked at the effect of poaching on seed dispersal. By using a number of methods the conclusions showed that there was a direct link between poaching and the reduction in seedling densities (Wright 2000). This is one example of what may occur if the Aguaje palm numbers were to be reduced significantly.

The harvesting of the Aguaje palm fruit is conducted by felling the entire tree. To avoid the loss of a good fruit source to a rival, Aguaje harvesters will locate a tree and fell it whilst the fruits are still ripening. This unripe state is a preferred choice by some of the terrestrial mammals; therefore the harvesting directly affects them. Due to the years of harvesting the suitable palms is becoming more and more difficult as the palms are becoming further away from human settlements. The selling of the fruit on the market whilst in its unripe state does not produce much income, therefore neither the environment nor the local community benefit from the harvest. (Penn & Neise 2008a).


Possible solutions

One way to reduce the impact of an unsustainable harvest on the Aguaje palm is by the use of Agroforestry. Planting Aguaje palms in a field evenly spaced will stunt the growth of the trunk, therefore allowing the fruits to be harvested sustainably. The Aguaje palm grows fairly quickly (maturing at 12-18 years). The fruit bunches hang as low as 2 metres off the ground meaning that the harvester doesn’t need to climb the trunk to collect the fruit. Agroforestry provides an economical incentive to the local communities as it allows the fruits to ripen before being harvested. This means that the fruit will have a greater economic value at the local markets (Penn & Neise 2008a).


Problems in the design of the research collection.

There were a few issues with the design of the methodology. Firstly there were a number of multiple visitations. This reduced the amount of data that could be used for the statistical analysis. One way to reduce this would be to check the stations twice a day. This would highlight the visitations by nocturnal and diurnal species, and reduce the amount of multiple visits.

Another issue of the project was the uneven amount of aguaje fruit between the three habitats. in varzea 625 fruits were placed on 25 stations, however only 500 over 20 stations were in terra firma and aguajal. This was due to the tough terrain and lack of moisture (within terra firma) therefore a reduced amount of stations were made. However all the results are proportionate of the overall amount, so it didn’t skew the data.

Another issue of the project was the lack of knowledge and resources of the footprints of terrestrial mammals in the rainforest. There was only one book that was used to identify the particular species. More sources of information may have lead to an increase success rate in identification of the species.


Further studies.

This research project has highlighted the need for spiny rat densities to be monitored in the Lago Preto conservation concession. It is also essential that the numbers of frugivores are monitored so if a reduction in numbers were to occur than a management plan can be put in place as soon as possible, to make sure the numbers don’t reduce any further. More studies into the palm harvesting techniques, to enable alternative methods of collection so that the harvest is no longer unsustainable. Finally education of the local people about their impacts on the rainforest, may provide an insight into community based management programmes.



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Figure3: Spiny Rat

www.mammalogy.org/mil_images/images/mid/1584.jpg
Figure 4: Paca http://animaldiversity.ummz.umich.edu/site/resources/mzm2/82.mr2.jpg/medium.jpg
Figure 5: Agouti:

http://magazine.naturspot.de/pict/agouti.gif



Appendix 1


The total number of fruits taken or eaten on site by each species in the Terra Firma habitat





Site Terra Firma



















Day 1

Day2

Day3

Day4

Day5

Total

Mean

SD

Spiny Rat

53

34

38

47

32

204

40.8

8.927485648

Agouti

0

10

1

8

12

31

6.2

5.403702434

Armadillo

0

0

8

3

5

16

3.2

3.420526275

Paca

0

0

1

9

0

10

2

3.937003937

Appendix 2


The total number of fruits taken or eaten on site by each species in the Varzea habitat.




Site Varzea






















Day1

Day2

Day3

Day4

Day5

Total

Mean

SD

Spiny Rat

49

81

70

73

50

323

64.6

14.36314729

Agouti

0

10

1

8

12

31

6.2

5.403702434

Armadillo

0

0

8

3

5

16

3.2

3.420526275

Paca

0

0

1

9

0

10

2

3.937003937



Appendix 3


The total number of fruits taken or eaten on site by each species in the Aguajal habitat.






Site Aguajal






















Day1

Day2

Day3

Day4

Day5

Total

Mean

SD

Spiny Rat

16

25

17

43

23

121

24.8

10.87198234

Agouti

18

8

12

16

10

64

12.8

4.147288271

Armadillo

3

0

0

0

0

3

0.6

1.341640786

Paca

0

5

5

0

0

10

2

2.738612788



Appendix 4


FR= Fruit removed from site FEO= Fruit eaten onsite

The proportion of fruit eaten of taken from each species related to identification of fruit taker/consumer







Spiny Rat




Agouti




Armadillo

Paca




Ants




Birds







F R

F E O

F R

F E O

F R

F E O

F R

F E O

F R

F E O

F R

F E O

Terra firma

179 (79%)

27 (71%)

24 (10%)

8 (21%)

15 (7%)

1 (2.6%)

8 (4%)

2 (5%)

0

0

0

0

Varzea

255 (89%)

62 (70%)

5 (2%)

22 (2%)

19 (7%)

0

5 (1.7%)

4 (5%)

0

20 (23%)

4 (1%)

0

Aguajal

94 (56%)

32 (91%)

61 (37%)

3 (9%)

10 (0.6%)

0

10 (6%)

0

0

0

0

0

Appendix 5





Site

No. Of Fruits

No. Of Fruits Taken

Identity of Fruit Taker

No. Of Fruits Eaten Onsite

Id of Fruit Eater

Terra Firma

500

331

226 (68.2%)

54

38 (70.4%)

Varzea

625

371

288 (77.6%)

126

88 (69.8%)

Aguajal

500

244

166 (68.0%)

43

35 (81.3%)







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