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Semi-annual Report

January-June 2018

LONG-TERM MONITORING PROJECT OF


BIRDS IN THE SOUTHERN AREA OF
TORTUGUERO NATIONAL PARK, COSTA
RICA

PRODUCED BY:

Thijs Claes – Avian Project Manager -2018– Present

IN COLLABORATION WITH:

FIELD COORDINATOR: Cormac Healy


FIELD STAFF: Danny Guy, Victoria Hawkins, Stephany Butera, Allen Bush Beaupré, Amber
Searcy, Daniel Sturgess, Emily Underhill

GVI COSTA RICA COUNTRY DIRECTOR: Cynthia Arochi

JALOVA, COSTA RICA


2018

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About this Report

Global Vision International has conducted this research. The findings and recommendations of
this report are based on the statistical analysis of the data collected in the context of the long-
term monitoring effort (2014-2018) with special attention to the period of 7th January – 7th June
2018.

Cited as
Claes, T., 2018. Semi-annual Report (January-June): Long-term monitoring project of birds in
the southern area of Tortuguero National Park, Costa Rica.

Acknowledgements:
This research has been conducted with the help of the following Global Vision International
volunteers:

Bandon Dembs, Corne Slemmer, Zoe McCallum, Jack Glover, Baley Good, Eoin Donohoe,
Jurgen Vincke, Alice Hirons, Emma Wagner, Danielle Small, Peter Lichtenthal, Timothy
Carlson, Brandon Dembs, Zarah Ehlin, Eleanor Green, Emma Brown, Diane Leishman, Amber
Searcy, Rebecca Barber, Tess Nelson, Emily Larmer, Freya Savage, George Buchanan, Amanda
Maretto, Jenna Scherger, Marlena Ryzyk, Emiel Zaandam, Daniela Campama, Lara Lehnen,
Kevin Goulart, Joris Giglio, Ethel Mery, Connor Dupree-Sood, Amanda Schultz, Alice
Douvillez, Tom Panneman, Kieran Bose Rosling, Alex Bartlett, Matt Smith, Elijah Denham,
Michelle Welch, Selina Mellin, Simon Thel, Brian Milne, Willem Van Doorninck, Krijn
Bresser, Matthew Webb, Madison McLatchie, Rachel Hetherington, Mara Ramirez, Lauren
Chappel, Ben Cozens, Alice Palmer, Christine Davison, Erin Gleason, Rob Harris, Ira Epstein,
Lucy Shephard, Leah Cole, Andrew Ujifusa, Jennifer Ujifusa, Christina Coyle, Sofia Hedman,
Karin Jermer, Spencer Kielar, Anna Westwood, Bethany Ball, Alex Willis, Zoe Mathurin,
Marcin Oksinski, Martin Schamberger

GVI Jalova, Costa Rica


E-mail: tortuguero@gviworld.com
Web page: http://www.gviworld.com

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Summary
Tortuguero National Park (TNP) is an important habitat for breeding, wintering and migrating
waterbirds in Costa Rica. The park is also an ecotourism hotspot and the canals within act as
main routes to Tortuguero village. GVI Jalova performed a long-term monitoring project to
evaluate the effects of boat traffic frequency and tourism on the rich avian diversity. After an
initial development phase (2010-2013), the protocols have remained constant. Since 2014, a
gradually more detailed and precise long-term analysis of the avifauna within the study area is
available. Differences in bird biodiversity, abundance and richness among the canals were
identified. The wide canals are particularly important to support large populations of migratory
species and common resident species within the study area. The narrow canals are a haven for
many less abundant species and are extremely important to sustain the high regional
biodiversity.

This study period, 53 surveys (approximately 1 hour) were conducted on 5 different canals
recording the bird community from a canoe. In total, 1957 individuals belonging to 95 bird
species were recorded. Comparable to previous results, the Shannon-Wiener index for TNP was
calculated to be 3.358 with an evenness of 0.737 considering all species on all canals for this
study period. All common target species, apart from the migratory ones, appear to have stable
populations within the park.

For the first time, several vegetation structures on the canals have been quantified. Large
variations in the vegetation structures among and within the canals are observed. Motorized boat
traffic passing during the surveys are shown to significantly increase the abundance of birds
identified during a section.

This report concludes with stressing the necessity of consistent methodologies, identifying areas
of interest for further studies and practical suggestions to enhance the management of TNP.

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Contents
Summary .................................................................................................................................................. 3
1. Introduction ...................................................................................................................................... 5
1.1 General and specific objectives ................................................................................................ 6
1.2 Duration of the project ............................................................................................................ 7
2. Methodology ........................................................................................................................................ 7
2.1 Study Area...................................................................................................................................... 7
2.2 Field Survey Method ...................................................................................................................... 7
2.3 Data Analysis ................................................................................................................................. 8
3. Results and Discusion .......................................................................................................................... 9
3.1 Avifauna from the canals in Tortuguero National Park. ................................................................ 9
3.2 Abundance, Richness and Diversity per Canal ............................................................................ 15
3.3 Abundance, Composition and Frequency of Occurrence of target species per canal .................. 20
3.4 Boat Traffic .................................................................................................................................. 29
4. Recommendations .............................................................................................................................. 37
4.1 Continuity of the project .............................................................................................................. 37
4.2 Recommendations to the management per canal ......................................................................... 39
5. Communication strategy and use of the results .................................................................................. 41
Reference List ........................................................................................................................................ 41
Attachments............................................................................................................................................ 45

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1. Introduction
Costa Rica is one of the most biodiverse locations in the world. Precipitation and temperatures
are impacted in the coastal lowlands by the inland mountain ranges they surround. This
geographical diversity provides the perfect conditions for a variety of vegetation to thrive, which
in turn can support a multitude of mammalian, reptile, amphibian, and avian species (Stiles and
Skutch 1989; Garrigues and Dean 2014). Approximately 903 bird species, representing 82
families, can be found in Costa Rica. This includes resident, migratory and those species classed
as “rarities” (Garrigues and Dean 2014). In spite of its small landmass size Costa Rica is host to
around 4% of all know species (Valerio 2006).

Birds in general have been widely used as bio-indicators of environmental change (Lower and
Kendall 1992; Kushlan 1993). Kushlan (1993) reviewed the use of colonial water birds as bio-
indicators; the results supported the use of bird behaviour, reproductive performance, presence,
absence, distribution, eggshell quality and other measurable characteristics as suitable factors
for assessing wider ecological conditions. Other studies have also shown that food availability,
hydrological variability, and vegetation composition of habitats can affect bird populations,
providing further evidence supporting the use of birds for the monitoring of biodiversity and
ecosystem health (Kushlan 1993; Alava and Haase 2011; Canepuccia et al. 2007).

In the last century there has been a 50% decline of wetlands habitats across the globe due to
human activities (Ma et al. 2010). This is an alarming fact, as well-managed wetlands can
provide alternative habitats for waterbirds such as feeding, breeding and resting grounds (Bibi
and Ali 2013; Kushlan 1993). In the early 1970’s Tortuguero National Park (TNP) located on
the North-Eastern Caribbean coast, was one of the first protected areas to be established in Costa
Rica (Boza 1993); TNP is now listed as a Wetland of International Importance (Ramsar site)
and an Important Bird and Biodiversity Area (IBA) (BirdLife International 2015; RAMSAR
2010). The park boasts 24 out of 350 wetlands found in Costa Rica (Alvarado Quesada 2006).

Although water birds in Costa Rica can occur in a wide range of humidity’s and elevations they
are typically specialist in terms of aquatic habitat type. Behind the Caribbean coast’s black sand
beaches and swamp forests lies a series of inland waterways connected by artificial canals, from
North Limón to Nicaragua (Stiles and Skutch 1989). The wetlands found within TNP are
essential for water birds in terms of breeding, wintering and stopover sites (Alvarado Quesada
2006). Considering that 19% (167 species, representing 26 families) of the avian species in
Costa Rica are water birds, many of which are seasonal migrants travelling from North America
(Alvarado Quesada 2006; Garrigues and Dean 2014) these waterways prove to be of vital
importance for not only local, but also regional avifauna. For the overall conservation scheme
of the habitats used by these migratory birds, the monitoring and conservation of the multiple
habitats used during their migration is of vital importance. Especially given that these species

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rely on multiple sites as habitats throughout the year (Sodhi et al. 2011). Furthermore, the North
American Waterbird Conservation Plan of 2002 noted 12 waterbird species as High Concern,
and in Costa Rica 6 species were listed as ‘at risk’ in 2006 (Kushlan et al. 2002; Alvarado
Quesada 2006).

There has been little research into the waterbirds of TNP, especially in a long-term context.
Groom (2011) listed 265 species of birds, which included the Agami Heron (Agamia agami), a
species listed as vulnerable by the IUCN (International Union for the Conservation of Nature)
redlist. Previous reports by GVI include those by Coupland et al. (2013), Barreto et al. (2014),
Vaca (2016) and Claes (2017a, 2017b) who respectively reported 81, 68, 64, 93 and 84 species
on the canals of TNP in a six month period. In total, 166 species have been recorded on the
canals (Apendix A). Other studies have also assessed the vegetation composition in the
Tortuguero area, such as that by Myers (1990) and Lewis et al. (2010) in Caño Palma Biological
Station. In the study area of this project however no such study has been completed before. The
vegetation structure of an ecosystem can directly affect the abundance and richness of bird
species (Bibi and Ali, 2013, Kushlan 1993). The number of habitats, or habitat heterogeneity, is
positively correlated with species richness as well (Gonzalez 2009). The introduction of plant
species can cause loss or alteration of habitats, resulting in changes to community structure as
well as threaten native species of birds (Alava and Haase 2006). Understanding community
composition of both flora and fauna can improve the ability to compare site quality over time
(Cohn-Haft et al. 1997). Moreover, changes in avian diversity need to be well understood for
adequate ecosystem management (Bibi and Ali 2013).

Because of its attractive biodiversity, Costa Rica is a well-known destination for ecotourism.
This is one of the biggest industries financially supporting many of the country’s economy.
Costa Rica politically supports protected area agencies and conservation on public and private
land (Buckley 2011). Maintaining Costa Rica’s rich biodiversity is becoming increasingly
important because hospitality to tourists and human pressures by expansion of agricultural lands
increases. Protected areas are thus more crucial than ever for the persistence of Costa Rica’s
rich biodiversity (Stiles and Skutch 1989). The management of such areas can only be
undertaken appropriately supported by long-term monitoring and assessments of both the
environment and its inhabitants.

1.1 General and specific objectives


The general objective of this project is to contemplate a long-term data base about the birds in
the canals in the southern part of Tortuguero National Park (TNP). This semi-annual report aims
to summarize the most recent findings in the context of the long-term trends. In addition, it aims
to assist the Ministry of the Environment and Energy of Costa Rica (MINAE) by providing a

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complete species list of the birds found in the southern part of TNP, with special attention to 30
target species.

The specific objective is to perform a valuable analysis of the collected data to allow
comparisons with previous results and conjure recommendations to improve the management
of Tortuguero National Park for the aquatic bird communities. Recommendations regarding boat
traffic within the canals and the health of the aquatic ecosystems, including fish communities,
vegetation structure and water quality are also included for reference as applicable to the results
obtained in this study.

1.2 Duration of the project

Global Vision International (GVI) began the Monitoring Project for the Canal Birds of
Tortuguero National Park in 2005. In 2010 the project was relocated to its current location, in
the Jalova Biological Research Station in the most southern area of TNP. Methodologies have
since been updated accordingly to monitor general bird diversity, richness, abundance and
habitat use over the long-term course of the program. Up to date, the methodologies of the
project have not been changed since 2014. No definite end of the project is set since constant
monitoring is required to allow adequate recommendations for the management of the national
park. The project is evaluated and the results and their analysis summarized every six months.

2. Methodology
Any modifications have been made in compliance with GVI Costa Rica Expedition’s
requirements. The following methodologies are very similar to those stated in Barreto et al.
(2014) and Levac (2015), with minor adjustments.

2.1 Study Area


The study area remained identical since January 2017. For a detailed description see Claes
(2017a).

2.2 Field Survey Method


A list of 30 target species from both resident and migratory populations was compiled according
to the species’ preferences for both feeding and breeding habitats (Apendix B). Together they
represent the total variation in the vegetation structure of the ecosystem particularly well. Data
was collected on abundance, richness, and when possible information on sex and life stage
(juvenile, adult or breeding phase) were recorded. All staff and volunteers were trained before

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taking part in any survey to ensure all possible information on target species could be recorded
with 100% accuracy. Since January 2017 the use of a digital camera has been encouraged to
identify species other than the target species. The weather during each survey is categorized into
different gradations of clear, cloudy and rainy conditions. Motorized boats passing the canoe
during surveys were also recorded together with the number of people on the boat, the purpose
of the boat (Tourism, Rangers, Local, and Commerce) and the power of the engine of the boat.

The surveys are conducted by 5 or 6 qualified observers in a canoe at a speed of 2 km/h lasting
approximately one hour. Moving transects were used rather than a point count methodology
considering that many of the target species are relatively stationary. Species identified solely by
vocalizations were also only recorded when believed to be within 25m of the canal’s edge.
Individuals flying above the canopy line were not recorded.

All canals, divided into 4 section of 500 meters, are aimed to be monitored twice times per
month. Once in both the morning (06:00-07:00) and afternoon (15:00-16:00). Sporadic
nocturnal surveys (20:00-21:00) are also performed on California (CAL), Caño Negro (CN),
Laguna Jalova (LJ). Given the presence of American Crocodiles (Crocodylus acutus) along the
canals, the methodology had to be adjusted for nocturnal surveys. A motorized boat with 115
HP was used instead of a canoe (the same speed was maintained). We also use one light on each
site of the boat and a strong spotlight in order to search and correctly identify each of the
encountered individuals. To minimise our impact on the night vision of the species encountered,
the spotlight only remains on any individual for a maximum of 10 seconds. Nocturnal surveys
on Rio Sierpe (RS) and Rio Sierpe Alto (RSA) are not achievable because of safety and logistical
reasons.

Since 2018, the vegetation structure on all the canals is also monitored once every three. For
every section of 500 meters, the amount of individual palms, big and small pieces of floating
debris are counted. Also the amount of floating vegetation, emergent vegetation, clear bank and
the width of the canal our estimated using the length of the canoe as a reference. This
information is collected following bird surveys in the morning.

Several point count surveys have also been performed this study period according to the protocol
of the PROALAS project (Programa de América Latina para Aves Silvestres).

2.3 Data Analysis


Microsoft Excel Software (2013) was used to create all the graphs and dynamic pivot tables
were used to summarize all the data. Shannon-Wiener Biodiversity Indices were calculated for
each survey and an average was calculated based on both the study periods and the canals. For
all target species the Frequency of Occurrences (FO) are calculated by dividing the number of
surveys on which the species has been seen divided by the amount of surveys on each canal as
first described by Bensizerara et al. (2013). An index varying from ‘Occasional’ to ‘Abundant’
is used to categorize the target species in five categories (Apendix C). To visualize the long-

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term trends of the FO, the different classifications were scored from 0 (Absent) to 5 (abundant).
The scores from every species in each canal, calculated from table like the one in Apendix D,
are then summed up to visualize the general presence of target species

Minitab 17.1.0 was used to perform statistical analysis and create models. The long-term trends
in average bird abundance per survey were analysed with linear regression model. The influence
of the canals, weather type and their interaction on the bird abundance data was analysed with
a general linear model after normalizing the data with a log-transformation. A Spearman Rho
correlation was used to analyse the relationship between floating and emergent vegetation. The
impact of motorized boat traffic on the abundance of target species seen on a section was
analysed with a paired t-test.

None of the information collected during the nocturnal survey are included in the analysis since
these would bias the data and are therefore discussed separately. All other information about the
avifauna in this report is collected during surveys following the standardized protocol, unless
mentioned otherwise.

3. Results and Discussion


3.1 Avian biodiversity

3.1.1 Regional

This study period, 53 surveys were conducted on the 5 canals (CAL=11, CN=11, LJ=11, RS=10,
RSA 10), including one night survey on CAL, CN and LJ. In total, 1957 individuals belonging to
95 species were recorded, of which 25 are target species. The Shannon-Wiener index for TNP for
this study period is 3.358 with an Evenness of 0.737 considering all species on all canals. The
Average Shannon-Wiener index and Average Evenness per survey are respectively 2.090 and
0.855 (Figure 1). The more intensive use of digital cameras during the surveys is reflected by the
increase in biodiversity since 2017a. The average species richness per survey follows an almost
identical pattern. Compensating for the increase use of digital cameras, the avian biodiversity in
the southern part of TNP appears to have remained constant since 2014. This is an encouraging
sign in the context of the current threats to tropical avian biodiversity. It appears that in the recent
past the management of TNP had the potential to protect the avifauna from causes of these threats
such as climate change, habitat fragmentation, habitat degradation and pollution. Actions can
nevertheless be taken to improve the protection of the habitat TNP provides for its inhabitants.

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Long-term trends of average biodiversity per survey
2.5

1.5

0.5

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months
Shannon-Wiener Index Evenness

Figure 1: Long-term trends of average Shannon-Wiener Index and average Evenness per survey for each study
period in Tortuguero National Park, Limón, Costa Rica (a=Januari-June, b=Juli-December).

With the exception of the year 2015, the surveys in the first study period of the year always result
in a higher biodiversity than the once in the second study period. This subtle alternating pattern
in the Shannon-Wiener Biodiversity Index could result from multiple factors. The consistently
lower sampling effort in December is one of them. This is partly compensated by a slightly lower
sampling effort in January as well however. Another explanation could be that the breeding period
for many species in TNP occurs from January until June, increasing their activity and detectability
on surveys. The breeding periods of the target species, which constitute more 68% of the total
amount of individuals recorded since 2015, are fairly evenly distributed throughout the year
however. Also percentage of records of breeding adult and juveniles is not markedly higher during
the first period of the year. A difference in weather pattern between study periods in the beginning
of the year compared to the end could be another explanation. Rain both decreases the activity of
birds and reduces our visibility, resulting in a decrease of the logarithm of the abundance, and
therefore very likely also the biodiversity of the survey (Figure 2). In general, rain was more
likely on surveys in the second study period of each year (Table 1). The fact that biodiversity was
higher in 2015b strengthens this hypothesis, since the typical weather pattern was reversed in
2015. The considerably lower amount of surveys performed in 2015a compared to 2015b, are
another undoubtable explanation for its low biodiversity index (Table 1). The influence of
migratory species, which would have an important influence on the biodiversity of the avifauna,
is evenly divided between different study periods. Although further analysis should confirm this,
the migration season is therefore not expected to create the alternating pattern. A change in arrival
and departure times of migratory species, as a response to climate change, could however result
in a pronounced pattern in the future however. A more detailed, monthly analysis would be
valuable to identify the importance of breeding and migration seasons. To date it is not possible
to perform enough surveys each month for such a detailed analysis.

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Influence of Canal, Weather type and their interaction
on the bird Abundance
1.8
1.7
Log(Abundance)

1.6
1.5
No Rain
1.4
1.3 Rain
1.2
1.1
1
AN CAL CN RS RSA
Canals

Figure 2: Influence of Canal, Weather type and their interaction on the logarithm of the abundance of birds.
ANOVA results: Canals: p < 0.001, Weather Type p < 0.001, R2(adj) = 44.86%. Tortuguero National Park,
Limón, Costa Rica (a=Januari-June, b=Juli-December).

Table 1: Comparison of the amount of surveys and the percentage of surveys performed in the between the first
and second study period of every year. Significantly lower percentages of rain during surveys compared between
study periods are marked with a green background and marked with a star (*). Tortuguero National Park, Limón,
Costa Rica (a= Januari-June, b= Juli-December).

Weather ratio for the whole study area


Year Study period a Study Period b
Surveys with rain # Surveys Surveys with rain # Surveys
2014 15.38%* 52 24.56% 57
2015 19.44% 36 11.32%* 53
2016 12.73% 55 12.00% 50
2017 1.61%* 62 12.00% 50
2018 15.09% 53

Although summarizing the total avian biodiversity in just one biodiversity index is helpful,
caution must be taking when comparing the results. It is not only sensitive to natural variations
such as the weather, but also influenced by the sampling effort as just demonstrated. The
differences in the amount of surveys performed every study period could therefore have a
significant impact on the biodiversity index. Since our surveys are performed by research
assistants with various expertise, we should not ignore these influences, but the consistency of
our results demonstrate the robustness of the current protocols. Some of this bias can be excluded
by using average abundance (Figure 3), but a different amount of surveys on a different time of

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day could still have an important impact on the results. Also occasional records of rare species,
or species that are difficult to identify, could still influence the average abundance like a
biodiversity index. The average abundance of target species is unlikely to be impacted by this
sampling bias however. Also the increased use of digital cameras is not likely to influence the
average abundance of target species.

Long-term trends of average abundance per survey


30
25
20
15
10
5
0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a

Periods of six months


Target Species Incidental Species

Figure 3: Average abundance per survey of target and incidental species for every 6-month study period divided
into target and incidental species in Tortuguero National Park, Limón, Costa Rica (a= Januari-June, b= Juli-
December).

Like the Shannon-Wiener Biodiversity Index, the average abundance of target species did not
change considerably since 2014 (Figure 3). The average abundance of incidental species did
increase significantly since 2015 (Incidental species abundance = -171.2 + 0.004247 Day, p-
value= 0.01). The regression model shows that, on average, there are approximately 1.5 more
incidental species recorded on surveys every year since 2015 (Figure 4). That equals to an increase
of approximately 50% from 2015 until now. This impressive increase is even including the night
surveys that started in 2016 and generally have a very low abundance of incidental species. Once
again, the use of digital cameras is the most likely explanation for this, greatly increasing
identification success. Growing expertise, especially concerning auditive recognition of bird
species, can also partly explain this increase. The increase in records of incidental species does
therefore not necessarily reflect an increase in actual abundance or biodiversity.
The alternating pattern in abundance between the first and second study period in the last two and
a half year could be the result of a gradual delay in arrival and departure times of migratory
species, but further data collection and analysis are required to confirm this hypothesis. Ideally,

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the increase of comparable data throughout the years will allow comparisons between full years
in the future, allowing a more intuitive analysis of long-term biodiversity and abundance trends.
We can assume that the averages shown in Figure 3 are consistently, however slightly,
underestimating the true average abundance of birds on the canals. This is partly because
throughout all the years of this long-term project a period with a high abundance of migratory
birds is overseen during December and January. More likely than not, the survey team was also
not able to identify all incidental species on most of the surveys.

Abundance of incidental species over time


80
70
60
50
40
30
20
10
0

Figure 4: Regression of abundance of incidental species per survey over time in Tortuguero National Park, Limón,
Costa Rica (Incidental species abundance = -171.2 + 0.004247 Day, p-value = 0.01, R2 (adj) = 2.6 %).

Eighteen new incidental species have been recorded this study period. Two of which were not
recorded during standardized surveys, the Pinnated Bittern (Botaurus Pinnatus) and the King
Vulture (Sarcoramphus papa). Even though the Pinnated Bittern (Botaurus Pinnatus) was only
recorded once, on RS, it is worth mentioning since it fits the description of aquatic bird species
and can therefore be valuable while considering the health of the canal ecosystem. Five
individuals of the King Vulture have been sighted during two separate sightings on LJ and RSA.
A continuous increase of this species can be expected to the example of the Turkey Vulture
(Cathartes aura) in the past (Reference). All the incidental species recorded during the
standardized surveys can be found in Apendix B.
The slight increase in biodiversity since the beginning of 2017 is associated with the addition of
52 species to those already recorded up until 2016. This is very remarkable since the project has
been running for almost 9 years. The slight change of the study area in the beginning of 2016
from AN to LJ, caused by the inaccessibility of AN by dominating floating vegetation, is one of
the explanation for this. LJ is not just another canal. Being an estuary, LJ provides extra habitat
types to the study area as a whole including shallow water and a higher salt concentration of the

PAGE13
water which support different fish communities (reference). It is also a resting and feeding area
for many migratory species. It is thus not surprising that many of the new species since 2016 were
recorded on LJ including typical sea and shorebirds like the Brown Pelican (Pelecanus
occidentalis), Laughing Gull (Leucophaeus atricilla), Royal Tern (Thalasseus maximus), Greater
Yellowlegs (Tringa melanoleuca) and Collared Plover (Charadrius collaris). With the first
sighting of the Least Grebe (Tachybaptus dominicus) in the last study period, CN also keeps
adding new interesting species despite the low bird bundance on this canal (See Section 3.2).
Some of the more conspicuous species found for the first time since 2016 are Chestnut-collared
Woodpecker (Celeus castaneus), Zone-tailed Hawk (Buteo albonotatus), Peregrine Falcon (Falco
peregrinus), Collared Forest-falcon (Micrastur semitorquatus), Hook-billed Kite (Chondrohierax
uncinatus), King Vulture (Sarcoramphus papa) and Tiny Hawk (Accipiter superciliosus). These
records can definitely be regarded as positive signs for the quality of the environment and increase
our understanding about it. The only recorded individual of the Tiny Hawk sighted to date for
example, is a juvenile. Confirming that the food supply, mainly hummingbirds, is abundant
enough for a successful breeding attempt of this very specialized bird species. In contrast, most
of the new species are small and/or inconspicuous members of the Parulidae, Trochilidae,
Tyrannidae and Thraupidae families, probably the result of an increased sampling effort rather
than an actual biodiversity increase with implications for the quality of the ecosystem.
The night surveys are not included the biodiversity and abundance averages, since they are hardly
comparable with the other surveys with a different methodology and bias towards some canals.
In total 227 individuals, encompassing 25 species, have been recorded during 18 night surveys.
The average abundance of target and incidental species is 9.17 and 0.29 respectively with little
variation between the canals. Because of their nocturnal foraging behaviour it is not surprising
that the Yellow-crowned Night-heron (Nyctanassa violacea), Agami Heron (Agamia agami) and
Boat-billed Heron (Cochlearius cochlearius) are more abundant during nocturnal surveys
compared to the ones in the morning and afternoon (Table 2). They are also among the ten most
observed species during night surveys. With one exception, the rest of the top ten is completed
species which are very abundant during the day surveys but show a decrease of average abundance
during night surveys. The Grey-necked Wood-rail (Aramides cajanea), despite not foraging at
night, seems to like to roost on palm leaves overhanging the water. This probably reduces the
threat of predation and explains the higher abundance recorded at night.

Table 2: Comparison between the average abundance per survey during night or day surveys for the top ten most
abundant species during night surveys. The highest average abundances for each species comparing night and day
surveys are marked with a green background and a star (*). Tortuguero National Park, Limón, Costa Rica

Species name Average abundance per survey


Night Day
Butorides virescens 1.79* 2.22
Nyctanassa violacea 1.33 0.58*

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Anhinga anhinga 1.25* 1.48
Aramides cajanea 1.04 0.07*
Tigrisoma mexicanum 0.88* 0.96
Agamia agami 0.79 0.00*
Jacana spinosa 0.46* 6.76
Cochlearius cochlearius 0.33 0.04*
Egretta caerulea 0.21* 3.75
Megaceryle torquata 0.21* 0.50

Completed …. Point count surveys. Rain in the morning has often prevented the possibilities.

3.1.1 Local
The conclusions from previous section are interesting to get an idea of the general trends in bird
abundance, richness and biodiversity. Reports of previous study periods have pointed out
differences in abundance, species richness and biodiversity indices between the different canals
however (Claes 2017a, Vaca 2016). This points out the importance to investigate the changes for
each canal separately. It also provides vital information allowing management recommendations
and plans to be specified to the level of the individual canals. The result of these plans will
therefore benefit both local and regional biodiversity in the best possible way.

The characteristics of the avian biodiversity for this study period show roughly the same patterns
that have been statistically proven in reports for the previous study periods (Figures 5, 6, 7, 8, 9
and 10). For now, all off the characteristics of the avian community on any of the canals seem to
be stable apart from the average Shannon-Wiener Biodiversity Index for each canal (Figure 9).
All the canals seem to contribute to the slight increase in for the whole study area seen in Figure
1.

Since 2017, LJ is the canal with the highest biodiversity, abundance and richness per survey
(Figure 5, 6, 7). It also has a high average Evenness, especially comparing it with the other wider
canals, CAL, RS and AN in the past (Figure 8). Its total biodiversity and richness for the whole
study period is also still among the highest of the five canals (Figure 9 and 10). This is especially
remarkable since only one of the banks is studied for half of its length. All these observations
reflect the unique variations of habitats for which estuaries like LJ are renowned for (Reference).
With only 2.9%, LJ is the canal with the lowest percentage of surveys with rain however, which
could favour its average abundance, richness and therefore biodiversity.

PAGE15
Long-term trends of average Shannon-Wiener
Index for each canal
2.75

2.5

2.25 AN
CAL
2
CN
1.75 LJ
RS
1.5
RSA
1.25
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 5: Average Shannon-Wiener Index per survey for each canal and study period in Tortuguero National Park,
Limón, Costa Rica

During 2018a the second highest average abundance, species richness and biodiversity index with
a moderate Evenness were recorded on CAL again. It has recovered from the drop in every
investigated characteristic in the previous study period (Figure 4, 5, 6, 7, 8 and 9). Again we can
find the most probably explanation for this unsuspected decrease with the weather. In 2017b,
27.27% of the surveys were performed with rain, while this was only 6.67% and even 0% in the
study periods 2017a and 2018a respectively. In general however, CAL seems to be the canal with
the most pronounced alternating pattern in average biodiversity between the first and second study
period of each year. With its high abundance and richness, CAL alone can explain most of the
alternating pattern for the whole study area (See Section 3.1 and Figure 1). The weather pattern
alone is not enough to explain this strong pattern (Table 3). The Frequency of Occurrence of the
target species on California show the same pattern for both migratory and resident species, ruling
out the migratory season as a cause of this pattern (See section 3.3). Also no clear difference in
the percentage of recorded breeding adult or juveniles is detected between the two study periods
in every year. The reason for the alternating pattern in California thus remains unknown.

Table 3: Comparison of the percentage of surveys performed on California between the first and second study
period of every year. Significantly lower percentages of rain compared between study periods are marked with a
green background and marked with a star (*) (a= Januari-June, b= Juli-December). Tortuguero National Park,
Limón, Costa Rica.

Percentage of surveys with rain on California


Year Study periods a Study periods b

PAGE16
2014 9.09% 25.00%*
2015 12.50%* 9.09%
2016 9.09% 9.09%
2017 6.67% 27.27%*
2018 0.00%

RS still shows an average bird abundance which can be related to the variety in habitat (Figure
5). Its’ first two sections are more like LJ and CAL. Broad canals with a great abundance of
emergent vegetation, floating vegetation and shallow waters. Going upstream, RS gradually
develops into a narrower canal representing the situation described for RSA and CN better (See
Section 3.4). A bit surprising is that, even with this variation in vegetation structure, the average
species richness and biodiversity per survey and per study period are not very high compared to
the other canals (Figure 4 and 6). It even has the lowest biodiversity index value for almost every
study period as a whole (Figure 8).

CN and RSA remain the canals with the least average abundance and richness per survey (Figure
4 and 5. Both have abundant gallery vegetation and floating debris, available for perchers as
resting and feeding habitat. They lack big patches of floating vegetation, emergent vegetation and
shallow water as important feeding habitat for some of the most abundant species like the Snowy
Egret (Egretta thula), Little Blue Heron (Egretta caerulea) and Northern Jacana (Jacana
spinosa). CN and RSA inhabit a lot of species that are rarely recorded other canals, including the
target species Sunbittern (Europyga helias), Green-and-rufous Kingfisher (Chloroceryle inda)
and Boat-billed Heron (Cochlearius chochlearius). In this way they provide suitable habitat for
different bird communities compared to the wider canals however, making them important for the
regional biodiversity. Especially CN proves this with its consistently high value for its Shannon-
Wiener Biodiversity Index per study period compared to the other canals (Figure 8).

PAGE17
Long-term trends of average abundance per
survey for each canal
70

60

50 AN
CAL
40
CN
30
LJ
20 RS
10 RSA
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 6: Average bird abundance of per survey for every canal and study period (a= Januari-June, b= Juli-
December). Tortuguero National Park, Limón, Costa Rica.

Long-term trends of average species richness per


survey for each canal
21
19
17
AN
15
CAL
13
11 CN

9 LJ
7 RS
5 RSA
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 7: Average bird species richness per survey for each canal and study period (a= Januari-June, b= Juli-
December). Tortuguero National Park, Limón, Costa Rica.

PAGE18
Long-term trends of average Evenness per study
period per canal
0.9

0.85

0.8 AN
CAL
0.75
CN
0.7
LJ
0.65 RS
0.6 RSA
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 8: Average Evenness per survey for each canal and study period (a= Januari-June, b= Juli-December).
Tortuguero National Park, Limón, Costa Rica.

TOTALS

Long-term trends of Shannon-Wiener Index per


study period per canal
3.25

3 AN
CAL
2.75
CN
2.5
LJ

2.25 RS
RSA
2
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 9: Shannon-Wiener Biodiversity Index for every canal and study period (a= Januari-June, b= Juli-December).
Tortuguero National Park, Limón, Costa Rica.

PAGE19
Long-term trends of species richness per study
period and canal
60

55

50

45 AN
CAL
40
CN
35
LJ
30
RS
25 RSA
20
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 10: Bird species richness for every canal and study period (a= Januari-June, b= Juli-December).
Tortuguero National Park, Limón, Costa Rica.

3.3 Target species


While the average abundance per survey and the FO are both viable methods to show trends in
target species populations, they should be interpreted differently. The FO is mostly sensitive to
rare target species since a few sightings on different surveys could increase its category rapidly.
It is also very sensitive to a change in the number of canals where the target species have been
seen during the study period. It is not a reliable way to visualize abundances however, since
abundant species could still be appointed the same score while their abundance has in fact halved
or doubled. The average abundance per survey is thus a better indicator of the population sizes,
while the FO provides us information about the temporal and spatial variation of the presence of
the target species in the study area. Together they provide us a detailed overview of the status of
every species which is used to evaluate the environmental situation in every canal and the study
area as a whole.

The FO and average abundance for all target species combined has stayed very stable on all the
canals. The highest FO is recorded for LJ (Figure 11). This means that we see the most different
target species on this canal while considering temporal variations. The highest abundance of target

PAGE20
species recorded this study period was on CAL, closely followed by LJ. After that, the usual pick
order of RS, RSA and CN follows for both the FO and the average abundance (Figure 11 and 12).

Sum of the Frequency of Occurrence score for


every study period in each canal
70

60

50

40

30

20

10

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

AN Cal CN LJ RS RSA

Figure 11: Sum of the Frequency of Occurrence score of target species for each canal and every study period (a=
Januari-June, b= Juli-December). Tortuguero National Park, Limón, Costa Rica.

PAGE21
Average abundance per survey for every study
period in each canal
60
50
40
30
20
10
0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

AN Cal CN LJ RS RSA

Figure 12: Average abundance of target species for each canal and every study period (a= Januari-June, b= Juli-
December). Tortuguero National Park, Limón, Costa Rica.

Also regarding the FO of the target species, the alternating pattern observed in CAL is the most
striking (Figure 11). The same pattern is not found when comparing the average abundance per
survey however (Figure 12). This indicates that some species are more consistently present in
CAL in the first study period compared to the second one. Keeping the average abundance in
mind, they are also more abundance when they are present in the second study period. There is
one event that comes to mind given these characteristics. Migration. It seems to have a bigger
influence on the bird community in CAL than on any other canal. A comparison of the sum of the
FO scores on CAL between the first study period and the second study period of each year are
shown in figure 13. Apart from some species with a very low FO score, none of which are
migratory, the FO score for all study periods in the first half of the year combined are higher than
the ones for the second half of the year. A lot of these species are also residents however. The
reason for their decline in the second half of the year is not yet clear.

PAGE22
Comparison of the Frequency of Occurence of Target
Species between study periods A and B on California

30
Sum of FO scores of periods

25

20

15

10

P. brasilianus
A. alba

C. amazona

M. torquata
E. rufescens

T. mexicanum
E. helias
H. fulica

N. violacea

T. lineatum
P. martinicus
B. ibis
B. virescens

C. inda
C. americana

C. cochlearius

E. tricolor
A. cajanea
A. guarauna

A. herodias

M. alcyon
A. agami

E. thula

M. cayennensis
A. anhinga

E. caerulea

I. exilis

L. albigularis
J. spinosa
C. aenea

Study periods A Study periods B

Figure 13: Average abundance of target species for each canal and every study period (a= Januari-June, b= Juli-
December). Tortuguero National Park, Limón, Costa Rica

The same tool that created to make figures like Figure 11, can be used to evaluate the long-term
trends of each species or species guilds in the separate canals or the whole study area. Graphs of
the FO scores are shown and information of the average abundance is provided given that it offers
extra insights. The most interesting findings are discussed below (Figure 14-20).

The Agami Heron (Agamia agami) is the only target species that has never been recorded during
surveys in the morning or afternoon. It is perfectly explained by the nocturnal behaviour of this
species. During night surveys, especially on CN and LJ, they are recorded almost half of the time,
albeit in low abundances. Even more encouraging are the two sightings on LJ and CAL by day.
It prefers to hunt for small fish and frogs along small streams (Stiles & Skutch, 1989). The small
tributaries of LJ seem to be ideal habitat for the Agami Heron to strengthen its populations.
Conversely, the large amount of boat traffic on LJ (See section 3.5) make that unlikely. Also the

A less optimistic trend is detected for the Limpkin (Aramus gaurauna) (Figure 14). It has not
been recorded on surveys since study period 2015b. And only one sighting outside of the
standardized surveys has occurred on the first section of CAL. Before the end of 2015, it was

PAGE23
occasionally sighted in RS and RSA. Local presence of Limpkins is known to be affected by water
levels and preferably eat apple snails (Pomacaea) (Stiles & Skutch, 1989).

FO scores Aramus guarauna


Sum of FO scores for all canals 2.5

1.5

0.5

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 14: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Limpkin (Aramus
guarauna) in Tortuguero National Park, Limón, Costa Rica.

A clear increase in the recordings of the Great Blue Heron (Arda Herodias) can be observed since
2017a (Figure 15). The same pattern is also observed for the Neotropical Cormorant
(Phalacrocorax brasilianus) and the Tricolored Heron (Egretta Tricolor), although their numbers
fell back down in the most recent study period. This is largely the consequence of the change in
the study area. Most recordings of these species are on LJ, the canal included in the study area
since 2017. The large area of shallow water on LJ and the abundant fish populations associated
with estuaries likely attracts these species (reference).

FO scores Arda Herodias


Sum of FO scores for all canals

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

PAGE24
Figure 15: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Great Blue Heron
(Arda Herodias) in Tortuguero National Park, Limón, Costa Rica.

Even though the Snowy Egret (Egretta thulla) is one of the most abundant species in the study
area, it showed a slowly decreasing FO score in the past (Figure 15). This was even true for all
the canals in the study area except for LJ where it could still be categorized as constant. In the
most recent study period it recovered slightly with the highest FO on LJ and also becoming
‘Common’ on California again. Meanwhile, the average abundance per survey for the Snowy
Egret is very variable. This could be a sample bias since this species is very abundance in a short
period of the year. The downward trend in the FO score for the solitary Green Heron (Butorides
virescens) is mainly the result of a decrease of sightings on the more narrow canals CN and RSA
(Figure 16). The average abundance of Green Herons per survey also started decreasing on RS,
while it stayed constant on LJ and CAL. On top of that, Green Herons were more abundant on
AN compared to LJ. Both of these species are known North American migrants. Loss of habitat
and its quality outside of TNP and less suitable migration routes could thus explain this decrease
too.

FO scores Egretta thula


Sum of FO scores for all canals

12

10

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 16: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Snowy Egret (Egretta
thula) in Tortuguero National Park, Limón, Costa Rica.

PAGE25
FO scores Butorides virescens

Sum of FO scores for all canals


25

20

15

10

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Figure 17: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Green Heron
(Butorides virescens) in Tortuguero National Park, Limón, Costa Rica.

Also the Green Ibis (Mesembrinibis cayennensis) has the lowest FO score since 2014 (Figure 18).
The decrease is not concerning yet however, especially since the average abundance of Green
Ibises per survey is variable but stable. Most sightings of this species occur outside of the
standardized surveys early in the early morning high in the canopy of the gallery vegetation.

Although already not very frequently sighted before, the Sungrebe (Heliornis fulica) seems to
become less common throughout the years as well (Figure 18). These unique birds hold linear
territories along the banks and like to stick around areas where dense vegetation hangs in the
water (Stiles & Skutch, 1989). It also avoids rapidly flowing water so it is surprising that it is
most frequently seen on CAL, the canal with the strongest current within our study area. The
specific ecology of the Sungrebe points out that its absence on LJ and its disappearance on all the
other canals could be caused by a variety of factors. It is easily alarmed (personal observation)
and could therefore avoid canals with a lot of boat traffic such as LJ and CN (See section 3.5).
The strong current on RSA could also deter this species. It is also possible that the amount of
vegetation hanging over the canal has decreased over the years, resulting in habitat loss. In the
most recent study period, the average abundance of Sungrebes per survey showed a slight increase
compared to the previous one.

PAGE26
FO scores Mesembrinibis cayennensis
and Heliornis Fulica

Sum of FO scores for all canals


8

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Heliornis fulica Mesembrinibis cayennensis

Figure 18: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Green
Heron (Butorides virescens) and the Sungrebe (Heliornis fulica) in Tortuguero National Park, Limón,
Costa Rica.

The FO scores of the two biggest neotropical kingfisher species, the Ringed Kingfisher
(Megaceryle torquata) and the Belted Kingfisher (Megaceryle alcyon), showed a distinctly
increased in the past. For the two last study periods, this trend has reversed however (Figure 19).
It is particularly interesting that both species show the same pattern since they are involved in
direct competition for the same food source. The Belted Kingfisher is a North American migrant
however, while the Ringed Kingfisher is the most frequently seen resident kingfisher species.
This could represent a general change in resource or habitat availability. A continuation of the
vegetation structure analysis would allow these hypotheses to be tested in the future (See section
3.4). Also the FO score of the American Pygmy Kingfisher (Ceryle aenea) is characterized by a
steady increase since 2015. Its distribution has become more widespread within the study area,
however staying in the more narrow canals.

PAGE27
FO scores Megaceryle alcyon and
Megaceryle torquata

Sum of FO scores for all canals


20

15

10

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Megaceryle alcyon Megaceryle torquata

Figure 19: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Belted Kingfisher
(Megaceryle alcyon) and the Ringed Kingfisher (Megaceryle torquata) in Tortuguero National Park, Limón, Costa
Rica.

The last interesting pattern is found in the negative correlation of the FO scores of the Green
Kingfisher (Chloroceryle americana) and Amazon Kingfisher (Chlococeryle amazon) (Figure
20). This pattern is even more pronounced in their average abundance per survey. This could
reflect direct competition between these two species of similar ecology and size.

FO scores Chloroceryle amazona and


Sum of FO scores for all

Chloroceryle americana
15
canals

10

0
2014a 2014b 2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Chloroceryle amazona Chloroceryle americana

Figure 20: Long-term trends in the sum of the Frequency of Occurrence for all canals of the Green Kingfisher
(Chloroceryle americana) and the Amazon Kingfisher (Chloroceryle amazona) in Tortuguero National Park, Limón,
Costa Rica.

PAGE28
The numerous target species that show a fairly constant FO score throughout the whole study are
the Great Egret (Arda alba), Little Blue Heron (Egretta caerulae), Northern Jacana (Jacana
spinosa), White-throated Crake (Laterallus albigularis), Purple Gallinule (Porphyrio martinicus),
Yellow-crowned Night-heron (Nyctanassa violacea) and Bare-throated Tiger-heron (Tigrisoma
Mexicanum). For all these species the average abundance per survey is also stable or has increased
in the most recent study period compared to the previous study period. The same is true for the
Anhinga (Anhinga anhinga), but it showed a decrease in the study periods from 2014b to 2015b
for unknown reasons.

Species with a lower FO score and average abundance per survey show a more variable pattern.
They include the Grey-necked Wood-rail (Aramides cajanea), Cattle Egret (Bubulcus ibis),
Green-and-rufous Kingfisher (Chloroceryle inda), Boat-billed Heron (Cohlearius cohlearius),
and Rufescent Tiger-heron (Tigrisoma lineatum). Just like the American Pygmy Kingfisher
(Chloroceryle aenea), most of these species prefer small streams and dense vegetation (Stiles &
Skutch, 1989). They are probably much more common than our results suggest, since their
preferred habitat is underrepresented in our study area. During non-standardized surveys for the
purpose of training, often performed on the tributaries of CAL, the average abundances for these
species is perceived to be much higher.

The last three target species have only been recorded during a standardized survey once. The
Sunbittern (Eurypyga helias) was seen brooding on a nest on CN in study period 2017b, the Least
Bittern (Ixobrychus exilis) on AN in 2016b and the Reddish Egret on (Egretta rufescens) on RSA
in 2014b are the target species. Apart from the standardized surveys, the Least Bittern has been
sighted twice and the Sunbittern about once or twice every year.

Future investigation should focus on the continuation of the analysis of long-term trends like the
ones just discussed. More information should be collected about the habitat preferences, diet,
behaviour, and interspecific interactions concerning the thirty target species so they can be used
as bio-indicators to a better extend. It would be especially interesting to extract information about
the fish and invertebrate communities and the water quality, since there is little information
available about these key components in the canals in the southern part of TNP.

3.4 Vegetation structure


GVI has increased their efforts to monitor the vegetation structure of the canals in the most recent
study period. Dense vegetation allocates habitat for invertebrates, increasing food availability for
waterbirds (Anderson and Smith 2000; Rehfisch 1994; Wiggins et al. 1980). Emergent vegetation
provides shelter and improves nest-building conditions for wading birds (Froneman et al. 2001).
Dense mats of floating vegetation provides foraging areas for members of the Jacanidae and
Rallidae families (Ekhande 2012). It can also cover the areas used by diving birds such as
cormorants, anhingas and kingfishers however. In their turn these species are beneficially

PAGE29
impacted by the presence of floating debris as resting and hunting habitat. Shallow water has been
shown to be one of the most important factors for high abundance of aquatic bird species
(Reference). Further investigations of the vegetation structure of the canals are therefore likely to
be the most important source of information to explain the spatial and temporal variation of the
aquatic bird communities in Tortuguero National Park.

The collected data concerning the habitat use of the target species reveals clear associations
between many of the target species and the vegetation structures they use. To just highlight a few
of these associations, Sungrebes (Heliornis fulica) are seen swimming during 85% of their
sightings. More than 75% of the recorded Northern Jacanas (Jacana spinosa) have been recorded
on floating vegetation. Raphia Palms (Raphia taedigera) are the preferred habitat for 47% of the
Yellow-crowned Night-herons (Nyctanassa violacea) as a resting place during the day. Anhingas
(Anhinga anhingaare recorded on floating debris more than 40% of the time, even increasing to
83% for Neotropical cormorant (Phalacrocorax brasilianus). Kingfingers clearly prefer the
understory (35%) and floating debris (17 %) as a perch over mid canopy (0.38%) and high canopy
(8%). Lastly, shallow water is good for just over 25% of the sightings of large heron species such
as Great Egret (Ardea alba) and Great Blue heron (Ardea Herodias).

Long-term dynamics in the vegetation structure of the canals will have to be analysed when more
data is collected. It is also too early to correlate temporal changes in vegetation structure with
variations in the bird community. For now, an overview of the comparisons of the vegetation
structure in each section is given in the following figures (Figures).

As expected, the river mouth of LJ is the widest section, quickly narrowing towards the first
section. The second section is covered by emergent vegetation, influencing the width of the open
water. Traveling more upstream, CAL also gets more narrow but with much less variation. RS
and RSA show a linear decrease in canal width traveling upstream, resulting in the smallest
section of the study area. The width of the section on CN stays fairly constant.

PAGE30
Width at the start of the sections
350
300

Width (m)
250
200
150
100
50
0

Section

Figure 21: Overview of the width of the start of each section within the study area in Tortuguero National Park,
Limón, Costa Rica.

The width of the sections is likely to have interaction width all other vegetation types. Trees
falling in the water do not immediately disrupt local transportation and are thus more likely to be
available as floating debris. Shallow water and subsequently emergent and floating vegetation
may than develop because of a change hydrodynamics. The effects of boat traffic like erosion and
flushing of birds could also vary dependent on the width of the sections (See Section 3.5).

The moderate current and width of the canal combined with little boat traffic seems to result in
section with high amounts of emergent vegetation on the first three sections of RS. Of all the
habitat types, the emergent vegetation is probably the most challenging to find birds. Only 0.51%
of all the sightings of target are on emergent vegetation. The most cryptic of the target species
such as the White-throated Crake (Laterallus albigularis) and the Least Bittern (Ixobrychus exilis)
are highly reliant on them. The abundance of emergent vegetation on RS could be the main reason
for the low biodiversity on these sections (See section 3.4), but also indicates that this could be a
result of a sampling bias.

PAGE31
Amount of emergent vegetation
700
600

Width (m)
500
400
300
200
100
0

Section

There is a strong positive correlation between the abundance of emergent and floating vegetation
(p-value <0.001, R-sq (adj) = 42.1%). Emergent vegetation provides a good structure for floating
vegetation to accumulate and they both profit from high nutrient availability in the water.

Correlation between emergent and floating vegetation


500
450
Floating vegetation (m)

400
350
300
250
200
150
100
50
0
0 100 200 300 400 500 600 700
Emergent vegetation (m)

P-Value = 0.000, R-sq 42.1

PAGE32
The presence of the floating Waterhyacinth (Eichhornia crassipes) was detected in low
abundance across the study area (CAL2, LJ1, LJ2, RSA2 and RSA4). Originally from South
America, this floating plant is known for its dominant and invasive character and causes
biodiversity and economic loss all over the world. The best defence against this threat is early
recognition and action, despite high initial costs (Little, 1966). In general, these plants cause more
problems on lakes however and the dynamics of the floating vegetation in our study area are
poorly understood. Fertilizers and insecticides flowing into the park could stimulate the growth
of floating vegetation, while pesticides could kill them of. Strong currents, created by persistent
rainfall or opening of upstream dams could flush the vegetation into the sea. Also the populations
of Manatees (Trichechus manatus), known for their appetite for floating vegetation influence the
amount of vegetation cover in the canals (Wainwright, 2007).

The amount of floating debris seems to be the vegetation type with the most variation between
the different sections within the same canal. More research is needed to determine whether this
is influences the local aquatic bird communities. Strong associations between certain target
species and this habitat type suggest they might.

Amount of small floating debris


20
18
16
14
Count

12
10
8
6
4
2
0

Section
.

The canals in the study area are known for their black water and also for the significant amount
of Raphia Palms (Raphia taedigera) along the waterways (Myers 1990). The Raphia Palm is of
African origin and is reported to have arrived in the neotropics over 2000 years ago (Lewis et al.
2010; Carney and Hiraoka 1997). This species is associated with environments where organic
accumulation and nutrient leaching occurs during high water levels. The most striking aspect
when comparing the number of palms in the section is the sharp decline approaching the river
mouth of LJ, likely caused by an increase in salinity. Apart from this, palms seem to be an
important part of the gallery vegetation in every section of the canals.

PAGE33
Amount of Palms
90
80
70
Count 60
50
40
30
20
10
0

Section

3.5 Boat Traffic


The effects of human disturbances on bird populations have been widely studied for many years.
Specifically, motorized boat traffic can be detrimental to aquatic habitats and their communities.
Mathews (1982) ranked noise and rapid water movements due to power boats as the activities
having the highest impacts on bird populations, and shore-based activities having the lowest.
According to Rodgers and Smith (2007) and Loong (2002) flushing or fleeing of individuals due
to noise, with the addition of shoreline erosion are some of the main direct impacts of boat traffic.
Some studies argue that birds can adjust to noise disturbance (Loong 2002). Regardless, this is an
altered natural behaviour and thus physiological stress may still be a factor. Continued disturbance
acts to increase energy expenditure when birds continually flee in flight (Hockin et al. 1992). This
is especially important when considering migratory species. During the pre-migration period,
birds should spend more time feeding and foraging than flying in order to increase fat reserves.
By increasing disturbances, energy use increases, and birds may change feeding site preferences
to less suitable ones. It could potentially even change migration routes (Loong 2002; Hockin et
al. 1992).

The habitat quality can also be affected by motor emissions and exhaust; the resulting turbulence
can severely impact the bottom substrate of canals, especially in shallower canals (Loong 2002;
Asplund 2000). Contaminants and potential oil spills lower food quality and can directly affect
birds in the case of ingestion or coating of feathers, reducing their water-repellent quality.
Alternatively, an increase in nutrients from a mixing of the substrate can cause eutrophication,
further leading to algal or floral blooms, a change in vegetation and aquatic fauna community
structure (Asplund 2000).

PAGE34
The boats we see during surveys are only a small fraction of the ones actually present on the
canals. A large variability by chance and a bias towards the surveys start times should be expected,
but long-term monitoring could reveal interesting trends about boat traffic quantity and
composition. With these limitations in mind, we find no evidence of a general trend in boat traffic
over the past three and a half years. This period, the most records of motorized boat traffic
occurred on LJ (25) followed by CAL (10), CN (9), RSA (4) y RS (1). The average amount of
boats per survey are in line with the general trends of earlier study periods. LJ receives by far the
most boat traffic including transport, tourism, local, ranger activities (Illustration 12). CAL and
CN experience moderate boat traffic. On CAL this is very seasonal with fishing and tourism as
the main contributors, while CN has constant traffic passing by as a main transportation route.
RS and RSA virtually no motorized boat traffic. This is encouraging since park regulations seem
to be adhered to. Some illegal canal passage on RS and illegal fishing on RS, CAL and LJ are still
sporadically observed.

Boat traffic in the southern area of


Tortuguero National Park
0.8
Average number of boats per survey

0.7

0.6

0.5

0.4

0.3

0.2

0.1

0
2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

AN CAL CN LJ RS RSA

Illustration 1: Long-term trend of average amount of boat traffic per 1-hour survey categorized per canal in
Tortuguero National Park, Limón, Costa Rica.

PAGE35
It is clear that most boat traffic originates for ecotourism, followed by local fishermen (Figure).
The activity of the park rangers has been increasing throughout the years, mainly on LJ. The data
does not include the boat journeys undertaken to complete GVI’s surveys. Ten boat journeys were
needed for the surveys on both RS and RSA. For CAL, CN and LJ, 33 journeys were undertaken
from the river mouth to the Jalova Ranger Station and back. Actions are undertaken to reduce
these boat journeys considerably.

Boat traffic in the southern area of Tortuguero


National Park
0.8
Average number of boats per survey

0.7

0.6

0.5

0.4

0.3

0.2

0.1

0
2015a 2015b 2016a 2016b 2017a 2017b 2018a
Periods of six months

Commerce Ecotourism Local Other/Unknown Rangers

Illustration 2: Long-term trend of average amount of boat traffic per 1-hour survey for all canals categorized in
traffic types in Tortuguero National Park, Limón, Costa Rica.

Since 2017 we have started gathering more detailed information about the possible influence
motorized boat traffic has on the amount of birds we see during the surveys. Out of a comparison
of the section that were surveyed in roughly the same period of the year with the same weather
type during the same time of day, the sections were motorized boat traffic was present during the
surveys, more target species were counted (N = 33, p-value = 0.028). At first sight, this seems to
be a positive sign, but it could be a result of the target species being flushed out.

Flushing of birds would be interesting to formally record during future GVI canal bird monitoring
surveys to assess the impact of various boat sizes on bird behaviour. These changes in behaviour
of birds reduces reproductive success and abundance, which can lead to an alteration in

PAGE36
community composition and lower diversity, factors to which ecosystem stability is directly
correlated (Bibi and Ali 2013; Sabine et al. 2008; Rodgers and Smith 2007; Loong 2002; Asplund
2000).

Motorized boats can alter habitats by shore erosion and consequently the destruction of vegetation
along the banks. In turn nesting cover and food availability are reduced as well. The abundance
of waders is low across CN in comparison to that of perchers, particularly in comparison to all
other canals whose composition mostly consisted of wading species. A trend which could be
directly caused by the high boat traffic on this canal. Interestingly the Agami heron (Agamia
agamia), a sensitive wader species, has only been recorded in CN during the night surveys when
there is virtually no impact from boat traffic. Additionally, we know that historically the canal
had a large amount of floating vegetation. The banks of this canal have undoubtedly been subject
to erosion because of the large amount of boat traffic, resulting in the absence of floating or
emergent vegetation (See section 3.4).

Despite the current efforts, it is still impossible to evaluate the long-term impact of motorized
boat traffic aquatic bird populations and communities. Serious thought should go into the
development of a proper experimental design to quantify the amount of boats using the canals.
Recommendations can be found in section 4.2.

4. Recommendations
4.1 Evaluation of the project
GVI has delivered one report explaining the development of the project from 2010 to 2016 (Vaca
2016). It shows that the methods of this project constantly developed until 2014. Since then, they
have been standardized and remained constant, apart from the addition of night surveys and the
change of AN to LJ. Despite consistency in the survey methods, reports were still mainly focused
on 6-month study periods until 2017. The overall conclusions did not change much for every
period. They did unravel the most important patterns of avian biodiversity on the canals of the
southern part of TNP. The wide canals are particularly important to support large populations of
migratory species and common resident species. The narrow canals are a haven for many less
abundant species and are extremely important for the regional biodiversity (Claes 2017a). The
biodiversity indices calculated every study period show that all the canals as well as the whole
study area contain a large avian biodiversity with a moderate evenness (Bibi y Ali 2013, Claes
2017).

The authors and readers of these reports should be aware of the following important factors to
ensure the quality of this project. Apart from presenting the current local and regional avian

PAGE37
diversity in the southern part of TNP, this report explores the dynamics of the bird populations
and communities since 2014 and evaluates the motorized boat traffic since 2015. This is the first
semi-annual report that includes an analysis of the vegetation structure. All of these monitoring
efforts are designed to facilitate the addition of the results from upcoming study periods. This
allows the early detection of long-term trends which can then be acted upon accordingly by the
management of the park. It is extremely important that the methodologies remain consistent to
ensure this approach to be valuable. Any addition or changes to the study area or protocol and
their impact on the collected data must therefore be carefully examined.

Following the same reasoning, the different methodologies of the night surveys performed on LJ,
CN and CAL make it impossible to compare them to the surveys by day. This does not mean they
are not valuable though. Conducting surveys after sunset has allowed us to increase our
understanding of the target species and their behaviour within our study area in TNP. I would like
to stress that there should be a constant evaluation of the sustainability of every aspect of the
project and efforts to reduce the impact it has on the environment. This report would lose all its
relevance when the process of making it has a negative impact on the environment it studies.

It is important to strive for an equal sampling effort for every canal, during both morning and
afternoon, on every month. This accounts for natural temporal variation in the collected data,
prevents a sampling bias and increases power to discover significant long-term trends. Upon the
detection of these trends, evaluation of their causes and effects should be the next form of action.
This can be achieved by actively stimulating research projects with the data collected by GVI as
a strong basis. The most cost-effective and therefore interesting partners to perform these studies
are students from local universities. The conclusions of these studies can then be used to develop
new approaches for the management of the national park to the benefit of the stability and
biodiversity of the ecosystem and the highly valued ecosystem services it provides.

4.2 Further research

The large amount of data collected concerning the aquatic bird community shows its’ temporal
and spatial variation. To truly use the target species as bio-indicators for the adjustment of
management practises, we need to understand what causes this variation. Of key importance now
is to unravel natural variations from those caused external influences such as the ecotourism
season, the use pesticides and fertilizers in the plantations or hydro-electric dams upstream. GVI
started collecting boat traffic data since 2015 and vegetation structure data since the most recent
study period to do exactly that. Comparing the different sections, it will soon be possible to
analyse specific associations between the availability of certain vegetation structures and the
abundance of a target species. On top of that, a better understanding of the influence the boat
traffic has on the amount of birds seen on a section is now available. The relationship between
boat traffic, the vegetation structure and the bird community is more complex however. Natural
rainfall, boat traffic, hydro-electrical dams and upstream farming practices can influence current,
erosion and consequently vegetation structures. Other crucial information to understand the

PAGE38
dynamic bird populations can also be found in the vegetation, fish and invertebrate communities.
They are all bounded by the physico-chemical properties of the water (Algea, oxygen, heavy
metals, pesticides, insecticide concentrations and pH) flowing in TNP. Based on these
observations, I recommend GVI, MINAA and other research organisations to evaluate the
possibility of following studies:

-Precise measurement of boat traffic the canals are subjected to. Speed activated cameras at the
entrance of the canals could be considered for this.

-Formally record flushing distances and the number of birds being flushed by boat traffic.

-Analysing water quality with multi-parameters on a regular basis. This could also prove useful
as an early warning system in case of a contamination.

-Investigating impact of farming practices and hydro-electrical dams on the currents in TNP.

4.3 Management of Tortuguero National Park


LJ and CAL have identified themself as the most important locations supporting large, healthy
populations of aquatic birds within our study area. Healthy estuaries like LJ and structurally
diverse canals like CAL serve as source-populations contributing to stability of the ecosystem.
Any change in the management taken here has large significance for both resident and migratory
species. The situation in this area is complex however, since it is located at the edge of the national
park, incorporated in the touristic zone and part of the main transportation route connecting the
communities surrounding the park. The large fish populations on this border of the national park
also make the socially and economically important fishing culture possible. It is of great
importance that local traffic and ecotourism, including tours and fishing, are monitored and
controlled to a more satisfying degree. This is more accomplished at the northern entrance of TNP
compared to our study area. Close to the village of Tortuguero, tours are scheduled at fixed times,
performed on non-motorized or electrically powered boats and led by professionally trained
guides. On top of that, the collection of visiting fees are organized better, providing resources for
the management of the park. In our study area, it is not uncommon to see five motorized boats
speeding through LJ with two visitors on each boat, despite their larger capacity. Also for local
traffic, the speed seems to be limited only by the power of the engine. Neither are the ¨no-fishing
zones¨ consistently respected.

The ranger station in Jalova is ideally situated to oversee these activities, but could use extra
organization and personnel to work towards the model in the northern part of the park. Speed
limits, set at 30 km/h, should be installed in this area and made clearly visible to all visitors of the
park. More specifically designated passage ways, as far away from perches available to aquatic
bird species, are also recommended to prevent birds from being flushed out of their resting and
feeding habitat. Randomized patrols and effective, direct consequences such as fines or access

PAGE39
restrictions (for a clearly defined time) should be in stall for when infractions of park regulations
are observed by rangers. A register of all the motorized boats entering the park and the infractions
they make would be a valuable tool to enforce regulations. GVI has a list of a large part of the
motorized boats using the canals available which could aid the initiation of such a register.

Clear communication to all actors and a transition period with a warning system preceding actual
penalties is recommended for every change in enforcement. Awareness of the general public on
sustainable management and the importance of wildlife are vital to create support amongst the
general public for these regulations. I strongly encourage awareness campaigns and informative
sessions available to local communities about the importance of estuaries and their fish
community for migratory and resident aquatic birds. These awareness campaigns could include a
yearly, large-scale beach clean on the southern part of TNP. This action not only supports shore-
and seabird species but also directly improves the habitat quality of endangered sea turtles. GVI,
local communities and even ecotourism lodges could be valuable partners to participate in or even
organize such an event.

CN not only contributes to the regional biodiversity with its own high diversity, but also with the
differences in its community compared to the other canals. Finding a good balance between this
supporting ecological role and its function as the main route connecting important communities
surrounding the park is a hard challenge. CN is especially rich in perching kingfishers and solitary
target species. Trees and vines falling in the water increase microhabitat and perch availability
for these species. Due to safety concerns, they are often removed for motorized boat traffic. These
actions should be carefully considered on a case to case basis and certainly not be used to increase
this canals’ capacity for boat traffic. The discovery of a nest of the Sunbittern (Europyga helias)
highlighted the difficult situation on CN. Even though this was probably the first confirmed
breeding attempt of this species in TNP, no action was taken to ensure the success of this attempt.
I propose Quick Reaction Protocols to be developed and distributed to ensure a more effective
reaction in similar circumstances, especially when it concerns endangered species. These
protocols can include a communication plan towards local communities, temporary speed and
noise limits, buffer zones and increased patrols. Research protocols which minimize disturbance
could result in new information to increase the efficiency of those Quick Reaction Protocols.
RS and RSA are currently well protected. With less boat traffic, these canals with diverse habitats
could provide space for species that are sensitive to disturbance. The abundant vegetation on these
canals means we are probably underestimating the true biodiversity of these canals. A diversity
of different canals should enjoy this complete protection. A combination of well conserved small
streams, narrow and wide canals ensures the conservation and recovery of the regional
biodiversity. All studies canals, regardless of abundance or richness, show important habitat
structures for some or many of the bird species. Establishing no-go zones will be the best
mitigation method for reducing human impacts on the ecosystems in TNP. It is important that
external influences in these areas are still monitored however. Sufficiently large buffer zones must
be in place reduce edge effect.

PAGE40
5. Communication strategy
Once this report is presented to the authorities of MINAE and ACTO, it will be published on the
social media canals of GVI with the goal of educating the general public about the fauna in TNP.
We also hope that this report will be published on the website of MINEA and ACTO as a way to
communicate our results with the general public. The results of this report will also be used as a
part of the yearly presentation to the guides of Tortuguero. A presentation with a summary of the
results will also be presented to the research assistants of GVI to strengthen the importance and
impact of the ecological studies performed by GVI. Finally, the results of this report are also
intended to stimulate further research by students of local universities

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Attachments
Appendix A. Alphabetic list of all the species seen in the study area since the start of the project.

Scientific Name Family Order


Accipiter superciliosus Accipitridae Accipitriformes
Actitis macularius Scolopacidae Charadriiformes
Agamia agami Ardeidae Pelecaniformes
Amazilia tzacatl Trochilidae Apodiformes
Amazona autumnalis Psittacidae Psittaciformes
Amazona farinosa Psittacidae Psittaciformes
Amblycercus holosericeus Icteridae Passeriformes
Anas discors Anatidae Anatiformes
Anhinga anhinga Anhingidae Suliformes
Ara ambiguus Psittacidae Psittaciformes
Aramides cajanea Rallidae Gruiformes
Aramus guarauna Aramidae Gruiformes
Aratinga finschi Psittacidae Psittaciformes
Aratinga nana Psittacidae Psittaciformes
Ardea alba Ardeidae Pelecaniformes
Ardea herodias Ardeidae Pelecaniformes
Attila spadiceus Tyrannidae Passeriformes
Botaurus Pinnatus Ardeidae Pelecaniformes
Brotogeris jugularis Psittacidae Psittaciformes
Bubulcus ibis Ardeidae Pelecaniformes
Busarellus nigricollis Accipitridae Accipitriformes
Buteo Albonotatus Accipitridae Accipitriformes
Buteo brachyurus Accipitridae Accipitriformes
Buteo platypterus Accipitridae Accipitriformes

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Buteogallus anthracinus Accipitridae Accipitriformes
Butorides virescens Ardeidae Pelecaniformes
Cacicus uropygialis Icteridae Passeriformes
Cairina moschata Anatidae Anatiformes
calidris minutilla Scolopacidae Charadriiformes
Calidris pusilla Scolopacidae Charadriiformes
Campephilus guatemalensis Picidae Piciformes
campylorhynchus zonatus Troglodytidae Passeriformes
Cantorchilus thoracius Troglodytidae Passeriformes
Carpodectes nitidus Cotingidae Passeriformes
Cathartes aura Cathartidae Ciconiiformes
Celeus castaneus Picidae Piciformes
Ceratopipra mentalis Pipridae Passeriformes
Chaetura pelagica Apodidae Apodiformes
Charadrius collaris Charadridae Charadriiformes
Charadrius wilsonia Charadridae Charadriiformes
Chloroceryle aenea Alcedinidae Coraciiformes
Chloroceryle amazona Alcedinidae Coraciiformes
Chloroceryle americana Alcedinidae Coraciiformes
Chloroceryle inda Alcedinidae Coraciiformes
Chlorophanes spiza Thraupidae Passeriformes
Chondrohierax uncinatus Accipitridae Accipitriformes
Cochlearius cochlearius Ardeidae Pelecaniformes
Contopus cinereus Tyrannidae Passeriformes
contopus cooperi tyrannidae Passeriformes
Contopus sordidulus Thamnophilidae Passeriformes
contopus virens Tyrannidae Passeriformes
Coragyps atratus Cathartidae Ciconiiformes
Crax rubra Cracidae Galliformes
Crotophaga sulcirostris Cuculidae Cuculiformes
Cyanerpes lucidus Thraupidae Passeriformes
Dacnis cayana Thraupidae Passeriformes
Dendrocincla fuliginosa Furnariidae Passeriformes
Dendrocolaptes sanctithomae Furnariidae Passeriformes
Dendrocygna autumnalis Anatidae Anatiformes
Dendroica pensylvanica Parulidae Passeriformes
Dives dives Icteridae Passeriformes
Drycopus lineatus Picidae Piciformes
Egretta caerulea Ardeidae Pelecaniformes
Egretta rufescens Ardeidae Pelecaniformes
Egretta thula Ardeidae Pelecaniformes

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Egretta tricolor Ardeidae Pelecaniformes
Elanoides forficatus Accipitridae Accipitriformes
Empidonax virescens Tyrannidae Passeriformes
Empidonax virescens Tyrannidae Passeriformes
Euphonia gouldi Fringillidae passeriformes
Eurypyga helias Eurypygidae Eurypygiformes
Falco peregrinus Falconidae Accipitriformes
Falco rufigularis Falconidae falconiformes
Florisuga mellivora Trochilidae Apodiformes
Fregata magnificens Fregatidae Pelecaniformes
Geothlypis philadelphia Parulidae Passeriformes
Geothlypis semiflava Parulidae Passeriformes
Geothlypis semiflava Parulidae Passeriformes
Glaucis aeneus Trochilidae Apodiformes
Harpagus bidentatus Accipitridae Accipitriformes
Heliornis fulica Heliornithidae Gruiformes
Heliothryx barroti Trochilidae Apodiformes
Herpetotheres cachinnans Falconidae Accipitriformes
Himantopus mexicanus Recurvirostridae Charadriiformes
Hirundo rustica Hirundinidae Passeriformes
Ixobrychus exilis Ardeidae Pelecaniformes
Jacana spinosa Rallidae Gruiformes
Laterallus albigularis Rallidae Gruiformes
Leptodon cayanensis Accipitridae Accipitriformes
Leucophaeus atricilla Laridae Charadriiformes
Leucopternis semiplumbeus Accipitridae Accipitriformes
Manacus candei Pipridae Passeriformes
Megaceryle alcyon Alcedinidae Coraciiformes
Megaceryle torquata Alcedinidae Coraciiformes
Megarhynchus pitangua Tyrannidae Passeriformes
Melanerpes pucherani Picidae Piciformes
Mesembrinibis cayennensis Threskiornithidae Pelecaniformes
Micrastur semitorquatus Falconidae Accipitriformes
Molothrus oryzivorus Icteridae Passeriformes
Myiarchus critinus Tyrannidae Passeriformes
myiarchus tuberculifer Tyrannidae Passeriformes
Myiozetetes granadensis Tyrannidae Passeriformes
Myiozetetes similis Tyrannidae Passeriformes
Myrmeciza exsul Thamnophilidae Passeriformes
Notharchus macrorhynchos Bucconidae Piciformes
Nyctanassa violacea Ardeidae Pelecaniformes

PAGE47
Nyctidromus albicollis Caprimulgidae Caprimulgifor
Pandion haliaetus Pandionidae Accipitriformes
Parkesia noveboracensis Parulidae Passeriformes
Patagioenas cayennensis Columbidae Columbiformes
Patagioenas flavirostris Columbidae Columbiformes
Patagioenas nigrirostris Columbidae Columbiformes
Pelecanus occidentalis Pelecanidae Pelecaniformes
Penelope purpurascens Cracidae Galliformes
Phaethornis longirostris Trochilidae Apodiformes
Phaethornis striigularis Trochilidae Apodiformes
Phalacrocorax brasilianus Phalacrocoracidae Suliformes
Piaya cayana Cuculidae Cuculiformes
Pionus senilis Psittacidae Psittaciformes
Piranga olivacea Cardinalidea Passeriformes
Piranga rubra Cardinalidea Passeriformes
Pitangus sulphuratus Tyrannidae Passeriformes
platalea ajaja Threskiornithidae Pelecaniformes
Pluvialis squatarola Scolopacidae Charadriiformes
Porphyrio martinicus Rallidae Gruiformes
Progne chalybea Hirundinidae Passeriformes
Protonotaria citrea Parulidae Passeriformes
Psarocolius montezuma Icteridae Passeriformes
Psarocolius wagleri Icteridae Passeriformes
Pteroglossus torquatus Ramphastidae Piciformes
Pulsatrix perspicillata Strigidae Strigiformes
Pyrilia haematotis Psittacidea Psittaciformes
Quiscalus mexicanus Icteridae Passeriformes
Ramphastos sulfuratus Ramphastidae Piciformes
Ramphastos swainsonii Ramphastidae Piciformes
Ramphocelus passerinii Thraupidae Passeriformes
Riparia riparia Hirundinidae Passeriformes
Rupornis magnirostris Accipitridae Accipitriformes
Sarcoramphus papa Cathartidae Ciconiiformes
Setophaga petchia Parulidae Passeriformes
Sporophila corvina Thraupidae Passeriformes
Stelgidopteryx ruficollis Hirundinidae Passeriformes
Stelgidopteryx serripennis Hirundinidae Passeriformes
Streptoprocne rutila Apodidae Apodiformes
Tachybaptus dominicus Podicipedidae Anatiformes
Tachycineta albilinea Hirundinidae Passeriformes
Tachycineta bicolor Hirundinidae Passeriformes

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Tangara larvata Thraupidae Passeriformes
Thalasseus maximus Laridae Charadriiformes
Thamnophilus atrinucha Thamnophilidae Passeriformes
Thryothorus nigricapillus Troglodytidae Passeriformes
Tigrisoma lineatum Ardeidae Pelecaniformes
Tigrisoma mexicanum Ardeidae Pelecaniformes
Tinamus major Tinamidae Tinamiformes
Tityra inquisitor Tityridae Passeriformes
Tityra semifaciata Tityridae Passeriformes
Todirostrum cinereum Tyrannidae Passeriformes
Tringa melanoleuca Scolopacidae Charadriiformes
Trogon massena Trogonidae Trogoniformes
Trogon rufus Trogonidae Trogoniformes
Trogon violaceus Trogonidae Trogoniformes
Turdus grayi Turdidae Passeriformes
Tyrannus dominicensis Tyrannidae Passeriformes
Tyrannus melancholicus Tyrannidae Passeriformes
Tyrannus Tyrannus Tyrannidae Passeriformes
Vireo Olivaceus Vireonidae Passeriformes

Appendix B. Migratory status and Guild association of target species.

Scientific name Common Name Migratory status Guild


Agamia agami Agami Heron Resident Wader
Anhinga anhinga Anhinga Resident Swimmer
Aramides cajanea Gray-necked Wood-Rail Resident Wader
Aramus guarauna Limpkin Resident Wader
Ardea alba Great Egret North American Wader
Migrant
Ardea Herodias Great Blue Heron North American Wader
Migrant
Bubulcus ibis Cattle Egret Resident Percher
Butorides virescens Green Heron Resident & North Wader
American Migrant

Chloroceryle aenea American-pygmy Kingfisher Resident Percher


Chloroceryle amazona Amazon Kingfisher Resident Percher
Chloroceryle americana Green Kingfisher Resident Percher
Chloroceryle inda Green and Rufous Kingfisher Resident Percher
Cochlearius cochlearius Boat-billed Heron Resident Wader
Egretta caerulea Little Blue Heron North American Wader
Migrant

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Egretta rufescens Reddish Egret North American Wader
Migrant
Egretta thula Snowy Egret North American Wader
Migrant
Egretta tricolor Tricolored Heron North American Wader
Migrant
Eurypyga helias Sunbittern Resident Wader
Heliornis fulcia Sungrebe Resident Swimmer
Ixobrychus exilis Least Bittern Resident Wader
Jacana spinosa Northern Jacana Resident Wader
Laterallus albigularis White-Throated Crake Resident Wader
Megaceryle alcyon Belted Kingfisher North American Percher
Migrant
Megaceryle torquata Ringed Kingfisher Resident Percher
Mesembrinibis cayennensis Green Ibis Resident Wader

Nyctanassa violacea Yellow-crowned night heron Resident & North Wader


American Migrant
Phalacrocorax brasilianus Neotropic Cormorant Resident Swimmer
Porphyrio martinica Purple Gallinule Resident Wader
Tigrisoma lineatum Rufescent Tiger-Heron Resident Wader
Tigrisoma mexicanum Bare-throated Tiger-Heron Resident Wader

Appendix C. Categorization of Frequency of Occurrence according to Bensizerara et al. (2013).

Clasificación FO (%)

I Occasional <20
II Rare 20-39
III Common 40-59
IV Constant 60-79
V Abundant ≥80

Appendix D. Summary of the composition and classification of the Frequency of Occurrence for all target
species per canal for the whole study in the most recent study period.

Cal CN LJ RS RSA Todos canales


Cuantidad de muestras 15 11 15 11 10 62
Agamia agami Occasional Occasional - - - Occasional

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Anhinga anhinga Abundante Raro Abundante Abundante Abundante Abundante
Aramides cajanea Occasional - - - Occasional Occasional
Aramus guarauna - - - - - -
Ardea alba - - Comun Occasional - Occasional
Ardea herodias - - Raro Occasional - Occasional
Bubulcus ibis - - Raro - - Occasional
Butorides virescens Constante Occasional Abundante Constante - Comun
Chloroceryle aenea Occasional Occasional - Occasional Raro Occasional
Chloroceryle amazona Occasional - Comun - Raro Occasional
Chloroceryle
americana Comun - Comun Occasional - Raro
Chloroceryle inda - - - - - -
Cochlearius
cochlearius Occasional - Occasional - Occasional Occasional
Egretta caerulea Constante Occasional Abundante Abundante Constante Constante
Egretta rufescens - - - - - -
Egretta thula Occasional - Constante Occasional - Raro
Egretta tricolor Raro - Abundante - - Raro
Eurypyga helias - Occasional - - - Occasional
Heliornis fulica Occasional - - - - Occasional
Ixobrychus exilis - - - - - -
Jacana spinosa Abundante - Abundante Abundante Abundante Constante
Laterallus albigularis - - Raro Abundante Comun Raro
Megaceryle alcyon Occasional - Occasional - Raro Occasional
Megaceryle torquata Comun Comun Raro Constante Constante Comun
Mesembrinibis
cayennensis Raro Occasional Raro Occasional Occasional Occasional
Nyctanassa violacea Constante Occasional Constante - Occasional Raro
Phalacrocorax
brasilianus - Occasional Constante - - Raro
Porphyrio martinicus - - - Occasional Comun Occasional
Tigrisoma lineatum - - - Occasional - Occasional
Tigrisoma mexicanum Constante Raro Constante Comun Comun Comun

Appendix E. Start and End GPS-Coordinates for all canals. Laguna Jalova (LJ), California (CAL),
Caño Negro (CN), Rio Sierpe (RS), Rio Sierpe Alto (RSA), Tortuguero National Park, Limon,
Costa Rica.

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Start End

LJ N 10°20.742' 10°20.614'
W 083°23'.917'' 083°24'085''

CAL N 10°20'.312'' 10°20'103''


W 083°23'826'' 083°24'461''

CN N 10°21'437'' 10°21'959''
W 083°24'230'' 083°24'681''

RS N 10°23'018'' 10°22'540''
W 083°25'658'' 083°26'328''

RSA N 10°22'726'' 10°23'181''


W 083°26'633'' 083°27'165''

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