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••
THE ARTHROPOD NEST FAUNA OF HOUSE SPARROWS
AND TREE SWALLOWS IN SOUTHERN QUEBEC
Cyrena Riley
Department ofNatural Resource Sciences,
McGill University, Montreal • May 2000
A thesis submitted to the Faculty ofGraduate Studies and Research
in partial fui fi Ilment ofthe requirements ofthe degree of
Master ofScience
• © Cyrena Riley, 2000 National Ubrary nationale 1+1 of Canada :a=r ~uilitiona and Acquisitiona et Bibliographie services services bibliographiques 315 wellitglDn StreM ••ru. W...' OIawaON K1A0N4 o...ON K1A0N4 c.n.da c.n.da
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Canadl TABLE OF CONTENTS • List ofTables iv List ofFigures v
Abstract vi
Résumé vii
Acknowledgments viii
1. Introduction 1
2. Literature Review 3
2.1. Ectoparasites ofBirds 3
2.1.1. Haematophagous Mites (Acari). . 3
2.1.2. Feather Mites (Acari) 4
2.1.3. Swal10w Bugs (Hemiptera: Cimicidae) 5 • 2.1.4. Lice (Phthiraptera) 6 2.1.5. Fleas (Siphonaptera) 6
2.1.6. Blow flies (Diptera: Calliphoridae) 7
2.2. Non-Ectoparasitic Nest Fauna 9
2.3. Arthropods Associated with Nestboxes Il
2.4. Nest Arthropods Associated with House Sparrows 13
2.5. Nest Arthropods Associated with Tree Swallows 14
3. Materials and Methods ...... 16
3.1. Study Site 16
3.2. Collection ofNest Fauna 16 • 11 3.3. Diversity and Abundance ofArthropod Nest Fauna 18 • 4. Results 24 4.1. Nesting Success ofHost Birds 24
4.2. Inventory ofArthropod Nest Fauna 24
4.2.1. Arthropod Diversity: House Sparrow Nests 25
4.2.2. Arthropod Diversity: Tree Swallow Nests 30
4.3. Comparison ofArthropod Nest Fauna Within Bird Species 34
4.3.1. House Sparrow Nests 34
4.3.2. Tree Swallow Nests 35
4.4. Comparison ofArthropod Nest Fauna Between Bird Species 35
5. Discussion 38
5.1. Nesting Success ofHost Birds 38 • 5.2. Inventory ofArthropod Nest Fauna 38 5.3. Comparison ofArthropod Nest Fauna Within Bird Species 47
5.4. Comparison ofArthropod Nest Fauna Between Bird Species 51
5.4.1. House Sparrow Nests 51
5.4.2. Trec Swallow Nests 53
5.5. Conclusion 55
6. References 57
7. Appendices 86
• Hi LIST OF TABLES • 1. Arthropod abundance in House Sparrow nests. . 66 2. ArthroPOd abundance in Tree Swallow nests. . 71
3. Abundance ofdominant arthroPOds in House Sparrow and Tree Swallow nests 75
•
• iv LIST OF FIGURES • 1. Cluster analysis ofspecies richness in House Sparrow nests in 1997 and 1998. . 76 2. Cluster analysis ofSimpson's index in House Sparrow nests in 1997 and 1998 77
3. Cluster analysis ofBrillouin's index in House Sparrow nests in 1997 and 1998 78
4. Cluster analysis ofspecies richness in Tree Swallow nests in 1997 and 1998 79
5. Cluster analysis ofSimpson's index in Tree Swallow nests in 1997 and 1998 80
6. Cluster analysis ofBrillouin's index in Tree Swallow nests in 1997 and 1998 81
7. Cluster analysis ofspecies richness in House Sparrow and Tree Swallow nests ( 1997·
1998) 82
8. Cluster analysis ofSimpson's index in House Sparrow and Tree Swallow nests (1997-
1998) 83
9. Cluster analysis of Brillouin's index in House Sparrow and Tree Swallow nests (1997-
• 1998) 84
10. Mean abundance ofdominant arthropods in House Sparrow and Tree Swallow nests .
...... 85
• v ABSTRACT • The diversity and abundance ofthe arthropod nest fauna ofHouse Sparrows (Passer domestieus (L.» and Tree Swallows (Taehycineta bie%r (Vieillot» in southem
Quebec were studied. Over 90,000 arthropods were extracted from the nests, including at
least 71 species (50 families) ofinsects and at least 11 species (8 families) ofmites. There
were no significant differences in the species richness or diversity ofnest arthropods from
year to year ( 1997-1998), or frOtTa nest to nest within either host species. There was no
significant difference in the overall species richness or diversity between House Sparrows
and Tree Swallows. Cluster analyses ofspecies richness and three diversity indices for ail
nests showed no clustering according to bird species. However, there were sorne
significant differences in the abundancc and diversity ofparticular arthropod taxa
between the two bird species, with different ectoparasitic and non-ectoparasitic species • dominant in the nests ofeach host species.
• vi RÉSUMÉ • La diversité et l'abondance des arthropodes des nids du moineau domestique {Passer domestieus (L.» et de l'hirondelle bicolore (Tachyeineta bie%r (Vieillot» ont
été étudiées dans le sud de la province du Québec. Plus de 90,000 arthropodes ont été
extraits des nids, incluant un minimum de 71 espèces (50 familles) d'insectes et un
minimum de II espèces (8 familles) de mites. Aucune différences significatives n'ont été
observées entre la richesse d'espèces et la diversité des arthropodes d'une année à l'autre
(1997-1998), ou d't un nid à l'tautre à l'intérieur d'tune même espèce d'oiseaux. Il n't y avait
aucune différence significative entre la richesse d'espèces et la diversité des arthropodes
de nids de moineau domestique et d'hirondelle bicolore. L'analyse de groupement sur la
richesse d'espèces et sur les trois indices de diversité sur l'ensemble des nids n'a pas
regroupé ceux-ci selon l'tespèce d'oiseaux. Cependant, certaines différences significatives • ont été observées sur l'abondance et la diversité de certains arthropodes entre chaque espèce d'oiseaux, et certaines espèces d't arthropodes ectoparasitiques et non
ectoparasitiques étaient dominantes dans chacune des espèces d'oiseaux.
• vii ACKNOWLEDGMENTS • 1 am truly grateful to my supervisor, Dr. Terry Wheeler, for his extensive help, encouragement and editing ofthe manuscript. Dr. Wheeler's guidance, patience and
extensive biological knowledge proved invaluable throughout this project.
1thank the other members ofmy supervisory committee Dr. David Lewis and Dr.
Rodger Titman, for the advice given throughout the project and for their help in editing
this manuscript. Dr. Titman was also extremely helpful with field related questions.
1am grateful to the following scientists from Agriculture and Agri-Food Canada
for help with identifications ofarthropod species: Dr. Valerie Behan-Pelletier, Dr. Evert
Lindquist and Mr. King Wu (mites); Dr. Gary Oibson (parasitoid wasps); Dr. Jean
François Landry (tineid moths). 1also thank Dr. Wheeler for help with the identification
ofthe Diptera and ofthe arthropods in general.
• 1am grateful to Dr. Pierre Dutilleul and Dominic Frigon who helped with the application ofthe statistical procedures.
1 would like to thank Dr. Luc-Alain Giraldeau (Concordia University) for
allowing me to band birds under his banding pennit and for allowing me the use ofthe
nestboxes at the Macdonald Campus Farm ofMcGill University.
1would also like to express my gratitude to those who helped me in my fieldwork:
lan Ritchie (Avian Science and Conservation Center, McGill University) for suggestions,
encouragement and providing me with necessary tools; and Elizabeth Wright for getting
up at dreadful hours to help me band birds.
Financial support during this project was provided by McGill University and by • viii research grants to Dr. Wheeler from the Natural Sciences and Engineering Research • Council ofCanada and the Fonds pour la Fonnation de Chercheurs et l'Aide à la Recherche.
Finally, 1would like to thank the Lyman Museum staffand graduate students for
help and support: Dr. Chia-Chi Hsiung, Frédéric Beaulieu, Patrice Bouchard, Stéphanie
Boucher, Scott Brooks, Vanessa Crecco, Joanne Mudd, Jade Savage, Cory Keeler and
Tilly N'Gaoh Abdouramane.
•
• ix 1. INTRODUCTION • There is a large and diverse fauna ofarthropods associated with birds and their nests (Woodroffe, 1953; Hicks, 1953, 1959, 1962, 1971; Krïstofik et al., (996), but other
than studies on a few well-known ectoparasitic arthropods deriving their food from their
host, the rest ofthe nest fauna has virtually been ignored.
There has been an increased interest in ectoparasites ofbirds in recent years,
prompted by the recognition oftheir possible role in avian natural selection (Priee, 1980;
Hamilton and Zuk, 1982) and more studies have examined the interactions between some
ofthese ectoparasitic species and their avian hosts (e.g., Brown and Brown, 1986;
Whitworth and Bennett, 1992; Clayton and Tompkins, 1995; Merino and Potti, (995).
Ectoparasitic species are only a smalt portion ofthe arthropod community present
in bird nests; the non-ectoparasitic nest fauna is much more diverse and sometimes more
• abundant (Danks, 1979). However, the diversity and ecological roles ofthe 000
ectoparasitic nest tàuna have been ignored in many studies. For example, it has been
estimated that less than 1% ofthe mites associated with bird nests in Canada have been
described or recorded in this country (Danks, (979). The ecology ofmany ofthese
species is completely unknown. Hicks (1953, 1959, 1962, (971) provided a checklist of
the arthropods associated with bird nests, and there have been a few recent studies on the
nest fauna ofselected bird species (Krumpâl et al., 1993; [wasa et al., (995).
Because ofthe lack ofknowledge ofboth the parasitic and non-parasitic arthropod
tàuna ofbirds' nests in Canada, the purpose ofmy study was to examine the diversity of • the arthropod fauna associated with the nests ofselected bird species. My study had three objectives. The tirst objective was to conduct an inventory of • the arthropods present in the nests oftwo passerine birds using nestboxes in southern Quebec: House Sparrows (Passer domesticus (L.), Passeridae); and Tree Swallows
(Tachycineta bicolor (Vieillot), Hirundinidae). This is the tirst comprehensive inventory
ofthe nest fauna ofeither host species and will provide baseline data for future studies on
the biodiversity ofarthropods associated with nests ofNorth American birds.
The second objective was to compare the patterns ofdiversity in the arthropod
fauna within these bird species. Assuming that the nest arthropod fauna is strongly
intluenced by a combination ofnest construction and host behaviour, the hypothesis was
that nests ofthe same bird species have similar nest fauna, and thus, the abundance and
diversity ofarthropod species would be similar between nests ofthe same bird species.
The third objective ofthis study was to compare the patterns ofdiversity in the
• arthropod fauna between the two bird species. The hypothesis was that nests ofdifferent
bird species have different Dest fauna; therefore, the total abundance and diversity of
arthropod species would be different in nests ofthe two host species.
• 2 2. LITERATURE REVIEW • 2.1. Ectoparasites of Birds Most studies on the arthropod fauna ofbird nests have focussed on ectoparasitic
taxa. This is not surprising because ectoparasites probably have a greater and more direct
effect on the health and fitness ofthe host than the rest ofthe nest fauna. These
ectoparasites often become very abundant in nests, and can represent a significant portion
ofthe nest fauna (Burtt et al., 1991; Krumpâl et al., 1993). Ectoparasites may affect the
host in several ways: direct intake ofblood from the bird can result in anemia, decreased
growth or death; ectoparasites such as feather lice can damage feathers and decrease their
insulating ability; ectoparasites can a150 transmit diseases or pathogens that can decrease
the host's health. Several authors have studied the effect ofselected ectoparasites on the
fitness of nestling birds (e.g., Gold and Dahlsten, 1983; Chapman and George, 1991; • Merino and Potti, 1996). Because many ofthe published studies have concentrated on single taxa ofectoparasites, the major groups ofectoparasitic nest arthropods are dealt
with individually in the following sections.
2.1.1. Haematopbagous Mites (Acari)
Several genera ofhaematophagous mites are associated with birds and their nest
material. Two well-known genera in the order Mesostigmata are DermanyssIIs Dugès
(Dennanyssidae) and Ornithonysslls Sambon (Macronyssidae) whose high engorgement
capacity and short generation time can make them lethal ectoparasites (Moss and Carnin,
(970). Female mites oviposit one to five eggs 48 hours after a blood meal. These mites • 3 have five developmental stages: egg, larva, protonymph, deutonymph and adult. The • larva and deutonymph do not feed on the host, but the protonymph and adult take blood meals. The complete cycle from the blood meal ofthe female to the next generation of
adults can take as little as five to seven days (Baker et al., 1956). As a result, populations
ofhaematophagous mites usually increase exponentially during the host's nesting period,
and can build up to severa) thousand mites per nest (Pacejka et al., 1996). They
apparently overwinter in the nest material (Rendell and Verbeek, 1996) and their
abundance can be greatly reduced by the removal ofold Dest material from nestboxes by
the male bird (Pacejka et al., 1996).
These mites parasitize a wide range ofhosts; tor example, the Northern fowl mite,
Ornithonysslls sy/vian4m (Canestrini and Fanzago) parasitizes several families ofbirds
throughout the Holarctic regjon, and Dermanyssus ga//inae (Degeer) is a common • parasite ofboth wild and domestic birds (Clayton and Tompkins, (995).
2.1.2. Featber Mites (Acari)
Feather mites are permanent ectoparasites which feed on the homy layers ofthe
skin and feathers (Rothschild and Clay, (952). Although the etTect ofthese mites on their
hosts is not weil documented in the literature, 1have observed that when they are present
in high enough numbers their feeding activity causes physical damage to feathers in
caged birds ofvarious species. This would decrease the feather's insulating ability and
result in increased stress and make the birds more susceptible to adverse environmental
conditions. • 4 Feather mites are very diverse, with 2000 described species in 32 families • (Janovy, 1997) and larger birds appear to host greater numbers offeather mites than smaller birds (Rôzsa, 1997). Many species offeather mites show extreme microhabitat
specificity on the body ofthe host bird (Clayton and Moore, 1997). For example, Atyeo
and Windingstadt (1979) found four species offeather mites occupying different regions
ofsingle primary feathers ofSandhill Cranes (Grus canadensis (L.». Poulin (1991) found
that colonial birds show a greater risk ofbeing parasitized by feather mites than solitary
birds.
2.1.3. Swallow Bugs (Hemiptera: Cimieidae)
The best known bird ectoparasites in the Hemiptera are swallow bugs ofthe genus
Oeciacus Stal (Hemiptera: Cimicidae). Oeciaclls vicarius Horvath is host specifie, being • associated almost exc1usively with the CliffSwallow (Hirundo pyrrhonota Vieillot), while other species in the same genus have a wider range ofpossible hosts. Adults and
nymphs ofswallow bugs are blood feeders on the host birds. Cimicid bugs can reproduce
more than once in their lifetime, and nymphs take a long lime to mature (Brown and
Brown, 1986). Adult bugs mate before overwintering and females lay eggs the following
spring. Except for a briefmigration period in the spring, they usually remain in the nest
or in nearby crevices. Although they are able to reach new nests adjacent to old ones by
crawling, it appears that the only way for them to reach new nesting sites several
kilometres away is by clinging to the body oftheir host and being transported
phoretically (Woodbridge and Olkowski, 1968). • 5 2.1.4. Lice (Phtbiraptera) • Chewing lice (Phthiraptera: suborders Amblycera and Ischnocera) are obligate ectoparasites which spend their entire life cycle on the host (Marshall, 1981). Several
species ofchewing lice are distinctive in their size, shape and rnethod ofattachment to
their host, which indicates the great variability in habits ofthese ectoparasites (Boyd,
1951). The Ischnocera are usually narrow bodied and grasp the host's feathers, their only
source ofnourishment. Their life cycle lasts 3-4 weeks and includes an egg, three
nymphal instars and the adult (Martin, 1934). The females probably rernain fertile for lite.
The Amblycera have squat bodies, are fast rooners and are found between the skin and
the teather quill; like Ischnocera they feed on feathers, but they also feed on blood and
skin secretions (Boyd, 1951). There is little published information on the effect ofthese
ectoparasites on their hosts. Lee and Clayton (1995) found that host reproductive success • and survival appear independent oflice abundance.
2.1.S. Fleas (Sipbonaptera)
The life cycle ofbird fleas (Siphonaptera: Ceratophyllidae) includes both blood
feeding and non-blood feeding stages. The adults feed on blood and mate while on their
host, and then drop to the nest rnaterial. Eggs are laid directiy in the nest material or on
the host and because they are not adhesive, they usually faH into the nest matenal. The
eggs hatch in 2 to 12 days (Rendell and Verbeek, 1996) and the larvae stay in the nest
material where they feed on a wide range ofdecayjng organic materials (Holland, 1985).
They then spin an irregular cocoon and overwinter as pupae (Ross et al., 1991). [n the • 6 spring, adults emerge from the cocoon, and can emigrate from the nest and live on the • ground or in vegetation until a new host is located (Bates and Rothschild, 1962; Harper et al., 1992).
Many species offleas are host specific, but sorne ofthe more abundant and
commonly collected species, such as Ceralophy//us ga//inae Schrank, are known to
parasitize a wide range ofhosts, including birds and mamrnals (Holland, (985).
Christe et al. (1994) found that Great Tits (Parus major L.) actively selected
nestboxes devoid offleas. Rendell and Verbeek (1996) observed a positive correlation
between the volume ofnest malerial and flea numbers.
2.1.6. Blow Oies (Diptera: Calliphoridae)
Probably the most thoroughly studied nest ectoparasites in North America are • blow flies ofthe genus Proloca/liphora Hough (Diptera: Calliphoridae), whose haematophagous larvae feed on avian blood, usually from the nestlings. Females of
Protocalliphora species lay eggs singly or in batches in nests, typically within a week
after the host's nestlings have hatched (Rendell and Verbeek, 1996). The fly eggs hatch
24 to 48 hours after oviposition, and the larvae go through three larval instars (Sabrosky
et al., (989). The first two are quite short, lasting a few days, and require only one blood
Meal each. However, the third instar lasts a minimum ofthree days (Bennett and
Whit\vorth, 1991) and requires two to three blood meals. This instar is believed to be the
one most detrimental to nestlings (Johnston and Albrecht, 1993). Rohy et al. (1992)
found that the Mean mass ofindividual third instar blow tly larvae increased from 10 mg • 1 to 75 mg. Third instar larvae stop feeding for a short period prior to pupation. Puparia are • brown and oval in shaPe, and can be found at any level in the nest material depending on the structure ofthe nest. Adults emerge from the puparia in 14 to 21 days, depending on
weather conditions (Sabrosky et aL., 1989). Emerging adults do not overwinter in the
nests (Sabrosky et aL, 1989) and emigrate from the nest to overwinter in crevices, cavities
or under tree bark (Rendell and Verbeek., 1996). The duration oflarval instars and
pupation varies from one Protocal/iphora species to another., and was summarized by
Bennett and Whitworth (1991).
The number ofblow fly larvae found in a nest varies greatly; Sabrosky et al.
(1989) reported over 1200 larvae found in one nest ofan unidentified host species., but
numbers are typically lower.
Protocal/iphora flies have been recorded from the nests ofa wide variety ofhosts • including many passerines, several birds ofprey and a few shorebirds (Sabrosky et aL, 1989). Several authors have noted the preference ofProtoca/liphora flies for birds with
sheltered nests. Mason (1944) found that although these flies are found in open nests of
several small passerines, there is evidence that bird species building open nests are not
preferred hosts; these nests will be hazardous to the larvae because there is a greater
danger offalling through the bottom ofthe nest. This was also observed by Bennett and
Whitworth (1992). Gold and Dahlsten (1989) stated that hosts ofProtocal/iphora are
primarily cavity-nesting birds in forests; attacks on cup-nesters were uncommon but also
associated with forested areas.
• 8 2.2. Non-Ectoparasitic Nest Fauna
• Non-ectoparasitic arthropods associated with birds y nests have been much less intensively studied than ectoparasitic species; most studies on nest fauna have been either
superficial or anecdotal. One ofthe tirst comprehensive studies on the arthropod fauna
associated with birds Y nests was by Nordberg (1936). He studied arthropods associated
with 422 nests of56 bird species from Finland and classified the arthropods on the basis
oftheir ecology and association with specifie types ofnests. He also discussed the
intluence ofenvironmental conditions (temperaturey humidity, nest volume, etc.) on the
insect community present in the nest. Woodroffe (1953) did a similar study in Britain,
although he did not indicate how Many nests or host species he examined. However, he
described in detail the arthropods extracted from the nest material and divided them into
ditferent categories according to their feeding preferences and trophie guilds: • ectoparasites, seavengers and predators. Woodroffe (1953) also described the environmental conditions present in the nests and their effeet on the nest arthropods.
The ehecklist by Hicks (1953, 1959, 1962, 1971) is the most extensive work on
the insects found in North American birds' nests. Hicks compiled a list based on
published records and personal observations ofthe insects round in nests. The ehecklist
focussed primarily on bird ectoparasites and did not discuss the relationship between the
bird and the insects; also, because the checklist was based extensively on previous
publications, it is likely that any errors or omissions in the original publications were not
corrected.
McAtee (1927) conducted a comprehensive study ofthe arthropod fauna present • 9 in several nestboxes occupied by different bird species" and made observations on the • ecology ofthe different categories ofinsects present. Hood and Welch (1980) conducted a detailed study ofthe nest fauna ofRed-winged Blackbirds (Agelaius phoeniceus (L.» in
Manitoba. They identified and listed ail arthropods present in five nests, and outlined the
differences in the arthropod fauna found by Woodroffe (1953) in dry nests and the fauna
found in their study. The nest fauna examined by Hood and Welch (1980) was dominated
by Diptera (65.60/0 ofail specimens) and there were very few mites, compared to
Woodroffe's study in which mites reached high numbers and the Diptera amounted to
only a few individuals.
Several studies have focussed on the seasonality and distribution ofarthropods in
birds' nests, but were often restricted to only one group ofarthropods. Burtt et al. (1991 )
studied the occurrence and demography ofmites in nests ofTree Swallows, House Wrens • (Troglodytes aedon Vieillot) and Eastern Bluebirds (Sia/ia sialis (L.», ail cavity nesters. They found three species ofmites in these nests: Dermanyssus hin,ndin;s (Hennann), a
haematophagous ectoparasite ofbirds; Dermatophagoides evansi Fain, a scavenger; and
Che/etomorpha lepidopteron,m (Shaw), a predator on small arthropods, including other
mites. They discussed the different guilds ofmites that were studied, and noted that not
ail mites present were ectoparasites ofbirds. Burtt et al. ( 1991) also showed tbat
predaceous mites may play a role in the control ofectoparasitic mites in birds' nests.
K.rumpâl et al. (1993) studied the distribution and seasonality ofarthropods associated
with Sand Martin (Riparia riparia L.) nests. The Siphonaptera were most abundant,
followed by larvae ofvarious insects; Formicidae and Diptera were considered • 10 subdominant. Similar work was done by Kristofik et al. (1995) on arthropods in • Penduline Tit (Remiz pendu/inus (L.» nests, where spiders and flies were most abundant. Krumpâl et al. (1995) focussed on the different species ofticks present in birds' nests
along with their distribution throughout the year in Slovakia. A similar study was done by
Kristofik et al. (1996) to investigate the arthroPOd composition of Bee-ealer (Merops
apiaster L.) nests, which consist ofground burrows, in Slovakia. The most abundant
ectoparasites were feather lice and the ectoparasitic fly Carnus hemapterus Nitzsch,
followed by mesostigmatid mites, ticks and beetles.
1wasa et al. (1995) studied the Diptera fauna ofbirds' nests in Japan from 69 nests
of 13 different bird host species. Different fly species were associated with different bird
species, with sorne flies only occurring in nests ofspecific architecture.
There have also been sorne less comprehensive surveys ofarthroPOds in birds' • nests. Nolan (1955) inventoried the arthropods associated with a Prairie Warbler (Dendroica disc%r Vieillot) nest, and ahhough this account is detailed in sorne ways,
most specimens were identified to family only. Whitehead (1988) studied the arthropod
fauna ofa European Starling (Sturnus vu/garis L.) nest situated under the roofofa
building and not surprisingly, most ofthe nest fauna, especially the Coleoptera, was more
characteristic ofthe fauna associated with a building than with the birds themselves.
2.3. Arthropods Associated with Nestboxes
Birds' nests may he characterized by the environmental conditions that are present
in them. Woodroffe (1953) divided nests into two different kinds: sheltered or dry nests, • Il and those open to the elements, or wet nests. The arthropod fauna ofthese nests differs • from one another since nests unprotected from rain can become saturated with water and undergo rapid bacterial and fungal decomposition. Therefore, much ofthe arthropod
fauna observed in these nests will be associated with the decaying plant matter, bacteria
and fungi. Organic material in dry nests decomposes comparatively slowly, and thus the
community ofscavenging mites and insects found in these nests will differ from that in
the wet nests (Woodroffe, 1953). Nests in nestboxes are in the dry category because they
offer a sheltered area which gives arthropods a dry habitat. The insulating properties of
the nest material create an ideal environment for arthropods and facilitates their
overwintering, which May partly explain their high abundance in these nests (Darolova
and Schleicher, 1997).
Sorne taxa ofarthropods appear to he more frequently associated with nestboxes; • Krumpal et al. (1993) found that the dominant group ofarthropods in Sand Martin nests and in bird shelters are Siphonaptera (adults and larvae) which find optimum living
conditions in these types ofnests. This was also observed by Tripet and Richner (1997),
who round that the prevalence offleas is highest in hole-nesting passerines, particularly
the hole-oesting Paridae. Gther arthropod groups that Krumpâl et al. ( 1993) round to be
abundant in these types ofnests were Araneae, unidentified insect larvae, Collembola and
Hymenoptera. Krumpâl et al. (1993) believed that the presence ofthis last group is due to
suitable conditions for the development ofparasitic wasp species and for ants who find
suitable feeding conditions. McAtee (1927) found mostly Diptera and Coleoptera
specimens in nestboxes used by various bird species. • 12 Usually nestbox studies focus on one specifie group ofectoparasites, without • mentioning the rest of the arthropod fauna (Rohy et al., 1992; Richner et al., (993).This is particularly true ofthe parasitic blow fly Protocalliphora. Gold and Dahlsten (1983)
discussed the preference ofProtocalliphora blow flies for cavity-nesting passerines rather
than cup-nesters or open nests. Furthennore, Bennett and Whitworth (1992) suggested
that nests in nestboxes are ideal for parasitic blow flies because ofthe solid nature ofthe
substrate. Iwasa et al. (1995) found that the nests ofhole-nesting birds have, on average, a
greater abundance and diversity offlies than those ofnon hole-nesting birds;
Protocalliphora species, for example, were only found in the nests ofhole-nesting
passerines. Gold and Dahlsten (1989) suggested that the hosts ofProtocalliphora flies are
primarily cavity-nesting birds in forests.
• 2.4. Nest Arthropods Associated with House Sparrows Hicks (1953, 1959, 1962, 1971) listOO over 200 species ofinsects associated with
House Sparrows, with Coleoptera and Siphonaptera being the two most diverse and
frequently collected groups. In addition to fleas, there are several records ofother
ectoparasites such as Il species ofProtoca//iphora, the ectoparasitic fly Carnus
hemapte",s Nitzsch (Camidae) and chewing lice. Hicks also recorded two non-parasitic
flies that are usually found only in bird nests: Leptometopa /atipes (Meigen) (Milichiidae)
and lVeossos mary/andiclls Malloch (Heleomyzidae).
Woodroffe and Southgate ( (951) recorded large numbers ofthe tlea
Ceratophy/lus ga/linae in the nests ofHouse Sparrows in Great Britain, but very few flies • 13 and mites. They found several other species ofsaprophagous arthropods; large numbers • ofbrown house moths (Hofmannophila pseudopretella (Staint) (Oecophoridae) and tenebrionid beetle larvae were present, along with a few beetle larvae in the families
Dennestidae and Ptinidae.
Woodroffe (1953) found that House Sparrow nests supported high numbers of
Tinea co/umbariella (Wocke) (Tineidae) and Anthrenus verbasci (L.) (Dermestidae).
There were also a few individuals ofptinid, tenebrionid and dennestid beetles other than
A. verbasci, and ofthe fly Fannia canicuJaris (L.) (Fanniidae). The most abundant mites
present were scavengers such as GJycyphagus domesticus (Degeer)., Tyrog(vphus farinae
(Degeer) and Tyrophagus tenlliclavlls (Zachvatkin). Other mites present were Acaropsis
docta (Berlese) and Cllna.."Ca capreo/lls (Berlese).
• 2.5. Nest Artbropods Associated witb Tree Swallows There have been far fewer arthropods recorded from nests ofTree Swallows than
House Sparrows. Hicks (1953, 1959, 1962, 1911) recorded only three species of
Protoca/liphora blow flies: Protocal/iphora splendida Macquart, P. 5ialia (Shannon and
Dobroscky), and P. azurea (Fallén). The only other arthropod sPecies recorded by Hicks
were the fleas Ceratophy/lus idius (Jordan) and C. ga//inae.
Woodroffe (1953) notOO that although ProtocaJ/iphora flies were widely
distributed in bird nests, they appear to thrive particularly in those ofswallows. This was
also observed by Pinkowski (1977), and Bennett and Whitworth (1992) noticed that
Protoca/liphora flies were common in the nests ofTree Swallows and European Starlings • 14 (Sturnus vulgaris L.). Rendell and Verbeek (1996) also found that fleas, Protocal/iphora • sialia and the ectoparasitic mite Ornithonyssus sylviarom were the most abundant ectoparasites ofTree Swallows in the nests they studiOO.
Woodroffe (1953) notOO that dennestid beetles often reached moderate abundance
in swallow nests, along with Tinea columbariel/a and a few species ofscavengjng mites.
Burtt et al. (1991) found large numbers ofthe haematophagous mite Dermanyssus
hin,ndin;s, along with the scavenging mite Dermatophagoides evansi and the predaceous
mite Che/etomorpha /epidoplerum in nests ofTree Swallows using nestboxes.
•
• 15 3. MATERIALS AND METHODS • 3.1. Study Site Field work was conducted at the Macdonald Campus Fann ofMcGill University
in Ste-Anne-de-Bellevue, Quebec (45°24.69'N, 73°56.59'W) during the summers of 1997
and 1998. The fann is composed ofseveral fann and storage buildings along with
cottages and is surrounded by pasture and corn fields.
New wooden nestboxes. 25 cm high, 10 cm deep and 10 cm wide were used in
this study. Twenty nestboxes were placed at a height ofapproximately 2 fi, tàcing east,
on telephone poles near the faon buildings in 1997. Nestboxes were emptied ofold Dest
matenal and thoroughly cleaned using a bleach solution before the 1998 field season and
10 additional identical nestboxes were added on telephone poles in the same area for the • second field season. 3.2. Collection of Nest Fauna
Ali nestboxes were checked every third day tor signs of nesling by hosts and to
observe the progression ofegg laying and incubation. The nest matenal was removed
from the nestboxes al least 17 days after the eggs had hatched, to ensure that ail nestlings
had fledged. In cases where the nestlings were still present after 17 days, nestboxes were
visited every day until the nestlings tledged and the nest material was collected the first
day after the birds left the nests. Observations on nestlings were done in accordance with
CCAC guidelines and under an Animal Care pennit from McGill University.
When dead nestlings were found in the nest material, the nest was omitted from • 16 the inventory and analysis. This was done because dead nestlings have a rapid and major • effect on the composition ofthe arthropod fauna in the nest; a wide variety ofcarrion feeding arthropods are attracted to the eareass by olfactory eues, often within a few hours
ofdeath. The nest material was discarded ifthe nestling appeared to have been dead for
more than a few hours. The approximate time ofdeath was estimated primarily on the
basis ofthe overall apPearanee ofthe nestling; discolouration ofthe skin and r;gor mortis
begin within a few hours after death. [n addition, the presence ofblow fly or flesh fly
maggots other than Protocalliphora larvae on dead nestlings was an indication that
tèmale carrion-feeding flies had enough time to deposit eggs on the carcass.
The nest material was placed in plastic bags and sealed for transportation to the
laboratory. [n 1997, the nest material was placed in a Berlese funnel with a 60 watt
lightbulb as a heat and light source for a minimum of48 hours and then observed under a • dissecting microscope to remove any remaining arthropods and inactive stages such as Diptera puparia. This step was necessary because large numbers ofarthropods, especially
mites, remained in the nest material even after several days in the funnels. Because of
this, ail ofthe nest material was removed from the Berlese funnel, preserved in 700/0
ethanol and later examined under a dissecting microscope. This Pennitted the removal of
ail arthropods, including inactive stages sueh as pupae and puparia and those that had
died before the nest material was collected. Although this method was labour intensive, it
provided a more complete inventory ofthe nest fauna. In 1998, the plastic bags
containing the nest malerial were placed in a refrigerator to decrease arthropod activity so
that Diptera puparia could be removed for rearing before preserving the nest material in • 17 70% ethanol. The preserved nest material was then placed into sealed plastic containers • for later sorting under a dissecting microscope. After the arthropods were removed, the nest material was placed in a large graduated cylinder and the volume ofnest materiai
was measured.
Because adult flies are much easier to identify than immature stages, Diptera
puparia were placed individually in pHI boules to allow them to complete the life cycle to
the adult stage. Adults that emerged were kept alive in the pHI bo,tle for two days to
allow body structures to become fully sclerotized and pigmented and were then preserved
in 70~/o ethanol. Ali arthropod specimens removed from nest material were preserved in
70% ethanol and later prepared for identification using appropriate techniques. Most adult
insects were dried and mounted on pins or points; fleas and scale insects were kept in
ethanol and cleared using warm 85 % lactic acid for identification. Immature insects and • ail mites were kept in 700/0 ethanol. Ali arthroPOds collected have been deposited in the collection ofthe Lyman
Entomological Museum, McGill University.
3.3.. Diversity and Abundance ofArthropod Nest Fauna
Prevalence ofarthroPOd taxa in the nests ofeach bird species was calculated by
dividing the number ofnests in which a specifie arthroPOd taxon was present by the total
number ofnests ofthat bird species. The prevalence was expressed as a percentage. The
intensity ofa given arthropod taxon present in infested nests was calculated by taking the
mean number ofspecimens ofthat taxon in infested nests ofa given host species. The • 18 intensity ofarthropod taxa was expressed as the mean ± standard deviation. When • arthropod abundance and diversity were compared between bird species, data from 1997 and 1998 were pooled together for each bird species.
The arthropod diversity ofnests ofHouse Sparrows and Tree Swallows was
analysed by calculating the species richness and three different indices ofdiversity for
each nest, with each nest representing an independent sample.
Species richness is defined as the number ofspecies present in a sample (Hayek
and Buzas, 1997). Magurran (1988) considered species richness an extremely useful
measure ofdiversity although obviously the relative abundance and population structure
ofthe fauna are not taken into consideration.
There are many diversity indices described and used in the literature, but almost
none are universally accepted. Furthennore, those that are widely accepted are not always • appropriate for the sampies to which they are sometimes applied (Magurran, 1988). The Shannon index ofdiversity Is an example ofsuch a popular index. The Shannon index
takes ioto account both the number ofspecies as weil as the species abundance, and
makes no assumptions upon the underlying distribution ofthe data (Hayek and Buzas,
1997). This index assumes that individuals are randomly sampled from an infinitely large
population (Pielou, 1975). However, in cases where the randomness ofa sample cannot
be guaranteed, or ifthe community is completely censussed with every individual
accounted for, the BriUouin index is more appropriate (Pielou, 1975; Magurran, 1988). In
my study, ail samples (nests) were taken from a specific site, and ail samples present in
the community were completely sampled, with aH arthropods in the nest accounted for. • 19 Another reason that the Brillouin index was preferred in my study is that the Shannon • index does not deal weil with sampies that contain many rare species; for example, ifthe value ofthe Shannon index is known for a specific sample, the addition ofmany rare
species to the sample does not change the value ofthe Shannon index appreciably. My
samples had a large number ofrare species represented by only one or a few individuals.
The Brillouin index (HB) was calculated as follows (Magurran, 1988):
HB = ln N!- L Innd N
where N is the total number ofindividuals recorded and ni is the number ofindividuals of
the ith species. As diversity increases, the value ofthe Brillouin index increases. • The second index ofdiversity used was the Simpson index, which is one of the most widely used dominance indices. A dominance index is weighted towards the most
abundant species in the sample while being less sensitive to species richness (Magurran,
1988). The value ofthe Simpson index (D) indicates the probability ofany two
individuals drawn at random from a community belongjng to ditTerent species, and thus
estimates to what point the community is dominated by a few sPeCies or ifthe abundance
ofspecimens is equally distributed among them. The Simpson index was calculated as
follows (Magurran, 1988):
D = L ( ni(ni - 1) ) N(N -1)
• 20 where ni is the number ofindividuals ofthe ith species and N is the total number of • individuals recorded. A higher value ofthe Simpson index indicates that most specimens in the sample belong to a few dominant species, and that the evenness and overall
diversity ofthe sample are low.
The third index ofdiversity used was the Morisita-Hom index, which measures
Seta diversity, or the degree ofsimilarity between two sites. Deta diversity is a measure
ofhow different a range ofhabitats or samples are in tenns ofthe variety ofspecies found
in them. The fewer species that the different communities share, the higher the Seta
diversity will be (Magurran, 1988). Although there are several indices that measure Beta
diversity, the Morisita-Hom index is one ofthe most satisfactory indicators (Wolda,
1981 ; Smith, (986) and was chosen because it is not intluenced by species richness and
sample size. A modified fonnula for the index (Wolda, 1983) that is less sensitive to the
• most abundant species was used in my analysis. The Morisita-Hom index (C.\(ff) was calculated as follows:
2I (ani x bni) C,"ffl = ---==------(da + db)aN x bN
where aN= the total number ofindividuals in site A and bN = the total number of
individuals in site B. Ifthe sites are completely similar, the Beta diversity is low and the
value ofthe Morisita-Horn index is equal to 1; in cases ofcomplete dissimilarity, the
Seta diversity is high and the value ofthe Morisita-Hom index is equal to O. • 21 The Brillouin, Simpson and Morisita-Hom indices were ail calculated by hand. • The distributions ofthe species richness value and indices were first tested for normality with a Kolmogorov-Smimov test, and because they did not follow a nonnal distribution,
a Kruskal-Wallis test was performed to detennine ifthere was a difference between the
species richness and the indices ofdiversity ofthe nest fauna ofHouse Sparrows and Tree
Swallows.
An Unweighted Average Cluster Analysis (SAS Institute, 1988) was used to
examine the grouping ofspecies richness and diversity indices ofnests within and
between bird species. Cluster analysis is reliable whether the data follow a nonnal curve
or not, and is an exploratory method for grouping data based on their similarity. The
cluster analysis was used to group nests within the same bird species based on the values
oftheir species richness and diversity indices. An unweighted method was used because • it assigns the same weight to each data point, on the assumption that the samples or communities are equally represented in the data. This was the case in my study since each
ofthe nests was only sampled once. The same cluster analysis was perfonned on ail nests
ofboth host species pooled to observe ifnests ofthe same bird species had similar
diversity and would therefore cluster together.
ln addition to the cluster analysis, the nest fauna within and between bird species
was also compared using the abundance ofdominant arthropods in nests. An arthropod
species was considered dominant ifthe individuals ofthe species constituted more than
20/0 ofthe total number ofarthropods. A Kolmogorov-Smimov test was first perfonned to
determine nonnality ofthe data; since the data were not nonnally distributed, a Kruskal- • 22 Wallis test (significance level set at 0.05) was used to compare the nest fauna ofHouse • Sparrows and Tree Swallows. The volume ofnest material was measured to detennine ifthere was a difference
in nest volume between bird sPecies. A Kolmogorov-Smimov test was used to detennine
whether nest volume followed a nonnal distribution and a Kruskal-Wallis test was used
to compare the nest volume between bird species. A Speannan 9 s correlation analysis was
perfonned to sec ifthere was a significant relationship between nest volume and
abundance ofarthropods.
•
• 23 4. RESULTS • 4.1. Nesting Sueeess of Host Birds There were nine breeding atternpts in 1997, ofwhich eight nests were successful
(six House Sparrow, two Tree Swallow). Thirteen of23 breeding attempts were
successful in 1998 (nine House Sparrow, four Tree Swallow). Overall, 21 nests (15
House Sparrow, six Tree Swallow) were successful during the course ofthis study.
4.2. Inventory of Artbropod Nest Fauna
A total of90,729 arthropods was extracted from the nests., with a mean intensity
of5349.6 ± 6511.51 (n = (5) per House Sparrow nest and 1555 ± 2227.49 (n = 7) per
Tree Swallow nest over both field seasons. At least 71 species (50 families) of 13 insect
orders and al least Il species (8 families) ofmites were extracted from the nests and • identified. The prevalence and abundance ofanhropod taxa identified in House Sparrow nests are given in Table 1 and those identitied in Tree Swallow nests are given in Table 2.
Complete data on the abundance ofarthropod species in individual nests are given in
Appendix 1 (House Sparrows) and 2 (Tree Swallows).
There was no significant ditTerence in the abundance ofarthropod taxa between
1997 and 1998 within each bird species, unless otherwise indicated in the text. Because of
this, the abundance data for each arthropod spccies were pooled over both years for each
host species in the tables and text. The mean and standard deviation given for each
arthropod sPeCies are the pooled values for both years.
• 24 4.2.1. Arthropod Divenity: Bouse Sparrow Nests • Araneae. Two spiders were found in the nests, one in 1997 and one in 1998. The specimens were in poor condition and could not be identified.
Acarina. Seven families ofmites were identified: Parasitidae; Macrochelidae;
Dennanyssidae; Uropodidae; Urodinychidae; Diplogyniidae; and Oribatulidae.
The most abundant family was the Dennanyssidae; the total number ofspecimens
from ail other mite families combined represented only 0.16% ofthe total mite
population. Dermanyssus hirundinis was the only species ofdermanyssid present. It was
found in ail nests and was the most abundant arthropod in the nests during both tield
seasons. Its abundance varied greatly between nests, from 38 to 21,351 specimens, with a • mean of 5037.6 ± 6718.77 D. hirondinis per nest.
Dermaptera. The ooly species ofearwig in the nest material was the European earwig
(Forficlila auricularia L.). The mean number ofearwigs per nest was 31.57 ± 31.6.
Homoptera. One species ofleathopper (family Cicadellidae), an unidentified species of
Euscelis Brullé, was present with 2 specimens in 1997 and 8 in 1998. There was also an
unidentified species ofscale insect (superfamily Coccoidea) present with a mean number
of 15.13 ± 19.64 per nest.
Coleoptera. Thirteen families ofbeetles were identified. Forty-eight beetles were • 25 collected in House Sparrow nests in 1997, and 124 in 1998. The proportion oflarvae was • higher in 1997 (66.7%) than in 1998 (2.4%). There was a Mean of4.24 ± 7.66 beetles Per nest. There were significantly more specimens ofthe genus AJeochara Gravenhorst
(Staphylinidae) in 1998 than in 1997 (p = 0.04). The thirteen families belonged to one of
three ecological guilds; the number in brackets indicates the total number ofindividuals
found in nests ofHouse Sparrows. The first group includes predaceous beetles:
Staphylinidae (67), Histeridae (30) and Cucujidae (1). The second category inc1udes
saprophagous species" feeding on plant and animal detritus and dung: Dennestidae (34)"
Scarabaeidae (8), Nitidulidae (4), Hydrophilidae (3), Anobiidae ( 1), Ptinidae (1) and
Cryptophagidae ( 1). The third category includes phytophagous families which feed on
live vegetation: Elateridae (3), Chrysomelidae (1) and Curculionidae (1).
• Siphonaptera. Fleas \vere the most abundant insect order in House Sparrow nests in
1997~ 893 specimens were found with a mean of 178.6 ± 156.55 per nest (Il ± 10.8
% adults per nest). Most specimens (95 ) were immature stages (1arvae and pupae) which
are scavengers in the nest material and do not feed on blood. There were 245 specimens
ofSiphonaptera in 1998, with a mean of30.62 ± 33.75 specimens per nest (7 ± 7.29
adults) and 83% were immature. Although the abundance was greater in 1997, the
diftèrence was not significant. Two sPecies were identified: CeratophyJ/us ga//inae and
Ceratoph.vllus idius. Only 1 adult specimen of C. idius was present. Adult tleas were
identified to species using published keys, and larvae were identified by association with
adults in nests where only one species ofadult tlea was identified. • 26 • Diptera. When both years were pooled together, the Mean number offlies per nest was 48.60 ± 184.4. Although there were noticeable differences in abundance from 1997 to
1998, with 29.6 ± 17.8 flies per nest in 1997 and 296.7 ± 422.99 in 1998, the di fference
was not significant (probably because ofthe high standard deviation and small sample
size). Flies were the most abundant insects in the nests in 1998. Most specimens offlies
(67% in 1997,990/0 in 1998) were immature stages (larvae and puparia). The Diptera was
also the Most diverse insect order in House Sparrow nests with 17 families and 21 species
for both years combined. The flies can be divided into three categories: families with
species that are closely associated with birds' nests; families that are saprophagous in a
range ofdecaying organic materials; and families that are apparently accidentai visitors.
Four families identified contain species that are closely associated with birds' • nests: Milichiidae; Carnidae; Heleomyzidae; and Calliphoridae. Leptometopa latipes was the only species of Milichiidae identified. Although
most Milichiidae are associated with a wide range ofhabitats, this species is usually
associated with birds' nests. It is rarely collected and titde is known of its biology. Nine
specimens were collected in House Sparrow nests, tour in 1997 and five in 1998.
The ectoparasitic fly Camus hemaptenls was the only species ofCarnidae
identified. Two specimens were collected in House Sparrow nests. The feeding habits of
adults ofthis species are unclear; they feed either on skin secretions or on the blood ofthe
nestlings. The larvae are scavengers (Grimaldi, 1997). This is the tirst record ofCamus
hemaptenls in Quebec. • 27 Neossos marylandicus was the only species of Heleomyzidae collected. Very little • is known ofthe biology ofthis fly, and specimens are very rare in museum collections. In 1997, this was the second most abundant fly species in the nests, with 34 specimens, ail
adults, representing 230/0 ofthe total fly fauna. In 1998, this species was the most
abundant fly with 2265 specimens including 942 larvae, 1302 puparia and 21 adults; this
represented 850/0 ofthe eolire fly fauna. The mean number ofN. marylandicus per nest
was 176.85 ± 365.03.
Protoca/liphora siaUa was the only species ofCalliphoridae present in the nests.
Thirty-three larvae and 98 puparia ofthis ectoparasitic species were collected. No adults
were collected in the nest malerial and no adults emerged from puparia taken to the lab
for rearing. Protocal/iphora sia/ia was the most abundant tly in the nests in 1997 with a
total of90 specimens (640/0 offly abundance) and a Mean of 18 ± 7.2 specimens per nest. • It was the third most abundant fly in 1998 with 41 specimens (1.50/0 oftotal fly
abundance) and a mean of 10.3 ± 6.5 specimens per nest. There was a significant
difference in the abundance ofP. sia/ia specimens between 1997 and 1998 (p = 0.04).
Most ofthe remaining fly families (10) are saprophagous and their presence in the
nests is not unusual because these nests have a lot oforganic material such as grass, dead
insects, bird feces, etc. in various stages ofdecay. The abundance ofspecies in these
tàmilies was low except for one Dest which contained 252 larvae ofScatopsidae.
Three families (Culicidae, Chironomidae, Dolichopodidae) are neither
saprophagous nor closely associated with bird nests. These flies were probably accidentai
visitors into the nests. Mosquitoes (Culicidae) were probably attracted by the nestlings • 28 and entered the nests to take a blood meal. Adult chironomids do not feed and • Dolichopodidae are predators ofother insects; both ofthese families may have been brought into the nest as food items by the birds.
Lepidoptera. The mean number of Lepidoptera specimens was 4.8 ± 4.98 per nest. Most
specimens (70%) were identified as a single species, Niditineafusce/la (L.) (Tineidae).
Other specimens were in poor condition and could not be identified.
Hymenoptera. Hymenoptera identified inc1uded parasitoid wasps and ants. Ail
parasitoids (12 in 1997, 5 in (998) in the nests were a single species, Nasonia vitripennis
(Walker) (Pteromalidae), which is a parasitoid ofProtoca//iphora puparia. Several tly
puparia were taken to the laboratory for rearing in 1998 but no Nasonia emerged from • these puparia. The ants (Fonnicidae) belonged to at least two genera, CamponotZls Mayr and Lasills Fabricius. Other genera were apparently present, but the specimens were
damaged and could not be identified. Carpenter ants (Campono/us species) are usually
found in decaying wood; members ofthe genus Lasius are known to tend cultures of
aphids and scale insects, and feed on their anal secretions. The ants were probably
accidentai visitors in the nests.
Other insect orders. Two other insect orders were present in very small numbers in
House Sparrow nests. One specimen ofPsocoptera was found, but its poor condition
prevented identification. Psocoptera are usually found in detritus. One specimen ofthe • 29 family Pentatornidae (Order Hemiptera) was collected; most rnembers ofthis farnily feed • on plants, although sorne species are predators.
4.2.2. Arthropod Divenity: Tree Swallow Nests
Acarina. Dennanyssidae and Macronyssidae were the only mite families identified ..
although there was one specimen ofanother family whose condition prevented
identification.
Dermanyssus hirondinis was the only species ofdennanyssid present, and
Ornithonyss"s sy/via11lm was the only species of Macronyssidae. Dermanyssus
hintndinis was the most abundant arthropod in the nests during both field seasons and
was present in ail nests except one, in which it was replaced by 1016 specimens of
Ornithon.vssus sy/viantm, an ectoparasitic mite ofsimilar habits. The abundance ofD.
• hiJ1tndinis ranged from 12 to 6427 specimens per nest, with a mean of 1348.67 ± 2538.52.
Dermaptera. Earwigs (Forficula auricularia) were much less abundant in 1997 (1
specimen) than in 1998 (22 specimens). Earwigs were only present in two nests in 1998.
Psocoptera. There was a total of38 Psocoptera (barklice), ail in a single nest. Ali
specimt:ns belonged to an unidentified species ofthe family Liposcelidae. Members of
this family are often associated with the nests ofbirds and mammals.
• 30 Phtbiraptera. Two specimens ofchewing lice were collected in the nests. Identification • beyond order was prevented by the poor condition ofthe specimens. These permanent ectoparasites usually do not leave the body ofthe adult bird, except to colonize the
nestlings.
Homoptera. The Cicadellidae was the only family present in the nests. The mean number
ofcicadellids per nest was 18 ± 25.65. Most specimens (141 of 144) were ElIsce/is sp.,
and the remaining three specimens were identified as Lala/lls ocellaris DeLong and
Sleesman. CicadllJa cype,.acea Osborn and one species whose condition prevented
identitication.
Coleoptera. Four tàmilies ofheetles were found in the nests. Staphylinidae (6 specimens) • and Histeridae (3) are predators, Dennestidae (3) are usually scavengers on dead plant material and Curculionidae (1) feed on live plants. Five specimens were not identified
because oftheir poor condition. There were fewer Coleoptera per nest in 1997 (3 in one
Dest) than in 1998 (15) and aIl specimens were adults.
Siphonaptera. Fleas were the most abundant iDsect order in 1997. Most specimens were
immature stages (980/0 in 1997; 81 % in 1998). There were significantly more specimens
per nest in 1997 (484 ± 627.91) than in 1998 (5.5 ± 4.95) (p = 0.04). The Mean number of
adult fleas was 2.0 in 1997 and 9.5 ± 2.12 in 1998. Two species ofadult fleas were
identified: CeralophY//lls gallinae and Ceralophy/lus idius. The great majority (95%) of • 31 adults were identified as C. idius. Larvae found in nests where only one species ofadult • tlea was identified were assumed to belong to the same species.
Diptera. Diptera was the Most abundant insect order in 1998, with 73.6 ± 66.12
specimens per nest. However, the mean number of ft ies per nest was 12.1 ± 31.67 when
both years were combined. The majority ofspecimens were immature stages (81 % in
1997, 91 % in (998), such as larvae and puparia. Diptera was also the most diverse order
with 18 families including at least 19 genera for both years. Among the families closely
associated with birds' nests, Camidae, Heleomyzidae and Calliphoridae were present.
One specimen ofCamus hemaplen~s was collected.
Neossos marylandicus was the second most abundant tly in the nests in both field
seasons, with 13 specimens in 1997 and 14 in 1998, representing 10% and 40/0 ofthe total • fly l'auna occurring in Tree Swallow nests. The mean number ofN. marylandicus per nest
for both years was 5.4 ± 2.61.
The only species ofCalliphoridae was Protocalliphora sialia. Over both years,
2571arvae, 177 puparia, and one adult were collected and a further ten adults were reared
l'rom puparia kept in the laboratory. This was the most abundant tly in the nests in both
1997 (104 specimens) and 1998 (331 specimens), which represented 81 % and 90 % of
the total fly abundance in Tree Swallow nests for each year, respectively. The mean
number per nest was 62.14 ± 56.08.
Ofthe 13 other families present in the nests, seven were saprophagous and six
were considered accidentai visitors. The six families that were probably accidentai • 32 visitors include predators ofinsects (Empididae and Dolichopodidae), biting flies • (Culicidae and Simuliidae) and families normally associated with aquatic habitats (Chironomidae and Platystomatidae). The abundance ofthese 13 families was low
compared to the Heleomyzidae and Calliphoridae with al most three specimens in a11
nests during each field season.
Lepidoptera. Most ofthe Lepidoptera (42 of52 specimens) were identified as Nidilinea
fllscella; the condition ofthe other larvae prevented their identification. Almost ail
Lepidoptera (50 specimens) were co11ected in 1998, and the Mean number of Lepidoptera
per nest was 8.67 ± 14.87.
Hymenoptera. Two tàmilies ofparasitoid Hymenoptera were identified: Pteromalidae • and Ichneumonidae. Ali pteromalid wasps identified were Nasonia vitripennis, which parasitizes Protoca//iphora puparia. Because ail puparia were preserved in 70% ethanol
in 1997, it was impossible to detennine the rate ofparasitism ofProtocalliphora by N.
vitripennis. However, in 1998 puparia were placed in emergence containers for rearing.
Twenty-four Prolocal/iphora puparia (120/0) were parasitized and a Mean of25.29 ±
15.71 individuals ofNasonia vitripennis emerged from parasitized puparia. The
ichneumonid wasp was not identifiable.
Otber insect orders. Four other insect orders occurred in sma11 numbers in the nests.
One specimen ofthe family Miridae (Hemiptera) was present. These are mostly plant • 33 feeders, but sorne are predators. There were also a few specimens ofEphemeroptera (2), • Neuroptera (1) and Trichoptera (8) that might have originally been carried inside the nestboxes as food items by the birds and fallen into the nest material.
4.3. Comparison ofArthropod Nest Fauna Within Bird Species
4.3.1. Bouse Sparrow Nests
There were few significant differences in arthroPOd diversity and abundance
between 1997 and 1998. The number ofarthropod species in House Sparrow nests varied
from 5 to 17 species per nest over both field seasons. The Brillouin index varied from
0.13 to 1.86, and the Simpson index varied from 1.05 to 6.02 when both tield seasons
were pooled together. There were no significant differences in the species richness and in
the indices ofdiversity between 1997 and 1998. In the cluster analyses ofspecies richness • (Figure 1), Simpson's index (Figure 2) and Brillouin's index (Figure 3), there were no c1ear divisions ofnests according to year, number ofnestlings or any other observed
factor. In tenns ofabundance, there were significantly more Prolocal/iphora sia/ia blow
nies in nests in 1997 than in 1998 (p = 0.04) and significantly more staphylinids ofthe
genus Aleochara in 1998 (p = 0.04).
Other observed differences from 1997 to 1998 such as higher numbers offleas
and Dennaptera in 1997, and ofectoparasitic mites, Neossos marylandiclis and
Coleoptera in 1998 were not significant, but this may be partly due to the small sample
size and high standard deviation ofthe samples. The abundance ofthe other arthropod
groups was consistent between years. • 34 • 4.3.2. Tree SwaUow Nests The number ofarthropod species present in Tree Swallow nests varied from 5 to
24 over both field seasons. The Brillouin index varied from 0.27 to 1.71, and the Simpson
index varied from 1.12 to 4.21. The cluster analyses ofspecies richness (Figure 4),
Simpson's index (Figure 5) and Brillouin's index (Figure 6) did not show anY trends, or
any separation ofnests according to year, number ofnestlings or other observed factors.
There were more Coleoptera species in 1998 than in 1997, although this
difference was not significant. The number ofother species remained similar between
years. Other than the Siphonaptera, there were no significant differences in the abundance
and diversity ofarthropods between years. The abundance ofthe remaining groups was • similar for both years. 4.4. Comparison of Arthropod Nest Fauns Between Bird Species
There were no significant differences between the species richness and diversity
indices of House Sparrows and Tree Swallows. When cluster analyses were perfonned on
the species richness (Figure 7), Simpson's index (Figure 8) and Brillouin's index (Figure
9) for ail nests ofboth species, the nests did not cluster according to bird species.
In contrast to the overall results, the value ofthe Morisita..Hom index was 0.05,
which indicates high Beta diversity (Iow similarity) between the nest fauna ofthe two
host species. This low similarity was due to several differences at the species level in the
arthropod fauna between House Sparrows and Tree Swallows. • 35 Although the total number ofarthropod species was not significantly different • between nests ofeach bird species (p = 0.83), there were significantly more species of Coleoptera in House Sparrow nests than in Tree Swallow nests (p =0.02). There were
significantly more total arthropod specimens in nests ofHouse Sparrows than in nests of
Tree Swallows (p = 0.04) and there was a significant difference in beetle abundance, with
the majority ofColeoptera specimens (910/0) in House Sparrow nests (p:::: 0.05).
The presence or absence, and mean abundance ofdominant arthropod species
(species whose individuals comprise more than 2% ofthe overall arthropod fauna) in
nests ofboth hosts are shown in Table 3; the mean abundance is compared in Figure 10.
There were several significant differences in the mean abundance ofdominant species
between House Sparrow and Tree Swallow nests; House Sparrow nests had significantly
higher numbers ofD. hinlndinis mites (p = 0.02) and C. gal/inae fleas (p = 0.01) than • Trec Swallow nests. There were signiticantly more specimens ofProtocal/iphora sia/ia (p = 0.0007), C. idills tleas (p = 0.03) and leathoppers in the genus Eusce/is (p = 0.01) in
Tree Swallow nests than in House Sparrow nests. Sorne ofthe dominant species were
only found in nests ofa single host species: scale insects were only present in House
Sparrow nests; O. sylviarnm mites were present only in one Tree Swallow nest.
The volume ofeight nests (four ofeach bird species) was measured and
3 compared; the mean nest volume for House Sparrow nests (1258.19 ± 278.49 cm ) was
3 aimost twice as large as nests ofTree Swallows (636.17 ± 213.47 cm ); this difference
was significant (p = 0.02). When a Spearman's correlation was performed on the nests of
each species separately, there was no significant correlation between the total Dest volume • 36 ofeither bird species and the abundance ofmites, fleas, P. sialia. N. marylandicus. or • total nest fauna. However, when House Sparrow and Tree Swallow nests were pooled together, there was a significant negative correlation between the abundance ofP. sialia
and nest volume (0 = 8, r = -0.72, P =0.05), meaning that nests with a greater volume had
fewer Protocal/iphora larvae.
•
• 37 5. DISCUSSION • 5.1. Nesting Sucees! of Host Birds In 1998, birds began breeding in early April, but due to the lack offood and short
periods ofcold weather, many nestlings died and several nests failed. Ail observed
nestIing deaths were attributed to cold weather, lack offood and subsequent nest
abandonment by the parents, except for one nest that was heavily parasitized by
Protocalliphora sia/ia larva. Adverse environmental conditions are probably the reason
that the nesling success of House Sparrows was lower in 1998 than in 1997. The oesling
success ofTree Swallows was higher in 1998, and this is probably because Tree
Swallows staft breeding later in the season, and therefore are less subject to seasonal
variations. There were probably more breeding attempts in 1998 because birds had time • to recognize the surrounding nestboxes erected in 1997. 5.2. Inventory of Arthropod Nest Fauna
The abundance and diversity ofnest arthropod fauna in my study were much
higher than in many other studies. However, it is often difficult to compare the absolute
values for arthropod diversity and abundance between different studies because of
differences in the host species examined, the type ofnest constructed, the numher ofnests
examined, and the age ofthe nests. Nests collected years or months after their
construction may contain more sPecimens and species or a ditTerent community
composition ofarthropods than those collected only a few weeks after their construction.
Ondrejkova et al. (1991) found 38.,817 arthropods (Acari excluded) in 217 nests used by • 38 five bird species and three mammal species, collected over four years. This gives a mean • of 178.8 insects per nest, which is lower than the Mean number ofinsects (287.3) in my nests, which were ail new. Hood and Welch (1980) found a mean of508 specimens per
Red-winged 81ackbird nest. Philips and Dindal ( 1990) found 22,991 arthropods in one
Screech Owl (Otus as;o (L.» nest and 26,553 invertebrates in a single American Kestrel
(Fa/co sparverius L.) nest, which are much higher numbers than those from my study.
However, these raptor nests were much larger than passerine nests and had been reused
for several years, and thus the arthropod population had time to increase in abundance
and change in composition.
The number ofinsect orders found in my study (13) was higher than in Many
previous studies; other authors have recorded between four and eight orders in nests of
single bird species (McAtee, 1927; Hood and Welch, 1980; Whitehead, 1988; Philips and • Dindal, 1990; Ondrejkovâ et al., 1991; Kristofik et al., 1994). The high abundance ofinsect orders and specimens in my nests May be due to
several factors, such as nest architecture and extraction method. The abundance ofcertain
arthropod taxa in nestboxes can be explained by the structure ofthe nestbox itself. For
example, earwigs were very abundant in my nests compared to other studies (e.g., Hood
and Welch, 1980; Whitehead, 1988; Kristofik et al., 1995) and this is probably because
nestboxes offer a shaded and dry environment. 1found hundreds ofearwigs in empty
nestboxes at my study site. This May have prevented the occupation ofthose nestboxes
by birds because these nestboxes were not used once the earwigs were abundant.
The low abundance ofspiders was unexpected, because most studies report • 39 greater numbers. Gold and Dahlsten (1989) reported spiders building webs in nestboxes • and feeding on emerging adult Protocalliphora. K.ristofik et al. (1996) identified 16 species ofspiders in their nests, and 15 in other studies (Kristofik et aL, 1994, 1995).
Although new, the nestboxes in this study had a lot ofinsects that could have been used
as prey for spiders. As with other groups ofarthropods, the spiders may not have had
sufficient time to colonize the new nestboxes. The studies by Kristofik et al. (1994, 1995)
examined nests that had been established for severa) years.
The overall mite abundance and diversity observed in my study was lower than in
other studies. This is probably because mites were newly established in the nestboxes;
ovcr time the diversity and abundance ofmites might increase. Members ofthe family
Parasitidae, Urodinychidae and Macrochelidae are predaceous on other mites and small
arthropods (E. Lindquist, pers. comm.); however, these mites were not abundant enough • in both years to have had an impact on the rest ofthe mite fauna. The Uropodidae also occur in a wide range ofhabitats and the family includes predators, scavengers and
detritivores, including species associated with bird nests and rodent galleries (Hirschman,
(972). The Diplogyniidae are common on passalid and histerid beetles in the New World
(KIantz, 1978), and were probably associated with the histerid beetles which were
numerous in the nests. Several ofthe Diplogyniidae specimens found in the nest material
were deutonymphs that attach themselves phoretically to the cuticle ofother animais
(Krantz, (978). Species ofOribatulidae feed on detritus in a wide range ofhabitats.
The abundance and diversity ofthese non-ectoparasitic mites was lower than
those reported by Burtt et al. (1991) who found thousands ofmites ofdifferent species • 40 (predators and scavengers) in the same nests. Krîstofik et al. (1996) reported 40 species • and 133,016 specimens ofmesostigmatic mites from 174 Bee-eater nests. The abundance ofectoparasitic mites in my study was generally lower than in
other studies. Pacejka et al. (1996) looked at Dermanyssus hirundinis population growth
from nests which were originally free ofmites. Twelve days after the inoculation ofthese
nests with mites, they found an average of21, 703 ± 5623.2 mites per nest. Not only are
these populations greater than those found in my study, they also demonstrate the
colonizing capabilities ofthese mites. Merino and Potti (1995) observed a mean of5280
DermanysslIs gallinoides Moss in highly parasitized nests. Johnson and Albrecht (1993)
reported hundreds to thousands ofmites per nestling in House Wren nests and Burtt et al.
( 1991) also found a high intensity ofD. hirundinis (9745 ± 978) in Tree Swallow nests.
The only other ectoparasitic mite species in my nests was Ornithonyssus sy/vianlm which • occurred only in one nest. The abundance ofcicadellids in the nests was much higher than in other studies,
and was concentrated in Tree Swallow nests. Hood and Welch (1980) found several
specimens in Red-winged Blackbird nests. The high abundance ofleatboppers in the
swallow nests might be explained by the composition ofthe nests. In 1997, when the
abundance ofleathoppers was highest, Tree Swallows started nest construction later
during the season, and lined their nest with green vegetation, esPecially grasses.
Cicadellids are herbivores which feed on live vegetation and were probably brought into
the nestboxes with their host plants. In 1998, the vegetation was quite dry when the
swallows constructed their nests and the abunda..'1ce ofcicadellids was much lower. Scale • 41 insects found in House Sparrow nests were obviously brought into the nestbox with their • host plants, because ail specimens were adult females which are sessile insect pennanently attached to their host plant. Because House Sparrows use more dried grass
and twigs in their nests, and because seale insects tend to be more abundant on twigs and
older grass stems and leaves, the higher numbers ofscale insects in House Sparrow nests
was not unexpected.
The number ofColeoptera families found in this study (13 families) is similar to
that found in previous studies. Hood and Welch (1980) found 7 families and 14 species.
Whitehead (1988) identified Il Coleoptera families in a starling's nest and Kristofik et al.
(1995) found a total of 10 families including 14 species ofColeoptera in Penduline Tit
nests. Kristofik et al. (1996) found that the number ofspecies and the abunùance of
Coleoptera in Bee-eaters nests increased every year the nests were used, eventually • reaching 25 species (15 fanlilies) and 2982 specimens. The Coleoptera families found in my study were also found in several other studies on bird nest fauna (Woodroffe, 1953~
Hood and Welch, 1980; Kristotik et al., 1994, 1996). They nonnally occur in detritus so
it was not surprising to find them in nest matenal, because nests represent an excellent
source offood for the beetles. Staphylinids and histerids are often associated with bird
nests (Woodroffe, 1953; Kristofik et al., 1994, 1996). Kristofik et al. (1994, 1996) found
that staphylinids were the most abundant and diverse group in the nests ofSand Martins
and Bee-eaters. Dermestid beetles were found in nests examined by WoodrotTe (1953)
and Kristofik et al. (1994, 1996). The beetles were probably attracted to the nests by the
odour ofthe decaying vegetation. • 42 The species diversity ofSiphonaptera in the nestboxes was similar to other studies • but the abundance ofadult fleas was comparatively low. Brown and Brown (1986) found up to 39 adult fleas per nest ofCliffSwallows, and Richner et al. (1993) found an average
of37 adults per Great Tit nest. Harper et al. (1992) cited several examples ofhighly
infested nests (up to 3793 tleas per nest) and Rheinwald (1968) identified 5754 C.
gallinae in the nest ofa double-brooded Coal Tit (Panls ater L.). The lower abundance of
tleas observed in my nests is probably due to the fact that they were first year nests. Fleas
overwinter in the nest material and thus their populations can increase in successive
breeding seasons.
Flies were the most abundant insects present in the nests ofboth bird species~
those with the greatest overall abundance were N. marylandicus and P. s;a/ia~ followed
by the Scatopsidae. Other than N. malylandicus and P. sialia, L. latipes and C. • hemaptenls are closcly associated with bird nests. The milichiid Leptomelopa /alipes has been reported from birds' nests (Ryckman,
1953) and large numbers have been collected from American Kestrel and Screech Owl
nests (Philips and Dindal, 1990). The species has also been reared from dung (Ferrar,
1987). The low numbers found in my nests might indicate that L. lalipes is in the early
stages ofcolonizing the nestboxes, or that the source population for colonization is very
low.
The abundance ofCarnus hemaplenls in my study was much lower than in
previous records. Kirkpatrick and Colvin (1989) found up to 22 C. hemaptenls per
nestling, and these flies apparently did not have any effect on the nestlings. Kristofik et • 43 al. (1996) found that C. hemaplerus was the only ectoparasitic fly encountered in nests of • Bee-eaters in Slovakia; nestlings had a Mean of7.7 specimens each~ and no detrimental effect on the nestlings was reported. Cannings (1986) found between 2 to 50 flies per visit
at nests ofSaw-whet Owls (Aegolius acadicus (Gmelin». He suspected that the death of
nestlings he observed was related to C. hemaplerus infestation, although he admitted that
the evidence was circumstantial. The difference in the abundance ofthis fly observed in
this study compared to other studies might be due to the fact that the nestboxes were
newly established and the flies had just begun to colonize them. AIso, most studies on
Carnus have been made in forested areas, and they may be more abundant in such
habitats. There might be a preference for large open nests such as those made by birds of
prey, but there is still too little ecological information on these flies to reach any
conclusions. • This is the tirst time that Neossos marylandicus has been reported in such large numbers, and this is also the tirst study in which immature stages have been identified.
Although the few known specimens ofN. malylandicus have been associated with birds'
nests (McAtee, 1927; Ferrar, (987), my study is the tirst to make a more definite
connection between the feeding habits ofthe larvae and the nests. The high abundance of
immature stages in my study suggests that the larvae are saprophagous in the nest
material, feeding on decayjng organic material.
The abundance ofP. sialia in nests was similar to that reported by other workers
(e.g., Johnson and Albrecht~ 1993; Matsuoka, 1997). The highest abundance in a single
nest in my study was 182 larvae, followed by another nest with 62 specimens. The • 44 intensity ofinfestation found in my study was much lower than that reported by • Whitworth and Bennett (1992) who found up to 600 larvae per Black-billed Magpie (Pica pica,(Sabine» nest, and 200 per Bank Swallow (Riparia riparia (L.» nest. while Roby et
al. (1992) found an average of 116 larvae per infested Eastern Bluebird nest.
It was not surprising to find several saprophagous fly families inside the nests;
these nests constitute ideal environments for larvae and adults who feed on detritus,
because nests in nestboxes are a sheltered environment containing a lot oforganic
material. In addition to the fly species mentioned above. several families of flies
identified in rny study have been reported in other studies. These are common tàmilies
such as Bibionidae. Scatopsidae, Chironomidae, Sphaeroceridae and Muscidae. Sorne
families, such as Empididae and Dolichopodidae are predators which rnay have fol1owed
other flics into the nestboxes. However, the majority offamilies were distinct from other • studies and rnay reflect the immediate environment ofthe nestboxes (e.g., the proximity ofaquatic habitats). Bloodfeeding flics such as mosquitoes and black flies were not found
in high numbers in the nestboxes, probably because they only enter the nestbox long
enough to feed on the birds and then leave.
The overall Diptera diversity at the family level was higher in my study (24
families) than in others. Nolan (1955) found four Diptera families in a Prairie Warbler's
nest. Hood and Welch (1980) found eight families in Red-winged Blackbird nests. [wasa
et al. ( (995) found eight families and 23 species of Diptera in 69 nests of 13 bird species.
Kristofik et al. (1995) found seven families and Il species offlies in 357 Penduline Tit
nests. Kristofik et al. (1996) identified eight families in 174 Bee-eater nests. One reason • 45 for the difference in diversity May be the collection techniques used. Hood and Welch • ( 1980) and Kristofik et al. (1995, 1996) used Tulgren funnels to colleet arthropods from the nest material; like Berlese funnels, Tulgren funnels do not pennit the collection of
inactive stages and dead specimens, and this might explain why the arthropod diversity in
those studies was lower than that in mine. 1wasa et al. (1995) used emergence bags placed
over the entrance ofthe nest and collected ail adult flies emerging from il. This method
does not allow the collection ofimmature or inactive stages and dead specimens.
Although there may be a biological reason for the higher diversity offlies in my study,
the most likely explanation is that the method ofextraction which 1used (i.e., microscope
examination ofpreserved nest material) allowed the extraction ofdead and immature
specimens which otherwise would have not been collected.
The only Lepidoptera that could be identified was the tineid moth Niditinea • fusce/la. Tineid moths are often found in birds' nests (Woodroffe, 1953; Nolan, 1955; Hood and Welch, 1980; Philips and Dindal, 1990) and probably feed on fungi or dead
organic matter found in the nests.
Il was not surprising to find the parasitoid wasp Nasonia vitripennis in nestboxes
because ofthe presence ofProtocal/iphora puparia. Parasitism by Nasonia species would
not have a significant effect on Protocal/iphora populations in the short term because
only the puparia are attacked. However, parasitism by Nasonia species may have a long
tenn effect in decreasing the local population ofadult Protocalliphora. The Mean number
ofNasonia per parasitized puparium found in my study was higher than that found by
Gold and Dahlsten (1989) who found between 15 and 20 Nasonia per puparium. • 46 Ants were the second most abundant Hymenoptera in the nest material after • Nasonia. Ants are often encountered in bird nests, especially in nestboxes with old nest material (C. Riley, personal observation). Nolan (1955) and Philips and Dindal (1990)
found ants in open nests, and Gold and Dahlsten (1989) noticed Carpenter ants making
tunnels in nest material. Their presence is probably due to the decaying organic material
and the sheltered habitat present in nestboxes.
Based on my data, it appears that the tirst year nest fauna in a nestbox is a
composite of: ectoparasites brought in by the birds; insects that are in the immediate
nestbox environment and colonize the nest themselves; and prey items that are lost in the
nest material. The high abundance ofavian ectoparasites (haematophagous mites, tleas)
during the initial stages ofnest construction and occupation was also observed by
Woodroffe and Southgate (1951) and Woodroffe (1953). Insects round in the immediate • environment, such as cicadellids and earwigs, can be brought inside the nest with materials used for construction, or enter the nestboxes on their own. Some possible
examples offallen prey items are the Ephemeroptera, Trichoptera and sorne Coleoptera
specimens.
S.3. Comparison of Artbropod Nest Faun. Within Bird SPfties
Nests within each host species were similar in the composition oftheir arthropod
fauna. There were sorne variations in species richness and diversity indices within each
bird species, but none ofthese were significant. This supports the hypothesis stated in my
second objective: nests ofthe same bird species have similar nest tàuna, al least during • 47 the first year. This was expected because the nests were first year nests ofthe same bird • species in identical nestboxes. The composition ofthe nests inside the nestboxes was similar, as Woodrotfe (1953) found in another study, and thus it is not surprising to see
no significant ditferences in their arthropod diversity.
First year nests are probably similar because the invasion sequence ofthe nest
fauna occurs in a similar fashion from one nest to another. Nests are colonized first by
arthropods brought into the nest by the bird host; theretore ectoparasites such as fleas,
haematophagous mites and other ectoparasites found on the bird will be among the
earliest colonizers. These will he followed by arthroPOds found in the incoming nest
material, such as cicadellids, scale insects and sorne Coleoptera. Once the nest
construction is completed, organic decay will probably reach the point where the matenal
attracls arthropods associated with detntus, such as scavengers. Given the variety of • decaying organic materials present in the nest (fresh and dried plant tissues, feces, feathers, dead arthropods), a variety ofphytophagous and saprophagous arthropods will
be able to exploit different niches in the nest. Once the ectoparasites and scavengers are
present in sufficient numbers, predators will colonize the nest. Fallen prey items will only
appear in the nest fauna once the eggs are present. The actual rate at which this
colonization will proceed will vary from one nest to another.
Although the same trophic categories ofarthropods are usually present in all
nests, the actual species composing them May be different from one nest to another.
Ectoparasites will vary according to their preferred host; for example, the tlea C. idius
was dominant in Tree Swallow nests while C. gallinae was more abundant in House • 48 Sparrow nests. The same was also observed for phytophagous species: cicadellids were • mostly found in Tree Swallow nests while scale insects were found in House Sparrow nests. The saprophagous fl y N. mary/alldicus was more abundant in sparrow than in
swallow nests. Although there were a few predaceous species ofbeetles in the nests, it
appears that by the time the nest collection was made in this case the predator community
had not yet had time to increase. This lack ofpredators in the arthropod fauna offirst year
nests was also observed by McAtee (1927).
Even though 010st ofthe nest fauna was similar, there were sorne variations in the
arthropod composition ofthe nests. These variations, whether in the number ofspecies or
ofspecimens, may be partly due to a variety ofexternal factors such as the habits and the
degree ofparasitism ofa specifie bird, yearly variations in environmental conditions and
the diversity and abundance ofthe arthropod fauna present in the immediate vicinity of • individual nestboxes. This last factor will not only influence what kind ofarthropod will get to the nest by their own means or by attachment to the Dest material, it will also
influence the type ofprey taken into the nestbox. The collection date ofthe nest material
may be a factor because many arthropod species are more active and abundant only
during a specifie time period in the summer. For example, the greater abundance of
Neossos mary/andicus in 1998 Olay be due to earlier colonization ofthe nest by the flies,
giving females the opportunity to oviposit and the larvae to mature before the collection
ofthe nest material. Another possibility is that the collection time ofthe nest coincided
with the seasonal peak ofabundance ofN. mary/andicus and this is why larvae and
puparia were only present during the second field season; this could also explain the • 49 extreme variation in the number ofspecimens found in different nests. • This variation observed in arthropod species and their abundance may also be dependent on internai factors, such as interspecific predation and interspecific
competition. Because ofthe small number ofpredators collected in the nest material, it
appears that they did not have a major effect on the structure ofthe nest fauna. However,
there is competition between arthropod species inside the microhabitat constituted by the
nestbox. This microhabitat is made up ofseveral ecological niches which are available to
the nest fauna, and the species which fill a particular niche in a nest May be those which
colonize and become established first, in the absence ofcomPetitors. This May explain
why, for example, there were so Many specimens ofN. marylandicus in sorne nests while
the species was absent in others; ifspecimens ofN. marylandiclls became established in a
particular nest and were able to substantially increase their population, then other small • saprophagous flies that are dependent on the same food source, such as milichiid or scatopsid flies would have little available resources in that nestbox, and would not be able
to expand their own population. This May also explain the scarcity ofC. hemapleros in
my nests compared to other studies; perhaps the access ofthis fly to the nestlings was
prevented by the high abundance ofectoparasitic species such as P. sialia and D.
hin,ndinis. 1suggest that one ofthe principal causes for the great variation in abundance
ofarthropod species from nest to nest in my study is interspecific competition.
Furthennore, this variation might be more pronounced in first year nests, because ail
species present in the nest are new colonizers.
• 50 5.4. ComparisoD of Arthropod Nest Fauns Between Bird Species • Although there were sorne significant differences in the abundance and diversity ofspecifie arthropod taxa, the overall diversity and abundance in House Sparrow and
Tree Swallow nests were not significantly different from each other. Therefore, the
hypothesis stated in my third objective was not supported: nests ofdifferent bird species
do not have a significantly different total nest fauna, especially during the first year.
The overalliack ofsignificant differences between host species may be an artifact
ofthe small sample size, or, as in the comparison within bird species, it may indicate that
the tàuna offirst year nests reflects the arthroPOd community composition ofthe
envirornnent immediately surrounding the nest and the degree ofparasitism ofeach bird,
regardless ofbird species. The arthropod abundance and diversity found in these nests
appear to be typical ofbirds' nests, because their composition is similar to that found in • other studies, except for the high abundance of Homoptera, Neossos marylandiclls and the absence ofspiders.
5.4.1. House Sparrow Nests
The significantly higher numbers ofmites, fleas and Coleoptera and the presence
ofscale insects in House Sparrow nests could be explained by the ditTerent nest
architecture ofthe two bird species, since nests ofswallows and sparrows differ in their
composition. House Sparrows completely fill the nestboxes with dried grass and twigs,
and incorporate feathers from other species along with a variety ofman-made materials
such as plastic and string. Their nest is usually loosely woven, creating volume and space, • 51 and is quite dusty which, according to Woodroffe (1953), could he important for mites • and immature stages ofinsects. Tree Swallows, on the other hand, use comparatively little material, mostly fresh and dried grass and small pieces ofwood. They usually line
the nest with feathers, which cao be their own or belong to other bird species. Their nests
are smalt, dense and thick, with very little space in which arthropods cao move. Also~
nests ofHouse Sparrows were significantly larger thao Tree Swallow nests in this study.
Mites, fleas and beetles May prefer the layered nests of House Sparrows where
there is a lot ofspace within the nest material. Ectoparasitic mites aggregate inside the
nest material placed at the roofofthe nestbox during the day, and move down to feed on
the nestlings at night. The dried grass at the top ofthe nestbox would offer the mites extra
space and easier access to the nestlings; this access is more difficult in Tree Swallow
nests because no malerial is placed above the nestlings. Since nestlings are the mites' • main source oftood and main limiting factor, easier access would Mean an increase in the mite population, which is known to increase exponentially. Also, the greater volume of
nest material in House Sparrow nests means a greater amount oforganic material as a
food source for saprophagous species such as tlea larvae and some species ofColeoptera.
Rendell and Verbeek (1996) found a positive correlation between the nest volume and the
number ofmites and t1eas, but this relation was not observed by Burtt et al. (1991). There
was no relationship between nest volume and the abudance ofmites, fleas or total
arthropods in my study.
Another factor that may explain the higher abundance oftleas is the behaviour of
the bird hosto For example, House Sparrows sPend comparatively more lime on the • S2 ground than Tree Swallows, which feed mostly on the wing. Therefore, House Sparrows • would be more accessible to adult fleas, which often rernain on the ground or low vegetation until a host is available.
The composition and volume ofHouse Sparrow nests could also explain why
more species ofColeoptera were present since the loosely woven nest material provides
beetles and earwigs (which usually gather in corners ofthe nestboxes) with shelter and a
large source offood.
The presence ofscale insects in sparrow nests may be explained by the large
amount ofdead plant material in the nestboxes; sessile female scale insects rernain
attached to the host plant, and will be brought inside the nest along with il.
5.4.2. Tree Swallow Nests • The presence ofcicadellids in Tree Swallow nests can be explained by the material used in nest building; cicadellids feed on fresh plant material, which is more
abundant in Tree Swallow nests, and they will, therefore, be brought in along with the
nest material.
The reason for the significantly greater abundance ofProtocaJ/iphora larvae in
Tree Swallow nests is unclear. Matsuoka et al. (1997) suggested that the degree of
parasitism by Prolocalliphora is dePerldent on the height ofthe nest, but ail nestboxes in
my study were at the same height. Sabrosky and Bennett (1958) suggested that most
Protoca/liphora show habitat or ecological preference rather than host preference or host
specificity. However, they did note a preference for P. sialia to parasitize Tree Swallow • 53 and Common Grackle (Quiscalus quiscula (L» nests. This was also observed by • Woodroffe (1953) for swallow nests. Because ofthis apparent host preference, Gold and Dahlsten ( (989) suggested that the nest volume and the odour and CO2 produced by the
nestlings were the two principal factors attracting Prolocal/iphora adults. Gold and
Dahlsten ( (989) also found that the number ofProtocal/iphora larvae in the nest was
positively correlated with nest volume; they suggested that the greater nest volume wouId
prevent contact between the larva and host feces, reduce exposure to low temperatures
and decrease detection by the parents who could remove them from the nest. My results
do not agree with those ofGodl and Dahlsten (1989); 1 round a strong negative
correlation between the nest volume and the number ofProtocallipltora larvae present.
Nests with more larvae (~53 larvae per nest) generally had a lower volume than others
J (s 1016 cm ). Furthennore, ifthere was a correlation between nest volume and • Prolocalliphora abundance, 1wouId expect House Sparrows to have higher numbers of tlies, because oftheir significantly larger nests. This was not the case.
Whitworth (1976) suggested that because blood is one ofthe limiting factors for
these tlies, more nestlings would mean a larger population ofProtocal/iphora. This might
be true with mites or fleas or any other ectoparasites whose adults are present in the nest
at ail time and feed on resources available in the nests. However, Prolocal/iphora females
laya certain number ofeggs and depart from the nest, so at most only the larvae born in
the nest will be present. More feeding space will not increase the total number oflarvae;
only a female ovipositing new eggs will.
A final possibility is that adult House Sparrows williocate Prolocalliphora • 54 larvae in the nestboxes and kill them, thus decreasing the abundance ofthis fly in their • nests. This was observed in nests ofBell Miners (Manorina melallophrys Latham) where females and nests attendants remove ectoparasitic arthropods (Clarke and Robertson,
1994). However, House Sparrows have not been reported exhibiting this behaviour.
1suggest that the greater numbers ofProlocal/iphora larvae in Tree Swallow
nests may be the result ofdifferences in host behaviour. Tree Swallows have a longer
incubation period ( 13.14 ± 1.46 days) and nestling period (about 15 days) than House
Sparrows (10.47 ± 0.99 and 12 days, respectively) which would provide flies with a
longer period oftime to find and parasitize the swallow nestlings. Tree Swallows also
have a more conspicuous nesting behaviour; they are very territorial, and there is one
parent near the nest al ail times. House Sparrow behaviour 1S more discreet, the female
being very secretive. The higher visibility ofthe swallows might have facilitated the • location ofthe nests by the flies. There probably were more C. idills fleas in Tree Swallow nests than in House
Sparrow nests because swallows are the natural host associated with these fleas.
s.s. Conclusion
My study has shown that the arthropod fauna ofbird nests may be much more
diverse than most studies ofsingle ectoparasite taxa would lead us to believe. Several
species often considered "rare", such as Neossos marylandicus, can be found in high
numbers in appropriate habitats. 1have also demonstrated that the arthropod fauna of
first year nests ofdifferent bird species is not significantly different from hast to hast, • 55 although the presence and abundance ofparticular arthropod taxa cao vary greatly, • probably as a result ofthe early colonization ofthe new nests. 1also have shown that the method ofarthropod extraction that 1have used May be
much more efficient at extracting the arthropods present in the nest material than
traditional methods like Berlese funnel extraction or emergence cages.
There are sorne logical further studies that would contribute to an understanding
ofthe diversity and ecology ofthe arthropod nest fauna ofnestboxes. Sampling the nest
fauna in subsequent years would allow an analysis ofthe competition and succession
between colonizing species from first year nests. Rather than retrieving the nests at the
end ofthe breeding season, leaving them tor the arthropods to overwinter and/or
recolonize the following spring might result in different patterns ofdiversity in the fauna.
Studies ofspecies abundance within specifie trophic guilds, such as ectoparasites or • scavengers, trom year to year could give an indication ofthe importance ofinterspecific competition in determining the structure ofthe nest community. Finally, sampling in
subsequent years might indicate whether unexpectedly abundant species such as Neossos
malylandicus remain dominant or whether my results were unusual.
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• 65 • • •
Table 1. Arthropod abundance in Bouse Sparrow nests. Dom. = dominant arthropod specics. OrderfFamily Gcnus/Spccics No. nests Prcvalence Range Total x ± s.d. Dom.
ARANEAE
Specics 1 1 7% 1 1 1
Specics 2 1 7% 1 1 1
ACARI
Parasitidae Para...ilus fimerolum 1 7% 1 1 1
Macrochelidae Ma,:roc/I1!/es nlIIscaedomeslicae 1 7% 1 1 1
Derrnanyssidae Dermanyssus hirundi"is 15 100% 38 - 21351 75564 5031.6 ± 6718.77 X
Uropodidae Uropoda sp. 2 13% 4-6 10 5 ± 1.41
0'1 Urodinychidae Uroobo\'ella sp. 1 7% 1 1 1 0'1 Diplogyniidae Diplogynïr.m sp. 1 7% 3 3 3
Oribatulidae Phuu/oppiu sp. 1 7% 1 1 1
Sche/oribulc!s sp. 3 20% 1 3 1
Species 1 6 40% 1·4 14 2.33 ± 1.37
Unknown mites 6 40% 1• J7 80 IJ.3 ± 13.72
DERMAPTERA
Forficulidae Fmficula uuriculuriu 7 47% 1-84 221 31.51 ± 31.6 X PSOCOPTERA
Species 1 2 13% 1-2 3 1.5 ± 0.71 • • •
Table 1. Continued.
Order/Family Genus/Species No. nests Prevalence Range Total x ± s.d. Dom.
HEMIPTERA
Pentatomidae Species 1 1 7% 1 1 1 HOMOPTERA
Cicadellidae Eusceli~' sp. 6 40% 1·4 10 1.67 ± 1.21 X
Coccoidea Species 1 8 1·57 121 15.13 ± 19.64 X COLEOPTERA
Staphylinidae A/eoc.:haru sp. 1 5 33% 1- 42 66 13.2 ± 16.36 X
A/eochara sp. 2 1 7% 1 1 1 0\ ...... a Hydrophilidae Sphaeridium sCllrabaeoide.'i 2 13% 1- 2 3 1.5 ± 0.71
Histeridae Species 1 4 27% 3·20 30 7.5 ± 8.35
Scarabaeidae Aphodius sp. 1 3 20% 1·4 8 2.67 ± 1.53
Elateridae Me/anotus sp. 3 20% 1 3 1
Denneslidae Dermestes /arJarill.f 2 13% 1·5 6 3 ± 2.83
Alicropepl.4S sp. 1 7% 1 1 1
Unknown denneslid larvae 3 20% 1·24 28 9.33 ± 12.74
Anobiidae Species 1 1 7% 1 1 1
Ptinidae Ptinus sp. 1 7% 1 1 1
NitiduHdae G/i.'~chn)(:',i/u.~ qlltld,.isignatwi 2 13% 2 4 2 • • •
Table 1. Continued.
Order/Family Gcnus/Specics No. ncsts Prevalence Range Total x ± s.d. Dom.
Cucujidae Species 1 1 7% 1 1 1 Cryptophagidae Species 1 1 7% 1 1 1
Chrysomelidae Species 1 1 7% 1 1 1
Curculionidae Species 1 1 7% 1 1 1
Unknown Coleoptera adult 5 33% 1- 5 Il 2.2 ± 1.64
Unknown Coleoptera larvae 4 27% 1- 2 7 1.75 ± 0.5 SIPHONAPTERA
CeratophyIIidae Ceratophyl/us gallinue 9 60% 12 - 440 1020 113.33 ± 137.47 X 0'1 00 Ceralophy/l,1S idius 1 7% 1 1 1 X
('erulOphyl/us larvae 3 20% 1- 68 117 39 ± 34.39 DIPTERA
Bibionidae Bibio sp. 1 7% 3 3 3
Psychodidae Species 1 1 7% 1 1 1
Scatopsidae St.:alOpj"U sp. 1 7% 1 1 1
Species 1 1 7% 252 252 252 X
Culicidae Species 1 1 7% 1 1 1
Chironomidae Species 1 1 7% 1 1 1
Dolichopodidae Species 1 1 7% 1 1 1 • • •
Table 1. Continued.
Order/Family Genus/Species No. nests Prevalence Range Total x ± s.d. Dom.
Syrphidae Species 1 1 7% 2 2 2
Milichiidae Leplomelopll Imipl's 4 27% 1- 4 9 2.25 ± 1.26
Camidae Carn..." hemaplenl.'i 2 13% 1 1 1
Sepsidae Sepsis sp. 1 7% 1 1 1
Themira sp. 1 7% 1 1 1
Unknown sepsid larvae 2 13% 1 2 1
Heleomyzidae NeoJJos mar)'landicuJ 13 87% 2· 1049 2299 176.85 ± 365.03 X
Sphaeroceridae Specics 1 1 7% 1 1 1 0\ \0 Chloropidae Species 1 1 7% 1 1 1
Anthomyiidae Species 1 1 7% 1 1 1
Muscidae MU.\·ca sp. 2 13% 1·3 4 2 ± 1.41
Species 1 2 13% 1·2 3 1.5 ± 0.71
Unknown muscid larvae 1 7% 40 40 40
Calliphoridae Prolocal/iphortl sia/ia 9 60% 4·28 131 14.56 ± 7.63 X
Unknown calliphorid larvae 1 7% 47 47 47
Unknown Diptera adult 3 20% 1·2 5 1.67 ± 0.58
Unknown Diptera larvae 6 40% 1- 5 10 1.67 ± 1.63 • ••
Table 1. Continued.
Order/Famil}' Genus/Species No. nests Prevalence Range Total x ± s.d. Dom.
LEPIDOPTERA
Tineidae Niditineu fu.ttcella 4 27% 5 - 18 34 8.5 ± 6.35 X
Unknown Lepidoptera larvae 6 40% 1- 5 14 2.33 ± 1.51
HYMENOPTERA
Ichneumonidae Species 1 1 7% 12 12 12
Pteromalidae Naso"i" vitripennis 5 33% 1 - Il 17 3.4 ± 4.34
Formicidae Cumpon()tus sp. 3 20% 1- 3 5 1.67 ± 1.15
Lusi,1.\' sp. 2 13% 1- 3 4 2 ± 1.41 ...... a o Species 1 2 13% 1 1 1
Unknown Hymenoptera adult 4 27% 1 - 2 5 1.25 ± 0.5 • • •
Table 2. Arthropod abundance in Tree Swallow nests. Dom. = dominant arthropod species. Order/Family Genus/Family No. nesls Prevalence Range Total x ± s.d. Dom.
ACARI
Dennanyssidae De,."umy.\'.'W.\' hil1l1u/ini.'i 6 86% 12·6427 8092 1348.67 ± 2538.5 X
Macronyssidae OrllilllO/~~'sj"ls .\)"l'iarum 1 14% 1016 1016 1016 X Unknown mites 1 14% 1 1 1 EPHEMEROPTERA Species 1 2 29% 1 2 1 DERMAPTERA
Forticulidae Forficula auricularia 3 43% 1· 19 23 7.67 ± 9.87 X
-..J PSOCOPTERA Liposcelidae Species 1 1 14% 38 38 38 PHTHIRAPTERA Species 1 1 14% 2 2 2 HEMIPTERA
Miridae Lygus sp. 1 14% 1 1 1
Species 1 1 14% 1 1 1
Unknown Hemiptera adult 1 14% 1 1 1
HOMOPTERA
Cicadellidae Cicudu/tl L:''1Ulrt.ll'('lI 1 14% 1 1 1 • • •
Table 2. Continued.
Order/Family Genus/Species No. nests Prevalence Range Total x ± s.d. Dom.
EU.'tcelis sp. 5 71% 4 -73 141 28.2 ± 28.37 X
Lululllj" ocel/aris 1 14% 1 1 1
Species 1 1 14% 1 1 1
NEUROPTERA
Species 1 1 14% 1 1 1
COLEOPTERA
Staphylinidae Species 1 1 14% 3 3 3
Species 2 1 14% 1 1 1 -.1 N Species 3 1 14% 2 2 2 Histeridae Species 1 1 14% 3 3 3
Dennestidae Trogodermu sp. 1 14% 3 3 3
Curculionidae Species 1 1 14% 1 1 1
Unknown Coleoptera adult 2 29% )-4 5 2.5 ± 2.12
SIPHONAPTERA
Ceratophyllidae Ceratophyllu.\· gll/linae 1 14% 8 8 8 X
Ceralophyl/wi idiuj" 3 43% 2 - 928 932 310.67 ± 534.63 X
Ceralophyl/u.'tlarvae 2 29% 9 - 30 39 13 ± 15.39 • • •
Table 2. Conlinued.
Order/Family GenuslSpc:cies 1 No. nesls Prcvalence Range Tolal x ± s.d. Dom.
DIPTERA
Mycelophilidae Species J 1 14% 1 1 1
Culicidae Species 1 1 14% 1 1 1
Simuliidae Species 1 1 14% 1 1 1
Chironomidae Species 1 1 14% 1 1 1
Stratiomyidae M;croL'hr)'~'a sp. 1 14% 1 1 1
Spccies 2 1 14% 1 1 1
Empididae P/atYPlIlplls Sp. 1 14% 1 1 1 .....,J w Dolichopodidae Cont/y/o.'îtY/lIs sp. 2 29% 1 2 1
Dolichopodinae species 1 1 14% 1 1 1
Species 1 4 570/0 1- 2 5 1.25 ± 0.5
Plalystomalidae Rivel/ia sp. 1 14% 1 1 1
Opomyzidae Geomy=tl triprmctata 1 14% 1 1 1
Milichiidae LeptomelOpll /utipes 1 14% 4 4 4
Carnidae Carnll~' hemaptenls 1 14% 1 1 1
Sepsidae Seps;.'î sp. 1 14% 1 1 1
Them;ra sp. 1 14% 1 1 1
lauxaniidae Species 1 1 14% 1 1 1 • • •
Table 2. Continued.
Order/Family Genus/Species No. nests Prcvalence Range Total x ± s.d. Dom.
Helcomyzidae NeoJ.m.'i mary/andicwi 5 71% 2-8 27 5.4 ± 2.61 X
Sphaeroceridae Ruc1li.\,/JOda /immw 1 14% 1 1 1
Spe/obiu sp. 1 14% 2 2 2 Specics 1 1 14% 1 1 1
Ephydridae Ephydrinae species 1 1 14% 1 1 1 Species 1 1 14% 1 1 1 Anthomyiidae Species 1 2 29% 1 2 1
Calliphoridae Prolocal/iphora sialitl 7 100% 14 - 182 435 62.14 ± 56.08 X -....1 ,&:l. Unknown Diptera adult 1 14% 1 1 1 TRICHOPTERA
Species 1 2 29% 1- 6 7 3.5 ± 3.54 LEPIDOPTERA
NidUineaJUscella 2 29% 3 - 39 42 21 ± 25.46 X Unknown Lepidoptera adult 1 14% 2 2 2
Unknown Lepidoptera larvae 3 43% 2-3 8 2.67 ± 0.58 HYMENOPTERA
Ichneumon idae Species 1 1 14% 1 1 1
Pteromalidac Na.wmiu 'Ii/ripe",,;... 2 29% 1- 3 4 2 ± 1.41
Unknown Hymenoptera adula 1 14% 1 1 1 Table 3. Abundance ofdominant arthropods in House Sparrow and Tree Swallow nests. N = number of nests infested; Diff. = significant difference (* - p < 0.05; ** -p < 0.005).
• House Sparrow Tree Swallow Diff. Species N x ± s.d. N x ± s.d. Dermanyssus hirundinis 15 5037.6 ± 6718.77 6 1348.67 ± 2538.5 * Ornilhon.vssus sy/viarum 0 1 1016
Forfieu/a aurieu/aria 7 3 LS7 ± 31.6 3 7.67 ± 9.87 Eusee/is sp. 6 1.67 ± 1.21 5 28.2 ± 28.37 * Coccoidea sp. 1 8 15.13 ± 19.64 0
Aleoehara sp.1 5 13.2 ± 16.36 0 CeralOphyllus gal/inae 9 113.33 ± 137.47 1 8 * Ceralophyllus idius 1 1 3 310.67 ± 534.63 * Scatopsidae sp. 1 1 252 0
Neo.'isos marylandieus 13 176.85 ± 365.03 5 5.4 ± 2.61 Proloeal/iphora sia/ia 9 14.56 ± 7.63 7 62.14 ± 56.08 ** • Nidilineafuseella 4 8.5 ± 6.35 2 21 ±25.46
• 75 • 1.25
1.00
0.75
• 0.50
0.25
0.00 -~ -~ -~ -~ 7 15 9 14 8 10 13 6 3 Il 2 4 5 12 Nests Figure 1. Cluster analysis ofspecies richness in House Sparrow nests in 1997 (nests 1-6) and 1998 (nests 7-15). Y axis represents the distance between clusters. • 76 • 1.25
1.00
0.75
• 0.50
0.25
0.00 7 6 4 13 l4 3 8 l2 9 II lO 2 5 15
Nests Figure 2. Cluster analysis ofSimpsoo's index in House Sparrow nests in 1997 (nests l-6) and 1998 (nests 7-l5). Y axis represents the distance between clusters. • 77 • 1.5
1.25
1.
• 0.i5
0.5
0.25
O. 7 4 6 8 10 15 9 II 2 5 12 13 14 3 Nests
Figure 3. Cluster analysis ofBrillouin's index in House Sparrow nests in 1997 (nests 1·6) and 1998 (nests 7-15). Y axis represents distance between clusters. • 78 • 2.00
1.50
• 1.00
0.5
0.00 7 6 3 5 4 2 Nests
Figure 4. Cluster analysis ofspecies richness in Tree Swallow nests in 1997 (1-2) and 1998 (3-7). Y axis equal distance between clusters. • 79 • 1.50
1.25
1.00
• 0.75 0.50
0.25
o. 1 1 7 6 5 3 4 2
Nests
Figure 5. Clusteranalysis ofSimpson's index in Tree Swallow nests in 1997 (1-2) and 1998 (3-7). Y axis represents distance between clusters. • 80 • 1.5
1.25
1.00
• 0.75 0.50
0.2
0.00 1 1 7 5 6 3 4 2
Nests
Figure 6. Cluster analysis ofBrillouin·s index in Tree Swallow nests in 1997 (1 2) and 1998 (3-7). Y axis represents distance belWeen clusters. • 81 • 2.50
2.
1.50
• 1.00
0.50
1 1
r-"-
0.00 _1000- -- -~ -- h7 hi5 t7 t6 t3 h9 hl4 t5 h8 hiO t4 hl3 h6 h3 t2 hll h2 h4 h5 hl2 hl tl
Nests
Figure 7. Cluster analysis ofspecies richness in House Sparrow and Tree Swallow nests (1997-1998). ft = House Sparrow nest, t =Tree Swallow nest. Y axis represents distance between clusters. • 82 • 2.00
1.50
• 1.00
0.50
0.00 h7 h6 h4 hl3 h14 hl h3 t2 t4 h9 tl t6 hll hlO t7 h5 h2 biS 15 13 h8 hl2
Nests
Figure 8. eluster analysis ofSimpson's index in House Sparrow and Tree Swallow nests (1997-1998). h = House Sparrow nest, t = Tree Swallow nest. Y axis represents distance between clusters. • 83 • 1.50
1.2
1.00
• 0.75
0.50
0.25
0.00 h7 h4 h6 h8 hlO hl5 t5 t7 h9 hll tl h2 t6 h5 t3 hl2 h13 hl4 hl h3 t4 t2
Nests
Figure 9. Cluster analysis ofBrillouin's index in House Sparrow and Tree Swallow nests (1997-1998). h = House Sparrow nest, t = Trec Swallow nest. Y axis represents distance between clusters. • 84 •
10000
1000 . --~--._------
!• House Sparrow_
-.5! i El Tree Swallow ca u Ut Cft ..2 -Gt 100 u 1: ca 'a 1: .a=' • C 10
1 derm scat neos gall fOff prat CQCC aieo niti eusc ami idiu
Figure 10. Mean abundance of dominant al1hropods in House Sp8rrow and Tree Swallow nests. derm - D. hirundinis; scat =scatopsidae sp. 1; neos =N. marytandieus; gall =C. gllllinlle ; fort =F. surieu/arill ; prat =P. sialia; cocc =CoccoïdH sp.1; aleo =A/eoehllfll sp.1; niti =N. fuseellll; eusc =Euseelis sp.; ami =O. sylvillrum; idiu =C. idius.
• 8S • • •
Appcndix 1. Arthropod spccics in nesls onlousc Sparro\\'s ~ 1997-19'JK).
Nest 1'1l\On 1 2 3 .. 5 6 7 K 9 10 Il 12 13 14 15 ARANEAE
Spccics 1 1
Specics 2 1
ACARI
Parasitidae
PartuÎlusJimerotum 1
Macrochclidac
MaCrtH:heles muscaedomestü'ae 1 00 0\ Dermanyssiduc
lJermtmyssus Irirundillis IHtIO 768 1765 661 4741 2351 21351 12356 1222 6705 38 625 1021 17749 2411
Uropodidae
Uropoda sp. .. 6 Urodinychidae
Uroobowl/a sp. 1 Diplogyniidae
OipJogJ'nium sp. 3
Oribatulidac
Phauloppitl sp. 1
.x'JIt!lorihtlles sp. 1 1 1 • • •
Appendix 1. Continued.
Nest Taxon 1 2 3 4 5 6 7 K 9 10 Il 12 13 14 15 Species 1 1 2 .. 2 1 4
Unknown mites 2 1 21 37 7 12 DERMAf)TERA
Forficulidae
ForJkula auricularia 84 46 5 1 53 31 1 PSOCOPTERA
Spech:s 1 1 2
JlEMIPTERA 00 -..) Pentatomidac
Species 1 1
HOMOPTERA
Cicadcllidae
Euscelis :!ip. 1 1 1 .. 2 1 Coccoidell
Species 1 1 8 1 3 24 25 2 57
cOtEOPTERA Slüphylinidae
AleochlITa sp. 1 1 8 42 7 8
Alelx:hllra sp. 2 1 • • •
Appendix l, Cuntinucd,
Nl."S1 Taxon 1 2 3 4 5 6 7 8 9 10 Il 12 13 14 15
Hydmphilidae
Sphaeridi"m .\'ctlrubueoitles 2 1
Histeridac
Species 1 J 3 20 4 Scardhacidac Aphodius sp. 1 " 3 1 Elaleridllc
Me/anolus sp. 1 1 1 00 OC Denncstidae
DermeJ'les lardarius 5 1
.\lit'r0l'eplus sp. 1
l1nknown denneslid larvue 1 3 24
Anohiidac
Species 1 1
P1inidae
PlÎnus sp. 1
Nilidulidac
GliJ(.'hrochiilu lIIUUlri!;iJ1:1JtJt"s 2 2
Cucujidllc • • •
Appcndix 1. Con'inu~J.
NLost Taxon 1 2 3 4 5 (, 7 8 9 lO II 12 13 14 15
Spœics 1 1
Cryptophagidac
Spedes 1 1
Chrysomelidae
Spccies 1 1
Curculionidae
Spccies 1 1
Unknown Coleoptcra aduh 2 1 1 CXl 2 S \0 Unknown Coleoptera larvae 2 2 2 1
SIPIIONAPTERA
Ccralophyllida~
CeralopIJylllls galli"ae 123 ....0 83 199 95 16 32 12 20
CeralOl,IJ.\'lIus idills 1
CerallJp'~~I/Ius tarvac 48 1 68
IJIPTERA
llibionidw:
8ibio sp. 3
Ilsychodidac
Spccic=s 1 1 • • •
Appendix 1. Conlinued.
Nesl Taxon 1 2 3 4 5 6 7 K 9 10 Il 12 13 14 15
Scatopsidac
s,..'UIOp.WI sp. 1
Spccies 1 252
CulicidllC
Specics 1 1
Chironomidac
Species 1 1
Uolichopodidac \0 o Specics 1 1 Syrphidac
Species 1 2
Milichiidac
Leplomelopa lalipes 1 4 2 2
Camidae
Camus hemaplenls 1 1
&.-psidac
Sepsi.\' sp. 1
Themimsp. 1
tJnknowll scpsid !arvac J 1 • ••
Appendix 1. Conlil1u~d,
NI."St Taxon 1 2 3 4 5 6 7 H 9 1() Il 12 13 14 15
Hclcorny;lidac
NI!(}.uo.\' nrury/andicl4!J' 3 l) 4 18 81 3 192 5 2 3 1049 928 2
Spha,..roccridac
Spt.'Cics 1 1
Chloropidac
Spt.'Cit.'S 1 1
Anthornyiidac
Spccil.'S 1 1 ..0 Muscidllc
Musca sp. 3 1
Specics 1 2 1
LJnknown rnuscid larvuc: 40
Calliphoridae
ProlOcalliphora sia/ia 10 20 12 20 28 6 13 18 4
Unknown calliphorid larvae 47
Unknown Dipll."fa "duit 1 2 2
Unknowl1 Diph..'ra l"I'Vae 1 5 1 1 1 1 • • •
Appcndix 1. Continucd.
Nesl Taxon 1 2 3 4 5 6 7 8 \) 10 Il 12 13 14 15 LEiJIDOPTERA
Tinciduc
NidilineaJUsL'ella 6 5 18 5
Unlmown I.cpidoptcra lurvuc 5 2 2 3 1 1
IIYMENOPTERA
Ichncumoniduc
S~cics 1 12
Iltcromillidilc 100 N Nusonia vitripe""is 1 II 1 1 3
Fonnicidac
ltlmpotlOlUS sp. J 1 1
IASius sp. 1 J
Spc:cies 1 1 1
Unknown lIymcnoplcra adult 2 1 1 1 Appendix 2. Anhropod species in nests ofTree Swallows (1997-1998).
Nest Taxon • 1 2 3 4 S 6 7 ACARI
Dcrmanyssidac
Demranyssus hinmdinis 1338 S8 147 110 12 6427
Macronyssidae
Ornithonyssus sylviarum 1016
Unknown mites 1
EPHEMEROPTERA
Specics 1 1 1
DERMAPTERA
Forticulidac
Forjicllia auricularia 1 3 19
PSOCOPTERA
Liposce! idae
Spccics 1 38 • PI-ITHIRAPTERA Spccics 1 2
HEMIPTERA
Miridac
Lygus sp. 1
Spccics 1 1
Unknown Hcmiptcm adult 1
HOMOPTERA
Cicadcllidae
Cicadu/a C}peracea 1
Eusce/is sp. 73 23 4 36 S
Lara/us ocellaris 1
Spccies 1 1
• 93 Appendix 2. Conrinued.
Nest Ta.xon • 1 2 3 5 6 7 " NEUROPTERA
Species 1 1
COLEOPTERA
Slaphylinidac
Spccics 1 3
Spccies 2 1
Spccic:s 3 2
Histcridac
Spc:cies 1 3
Dcrrnc:stidae
Trogoderma sp. 3
Curculionidae
Specics 1 1
Unknown Coleoptera aduh 1 4 • SIPHONAPTERA Ccratophyllidae
Ceratophyllus gal/inae 8
Ceralophylllls idius 2 928 2
Ceraloplryl/us larvae 30 9
DlPTERA
Mycctophilidae
Spccics t 1
Culicidae
Species 1 1
Simuliidae
Species 1 1
Chironomidae
Species 1 1
• 94 Appendix 2. Continued.
Nest Ta.xon • 1 2 3 4 5 6 7 Slraliomyidae
J/icrochrysa sp. 1
Spccics 2 1
Empididae
Plarypalpus sp. 1
Dol ichopodidae
ContJ.vlostylus sp. 1 1
Dolichopodinac sp. 1
Spccies 1 2 1 1 1
Plalystomatidae
Rive/lia sp. 1
Opomyzidac
Geom.v::a Iripunctala 1
Milichiidac • Leptomelopa lalipes 4 Camidae
Carnus he",aplenls 1
S~"sidae
Sep:l"is sp. 1
Themira sp. 1
Lauxaniidae
Spccies 1 1
Heleomyzidae
Neossos marylandicllS S 8 4 8 2
Sphaeroceridae
RaclJispoda Iimosa !
Spelobia sp. 2
Species 1 1
• 95 Appendix 2. Continued.
Nest Taxon • 1 2 3 4 S 6 7 Ephydridae
Ephydrinae sp. 1
Species 1 1
Anthomyiidae
Sp..-cics 1 1 1
Caiii phoridae
Protocalliphora sia/ia 62 42 61 182 53 21 14
Unknown Diptera adult 1
TRICf-IOPTERA
Spccics 1 6 1
LEPIDOPTERA
Tincidae
.\ïditineafusce//a 3 39
Unknown Lepidoptcra adult 2 • Unknown Lepidoptera larvac 2 3 3 I-IYMENOPTERA
Ichncumonidae
Spccics 1 1
Pteromalidae
.\'asonia \'itripennis 3 1
Unknown Hyrncnoptera adult 1
• 96