ferguson 1 - digital.lib.usf.edu

16
Ferguson 1 Ectoparasites and their Avian Hosts in the Forests of Monteverde, Costa Rica Grace E. Ferguson Department of Environmental Science, Policy, and Management, College of Natural Resources University of California, Berkeley EAP Tropical Biology and Conservation Spring 2019 7 June 2019 ______________________________________________________________________________ ABSTRACT The term ectoparasite is defined as any parasite that lives on the outside of its host. On avian hosts, they occupy either the skin or feathers, consuming blood and tissue to provide the necessary nutrients to live and reproduce. In return, these tiny residents can cause harm to their hosts by transmitting disease and damaging important tissues. What is yet to be fully understood about this relationship between these parasites and their hosts is whether or not there is a clear correlation between characteristics of the host and the number and type of ectoparasites it carries. The main question of this study was whether or not it is possible to predict the abundance of ectoparasites on birds based on a number of characteristics of the host including their diet, the level of forest the inhabit, their social interactions, and their taxonomy. I mist-netted birds at three forested sites in the Monteverde area, examining each bird caught for the number and type of parasites it carried, and compiled that information with other research on the species of birds caught to answer this central question. My findings indicate that birds who spend most of their time in pairs have fewer parasites and that birds who consume mostly fruits and insects are likely to carry more parasites on average, however the abundance of ectoparasites is generally very variable. In addition, birds in pairs are far less likely to have ectoparasites present, which suggests an ecological benefit to a paired behavior strategy. Ectoparásitos y sus aves hospederas en Monteverde, Costa Rica RESUMEN Los ectoparásitos son parásitos que están fuera del cuerpo del organismo hospedero. En las aves ellos se encuentran en la piel o en las plumas, consumiendo sangre y otros tejidos que proveen los nutrientes necesarios para su vida y reproducción. Los ectoparásitos pueden causar daños a sus hospederos al transmitir enfermedades y dañar sus tejidos. Lo que todavía no se entiende bien acerca de la relación entre ectoparásitos y aves es si existe alguna correlación entre las características de las aves con la cantidad y el tipo de ectoparásitos que llevan. La pregunta principal de este estudio fue si se puede predecir la abundancia de ectoparásitos en aves según ciertas características de los pájaros como su dieta, estrato del bosque en el que habita, interacciones sociales, y grupo taxonómico. Capturé aves en redes de niebla en tres sitios en Monteverde, examiné cada pájaro observando el número y tipo de parásitos que llevaba, y compilé esa información con otras fuentes sobre las especies capturadas. Mis hallazgos indican que los pájaros que permanecen en parejas tienen menos ectoparásitos y los pájaros que

Upload: others

Post on 14-Feb-2022

1 views

Category:

Documents


0 download

TRANSCRIPT

Ferguson 1

Ectoparasites and their Avian Hosts in the Forests of Monteverde, Costa Rica

Grace E. Ferguson

Department of Environmental Science, Policy, and Management, College of Natural Resources

University of California, Berkeley

EAP Tropical Biology and Conservation Spring 2019

7 June 2019

______________________________________________________________________________

ABSTRACT

The term ectoparasite is defined as any parasite that lives on the outside of its host. On

avian hosts, they occupy either the skin or feathers, consuming blood and tissue to provide the

necessary nutrients to live and reproduce. In return, these tiny residents can cause harm to their

hosts by transmitting disease and damaging important tissues. What is yet to be fully understood

about this relationship between these parasites and their hosts is whether or not there is a clear

correlation between characteristics of the host and the number and type of ectoparasites it carries.

The main question of this study was whether or not it is possible to predict the abundance of

ectoparasites on birds based on a number of characteristics of the host including their diet, the

level of forest the inhabit, their social interactions, and their taxonomy. I mist-netted birds at

three forested sites in the Monteverde area, examining each bird caught for the number and type

of parasites it carried, and compiled that information with other research on the species of birds

caught to answer this central question. My findings indicate that birds who spend most of their

time in pairs have fewer parasites and that birds who consume mostly fruits and insects are likely

to carry more parasites on average, however the abundance of ectoparasites is generally very

variable. In addition, birds in pairs are far less likely to have ectoparasites present, which

suggests an ecological benefit to a paired behavior strategy.

Ectoparásitos y sus aves hospederas en Monteverde, Costa Rica

RESUMEN

Los ectoparásitos son parásitos que están fuera del cuerpo del organismo hospedero. En

las aves ellos se encuentran en la piel o en las plumas, consumiendo sangre y otros tejidos que

proveen los nutrientes necesarios para su vida y reproducción. Los ectoparásitos pueden causar

daños a sus hospederos al transmitir enfermedades y dañar sus tejidos. Lo que todavía no se

entiende bien acerca de la relación entre ectoparásitos y aves es si existe alguna correlación entre

las características de las aves con la cantidad y el tipo de ectoparásitos que llevan. La pregunta

principal de este estudio fue si se puede predecir la abundancia de ectoparásitos en aves según

ciertas características de los pájaros como su dieta, estrato del bosque en el que habita,

interacciones sociales, y grupo taxonómico. Capturé aves en redes de niebla en tres sitios en

Monteverde, examiné cada pájaro observando el número y tipo de parásitos que llevaba, y

compilé esa información con otras fuentes sobre las especies capturadas. Mis hallazgos indican

que los pájaros que permanecen en parejas tienen menos ectoparásitos y los pájaros que

Ectoparasites and Avian Hosts in Monteverde Ferguson 2

consumen principalmente frutos e insectos también llevan en promedio más ectoparásitos, sin

embargo, la abundancia de ectoparásitos es por lo general muy variable. Además, los pájaros en

parejas son menos propensos a llevar ectoparásitos, lo que sugiere un beneficio ecológico para

este comportamiento.

Ectoparasites have accompanied their avian hosts for many millions of years, relying on

their blood, feather, and skin tissue for survival and reproduction (Proctor and Owens 2000).

Most ectoparasites have specialized to their hosts to the point that some can only survive without

their host for short intervals, if at all (Schmidt and Roberts 2000). This not only strongly affect

the diversity of ectoparasites, causing rapid speciation and limiting genetic mixing between

populations, but also reducing the ways in which ectoparasites can be transmitted between hosts

(Schmidt and Roberts 2000). While ectoparasites can have major effects on their hosts by

transmitting disease, reducing vigor by damaging feather and skin tissue, and even consuming

nestlings in their entirety, they do not necessarily inhibit normal behavior or life processes and

live as many as 80% of wild birds (Proctor and Owens 2000; McFarlin and Robinson 2011).

The ectoparasites that specialize on these wild birds can come in many different shapes

and sizes and each has a unique niche that it inhabits (Schmidt and Roberts 2000). Diptera from

the Hippoboscidae family are one of these common avian ectoparasites, and tend to live near the

skin of the abdomen of birds to feed off of blood and are therefore also transmitters of blood

borne diseases (Arnold 1968). In addition, there are Acarid mites, mostly in the family

Proctophyllodidae which is a family of feather mite that is predominantly associated with

Passerine birds, though they have been documented on many orders (Mirnov et al. 2017). They

live on feathers with strong veins in the area between barbs, feeding on feather tissue and debris

(Mirnov et al. 2017). There are also ticks (Acarina: Ixodidae) and avian chewing lice (Insecta:

Phthiraptera) which occur in lower densities, but remain important members of the avian

ectoparasite community (Arnold 1970; Clayton et al. 1992). The prevalence and role they play in

the lives of birds around the world makes them a subject worthy of study, especially since the

population distribution of avian ectoparasites is highly variable. Furthermore, understanding how

ectoparasites affect bird communities could make it easier to understand how these interactions

might change in a future where there is a general decline in insect abundance and birds continue

to face threats from habitat loss and changing climates.

To help better understand this dynamic relationship, I asked the question: is it possible to

predict ectoparasite presence and abundance on the birds around Monteverde based on the

characteristics of their hosts? For many of these ectoparasites, they cannot survive without their

host (the avian chewing lice are an example of this according to Clayton and coworkers (1992)),

so one of my hypotheses is that contact with nestlings and other birds in a flock will affect the

number and diversity of ectoparasites on the birds. My other hypothesis rests with the ability of

some ectoparasites to survive in leaf litter, nest material, and inside flowers (Schmidt and

Roberts 2000), so therefore I hypothesized that the time birds spent in contact with these surfaces

would impact the number and type of parasites on their bodies.

Ectoparasites and Avian Hosts in Monteverde Ferguson 3

MATERIALS AND METHODS

Mist netting - I captured all of my birds for sampling using mist nets in 3 different forests at different elevations in the Monteverde area: the Rachel and Dwight Crandell Reserve trails, the Monteverde Ecological Sanctuary, and the Santa Elena Reserve. The timing of opening mist nets was very much dependent on weather conditions such as rain or lighting, making the actualized timing of opening and closing variable. In general, nets were opened between 5:15 am and 7:00am and closed sometime between 10 am and 11:00 am on ten days over the course of two weeks from May 12th to May 26th, accumulating approximately 50 hours spent mist-netting. The majority of the time, four 12-meter nets were used, sometimes supplemented by a 6-meter net and sometimes by an additional three 12-meter nets. The placement of the nets was deliberate with the intention of catching the most birds, so nets were placed along parts of the trail where more birds were observed, or where nets had been successful in the past.

Collection and Identification - Every bird was first identified using The Second Edition of The

Birds of Costa Rica by Richard Garrigues and Robert Dean, weighed using a spring scale. The

examination of the bird included all parts of the bird, first with the naked eye and later with a

magnifying visor to better view the smaller or more obscured ectoparasites. There was no strict

allotted time, but each bird took approximately 5-10 minutes to examine fully. Here again,

priority was placed on location of parasites rather than following a strict time regimen. In the

case of ectoparasite presence, I used tweezers to remove a few individuals from the bird and

placed them in a vial with 70% Ethanol solution to preserved. In some cases, the parasites were

too scarce to sample, in which case I simply made a note of their presence and number. In

addition to parasites, data was taken on feather condition as an indication of timing of most

recent molt, presence of brood patch to indicate whether the bird was currently sitting on a nest.

Mite Identification - I placed the mites in the 70% Ethanol solution in small sealed vials until

they could be identified. The identification process included extracting the mites from the vial

with tweezer and placing them on a slide to be viewed under a compound microscope. The

majority of the mites were clearly visible under 100x magnification. I then took pictures of each

mite so that the photos could be grouped into fifteen different morphotypes plus at least one

species of lice, one tick, and Hippoboscid flies. Morphotypes were defined based on body shape,

length, and hairs present.

Analysis - After accumulating all of the data and information, I compared mite presence and

abundance on the birds based on diet, habitat, feather condition, and presence of brood patch. To

analyze the number of ectoparasites on each bird beyond presence or absence, I assigned scores

based on the estimated quantity of ectoparasites on each designated part of the body (head, body,

tail, and wings). Anything under ten parasites received a score of “1”, while every bird with a

parasite load between ten and twenty-five received a “2”, and parasite loads larger than 25

received a “3” to indicate a high density of parasites on that respective body part. I then

compared the average scores for each body part against the other factors such as forest level

inhabited, diet, brood patch, and species. Any relevant information that could not be gathered in

the field, such as behavior, diet, and niche occupied was gathered from A Guide to the Birds of

Costa Rica by Stiles and Skutch (1989).

Ectoparasites and Avian Hosts in Monteverde Ferguson 4

RESULTS

In total, I captured 84 birds of 36 species from 14 families. Of the birds captured, there

were six general diets, eight described niche categories, and three categories of social behavior.

Out of 84 individuals, 65 (77.4%) were observed to have at least one of the four different types

of ectoparasites. Further breaking it down into species, there were only four species with no

observed parasites and a total of 11 species with less than 50% of individuals having

ectoparasites (Table 1). There were five birds (6.0%) with flies and another five birds (6.0%)

with lice (identified after collection and observation under a microscope), while the rest of the

ectoparasites observed were mites on the feathers of the wings, back of the head or the body.

Wings were the most likely to be infested with these mites, with 62 (73.8%) birds observed to

have mites on the wings versus only eight birds (9.5%) having parasites on their bodies, 25 birds

(29.8%) with mites on their heads, and 16 birds (19.0%) with mites on their tails.

Table 1: The percent frequency of presence of ectoparasites on each species of bird included in the study. The table

is ordered alphabetically by family, with the number of individuals included in the average in the column labeled

“Sample Size”

Family Species Ectoparasites Sample Size

Cardinalidae Habia fuscicauda 100% 2

Cracidae Chamaepetes unicolor 100% 1

Emberizidae Arremon brunneinucha 100% 2

Emberizidae Chlorospingus flavopectus 50% 2

Emberizidae Melozone leucotis 50% 2

Furnariidae Dendrocincla homochroa 100% 2

Furnariidae Margorornis rubiginosus 100% 1

Furnariidae Premnoplex brunnescens 25% 4

Furnariidae Sclerurus mexicanus 50% 2

Grallariidae Grallaria guatimalensis 100% 1

Momotidae Momotus coeruliceps 100% 1

Parulidae Basileuterus culicivorus 100% 1

Parulidae Basileuterus rufifrons 100% 1

Parulidae Basileuterus tristriatus 100% 1

Parulidae Myioborus torquatus 100% 1

Parulidae Seiurus aurocapilla 100% 1

Picidae Picoides fumigatus 100% 1

Pipridae Chiroxiphia linearis 67% 9

Strigidae Ciccaba virgata 100% 1

Trochilidae Amazilia saucerrottei 100% 3

Trochilidae Amazilia tzacatl 50% 2

Trochilidae Campylopters hemileucurus 100% 2

Trochilidae Eupherusa eximia 80% 5

Trochilidae Heliodoxa jacula 100% 1

Ectoparasites and Avian Hosts in Monteverde Ferguson 5

Trochilidae Lampornis calolaemus 100% 5

Trochilidae Phaethornis guy 100% 3

Troglodytidae Henicorhina leucophrys 33% 3

Troglodytidae Henicorhina leucosticta 0% 2

Troglodytidae Thryophilus rufalbus 43% 7

Turdidae Catharus aurantiirostris 100% 1

Turdidae Catharus frantzii 100% 2

Turdidae Catharus fuscater 100% 1

Turdidae Catharus mexicanus 100% 3

Turdidae Myadastes melanops 0% 1

Turdidae Turdus assimilis 100% 1

Tyrannidae Empidonax flavescens 0% 1

Tyrannidae Mionectes olivaceus 100% 5

Only eleven birds had a brood patch present, but eight of those eleven had parasites

(72.7%). Of those with no brood patch, 57 out of 73 birds had parasites (78.1%). Looking at the

social behavior, 34 out of 39 (87.2%) of solitary birds and 22 out of 27 (81.5%) social birds had

ectoparasites while only 9 of 18 (50%) of paired birds were positive for ectoparasites. In diets,

the carnivore and 100% of the eight Fructivore/Insectivores were observed to have ectoparasites

while only 31 of the 45 Insectivores (68.9%) and two of the four Omnivores (50%) had any

ectoparasites. Of the 19 birds with feathers in excellent condition, only 10 had parasites (52.6%),

while birds who were molting or with feather in good or fair condition were 88.9%, 82.8% and

84.4% likely to have parasites, respectively. Lastly, nine of the 12 birds who regularly come into

contact with the ground had ectoparasites (75%), while 23 of the 32 birds spending most of their

time in the understory (71.9%) and 33 of the 40 birds living between the understory and the

upper canopy (82.5%) had ectoparasites.

After the identification of 15 Acarid mite morphotypes and one Louse morphotype, there

was no morphotype that correlated directly to a species of bird, however the data indicate that

there is some connection between family and morphotype (Table 2). There is a morphotype

called “Mite 6”, for instance, which was only found on two families. Another example is “Mite

14”, which was indeed found on four different families, but 71% of the birds carrying this

morphotype belonged to the same family: Trochilidae (Table 2).

Table 2: Morphotype of Acarid mite (and one louse) on left with the family and species of bird from which the

ectoparasite was extracted.

Morph Family Species

Louse Trochilidae Lampornis calolaemus

Turdidae Catharus mexicanus

Emberizidae Arremon brunneinucha

Parulidae Myioborus torquatus

Grallariidae Grallaria guatimalensis

Mite 1 Troglodytidae Henicorhina leucophrys

Ectoparasites and Avian Hosts in Monteverde Ferguson 6

Parulidae Basileuterus culicivorus

Mite 2 Trochilidae Campylopters hemileucurus

Emberizidae Arremon brunneinucha

Turdidae Catharus mexicanus

Turdidae Turdus assimilis

Parulidae Basileuterus tristriatus

Cardinalidae Habia fuscicauda

Mite 3 Emberizidae Arremon brunneinucha

Parulidae Basileuterus tristriatus

Parulidae Seiurus aurocapilla

Mite 4 Trochilidae Lampornis calolaemus

Turdidae Turdus assimilis

Mite 5 Trochilidae Campylopters hemileucurus

Tyrannidae Mionectes olivaceus

Mite 6 Pipridae Chiroxiphia linearis

Trochilidae Campylopters hemileucurus

Trochilidae Lampornis calolaemus

Trochilidae Phaethornis guy

Tyrannidae Mionectes olivaceus

Pipridae Chiroxiphia linearis

Pipridae Chiroxiphia linearis

Pipridae Chiroxiphia linearis

Pipridae Chiroxiphia linearis

Mite 7 Trochilidae Lampornis calolaemus

Trochilidae Amazilia tzacatl

Furnariidae Dendrocincla homochroa

Mite 8 Turdidae Catharus mexicanus

Tyrannidae Mionectes olivaceus

Mite 9 Furnariidae Sclerurus mexicanus

Cardinalidae Habia fuscicauda

Mite 10 Parulidae Basileuterus rufifrons

Trochilidae Amazilia saucerrottei

Mite 11 Furnariidae Sclerurus mexicanus

Mite 12 Furnariidae Sclerurus mexicanus

Tyrannidae Mionectes olivaceus

Mite 13 Parulidae Myioborus torquatus

Trochilidae Eupherusa eximia

Trochilidae Eupherusa eximia

Turdidae Catharus aurantiirostris

Ectoparasites and Avian Hosts in Monteverde Ferguson 7

Mite 14 Trochilidae Campylopters hemileucurus

Trochilidae Phaethornis guy

Trochilidae Eupherusa eximia

Emberizidae Arremon brunneinucha

Troglodytidae Thryophilus rufalbus

Trochilidae Amazilia tzacatl

Trochilidae Amazilia saucerrottei

Mite 15 Troglodytidae Thryophilus rufalbus

Troglodytidae Henicorhina leucophrys

Cracidae Chamaepetes unicolor

Turdidae Catharus fuscater

Trochilidae Phaethornis guy

Trochilidae Heliodoxa jacula

Troglodytidae Thryophilus rufalbus

Trochilidae Amazilia saucerrottei

Turdidae Catharus aurantiirostris

In the next section, bird characteristics were compared against the average scores of

approximate parasite number (from zero to three) to reveal patterns in the number of parasites on

each bird. When these average scores were compared against bird diet, birds eating

predominantly fruits and insects had, on average, more parasites between the four parts of their

bodies, followed by the birds only eating nectar (Fig. 1). In a comparison of average scores to

niche space, birds who came regularly come into contact with the ground had fewer average

parasites than those who spent much of their lives in shrubs or trees (Fig. 2). In the final

comparison of average scores to social behavior, birds living in pairs had fewer parasites than

both birds who are solitary or social (meaning living in flocks of 3 or more) (Fig. 3).

Ectoparasites and Avian Hosts in Monteverde Ferguson 8

Fig. 1: Comparison of bird diet categories to the average score of each body part to reveal the total average score for

each diet category. The Fructivores/Insectivores have a cumulative average score of 4.25, which would correspond

to a number of parasites over 50. Meanwhile, the Fructivores have a cumulative average score of 1.8, which

corresponds to between 15 and 20 parasites per bird on average.

Fig. 2: Comparison of bird niche categories to the average score of each body part to reveal the total average score

for each niche category. The cumulative average score of birds spending significant time in contact with the ground

is significantly lower than birds of both other categories.

1

0,1778 0,0952

1,375

0,4222 0,6190,5

1,2

2,375

1,2222

1,8095

1

0,6

0,5

0,1778

0,381

0

0,5

1

1,5

2

2,5

3

3,5

4

4,5

5

Carnivore

(n=1)

Fructivore

(n=5)

Fructivore/

Insectivore

(n=8)

Insectivore

(n=45)

Nectivore

(n=21)

Omnivore

(n=4)

Aver

age

Cu

mu

lati

ve

Sco

re

Dominant Diet

Tail

Wings

Head

Body

0,12 0,08 0,21

0,41 0,62 0,5

1,29

1,54 1,43

0,12

0,41

0,18

0

0,5

1

1,5

2

2,5

3

Ground to Lower Canopy Understory Understory to Middle Canopy

Aver

age

Cum

ula

tive

Sco

re

Niche

Tail

Wings

Head

Body

Ectoparasites and Avian Hosts in Monteverde Ferguson 9

Fig. 3: Comparison of bird social behavior categories to the average score of each body part to reveal the total

average score for each niche category. The cumulative average score of paired birds is significantly lower than both

the social and solitary birds, with a score less than two.

DISCUSSION

To review, the main question of the study was whether or not it is possible to predict

presence and approximate number of ectoparasites on birds in the Monteverde area based on

physical, behavioral, and taxonomic characteristics of the birds themselves. According to a past

study done in this area, ectoparasites were more likely found on species living in forested

habitats and species with life strategies that included excessive contact with leaf litter or flowers

(such as nectivorous hummingbirds and ground dwelling birds) (McFarlin and Robinson 2011).

In addition, the natural history of many of the families of ectoparasites indicate that their

presence on birds would be related to the amount of time birds spent in contact either with other

birds or with substrates that could provide habitat for the ectoparasites between hosts. For these

reasons, I hypothesized that it would indeed be possible to find connections between the

physical, behavioral, and taxonomic characteristics of the birds and the number of ectoparasites.

My results indicate that while ectoparasite populations are varied and diverse on the birds

included in this study, there are some avian characteristics that are indeed correlated to

ectoparasite populations. First, there were a number of species of which none of the individuals

captured had ectoparasites, while there were also a majority of species in which every bird

captured had ectoparasites present. This indicates that there is at least some connection between

species and ectoparasite populations. The additional analysis of mite morphotype and bird

species or family showed no strong correlation in the majority of cases. This could indicate that

mite species are less closely associated to specific species than I predicted, but it could also be

the result of difficulty in properly classifying morphotypes.

When considering avian diet, niche, and contact with nests, there was less clear of a

correlation between these factors and presence or number of ectoparasites. This suggests that the

0,125 0,111 0,152

0,375 0,4820,697

1,208

1,519

1,5760,125

0,222

0,424

0

0,5

1

1,5

2

2,5

3

Pairs Social Solitary

Aver

age

Cum

ula

tive

Sco

re

Social Category

Tail

Wings

Head

Body

Ectoparasites and Avian Hosts in Monteverde Ferguson 10

ectoparasites I observed do not spend a significant amount of time apart from their hosts. In fact,

there were, on average, more parasites observed on birds that rarely come into contact with the

ground, which indicates that my prediction that ectoparasites might be transferred through leaf

litter was not supported. Going on, birds with new feathers that had little to no sign of damage

experienced fewer instances of parasitism when compared to birds in the process of molting or

with more damage to their feathers. This result could either indicate that newer feathers are less

likely to have ectoparasites or that ectoparasites contribute to feather damage, both of which

have scientific support (Jovani and Serrano 2001; Schmidt and Roberts 2000).

The strongest connection found was between the social behavior of the bird and presence

and abundance of ectoparasites. In this case, both solitary and social birds experienced similar

amounts of parasites at similar rates, while birds who lived in pairs not only have fewer average

parasites but were also less likely to have parasites in general. This suggest a connection between

pair behavior and protection from parasites and may indicate grooming as an important function

of the pair strategy. A future study might further analyze this pair relationship.

In conclusion, my hypothesis that it would be possible to predict ectoparasite presence

and number based on physical, behavioral, and taxonomic characteristics of the birds of

Monteverde, Costa Rica, was only partially supported with only a few of the characteristics

studied showing strong correlations to parasite distribution. In a future study, it might be

beneficial to consider other factors specific to individual birds to assess variation between

individuals of the same species or with similar natural histories. Going forward, this study has

shed some light on the connection between the birds of Monteverde and their ectoparasites, but

there is still much to learn and many more factors involved in this complicated relationship.

ACKNOWLEDGEMENTS

I would very much like to thank Federico Chinchilla for assisting me every day with the

mist-netting and parasite collection. His presence (and coffee) was an enormous help and I really

couldn’t have done it without him. In addition, thank you to the Monteverde Institute and

Santuario Ecológico Monteverde for allowing me to use their space for mist netting and a big

thanks to Luisa Moreno and the high school students of the Amigos del Ambiente for letting me

join them at the Santa Elena Reserve for mist-netting in the cloud forest. Thanks to Frank Joyce

and Katy VanDeusen for introducing me to the world of Costa Rican birds. Lastly, thank you to

UC EAP for such a wonderful experience.

LITERATURE CITED

Arnold, A., K. 1970. Notes on avian ectoparasites from Costa Rica. l. Acarina and Diptera. Revista de Biología Tropical, 16(2): 259-265.

Clayton, D., G., R., & Price, R. 1992. Comparative ecology of Neotropical bird lice (Insecta:

Phthiraptera). The Journal of Animal Ecology,61(3): 781. Dube, W. C., Hund, A. K., Turbek, S. P., & Safran, R. J. 2018. Microclimate and host body

condition influence mite population growth in a wild bird-ectoparasite system. International Journal for Parasitology: Parasites and Wildlife,7(3), 301-308.

Ectoparasites and Avian Hosts in Monteverde Ferguson 11

Evans, B., & Taylor, E. 2000. Factors affecting parasite abundance on birds. UCEAP Spring

2000 Course Book. Jovani, R., & Serrano, D. 2001. Feather mites (Astigmata) avoid mounting wing feathers of

passerine birds. Animal Behaviour,62(4): 723-727. Knee, W. 2006. Keys to the families and genera of blood and tissue feeding mites associated

with Albertan birds. Canadian Journal of Arthropod Identification. Mironov, S. V., Literak, I., Sychra, O., & Capek, M. 2017. Feather mites of the subfamily

Proctophyllodinae (Acari: Proctophyllodidae) from passerines (Aves: Passeriformes) in Costa Rica. Zootaxa,4297(1).

McFarlin, M., & Robinson, J. 2011. Bird ectoparasitic survey and possible community

implications for altered habitats in San Luis, Costa Rica. UCEAP Spring 2011 Course Book.

Proctor, H., & Owens, I. 2000. Mites and birds: Diversity, parasitism and

coevolution. Trends in Ecology & Evolution, 15(9): 358-364. Prahl, L. 2006. Determining avian vulnerability to ectoparasites using morphological and

natural history traits. CIEE Fall 2006 Course Book. Schmidt, D.G., & Roberts, S.L. 2000. Foundations of Parasitology. New York: Mc McGraw

Hill.

Ectoparasites and Avian Hosts in Monteverde Ferguson 12

APPENDIX

Table (Fig). 4: A list of the species collected, their collection date, the bird weight, and the site in which they were

collected. (MVI= Monteverde Institute, Rachel and Dwight Crandell Reserve trails, SM= Santuario Ecológico

Monteverde, and SER= Santa Elena Reserve)

Date Collected Bird Number weight (g) Species Name

5/14/19 MVI 01 27.50 Thryophilus rufalbus

5/16/19 MVI 02 30.00 Thryophilus rufalbus

5/16/19 MVI 03 4.50 Eupherusa eximia

5/16/19 MVI 04 18.00 Chiroxiphia linearis

5/16/19 MVI 05 5.00 Lampornis calolaemus

5/16/19 MVI 06 42.50 Melozone leucotis

5/16/19 MVI 07 25.00 Thryophilus rufalbus

5/16/19 MVI 08 4.75 Lampornis calolaemus

5/16/19 MVI 09 162.00 Momotus lessonii

5/16/19 MVI 10 32.00 Catharus mexicanus

5/16/19 MVI 11 27.50 Thryophilus rufalbus

5/16/19 MVI 12 28.00 Thryophilus rufalbus

5/16/19 MVI 13 20.00 Chiroxiphia linearis

5/16/19 MVI 14 31.00 Catharus mexicanus

5/16/19 MVI 15 13.00 Campylopters hemileucurus

5/16/19 MVI 16 57.00 Arremon brunneinucha

5/16/19 MVI 17 22.00 Thryophilus rufalbus

5/18/19 SM 01 18.00 Chiroxiphia linearis

5/18/19 SM 02 27.50 Thryophilus rufalbus

5/18/19 SM 03 4.50 Amazilia tzacatl

5/18/19 SM 04 5.50 Amazilia tzacatl

5/18/19 SM 05 3.50 Eupherusa eximia

5/18/19 SM 06 16.00 Henicorhina leucosticta

5/18/19 SM 07 13.00 Basileuterus rufifrons

5/18/19 SM 08 18.00 Chiroxiphia linearis

5/18/19 SM 09 16.00 Chiroxiphia linearis

5/18/19 SM 10 36.50 Habia fuscicauda

5/18/19 SM 11 35.50 Habia fuscicauda

5/18/19 SM 12 450.00 Ciccaba virgata

5/21/19 SM 13 17.00 Chiroxiphia linearis

5/21/19 SM 14 38.50 Dendrocincla homochroa

5/23/19 SM 15 16.00 Henicorhina leucosticta

Ectoparasites and Avian Hosts in Monteverde Ferguson 13

5/23/19 SM 16 18.50 Chiroxiphia linearis

5/23/19 SM 17 5.00 Amazilia saucerrottei

5/23/19 SM 18 5.50 Amazilia saucerrottei

5/23/19 SM 19 47.50 Melozone leucotis

5/23/19 SM 20 17.50 Chiroxiphia linearis

5/23/19 SM 21 4.50 Eupherusa eximia

5/23/19 SM 22 5.00 Amazilia saucerrottei

5/23/19 SM 23 11.50 Basileuterus culicivorus

5/23/19 SM 24 18.00 Chiroxiphia linearis

5/23/19 SM 25 45.50 Dendrocincla homochroa

5/23/19 SM 26 5.00 Eupherusa eximia

5/24/19 SM 27 17.00 Seiurus aurocapilla

5/24/19 SM 28 28.00 Catharus aurantiirostris

5/25/19 SER 01 18.50 Chlorospingus flavopectus

5/25/19 SER 02 9.20 Campylopters hemileucurus

5/25/19 SER 03 6.40 Phaethornis guy

5/25/19 SER 04 4.70 Lampornis calolaemus

5/25/19 SER 05 5.30 Lampornis calolaemus

5/25/19 SER 06 6.40 Phaethornis guy

5/25/19 SER 07 5.00 Eupherusa eximia

5/25/19 SER 08 17.50 Premnoplex brunnescens

5/25/19 SER 09 45.90 Arremon brunneinucha

5/25/19 SER 10 30.00 Sclerurus mexicanus

5/25/19 SER 11 4.30 Lampornis calolaemus

5/25/19 SER 12 11.90 Mionectes olivaceus

5/25/19 SER 13 30.50 Catharus mexicanus

5/25/19 SER 14 10.20 Catharus mexicanus

5/25/19 SER 15 27.00 Catharus frantzii

5/25/19 SER 16 69.30 Turdus assimilis

5/25/19 SER 17 15.50 Henicorhina leucophrys

5/25/19 SER 18 33.10 Myadastes melanops

5/25/19 SER 19 14.50 Premnoplex brunnescens

5/25/19 SER 20 18.70 Henicorhina leucophrys

5/25/19 SER 21 12.00 Basileuterus melanotis

5/25/19 SER 22 600.00 Chamaepetes unicolor

5/25/19 SER 23 98.00 Grallaria guatimalensis

5/25/19 SER 24 30.00 Catharus fuscater

5/26/19 SER 25 5.00 Phaethornis guy

5/26/19 SER 26 5.00 Heliodoxa jacula

Ectoparasites and Avian Hosts in Monteverde Ferguson 14

5/26/19 SER 27 20.00 Mionectes olivaceus

5/26/19 SER 28 20.00 Mionectes olivaceus

5/26/19 SER 29 30.00 Margorornis rubiginosus

5/26/19 SER 30 28.00 Sclerurus mexicanus

5/26/19 SER 31 15.00 Mionectes olivaceus

5/26/19 SER 32 17.50 Premnoplex brunnescens

5/26/19 SER 33 17.50 Premnoplex brunnescens

5/26/19 SER 34 20.50 Mionectes olivaceus

5/26/19 SER 35 14.50 Chlorospingus flavopectus

5/26/19 SER 36 18.70 Henicorhina leucophrys

5/26/19 SER 37 35.00 Picoides fumigatus

5/26/19 SER 38 30.50 Catharus frantzii

5/26/19 SER 39 20.00 Empidonax flavescens

Table (Fig.) 5: Sample name, family, common name, and score assigned based on approximate number of parasites

found on each section of the bird’s body.

Sample Name

Family Species Common name Body Head Wings Tail

MVI 01 Troglodytidae Rufous-and-white Wren 0 0 0 0

MVI 02 Troglodytidae Rufous-and-white Wren 0 0 0 0

MVI 03 Trochilidae Stripe-tailed Hummingbird 0 0 0 0

MVI 04 Pipridae Long-tailed Manakin 0 0 2 0

MVI 05 Trochilidae Purple-throated Mountain-gem 0 0 2 0

MVI 06 Emberizidae White-eared Ground-Sparrow 0 0 0 0

MVI 08 Trochilidae Purple-throated Mountain-gem 0 3 3 0

MVI 09 Momotidae Blue-crowned Motmot 0 0 2 1

MVI 07 Troglodytidae Rufous-and-white Wren 0 0 0 0

MVI 10 Turdidae Black-headed Nightingale-Thrush 0 2 3 0

MVI 11 Troglodytidae Rufous-and-white Wren 0 0 0 0

MVI 12 Troglodytidae Rufous-and-white Wren 1 2 2 0

MVI 13 Pipridae Long-tailed Manakin 0 0 2 0

MVI 14 Turdidae Black-headed Nightingale-Thrush 0 3 3 2

MVI 15 Trochilidae Violet Sabrewing 0 0 3 2

MVI 16 Emberizidae Chestnut-capped Brush-Finch 1 1 2 1

MVI 17 Troglodytidae Rufous-and-white Wren 0 0 1 1

SER 01 Emberizidae Common Chlorospingus 0 0 0 0

Ectoparasites and Avian Hosts in Monteverde Ferguson 15

SER 02 Trochilidae Violet Sabrewing 0 1 2 0

SER 03 Trochilidae Green Hermit 0 1 3 0

SER 04 Trochilidae Purple-throated Mountain-gem 0 0 3 0

SER 05 Trochilidae Purple-throated Mountain-gem 0 0 3 0

SER 06 Trochilidae Green Hermit 0 0 1 0

SER 07 Trochilidae Stripe-tailed Hummingbird 0 1 1 0

SER 08 Furnariidae Spotted Barbtail 0 0 0 0

SER 09 Emberizidae Chestnut-capped Brush-Finch 0 2 1 0

SER 10 Furnariidae Tawny-throated Leaftosser 0 0 0 0

SER 11 Trochilidae Purple-throated Mountain-gem 0 0 2 0

SER 12 Tyrannidae Olive-striped Flycatcher 0 0 3 0

SER 13 Turdidae Black-headed Nightingale-Thrush 0 2 2 0

SER 14 Parulidae Collared Redstart 0 2 3 0

SER 15 Turdidae Ruddy-capped Nightingale-Thrush 0 1 2 0

SER 16 Turdidae White-throated Thrush 0 1 2 0

SER 17 Troglodytidae Grey-breasted Woodwren 0 2 2 0

SER 18 Turdidae Black-faced Solitaire 0 0 0 0

SER 19 Furnariidae Spotted Barbtail 0 0 3 0

SER 20 Troglodytidae Grey-breasted Woodwren 0 0 0 0

SER 21 Parulidae Three-striped Warbler 0 0 3 1

SER 22 Cracidae Black Guan 0 0 0 3

SER 23 Grallariidae Scaled Antpitta 1 2 3 0

SER 24 Turdidae Slaty-backed Nightingale-Thrush 0 2 2 2

SER 25 Trochilidae Green Hermit 0 2 1 1

SER 26 Trochilidae Green-crowned Brilliant 2 0 1 0

SER 27 Tyrannidae Olive-striped Flycatcher 0 1 2 0

SER 28 Tyrannidae Olive-striped Flycatcher 0 0 3 0

SER 29 Furnariidae Ruddy Treerunner 0 0 3 0

SER 30 Furnariidae Tawny-throated Leaftosser 0 0 3 0

SER 31 Tyrannidae Olive-striped Flycatcher 0 2 0 0

SER 32 Furnariidae Spotted Barbtail 0 2 0 0

SER 33 Furnariidae Spotted Barbtail 0 0 0 0

SER 34 Tyrannidae Olive-striped Flycatcher 0 0 0 0

SER 35 Emberizidae Common Chlorospingus 0 2 3 0

SER 36 Troglodytidae Grey-breasted Woodwren 0 0 0 0

SER 37 Picidae Smokey-Brown Woodpecker 2 0 1 0

SER 38 Turdidae Ruddy-capped Nightingale-Thrush 0 0 2 0

SER 39 Tyrannidae Yellowish Flycatcher 0 0 0 0

SM 01 Pipridae Long-tailed Manakin 0 0 2 0

Ectoparasites and Avian Hosts in Monteverde Ferguson 16

SM 02 Troglodytidae Rufous-and-white Wren 0 2 1 1

SM 03 Trochilidae Rufous-tailed Hummingbird 0 0 0 0

SM 04 Trochilidae Rufous-tailed Hummingbird 0 0 1 1

SM 05 Trochilidae Stripe-tailed Hummingbird 0 0 2 1

SM 06 Troglodytidae White-breasted Woodwren 0 0 0 0

SM 07 Parulidae Rufous-capped Warbler 0 0 3 0

SM 08 Pipridae Long-tailed Manakin 0 0 0 0

SM 09 Pipridae Long-tailed Manakin 0 0 2 0

SM 10 Cardinalidae Red-throated Ant-Tanager 0 1 2 2

SM 11 Cardinalidae Red-throated Ant-Tanager 0 0 1 0

SM 12 Strigidae Mottled Owl 1 0 0 0

SM 13 Pipridae Long-tailed Manakin 0 0 0 0

SM 14 Furnariidae Ruddy Woodcreeper 1 0 0 0

SM 15 Troglodytidae White-breasted Woodwren 0 0 0 0

SM 16 Pipridae Long-tailed Manakin 0 0 0 0

SM 17 Trochilidae Steely-vented Hummingbird 0 2 1 0

SM 18 Trochilidae Steely-vented Hummingbird 0 0 3 1

SM 19 Emberizidae White-eared Ground-Sparrow 0 0 1 0

SM 20 Pipridae Long-tailed Manakin 0 0 2 0

SM 21 Trochilidae Stripe-tailed Hummingbird 0 0 2 0

SM 22 Trochilidae Steely-vented Hummingbird 0 3 1 0

SM 23 Parulidae Golden-crowned Warbler 0 0 2 1

SM 24 Pipridae Long-tailed Manakin 2 0 2 0

SM 25 Furnariidae Ruddy Woodcreeper 0 0 2 0

SM 26 Trochilidae Stripe-tailed Hummingbird 0 0 3 2

SM 27 Parulidae Ovenbird 0 0 1 0

SM 28 Turdidae Orange-billed Nightingale-Thrush 0 0 3 0