Text

Forwards Addl Matl to Support 901220 Amend Request for Field Study
	ML20070S088
Person / Time
Site:	Purdue University
Issue date:	03/04/1991
From:	Schweitzer J; PURDUE UNIV., WEST LAFAYETTE, IN
To:	; NRC OFFICE OF INSPECTION & ENFORCEMENT (IE REGION III)
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ML20070S086	List: ML20070S083; ML20070S088; ML20070S103;
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PURDUE UNIVERSITY f;p?g k/

r OEPARTMENT OF R AOiot OGIC AL AND ENVIRONMENT AL M AN AGEMENT s March 1991 U.S. 11uclear Regulatory Commission Materials Licensing Section 799 Roosevelt Road Glen Ellyn, Illinois 60137 Gentlemen:

Please find enclosed additional material to support our amendment request for a field study dated December 20, 1990. This material has also ' en submitted to Dr. Donna-Beth flowe, tiMSS, Washington.

The total amendment request should include:

Original Letter - Dated December 20, 1990 Attachment #2 - Doubly labeled water method (llagy)

Attachment #3 - Permission from 11WSCC Attachment #4 -

ISBli permission March 4, 1991 Letter Attachment #1A - Environmental Assessment (5 pages)

Attachment #2A - Background and map (11 pages) g Attachment #3A - Large map ,

If any of this material is missing or there are any questions, please contact me as soon as possible at (317)494-2350. Due to the anticipated beginning of the nesting perlsd in Mid-March, we would appreciate expedited handling of this request. Thank you in advance for this concideration of this matter.

Sincerely, 1

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mes F. Schweitzer, Ph.D.

Radiation Safety Officer JFS/gmt Enclosure 9104020127 cic318 PDR A D f"^ K O '5 0 0 0 1 8 2 P PDR Civit ENGINEERING BUILDING. B173

WEST LAFAYE T7E. (N .l7907 9

AT T A CilH ENT #IA ENVIRONMENTAL ASSESSMENT Identification of the Proposed Action The proposed action is to amend NRC Byproduct Material License No.

13-02812-04 1987.

issued to Purdue University and renewed on May The license authorites- use of byproduct material but 28, prohibits the release of material for field studies. The proposed amendment would authorize Purdue -personnel to inject tritiated water in approximately 80 eastern phoebes (Sayornis phoebe) . The birds will be located on U.S. Government Property at the Naval Weapons Support Center (NWSCC) in - Crane, IN. The study involves the release of tritiated water and water vapor directly into the environment. The proposed amendment would allow injection of tritiu.,into the birds as part of an ongoing study of this species.

Accordingly, pursuant to 10 CFR Saction 51.21 HRC has prepared this assessment to the resulting environmental impact.

The non-site-specific aspects of the study, i.e. possession, use and subsequent laboratory analysis of radioactive samples, waste disposal and health and safety aspects of use are performed under a

, broad scope license (13-02812-04) issued by the NRC.

Background

By letter dated December 20, 1990 (and attachments) purdue University (Purdue or applicant) Radiological and Environmental Management office requested an NRC amendment to perforn Hydrogen-3 labeled water (tritiated water) ~ studies on the eastern phoebe (Sayornis phoebe) in their natural habitat. These studies have L

received tentative approval - by the Indiana State Board of Health and NW5CC Radiation Safety office pending NRC approval.

The field study will extend over several months (March-August 1991) and may continue again in March-August 1992 if the data obtained in

,. -1991 is not' sufficient. The phoebes will -be . injected -with tritiated water, held until- the water equilibretes, sampled, and released into the environment. The birds will be recaptured and sampled 24 to -48 hours later to measure- body water turnover au relateu to energy-expenditure.

Purdue has been authorized by the U.S. Fish and Wildlife Servh:e under master permit 20439 to band the eastern phoebe with aluminum and plastic leg bands.

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0 Need for the Proposed Action The eastern phoebe (Sayornis phoebe) is a small double brooded bird weighing approximately 25 grams. As man changes the environment in which the bird lives, it must adapt to assure significant avopagation. Studies using tritiated water are intended to identify energy expenditure during the breeding period of the phoebe. Birds will be chosen to include those building new nests in pipe cuiverts, structures, and rocks and those utilizing existing nests in those features. The energy expenditures will be measured to relate this to the reproductive output of the phoebe using the six different study groups.

This study may enable researchers to identify factors that would allow the phoebe to propagate more efficiently.

Environmental Impacts of the ProDoa M Aotion The affected environment. The study site, Naval. Weapons-Support Center Crane (NWSCC), is located in southern Indiana in Martin County and consists of approximately 63,000 acres of land, 100 square miles. NWSCC consists of approximately 2,800 p'rmanent structures, the majority being explosive storage mattzines, serviced by 475 miles of roads, 175 miles of railroads, i 52 miles of cleared power lines. There are approximately- *~

residences on base along with a Bachelors Enlisted Quarters (BEQ) and Bachelor officer Quarters (B00), which are located near the residential area. There are approximately 130 residents on the base area. The NWSCC. is mostly forested land (75%) with maintained roadsides, power line right-of-ways, water lines, and-safety zones around storage bunkers. The surrounding off-base area is mainly farmland. Since NWSCC is an active ~ military base, access by the general public is severely limited. There are 4 main gates by which to access the facility is gained. They are I manned by security guardo and you must have a badge or visitor pass to obtain admittance. The perimeter is completely fenced and is patrolled along with the rest of the base on a 24 hour:

basis. The 4000 employees are not allowed to " freely" roam the l base outside the main industrial area. Approximately 100 employees travel on service roads and perform duties in the area outside the industrie.1 complex. The public may obtain -fishing -

permics for Lake Greenwood, but may not go anywhere else on base.

Due to the gulf crisis, at this time all cutdoor recreation activities have been suspended at NWSCC.

Nests will be chosen to avoid those l'o c a t e d n e a r a population center (greater than 20 houses). No nest will be chosen that is closer than 1.5 miles to a population center, including the base housing area. Also nc nest will be chosen that is closer than 0.5 miles to the fenced perimeter. - The nearest significant population center is to the northwest,-Crano Village (pop. 339) which is over 5 miles to the closest nest. There is no housing that would be located within the primary travel area of the bird which is a maximum of 2 acres (a circle with a radius of=

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approximately 250 feet). The closest residence is 0.8 miles from the closest nest (see large map).

Study subjects. The phoebes included in the tritiated water study will be adult females actively involved in nest building. Bir ds will be banded upon initial capture so that each bird may be identified upon subsequent recapture. Birds will be banded or identified to indicate that they contain radioactive material and will also have a U.S. Fish and Wildlife band attached.

Recapture of the birds will be accomplished by placing mist nets in the vicinity of the nest. While rearing young or incubating eggs, fidelity to the nest area is very high. Recapture rates of the female should exceed 75% during these periods. The rates for recapture may be somewhat less during the phases where eggs or young are not present. The recapture rate during these periods should exceed 50%. The half-life of tritium in the eastern phoebe is estimated to be approximately 0.5 days due to the high metabolic rate of the bird. Therefore the tritium would be eliminated from the bird in the period of approximately one week.

The range of these birds is very short-and it is not probable that any birds would range off the NWSCC property for significant periods while containing tritium. A typical home range for the birds during this period is 1 to 2 acres. Movements of large scales during the reproductive season are very rare. These large movements are usually only seen from year to year. The females (the only sex injected) spend nearly 90% of the time at the nest when eggs or young are present. Efforts will be made to choose subjects that inhabit the central portion of the facility to minimize any travel in unrestricted areas.

Study protocol. Oxygen-18, a naturally occurring non-radioactive isotope of oxygen, is added to tritium to produce tritiated water with a higher oxygen-18 level than found naturally in air (0.2%

l oxygen-18 naturally). This permits double-labeled studies on the blood samples and better quantification of body water turnover data. The oxygen-18 is activated to fluorine-18 in a cyclotron after the blood sample are collected and returned to Purdue. The oxygen-18 analysis will be done in an enclosed laboratory setting. This does not involve the use or release of by-product material at the study site and will not be discussed further.

The study protocol calls for initial injections of approximately 25 microcuries per bird in up to 80 birds. An injection of approximately 25 microcuries was chosen to minimize the amount of radioactive material used and maximize the probability that the blood sample would contain enough activity for purposes of counting statistics.

Each bird may receive up to 5 injections during each breeding !

cycle (there are two) for a total of 250 microcuries for the L _ . , . . _ __ , _ ._ _

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4 entire study period. Injections will be performed during nest building, after nest completion, during egg laying, during the incubation period and after egg hatching (rearing). The breeding cycles occur during March / April and June / July of each year. The maximum amount of tritium present in all birds at any one time would be ? millicuries. This assumes all birds are injected at the same time and all birds are recaptured. Recapture will not occur simultaneously due to variations in the breeding cycle and all birds will not be recaptured and injected for every period of the breeding cycle. There will be no recruitment of birds during the study unless the number of birds falls below eight-in any of the six study groups. The amount of tritium used should be significantly less than the maximum cited above.

Pathways to the environment. The phoebes lose tritiated water by respiration (exhaled water vapor) and direct losses in feces. It

- is thought that the most significant loss would be in water vapor in the exhaled breath. The average loss per day veries with the amount of tritium in the bird. The maximum loss is during the first day after injection and the amount eliminated in that day '

may be up to 18 microcuries. The total loss for all birds (assuming 80 birds) could be up to 1.5 millicuries per day. The primary study will take place from March 1991 to August 1991, however the study may continue in 1992 if data-collected is not sufficient. Assuming that every bird is recaptured and that each bird receives every possible injection the total release to the-environment per year would be 20 millicuries maximum. The total actual release is expected to be between 10 and 15 millicuries per-year.

Due to the dynemic environment in which the tritiated water is released the tritium will be distributed as water vapor, found!in streams, and deposited uniformly over the area.- Sun, wind, and rain will act to distribute the tritium sc that excess levels could not be measured above background.

Pathway.to humans.. Several factors such as security of the area, desire of birds to avoid contact with humans,-and remoteness of the area would minimize the chances of-human interaction with the-birds. Even if humans were in the area the releases are such that exposure would be below the limits in- 10CFR20, " Standards for Protection Against- Radiation" --limits for releases -to unrestricted areas, personnel on-the base area will-be notified regarding study and appropriate action to take if a bird is found.

The birds are small (<25 grams) and therefore lack food value.

- In the unlikely event a bird was consumed immediately af ter injection the whole body dose to the individual would be less ,

than 2 millirems. This dose is less than one percent of the maximum permissible exposure allowed-to a member of the general public in 10CFR20.105.

t The tritium released as tritiated water is expected to evaporate and mix with water and water vapor present in the environment.

Therefore concentration in any particular segment of the environment is unlikely. Entry into the food chain by consumption of plants or animals is unlikely to- result in-any measurable dose to the general public. ;

Personnel who handle the tritiated water, collect samples and inject birds have been provided with procedures and equipment to minimize their exposure to below 10rFR20 levels. Procedures include minimizing the release to the environment during routine and emergency situations.

Effects on other species. Other organisms, both plant and animal, in the area should not be affected by the release of tritiated water into the environment. The birds are not readily subject to capture by predators so no animal species should be

< affected. Mortality to the eastern phoebe occurs over the winter period where up to 50% _ may die. The birds will contain no tritium during the winter period. There are also no endangered species in the area that may be affected. -The low levels of tritium found in the environment should have no .ef fect on any species in the study area.

Dose and etfect on the eastern phoebe. The internal beta dt.3 from tritiated water to each bird will be approximately s00 mi.llirads per injection. If a bird were to-receive the maximum of 20 injections the dose would be 4 rads. The LD S o for the-song starrow Ls approximately 800 rads (gamma) so this amount is far below the dose that would cause death.

It is also far below the 30 roentgens (gamma)- that may cause reduced reproductive potential ,in; mammals. -

Acute radiation affects would not-be expected since-this dose _is also distributed over a five month period. Since this dose is below the acceptable occupational dose f or h" mans--(5 rems /yr ) -we - would- expect that no effects would be seen in the bird population.

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f ATTACHMENT #2A EXPERIMENTAL PROTOCOL The protocol followed will be that of Nagy (UCLA publication 12-147).

The birds will be injected in the pectoral muscle and held in black bags for one hour to allow equilibration of tritium in the body water.

Samples of 50 microliters will then be taken from the brtchial vein to estimate the total body water. Recapture and sampling will be attempted at 24 hours2.777778e-4 days 0.00667 hours 3.968254e-5 weeks 9.132e-6 months . If this is unsuccessful recapture will be attempted at 48 hours5.555556e-4 days 0.0133 hours 7.936508e-5 weeks 1.8264e-5 months . Sampling procedures will remain the same.

It is estimated that the tritium dose will be one microcurie per gram of body weight. The tritium is expected to distribute equally throughout the body water which is estimated to be 630 mL per kilogram of body weigut. Therefore, a 20 gram bird woulci have 12.6 mL of body water. The sample size used for counting is 0.01 mL so a sample taken just after injection would contain approximately 14000 counts (at 40%

counter efficiency).

2 0 microcu ries x . QJ mL sample x 2. 2 R.10E6_ dpq 'x 9 4 cp3 = 14 09 5 cpm 12.6 mL water microcurie dpm i

At 48 hours5.555556e-4 days 0.0133 hours 7.936508e-5 weeks 1.8264e-5 months approximately 900 cpm per sample of blood would remain in the bird. This count rate would allow for 1.5% error at Lne 95%

confidence level if the sample is counted for 20 minutes. Although a

20 minute count time is long, it is acceptTble for this study.

The study will investigate three groups and two treatments as indicated below:

Group Treatment Llynber of Birds Structures Building new nects 30 ( 2 x15 )'

(Bridges, Using old nests square culverts)

Pipe culverts same 30 (2x15)

(corrugated round pipe)

Rock outcrops Same 20 (2x10)

The rock outcrop group has fewer subjects since fewer inhabitants are expected to be found in this habitat. We expect the only recruitment cf new birds would be in this study group.

Study S ub_j_ec t;p The phoebes will be identified as radioactive in one of two ways. The first would be using a stencil and spray paint to print a trefoil (radioactive symbol) on the bird. The paint would not mat the feathers and would wear of f af ter the bird is no longer radioactive.

The second method would utilize an identifying band (such as yellow or orange) on birds containing tritium. Personnel in the area wcula be notified regarding the significance of the band. A cample notification is below:

"A study of the Eastern phoebe (a small bird) is being conducted on NWSCC property. Small amounts of tritium (radioactive hydrogen) is injected into the birds to study water turnover. The birds are identified by a radioacti r symbol spray painted on the back. The p birds will also have log bands attached. If the birds are found they j cf - should not be hcndled. Call L.E. Andrews at 854-1165 for N~

instructions."

The Af f ected Environment aDg_Paillway. to ILumAnsa As previously mentioned, the tritium will be dispersed evenly throughout the environment. It is thought that the water shed would be the mosi likely place for accumulation. The body of water known as Lake Greenwood is open for public fishing. The lake contains 8000 acre feet of water for a total of 2.6 billion gallons.

Assuming that all tritium releac 1. i the study resided in the lake the concentration would be 2x10'Yedmicrocuries/mL. '

20 millicuries x gallo_a x 1000 pi rocuries = 2x10-9 microcuries/mL 2.6 x10V gallons 3780mL millicurie Since tritium does not bloconcentrate the concentration in the fish would never exceed the lake concentration. The calculation below assumes a person ate one pound of fish per day for a year at the maximum possible tritium concentration in the fish.

2x10-9 uCi/mL x 454mL x 1 poune x 7 days x 52 wee};n = 3. 3 x10-4 uCi/yr.

Ib. day week year If a dose conversion f actor of p.064 millirem is used the individual would receive a 2x10 3 millirem comm/ itted microcurie dose equivalent.

If an individual drank 3 liters of the water per day he wciuld roceive 1.4x10-4 millirem / year.

3000 mL x 365 days x _2x179 microcuries x .064 mrom = 1,4x10-4 mrom day year mL microcurie If assumptions are made that 10 birds inhabit a s qt.7 re mile and all the tritium is deposited in that square Ic the cor. centration would be 2.5 millicuries /sq. mile. '

2.5 mi)l1 curies = 2.5 mci / square mile 1 r 'are mile The area is inhabited by deer that would graze on grasses and plants that would contain tritium. The deer population is approximately 2600 and it can be assumed that the deer will eat all the vegetation in the territory. The property contains 100 square miles and if you divide that by 2600 deer, each would eat .038 square miles of vegetation.

The total activity ingested by a deer could then be a maximum of 95 microcurles.

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2.5 mil 11 curies x .038 mi2 = .3 095 milli.gurief m:M deer deer If half of the deer could be eaten the total dose equivalent to the person would be 3 millirem (less than one percent of limits).

0.5x95 microcuries x .064 millirem /microcuries = 3 millirem Finally, assume that crops for human consumption are grown at this concentration of 2.5 millicuries per square mile. If one individual were to consume an acre of crops then the dose to the individual would be 0.25 millirem.

2.5 millicuries x mi2 x 1 acre = 3.9x10-3 millicuries / person 3.9 microcuries x .064 millirem / microcurie = .25 millirem Many of these examples are mutually exclusive and are shown to be the worst case in every example. This would demonstrate that any dose from the proposed study would be very low.

The ' Bird Banding Laboratory, Office of Migratory Bird Management, U.S.

Fish and Wildlife Service has authorined Dr. liarmon Weeks to caiture and band Eastern phoebes with aluminum and color bands. Dr. Weeks has l studied the Eastern phoebe at flWSCC since 1970 and has banded 2000-3000 birds. lie has studied breeding chronology, homing or site fidelity, nest success related to age, and various other factors.

l Theresa Rohling, a graduate student and subpermittee, will be conducting the work in this study.

Ms. Rohling will perform field work in trays lined with absorbent paper. All contaminated and potentially contaminated material will be collected and disposed of as radioactive waste. Any material spilled on the ground will be picked up and' disposed of as waste. Injection solution volume for a gi n day should be less than 20 milliliters so no large scale spills are anticipated. There are no bicassays planned for Ms. Rohling unless there is reason to believe a large exposure has occurred. Only individuals using 100 mci at any one time are required

) to submit to bloassay.

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There are approximately 130 people in residence on NWSCC in housing, the BEQ and 90Q.

Bis changes on a fairly regular basis with the movement of military personnel to new duty assignments or retirement.

There is a public railroad line that runs through the property and stops to load and unload cars. The load / unload area is to the north east of the Burns City gate. These are freight trains not passenger trains and due to the present world situation are moving through on a daily basis transporting ammunition and supplies on and off base. The on base use of railroads is to facilitate access to (xplosive magazines and case of moving materials.

There are around 4000 people working at NWSCC,;nalnly in the industrial area complex. The only people who are out in the area where the birds are a;c explosive handlers / workers. Their job is to move and store explosives in the storage magazines. Lynn Andrews, Natural Resource Manager, estimates that the number of people in these various crews number about 100/ day, These people work throughout the entire base area. Since phoebes do not nest in the bunkers and would avoid humans and their disturbance, the chance of one of these workers encountering a bird is very minimal.

Habitat:

Phoebes make extensive use of artificial nest sites, bridges and culverts, and natural rock outcrops when they are available. There is a very specific microclimate associated with phoebe nests.

The area around bridges and culverts is a maintained roadside approxinutely 510 m wide on either side of the road. All of the nest structures re associated with a source of water and thus insects for food. The territory that the bird establishes must also contain adequate nest building materials such as moss and mud. Nest sites are never found that are further than 25m from a forested area.

Population and density estimates:

Rarely are two active nests found on a structure at the same time. This may occur on extremely long rock outcrops, but even then they tend to be as far a part as possible (approximately l

100mk Phoebes are very territorial birds and almost invariably defend the whole structure from conspecifics. We estimate that we know the location of at least half of the nesting pairs on base.

Based on this assumption, a population estimate of adult breeding phoebes on NWSCC would be 200 pairs. This is the only war we can obtain the information and our estimate is just that, an estimate.

Migration:

Phoebes are a migratory bird that spends its summers in the Eastern United States and winters in Mexico and some of the Southern states.

Alternatives:

An alternative would be to use time-energy budgets (TED) which we have, but without the i laboratory ability to measure on major element of interest, nest building in adults, no breakdown of I

energy partitioning is possible. TEDS require you to know the percent of time a bird spends in a specific activity and an associated energy cost. This has been done in a captive environment with birds in environmental chambers, usually for resting metabolism. Forcing the birds to nm on a tread mill has been donc and some flight work through this laboratory method. We would not be able to l simulate nest building conditions within the confines of a laboratory.

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JusWicatlom These following excerpts are taken from the proposal for this study.

BACKGROUND The castem phoctw (Sayornis phoebe)is a common summer resident at Naval Weapons Support I'

Center Crane (NWSC Crane), Indiana. NWSC Crane is located in southern Indiana in Martin and Greene Counties. Phoebes use the man made structures and available natural substrates for nesting.

The female castern phoebe builds the nest without any physical help from the male. She is usually double brooded and incubates alone; the male does defend their territory and helps in the feeding of the nestlings. Females may either reuse an old nest, whether its own or another species, by modifying them by adding fresh moss and reforming and lining the nest cup, or they build a new nest.

The complex nest building of most avians plays a central role in their reproduct!ve success, but directly measuring the energetic cost of nest building has not been done (W!thers 1977). Ilowever, since the development of the doubly labelled water technique (Lifson & McClintock 1966, Nagy 1983),

measuring the eacrry budget of free-living birds has become feasible (Bryant 1988, Bryant &

Westerterp 1980,1? Bryant et al 1984, Flint & Nagy 1984, liails 1979, llalls & Bryant 1979, Ricklefs &

Williams 1984, Utter & ' -: .'c- ce 1973, Weathers & Nagy 1980, Westerterp & Bryant 1984, Williams 1987,1988, Williams & Nagy 1984,1985, Withers 1977). By examining the parental costs of reproduction, we gain a better understanding of the limits affecting reproductive output (llails &

Bryant 1979). >

Thus, by measuring energy gains and expenditures, examination of the energetic constraints of successful breeding at various stages of the reproductive cycle are possible (Dryant & Westerterp 1980). King (1974) suggests that allocation patterns of time and energy are intertwined with the animals life history strategies. The evolution of life history strategies may le better understood by looking at the fundamental constraints of food acquisition and efficient energy expenditure (Williams 1987).

PURPOSE The purpose of this study is to determine the energetic expenditure of fenv le castern phoebes

' using old nests and those building completely new nests, as it relates to their reproductive output.

The premise is that use of old nests confers an energy saving cn the user, which may subsequently be crpressed as increased production. It is from the previous studies that have shown some difference in reproductive success between birds using old or new nests that this study was conceived.

JUSTil'ICATION Pianka (1976) and Calow (1978) proposed the use of energy as a valid quantity for evaluation of the costs and benefits of reproduction. Patterns of energy allocation lead us to a better understanding of some life-history adaptations (Cody 1966, King 1974). These patterns are governed by the acquisition of food and energy expenditures, which are themselves fundamental constraints on the evolution of life-history t;alts (Williams 1987). Short lived passerine species should have a high reproductive level with a comparably high energy expenditure (Williams 1987). The energetic costs the parent incurs not only reflect on the current reproductive output (Cody 1966), but may influence

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the future reproductive output of the pare..- i. Williams 19M). Energy flows through principle populations within an ecosystem allow us - analyze the functkniing of the ecosystem. The metabo19 heat loss is a ma}or part of this energy flo,v and the use of doubly labelled water allows us new approaches to measuring this important parameter (Utter & LeFebvre 1973).

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Bibliography

1. Bryant, D. ht.1979. Reproductive costs in the house martin (Delichon urbica). Journal of Animal Ecology 48:655 675.

2. Bryant, D. hl., C. f. I bits, and P. Tatner.1984. Reproductive energetics of two tropical bird species. Auk 101:25 37.

3. Bryant, D. hi., and K. R. Westerterp.1983. Short. term variability in energy tumover by breeding house martins Delichon urbica: a study using doubly-labelled water (D2 OI8).

Journal of Animal Ecology 52:525 543.

4. Buttemer, W. A., A. M. Ilayworth, W. W. Weathers, and K. A. Nagy,1986. Time-budget estimates of avian energy expenditure: physiological and meteorological considerations. Physiol. Zool. 59:131-149.

5. Cettinger, R. D., W. W. Weathers, and K. A. Nagy.1985. Energetics of free-living nestling house (inches: measurements with doubly labelat water. Auk 102:683 684.

6. Ilails, C. J., and D. M. Bryant.1979. Reproductive energetics of a free-living bird. Journal of Animal Ecology 48:471-182.

7. King, J. R.1973. Energetics of reproduction in birds.(Farner, D. S.). National Academy of Sciences, Washington, D.C.

8. Lefebvre, E. A.1968. The use of D OI8 for measuring energy metabolism in 2

Columba liviq at rest in flight. Auk 81:403-416.

9. Lifson, N., and R. hicClintock.1966. Theory of use of the turnover rates of body water for measuring energy and material balance. Journal of Theoretical Biology 12:46 74.

10. Lifson, N., G. B. Gordon, and R. NicClintock.1955. hicasurement of total carbon dioxide productien by means of D2OI8. J. Appl. Physiol. 7:701 710.

11. hiullen, R. K. The D Ol8 method of measuring energy metabolism of free-living 2

! animals.(Gessaman, J.,). Utah State University hionographs.

12. 2 Nagy, K. A.1980. CO production in animals: analysis of potential errors in the doubi labeled water method. American Journal of Physiology 238:R466.R473.

13. Nagy, K. A 1983. The doubly labelled water (31111180) meth-d: a guide to its use.

Laboratory of Biomedical and Environmental Sciences, Los Angeles, California, LCLA Publication No. 12-1417, 45.

14. Nagy, K. A.1989. Field bioenergetics: accuracy of models and methods. Physiological Zoology 62:237 252.

I

15. Nagy, K. A.1975. Water and energy budgets of free-living animals: measurement usin ;

isotopically labeled water.(Itadley, N. F.). Dowden, Hutchinson, and Ross INC.

j Stroudsburg, PA.

1 i 16. Ricklefs, R. E., and J. B. Williams.1984. Daily energy expenditure and water-turnove-

-.- - -. =- - - - - - -. - - .. .- _ _

'-I rate of adult European Starlings (Sturnue vulcaris) during the nesting cycle. Auk 101:707 716.

17. Schoeller, D. A., R F. Kushner, and P. J. Jones.19'x,. Validation of doubly labeled water for measvang energy expenditure during parenteral nutrition. Am.;r. J. Clinical Nutrition 44:201298.

18.

Utter, J. M., and E. A. I.ePebvre.1973. Daily energy expenditurgof I purple martins (Pronne subh-) during the breeding <cason: estimates using2D O and time budget methods.

Ecolo r,y 54:597-6N.

19. Walsberg, G. E.1983. Avian ecological energetics.(Farner, D. S., .l. R. King, and K. C. Parkes).

Academic Dress, New York.

20. Weathers, W. W., W. A. Buttemer, A. M. Ilayworth, and K. A. Nagy.1984. An evaluation of time budget estimates of daily energy expenditure in birds. Auk 101:459 472.

21. Weathers, W. A., and K. A. Nagy.1934. Daily energy expenditure and water flux in black rumped waxbills (Estrilda troclodytes). Comparative Biochemistry and Physiology 77A:453-458.

22. Weathers, W. W., and K. A. Nagy.1980. Simultaneous doubly labeled water (31111180) and time budget estimates of daily energy expenditures in Phainopenia nitens. Auk 97:861-867.

23. Weeks, H. P., Jr.1979. Nesting ecology of the Eastern phoebe in Southern Indiana. Wilson Bulletin 91:441-454.

24. Westerterp, K. R., and D. M. Bryant. 1984. Energetics of free existence in swallows and martins (Ilirundinidae) during breeding: a comparative study using doubly labeled water. Oecologia 62:376 381,

25. Williams, J. B.1987. Field metabolism and food consumption of savannah sparrows during the brmding season. Auk 1N: 277-289.

26. Williams, J. B.1988. Ficht metabolism of tree swallows durine; the breeding season Auk 105:706 714.

27. Williams, J. B.1985. Validation of the doubly labeled water technique for measuring energy metabolism in starlings and sparrows. Comparative Biochemistry and Physiology 80A:

349 353.

l j 28. Williams, J. B., and H. Ilansell.1981. Bioenergetics of captive Belding's savannah sparrows I

(D)sserculus sanwichensis Belding). Comparative Biochemistry and Physiology

{ 69A:7b' 787.

29. Williams, J. B., and K. A. Nagy.1984. Daily energy expenditure of savannah sparrows:

comparison of time-energy budget and doubly labeled water estimates. Auk 101:221 229.

30. Williams, J. B., and K. A. Nagy.1985. Daily energy expenditure by female savannah sparrows feeding nestlings. Auk 102:187-190.

31. Wi. ima, J. B., and K. A. Nagy. IMt. Validation of the doubly labeled water

technique for measuring energy metabolism in savannah sparrows. Physiological Zoology: 325 328,

32. Williams, J. B., and K. A. Nagy.1985. Water flux and energetics of nestling savannah sparrows in the field. Physiological Zoology 58:515 525.

33. Williams, J. B., and A. Prints.1986. Energetics of growth in nestling savannah sparrows: a comparison of doubly labekd water and laboratory estimates. Condor 88:74 83.

l i

l

. , -- , , - - - , - . - . , . . ~ . . . -- -- .

Weeks, II. P., Jr.1970. Eastern phocle nesting in old barn swallow nest. Wilson Bulletin 82(4):463-if>l.

Weeks,11. P., Jr.1977. Nest reef p.ocity in Eastern ., .tes and barn swallows. Wilson Bulletin 89(4):632 635.

Weeks, II. P., Jr.1977, Abnornul nest building in the Eastern phoebe. Auk 91(2):367-36%

Weeks, i1. P., Jr.1979. Nesting ecology of the Eastern phocle in Southern Indiana. Wilson Dulletin 91(3):441-451.

Weeks, II. P., Jr.1978. Clutch site variation in the Eastern plumbe in Southern Indiana. Auk 95:656 6(4 l

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NESTING ECOLOGY OF THE EASTEltN l'HOEBE

, IN SOUTIIEllN INDIANA l lhn\ios P. WEtu, Jn.

.a l e The En tern l'hoebe Su3 ornn phoebe n a comn on summer re<ident in

.outhe n Indiana, make< extensise use of bridges und culverts for nesting throu; .ut mo+t of the eastern United 5 ates. This adoption of man made 91 struct t. - has allowed espan ion of breeding populations far from their pimesal ne ting habitati and may offer increa ed security from terrestrial i pred an u .. \est ucces4 how es er, ma3 he adsersely influenced by human and lis e inck r 4t u r ba nce. Nest < on artificial +itucture* frequently must 3 adhere to sertir al elements, with the re ulting increased likelihood of f alling. (,e Although Ea icrn l'hoebe nests are ea il> detected and readily acces<ible, rnany a int- of phorhe nesting bioloc 3 aie poorl> described. Kendeigh 1932, ronunented on thi4 parados. set little has hoen accompli-hed since th en, an attempt to fill ome information soidi on the breeding biology of thi- pri ic, m the \lidwe<t ! tudied a ne, ting population of Ea< tern Phr.che in an unglaciated portion of outhern Indiana in 1970 and 1971.

special empha i wa. phk ed on esaluating u e of and riuantif> ing s arious acce p.n ameter in ne t4 on hiidge an 1 (tdsert* in a region of abundant natural nr t- ite- An ,nhhtional ohjei tis e oa< 10 identify the im portance of old ne-t iemaining from presiou+ ea un and ne4t placement on oc< ting I ,dtte.-

()nly a few ne t- in nat ut al 13 cation w ele esamined. 30 birds were marke+

l 1

i \t LTil oD-Thn ook w a. ..ndu o-d on Cr.mr Naul U , icen Support Cent er ' Cran + NW5Ci.

a 211 Loc arra ui, upsing the northern third ot \tarun County in u.uth-i entral Indiana, but &; .d the area i i ompo ed t .ad held , ma.mained road 4idr* and other op nino ad ind t nial inmplese : the remainmg area i. wo ,ded. The rombination of 3 ilendritir cr-am n icm. and GO km id readi pnnel. nuns bridp. and ruhert, for poiential nemite-In frHury pCO. I ir perted all bndes and iuhcrt4 on the study area for old Eastern gg)

Ibebe t , numbend each nest found. and re< orded it< rondition and bwation. Other i i

> tenu. .,i.4ites such a4 building. and rewL outcnip4 near roads ahu were eumined.

I skiteo ar, at 54 das intersah after the br+t phoebes arrived and recorded nest : a.

o ntent.. ..s id. m es of new buildirg, and lwhanoral data. 5ites of cumined nest < j undudnt : o.nt rete brides creumted wood bndg>. wood and I beam bridges, tuntrete

s. whert . pipe i uherts, rotL ruhert4 buildmgs, nataral t ruck outcrop >. and miscellaneous.

' I o>n4;d red ans pan longer ti.e.. linear road di iances than 4 m a bridr and less than h a i uhrrt. After the pKO nc ong sea on I remmni all old nr<ti to ane*s the effect

'd a lat L of esi ting nc t+ on produ+.tisily and ne,t plarrment parameter

in 1971. This 4Il

8 I

[ .

M2 Ti!E u ll. SON Dl!!1ETIN Id 91, h 1, Sric"act 19'o rtnonal ebndnated the im-dlulin of rparating the ffert cf pre crue of ohl m the elb-.-t of time in the 19'l nonn; .a4 n. . f r%

llata . olb t tion it. 101 wa imH 19 q Ib au c phoebe. occa<ionalh began metal nc t* st difierrnt lo, ation nn4 or completed a nest without depaiting rgp, all nc t duccc .r i, y ,,

4 ulathn wro , v d ,,,

nest < known to base receised at lwt I egg. A aun .wid not i dehned e w thu produced at lmt I fledgling. Lt dau were *orted lo parame ter* to be i ony ..o d. a) because of n te with mining data. totah in mm. . ompari on are not the e a ,3

t nthers. All ne+t$ with clutches initiatM on or bef" e l*> Lv w ere s laaihs 6

season not+; the remainder were late -a.-n not e. .M urh

l. Signifi. ara e in ne t tru-e and ( h.t, h sie was te3ted with the stus uti i te t

I d

I compariwns werr wuh ilo t '. 41(4 ,7

- 1 libt LT- \ N D tilaa t einN

, hi clrrit al om/ nest buiblin,L-l fir +1 ob ers ed Eastern Phoches on tio

]f area of. 17 ?. larch 147tl 0: d 12 brrh 1971, when inales 4tuh were sin.a2 on e territory at bridae, that weir occupied enntinuously thereafter, l found the earliest date, for the hevinning oi ne+t* in 1970 and 1971 to he 2 bil a.w[

27 March. rr pertit elt. Earh -ea on con truction of a new ne t or irr-osation of an old one u ually took at lea 1 7-il da>

and at time- a lonaa 10-12 f(I ilau Temperaiuie cemed to be a tron conttolliu factot duiina ihh -riod :

1 reg.u dle , of con truction taue. huiMind wouhl often rea. . < otnplete ning p . . .

dau u hen temperatuie- were h4m ca. lO'C Sim e nr i huihiin.

m to

( he under partial control of 1ctul eratuie. -ra...nal s ariation in oc '

[

could he con iderahic. bi that w eir es identl> e ompiere iceni n : 4tood

.ation 1: em pt > ior IM da> < piior to ree i n in: # L eks lo!"' \c 1. on ' ction aller mi.IdlJy look a- Iittle a? 3 for complete con *lt u tion an ! dau for tenm ation of cii tina ne t-b!

I elutch inination),

- 11) 3 das, het u een fleik jen a:e! new

'f hr pim ement um/ r y.c.-l'F - huih hoth tatara and adhei, ne tm

{ 'l he> often depended on otnr ' ne ir regularit y i e.a. nail- wi l

ou;h l co ncret e. w ood splinter- mud o d

et [Tr> poi >lon pohtum ni 3 -/ ren h

enementarium ) ne-t< l for initial Luchment of nest material for nerent

{ ties t 4, yet the early *taue< of huibh: in the-e ne<t* wete mole diffi' 't than F for *tatant ne,t.4 The femalr. for My femah, build 6 5mith 1012'. had to

)

hus er at the nest.4ite ain! " throw ' ud again't the urfai r w ith a $/k oi her head. lhis continued until r mh of a ha4e w a buih up to alkin perching. \dh rent nest < werr th

haracterited b) a s iirle of n-i l mud spla*hes radiating from the ne l.

Adherent nests were rnore comn. , th ri 3tatant nc t* nndrr hoit idze$

6 56.1% \

M.99 ) and cuhert. 4 5 53 11.E n , hserthele *. it anut h he concluded that Eastern Phoehe- rrierred adherent to <taiant nem +ince f availability of suitahic sites had an wrridinu effect. A hiuh pere:rotae of 4

J f

.f l t l

I o

i, -

i frech

ETSTI'11N Pil0EDi' NESTING ECOLOGY I43 cuhert n,uts weie adherent, but these were scry few sites for statant nests.

i \lmo t all bridges had many potential etatant sites, but frequenth not in i preieved h*alion*. In man 3 conciete and creo*oted wood bridges structural l demeo at bridge upports prmided nmst of the " helf" situation but these were t el> u4cd Ne is placed in the*e locations faced across cream flow

.ind p allel with the roadway. Where po*+ible phoches strongly preferred

io face ne t inu ant the bridge edas. Drainnipes from the road surface, j protruding at "o' to 133 from sertical, were the most comman +upport

.It of l!. for tatant ne ts under concrete brid;es, in 1071, only 31.7% l9l of ne t- built under brid2es were adherent. The arpro\imately equal dis. ,

tribution of the 2 i> pr. of nc+ts may base tesulted 'rorn the need to con 4truct fntir*h neu nr l* and the saCancy of a nuinber of preferred stalant site

d.eca u - of eld ne t lemm al l, a- well as the apparent relati e ea+c of

)

4tatam .n n uction.

Nesi. m eir built on a t ariety of n urture- alth"uch the majorit> were b

en h:i&e and cuheit- < Talde l !. All hrike- and cuherts kete a ociated a ah mamiained roadside segetation: hourser. no bridge < ontainine a nest wa, mme ihc.n 25 m '.om a ital.le aiea of woodland. Such an obligatory nmi-w o"J!and ielation* hip w a uege teil h3 Gruber et al. i 1971 i in Illinois, and Klaa i pGn foumi in Kan as that +ome woody segetation w a- neces.

.ary nr.o nr-i.bri4c3 Cuh. - t ar> Ir - in de ign than I iid e- but the smooth sertical ides I ..' con ie < uheit icquired adherent ne t . Cukert, of coituuated metal 5

are of i 2 m di.unctri weie generails ura ahle by phoebes unle+s a con-4truction Rin i e.g.. incial tiip hangin from lupi allowed nest attachtnent.

f'or no t hiidge< and cuheits, with the e\ception of pipe cuherts, the N"lber of firste and ne* lins attempl4 ha* lalgeh a function of the number 0I as aildlele tiuctures. Onl) 3 of 21 (?\ainitied Cortuge!"d inetal pipe culserts eser had a nest. Theie was neser simultaerusl> more than 1 acti e ne4t on an> structure: Ohlendorf i1976) found the same situation in both Sa>'.s daror i uoa j and Illack phoebes (S. nigricans) in Texas, in n i ituations there was *omewhat nmre than 1 nesting attempt per ne+t i luble 1i, which indicates the reu+e of ohl nests for a second clutch; but since a few nests especially in 1970, recei ed 3 clutches. these data do

., 311 "M accuiately ieflect the percentage of nests that were reused. Seseral trends .j here evident, honeser. There was significantly rnore reu e of ne<ts in cuherts M w.

than on bridees in both 1970 a t = 6.27, dl = 60, l' < 0.001) and 1971 it =

l 119. di = P>, l' < 0.01 ) . This perhaps reflect

differences in availability of ,

dd ne*'- from which to choose for a <econd n~ ting thigher under bridges g than t ler cuherts in 1970) and the number of available sites for the

">n+tro tion of a second nest t judged considerahly higher under hrikes t.

)

l I __

I ut Ti!E WIL5ON l10LLETi.%

I d. 91, h t sqiiemi-r l'o g %

Ta nt.t 1 EuttnN Puotnr. Ntsri sso No.itsc Arrturrs Arconnisc rii L-stiosi -

ume tu o ^

eme n t Nest I..( omn Nem Nest m4 a ttoneis >

attempts ne t Nesh w o u i,Z Q

.in em pt,

.- wo Ilrike, 38 00.76 Cuhen, 17 i t ? "> i 1.2 P 30 m ot 44' u ti tw.

24432 0i M ilfvt s 1..W 21:350, Ali kli lhaildmp 6180 7 f 7.1 )

tr 1.16 3 OUh R 4 3' tw Mw. ttvtures 7 ( 9.3 i 9 <9 li 1.28 4' 76 b i fs)

TOT.\L \RTiflCI \L 75 99 1. 12 (o H 1 13P Natural 8 10 IS 5 7 i la TOT 5L 81 Kn 1.31 fi5 9H [,M i pai.. t ,r .in nem. .m hulmv thme tousut at oam .oirr mo in.c. mm A., m 4 mi h.- 1,.- .i ;

[wri e nt f sign'.f u mth i t r 6 72, di a 1.11. P <* O out for al 6ttmturn. t : El M. at n i4 P 4 D IK4 p, r hWIJes - l if it*f th tf1 t uIO % line

,I ' sie ms cenin,anth aitierent e r r 6 27. at : fio P C 0001+

' \l..an s enta mik aaterent i f = 2 DJ at : W. l' o ut .

I '

further indication of the po* ihle effect at allahle old ne t- ma a.a e on i j

(,, nest reu e i4 re'.ealed h) compari on of data from 1970. s hen old ne t* weie i

as ailable. and 1971, when all old nc t had been irmosed piion to the nestin:

ra*on > Talde t i. Significamh more attempt

per nest ucie made in 1971 than in l'40, both in total n on man made sites a = 6,72. di = 1:13, P

< 0.ml i and in bridge not. 9.bR d ( = 66, l' < 0.001 . I hi diffnence i

is not reflected in ruhert- a- s ould he esperteil h om the Jfoic!r+nlioned high reu e in 1970 ;

)

I [a tern Phoelies on Crane M\ { u ed ohl ne-!* not onl) for e> cood-hut aho for earlier attempt *, Of M obl ne t* in saryim: ronditi. i - uarked before the 1970 season.1:1 i ?n3 i wnr *uh equently renos a:ed and u+ed at some time iluring the season. lleu e of the 3ame ne-t after an un ucces<ful l first clutch n as not uncommon. \lo*t researchers agree that Ea tern Phoebe

L

(;

occasionally or egen freiluently n u*e a nest for econd brood t hlaa 19 R Mengel 19631, but no exact f m tuencies ha)e been provided. I found 31

! f sites in 1970 and 33 in 1971 on which 2 or more ne*iine attempt

ue e made:

16 i47,19 i in 1970 and 17 i31 D i in 1971 were in the same nec f f only h j, instances in which the first ne ' was succeuful are considered. - 1 (10 4

[ of 27# and Ul.07 117 of 211 ,i the second nesting attempt

weie in the same nett in 1970 and 1971, re rectively. The higher reu*e in Ita!! was at least partially attributable to the- pre season remosal of 1970 nests, which 4 eliminated most eld alternate site j preclude

statistical comparison. . although the removal of all ne ts in 197 k

i

l A

T rdi

E 61ERN PHOEllE NE TING ROI.tiGY 115 1 found httle es klence to upinirt the po talate that plu.ebe often buihl new ne i- in ucieeding utiempts to asoid mite infe tation illent 19121 fliere u a. ome Inortahty of tiestlings frorn the northern foul mite IOr-

%/ min on. oh mrum i hui hirds at time < ieu ed nc is in which the eailier i;ege.' u.una ucie beasih infe ted. In 10 0 i found that in in-tance where l' inh ed a different ne<t for a sulsequent clutch. 62; renmated an ohl ne,t r.uhei than buikt a new one, since miic sursise acil frcm sea on to

,f A40n in Id nests t hlaa* l9Ibi. little reduction in ri k of mite infe' talion imuld I.c cained from this nc t shift. [gl A tacit a umption here i< that a pair of bird- that occupied a tructure yntmuitu h liif 46ughitut IIIe ne tiltu ca on h as lbe

a me pair, behasior '

and esent .quence indh ated thi< w a* u<ually the ca e: although t'.ere w a<

!Ael3 -mr wpb< cment, the frequenes of urb wplai ement could not he

!cte r n,

. ! hi au e the hird were unmarked. klaa I1970 8 found that 031 mi 1,a n 1:a to n 1 % che- iemained et ihr ame -ite < brideri throu;hout

he hm Jnu ca on. ( nhlendoif ! 1976i inund -imilar liehas ior w ith pa3%

g and i:la,L pim,he- but al o found ome r apid equential r eplac ement of Q i4eeden Ih.n em . in nn -ui ereding e on ide,atio,- the a <umption of non.

eplacem. M id hiccilet - i- not critical to the s alidit s of the pt e eutation.

bf -/d i ' Hr.t[ 4 [f4 /( b a[ c, -l$ggdd)ing an ! 8 lut8 b+i/c s.MiJtion base been Wtailed ' m l rie Week- 1970 ' The cai!+,t c;. w a. 10 \piil in lo o r and o ' , o in 1o:1. In hoih scan 2 definite prak- in egwlasing illu trated I de c h. M i li. Joul ;r.lirondedne-. of the pecie, the early peak for botn

' war < s m mid. \piii. anil the late. ca on pcA in 14 0 in late \la>. Th.

m und p, a in Piil n a. inrxplicahis delased in about lo dar. T:ir late <t sa: a chu, h na- hesuo wa- 23, June in loro and 21 June in 1971.

Ihe aun a;c ize foi 1:1 t ompb te club het u a- 1.73 0.012 s Ei range L6 cu e. 6 .t = 5 reut 91.n = -1 3 em i w ith no significant dif fer.

T** in 4 !atell 4lle IWtueen scars. I f(>Und. !Miu es er, a 4 gnkfiCaill lice}ine t : .I a di r 165, li < o.not i in tlutch.<iee in late neit iclutche begun 3her 13 \l.n i. Clan h-ize were ub+tantially lower in ome in4tance, where hirds 1 : new iather than u ing old nest

and where bird buik adherent UIber Ioatl =latant nest 9 l keeks [9 Ol, Ibilc/o n ; fledging and nesting Jucce33.-$ince I e\amined mo t potential , ll]

Ul" prior to ne t initiation, nearl) all nest $ nere know n from inteption, t I therefore include all nc tA in in) measure of ne-ting sucte*< and ha+e the

Zing 3u ce+4 calculation 3 va lutal eggs MUhth more ne* ting attempts %ere made and eggs laid in 1970 than in I i l able 2 i. \e4ling succes wa$ higher in l"To. and it is esident that e

'"' I f nu alone cau<ed t:v decreawd 1971 un e 4 Esery facet of the

'%d' ih e e> cir appeared depre<*ed in 1971 i ompared to 1970. Perhaps g I

VI .-

5 l.

416 Tile WILSON lil1LETIN IV. of. .Yo. 3, September Ivm l'

Tanto 2 !

Stilst sRY OF Nr.3rtNG D si g roR rnt Eastuis l'uotni. os Crisu .NWsi._ I, ,sg 1970 1971 ma

~ ~ . - _ _ _

NeAct Pe rs ent N n.n he t Pers ent~--~-% nder O

--myt .~.

g .L. e.l ar ti s e n. t ~ 94 91 p.A l

,f N.t ed cornplete clatchett 89 82 171 .

Total era laid 4M 117 8 .-

Ens per actiu n. 4: lii 439 I 6; Egp per complete cluuh i 75 430 i: I

Esp hatched 340 '56 281 67.6 2 i j, i] Egg hatched per ai tiir ne4t 1 13 2.96 .

I Em iniertile ur addled 1 0 th :2 ;-

Egp k. t or abandoned 93 21 8 lin 278 211 !

21. -

Young fledgni m: 226 3 11 l Your.g fledyd per actise ne+t 110 236 21 3

Young finized of em 12i.1 <&2 at; c.y

} bug flehed . f egg, iwi hed 9n ! 79 1 81; Ynung I.. t b" fore firhing n 37 41 suo tnful n Ns 'l 70. ; 5; 'i ?. i 12 <> l o i uu., n.4, i....na an , i,a. t.m . I i

p-4 the hiah producthity in loin re-ultnl in a 1971 p+ pulanon ahn..un.dly hi h in fir l ) ear leirfl . W hirli are tang' rally conct*tlett to lit

Ir%* siU f U--l u t bl ertl8*r.

I t.ian espe:iented aduk. i son !!aartman 1971, l.ack lo on s, in a.i lition. ihr kark of ohk nesti in k9Il ro hase had an influent e.

.\ls limited number i N = lo s.i esact determination- of incubati~n period p ii.e., la4t egg laid to la t ege i a. hnl > and ontline perio.1 i N = 1 h aner i *

, with most puldished repoit- i n al er et al.1971. tonri 19.19 ' t t

o d.n .

wa4 the modal len th for each: istaa- il975 foumi an as eiag, io period i

, of al! dan for phoelie<. In m .t in,tances all eggs hatched aiuun a 21.h il perioil, llatchability of epp that sunised the ineuhation period wa 911' for combined sears: a4 with ther f actor +. it was hiuher in loTo i%n': '

than 197 t (93. L'; i .

. I found a significantly hiah. - 2i z = 1:;i.6, df = 1, P < o.noli 4unhal

! : rate for nestline than for egn. ,ith !!.6'i of al! eggs laid hatchin and W '

'.' of all young that hatched fled; ns. (Jsing 21 dan iincluding layinai as th-aserage time that eggs remaiix in ne<ts and 16 dan as the aseta. e+ din 2 I 4

petiod. I found a loss rate of 1 Y day for eggs and 0.91'; day L rstlin2-j Both rates are substantiall> he r than mo+t renort< for open ne< ting altricial i, bird 4 iKnupp et al.1977. R. - herry and Klim<tra 197tn. The higher r de

of egg than nestling lou is esiJently normal in ahricial specic+ Nice 193 **

t i

f

!i

tr+ch E WTDW PHoEltE NE4 TING H 01.00 14 7 i

31 wouhl be espected, since almo*t all ne*tline 10 % are to predator < but l ran> agent cau e egu los.es.

I similatly. lightly moie ne-t* f ailed in the ege i30.14 ) than the ne4tling l . !!.6' i t a ac. The pattern of nest failure sug2eited that oiice the eggda>ing alr h iv uhalion period was completed, the ne+t w as r elath el) eeure

' Gr.ti} lodeliini. hea!!y tYi of un Ucle% ful nev!$ f ailed duiing eggdU kng or i sery eails incubation, while onl> 11', of the f ailuir occurred from that point

- inrough hatchin.:. This to + pauern might be piedicted sinte abandonment I a nn.,t cononon in thi- eads tnge. and the mo4t readily een and acce sible gl te t, wouhl be taken irtickl> l.3 opportunistic predatois. Af ter hatchina, the

icedina actis it) of the aduh couhl auract piedatois to the more concealed r.est s. I he slaih nest muttality t ale w a U.'A' ba-ed on a 37 day period mclu w Ll .n layin2 peind i, con iderabl> le

than the 2.15 .o. s ;- )

kul !Is lucklef i l"m i for open-nc ting pa erines.

N im i 17. found that ne-ling -uctr-- in open ne* tine abrhial bird 4 in

~

f S b uh I..m em ate Zone a wrage- Pr. i im ib dginn succe- i. Though

.he gaw no mean. he tated that nc tine suur - in lude nc-ting biid was p uh4taniiah hiahi r with a I!cdgine -u tiu ( os er. \e linu <ua e- for both 3 ear cond ined

. as bl ou -hehih loo m than the on.!' < figuir for dattd nests t hi- ucceu I

u sim' to the mean -ucce for hole.ne ter- gn en by Nice 41937 i. which itse ri<ahlv an infhant salue i Alayfield l+1 acceo ha ed on dated i nesting w a 4ienificanil3 .g = 13.L dl = 1. P c O.001 ) hinher in 1970 than lo!1 l'ahle 16. Graber et al. t 19:13 i--und +ubstantiall> lower ne t nret ew foi c ent ral and ou. r n llhnoi* pho & - 436-579 ), a+ did Klaas P'70, in Kan,a i 3." i for ne t not parasitied by llrow n. headed Cowbirds Voloilans mer ) . Ib ihle rea ons for the dificience in succe on Crane NW5C and other aiea. include s ariation4 in ne t. ites and predator popu.

btion- The lack of human and the tock inimierence on Crane NW5C un.

khte S al o played a role As ob ersed by Nice i1957), fledging succeu, ha ed iotal eggs laid. w as lower than ne4 tin 2 succen based on total nests Table J i. Ilowewr, differences were , mall becau e infertility was low. mendt acces wa4 hich. and predators tended to take completa clutches or brood 4. '

ln Sucrew parameter

ihatching, ne4tling and fl dainu) were all significantly t hi;her T = 13.0. x 2= 23 6, x 2= an.1i df = 1. P < o.001, respecti ely) in 3 i

P40 than 1971 (Table 2 6 Additionally, nestlina .ucce* w a4 highet ix =

112.0. p = 30.0: df = 1. P < 0.001 : 1970,lo1. respecti ely) than hatching

'dc*s in 1.oth > ears. When only succe45ful ne-t< are considered, the infre.

m nc of partial ne ting failure is evident: or combined years, hatching g

"x ce- w as 92.uq and n tling succes. 03.01 Esen in the relathel> lower g 4

t

} 4 11)

Tile WILdON 111'LLETIN

IV. 91, L J. Scienhn 197o TAotr 3 Ntstnr, Sertts, ron Eisnas Puor.ns <.s Cn ot NW C. hm n n Citwmrno av Srmn no Nur Snt' 1970 ~' ^

wit l'atly (4i tate ini T atal U i Ea<h ('; i tate 4 <

37~ ~

liridges Cuberti 16/21(76.2) 15/2h 71.4) 31 '42(718) 11/22@3A) 1121go 27. k 13/18(*2.21 12/16i710) 25 '34(73.5) 10/17(*.8.8) 4 11ao.8) It :n p<-

Misc. et r uct u res 5 3(100 01 547 Gl.O 10 '12:313) 3 6 (8 L3) 2 I po ol : Ic9 n Natural

1. 3 (n3! 4 5 .80.0) 5 8 (62.5) 2'3 ((C) 3.1 i ?3 u - :ta TOTAL 5 33 47J1.5) 36 4M15) 71,96G10) 31 W l 6) 22 41 31 1 i o p .,

each sc uon m 1 apra-15 M o , i.t's.imrac

+ oo r 16ece..bwm iwi ae n rmo i"n ma w re hau a oua hme m si hn h a av Ma -in lime.

'snu eu ugnd n anth lughet i 18 e 61, af : 1. P < 0 Oli th.m for 1971 a uh ert won p

hatching succe4s figuie<. most of the failure to hatch was attiibutahh to eg; infertility rather than egg di4aitcarance.

Nesting succe** for both year + combined diffrred \ cry little beturn earh 169.D ) and late (bl.IS .) f ea *ons, he flifference 54 allno t totab.

tidnd.

able to a moderate late ea on decline in *urte** in 1971; succe-- t

' l almot

I hientical in both ,cason+ of 1970 iTable 3i. Giah-r et al. 1"71 al o fou nd

{ a decline in ne4 ting succe-4 (i ',' in the late 3ea,on in southrin lihnok ith.nr

{ dhi+ ion of seasons wa+ identical to mine). This appaient decline of ne4tnu succeu with time is contran

(- o the *ituation found in mo t ahrivial i-le-i blan 1963, Longcore ar. I Imnes I"69, Ro cherry and Niitu n a 1"70,. h:

addition, fledging sucre a'

. tended to be higher in the e,u h than lxe sea 3on; no general montur <ierea c in fledging sucee-- w a4 es ; t. wi:h Slay having the highe,t tuo for an> month in 1970 and he i 1" 1

. i iTable 4). l'leduing succe34 e e, hones er, ahia>

highe t in the n. ah4 aml season

when the greate+t nund er of eggs w re laid.

i Longcore and Jonr4 i l9(& citing entomolouical data illotting twl.

suggested that a partial explan aion for low succes, in Lod Tin u bes 4 /6/+

cichla mustelina) in the earl > -a.on when many nest

were aethe might 14

$ - a functional respon+e by pred: sors, i.e., to take a higher propottion of pre) when more are available. T}n, concept cannot he accepted as an axiom in I

settebrate communities e5mith 1974i, Phoehrs do not depend on metatise

[

I corcealment for nests, and it @ pears that predator re*ponses to y i en and young) density differ bet 1 een nesters in this niche and sp~i- ne< tin:

f in vegetation. \ lost parerine* . e segetation for nesting cmer ubich +hou!.I i

3 ,

l afford greater concealment a, the season advances (Lanyon 1957 Longcore I f and Jones 1969,: cover for ph1 ehe nests does not change appreciably. Culb 4

I

I Tnha E \3 TERN PHOEllE .\ ESTING Ecol.OGY 419 Tsut.t.I

,\s u wh .a .ht susc St u to Courost Nr5 in Tiut Potou ton L ottas Pnot ou os Cu n t N \\ 3C. t wi n 4

'~ ~

Han bmg .nu ce. Nothnu usucu I lea an un c e s wr ik mon r it mi <

ratna r.a t . .

I Ico n.; ul '177 68 I 116 121 n9 1in 177 (as

ir lu ni al lu tri 79.9 tu Is9 70 4 g Jane 68 86 31.0 38 Wi 83.1 58 88 69 0 bJIb 'I kprd-kb \$Jti !!O 226 73.2 IW 170 41,1 t 100 224 70 8 1.a t.- In \l,n - 10 June i 170 221 73 9 147 170 86.5 147 Oli 63.6 lrl \n 136 210 7 4..! 136 1% 83.9 131 210 6 LH 1 h 31 66 WO 2.t "U 09.7 21 h6 368 Jun- 92 lli (12 ti! 90 72.8 67 lil 17,3 E. I \rri!~ 13 \la, i 163 211 71.1 1i4 163 HI2 1;9 211 ( 0.0 in h to lunci llo 186 62 4 t., i ce ., tan hva n 83 lin 7L1 M l86 43.7 p no ..

i.

L ,i.

e em 1.na i r ne ime; .1n; uni. ea e.. ci

.htt.i , : m a . ..' n g N a ;nt n. of e - nd

' [.il ?!J 1 U p ,1 la\ ilig at tbt* peak of Ibe nc-iknp *ca*on are moIe Su(Cenful than tho-e hn io e.nlici ni later i t'atter on loni lirow n 1"67 8 Though it h difih uh ! -enegate < don > ef fects. the similarity of these gull and phoche I

'iata m.u 1.e more than coincidental. Gisen specic* that depend on 4tructural or ::co -

blogical (oncealment and or irlatit e inaccewihilit> of nest

as n-dat .+ o e- and gi.cn a relatisel> *tal le predator population with hmileu .oclidit ) . a con-tam numhet of nests would he taken reganllen of the nond-r .n ailable. i.e.. a maller proper tion as numbers increa e. Thi4 nidenth o.cui, in thi4 Ea tern 1%ebe population.

For h"th > nn , combined, nesting +ucce - appeared higher in statant

'WO'; i than adherent 161.:19 P nests though the difference was not statis.

tically si;nificant: this apparent eleuted succes was general in sea $onal and warly lo caidow ns. In l'Cl. Succe" for statant nests i63.29) was signifi.

t.u tly x = l.1, di = 1. l' < 0.0h higher than fo. adherent i s !.191.

I 5tatan- it. had < lightly higher oserall fledging succen than adherent nests, ll) 61.01.,nd 00.10, re,pectis ely. This difference was not con 4istent between 5tu* with adherent success higher in 1970 i 70.69 u 66.7%) and statant j,

L.

in l'nt i c1.0G u 47.:19 ) . For all successful ne<ts, however, fledging success

% highe3t q; = 0.0. df = 1, P < 0.01) in adherent nests 193.4% ss 80.19 ). '

Thi4 eles ated <uccew was consistent between > ears and was contributed to h bth hatching and nestling succen. This ma> indicate that nest predators that u nalh take single eggs ie.g.. Peromycu.i spp.) can more easil> reach

LiO Till' Wil 50N liULLETIN

la 91. L. :. Sevimber m.N statant nests, as would be espected. Other losses in adherent nests proportionately higher. muti Although adherent outnumbered statant nestings il(M s 371, ni) osera d impreuion was that il a suitable statant site was asailable it was pr e fe rre, k These succe3s figures, as well as clutch. site data t Weeks l'CI, indicate th,

' a preference for statant sites has definite adapthe advantages, although the-f remain unknown.

i

~,

]* 14ecause of numerous factors, most related to susceptibility to ne,t iti<n pi certain neit sites seern to be more secure than otheis ! Nire 1057, Cot g Knupp et al.1977 i. With phoebes, one micht espect nest un eu to be hi:he 1971 D on bridges than in cuherts, l.ecause cuhert nests userage much lower an, i1

{

cl cr to the stream, thus increasina their asailability to opportuni4 tie groun predators, This does not seem to hase been the ca e in 10I0. for sucte-g appeats almost equal t Tabh 3i. In 1971, honeser, sucee" w a 41snificant!

[ ,

bmer ix" = 6.7, dl = 1, P < 0.01 e in cuherts; the difference wa c peciall,

[ es ident late in the sea,on.

'p fh dging succew w as slightly higher on bridge, u.o.ir: , th an i h en.

( bI.37 ) . Ilou t% er, lbe duece*4 b as mil eiinsislt*nt l'elw et'n } edl* n i. uhet:

un. css higher i '.1970 i 71.10 u 6 l.19 ) and bridge iuct eu highri m 197

, i ,- $ 50% u 11.0W. For sucreoful ne4ts, fledging ucrew wa hi.:hn ip-17.7, di = 1, P < 0.001 > in euh est 4 9 6.37 o U0.74 ) ; thi diffnem e wa.

f[

con i lent betw een S ear , un! was reflected in hoth hatchn . and nc tlin, p ,, ucce 3. It appeais them

C,

m. ihat while to-e, of complete ilun he i bn.ud i

rather than partial lone- i- the rule this tendency i, no..t pionoumed in

( i cuh ert nests. Partial lo- apprar more ronunon in hiidue ne t-

!thu-other lo-ses must be con. por. linch tigher in ruhett no i-

, I found nesting ,ucce" ignificantl> lower t x- = ..a. df = 1. F 0 n!-

,' on ciemoted wood than wm irte bridges, 35.07 u Ta.n hu L,un and j ilurchfield (1973) found ti.at flarn Swallows (#irundo rushvii rarely used I

creo4oted w ood bridges in \li ,i%pi and speculated about nosiou elicos a

of creosote or increased - certibiiity to predation. Ilecan e the increerd l los,es i found on creosoted moc 1 bridge 3 were not from lowered hatchabilas J

or nestling sursivah I lo mt think lowered succeu was attributalde to an) chemical characteristics oi . ie a e. Wooden bridges base man > uctural

.., upport elements. e.g. brat .

bolts, etc., to *upply panage for pie rs and t

t

3. wood presents an easier ch-bing surface than conclete.

No information is as allaldr on post. fledging and adult surs h al in phoch" j if atemperate for population remains pawerines, stal le. howeser, and adult mortality i , 507 t Ricklefs 1973), post.lledgimr jusenne i* abo mortality is high A rough ~tirnate of > oung fledged pair mas be obtaind using the method of Nicr il9371 and the conservatise e timate oI N t &I E

,1 e

I

nh

EASTEltN MlOEllE NE5 TING ECOI.OGY 451 atletnpts pair: 1970 - 6.76 young pair: 1971-5.1-1 )oung pair: hoth years

.5.97 soung pair. Therefore, the post.fledgine vursisal reviuired to halance adult lo,*e, wouhl he im in 1970,19.59 in 1971, and 16.79 metall.

I fince phoebe 4 mme north sery early i are periodically sirtims of spring In uch rara, increased first.3r urshal couhl present a prolonged g stor m- 3 popul on depreuion.

Can a nnd wuc of nest Juilures.-Causes of ne<t loues on Crane NW5C were s aii.~h lini no lones in the 2 3 ears were attributnl to human or the<tock interfei enc et \lmmt half (49.34 i of the louet constitutinu 1R29 of all 91 anempi+. u a+ attributed to nest predatoin 3 timea a* many loues were from ,

} redation a. fioni lbe lle\t !rading factor. }!oweser, both tile peIcenta2e of total nc in .: anempl* lo+t to predators and the proportion of total loues attributed to piedation were lower in the3e data than in many 6tudies of I r.idici u en ie*. Llan il%h attribuied tr of all failures in deciduous

+Lrub Lui in piedation, while I.ack ilo5li thought that 75'" of failures p in op. tw ling specie = wete alltibutalde to ] tedation.

Situ e timehe ne line *ueceu wa generalh hizher than that for open-l nestin: Unicial biol- and the peirentace of totallo >es attributed to predators va, le - I , on. lude that ne t. oe -ch rtion b3 the Eastern 1%ebe afford, I them a i, un deuter of ermits 'n,m piedation. I beliese the principal ne<t i

predatol- u ere the blaf k tal nake lllophe o. olooleto), raccoon il' roc)on Ia!or). .m l w hite Iooled mou e il'eromj seu s leu ropN t t . llecau e no Ilouse i Wren. 7 Jodun nedon i, llou e sparrow

il'ouer domesneus i, or Com.

l rnon < Ue* - Quisenlus qu Senla i ne*ted on tructure< w ith 1,hoebes, and

since b at Jay iCynnocitta erittu,a i were nes er ob crsrd at bridge *. I beliese that mian ne.t piedation was minor if it occuned at all

.kbalubhmelil,alwa)4 in the eg" 9tage, accounted for user $59 of lowes, 3 but it. cau e wa* 2enerally unknow n. l.o u e , iiom ne4u falling il2.!G i or from Ih.eding i;.01) were lower than espected. Various mites are common I in Phoel-* and frequently infe<t soung but in only 4 infestations was the Lara<ite l..ad *ufficient to cause nestling death, it. 3 of 11 instances of clutch aband nrnt. nest

were beasily infested with mites, and the egn were ms er, u nh dark stains from mite droppings. The+e large population 4 may l hate t aumi the abandonment. , 11 Ahhough cowbird 4 are common on Crane NW5C, only 7 phoche nests .i

! 461 of a!! ne4 ting attemptal failed from cowbird interference: 3 additional d' resta reemed at least I cowbird egg. hut 2 of thne were succeniul and the 6ther w a de4tro> ed by a predator. This yiehb a parasitism frequency of #

315 \ithough it appears that cow hird interference in this phoebe popu. f lation h li ht i .\layfiehl 1965i. I'riedmann 19M ) identified the Eastern ""

l Ph* as a sery important cowhird host. In a hiition, Klaas (1975i found g

3 l

t E.

i 15 2 Tile WILWN Dt L!1: TIN . l'el. H. L.1. September 197" parasitism frequency of 2 Fi in his stud); atul Graber et al. il97 t Central I and soudiern Idinoi5 with die PNCt; 1011 Of Nit. CNtremein Coun! ca, fourMI :109 of all na not,octn 6'

sis affecteil. A more critical esamination n(

my 3lr data snealed that 5 of 15 i3M > ne ts in natural locations tecched s

j f*M M 4 h li ta on ) 6 O( C6 13 I t>n attificial strueluee4

' h t.f t" liara itifri[

, This differente is signific int ip = n6. di = 1. I' < 0.001 i. Aithouzh i g Graber et al, t 19711 nase r o ne+t site breakdow n. their discua.

I that a con <iderable number were in naturalKlaa4 locations.

t 1973 i. on th,.

i S other hand, e\amined Only nett9 On brid24 and Cuhert51 On hi* slu.h area theie were no natural site 4 for nnts. lie stated that

}qeeder, lbe Phoebe "reCeksM mMt of ibe Cohbkrdh attention %" .arh becau+e i i

d carb ,a the

, o season. Although only a small number of ne t* in natural sites

_c_ mc.it.~,.,ua_e_o _ fo,d,_.m e were etun

outhern Indiana, i) lieduced rowbird para *iti+m abine could base been a

'. +ignihcant elerthe factor in the substantial adoption of man made .tiudon, a4 l

.! ne414ites, esen in aleas of phantiful natural *ites.

failure p rate a}'peaied higher (or adltelent than statant ne a t , brit l}m l, apportionment of loue, among the sariou cau.e4 the 7 $. . .es w a, sers imil.o . ' 3 r c.i y an indication that hi2hrt 19.5 7. t o ll.P

[ encruv deman I, of adheient ne,i con.

uruction i Week.19Tu. Klaa, lo!ni louer, the enem re en e of irma!c+

J I

'h 1

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.minra in , ,ulo rnua huh.r, m !

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b-t I snhbng h.gan about ! L . m.a sun.

.p m .u,. h+n m.ugn, b. , b4 512 An and i omph comn n. d nro.

h.T a wu t-ne a,..n m ana .n .. ,

O b.nn.4 hang orger .na .ippaomn o. ih-new, fr.an prni..u+ sean and tru iro , m m armanao, n-. n. . none .# e.

tu for a .. snd 1.rnod. ..nne n r.t .!!.1 % in 19:0 and 31'. in lWl' Noting surres. in 1970 <!0 L i

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n was f!rdging su4 te.t Suo ro lun .

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and saried arronhn,s to nnt ar . unie or dm hned shghtly a. tne .rmn a.han el N)) '

u fb-d,:ing Sui t n* w as highe t in -

.urve . aai 4hghtly higher m naura rahi when nening intensin was highen. Fledu -

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.u. . .... m. .-w ha u., .n n ' ' .h u4" tubert., but pn.l.ahdin of panu! , wa greater ..n
)oung pair par w a= flehrd. I.ri.f a s An awra- "I M vn.dai..n u, ihr nuj.., on.e s i ?
Ifl 5 .,f all neu attempo. attribe i f.,aure wah nm a an b..... , ...n-on n":
caun, with Drown headed Cowlont ! , predanon. AWndonrn< nt wa ihr ned lehN
(+ f:owinnh para +itierd a *ignihrantly
- tirrente i ming bibire of ..nh :tK: of attent" Arr perant el ne ts in natual ilun artifn ial H'r'
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C \STLilN l'il0EllE SE5 TING EcolmGY 153 Nest loar- "rrr ErtJtest in early egg or rnid. to late ne.thng stago, Nr.4% nmrtahty rate
..a 0 979 das. \lortahty rate was higher fer eggs (1.337 day) than nnilings '0 919 /
Ja) h A(D O W Li'.hG 1! E.N Tii i ani prattiul to the Indi.ana ()rpartment of Natural He+ources for support durir'g the penod of thi4 tudy and to the per.onnel of Crane Nasal Weapone support Center for their < pi rau-n. C M Kirkpairii k,11. 31. Ohlendorf. G.11. l'arker, and W. J. 5mith effere. ,riptal suggr+tions on the manu+cript. Thi, is Jourcal l' aper No. 69% from the :
runlu, \gre uliural Esperiment station. 9I a
LITElt\TL'It0 0110D 007, \. C. 1912. Life hi tori-s of North Amencars f!u atchers, larks. sw allow s, and 3
their alh .. I'.5. N att. M ui. llull 179.
I;m 3.1; t. il 19td. Ilrreding suctre, and populaii..n growth in a volony of l{ctring and tr4.. r Illuk backed Gull 4, Laru argeniaru and !.. inum. Ibis 10930M15 h
Coot 't i 19 4. E, ologb al opret, of reproducto.n. Pp.161-312 iri .bian bb.h gy, e
l) 3. } arner and J.11. King, eds. ). .\< a&mii l're,$, L w Wrk. p har a 11 lo a llo t n lanon, of th. paranut ..wbud,1 A Ltl~ Mu . Dalt 2.it Qt m n IL M J. W. Gn ua n o n E. [.. K nm 19?i Imrmi, h rds: T s ran mdae. Ill. .
Lt lbs s un. 1od Lt. L A l llott.nc. ( .- 19-,1 Pon. iphs of in,e< t pn.date.n. \ u. lin . Eni..no I. k lt L IR2.
1 so; u s. I. \ on P. G lh nun n t n. 1973. S e,1 m chroon of llarn wallow. in wi . . rml Tb u-ippt .\m. \lnll. .ht. 9 6 00 . W Krwunn. - L 1930. l'an ntal can an ! n, cu.luti-n i ldni - i.. Ih.L W m gr. 22; I- A kt u- 1 L PG \ popolati"n unh of the Ea n rr h +hr anu n + *o. ial r. iati-n hip w n o. in..w n lo ad, d Low hinL Ph.D. thi-iv. l m . Kan a . Lawrence.
1973 Cow hnd psu a mm ani n. ung u. . i + m the Ea irrn Phoehr, t L i . Pap.
l: s H On a Tho. N at. lli t.11:1-16.
l Ku t it it \L lb 11 un ts jn. no 1, 11 linioso 197* liepn.dm th e i+ 1-g3 of Ano n. an 1;..lon4 in n..rtho n \laine. \uk 9 8 Et.C ln IL IMI. 'lhe natural regulation of animal numia r Osford I nn. l're . lendon.
1% l'opuanon t mhe- of 1.;rd,. t hfor i 1 na hen. London.
l m os. E . E. 19i7 The comparathe biology of the ma lowlark iSturnellui in Wi+-
con < i n. Puhl huttall Urmthol, Club 1.
Imu mn.. J.11. no 11. E. Jom 1909. Flern du, t n e u, e e*+ cf the Wood Thru4h in a +!aw are w oodiot. Wil4on llult 81:iW406.
hu Nr4 ting + us ten calculated from espmure. Wil on llull. 73: I i t,. Il F. 1961 2 = > 261. j

. 19o3. The lin.w n headed Cow bird, with old and new host. Lhing Pord 4:lM8. *'

hui. IL M 1963 The birds of Kentutly. AOl~ ornithot Enogr. L. 3.
NKr, .\L M. 1917. siudies in the life histor3 of the ong Sparrow, uit 1. Trans.
Linnean Soc. 4:1-2 17.
. 1937. Ne.ung sucre. in altririal birds. Auk 71:305-321. llf Lux V.. jn. 1 94. 1 I;eproducove sucren of Lirds in a deciduous scrub habitat. Ecol, m4LE311 g I
n >
fi 434 TiiE WILSON til'LLETIN a IM 91. W.1. Septemhn to 7 O uu wont, 11. M.
' 1976. Comparatiir breeding eenlogy of Teus. Wilu,n llull. 8811M71. t horbn in Trs ' n.
Parnnsos L J. 19(& Timing and ndsbraidus. Ibi. 107:43 5 439 9 acing of biouds in the liia k-lu a ini Cult. L,,rra, llicKurs, it E.
Znvl 9:1 48. 1969. An anal > $is ul ne stha mr.rtality in birds. Smith.onian G,ntra.
,j h . 1973. Fecundity, mortaht), and asian demograph3 Pp. 36(.-4 G in tire eilq
. ! biol"gy ni birds I D & Farner, ed). Natt. Arad. Sci.. Washington. D C.
liosuunt, J. L or) W. D. Ku us ta s.
1970. The nesting endog3 and repr..ductio f '
prtformance of the Eastern Wadowlark Wilson llutL 82:2 4 267 i s ume. IL L 1974. E ologs and behl bh logs. llarpe r anel Row. New b rk.
S u n n W P.
4 1962. Nesting habits of the Eastern Phoebe. Auk .io:lio-il?
r m a. D. 1939.
Temperature, growth and other studies on the Easinn 1% i State Mus. Cire, .22:1-12. M p, m li s sa t M s N L. 1971. Population dy nai
] 4 D. 5. Farner and J. IL King. rdu. Academn- l'rroPp, 391-09 in Misn biobigy
% t t >.s. ll. P,, j a. . New \ ock.
1978 L; ,
Tuk$6:fdl 464 Clutth she uriation in the Ea ttrn Phoi he in -uthern India Derf. Ol' I OltEsiltY AND \ \Tt 11 \!. I: dot m.b. PL liliUE F .1\ ,
% FvT I O E TTE, l%DI \ \ \ ( ?)il!. ALLEp t't.D l I \t G. 19 Ul.
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' lairwlildan 86 l If F*i 8 l il li -
k .1'lMlf t k t l.'t f t.r '.I h hi h e h k. M %,
tor) ,d j tJ ill . . .i O ra -
..t ornithologs. Corncu l~nmtats hhc 4 bw T nrk 19BD. ins m. mi r w ho k na ,
of an i,utr upon wlu h it would be appropriatr for the H d .,n 4
4
4... o t y t.. t Ae an oth bl 1
poutien h. uld i in Wr, h 19N. Th. rewlanom et ommuni~ ate thi Srtuaden to L 1.aphon I.. f"ri the annual matin; U
the membrr+ hip van unite to espn - ed at each annual m.,tma an the onb u.iy in whh 1 i ,
nu t t e r +.
t hn. ugh the 5.wiety. a 0+rmal pi..ni..n on . n .:rutis a i
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l j CLUTCH SIZE VARIATION IN THE EASTERN PHOEBE IN SOUTHERN INDIANA
!!ARMON P. WEEKS, JR.
l Department of Forestru and Natural Rnources. Purdue Univeruty.
}, Wnt 1.4fayette. Indsana 47w? CSA lJ Asstiuct --Clutch sizes were examined in Eastern 1%ebe nests in $nuthern Indian' m irgo
., and 1971. The majooty of the nesu was en bridges and cuherts. Nest budding began m 1.ue 1 March with birds regularly using old nests and budding both statant arid adherent nra nesta i 1%ebes raned twn brmis w1th peaks m clutch inmation in mid Aptd and m late Mas -carly d June. Avrag, dutch site w as 4.73 with little s arianon between years. the most common clutch L size was fne. Withm a year, clutch site dedined ugnificanti) late in the season Mean cuuh me h!! was lower m auherent than in statant and iao as oppo<cd to pre emung nests the enne (ost of nc-t buildmg a dncussed as a pmuble cause of these reducuens Rn ntrd N Sq ot N77, G aarried 9 Atrd his.
h ..
j Titt Eastern Phoebe (Sayernis p/webri13 a common summer resident throuch, .t r much of the eastern United States, including Indiana and surrounding stf es t Butler H%. Mencel 1%L Graber et al.1974L In its phylocenetic development the Ty-rannidae experienced an incredible degree of divereence in nest site selection in this proi es the Eastern Phoebe developed a nest site requirement (i.e. rocky bhdf and outcrops, mouths of caves, or streambanks with overhangs) that undoubtedly made
it an uncommon and local breeder in pr tine times. Although the Ea3 tern Phoebe has u idely accepted man's dwelbngs, bndues. and ridverts a5 nesting subitrates and
thereby extended its di3tribution conMerably, these birds are at the mercy of i
tinually chancing engineering desien (Whitaker 1974L For example. thi 3petie- .n nesting on vertical elements of concrete bridees as opposed to older wuden or I.
beam structures, may have to divert encrey from eug production into construction ot adherent nests. This adaptive behavior micht affect clutch size and. ultimately, population numbers, The evolution of clutch size in various species and the influence of estrtn3ic and 4
intrinsic factors on intraspecific variation was treated in detail by 1,ack tlW41 and
- von Haartman (1967L in their research. predominantly with European cavity ne -
""*" ters they had the opportunity to examm large numbers of easily available ne -
g The nest site selection of the Eastern Pkebe should make it an equally attractn e research subject. Because of its widespread use of bridges and culvert > for nestine fu substrate, nests ara easy to find, often evily accessible, and, in many areas, nu-73,
' merous. Kendeigh (1952) commented on the surprising paucity of information on j ,,, g%
.,s the breeding biology of this common species. a dearth that continues to the present.
4 I studied Eastern Phoebe nesting biology in southern Indiana in 1970 and 1471.
i Emphasis was on birds nesting under bridges and in culverts. although some nests on other structures and in natural sites w cre included. On many of the larger bridm Eastern Phoebes nested synchronously with Barn Swallows (Hirrando not .t )
4 thrnuuhout much of the season.
I h
MFra %
The umh area w ai Crane Nat al Weapons Supycn Center. a M14m area occupying the northern part of Martin County in ungianated moth centra! iruhana -\pprmimatQ 7 M of the area u umded and the remamder is old fiel15 mamtamed roadsidr and other openings, and industrial The comdeset d
6% The Auk 05. 6M-u6 O noter t d
I if a ner 6%) Phocbr Church Sur Unanen b e, ;
t , m t- o dramed by (bree major treams ta-h with a dendutic drainage pattern that grades from deep pe r - 'nu s alle> m the north and tast into t> road ahm1 fholplains flanked by w albhke bhdis in the
,~
d,
' wi w est Tto, pattern viupled u nh MO im of roadw as s remlu m a large number and good
' ton ai tindges and ruh erti lt F el ruary ICO l tr@cctrd aM bndre$ and (uh ern en the study area 1H tJd I.a,trrn Phoebe nests, ne N'd t a< n not found and remrdni its conditmn and lot atmn OO;er potential nest Mtes inchuhng la o and other structures and natura! utes prmmate to road + uere av esanuned After the arrb al
( /
d et : t phaebes. I s iuted each nie at M das inten ah and rrentded ne 1 o>nwnts new construction, an w mral tniormatmn Aner the UJa nc-ang cawn all phoebe nc-t were remmed in order to a- 'hr dicct ni a lac k of emtmg not3 on s armu+ produrin it) and ne t plac emt ni parameters in the
r }."E scar All data (t U* Cthr! In j C l u al identh al tu 14' Un!) t juiches knnv n to be CO'Tip!cte
w - ad in anah se, o camer data w ere +a ned f rom a smali weawr canon on t rane NMC.
f Cm n the popubimn of dutch me+ u ere esamned 9:th the d test WN r uimparmns, includmg b c luttk me thaneew were w oh the Nalenn Diest RDt't/rs e:11 lectSSION L 't types and tlaamen!5 -In both 3 ears the phoebes arrived on the study area in mid Alarch and nest building beean in late Starch-carly Aptd. Construction of
. e s nc ts n a+ lou with time ot' con truction varyine between 7 and 12 days.
T re u a an mters al of M days betw een nest completion and laying of the nr31 t.. a i haracterstic pattern m La tern Pnoebes #Graber et al.1974. Klaas 1970L h .. , Nm found this mters al to be loneer in adherent than statant nests and -
I c ,c-noned u hether adherent nev baldme u a, an estra phpioloeical drain leadmt i a delay in m ulation. \ly observatsn are not utnciently mten-e to allow this tapan,on but my clutch - /c data mdn ate a pomble enero cmt of nest bud hne Phoehes constructed both adherent and 4 atant nestsiFit li statant nests as craced e Her dro than NO ui than adherent iu ually creater than C; nest < The vport '
fr a tatant nest also prm ided percune for the female durine con <truvion In a uon to heme larger. a ihetent nert- appeared more ddncult to con (truct a birds h o hover to apply nestme matenal thraunhout the early +tates p
webe trequently u3ed ohl nests rather than constructm: completely new onei Ere than 7W of pre-exi3tme ne3t3 w ere u3ed by phoeber during the 1970 sea on- ,
O? ne+t3 w ere always renovated before u'c 3b the addition of new moss to tht rim n! a new lininc Old liarn Sw allou nt-ts were occasionally modined in the ame )
manner and used by phoebe- (Weeks Um About 509 of the econd clutche, w ere la!! m the same nest as the nrst clutch in mstances where the nrst brood fledeed s cessfully. 607 in 1970 and A01 in 1471 of the second ne tme attempts ne re in I tl ame nest as tne nrst Meeks unpubbshed AISL 3R Neits were found in many situations mcluding concrete bridges creosol-d 1ood bn tees, wood and 1 beam bridges, <quare-formerl , crete culverts, round 1 orru-cated metal pipe culverts. rock cuh erto build 'iscellaneous structures, and natural rock outcrops iFig. It Alost of the ned were under bndres or m culeerts. Alost bridges had numerous potu for statant nests but cul-serts often had none, Alan 3 of the pipe culverts were unusable unless a construction f.au allowed the attachment of a statant nest In the follou me clutch size considerations I assume that birds continuously oc-n eing a civen structure u ere the same pair. Behavior and es ent sequence indicated l t! this was usually the case, and Klaas (1970) found that wJ of his banded L tern Phoebe < remained at the same brid:e throughout the breedine season. Ilow-es er. the vali<hty of the presentation doe 3 not hinae on the correctness of the as-sumption
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Tu uni: ot' ouuatmn of clutche4 of the Ea h Po el,e in wuth-sentral in I.ana in Itc0 anct h'l 3R immediately and laying shortly thereafter, centributing to the ses cral early May
, dut:iies : Fig. 2t There was a de6 nite lull in clutch initiation in early and mid May of both y ear > and 15 May was chosen a, the dit ithng point betu een nrst and second
-[ clutches in subsequent clutch size consideration, The second peak in clutch initiation wa> not as disting as the 6rst. owing in large part to the damping effect of nest failures and resulting renestines. This was most eendent in 197 I w hen overall nesting succest u a3 lou er 138.P.i) than in 1970 t?2.Fi).
In 1470. the second peak in clutch initiation uas separated from the nrs,t by about I
40 days t Fig. 2L about the minimum recychne time from the nrst egg of one sue-(essf ul clutch to the first egg of the second o e 4 days laying + 16 days incubation
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660 flAnuos P wtexs.Ja. (Auk. Wt y TABLE and 107)I.inYafub0n wuth central inIndana he of f astern Phoebe dut(hes uithJn and betw een r- the breed i
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Nurnber of em 5easonal a wn 3 4 5 6 N i* E 1970 Total 3 is 66 1 su Aprd 4 ;; t < i s l - i 4 J7 0 May 31 4 77 n%
0 4 32 1 37 l l June 2 10 '
O to 4 92 . c x l April-It Mas 36 4 26 ? ^ ::)
j lb May-10 Jee 2 1 3 0 al 4.43 : 0 <
13 30 1 4h 4 65 :
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16daysne'W 3 day nest repair r 42 da,M in 1971 hourva the 3eand peak wa3 absat day s later than the tire This approsiamtely 1 ucek delay m M rene3 tine ir. !:.71 u a< confirmed throuch exannanon . ' the mean interval betu re
-ucce,4ful : led;ine of the first brond and the beginmne M 3e second clutch m the
- Il g -ame nest in the 2 yeah in 1970 th interval as eraged 9 day 3 in = 21t a um' icantly IP < 0 001.1 testi shorter permd than the 13.3 day mean in = .'.h in . 71
..M d An explanation for this delay is not readily discernable Althouch late Slay. u hen the second peak would have begun if the 1971 interval was the same a3 lo- w a-f"'1 relatively cool with hiehs in the 1st rance and lows near 7T. thi3 3houhl n. ;ase been sufficient to delay avulation 3 Although the mean interval between nestmes was relatis ely lone, ses eralin3tance>
1 of an interval a3 short as 1-3 day. uere ob3erved, both in the <ame nests and 4 mm e.,
renovated. pre existing ones, in one mstance, renontion of a second nest w as be;un w hile large young were still present m the first nest. Klaas (1070s found a mean X{3 (Q interval between termination of succruful tir3t nests and the beeinrine of c.nothe to be 7 5 days lie commented that 3
A um
. ce this interval u as 3horter than his eb crs ed A ",3 physiological recycling times ti.e. time necessary to produce eggs alter loss of clutch or brood durine nesting) of 9-10 day 4 the inhibition to ova development mu,t end
,, "' k C w hile yount of previous successful br-ds are still in nests. Sly tindings generally concur with his observations.
Clutch sm :ariation.-A3 with most stadies (Davis 1953h clutch sizes here reter to the number of eggs found in nests ;,re3umed to contain complete clutches. Elo"~
ever. as these nestines were highly localized and most nests w ere observed frt'rn inception, my counts should b very accurate. Nevertheless, the powibility exb: 3 that single ezes were removed by predators durine laying and thus not recorded No clutch size was assumed for the few nests containing young at discos er>The mean clutch size for 171 ne3 tings na> 4.73 : 0 042 (Table n. Little variatmn be-turen years nas evident in clutch size with 1470 and 1971 averaging 4.73 2 0 OSS
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n= $5 15 9 8 27 32 14 (7 10 20 30 10 20 30 9 19 29 I april. MAY JUNE h sc an , Lao, u. m the L u rn tu n, u i -.u. e noit. m iu ': m -"otu rniral u l.a aii i - h prt u ni ent tan lanj t tror i
I
' sE',m i . To r o ont respectis ety . I cu comparatn e data are avadable, but Klaa-i U. + :mi a mean cluu h 3 ire of am m nr3t3 non-para itized by cou birds in Kan.a 3 in .Tio lucan studies. u here ob3cn anon mtensity may not has e been equally ihstr d through the year, Cuthbert (l'n,'t found a mean of 4 64 and .\liddleton and ton i1W whose data w ere pnman!3 on tirst ne tinc+. observed 4.49 eec-per . O n ih e i- me nunt common (lutc h ,ize u a u nh 74.."i of UC0 and 39a'; of 1471 J' i clutcht - of thi., sve : Table i L Graber et al < l(G4) found 3imilar percentaces for ,,
I ris e.ece ciutchr3 in Illmoi, but other tudie-iCuthbsrt 10nL .\liddletua and Johnston lh Klaa,10:5 > show ed rive-ece clutche4 to compose approximately 30'i of total
)
a clutche, Clutch si/c appears to be closely controlled in the Eastern Phoebe; I found approximately 95'; of all clutches to be of four or fis e cues. Clutche< of three or six eens ",ere relatively rare. I found three succcuful nests that contained tuo-ege clut<
in ICCl. but in two of these locations previous events indicated partial BR tlutch predation, probably by Peromyscus Irio cpus, was likely and the third wa3 probabl3 affected by pre-existing abandoned ceg.s. None of the e was considered normal and all were excluded from clutch 3ize compilations. Graber et al. (1974>
'imilarly obsen ed that all of their one- and two-eeg clutches may have been the rnuit of predation. Klaa.s t IcC3) recorded eicht u 7'N nests w ith tu o rees; although l these ne,ts were presumably non parasitued by cowbirdt the hich incidence ot para itt m found in his study area (249 ) leace3 open the possibility that cou birds remm ed eces from these nests without depo itin;: any of their own.
Cl nt h me was found to vary between pha es within the bre edine seasons (Table l' Low er mean rlutch size in the late season was found in oath years. with these
'hifere ce> 4tenificant IP < 0 001, t.te3t! in 1971 and when years were combined This dechne in clutch size with 3ea3on is a generally obsen ed phenomenon (Davis IH3) Such a decline has been hypothesized as being adaptive iLack 1954. flaukioja 1
_ \
e A
4 d
J 662 il ut slos P. % t t Ks. Jit I ban Vul y I ,
q >
Trea 1. Clutih iue, in stern Phorbr$ in +outh.ientral Inihana. rategorurd bs 13 pc or ne I t on-tra, tmn 4 ; . .
J t Can- Clutth $de e .stra tmn - - - - - ~ - - -
f Sea onal < ategon t.i pr N 3 a ( S s: 4 4 1970 Total
, c\dherent 5t 1 11 H 0 4.*1 - 0.u:5 f 5tatant G ! 1* '
a* 2 n g41 J ) Aprd ll \le \dherent 20 I i 11 0 4' : O pH 5tatant II O I b 1 4 Oc - a go, r 16118 -30 Jane Adherent 25 I 1; O 4 S .. 7. 0 114 statant 20 t 6 12 1 49: .i !! ; ,
l '
i re t Tc,tal Adherent 41 o 14 : o 4x: oc
?talant 34 O IA 20 4 '4 : _t 4
I Aptd- ti \la Tdherent 13 0 4 I r# o 3, '
) ' tatant 1.< 9 4 m 4 t _'- e j
, p> Nin - m jan, Adherent is o o x o 4 .. ro_.
} 's tata nt 20 0 19 0 4*- : 0 ttt
, .- B..' b :, car i..i a! ' Adherent c. : + u o 4- -ou3 5 tatant !! 1 !i 47 6 4 ' : a 0* 7 i j i Aprd m sim . -\.nier e nt a t m o 4- - 0w
g

tatant 3t u t
t 4
-am O v> Alm - o junt Adhereni a !
= > 4 " an- ,
yl Statant a i h , 1 .
0>..
I '
t e nii e i t . 6. o .n .: , , it nv g s 2
I at an i s n-
> .1:
! IC P suth the ulumate controlhiie tactor beine a dechne m late ca on ka.!ab;h:.
' os iond for youne
, . . =
eeam .\hhough the ca3onal dhference- that I found in mean cluti h sue w ere .remtican' "6" the decline was not a tradual one thro .cwut the 3eason a3 iliu trated for p . Eastern i
.\leadou lark iS!:o nella mapia 3 by R 1 berry aml Klim tra i UJm Rmer it u;
gg I marked by a conet..no through mes the peruul with a 3uh tantial !an ca-
_ed dn hne (I-ie. 'o imdar to that demon anted for . erd Buntme- iEmN i '
w>
f P"*j nIzoi by llaukmja il970t .\ lost of u; 'e late clutthe w ere tlur. (lu: i an
tailure of an earlier attempt .\lthough the os erall mean clutch ize tor .\ pol an.!
h:
hh M y h[ .\ lay w ere almort hientical tTable It there u a no con i3 tent c betu ren s ear 3 rince the May mean was substantially hieher in 197u and lower in 1071 than the Aprl 7 y,, mean. Simdarly a ben the mean clutch - zes for the peak lapne period, o: cat h > car i g . ,,,,, Jiu. 2i are compared ti e. periods u hen '.he majority of the second ilutche represent 5 @ normal second attempts tollowine succc tul completion of a lirsti. In!O calue3 u tre
.i (y not appre iably <htferent (4.85 vs. 4.mn , hile lat clutches w ere con 3hlerably 4maller
'M use m 1971 (4.96 v3 4. m .
Spurred by the ob3erved thfticulty in onstructio . of adherent a3 oppe ed to sta-o ,,f,[
tant nests and the strone tendency of crds to use ohl ne t3 when as anable. I ev up* amined clutch >ve as it varied with thes ,ets of factor 3 (Table 3 2 and M Interaction i ri of factors is of course po3sible, but there was no evident trend in adherent or 3tatant 5"!
~
placement of new nests under bridee and culverts, with eight early 3eason IC l, } .
new ne3ts dicided equally and eight of m late season ne t3 adherent in M71. *27
_ . .g of 70 carly-season new nests were adherent while 67'i of 12 late,ca. on new nt3t gi r e
w ere 3tatant.
y i The mean clutch size for statant ne, u a, con +istentle larger toan that f or ad-p,o .
i herent nest + iTable 2L although this ihtference was sub3

antial only in thi car;>
-ca,on of both M70 and 1071. In neither instance was this ditference < tat!3thah)
'y 20 m}.
a
AA 4
l i
(t< i .t - ./*l l'hes Ar (M h \Gr Il Mf f e s s 66.1 l tmi Chm h ute, in horn hn.rbe. in " aihmental Inhna ' n temed i N,. m ,_a ,na , no,oe i~. mem ua o.,mi,e,n bol.in em-,,u of mnot s r a on i oniou h 1. anal3 e of ( a,cs rnar ke I .iit o an aatt o.k *
s. , _
um ._
I'
<a a iaternes hi+1oh N t 4 t f, g t $1:
w I .a h, ru tmv +v '
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1 i 6 U 4 M f 0 .41 I *R : 4 ,l a t- he's 06 .ig u '
14 1 4 71
O b W '
New 11
b t. 0 4.ta 1 O lii J4'l ! [ he t'%lilin g
Nu t * *
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M .d > n e to t wo to M P O :." and I belic\ c toe tidferenic, to be real in i ad in '
( tht tren ! ton ard larccr ilutcht. i tatant at 31 u a cenitiran' in t he i a+
,Ca o P O M. p s di and in the t :ai ra un if - nM p s O(n lor b. '
\raf.
91} Hlaril l he l o' I \ alues air U ja t d(\ llHin ati'.i sinrt all rarl) nc 14 u i 'i hrU 's*I- Illiu t'\ f r innl\ t Ig} t ad !!lt es i a!l) t'a in l 47() lng t
u ere ( of).t rth ti .]
lhil' idr 4
.Ili IfMIH alltin that lai t(its in .O Hn bin to (hlth idt) of nt t ron trut titin ng br ratmg
\
nc -t > tt tho e t ongdett i) hulh Ju t prior to oilpo ittun. had louer mean slun we in .11 ca on, w here wmpart on u. re \ ahd tTable h Again. hon n er th0 dd!t rt ne rs V ithin indleblual ta' tin 3 'd lY70 w ere not stati tit ally (tenttirant largth ht i ail t' p.
are c apared a of lou number 3 of neu nt i- arul w hen incan lor the w hole s tar 1.ouL n - Mn the normal 3caional difference m ilutch 3in com.
phtatt. ;he c<tabh hment of stati tit al dif ferentet Agam, hou e\ er, I behew tht "
f dihtit n(r are real since the trend tou ard larger (lutches in pre-exotine nests io m ~. m mfonnP. om. e = m sin m dinem m,eremomd p,iorto l the
13rason, no pre existing ne313 u t re available to afford a comparison s
hh debate vill exists as to the ultimate factono in the n olution of dutt h ta '
and it \ ariatmn with season, age. latitude. and other factors (Welty 19751. Primary theoric3 imlude I.acki(105-H ideas that the (ontrolhng factor is the maxircun' num- p ber that the parentis) can adequatel\ nouri ii, that of Skutch (19671 sb re clutt h sin a adju ted to compensate for mortahty, and that birds tend to la) the large t dutchr that they can produce s\ on Haartman 147ll The latter suggestion i3 refuted 14 a decret by several observat.ah8. especiall) n ork u ith indeterminate layers (\ on I
haa"! man 1967). Regardless on the ultimate factorist that contributed to dt ternti-Date d l
' of (lutch i/e, 3nnie proximate factt ri 63 operates to control \ ariation seen
%Ithm a species population It ocem> mo t hkle) that this proximate factor re\ oh e-Aroulld the liioenergetic$ O[ bret ding, inclwhng ll1e effect) of nutrition und operty g budert.
Although an increasine amount of work is beine done on the enercetics of a\ian l teproduction. little has yet been conduded recarding energy partitioning and rela-I lion htp> in wild populationt King 097M conduded that then ti sienificant energy
\ l,
6M liaruoxP wn u,ja i wt, y,a y cost (prmr to hatching) only in the production of the ova. and Zwickel t1WD belin eri reproduction not to be sigmhcantly curtaded by calonc $Lortage under natural (on.
ditiont Kendeich (107M (onsidered incubation an important reproduction encrev
} co>t, and Drent 11470 presented esidence to support Kendrighi position. Addition.
} ally, King ilo7M (oncluded that the period of feeding and parental care of yo anz
^
is the part of the breeding season in u hi(h caloric short?ce eserts its major ettuts 2
on reproductne potential. Zwickel (107M how ever, pointed out that the timine of 1! the breeding season had evolved to coi. spond uith peak fomt abunilame for >oung
= ,
and cited Cmlyi t1971) ebservation that the incidence of young dying M orvation C 1 in the nest w as surprisingly low, in these consideratmns of energy lir m,em.
k phasis is on the energy cost to the female up to the time of hatching, nr hatchmr the number of calories re(cived by developing younc is the major ancern Nour, j
q i+hment of soung is of course important to the population dynamio of the 3pi. n but also important is the drain on the feeding task on tha energy reserves < the j female. w hic h must often sub3equently produce replaument or second slutt hes L The activits of nest building is very 3imilar to feeding of >ounu and,if the number
of trip, is simdar, is probably equally cosdy energetically. There is of course a
<htference in material collet ted, a fac tor w hich may make nea budding more n.
pensis e than G e dine young The tollectmn of food for youne may be mereb an j
esten ion of the birdi normal feedme ac uvities an:l the bird is free to eat oct a*ional 4 catches and probably do;> Gatherine ne>t material require, different < catch imaer-and probabh a not done in concert uith normal feeding 30 that enerey roers c. mas
[*- be utdi/ed durm; periods of active nest con tructiom
%=4 se si A,,9l f" The female Pied Flycatt her tFicrdula hypol, ura) in Finland ha ben hou n to h .- 174 of her body w eight by the end of the ne tline feedine pe:iod Mehy Umi bu' r,o dut h measurements bas e been made f or any pnir uith respet t to nes'

[Y 9 budding. KendeiLh il97M indicate l irom the w ork of ElMaill 3 iUm uith Zebra

%j. .g..J !g Fmt he, Tarnier 3gia e astanotir that the ent rey cmt of ese pro 4hn hon (annet
'M ra dy be separated from the to<t oi rest buihhne and early mcubation !!ou n ct.
q
" me
. .. di 3peties wmmonly u ed in capn tudice budd s er3 smple nnt, k llin !c
'197M pinnted out, construc tion of Ven simph* nnts requirn beha\ ior of 2rea!
(
i comptnity but not, I think. an eners ou'put approaching that a wild birl met 1 ,.
1 s s pend m buddine a laree ne3t Thi3 energy espenditure immediate!3 prmedn an i in mo4t specte+ overlaps with a 3emn I, established energy output, the produition
.im es.
of eggs in >pecies that regularly raise a econd brood, as does the phoebe. ton 3truc-U tion of the second nest is nearl> a conunuum in time w hh the encrectically (ocl>
MD
mtivity of feeding young and may acu t succeeding reproduttive performant e.
.4W There are feveral other aspects of the Eastern Phoebei nestine ecoloey that in-
" # '.' l 7 dicate that the enerev co3t of nest con truction may be important. The phoebe budd>
a relatively large ne[t, requirine several days. The reduced (lutch site in neu!) budt
_{ ."' M nots and in the larger adherent nuts meents energy partitioning Thi, enere co t N may be a dominant factor in the evol.. tion of <e eral nesting traits noted above. l'
,j , the use of old conspecific nuts if avaibble. cven occasionall> thoie of other species (j;-
l' in preference to building new ones (Table M, h the reu>e of the >ame not for second broods, even though the risk of para ae problems may be greater; and M the con.
struction of a >tatant nest when support in a suitab!c hication is available (Table D-
%]' =;
Additionally, the delay between nnt completion and clutch initiation in the Ea>t.
' ern Phoebe may be energy related Klaas (100) found this delay to be longer in phoebes that constructed adherent. a3 opposed to statant, nests. This compliments i
<l 4
i
M -
e m mes.a.6, .4 e m h a. 4,m.44w--
s diim m ar-mm - h-4saa-d-m -+.6-+4ep--.m-m a wse-. L--- --
! 6 7 l
l 1
l l
l I
I l
o , ,+ tr i e q /%ee (!unh u:t l'ao uma bM i
I my Nnding of reduf ed (lutch sue m adherent nests and suggest= a larger drain on I enerc> reten es in more exten n e and ddfu ult nest oin trut ta.n King it o indi-i catrd that follu ular grms th. and aoouated energy drain, begno m small 3ongbirds N l j about a da3 3 prior :o ovipo ition This mtans that in almost all instarn es with the ,
I Ea tern l'horbe nr3t buih!ing (ca ed prior to the beeinmn;' ol foHu tdat de\ tlop- I
. nu nt. thus pren nting the conc urrent impontion of tuo e nere) drains 8' i
((W(Ll.pl(3'sb l =
Thnouch Ea tern l'hoebes on Crane NWH' t ommon13ut t ulvert and brid:c3 for at tmg and enmy a high dreree of nc tme m ter. tht> appear 3 to be at w me B l ui t in term of reg production The durabihty of esisting nests frorn season to l
3ea on on the c -Inn ture, u of real ht ntht. as it relit ves the birds of the entrev l m th ta k of ton nuttvic completel> neu nests floucs er ne t freriueritly rnu t
! be rel udt in i uh er' w hen destrm ed bs high n ator or predators, and neu nest are oo .-n.dh hmh be nrath briders tor unL now n behas wral n a on The c ne t are 6 l oft .o HE nt 'I he at t of buihhne < -pn iaH3 of adherent no t- es identl3 ha an j
.I t luttfi l/c b
eth .,i ( i '*l Ii t Iki r }(ma!!'. l8d hi In n f U1, t'i ,
O baakt t 'ldi tri onunended an .:rm tal nesting 3htlf m new pipe (uh t it o
=
1 that I a It rn l'horbr* and l tarn 'u.Ilh m .ouhl U c thr e trm ture3 .NW h t hr h e *i ll u mih! undoubtoHy initra r population pr slui tivit3 on Cranc. for leu pipe i uh rt-f h.I'l -nitalde ne,t 4tt' a f ul an nit et t '. rr) i al\ rrt tir l>rldge that <lUl has e a - i 'I t no 1 ott u a tu t uph d Tl> hndm o: thi unportatu e of prt t Ai ling nr t ar ac pnJ.ahlr t nere3 ro i of .r!ht rent ion um ton mdu ate that no tuiu 3 helve mm ' se her ht on other -trut ture a ui d Pla< ed m preit trt d lo( ato n' on .itty trus ' re, j tht a ould allow t on trut tmn ot t a t.o.: nr3t* and sunultanrou 13 amure m . ter dur -ht3 Prefi rred ite ma3 b. n/ ntn ily rit oem/ed by an indn idual tano ar u no phothe nestme builoe3. or ma3 he <h tcrmined by Croctme es t ral hrh t -
. . mn u, l
i n no.uoiintw inhanai m +r : t,a r i u da:a, e nn n a ni m ard to nn u , ., on , m ,
m - _i ,4 i ~ ,,m,o emn ~ _~ n._ su . _ ,a.~ x ),
e s pr . i t i la riv > I ho nw R l' ilom6 rd C \1 h rkt.atru k ana l I k! .a ot h rul u i t u! < 'n -m-bl t o m.ou u opt T !u o Jm na! pant t L
3: .,n t he PorCr Acri. Jino al la pi rime nt >-t . n i
i f! b % I' bi lYh $3 lu ais A W i s .* he tard of imbana \r M Rrpt inct thpt Gi "I Nat Rn N tit. BR com. u t wi tu .a a.n ~ re, wn en m m .,, u an sam uaios i entt A .I R Kmg. l .1 i Nro York La <m. Pre. +
Ct 't u t i h r . N L t v .' The \1u mgan Au l 54 n m wt> pt orbe sta.13 <part !b Jam Piny M art + ,
,i bM l- tb l~ lult h rryd:r:g bo.l.gy < ' bird- l '; + l6 4 - h in Ret tr;t a h an. r in a t ian I,ivili. (
u olt on idi l'rbana Unn t.t Ilhnoo Pn Uh s t R im i na.liat mn Pp 4.i b 4l
O M anes u.! 3'D 5 f a'ner A J R kinc i ,i
% u York Acadtno hisi I W ui t y. .\ le i nctn rourtmm . te me and inc abawir. .n the L bra i mr b ? ;.
, P pa rTruwfo Corular f
S tl- ? 4 U nik. R R J W Gkort &t,t Kat hhm tor <!, T > rav elat O! Nat flot ,; r s N.1 Lit. Ee U ' Miil A I IC u Clshh Httofti( Re id l- ' . ! m h e t:a o t w in t se (UnalihhKa4I 10 t Mbi, R A 107.1 lichas mr Pp 4 N utos \ m i . 4m i el t 'D
I arnrr & J R King 1q Ne u Ymt Acadenig Prn-yp . _ .
i
M6 lirastos l' Wi t Kl. Ju i\ut Voi a KtNDilGH. $ C lW Parental (are and it3 fielution in lords ll! lhol SIonogt .2 .- @
10 0 Energruu of tt production in birds dncumon Pp t il-lit in tirerdire 1-log 3 of i e-L
< D 5 F ather Li L hhington. D C Nat Atatt 5a Kisc. ) R 1W Entreetit s of reproductmn in tntd* Pp N 10' = lireeding tehr et tord. ,D 5 Farner.Id % a hingtori. D. C . Nat b a i Sa Kt. An E E 14?J A population Studs of the latterr> Phorbe ar.1 its no:ial erlatior Mp. uith the i llrow n ht a' led Com bird l'npubh hed Ph D thrin. L.tu rem e Kansas. U na of Kan..u i
14 L C a turd parautum and nc4tirig suuen in the Ea> tern 1%ebe Oc t Par t nn Kan ljus Nat 11.t 41 1 11 tw L D lati The natural rtgulation of anirnal nurnt ers London. Osford t'nn Prr..
W sc s t.. R .\1 iM Thr birds ol Kentut Ls Ornithol Monogt No 3 Mllpl81l Ti>N. D * & ll ) JOHNstos lo% The Mii h>xan Audabi.n so< irt> i. hor t r ,tm ! . rt 1 A 4.wh of ite phort.e m M.n omb Count 3 Jas k.Pme % arbirr 34 6.4-rA IW i e s m . J 1. & W D Ki r sn T ka 10:0 T he nestmg h olos3 and treroibutn r per " nan, r M tv f.a tvrn Mt.i salar k Wihon Etutt <2 la t- 147 Mi a n. A F ie Adaptn e hmitanon of the reprodutin e rate of bir l Ims im t- as l \ N } l 4 4 9 t \l 4. N [. l%I Clutt h mitt 11. thr hed k ls t alC ht r Irlif lnt rr r.at S h t ; h r il C . ,r r '4 [ 1.( .
'{ 164 1 --
l'.71 P wh3 tion ih namit i. Pp J L l - s ' 4 of A t ia n biolvo i nI I ill $ lorne r & J R Kirg
' ds New Y rk. k adrnut Ptr ,
% t i M.. Il P JP 19? Ni >t rriiprm oy m i a ttrn Phot br .an I 1:arn '.w aks. %. - P ?! * <
l0 r. t:-U n
%ith.J C 1e Iht b'r of lord, Phda le Iphia % li 'annder. Co i M inia u s. U \ 74 Phorbe and flarn u alhm m .t ng .trm tmt, 'i r a n. \! P. : %M
', Cont 3i **
l /u n hit l' ( l I:ne rgi tn of trprodu;' i i nn bird
tb u iP U n Ip M 1 11- N I' : f'i l o i 6
)- ofInnh li tarner I d > M a.hmet r it C Nat tai vi emer
>er%
asens tem --
h m
ne u...n 0,n,o a..we+,, m, a ms ne, o mdm idum, e,e ..,_
t he ,uurts f our+1:n i wt. t he m irn pndt. it u d in the n as bot h am.orut and emfr- 1 m
i
. .M-take an acn e pr .n n ory im, anon ana oper# .n Dui.or m m ,rrreutuar e . " - 1 %
t ude nts 1.de tr e?I rr hip, tan be obtained ur ' I il Dec ember 19'at ha .S r u f or .r + m n.ind M
.ipphcanon form, a rne Robert C % h:tmore Membership Ch.urman. B d.on OrM d with E" Dat ion of f oree % e t Viroma l'r+n er.in \lorg.u.to a n. West Viremia N 0 4 1t th m A \t Hipet*5titti of) Dirdt p(lbr hra and 'betre u nli be hrht at the Cnn tr*in Ut Ca;t I't 'od h og Afura, on 14 ? t .% ember 1979 Iloth forrr . -emon an t eu ursion, are plannel.snt' ed phg ged;ng glU be pub!bhrd as a tupplernent ! Ibc CUF8'tP8W Hf thulletin inf the 50uthr, 's ab'I I
'4 ',:W Group Perwns iraerestedin attend ng the sy rnt Nm shoul.l u rne to the Organising *ct retar3, \lr.G.
\!' .i l
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1 l [fnhrrsil3 of Cape Tow n,Itundcho.ch 7 700, South Africa, gmne deta h of their ; m e<! pater 1,
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