Vertical Distribution of Avifauna in the Cocha Cashu

Understory
Danielle Blick, Ross Furbush, Chris Stafford
Dept. of Biology, University of Washington

Abstract:
Understory birds were sampled in Cocha Cashu Biological Station using mist-nets and field
identification. We used capture height as a metric for vertical niche partitioning of understory
rainforest birds. Our data supports the hypothesis that bird Families use the understory preferentially.
This preference can be partially explained by morphological variation using a wing-to-weight ratio.
Other possible explanations for family strata level preference are discussed.
Abstracto:
La muestra de la diversidad de aves tomada en Cocha Cashu Estacin Biolgica, usando
redes de captura e identificacin de campo. Como paso adicional, fue determinado que de acuerdo
a un cierto peso, usan el nivel inferior del bosque llevando un record de el lugar de captura. Fue
descubierto que a ciertos niveles en la red puede representar una abertura tpica del la parte inferior
del bosque, que permite una mayor cantidad de especies usarlo para viajar, una especie de
carretera para las aves. Familias particulares son tambin ms frecuentemente capturadas en
ciertas redes. En un paso final, el radio del peso de las alas, fue calculado para cada especie
mostrando una correlacin baja con el lugar de captura en la red.

Introduction:
Manu National Park is considered one of the largest and richest natural reserves in the world.
A total of 800 birds species have been identified there with 500 of those being found in the lowland
forest around Cocha Cashu Biological Station alone (Schulenberg et al 2007). Despite such an
enormous amount of concentrated biodiversity, there seems to be little understanding or research
about the movement of the birds through the understory. Tropical species tend to be specialized in
horizontal and vertical habitat selection, and had lower "niche breadths" in foraging substrate and in
foraging height. In order to explore this area of research further, we first had to ask what is the
diversity of birds at different strata levels within the understory of Cocha Cashu?
Thamnophilidae (Antbirds) , Pipridae (Manakins) , Dendrocolaptidae (Woodcreepers),
Trochilidae (Hummingbirds) were amongst the four most common families we captured during the
course of our experiment. Often birds that are closely related in the same family share natural history
behaviors. These behaviors relate very closely to how these birds travel, forage and move
throughout the understory strata. This leads us to the second research question; At what height are
different birds potentially using the lower strata at Cocha Cashu. Hopefully we will begin to
understand how such a great amount of diversity is able to coexist. Gaining an understanding of
species coexistence and ecology has even greater conservational implications for the species and
their habitats alike.

Methods:
Our study took place at the neotropic research station Cocha Cashu, located on the Madre
de Dios River in the Peruvian Amazon (Latitude: -11.900000 Longitude: -71.366670). We captured
birds using mist-nets divided into four horizontal trammels, A, B, C, and D. The trammels ranged
approximately from ground level-0.625m, 0.625m-1.250m, 1.250m-1.875m, and 1.875m-2.500m,
2.500m-1.875m, respectively. A careful record was maintained on the trammel capture location for
each individual bird including recaptures.
We sampled for five consecutive days from 6:00am-11:00am as well as an afternoon session
from 3:30pm-5:00pm, excluding periods of heavy rain. Seven mist-nets were set up simultaneously
along pre-existing forest trails and were checked at 30 minute intervals. Two locations were
surveyed. The first location was a continuous old-growth forest, with little to no treefall gaps. The
second location was also an old-growth forest however was heavily influenced by tree falls leaving
large forest gaps. Nets at the second location strayed slightly from the forest trails.
Captured birds were identified using the most comprehensive field guide available for Peru,
Schulenbergs Birds of Peru, and processed by at least four researchers. We recorded body weight,
wing length, and height captured. Captured birds were marked with a clip of the outer tail feather.
Recaptured birds were released following their identification. Pictures were taken of the dorsal and
ventral side of the wings, as well as the tail plumage.
Figure 1
Results: Our team captured a total of 66
individuals. Within this sample 26 unique
species were identified, separated into
12 unique families. Trammels A and B
have more families and species than C
and D with B having more species
representation (Figure 1). Trammel B
alone, yielded the greatest diversity with
14 different species and 8 separate bird
families. Trammel A also showed a
higher amount of diversity than trammels
C and D. In Trammel A, 12 unique
species were identified belonging to 8
different families. A general decreasing
trend can be seen below trammel B.
Figure 1: Avifauna were captured and identified. Individual species
and families were plotted against corresponding capture height.

Trammel C produced 8 species of birds and 5 different families. Trammel D, the highest on
the net, captured the fewest amount of birds with only 5 species and 4 families. The graph allows us
to see that the family and species follow the same trend as trammel height increases.
Figure 2

Dividing the species into families

and comparing the four most frequently
caught shows us that 3 of the 4 families
were most frequently caught in trammel C:
Pipridae, Dendrocalaptidae, and
Trochilidae, but the fourth family,
Thamnophilidae, was more frequently
caught in the lower strata in trammels A
and B. The total number of occurrences
follows the trend of the 3 families with a
climax in C, but this is actually only a result
of high occurrences of Dendrocalaptidae
(n=10), Pipridae (n=15), and
Trochilidae(n=6). The remaining families
actually follow the trend of the fourth family,
Thamnophilidae (n=14). Trammel D had the
least occurrences by comparison.
Figure 2: Captured avifauna were sorted into family groups.
Families with a sample size of 5 or larger capture occurrences
were plotted against corresponding capture height. Families with
less than 5 capture occurrences were combined into other and
plotted against height captured.

Wing-to-weight ratio shows a loose correlating

trend with trammel level. The species of the
lowest trammel (A) are clumped with relatively
small wing-to-weight ratio variation. Trammel B
had the most variation in wing to weight ratio.
We didnt catch anything below 3.0 wing to
weight ratio in trammel D.

Discussion:
We observed a higher diversity in the lower strata levels than the higher strata levels of the
understory (Figure 1). This may be a result of trapping error or mist net efficiency. If sun casts on the
net then birds are more likely to see and avoid the net. The lower strata level will most likely be the
darkest part of the forest as a result of abundant leaf cover. With less sun cast, the net is more
efficiently hidden resulting in more capture occurrences (Jenni et. al). If biodiversity is a function of
number of capture occurrences, more occurrences increases the likelihood of capturing different
species; then this sun-cast effect would affect our biodiversity results.

It would appear that our

data does not follow the sun-cast
effect because total capture
Figure 1
occurrences is highest in trammel
C (22), followed by trammel B
and A (16 and 14) with the lowest
capture occurrences in trammel
D (5) (Figure 2). But we would
argue that trammel C is highly
skewed by the large number of
Pipra fasciicauda captured (11)
and that this species is
behaviorally biased towards
capture in trammel C.
Mist net can also be a poor reflection of
understory biodiversity because it is has
been observed that different species and
wind conditions can alter capture success
and escape proportions (Jenni et al). Without
monitoring the nets, via observation or
camera, we are unable to determine the
escape rate. Perhaps one species was
caught more frequently than others, but the
particular species is capable of escaping
before retrieval. Escape rate could also vary
between trammel heights, the birds caught
higher in trammel D are better at escaping or
more birds are able to escape from trammel
D.
Rainforest understory vegetation is much different than the open, dry, high canopy, but it has
variation of its own. This variation in the understory environment could be reflected by different
biodiversity per trammel. Perhaps the complexity of ground niches allows for more biodiversity with
more food availability, hiding spots, and shade from sun. In contrast, the portion of understory that
Figure 3: Captured avifauna were measured for weight and exists between 1.875 and 2.5 meters (D) could have an
wing length. Wing-to-weight ratio was calculated and
environment that is less conducive for high biodiversity.
plotted against height captured. Species with more than
Thamnophilidae is one of the most species
one individual caught at a certain height had their wing-todiverse families found in environments similar to Cocha
weight ratio averaged and plotted as a single point.
Cashu (Schulenberg et al 2007) and these birds are
classified as ground feeders. A family with high diversity and a strong association to the ground
supports our data results: the lower trammels, A and B, having the highest diversity.
Of the 12 families sampled, only 4 of them had high enough of a sample size to see trends.
These four families showed strong trends towards particular trammel levels: Trochilidae, Pipridae,
and Dendrocalaptidae were most frequently caught in trammel C and Thamnophilidae in the low

trammels, A and B. Particular families are more frequently caught in a particular trammel suggesting
they use different understory strata preferentially.
The trammel height most closely associated with a certain family could be congruent with the
level of the forest they use the most for feeding, nesting, or other behaviors. The previously
mentioned family of Thamnophilidae has a natural history more closely related to the ground,
consistent with the findings in trammels A and B. Trochilidae, hummingbirds, primarily feed on nectar
from flowers and there are relatively few flowers found low in the low understory near trammels A
and B.
We had a high capture occurrence of Pipra fasciicauda because our nets were set up in
close proximity to an active lek. Trammel C (appears to be the optimal height level to capture Pipra
fasciicauda. The height of their main displaying perch in the lek ranges from 3-5 m (Robbins 1983).
This seems too high to have an influence on where in the understory they are captured, trammel C
lies from 1.25-1.875 m, informing us there is a different behavior that is going on between 1.25 and
1.875 meters of the understory.
Black-spotted Bare-eyes, along with other species in the Thamnophilidae family, nest in tree
stumps on the ground and feed on the ground (Carlos et al 2000) so their most used strata level of
the understory is consistent with our capture findings because they were caught only in trammels A
and B (0-1.25m).
Looking at natural history by family is important because the optimal capture trammel is
different between Thamnophilidae and the other families of interest. But if we instead look at all the
families with highest capture occurrences in C (Dendrocolaptidae, Trochilidae, and Pipridae) as one
population, then we could speculate that they are all using this portion of the understory for the same
reason. The strata level from 1.25-1.875 meters could be the optimal traveling portion of the
understory for these families. Optimal traveling conditions for these three families could consist of:
low amounts of vegetation for easy flying, high twig proportions for lots of short bursts and rest, a
balance, or other combinations of optimal understory travel.
If trammel does not accurately represent the content and use of the corresponding
understory strata height level, then family height and use behavior cannot be derived from our
results. As mentioned previously, this could be a sampling bias from using mist nets. We also only
sampled two different sites and our mist nets stayed in the same specific location at each of the
sites. This may reduce our ability to make generalized statements about all of the understory. A
perfect window, with easy travel-ability and access, or other natural structure in the undergrowth,
such as an optimal perching branch or significant cover shade, could influence bird behavior around
varying strata level at different nets and sites. Trammel C could have captured a majority of the birds
because the birds were reacting to a specific natural window or structure only present at the few net
sites we sampled.
Despite using only two sites and one net location per net, our data still supports the idea of
family preference to trammel and strata level height. If mist nets had an effect on family trammel
capture occurrences, then Thamnophilidae would not have shown such a strong difference from the
other three families. When the four families are removed from the total occurrences (Figure 2) we
can see the remaining families follow the same trend as Thamnophilidae which also supports the
idea of family preference.
Bird families appear to show preference to a particular understory strata level. As explored
previously, this could be a function of mist net capture success, or family behavior and natural

history, but this trend could also be a function of varying morphological features. This corresponds
well with the idea of family preference, because birds in the same family tend to have similar
morphological traits: members of Trochilidae tend to be small and light whereas Columbidae
(pigeons and doves) tend to be large and stubby.
We used a wing-to-weight ratio as a morphological quantifier, low numbers representing a
long-winged light bird and large numbers for stubby, heavy birds. We isolated each species
according to trammel level to see if wing-to-weight ratio variation within a species varied between
strata levels: short-winged relatively heavy P. fasciicauda perhaps caught in a lower trammel than
long-winged relatively light weight P. fasciicauda.
Under the assumption that ground birds would spend less time flying, we would expect their
wings to be short and stubby relative to their weight. We see this trend in our data as 32.8 percent of
the variation in wing to weight ratio is explained by trammel level (Figure 3).
Some other trends not captured quantitatively show that the species of the lowest trammel
(A) are clumped with relatively small wing-to-weight ratio variation whereas trammel (B) have the
most variation. Despite having a low sample size for the highest trammel (D), we see that there is
not one species wing-to-weight ratio that exceeds 3.0 (Figure 3). A wing-to-weight ratio of 3.0 has no
biological significance, but it is noteworthy that all other trammels show capture occurrences of
species with less than 3.0 wing-to-weight ratio.
Future Implications:
Our preliminary data shows a loose trend between each trammel level in terms of biodiversity
and by family and wing-to-weight ratio. Further, extensive sampling should be done to fill in the
outlined trends from this pilot study. Along with more sampling efforts, there are other variables we
would like to consider adding to our sampling method to better capture bird behavior at varying
height levels in the understory at Cocha Cachu.
We would hope to better record the surrounding vegetation at each site and at a certain
diameter around each net, whether it is a complete survey of vegetation species, height, and
distance from net or a general estimate of percent vegetation cover at corresponding trammel height.
An array of natural history literature reveals important symbiotic relationships between specific birds
and plants. More information on the vegetation at our sites would help us better analyze possible
family, and even species interactions around the net. Common nesting materials, nest sites, or
feeding sites such as snags with many insects or a specific fruit bearing tree, all may influence which
trammel captures individuals and which part of the understory each species is using. An analysis of
surrounding vegetation would also help us better understand the bird highway hypothesis, perhaps
significant obstacles of natural structures would correspond with trammel capture success.
We never banded the birds, only snipped the tail feathers as a recap indicator, but bands
would help us track whether, not only species but individuals were showing preference to a certain
trammel height and therefore a certain understory height. This could be a result of a nearby nest at a
certain height, an individuals territory, a particular forage sites, or traveling channels. This would also
help us distinguish between resident birds and migrating or long distance traveling birds. Resident
birds might show a tendency to be caught lower than long distance traveling birds because flying
high can efficiently avoid obstacles and they would stop to feed less compared to residents.

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